Context-dependent specialisation drives temporal dynamics in intra- and inter-individual variation in foraging behaviour within a generalist bird population

Individual niche variation is common within animal populations, and has significant implications for a wide range of ecological and evolutionary processes. However, individual niche differences may also temporally vary as a result of behavioural plasticity. While it is well understood how niche variation is affected by changes in resource availability, comparatively little is known about the extent to which individual niche differences may vary within the annual cycle due to internal drivers. Here, we assess how time- and energy-constraints imposed by incubating and brood rearing affect inter- and intra-individual variation in the foraging behaviour of lesser black-backed gulls, a generalist seabird with strong individual niche variation. To this end, we compared daily foraging trips of 22 breeding and 23 non-breeding GPS-tracked adult gulls from two colonies in the Southern Bight of the North Sea over the course of the breeding season. We find that breeding birds, unlike non-breeding ones, did indeed alter their foraging behaviour during the breeding season. Both sexes reduced their searching effort by increasingly revisiting earlier foraging locations, allowing for shorter and more frequent foraging trips. Breeding females also showed pronounced shifts in their habitat use and strongly specialised on urbanised foraging habitats throughout the breeding season. Hence, while individual variation in habitat use remained largely consistent within non-breeders and in breeding males, individual variation among breeding females almost completely disappeared. Female lesser black-backed gulls are on average smaller, and therefore often outcompeted by males for the most profitable food sources. The temporal specialisation on spatially reliable anthropogenic food sources during breeding hence suggests a complex interplay between intrinsic competitive constraints, resource reliability and shifting time- and energy budges in shaping temporal dynamics in individual niche variation within our study population.     

The effect of habitat quality on foraging patterns, provisioning rate and nestling growth in Corsican Blue Tits Parus caeruleus

Many bird species face seasonal and spatial variation in the availability of the specific food required to rear chicks. Caterpillar availability is often identified as the most important factor determining chick quality and breeding success in forest birds, such as tits Parus spp. It is assumed that parents play an important role in mediating the effect of environment on chick development. A reduction in prey availability should therefore result in increased foraging effort to maintain the amount of food required for optimal chick development. To investigate the capacity of adults to compensate for a reduction in food supply, we compared the foraging behaviour of Blue Tits Parus caeruleus breeding in rich and poor habitats in Corsica. We monitored the foraging effort of adults using radiotelemetry. We also identified and quantified prey items provided to nestlings by using a video camera mounted on the nest. We found that the mean travelling distance of adults was twice as great in the poor habitat as it was in the rich. Despite the marked difference in foraging distance, the proportion of optimal prey (caterpillars) in the diet of the chicks and the total biomass per hour per chick did not differ between the two habitats. We argue that relationships between habitat richness, offspring quality and breeding success cannot be understood adequately without quantifying parental effort.     

Species conservation on human-dominated landscapes: the case of crowned crane breeding and distribution outside protected areas in Uganda

Species primarily dependent on habitats outside protected areas have to cope with the direct pressure of human utilization or harassment, and the indirect pressure of habitat modification. In Uganda, wetlands mainly used by cranes occur mostly outside protected areas. The country's wetlands are however under heavy modification, and this and other pressures are thought to have reduced crane populations to 10–30% of the number in the early 1970s. From January 2005 to January 2006, we assessed the status of crane breeding and distribution in the country. Breeding and foraging sites were assessed through a nationwide newspaper survey, and visits to wetlands. We established occurrence of 21 nests during the study period, and crane use of 27 out of 30 districts surveyed. Crane harassment and trapping were common during breeding, as was crop damage by cranes. This is the first attempt to show the distribution of crane breeding outside protected areas throughout the country. In addition to maintaining suitable breeding habitat, survival of cranes in Uganda lies in part in community engagement and punishing crimes related to hunting and destruction of nests.     

