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.     

Effects of supplemental food on intra-and inter-specific behaviour of the Varied Tit Parus varius

Varied Tits Parus varius lived in flocks containing a pair, or a pair with one or two unpaired birds. To test whether the sociality of Varied Tits or their participation in mixed-species flocks is sensitive to additional food (sunflower seeds), we compared the following parameters between fed and unfed periods at the individual level: population size, degree of site fidelity, mono-specific flock size and the stability of its membership, home-range size and distribution, attendance rate with mixed-species flocks, and flock size. Neither the size of the mono-specific flock nor its membership was affected by food supply. Whether food was added or not, individuals exhibited a strong site fidelity resulting in stable population size. Supplemental feeding had no effect on home-range size or distribution. In the presence of extra food, Varied Tits were observed in mono-specific flocks but rarely in mixed-species flocks. However, when we stopped feeding, they shifted to mixed-species flocking. After adding food, mixed-species flocks were significantly smaller than in the control samples. Varied Tits were more likely to join mixed-species flocks as temperatures dropped and wind speed increased. These results suggest that intra-specific sociality of Varied Tits is relatively insensitive to food supply, but they easily shift to mixed-species flocking in relation to food and weather conditions. We conclude that Varied Tits participate in mixed-species flocks to obtain short-term benefits e.g. increased foraging efficiency but they also obtain long-term benefits from stability of pair bonds and strong site fidelity, which did not respond to supplemental food.     

Composition and foraging behaviour of mixed-species flocks led by the Grey-cheeked Fulvetta in Fushan Experimental Forest, Taiwan

Thirty-two species were recorded in mixed-species bird flocks led by the Grey-cheeked Fulvetta Alcippe morrisonia in Fushan Experimental Forest, Taiwan. Flocks averaged (± se) 5.8 ± 0.2 species and 51.4 ± 2.7 birds. Most participants were resident species (86.3%), some were elevational migrants (12.6%) and a few were latitudinal migrants (1.1%). Flock size was determined primarily by the abundance of Grey-cheeked Fulvettas, the most abundant species (68.1%). Flocks moved at an average rate of 10.8 ± 0.7 m/min, with larger flocks moving faster than smaller flocks. In moving flocks, canopy species were usually near the front, while understorey species usually followed. Fulvettas gave higher-intensity alarm calls and dived down more frequently in response to avian threats, especially raptors, than to non-avian threats. The overall foraging niche-breadth of the fulvetta was greater than that of any attendant species. Each species in a flock had a unique foraging niche. Most attendant species exhibited low foraging niche-overlap with the Grey-cheeked Fulvetta. Both the predator avoidance and the foraging efficiency hypotheses for mixed-species flocking were supported. The Grey-cheeked Fulvetta plays a critical role in the function of mixed-species flocks. A large flock formed around the Grey-cheeked Fulvetta provides attendant species with numerous opportunities for obtaining food and protection from predators.     

Effects of snow cover on the social and foraging behavior of the great tit Parus major

In the Joetsu region of central Honshu, Japan, snow lies on the ground 2–3 m deep from January to February every year. To test the effects of snow cover on the social and foraging behaviors of the great tit (Parus major Linnaeus) in the region, the following parameters were compared between the pre-snowy period (November and December) and the snowy period (January and February) at the individual level: population size, monospecific flock size and the stability of its membership, home range size, attendance rate with mixed-species flocks, and foraging height. Great tits lived alone or in pairs, but often joined mixed-species flocks. Throughout the study period, individuals exhibited strong site fidelity, which resulted in a stable population size. Neither the size of a monospecific flock nor its membership was affected by snow cover. The home range expanded when birds joined mixed-species flocks during both periods. Birds more frequently joined mixed-species flocks during the snowy period, and the size of mixed-species flocks was significantly larger than during the pre-snowy period. During the snowy period, birds shifted their foraging position from the ground to the upper parts of trees. They also used the upper parts of trees when they joined mixed-species flocks. These results suggest that the intraspecific sociality of great tits is relatively insensitive to snow cover, but that the home range size and foraging positions are affected by joining mixed-species flocks, rather than by snow cover.     

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.     

