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.     

Role of mixed‐species flocks in the use of adjacent savannas by forest birds in the central Cerrado,Brazil

Abstract: We examined the role of mixed‐species flocks for forest birds during their breeding and non‐breeding seasons in the use of savannas adjacent to forests in central Cerrado, Brazil. Transect surveys (n = 64) were conducted in eight savanna patches. Distances of birds from forests were estimated. Recorded birds were classified as members or not of mixed‐species flocks. About half of the bird species recorded in savannas were found in at least one mixed‐species flock. As distance from the forest increased, the number of species in mixed‐species flocks tended not to vary, while the number of species foraging alone or in mono‐specific groups decreased. Thus, for some forest species, participation in mixed‐species flocks allowed a greater use of more distant savannas. This tendency of being in mixed‐species flocks at greater distances from forests also can be interpreted as a reluctance to forage alone or in mono‐specific groups due to higher predation risk in less protective vegetation distant from cover. There was strong seasonal variation in the participation of bird species in mixed‐species flocks. There were significantly more species in mixed‐species flocks than out of these associations in the non‐breeding season, while differences in the breeding season were not significant. These patterns occurred, in part because mixed‐species flocks tended to be more frequent, to have more species and to forage at greater distances from forests during the early non‐breeding season than in other periods. This study suggests that the formation of mixed‐species flocks plays an important role in promoting the use of adjacent savannas by forest birds at forest/savanna boundaries in Cerrado. It also pointed out a novel advantage gained by birds with participation in mixed‐species flocks – greater use of adjacent vegetation patches.     

Intraspecific differences in benefits from feeding in mixed-species flocks

The Madagascar Paradise Flycatcher Terpsiphone mutata and Common Newtonia Newtonia brunneicauda frequently form two-species flocks in the deciduous dry forest of western Madagascar. In T. mutata , some males have long tails, while other males and females have short tails. When foraging in mixed flocks, each type of bird captured prey more rapidly than otherwise, but the degree of increase in feeding rate was smaller in long-tailed males. When in mixed flocks, all T. mutata caught prey on leaves in the canopy where N. brunneicauda foraged. Long-tailed males changed feeding habits from sallying when not in mixed flocks, whereas short-tailed birds showed no change of feeding habit. The elongated tails of long-tailed males may have made their foraging less efficient owing to decreased agility in the canopy. N. brunneicauda is monomorphic and often formed groups of three to five individuals. In monospecific flocks, subordinates fed at low rates on branches owing to frequent hostile encounters. When foraging in mixed flocks, however, subordinates foraged among leaves, and their feeding rates increased because the frequency of intraspecific interference decreased greatly. Dominants did not show any difference in feeding pattern with social situation. Thus, heterospecific flocking was more advantageous for subordinates.     

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.     

Mixed species flocking of tits (Parus spp.): a field experiment

Summary We tested two general models of flocking behaviour, namely the antipredation model and foraging efficiency model on mixed-species tit flocks (Parus spp.). After food addition the size of mixed-species flocks was significantly less than in the control samples. In the presence of extra food significantly more birds were observed either in monospecific flocks or solitary, than during the control observations. In the presence of a living predator the birds foraged in larger mixed-specifies flocks than during the control observations. In addition, the social behaviour of Great Spotted Woodpecker, Middle Spotted Woodpecker and Nuthatch shifted to mixed-specific flocking. The size of monospecific flocks was independent of both treatments. The density of birds increased significantly after food addition, while in the predator presence the birds tended to leave the forest. These results support the view that both the antipredation model and foraging efficiency model seem to be valid for mixed-species flocking. However, in the case of monospecific flocks, the territory maintenance could be the most important factor.     

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.     

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.     

