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.     

The Effect of Within-Flock Spatial Position on the Use of Social Foraging Tactics in Free-Living Tree Sparrows

The benefit of producer (searches for own food) or scrounger (exploits the others’ food discoveries) foraging tactic in a group of socially feeding animals may depend on where the individual searches for food within the group. Scrounging may be more advantageous in the centre of the group, having more individuals around to join, while producing may be more beneficial at the edges, where more unexplored food patches may be found. This study shows within‐flock position correlates with foraging tactic use of feeding birds in socially foraging tree sparrows, Passer montanus. Sparrows staying closer to the centre of the flock found their food patches more frequently by joining (i.e. use more frequently the scrounging tactic) than those staying toward the edges. To our knowledge this is the first field study demonstrating the relationship between spatial position and foraging tactic use. We investigated this relationship under different perceived predation hazard, and found that under elevated risk of predation, central individuals may increase their use of joining more than individuals on the periphery of the flock. Moreover, we show that extremely specialized use of searching tactics may be very infrequent in tree sparrows. As both within‐flock position and search tactic use can be altered very quickly and without leaving the flock, individuals may easily alter them in order to adjust their behaviour.     

Factors affecting flock size in the Alpine Chough Pyrrhocorax graculus

The Alpine Chough Pyrrhocorax graculus is a social corvid which now uses food provided by tourist activities in mountain regions (e.g. at ski stations, refuse dumps, picnic areas). Foraging flock size, flock distribution and flock structure of Alpine Choughs were studied in the northern French Alps between 1988 and 1992. Flock size varied during the year, being larger in winter than in summer. Alpine Choughs were not evenly distributed in space, and their numbers depended principally on site-specific factors, such as seasonal human presence. A significant positive correlation existed between Alpine Chough foraging flock size and the local human population. Immature birds gathered in winter in the largest flocks and principally frequented sites with the greatest food availability (e.g. refuse dump). In summer, immatures dispersed, joined small breeding groups and were more homogeneously distributed.     

Foraging behaviour of individual coal tits,Parus ater,in relation to their age,sex and morphology

The aim of this study was to reveal the causes and consequences of individual differences in foraging behaviour of coal tits, Parus ater, on the island of Gotland in the Baltic. On Gotland the willow tit, P. montanus, and the crested tit, P. cristatus, are absent, while these larger species are abundant competitors on the mainland. The Gotland population of coal tits exploited parts of the foraging niches of their absent competitors. Gotland coal tits are bigger than those on the mainland; they have thus evolved towards the size of the missing competitors. Individual differences in niche use were related to their morphology. Large birds more often used the inner parts of trees whereas small individuals foraged more on needles in the outer parts of the trees. Also, individuals with wing shapes more suitable for hovering and slow flight foraged more on needles. It is argued that morphology affects the profitability of different foraging sites. However, for foraging site selection, age was more important than morphology. Older, and probably more dominant, individuals occupied the most profitable foraging sites.     

Seasonal dynamics of resource partitioning among foliage-gleaning passerines in northern Finland

Summary The five most abundant species were included in a year-round study with respect to six foraging niche dimensions. Approximately full multidimensional utilization functions were used for niche metrics. During summer foraging overlaps were invariably high, but in other seasons periodically lower. Foraging site breadth was lower in winter, when fewer sites are profitable for foraging than in summer. Feeding posture versatility, by contrast, was highest in winter. Seasonal foraging shifts were very prominent, as great in fact as between-species differences. Often seasonal trends were parallel in different species. Niche axes of macrohabitat-type (e.g. tree) were more open for foraging variation and axes of microhabitat-type (tree part) more rigid. Among the resident species seasonal variation in foraging was greatest in Regulus regulus and Parus cristatus, whereas the foraging behaviour was more stable for P. montanus (an abundant species with broad niche), perhaps owing to greater intraspecific competition. In these northern forests foliage-gleaners must be versatile generalists to cope with the unpredictable resources, and thus they overlap broadly in their general resource niches which are determined by their genetically fixed cost and benefit relations to each resource type. Anyhow, presumably during periodical food shortage, the actual resource uses adjusted by resource availability and competition may overlap narrowly.     