The social organization of non-breeding Magpies Pica pica

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

Feeding time strategies of the fringe-toed lizard,Uma inornata,during breeding and non-breeding seasons

Summary I examined the foraging behavior during the breeding and non-breeding seasons, May and July 1986, of the fringe-toed lizard Uma inornata (Iguanidae). During the breeding season males differ from females in their diet and in their foraging time strategy, males exhibiting time minimization and females energy maximization. In May, plant associated foods were selectively eaten. Males concentrated on flowers, a readily available quick energy food, which reduced foraging time and increased time for reproductive activities. Time budgets indicate that males spend over twice as much time in the open and in movement in May than do females. Females at this time restrict their activities to the cover of perennial bushes, and feed primarily on plant foods (flowers and arthropods). Energy maximization appears to be maintained by both sexes in the non-breeding season when food resources diminished to one-half of those in the breeding season. The lizards were less selective in their July feeding habits, broadening their diets to include ground-dwelling arthropods and foliage. Predation by these lizards follows a wait-ambush mode of foraging.     

Differential Range Use between Age Classes of Southern African Bearded Vultures Gypaetus barbatus

Bearded Vulture Gypaetus barbatus movements were investigated in southern Africa to determine whether an individual''s age, sex or breeding status influenced its ranging behaviour and to provide the information required to guide conservation activities. Data from satellite transmitters fitted to 18 individuals of four age classes were used to determine range size and use. Because of the nature of the movements of marked individuals, these data could be used to determine the overall foraging range of the entire population, which was estimated to be 51 767 km². Although juvenile, immature and sub-adult birds used different parts of the overall range, their combined foraging range was 65% (33 636 km²) of the overall range. Average adult home ranges (286 km²) were only around 1% the size of the average foraging ranges of non-adults (10 540 –25 985 km²), with those of breeding adults being even smaller (95 km²). Home ranges of breeding adults did not vary in size between seasons but adults utilized their home range more intensively whilst breeding, moving greater distances during the incubation and chick hatching period. Range size and use increased as non-adults aged. Immatures and sub-adults had larger range sizes during winter, but range use of non-adults did not vary seasonally. Range size and use did not differ between the sexes in any of the age classes. Information on home range size and use enables specific areas within the species'' range to be targeted for management planning, education and conservation action.     

Changes in the Montagu’s Harrier Circus pygargus diet in Eastern Poland across decades promote insects and reptilians,but not birds and rodents

We investigated temporal changes in diet composition of the Montagu''s Harrier Circus pygargus breeding in natural habitat (calcareous peat bog) in SE Poland. We characterized diet composition in a three‐year period (2007–2009), based on pellet analyses. We investigated whether diet composition was affected by years or stage of breeding. We compared diet of the studied population between 2000s and 1990s and with other populations. We found that the food of the studied population was dominated by insects and mammals (by number) and mammals and birds (by biomass). Biomass and abundance of main prey items differed between studied years because of different air temperatures. We found some interannual differences in contribution of some prey items including higher number of thermophilic prey (insects and amphibians) in warmer years. Comparison of pellet composition in the 1990s and 2000s revealed significant increase in the abundance of thermophilic prey (insects and reptiles) and decrease of mammals including Microtus voles and birds. Those changes may be linked to habitat changes in areas neighboring peat bogs and climate change‐induced changes in prey communities. The studied population was able to respond to changes in foraging habitats and prey composition by opportunistic foraging on easily available prey. The diet of the studied population is the most similar to the geographically closest populations foraging in similar habitats and characterized by high contribution of insects.     

Selection of foraging habitat and nestling diet by Meadow Pipits Anthus pratensis breeding on intensively grazed moorland

Capsule Foraging sites with low vegetation height and density, but with high arthropod biomass, are selected.

Aims To test the hypothesis that on intensively grazed moorland, breeding Meadow Pipits forage for nestling food where arthropod prey are most readily available, and therefore that foraging site choice is a function of prey abundance and vegetation structure.

