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.     

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.     

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.     

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.     

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.     

Effect of Group Size on Feeding Rate when Patches are Exhaustible

One benefit of group foraging is that individual foragers can join the food discoveries of companions and thus increase encounter rate with food patches. When food patches are exhaustible, however, individual shares of each patch will decrease with group size negating the effect of increased encounter rate. Mean feeding rate may actually decrease with group size as a result of aggression or time wasted joining already depleted patches, or when searching to join the food discoveries of others, which is referred to as scrounging, precludes finding food. I examined the relationship between mean feeding rate and group size in captive flocks of zebra finches (Taenopygia guttata) foraging for small clumps of seeds. Finches in groups of two or four fared better than solitary birds in terms of mean feeding rate despite the fact that birds in groups scrounged a large proportion of their food. Solitary birds initiated feeding activity after a longer delay, which led to their lower success. Early departures by food finders from food patches joined by others may have lessened the impact of scrounging on mean feeding rate. As a result of benefits from the presence of companions, group foraging in zebra finches appears a viable alternative to foraging alone despite the cost of sharing resources.     

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.     

The effect of human disturbance and flock composition on the flight distances of waterfowl species

Flocking bird species tolerate an approaching human up to a certain distance. We measured this distance, i.e., flight distance, to an approaching small boat for 11 waterfowl species. The flight distances correlated positively with flock size and species diversity (Shannon index H′) in species that showed relatively short flight distances when they were in a single-species flock. However, we did not observe such a correlation for single-species flocks that showed relatively long flight distances. Only pochards (Aythya ferina), a species with large individual variation in flight distances, showed a positive correlation between flight distance and flock size in both single- and multispecies flocks. Flight distance seemed to be affected by usage of the water area: flight distances tended to be longer for waterfowl species that use a water area for foraging than for those that use it primarily for resting. Thus, the behavior of actively foraging species may be more affected by human disturbances than that of resting species. Received: March 10, 2001 / Accepted: May 22, 2001     

ABUNDANCE AND SEASONALITY OF THE SAVANNA AVIFAUNA AT PORT MORESBY, PAPUA NEW GUINEA

The results of 156 transect counts carried out over 30 months in savanna of the Coastal Hill Zone, Port Moresby, Papua New Guinea, are analysed. The avifauna is depauperate compared to similar habitats in New Guinea and temperate Australia. Twelve common species of the Port Moresby savannas were not so in the Coastal Hill Zone and reasons for their rarity are given. Most species present are solely savanna forms but some are dependent upon the presence of small patches of monsoon forest. Average density of birds was estimated at 55 per 10ha, a very low figure compared to New Guinean rainforests. Carnivores and mixed-feeders make up two thirds of numbers present but frugivores, because of their large individual size, are a significant proportion of biomass. There was little significant seasonal difference in overall numbers except for granivores which decline in numbers during the wet season. However, individual species show regular seasonal fluctuations reasons for which are postulated. A correlation exists between flowering of trees and numbers of certain species, and also between the annual grass-fires and numbers of certain grassland species. Merops ornatus, Lalage sueurii and Coracina novaehollandiae, considered in the literature to be wintering migrants, are also shown to be breeding residents. Breeding records for 462 species were analysed. Most breeding of insectivores and mixed-feeders occurs in the Austral spring during the dry season and ends before the onset of heavy rains. Granivores breed in the late wet season. The few records for frugivores indicate a wet season peak.     

The ecology of mangrove forest birds in Peninsular Malaysia

The density, distribution and feeding ecology of birds of mangrove forests in Selangor on the west coast of Peninsular Malaysia were investigated between February and May 1989. Excluding aerial hunters and wading birds, 47 species were recorded from four mangrove forest sites in Selangor, though undoubtedly some rarer species were missed due to the brevity of the study. The estimated density of birds, using fixed-width transects, ranged from 15 to 26 birds per ha. One quarter of all birds counted belonged to just one species, the Ashy Tailorbird Orthotomus ruficeps. Excluding ground feeders (three species), four foraging guilds were recognized among 17 of the commonest bird species: obligate foliage-foraging insectivores (seven species), bark-foraging insectivores (four), facultative nectar-ivores (five) and aerial hawkers (one). Species within the first and third guilds apparently were partitioned by foraging technique and heights, while those of the second guild (woodpeckers) differed mainly in their selection of plant species. The horizontal distribution of several species corresponded to the zonation of mangroves. Avifaunal differences between sites reflected differences in forest floristics. The greatest number of species occurred at the dry landward edge site, which supported seven bird species not encountered elsewhere in Selangor. This habitat has almost completely disappeared from much of the peninsula owing to land reclamation. Four of the six resident mangrove-specialized bird species in Selangor were regularly recorded at two or more sites, another (Greater Gol-denback Woodpecker Chrysocolaptes lucidus) occurred at only one site and the last (Ruddy Kingfisher Halcyon coromanda) was never recorded. Two species of sunbirds probably play an important role in the pollination of several species of mangroves. The colonization of anthropogenic habitats by many mangrove-dwelling bird species is attributed partly to their generalized foraging niches, which may have evolved in response to regular fluctuations in food availability and the dynamic physiography of mangrove communities.     