Space and habitat use by Red Kites Milvus milvus during winter in the Guadalquivir marshes: a comparison between resident and wintering populations

Differences in habitat selection, diet and behaviour of resident and wintering Red Kites Milvus milvus were studied in Donana National Park, southwest Spain. Adult resident Red Kites roosted at their nests, while immature residents and wintering birds gathered at communal roosts. Individuals remained on average for two consecutive nights (range 1–8) at the same communal roost. Wintering kites spent significantly more time foraging gregariously than residents. The use of the marsh was greater by wintering kites while residents more often used the forest. Wintering birds consumed more goose carrion than the residents. Wintering kites had larger core areas, moved farther from their roost sites to feed and changed foraging areas more frequently than residents. Among residents, adult females had the smallest home ranges (core area, distances travelled and time spent flying). We suggest that Red Kites wintering in Donana occupy the marsh because it provides high food availability, and is vacated by Black Kites M. migrans which exploit this profitable habitat during the breeding season.     

Use of winter habitat by roe deer at a northern latitude where Eurasian lynx are present

Winter climate at northern latitudes is a challenge to small-bodied ungulates, and they modify behaviour to save energy and to increase the likelihood of survival. Also, the ongoing expansion of large carnivores in several European countries can lead to the recovery of (potentially energetically costly) anti-predator behaviours. In an area recently recolonized by Eurasian lynx Lynx lynx , we snow-tracked radio-collared roe deer Capreolus capreolus in order to investigate their bedding and feeding behaviour during winter, and assess how environmental factors affect their habitat use. We also tested the prediction that roe deer use more open sites than locally available in areas with a stalking predator such as the lynx. Our results showed that both bed sites and foraging sites had more cover, compared with random sites. Most of the variation in canopy cover and in the distance and foraging sites between bed sites and foraging sites was explained by prevailing weather. As the winter progressed, the presumed depletion of fat reserves promoted the use of more canopy cover at foraging sites by night, less by day and a decrease in the distance between beds, foraging sites and human activities. Males used artificial feeding sites less often and bedded further from humans than females. The data fit the hypothesis of tighter energy budgets for family groups (females with fawns) or that males are more cautious towards humans. There was no support for the hypothesis that roe deer used more open habitat than locally available in order to reduce their vulnerability to lynx predation. Owing to severe winter conditions and the danger of starvation, roe deer seem to be forced to accept a high risk when predators are present, not changing their main pattern of habitat use from comparative areas where predators are absent.     

Effects of position and flock size on vigilance and foraging behaviour of the scaled dove Columbina squammata

Amongst the benefits of foraging in flocks are the enhancement of food finding and predation avoidance. Characteristics such as size, individual position, as well as position and distance between members are factors that may influence vigilance and foraging. In a study using scaled doves, Columbina squammata, I observed a negative correlation between group size and vigilance and a positive correlation with time spent foraging, which suggests a reduction of costs and an increase of benefits as a consequence of larger group sizes. Individual position in the flock appeared to be an important factor in this trade-off. Peripheral individuals were more vigilant and foraged less than central ones, suggesting an edge effect in flocks of this species. The clustering of conspecifics may be related with fast transmission of information. Overall, aggressive interactions were rarely observed; when registered, they occurred mostly in larger groups, suggesting an effect of interference competition. These results imply that predation may be a strong pressure on the scaled dove's flock formation and behaviour.     

Relation of group size and daily activity patterns to southern lapwing (Vanellus chilensis) behaviour

Behavioural patterns of birds commonly vary according to flock size and daily activity pattern. Southern lapwing behaviours and their relation with flock size were studied, as well as the relationship between the frequency of behaviours and the period of the day. Results showed that the proportion of time spent in foraging and vigilance was higher during the morning, when small groups were more common, and decreased from midday on, when group size increased. Maintenance and inactivity (sleeping) behaviours presented the opposite pattern. Correlation between flock size and period of the day, and their similar effects on bird behaviour may be evidence that groups of different sizes could have different functions throughout the day. Our results show that southern lapwings seem to form smaller flocks for feeding in the beginning of the day and larger flocks later for different activities (e.g., maintenance and sleeping). In this sense, it is possible that group size variation throughout the day is related to different demands for specific behaviours (functions) of groups, according to the daily activity patterns of the birds.     