Methods Observations of adults provisioning nestlings were made from hides positioned close to 19 nests within grazed, 3.3-hectare experimental plots at Glen Finglas, Scotland. Vegetation height and density and arthropod abundance from mapped foraging sites were compared with control sites. Prey items fed to nestlings were quantified and compared with their relative abundance.

Results Meadow Pipits selected foraging sites with significantly lower vegetation height and density, but with significantly higher arthropod biomass. Our data suggest that within foraging sites, Meadow Pipits select particular prey types to provision nestlings, in particular, Lepidoptera larvae, adult Tipulidae and Arachnida.

Conclusions In intensively grazed upland systems, it appears that Meadow Pipits select foraging sites that optimize total food abundance and accessibility. In order to understand how anticipated changes to livestock farming in Europe will affect grassland birds, we recommend that future studies should investigate the foraging and vigilance behaviour, diet composition and breeding success of a variety of bird species provisioning nestlings under a range of livestock management scenarios.     

Foraging behaviour and prey size of the painted stork

The foraging behaviour of painted stork Mycteria leucocephala was studied during 2004–2006 at 14 different sites in the Delhi region, India. Observations were recorded on 131 individuals, including 29 juvenile birds using a video camera. Recordings were also made at the nesting colony in Delhi zoo to study the prey sizes regurgitated to nestlings. The results confirm that the painted stork is a tactile forager and exclusively piscivorous. Foraging group size ranged from 1 to 18 individuals. Per 5 min foot stirring rates in the vegetated habitats were significantly higher than in non-vegetated habitats. The attempt rate and feeding rate in the breeding season were significantly higher than that in the non-breeding season. Prey sizes taken in the breeding season were significantly smaller than those taken in the non-breeding season. About 80% fish fed to the chicks were smaller than 10 cm. Young chicks were offered smaller prey compared with older chicks. The variations in foraging parameters are discussed in relation to habitats and their conservation in the Delhi region.     

Post-mortem examinations of New Zealand birds. 2. Long-tailed cuckoos (Eudynamys taitensis,Aves: Cuculinae)*

Little is known about the biology of long-tailed cuckoos (Eudynamys taitensis). In this study, 79 dead cuckoos, mostly from the wider Auckland region, New Zealand, were examined and dissected to shed light on the sex ratio, morphology, breeding season and diet. There were no statistically significant sexual differences in the means for weight or body measurements. However, immature birds had significantly shorter bill, wing and tail lengths than adults. In adults, gonads were enlarged (for breeding) from October to January, while all cuckoos identified (from plumage) as immatures had small gonads. Of 888 food items identified from 62 gizzards, 94% were insects. The main foods were cicadas and shield-bugs (Hemiptera, 48% of food items), stick-insects (Phasmatodea, 19%) and wētās and katydids (Orthoptera, 13%). Small vertebrates (lizards and birds’ eggs and nestlings) were a minor dietary element (1% of food items; 13% of stomachs). Cicadas, stick-insects and praying mantids (large insects abundant in late summer) made up 57% of the immature diet; in adults these three categories made up just 13% of food items. Immatures leave New Zealand on migration several months after adults, and this delay in late summer and autumn coincides with an opportunity to exploit a seasonal abundance of large insects.     

Seasonal and annual variation in the diet of breeding and non-breeding Wilson's storm-petrels on King George Island, South Shetland Islands

The diet of breeding and non-breeding Wilson's storm-petrels (Oceanites oceanicus) was analysed over four breeding seasons on King George Island, South Shetland Islands, in order to test if there are changes in diet composition within and between seasons. Krill was the most important food item, followed by myctophid fish and amphipods. Breeding birds showed a clear seasonal pattern of diet, with krill decreasing and alternative prey increasing from the incubation to the chick-rearing period. Non-breeders were not found to change their diet composition. The data suggest that Wilson's storm-petrels selectively choose alternative prey to krill in order to meet the nutrient demands of their chicks. In years of low food availability, Wilson's storm-petrels may have a limited option of prey choice.     