Flocking behaviour of foraging birds in a neotropical rain forest and the antipredator defence hypothesis

Randomly encountered foraging birds were recorded in a primary rain forest of French Guiana (13,550 records of 216 species), together with their size, diet and habitat use, to assess the relative frequencies of different types of flocking behaviour and some of their ecological correlates. Overall, 42% of birds foraged singly, primarily carnivores (raptors), nectarivores (hummingbirds) and lek-mating frugivores (manakins, some cotingas). For-aging in pairs (26.6%) was widespread, notably among insectivores in the 17–32-g size class. The remaining 31.4% of records were birds in groups of different composition and function, including, in order of decreasing frequency, (1) multispecies upper canopy flocks (83 member species identified)—the largest and most mobile associations of small insectivores, nectarivores and frugivores, mostly tanagers; (2) understorey mixed species flocks of small insectivores, at midlevels of closed forest interior, with 12 core, obligate members and 74 occasional species, mostly active foliage or bark gleaners and probers sharing a unique set of ecological characteristics; (3) monospecific groups (29 species), either gregarious foragers but solitary breeders (large frugivores in canopy) or also breeding colonially or several permanently group living cooperative breeders; (4) opportunistic gatherings of frugivores at fruiting trees (at least 40 species); (5) army ant followers near ground of closed understorey (29 species of mid- to large-sized insectivores); (6) followers of Red-throated Caracaras Daptrius americanus (23 species, usually canopy frugivores entering understorey with caracaras); (7) two raptors following monkeys. Attributes of vulnerability to predators defined by habitat structure (vegetation density or openness) and foraging behaviour (conspicuousness, speed, degree of vigilance) were important determinants of flocking propensity, at least in flocks that were not attracted by a particular food source. The results suggest that the permanent mixed-species flocks in the mature forest under-storey may be an antipredator defence to compensate for the conspicuousness and reduced vigilance resulting from active foraging behaviour in semi-open vegetation, where early detection of predators is difficult.     

Advantages of social foraging of Willow Tits Parus montanus

The mean number of Willow Tits Parus montanus in single-species flocks was significantly larger than in mixed-species flocks of Willow and Coal Tits P. ater. Both flock size and the tendency of Willow Tits to join mixed-species flocks were negatively correlated with ambient temperature, probably because each bird, when the metabolic rate of the birds increased, could allocate more time to foraging due to improved predator detection by many eyes. The vigilance time of Willow Tits decreased with flock size and was determined by the total number of individuals in a flock rather than by the number of Willow Tits in mixed-species flocks of Willow and Coal Tits.     

ADAPTIVE ADVANTAGES OF MIXED-SPECIES FEEDING FLOCKS AMONG SEED-EATING FINCHES IN COSTA RICA

The feeding ecology of three Costa Rican finches occurring in mixed flocks, Tiaris olivacea, Sporophila aurita corvina and Sporophila torquella , was investigated by measuring both behavioural and ecological variables. Observations on foraging height, rate of hopping and pecking rate, as well as the identity, proximity and number of nearest neighbours were recorded. In addition the duration of all feeding and perching episodes were timed. Comparisons were also made between the abundances of food items (grass seeds) consumed and those potentially available for consumption.
The analysis of variance of the feeding behaviour revealed that the presence or absence of neighbouring birds, whether of the same or different species, influenced the duration of feeding bouts more significantly than did either differences in habitat or species-characteristic behaviour. In addition the dietary comparisons revealed overlap in both species and size of seed consumed. Such similarities suggest that these species are not partitioning fields in the classical sense.
We propose that the increase in the duration of the feeding bout associated with the presence of mixed species aggregations leads to increased feeding efficiency and is the result of intra- and inter-specific social learning. Certainly flocking is often advantageous, since searching in a group facilitates finding clumped resources; mixed species flocking, by increasing exposure to a diversity of foraging places and patterns, can further augment feeding 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.     

Foraging strategy in a social bird, the alpine chough: effect of variation in quantity and distribution of food

I conducted feeding experiments to determine the effect of variations in food availability on individual flocking behaviour and foraging efficiency in a social corvid, the alpine chough Pyrrhocorax graculus, which lives in large flocks all year round. In 37 trials I varied both food quantity and the number of food patches. A decrease in the amount of available food reduced the mean flock size, the proportion of birds that had access to food, and their mean pecking rate. A decrease in the number of patches, on the other hand, reduced only the proportion of birds that had access to food. The number of choughs foraging was not influenced by food competition but depended only on the number visiting the site. Females competed less well than males: when food was made scarce, they frequented the site in the same proportion as did males, but had less access to food. I suggest that in this social corvid, long-term advantages to flocking related to social bonds, such as the maintenance of pair bonds, may compensate for short-term costs such as a reduction in foraging efficiency. Copyright 1999 The Association for the Study of Animal Behaviour.     