Winter territoriality and flock cohesion in the black-capped chickadee Parus atricapillus

In northwest Massachusetts, black-capped chickadees shifted foraging groups repeatedly each day during the winter. As a result, there was no clear distinction between flock members and floaters, but instead a continuous gradation from birds that formed relatively stable associations to those that shifted groups more frequently. Group territories were absent and individual territories were at most poorly defended, if present at all. This picture differs from other recently studied populations, where flock membership is stable and flock territories are clear. Comparison with other populations indicates that winter social structure in the black-capped chickadee may be sensitive to both winter food levels and the proportion of year-round residents. This indicates that cost-benefit analyses of winter social organization should consider factors acting throughout the year as well as the residence status of the wintering birds. The data suggest that flock size, population density and home range size are sensitive to winter food on the study area, whereas territoriality and flock cohesion may respond more clearly to the local availability of breeding resources or the proportion of year-round residents.     

Foraging behaviour and home-range utilization in a suburban Badger (Meles tneles) population

The size and distribution of home ranges, and the dispersion of latrines, was investigated to describe the use of space and degree of territoriality within a population of Badgers ( Meles meles ) living in the suburbs of Bristol. The movement patterns of different age and sex classes were studied, along with seasonal changes in various measures of activity. Part of the study area was surveyed in detail to compare the Badger's use of space with a variety of habitat parameters, and to examine the predictability of their foraging behaviour. Many of the results are in contrast to earlier studies on rural Badgers. Range sizes were variable, and inconsistent over time, many group ranges were not contiguous and those that were overlapped. Territorial boundaries were not obvious and latrines were clumped around sett sites. Age- and sex-specific differences in movement activity were observed, along with an expansion of ranges in spring and a decrease in all forms of activity over the winter. Using multiple regression analysis, it was not easy to predict the foraging behaviour of suburban Badgers on the basis of 37 habitat variables sampled. There was some evidence of more structured foraging in the autumn, when more predictable food resources were available. The relevance of these observations to other studies on the ecology of Badger populations at varying densities and in different habitats is discussed.     

Characterizing complex mixed-species bird flocks using an objective method for determining species participation

We developed sampling methods to characterize the participation of bird species in foraging flocks led by the Eastern Tufted Titmouse (Baeolophus bicolor) in North-central Florida during winter, because standard field methods, developed primarily for permanent resident Neotropical flocks, were intractable in our system. During January–February 2004 and November 2004–March 2005, we observed 55 mixed-species flocks, recorded 40 potential flocking species [mean of 12.4 species (SD = 3.8; range 3–20), 26.3 individuals (SD = 12.2; range 8–60), and 3.1 titmice (SD = 1.4; range 1–7), per flock]. Twenty-six species were observed frequently enough (>10% of observations) to be included in analyses. We paired 60-min flock observations with 10-min point counts conducted in locations used by flocks, but after flocks had moved more than 100 m away. This method yielded a measure of flocking propensity: the ratio of the number of individuals observed in the flock versus during the point count for each species. We used regression tree (RT) analysis to classify species into groupings according to their levels of flock participation, and to investigate relationships between flocking propensity and various environmental and social factors that we measured. Our analysis identified three clear species groups; “Nuclear/Regular Associate” (12 spp.; high/moderate), “Occasional Associate” (four spp.; moderate/low), and “Non-joiner/Accidental” (ten spp.; low/no flocking propensity). Groupings were similar to schemes produced via more time-intensive field methods. In order to contextualize grouping categories, we conducted a review of flocking group definitions and relevant autecological information (e.g., interspecific sociality) about our study species. We found this method to be useful for geographically extensive sampling of species’ participation in mixed-species flocks, despite high inter-flock variability in species composition and limited labor.     

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.     

