Arrival and habitat use by Lapwings Vanellus vanellus in the early breeding season

The arrival on the breeding grounds and habitat use in relation to agricultural land use by Lapwings Vanellus vanellus was compared in rough grazing and arable study areas in the Midland Valley of Scotland. First arrivals took place in early February. Males preceded females and territory occupation by the males began soon after first arrival. During the pre-breeding period birds remained in flocks and day time habitat use and activity was strongly influenced by the lunar cycle. Birds fed mainly in those habitats in which prey (leatherjackets and earthworms) were most plentiful, and in which their feeding success was high. The choice of nesting habitat was not influenced by food availability in the immediate vicinity of the nest site but by the crypticity of the clutch and incubating adults. On rough grazing birds preferred unimproved land while on arable land spring cereal was the preferred habitat. The choice of actual nesting field on the arable land was influenced by the risk of predation and the proximity of suitable feeding fields for the adults and chicks.     

Invasion of an exotic forb impacts reproductive success and site fidelity of a migratory songbird

Although exotic plant invasions threaten natural systems worldwide, we know little about the specific ecological impacts of invaders, including the magnitude of effects and underlying mechanisms. Exotic plants are likely to impact higher trophic levels when they overrun native plant communities, affecting habitat quality for breeding songbirds by altering food availability and/or nest predation levels. We studied chipping sparrows (Spizella passerina) breeding in savannas that were either dominated by native vegetation or invaded by spotted knapweed (Centaurea maculosa), an exotic forb that substantially reduces diversity and abundance of native herbaceous plant species. Chipping sparrows primarily nest in trees but forage on the ground, consuming seeds and arthropods. We found that predation rates did not differ between nests at knapweed and native sites. However, initiation of first nests was delayed at knapweed versus native sites, an effect frequently associated with low food availability. Our seasonal fecundity model indicated that breeding delays could translate to diminished fecundity, including dramatic declines in the incidence of double brooding. Site fidelity of breeding adults was also substantially reduced in knapweed compared to native habitats, as measured by return rates and shifts in territory locations between years. Declines in reproductive success and site fidelity were greater for yearling versus older birds, and knapweed invasion appeared to exacerbate differences between age classes. In addition, grasshoppers, which represent an important prey resource, were substantially reduced in knapweed versus native habitats. Our results strongly suggest that knapweed invasion can impact chipping sparrow populations by reducing food availability. Food chain effects may be an important mechanism by which strong plant invaders impact songbirds and other consumers.     

Supplemental food alters nest defence and incubation behaviour of an open‐nesting wetland songbird

Climate‐driven increases in spring temperatures are expected to result in higher prey availability earlier in the breeding season for insectivorous birds breeding in wetland habitats. Predation during the incubation phase is a major cause of nesting failure in open‐nesting altricial birds such as the Eurasian reed warbler. The nest predation rate in this species has recently been shown to be substantially reduced under conditions of experimentally elevated invertebrate prey availability. Food availability near the nest may be an important determinant of adult incubation and nest defence behaviours during the incubation period. We used two experimental studies to compare incubation behaviour and nest defence in food‐supplemented and unsupplemented adult Eurasian reed warblers during the incubation phase. In the first study we measured nest defence behavioural responses to a taxidermic mount of a native predator (stoat Mustela erminea). In the second study we used temperature loggers installed in nests to measure breaks in incubation as a measure of nest vulnerability. Food‐supplemented birds responded aggressively to the presence of a predator more quickly than those in the unsupplemented group, suggesting they are closer to their nest and can more quickly detect a predator in the vicinity. Food‐supplemented birds also had shorter breaks in incubation (both in terms of maximum and mean off‐bout durations), presumably because they were foraging for shorter periods or over shorter distances from the nest. This study therefore identifies the behavioural mechanisms by which changes in food availability may lead to changes in nest survival and thus breeding productivity, in open‐nesting insectivorous birds.     