Sharing food in the stomachs of seabirds between adults and chicks--a case for delayed gastric emptying

1. Retention times of food in stomachs of breeding and non-breeding African pengiuns (Spheniscus demersus) were examined. 2. Rates of gastric emptying in non-breeding birds were linearly related to the mass of food in the stomach. 3. Breeding birds returned to the nest with more food in their stomachs when chicks were larger. The rate of regurgitation of food was linearly related to chick mass. 4. Retention times of food in the stomachs of breeding penguins can only be explain by cognisance of observed regurgitation rates and by assuming that there is no gastric emptying through the pyloric sphincter.     

Habitat use by breeding Tree Sparrows Passer montanus

Tree Sparrows Passer montanus showed a strong preference for nest-sites adjacent to wetland habitats, and avoidance of sites on intensively managed mixed farmland distant from Rutland Water. A variety of wetland edge habitats were frequently used by foraging adult birds when provisioning nestlings, and invertebrate prey of aquatic origin was frequently recorded in chick diet. We suggest that the wetland edge habitats at the study sites played a key role in providing adequate diversity and availability of suitable invertebrate prey to allow successful chick rearing throughout the long breeding season of this multi-brooded passerine. It is possible that large areas of formerly occupied farmland in the UK no longer provide these invertebrate resources due to the effects of intensification in the late 20th century. There was no evidence that the availability of supplementary seed food within 1 km of the nest-site influenced nest-site choice, or affected productivity. It is probable that the importance of seed food resources to the persistence of Tree Sparrow populations operates on a larger spatial scale than that tested in this study, due to the greater mobility of Tree Sparrows in the non-breeding season, when seed resources are most likely to be a key limiting factor. When considering conservation management aimed at recovering breeding populations of Tree Sparrows, the key resources of seed food, invertebrate food and nest-sites must be made available at appropriate spatial scales.     

Dung‐associated arthropods influence foraging ecology and habitat selection in Black‐necked Cranes (Grus nigricollis) on the Qinghai–Tibet Plateau

Variation in grassland vegetation structure influences the habitat selection of insectivorous birds. This variation presents a trade‐off for insectivorous predators: Arthropod abundance increases with vegetation height and heterogeneity, but access to arthropod prey items decreases. In contrast, grazing by large herbivores reduces and homogenizes vegetation, decreasing total arthropod abundance and diversity. However, the presence of livestock dung may help counteract the overall reduction in invertebrates by increasing arthropods associated with dung. It is unclear, however, how the presence of arthropod prey in dung contributes to overall habitat selection for insectivorous birds or how dung‐associated arthropods affect trade‐offs between vegetation structure, arthropod abundance, and access to prey. To explore these relationships, we studied habitat selection of the Black‐necked Crane (Grus nigricollis), a large omnivorous bird that breeds on the Qinghai–Tibet Plateau. We assessed the relationships between habitat selection of cranes and vegetation structure, arthropod abundance, and the presence of yak dung. We found that Black‐necked Cranes disproportionately foraged in grassland patches with short sward height, low sward height heterogeneity, and high numbers of dry yak dung, despite these habitats having lower total arthropod abundance. Although total arthropod abundance is lower, these habitats are supplemented with dry yak dung, which are associated with coleopteran larvae, making dung pats an indicator of food resources for breeding Black‐necked Cranes. Coleopteran adults and larvae in yak dung appear to be an important factor influencing the habitat selection of Black‐necked Cranes and should be considered when assessing grassland foraging trade‐offs of insectivorous birds. This research provides new insights into the role of livestock dung in defining foraging habitats and resources for insectivorous predators.     