The social organization of non-breeding Magpies Pica pica

The social behaviour of non-breeding individuals in a colour-marked population of Magpies was studied. In early autumn most non-breeders began to forage in a common area, the 'Non-breeding Flock Area'. A few individuals remained on their natal territories away from the flock area as solitary non-breeders. Most non-breeders were first-year birds, but some were second years or adults which no longer held a territory. Birds foraged in groups, their food intake rate differing with both group size and location. It appeared that the area in which birds foraged had a significant effect on food intake rate; foraging groups tended to form at sites rich in food. There was a dominance hierarchy amongst non-breeders; an individual's foraging behaviour, survivorship and chances of breeding were status dependent, with subordinates feeding less in groups and being less likely to survive and breed. Solitary individuals' chances of breeding were similar to those of high status birds, although their survivorship to breeding age may have been lower. Non-breeding Magpies are compared with non-breeders of other species and the factors which may influence their social behaviour are discussed. It is suggested that remaining as a solitary non-breeder is a viable alternative to becoming a low-status flock member for some birds.     

Effect of increased food abundance near forest edges on flocking patterns of Coal Tit Parus ater winter groups in mountain coniferous forests

CapsuleIndividuals concentrated near forest edges in bigger social groups than in forest interiors and foraged more on pine cones which were more abundant there.

Aims To evaluate differences in food distribution between forest edges and forest interiors and their effects on the non-breeding flocking patterns of Coal Tit populations inhabiting mountain coniferous forests.

Methods We collected cone production data at forests edges and interiors in mountain pine forests located in the Pyrenees (northeast Iberian peninsula). At the same sites, we also quantified Coal Tit abundance, flocking patterns and foraging behaviour by means of paired bird surveys during autumn and early winter.

Results We recorded a larger abundance of pine cones available on trees along forest edges compared with forest interiors. Coal Tit groups were of bigger size along forest edges, although the number of social groups detected did not differ from forest interiors. Our observations on foraging behaviour supported the hypothesis that differences in flock sizes and overall abundances associated with distance to the edge are due to differences in the availability of pine cones and to the heavier use of these foraging substrates by birds along forest edges.

Conclusions Our results suggest that by changing food distribution, edge effects on pine cone production may be significantly involved in local changes in the social structure of the Coal Tit. An increase in resource heterogeneity and local population density may have important implications at a population level, such as favouring mobility of individuals searching for food resources and thus a transient life, and increasing the costs of territory defence to resident individuals.     

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     

Individual food-hoarding decisions in a nonterritorial coal tit population: the role of social context

Among the Paridae, food hoarding is thought to be strongly associated with the exclusive use of territories by winter groups, although it has also been described in populations with loose social systems. However, detailed data on such populations are scarce. To identify the mechanisms underlying individual storing decisions, I studied hoarding behaviour in a nonterritorial, high-density coal tit, Parus ater, population in a subalpine forest. The presence of close neighbours (within 5 m) had the strongest, negative influence on caching probability, whereas more distant neighbours foraging in the same flock did not affect the probability of caching. Adults concentrated their stores in the centre of their home ranges. Caching location, but not caching intensity, was affected by the time of day. Coal tits took longer, and travelled further, to cache food in the inner parts of trees than in the outer parts. Resident adults stored in inner, safer caching locations more often than juvenile residents and transients. Differences in hoarding effort according to the storing substrate, and the daily storing location pattern of juvenile residents and transients, suggest that stores have a different use depending on where they are located. Therefore, my results show that hoarding is compatible with a nonterritorial, nonbreeding social system; hoarding could thus have originated in an ancient, nonterritorial, but sedentary, tit species. Copyright 2000 The Association for the Study of Animal Behaviour.     