Observations of mixed-species bird flocks at Kichwa Tembo Camp,Kenya

Mixed-species foraging flocks were studied at Kichwa Tembo Camp on the edge of the Masai Mara National Reserve in Kenya between July and September 2004. Observations were made on 29 mixed-species flocks, in which 24 species participated. African Paradise-Flycatcher Terpsiphone viridis, Black-backed Puffback Dryoscopus cubla, Grey-backed Camaroptera Camaroptera brachyura, Collared Sunbird Hedydipna collars and Cabanis's Greenbul Phyllastrephus cabanisi were the most common participants in mixed-species flocks, as well as among the most frequently encountered bird species overall. The Black-backed Puffback was identified as the nuclear species in flocks due to their abundance and frequency with which they were followed by other species. Mixed-species flocks represent another niche dimension in this diverse bird community, but few of these species could be described as flock specialists; most of the birds observed in mixed-species flocks in this study were opportunistic attendant species, including the African Pygmy-Kingfisher Ispidina picta, not previously described as joining mixed-species flocks.     

Flock size and habitat-dependent food and energy intake of foraging Goldfinches

Summary During the breeding season Goldfinches (Carduelis carduelis L.) feed on milky ripe seeds of about 20 food plants. Individual Goldfinches joining a flock reduce the time spent vigilant with increasing flock size. Therefore birds feeding in flocks get an increased intake of kernels per time unit. This was measured for five different food plants (Dactylis glomerata (Gramineae), Knautia arvensis (Dipsacaceae), Senecio vulgaris, Taraxacum officinale, Tragopogon pratensis (Compositae)). In large-sized flocks, birds fed up to 2.3 times more kernels, than when feeding solitarily. In addition, visibility in the vegetation leads to a further increase of kernel intake. Thus feeding under good conditions as in recently mown areas, can raise kernel intake to the seven fold per time unit as compared to solitary feeding. The maximum ingestion rate of kernels was 98 per min which implies a head up-and-down movement every 0.6 s. The calculated energy intake of birds per time unit is lowest in Senecio and highest in Tragopogon. Thus the birds, when feeding on Tragopogon in larger flocks, can gain an energy intake 16 times higher than that reached when feeding on Senecio, despite of a higher kernel intake rate. The energy intake individual Goldfinches gain at the particular plant species is markedly increased with flock sizes up to eight birds, with larger flocks the intake increases only slightly.     

Can spatial and temporal food variability explain the winter foraging movements of a threatened saltmarsh insectivore?

The white‐fronted chat (Epthianura albifrons) is a small, insectivorous passerine that is threatened with extinction in the north‐eastern part of its range, partially due to loss and degradation of its saltmarsh habitat. Food availability is a potential limiting factor for the disjunct populations that survive in saltmarsh refugia, surrounded by urbanized land, because the anthropogenic matrix reduces the capacity of birds to commute to alternative grassland habitat to exploit temporary insect outbreaks. This limitation is likely to be exacerbated during the winter months when local arthropod abundance in saltmarsh is reduced. This study measured temporal and spatial variation in the abundance of saltmarsh arthropods to determine whether patch switching by foraging flocks can be explained by variation in food availability. Arthropods in the size range known to be important in the diet of white‐fronted chats were vacuum‐sampled from six patches within a continuous area of Sarcocornia‐dominated saltmarsh over a four‐month period. The location of foraging birds was recorded during the same period. Despite superficial similarity in vegetation composition and structure, there was significant variation in arthropod biomass among sites through time, such that the patches with the highest food abundance changed from month to month. There was little evidence, however, to suggest that white‐fronted chats foraged in saltmarsh patches with the highest overall food abundance. During the course of the study, birds were discovered flying 2 km from the saltmarsh to a development site where they foraged in weedy grassland. Arthropod samples collected from this site contained an extremely high abundance of Hemiptera and Neuroptera larvae, supporting previous research indicating that white‐fronted chats forage on irruptions of particular arthropod taxa. These findings indicate that food abundance is unlikely to be the main determinant of foraging site selection within saltmarsh, but highlights the potential importance of alternative foraging habitat types for this species.     