Linking vegetation structure and bird organization: response of mixed-species bird flocks to forest succession in subtropical China

As forests undergo natural succession following artificial afforestation, their bird assemblages also change. However, interspecific avian social organization associated with forest succession has not been fully understood, particularly for mixed-species bird flocks. To disentangle how mixed-species flocks change as a function of local forest structure, we analyzed flock characteristics (particularly species richness, flocking frequency and propensity) and vegetation physiognomies along a presumed successional series (early, middle, and advanced) simultaneously in subtropical forests in southern China. As hypothesized, monthly point counts demonstrated that complexity of flocks increases with the progression of natural forest succession at a local scale. Advanced forests differed significantly from pioneering plantations with respect to vegetation structure, flock characteristics and constituents (especially for understory specialists). Importantly, forest succession affected flock patterns particularly in relation to the flocking propensity of regular species, and the frequency of nuclear species (Huet’s fulvetta Alcippe hueti), which in turn determined flocking occurrence at different successional stands. Canonical correspondence analysis indicated that understory flocking species (mainly Timaliidae babblers) were significantly associated with intact native canopy cover, complex DBH diversity, as well as high densities of dead trees and large trees, representing a maturity level of successional stands. Our study reveals that the effect of natural forest succession on mixed-species bird flocks is species-specific and guild-dependent. From a conservation perspective, despite a high proliferation of pine plantation in southern China, priority should be placed on protecting the advanced forest with a rich collection of understory flocking specialists.     

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 evolution of communal roosting in birds: origin and secondary losses

Three main benefits are thought to underlie communal roostingin birds: a reduction in thermoregulation demands, a decreasein predation risk, and an increase in foraging efficiency. Iinvestigated interspecific variation in communal roosting tendenciesacross categories of several ecological factors to examine therelevance of each functional hypothesis in the evolutionary transitionto communal roosting and the secondary reversal to solitary roostinghabits. The study phylogenetic tree included 30 families and437 species. Evolutionary transitions to communal roosting occurredmore often on branches with flocking species and with largerspecies but were not associated with diet, territoriality, geographicalarea, or time of day. The association with flocking activitiessuggests that increased foraging efficiency, a factor thoughtto operate through the formation of flocks, may have been akey factor in the origin of avian communal roosting. However,several transitions to communal roosting occurred on brancheswith nonflocking species, indicating that foraging efficiencymay not be the only factor involved in the evolution of communalroosting. Secondary losses of communal roosting habits occurredon several branches, with a concomitant loss of flocking behaviorand a tendency to exhibit territorial behavior and nocturnalforaging. Secondary losses suggest that communal roosting iscostly to perform and maintain and may be lost when an asocialselection regime operates. The large number of exceptions tothe above patterns may force a reevaluation of current functional hypothesesabout communal roosting in birds.     

Behavioural and hormonal effects of social isolation and neophobia in a gregarious bird species, the European starling (Sturnus vulgaris)

Separating gregarious individuals from their group members often results in behavioural and physiological changes, like increased levels of corticosterone. Testosterone and corticosterone, in particular, have been implicated in the response of mammals to novelty. Data in birds are, however, rare. The presence or absence of group members may also influence an individual's response to novel stimuli. We assessed the behaviour and hormonal response of European starlings (Sturnus vulgaris) to a novel object in two different situations and seasons: each starling was tested when separated and when in contact with its group members in May/June (breeding season) and again in September/October (non-breeding season). Starlings are gregarious throughout the year, but as foraging flocks are small during the breeding season and large during the non-breeding season, we assumed that non-breeding starlings would be more affected by social isolation. Overall, starlings had higher levels of corticosterone, lost more body mass, and were more active when they were separated from their group. Isolated individuals, however, did not show a greater neophobic response than individuals in the presence of their group members in either season. Circulating levels of testosterone and corticosterone were higher after a test with novel object than after a test with only the familiar feeding dish in both sexes and seasons. However, control tests for handling effects confirmed only the increase in testosterone. Our study shows that social isolation is stressful for unrelated and unpaired members of a wild flocking bird species and demonstrates that novelty can lead to a rise in testosterone in birds.     