鄱阳湖围垦区藕塘生境小天鹅与白鹤的种间关系

鄱阳湖是小天鹅（Cygnus columbianus）和白鹤（Leucogeranus leucogeranus）极为重要的越冬地,它们均主要以沉水植物苦草（Vallisneriaspp.）冬芽为食,且通过觅食空间生态位分化减少种间竞争。近年来,鄱阳湖苦草冬芽锐减导致大量小天鹅和白鹤由自然生境转移到南昌五星白鹤保护小区的藕塘觅食。大量小天鹅和白鹤集中在小片藕塘觅食可能导致种间竞争强度增加。因此,本研究以五星白鹤保护小区藕塘为研究地点,于2021年11月10日至25日,采用瞬时扫描法和焦点动物法调查了藕塘与白鹤混群和不与白鹤混群时小天鹅的日间行为、单次取食时间和每分钟摄食成功频次,并采用单因素方差分析或Mann-WhitneyU检验对数据进行统计分析。结果表明,小天鹅日间行为主要以觅食（45.59%）、运动（17.05%）和休息（15.92%）为主。小天鹅和白鹤混群时的觅食行为比例和单次觅食时间显著高于不混群时,表明小天鹅主要通过增加觅食时间以应对种间竞争的负面影响,满足能量需求。小天鹅混群时的每分钟摄食成功频次显著高于不混群时,这可能是由于小天鹅通过摆动脚蹼或者扁平喙啄食的方式较难取食...     

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.     

Foraging strategy and predator avoidance behaviour: an intraspecific approach

Relationships between predator avoidance behaviour (scanning and flocking) and foraging were studied in Calidris alpina, to test predictions regarding the effect of foraging techniques on such behaviours. The scanning hypothesis predicts that individuals with a tactile hunting technique and individuals with a visual hunting technique (both continuous searchers) do not differ in any variable related to scanning behaviour. The flocking hypothesis predicts that visually hunting individuals witl tend to form smaller flocks than tactile-foraging individuals. The two continuous feeding strategies did not differ among individuals in vigilance rate, nor in vigilance time or mean scan duration. However, with respect to flocking behaviour, visual foragers differed from tactile foragers in foraging flock size. The relationships between flocking behaviour and foraging strategy are discussed. The pattern found at the intraspecific level are the same as those found at interspecific level.     

Relationships between frequency of mixed-species flocks, weather and insect activity in a montane cloud forest in Ecuador

The frequency of mixed-species flocks of birds, weather and insect activity were studied in a high-altitude cloud forest in Ecuador. It is generally accepted that participation in mixed-species flocks improves foraging efficiency. If this is true, more flock activity may be expected when food is less available, which may happen during long-lasting periods of rain and otherwise adverse weather conditions. The total number and mean size of flying insects decreased as rainfall increased. The number of flocks observed increased with decreasing number and biomass of insects. Relatively more flocks were seen during rain than during dry weather. No flocks stopped foraging during rain. The flock activity pattern appeared to be the opposite of that found in humid lowlands, probably because of different weather regimes in the two zones. In the lowlands, heavy rain is typically of short duration. In high-altitude cloud forest, the rainfall is often less intense but persists for prolonged periods. Hence, from an energy point of view, cloud forest birds cannot afford to stop foraging during adverse weather conditions when insect availability is low. The results suggest that some mixed feeding parties have evolved in response to low tropical insect availability, necessitating long feeding excursions outside the territory.     

斑块质量对大斑啄木鸟冬季觅食行为的影响

为了解大斑啄木鸟(Dendrocopos major)冬季对食物斑块的利用对策,2011年1月和2012年2～3月,在内蒙古乌拉特前旗的农田防护林中,采用目标动物取样法和全事件记录法,观察了大斑啄木鸟在食物斑块的觅食行为,利用主成分分析方法对斑块质量进行评价,通过比较不同质量斑块中大斑啄木鸟的觅食频次、停留时间、觅食成功频次及觅食成功率等指标,分析斑块质量对其觅食行为的影响。结果显示,在不同质量斑块中大斑啄木鸟的觅食频次、停留时间、觅食成功频次差异都极显著,但觅食成功率差异不显著;除停留时间外,不同性别间觅食差异不显著。大斑啄木鸟的觅食频次、停留时间、觅食成功频次与斑块质量呈显著正相关,觅食成功率与斑块质量相关性不显著。大斑啄木鸟倾向于在质量水平高的斑块觅食,表现为在这些斑块停留时间更长、往返次数更频繁;但觅食成功率不受斑块质量影响,这可能是大斑啄木鸟适应不同觅食环境的一种生存本能。     

The composition and foraging behaviour of mixed-species flocks of forest-living birds in Madagascar

Mixed-species flocks of forest living birds were investigated in a rainforest at Perinet/ Analamazaotra (Andasibe), Madagascar. Most insectivorous birds in the area participated in mixed-species flocks. Flocks were composed of foliage gleaners, foliage-branch gleaners, trunk gleaners and flycatchers. Species whose foraging techniques and foraging locations in vegetation were similar were different from each other in foraging heights. The nuclear species often occurred in small conspecific flocks in and out of mixed-species flocks.     