Low Foraging Success of Semipalmated Sandpipers at the Edges of Groups

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

Experimental manipulation of termites (Isoptera,Macrotermitinae) foraging patterns in a Sahelo-Sudanese savanna: effect of litter quality

Summary The foraging pattern of four sympatric fungus-growing termite species was followed for five months of the dry season in a semi-arid savanna (Senegal). Experimental plots were established with and without termites (exclusion with insecticide) and amended with different litter quality: Acacia leaves, millet canes, ground millet (<500 µm), Combretum wood and cattle manure. Results showed different foraging patterns between the fungus-growing species. Odontotermes nilensis, for example, rapidly developed its foraging activity on the litter. In contrast, Macrotermes subhyalinus delayed its foraging activity but, extensively collected the food after its discovery. Differences in food preference were evident between the termite species: Odonto-termes nilensis foraged preferentially on millet, canes and Acacia leaves, Macrotermes subhyalinus preferred cane and ground millet. Ancistrotermes guineensis selected the millet, and Microtermes sp. was mostly recorded foraging on pieces of dead wood. Relationships between different foraging parameters were established to assess the relative intensity of this activity over the dry season. The comparison between untreated plots and plots treated with Fipronil® (Aventis) clearly demonstrated that the exclusion of termites from part of the savanna is possible and that this possibility can be used to study their role on the soil and organic matter cycle in situ.     

Microhabitat associations of terrestrial insectivorous birds in Amazonian rainforest and second‐growth forests

ABSTRACT Across the Neotropics, small‐bodied terrestrial insectivores are sensitive to forest fragmentation and are largely absent from second‐growth forests. Despite their sensitivity to forest structure, the microhabitat relationships of these birds have not been quantified. From July 1994 to January 1995 in central Amazonia, we characterized habitat at sites where nine species of terrestrial insectivores were observed foraging, as well as at randomly selected sites in continuous forest and two types of 10–15‐yr‐old second‐growth forest common in Amazonia (Vismia‐ and Cecropia‐dominated). We used factor analysis to find suites of correlated variables. From each factor, we selected a representative variable that was relatively easy to measure. We used Bayesian analysis to estimate means and standard deviations of these variables for each species and for each type of habitat. All nine focal species were associated with ranges of microhabitat variables, such as leaf litter depth and tree densities, often absent in second‐growth forests. At least in the early stages of regeneration, neither type of second‐growth forest provides suitable structure for the terrestrial insectivores in our study. The large leaves of Cecropia trees that make up the thick leaf litter may preclude the use of Cecropia‐dominated second growth by our focal species, many of which manipulate leaves when foraging. The leaf litter in Vismia‐dominated second growth was also thicker than sites used for foraging by our focal species. In addition, Vismia‐dominated growth had more small trees and small nonwoody vegetation, perhaps impeding movement by terrestrial birds. In continuous forest, our focal species foraged in microhabitats with characteristics that generally overlapped those of randomly selected sites. Thus, our results are consistent with the hypothesis that microhabitat differences make second‐growth forests unsuitable for our focal species.     

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.     

Foraging behavior of migrant warblers in mixed‐species flocks in Venezuelan shade coffee: interspecific differences,tree species selection,and effects of drought