THE FORMATION, STRUCTURE AND FUNCTION OF MIXED-SPECIES INSECTIVOROUS BIRD FLOCKS IN WEST AFRICAN SAVANNA WOODLAND

Mixed-species flocks of birds were observed during the wet season (July to September 1975) in savanna woodland in Ghana. Thirty-four flocks contained birds of 56 species in 20 families, including insectivorous, granivorous, and nectarivorous species, using a wide range of foraging methods. Only two species occurred in more than half the flocks. There was no correlation between the number of flocks joined by a species and its abundance in the community. Among insectivores, but not granivores, the species which joined most flocks were those which habitually occurred in the largest single-species groups. All stages of breeding activity were represented by the various members. Some species joined flocks only while these were passing through their territories. Of the two species which were present most frequently, there were no differences between mixed and single-species flocks for Eremomela pusilla, but Parus leucomelas foraged and called on more occasions in mixed flocks than single-species flocks, though the rates of foraging and calling were related only to the number of P. leucomelas present. Groups of P. leucomelas appeared to initiate some flocks by attraction due to their conspicuous wing-bars, active movement, and loud calls. Black-and-white species joined them first, followed by birds of other plumage patterns. The advantages of mixed flocking are thought to be connected with finding patches of the food of bark- and foliage-searching insectivores, which were the only species regularly seen foraging in the flocks. Because of dry season burning which leaves small unburnt patches of savanna, these insect species may share a common, patchy distribution. Birds may also gain protection from predators, and some species probably gain no advantages. The species composition and behaviour of flocks previously recorded elsewhere in African savannas are similar to the Ghana flocks.     

Exclusion of deer affects responses of birds to woodland regeneration in winter and summer

Using an exclosure experiment in managed woodland in eastern England, we examined species and guild responses to vegetation growth and its modification by deer herbivory, contrasting winter and the breeding season over 4 years. Species and guild responses, in terms of seasonal presence recorded by multiple point counts, were examined using generalized linear mixed models. Several guilds or migrant species responded positively to deer exclusion and none responded negatively. The shrub‐layer foraging guild was recorded less frequently in older and browsed vegetation, in both winter and spring. Exclusion of deer also increased the occurrence of ground‐foraging species in both seasons, although these species showed no strong response to vegetation age. The canopy‐foraging guild was unaffected by deer exclusion or vegetation age in either season. There was seasonal variation in the responses of some individual resident species, including a significantly lower occurrence of Eurasian Wren Troglodytes troglodytes and European Robin Erithacus rubecula in browsed vegetation in winter, but no effect of browsing on those species in spring. Ordinations of bird assemblage compositions also revealed seasonal differences in response to gradients of vegetation structure generated by canopy‐closure and exclusion of deer. Positive impacts of deer exclusion in winter are probably linked to reduced thermal cover and predator protection afforded by browsed vegetation, whereas species that responded positively in spring were also dependent on a dense understorey for nesting. The effects on birds of vegetation development and its modification by herbivores extend beyond breeding assemblages, with different mechanisms implicated and different species affected in winter.     

Long‐Distance Calling by the Willow Tit,Poecile montanus,Facilitates Formation of Mixed‐Species Foraging Flocks

The occurrence of mixed‐species foraging flocks is a worldwide phenomenon in terrestrial bird communities. Previous studies suggest that individuals participating in flocks might derive benefits in terms of improved feeding efficiency and/or reduced risk of predation. However, very little is known about how individuals establish mixed‐species flocks. Here, I provide the first experimental evidence that long‐distance calling by the willow tit, Poecile montanus, facilitates the establishment of mixed‐species flocks at a foraging patch. Observations at experimental foraging patches showed that willow tits that find a food source produce long‐distance calls, particularly when they are isolated from conspecific flockmates. The probability of long‐distance calling was negatively correlated with the number of heterospecific foraging individuals near the food source. A playback experiment confirmed that calls attract both conspecific and heterospecific members of foraging flocks. This study demonstrates that willow tits use long‐distance calls to attract conspecific flockmates to foraging patches, and these calls can also facilitate the formation of mixed‐species flocks on patches.     

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.     

The formation of “mega‐flocks” depends on vegetation structure in montane coniferous forests of Taiwan

A mixed‐species bird flock is a social assemblage where two or more bird species are moving together while foraging and might benefit from increased foraging efficiency and antipredator vigilance. A “mega‐flock,” which includes flocking species from different vegetation strata, often exhibits high species diversity. Mechanisms for the formation of mega‐flocks have not yet been explored. In this study, we evaluated the influence of vegetation structure and bird species diversity in driving the occurrence of mega‐flocks. We investigated the composition of mixed‐species flocks, local bird communities, and vegetation structure in five vegetation types of two high‐elevation sites in central Taiwan. Mega‐flocks occurred more frequently in pine woodland than later successional stages of coniferous forests. However, species richness/diversity of local bird communities increased along successional stages. Therefore, vegetation variables exhibit more influence on the occurrence of mega‐flocks than local bird communities. Besides foliage height diversity, understory coverage also showed positive effects on flock size of mixed‐species flocks. Our results indicated that pine woodlands with more evenly distributed vegetation layers could facilitate the interactions of canopy and understory flocks and increase the formation of mega‐flocks and thus the complexity of mixed‐species flocks.     