Immature gannets follow adults in commuting flocks providing a potential mechanism for social learning

Group travel is a familiar phenomenon among birds but the causes of this mode of movement are often unclear. For example, flocking flight may reduce flight costs, enhance predator avoidance or increase foraging efficiency. In addition, naive individuals may also follow older, more experienced conspecifics as a learning strategy. However, younger birds may be slower than adults so biomechanical and social effects on flock structure may be difficult to separate. Gannets are wide‐ranging (100s–1000s km) colonial seabirds that often travel in V or echelon‐shaped flocks. Tracking suggests that breeding gannets use memory to return repeatedly to prey patches 10s–100s km wide but it is unclear how these are initially discovered. Public information gained at the colony or by following conspecifics has been hypothesised to play a role, especially during early life. Here, we address two hypotheses: 1) flocking reduces flight costs and 2) young gannets follow older ones in order to locate prey. To do so, we recorded flocks of northern gannets commuting to and from a large colony and passing locations offshore and used a biomechanical model to test for age differences in flight speeds. Consistent with the aerodynamic hypothesis, returning flocks were significantly larger than departing flocks, while, consistent with the information gathering hypothesis, immatures travelled in flocks more frequently than adults and these flocks were more likely to be led by adults than expected by chance. Immatures did not systematically occupy the last position in flocks and had similar theoretical airspeeds to adults, making it unlikely that they follow, rather than lead, for biomechanical reasons. We therefore conclude that while gannets are likely to travel in flocks in part to reduce flight costs, the positions of immatures in those flocks may result in a flow of information from adults to immatures, potentially leading to social learning.     

Mixed-species flocking of forest birds on Little Barrier Island

Abstract

Three aspects of mixed-species flocking of forest birds on Little Barrier Island were investigated. Whiteheads, fantails, parakeets, and grey warblers occurred more often in flocks than in “non-flocking” situations. Whiteheads were the main lead species, although parakeets formed groups within flocks and occasionally appeared to lead. Whitehead clumps defined the flock centre; only fantails were found commonly in the centre with whiteheads. Birds other than whiteheads generally orientated below or to the side of their nearest neighbours. We suggest that mixed-species flocking is a significant factor influencing the structure of forest bird communities in New Zealand during winter.     

FLOCKING AND FEEDING IN THE GREAT TIT PARUS MAJOR–AN EXPERIMENTAL STUDY

Some aviary experiments designed to investigate the survival value of flocking in the Great Tit Parus major are described.
An individual searching for a source of food is less likely to find it if he is on his own or in a pair than when in a flock of four. This is a result of local enhancement: with more pairs of eyes searching, some member of the flock is likely to come across the food sooner than would a lone bird, and once one finds the food, others in the flock more or less immediately assemble at the food source. Once the birds have found a source of food, dominance interactions play a role in determining which individuals obtain the most food.
If one bird finds a single item of food, both it and the other members of the flock rapidly alter their searching behaviour, concentrating their attention in the general area and /or type of place where the food item was found.
Even if a bird searches in a place and does not find food, the others in the group show some increase in searching effort in the same area or type of place. The extent to which they do this depends on whether they are used to finding food in clumps or dispersed.
The relevance of these findings to birds in the wild is discussed. It seems likely that local enhancement does occur in natural flocks, and the flocking in Great Tits is of benefit to the participants because they increase their effectiveness in finding food. However, the question of the survival value of mixed-species flocks of titmice remains unresolved.     

Nahrungserwerb und Konkurrenz des Blutspechts,Picoides (Dendrocopos) syriacus

Summary The Syrian Woodpecker, Picoides syriacus, acquires its food by excavating, probing, gleaning and fly-catching (Fig. 1). The food-acquisition is carried out from ground level to the crown layer of trees and beyond (Fig. 2).In an ecological sense the Syrian Woodpecker occupies an intermediate position between the Great Spotted and the Middle Spotted Woodpecker (Picoides major and medius). That is true for food-acquisition since it excavates more than medius and less than major whilst probing more than major and less than medius. The species is also intermediate in the choise of base from which foraging is done. Great Spotted Woodpeckers are found frequently on thin twigs and Middle Spotted Woodpeckers on thick stems.Between Great Spotted and Syrian Woodpecker there exist interspecific territoriality. Both are neutral with respect to the Middle Spotted Woodpecker. In spite of strong competition for nesting holes and frequent aggression the territories of Syrian Woodpecker and Starling (Sturnus vulgaris) overlap. These findings are discussed.