Shade coffee has been identified as an important habitat for Nearctic‐Neotropical migrants during the non‐breeding season, including species of conservation concern such as Cerulean Warblers (Setophaga cerulea). To better understand habitat features important for migrants in shade coffee, we studied the foraging behavior of migrants in mixed‐species flocks at six shade‐coffee farms in the Cordillera de Merida, Venezuela, in 2008–2009 and the El Niño drought year of 2009–2010. We examined interspecific differences in foraging behavior and tree species selection of three foliage‐gleaning migrants, Blackburnian (Setophaga fusca), Cerulean, and Tennessee (Oreothlypis peregrina) warblers, and aerial‐foraging American Redstarts (Setophaga ruticilla). For morphologically similar Blackburnian and Cerulean warblers, we also examined factors influencing foraging rates (attack and movement rates), capture of large prey, and maneuver/substrate type. We found that aerial‐foraging American Redstarts foraged lower, used more aerial maneuvers, showed no tree species selection, and were less likely to forage in flocks than foliage‐gleaners. Although foraging rates were similar for Blackburnian and Cerulean warblers, the three foliage‐gleaners differed in foraging height and use of maneuvers. Cerulean Warblers foraged lower than the other two species, whereas Blackburnian Warblers used the greatest proportion of woody gleans. All three foliage‐gleaners selected Inga spp. (a commonly planted shade tree in shade‐coffee farms) for foraging, and Blackburnian and Cerulean warblers captured a greater proportion of large prey in Inga spp. than in other tree species. During the drought year, Blackburnian and Cerulean warblers captured half as many large prey and used a greater proportion of woody‐gleans. We found that interactions among behavioral, floristic, and environmental drivers influenced the foraging behavior of migrants wintering in shade coffee. Our results support those of previous studies suggesting that migrants partition resources behaviorally during the non‐breeding season, that foliage‐gleaners may benefit from the presence of shade trees, especially Inga spp., in agroforestry systems, and that drought may influence the foraging behavior of foliage‐gleaning migrants, presumably due to reduced prey availability.     

Parid foraging choices in urban habitat and their consequences for fitness

Urban environments are habitat mosaics, often with an abundance of exotic flora, and represent complex problems for foraging arboreal birds. In this study, we used compositional analysis to assess how Blue Tits Cyanistes caeruleus and Great Tits Parus major use heterogeneous urban habitat, with the aim of establishing whether breeding birds were selective in the habitat they used when foraging and how they responded to non‐native trees and shrubs. We also assessed whether they showed foraging preferences for certain plant taxa, such as oak Quercus, that are important to their breeding performance in native woodland. Additionally, we used mixed models to assess the impact of these different habitat types on breeding success (expressed as mean nestling mass). Blue Tits foraged significantly more in native than non‐native deciduous trees during incubation and when feeding fledglings, and significantly more in deciduous than evergreen plants throughout the breeding season. Great Tits used deciduous trees more than expected by chance when feeding nestlings, and a positive relationship was found between the availability of deciduous trees and mean nestling mass. Overall, the breeding performance of both species was poor and highly variable. Positive relationships were found between mean nestling mass and the abundance of Quercus for Great Tits, but not for Blue Tits. Our study shows the importance of native vegetation in the complex habitat matrix found in urban environments. The capacity of some, but not all, species to locate and benefit from isolated patches of native trees suggests that species vary in their response to urbanization and this has implications for urban ecosystem function.     

Breeding limits foraging time: evidence of interrupted foraging response from body mass variation in a tropical environment

Birds should store body reserves if starvation risk is anticipated; this is known as an ‘interrupted foraging response’. If foraging remains unrestricted, however, body mass should remain low to limit the predation risk that gaining and carrying body reserves entails. In temperate environments mass gain in female birds during breeding is often attributed to egg formation and mass loss after incubation to flight adaptation or the effect of reproductive workload, rather than as a result of an adaptive interrupted foraging response to the limited foraging time or unpredictable foraging conditions that breeding demands. In tropical environments, foraging conditions vary more within the breeding season than in temperate environments, and so studies in tropical environments are more suited to decouple the potentially confounded effects of increase in body reserves versus egg formation on the body mass of breeding birds. In this study, we test whether breeding results in an interrupted foraging response in a tropical savannah system using body mass data collected over a 15‐year period from female common bulbuls Pycnonotus barbatus. This species breeds both in the wet and dry season, despite fewer resources being available in the dry season. Breeding stage predicted female body mass: body mass peaked abruptly during incubation, but was not closely associated with the egg‐laying stage, and declined during brood rearing. Breeding females were heavier in the dry season than in the wet season. In the dry season, heavier birds were more likely to incubate eggs or brood chicks. These observations suggest that increased body reserves are required to buffer the consequence of limited foraging time or impoverished foraging conditions, which may be most pronounced during incubation and in the dry season, respectively. Such mass increases are consistent with an interrupted foraging response, which may apply to temperate zone birds experiencing foraging restrictions during breeding.