Movements of radio-tagged Grey Herons Ardea cinerea during the breeding season in a large pond area

This paper describes the strategies of resource utilization in the course of the breeding season by five radio-tagged Grey Herons Ardea cinerea. The seasonal changes in exploitation of the environment by two breeding adults, one non-breeding adult and two non-breeding first-year birds were studied from March to August 1982, near Zonhoven in Belgium. Two adult breeding birds could be followed continuously from the end of March until the middle of June. During the first month they explored an extended area all around the colony, but each concentrated its search in a specific direction. From the end of April until the beginning of June, most probably from egg-hatching until the end of breeding activities, each bird spent a very large proportion of its time at a particular feeding site, from which other herons were actively excluded. In the first part of June they again visited different sites, each maintaining its preferred direction. From the middle of June onwards they seemed to have left the fish-pond area. The pattern of movements of the first-year birds differed markedly from that of the breeding adults. In April, although both non-breeding and breeding birds explored large areas, only the areas used by non-breeders were centred on the colony. From the end of April onwards, probably after general egg-hatching in the colony, the non-breeders very rarely revisited the colony, and from May till August their ranges became more and more restricted to very small areas at an increasing distance from the colony. They were never observed defending particular sites. The results are discussed with regard to recent speculations about the evolution of colonies as an adaptation for the exploitation of food resources. Breeding herons seem to explore a large part of the environment during incubation and defend a particular site while feeding their young. Choice of feeding site by non-breeding birds may be influenced by the site defence of the breeding birds. Non-breeding birds exploit a large area when breeding birds occupy feeding territories. Perhaps they are forced to forage in less suitable places at this time. Colonies might have evolved as a strategy to minimize effort in resource esploitation as, especially at the beginning of the breeding season, the colony could act as an assembly point in the exploration of the environment. However, its importance as an assembly point diminishes in the course of the season, as non-breeding birds no longer visit the colony and adults defend territories.     

Foraging Ecology of Fall-Migrating Shorebirds in the Illinois River Valley

Populations of many shorebird species appear to be declining in North America, and food resources at stopover habitats may limit migratory bird populations. We investigated body condition of, and foraging habitat and diet selection by 4 species of shorebirds in the central Illinois River valley during fall migrations 2007 and 2008 (Killdeer [Charadrius vociferus], Least Sandpiper [Calidris minutilla], Pectoral Sandpiper [Calidris melanotos], and Lesser Yellowlegs [Tringa flavipes]). All species except Killdeer were in good to excellent condition, based on size-corrected body mass and fat scores. Shorebird diets were dominated by invertebrate taxa from Orders Diptera and Coleoptera. Additionally, Isopoda, Hemiptera, Hirudinea, Nematoda, and Cyprinodontiformes contribution to diets varied by shorebird species and year. We evaluated diet and foraging habitat selection by comparing aggregate percent dry mass of food items in shorebird diets and core samples from foraging substrates. Invertebrate abundances at shorebird collection sites and random sites were generally similar, indicating that birds did not select foraging patches within wetlands based on invertebrate abundance. Conversely, we found considerable evidence for selection of some diet items within particular foraging sites, and consistent avoidance of Oligochaeta. We suspect the diet selectivity we observed was a function of overall invertebrate biomass (51.2±4.4 [SE] kg/ha; dry mass) at our study sites, which was greater than estimates reported in most other food selection studies. Diet selectivity in shorebirds may follow tenants of optimal foraging theory; that is, at low food abundances shorebirds forage opportunistically, with the likelihood of selectivity increasing as food availability increases. Nonetheless, relationships between the abundance, availability, and consumption of Oligochaetes for and by waterbirds should be the focus of future research, because estimates of foraging carrying capacity would need to be revised downward if Oligochaetes are truly avoided or unavailable for consumption.     