Evidence for facilitation among avian army‐ant attendants: specialization and species associations across elevations

Mixed‐species assemblages can involve positive and negative interactions, but uncertainty about high‐value patchy resources can increase the value of information sharing among heterospecific co‐foragers. I sampled species composition of bird‐flocks attending army‐ant raids in three adjacent elevation zones in Costa Rica, across multiple years, to test for positive and negative associations among raid‐attending bird species. My goal was to test whether the most frequent and specialized raid‐attending species showed evidence of facilitating or excluding other bird species. I quantified elevational variation in avian community composition at raids, then asked whether species composition was associated with variation in flock characteristics (flock size and species richness). I identified the most frequent raid‐attending species (those that attended raids most frequently relative to their mist‐net capture rates), and bird species that performed specialized army ant‐following behavior (bivouac‐checking, which allows birds to memorize and track mobile army‐ant colonies). There was significant turnover of bird species among zones (including the frequent and specialized attendants); patterns of species overlap suggested a gradual transition from a Pacific‐slope to an Atlantic‐slope raid‐attending bird fauna. Raid‐attendance frequency was positively correlated with bivouac‐checking behavior. With few exceptions, the most frequent raid‐attending bird species, and the bivouac‐checking species, also participated in the most species‐rich flocks. High species‐gregariousness suggests many of the frequently attending and/or bivouac‐checking species functioned as core flock members. However, some bird species pairs were significantly negatively associated at raids. Despite species turnover, per‐flock numbers of birds at raids did not differ among geographic zones, but flocks on the Pacific‐slope were heavier because larger bodied bird species attended raids. Previous studies showed that the size (biomass) of bird‐flocks corresponds to the amount of food the birds kleptoparasitize from ant raids, and the heavier Pacific‐slope bird‐flocks could have greater negative kleptoparasitic impacts.     

Vigilance towards raptors by nuclear species in bird mixed flocks in a Brazilian savannah

A nuclear or leader species is the one around which foraging activity is organized. In the campo-cerrado (Brazilian savannah) up to four bird species (Saltator atricollis, Cypsnagra hirundinacea, Mimus saturninus, and Neothraupis fasciata) may function as nuclear or leader species in mixed species flocks. The aim of this study was to assess the features shown by these nuclear species. I quantified parameters of sociality, communication and alertness of nuclear bird species in mixed flocks with different composition. Parameters related to sociality (mean intraspecific group size) and communication (frequency of contact calls) were not correlated with the leadership. On the other hand, the most alert species was in the front of a given mixed flock most of the time. The leader species spent more time in vigilance and gave most alarm calls due to approaching raptors earlier. The results of this study strongly suggest that the alertness of a species is the major character of nuclear bird species in mixed flocks of the campo-cerrado.     

An appraisal of seed enumeration and videographic techniques for determining seed removal rates by birds

We examined the efficacy of seed enumeration and videographic techniques for determining seed removal by birds from indigenous (Chrysanthemoides monilifera and Olea europaea subsp africana) and alien (Lantana camara and Solanum mauritianum) shrubs at different study sites in the Cape Floristic Region. The seed enumeration technique involved counting the numbers of fruits and associated seeds removed monthly by birds, excluding those naturally abscised, from the shrub canopy. The videographic technique involved visual counts from images of the numbers of fruits and associated seeds consumed by birds over specific time intervals captured by a digital camcorder. Daily seed removal rates by all birds, irrespective of species, measured by both techniques were similar with no significant interactions evident between measuring techniques, site and shrub species. Both techniques displayed higher seed removal from tiny‐seeded S. mauritianum than other shrub species; this was also evident among individual bird species. However, the seed enumeration technique was unable to discriminate between foraging organisms, contamination of traps by wind‐blown fruits abscised from neighbouring branches and fruit theft from the canopy and the traps. In contrast, the videographic technique provided permanent visual and time‐lapse records for individual foraging bird species allowing greater measurement precision and interpretation of fruit removal behaviour by birds. We recommend use of the videographic technique over the seed enumeration technique for studying vertebrates’ seed removal in a detailed manner.