Foraging behaviour of non-breeding Western Sandpipers Calidris mauri as a function of sex, habitat and flocking

Individuals within a population may vary considerably in the way they exploit available food resources. If the sexes differ in the size of their feeding apparatus, there can be differences in foraging behaviour and habitat use, hence one sex may be more susceptible to competition. We examined relationships between sexual dimorphism in bill size and foraging behaviour, and habitat and microhabitat use of non-breeding Western Sandpipers Calidris mauri at Bahía Santa María, northwestern Mexico. Western Sandpipers are sexually dimorphic, with females about 15% longer-billed than males. Males used a higher proportion of pecks, had a lower probing–pecking rate, walked at higher rates, foraged at sites with lower water content, and had greater variation in foraging technique than females. Moreover, males decreased their proportion of pecks and foraged at a higher rate than females when they changed from feeding alone to feeding in flocks, suggesting a greater safety advantage or susceptibility to conspecific interference when flock feeding. We compared behaviour and microhabitat usage in three habitats: brackish flats, mangroves, and cattail marshes. Sex-related differences in foraging behaviour and microhabitat use were consistent among habitats. Birds in brackish flats and mangroves used a higher proportion of pecks, foraged at lower rates and walked at higher rates, and foraged at deeper sites, with a lower proportion of water cover, than those in cattail marshes. Sex-related differences in foraging behaviour and microhabitat should reduce the level of competition between sexes, and may account for sex differences in Western Sandpiper distribution observed between habitats in Bahía Santa María.     

Resting site selection by large herbivores – The case of European bison (Bison bonasus) in Białowieża Primeval Forest

Resting site selection by European bison (Bison bonasus, L., 1758), the largest terrestrial mammal of Europe, was studied in the free-ranging population in Bia?owie?a Primeval Forest (Poland) in 2009–2010. In total, 104 sites of 21 bison (both collared and uncollared) were analysed to determine the most important microhabitat characteristics selected by resting bison during summer and winter and to study the influence of supplementary feeding on resting behaviour of this herbivore. Resting sites were identified on the basis of GPS locations and activity records collected by GPS collars, as well as direct observations of bison, and were compared with control sites. Microhabitat selection by bison did not differ significantly between the sexes. During summer and winter, bison resting sites displayed a high tree density, low visibility and high complexity (structures providing cover). Summer resting sites were also characterised by a significantly lower abundance of blood-sucking insects and denser canopy than control sites. Winter resting sites showed a lower complexity and higher visibility than summer sites, and were less often located in mixed forest habitats. During winter, bison rested more frequently in forest below 50 years of age than in older forest. Resting sites of non-fed bison were more often located in young coniferous forests, were lower in visibility and situated closer to open areas than sites of bison using supplementary winter feeding, suggesting a trade-off between food and cover. The results indicate that European bison select their resting sites in areas of mosaic habitat structure providing cover from disturbances with access to profitable natural forage grounds.     

Number of neighbors instead of group size significantly affects individual vigilance levels in large animal aggregations

The group size effect states that animals living in groups gain anti‐predator benefits through reducing vigilance levels as group size increases. A basic assumption of group size effect is that all individuals are equally important for a focal individual, who may adjust its vigilance levels according to social information acquired from them. However, some studies have indicated that neighbors pose greater influences on an individual's vigilance decisions than other group members, especially in large aggregations. Vigilance has also been found to be directed to both predators (anti‐predation vigilance) and conspecifics (social vigilance). Central individuals might rely more on social vigilance than peripheral individuals. To test these hypotheses, we examined the effects of flock size, number of neighbors and position within a flock on vigilance and competition of greater white‐fronted goose Anser albifrons that form large foraging flocks in winter, controlling the effects of other variables (group identity, winter period and site). We found that individual vigilance levels were significantly affected by number of neighbors and position within a flock, whereas flock size showed no effect. Individuals devoted a large component of vigilance to nearby flock mates. Central individuals directed a relatively larger proportion of vigilance to monitor neighbors than peripheral ones, indicating that central individuals more relied on social information acquired from neighbors, possibly caused by the more blocked visual field of central individuals. Moreover, some social vigilance may function as conducting or preventing agonistic interactions since competition intensity was positively correlated with number of neighbors. Our study therefore demonstrate that the number of neighbors is more important than group size in determining individual vigilance in large animal groups. Further studies are still needed to unravel which neighbors pose greater influence on individual vigilance, and the factors that influence individuals to acquire information from their neighbors to adjust vigilance behaviors.