Sex-Specific Habitat Utilization and Differential Breeding Investments in Christmas Island Frigatebirds throughout the Breeding Cycle

In seabirds, equal bi-parental care is the rule, as it is considered crucial for raising chicks successfully because seabirds forage in an environment with unpredictable and highly variable food supply. Frigatebirds forage in poor tropical waters, yet males reduce and even stop parental care soon after chick brooding, leaving the female to provision the chick alone for an extended fledging period. Using bird-borne tracking devices, male and female Christmas Island Frigatebirds (Fregata andrewsi) were investigated during the brooding, late chick rearing and post-fledging period to examine whether sexes exhibit foraging strategies that may be linked to differential breeding investments. During brooding, males and females showed similar foraging behaviour under average marine productivity of oceanic waters close to the colony, but males shifted to more distant and more productive habitats when conditions deteriorated to continue with reduced chick provisioning. During the late chick rearing period, females progressively increased their foraging range to the more distant but productive marine areas that only males had visited during brooding. Birds spent the non-breeding period roosting in highly productive waters of the Sunda Shelf. The sex-specific utilisation of three different foraging habitats with different primary productivity (oceanic, coastal, and shelf areas) allowed for temporal and spatial segregation in the exploitation of favourable habitats which seems to enable each sex to optimise its foraging profitability. In addition, post-fledging foraging movements of females suggest a biennial breeding cycle, while limited information on males suggests the possibility of an annual breeding cycle.     

Co-operative Rearing by Slender-tailed Meerkats (Suricata suricatta) in the Southern Kalahari

Slender-tailed meerkats or suricates, Suricata suricatta, are small, gregarious and largely insectivorous mongooses which inhabit the arid and semi-arid areas of southern Africa. We describe four elements of co-operative rearing of the young: 1. babysitting at the den; 2. creching on foraging trips; 3. provisioning with prey items away from the den; and 4. allonursing , including spontaneous lactation. Individual band members differed markedly in contributions to co-operative rearing. All adults and yearlings guarded the den and helpers actively defended young against predators. Breeders babysat significantly less than did non-breeding adults. Differences in age–sex class contributions were also evident in creching behaviour. The feeding of kittens by provisioners extended from soon after their first emergence from the den at 2–3 wk of age to effective foraging independence at 10–12 wk. Breeding females initially contributed little care other than nursing. Allonursing occurred in six of 25 closely observed litters, including three incidents of spontaneous lactation when subordinates nursed the young of higher-ranking females. Such co-operation is likely to be critical to the survival of these highly sociable mongooses in a semi-arid environment such as the Kalahari where food availability can fluctuate enormously.     

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

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

Habitat Use and Body Mass Regulation among Warblers in the Sahel Region during the Non-Breeding Season

Migratory birds face significant challenges across their annual cycle, including occupying an appropriate non-breeding home range with sufficient foraging resources. This can affect demographic processes such as over-winter survival, migration mortality and subsequent breeding success. In the Sahel region of Africa, where millions of migratory songbirds attempt to survive the winter, some species of insectivorous warblers occupy both wetland and dry-scrubland habitats, whereas other species are wetland or dry-scrubland specialists. In this study we examine evidence for strategic regulation of body reserves and competition-driven habitat selection, by comparing invertebrate prey activity-density, warbler body size and extent of fat and pectoral muscle deposits, in each habitat type during the non-breeding season. Invertebrate activity-density was substantially higher in wetland habitats than in dry-scrubland. Eurasian reed warblers Acrocephalus scirpaceus occupying wetland habitats maintained lower body reserves than conspecifics occupying dry-scrub habitats, consistent with buffering of reserves against starvation in food-poor habitat. A similar, but smaller, difference in body reserves between wet and dry habitat was found among subalpine warblers Sylvia cantillans but not in chiffchaffs Phylloscopus collybita inhabiting dry-scrub and scrub fringing wetlands. Body reserves were relatively low among habitat specialist species; resident African reed warbler A. baeticatus and migratory sedge warbler A. schoenobaenus exclusively occupying wetland habitats, and Western olivaceous warblers Iduna opaca exclusively occupying dry habitats. These results suggest that specialists in preferred habitats and generalists occupying prey-rich habitats can reduce body reserves, whereas generalists occupying prey-poor habitats carry an increased level of body reserves as a strategic buffer against starvation.