A comparative study of insect abundance and reproductive success of barn swallows Hirundo rustica in two urban habitats

Few studies have quantified the relative reproductive success of passerines in urban habitats. I studied food availability and reproductive success of barn swallows Hirundo rustica in two urban habitats during 2012–2015. Barn swallows breeding in the town center experienced lower insect densities than those in the town periphery. Lower food availability resulted in reduced feeding rates per capita, lower nestling body mass, longer nestling periods, longer inter‐clutch intervals, fewer first and second brood fledglings and a lower total number of fledglings produced during the breeding season in comparison to barn swallows breeding in the town periphery. I hypothesize that the lower intra‐specific competition for nest sites and fitness advantages linked to the solitary breeding in urban habitats balanced the apparent costs of reproduction in more urbanized habitats.     

BREEDING ECOLOGY OF TURNSTONES ARENARIA INTERPRES AT HAZEN CAMP, ELLESMERE ISLAND, N.W.T.

Turnstones arrive on Ellesmere Island in late May or early June. Pair-formation takes place during migration, or after arrival in courtship groups along the beaches, or on the nest territory, depending on weather conditions. Pair-formation was not observed at Hazen Camp in 1966 as most birds were already paired when observations began on 3 June. The preferred nesting habitat was Dryas-hummocked tundra closely associated with a marsh, stream, or pond. A census area of 240 ha supported 13 breeding pairs, possibly 14; the total number of pairs breeding in the Hazen Camp study area was estimated to be 70 (3.04 pairs/km²). Egg-laying began on 10 June, with 46% of the first eggs laid 13–15 June. 62% of the sets were completed between 19 and 21 June. Both sexes incubated, the female regularly and the male sporadically. Hatching was also well synchronised; most clutches hatched between 7 and 14 July. Nest success was high (79%). After hatching territories dissolved and family groups moved freely over the tundra, concentrating at ponds where food was readily available for the young. Both adults attended the young during the pre-fledging period, but the females apparently departed long before the young fledged. Males left once the young could fly and the adult fall migration was complete by early August with the exception of late breeders. Most of the young departed in the second half of August. Fall migration is complete by late August or early September. The breeding season appears to be timed so that the young are raised when food is most abundant. Food supply (dipterous insects, especially adult chironomids) was highest from 8 to 12 July at the peak of the hatching period. While food supply for the young declined during the growing period, the early departure of half the adult population, and family movements over the tundra, appeared to reduce the food demand correspondingly. Food appeared to influence the distribution of breeding pairs markedly, restricting them to the vicinity of marshes, streams, and ponds. Territoriality displayed by Turnstones is believed to be closely associated with the protection of the nest against predators and adult food requirements during incubation; it also seems likely that territory has at least a local effect in regulating the number of breeding pairs.     

Heavy and persistent rainfall leads to brood reduction and nest failure in a passerine bird

The process of reproduction is a sensitive period for bird offspring, since they are exposed to environmental conditions for several weeks. Within the long reproductive period, adult birds do not only have to maintain their own body condition, they also have to ensure nutritional needs of their nestlings. In the worst case, if weather conditions are very harsh, breeding is not successful and fails. This suggests that weather conditions might be an important driver of breeding success. Here, we studied the effect of weather conditions (temperature and precipitation) on the survival of 350 nests of great tits Parus major during nestling period in six years (2010–2015). We used traditional generalized linear mixed model (binomial response variable) and a dedicated survival-analysis program (MARK) to analyze the data. The two approaches allowed us to highlight and compare the effect of weather conditions on a fine (nestling survival) and coarse (nest fate) scale. Both methods showed that precipitation explained most of the variation in individual nestling and overall nest survival during the 15-day nestling period. Heavy and persistent rainfall did not only lead to brood reduction, it ultimately also led to losses of the entire brood. Likely causes were the negative effects of precipitation on thermoregulation, food availability and predation risk. However, while reduced food availability most likely might have led to brood reduction through selective individual nestling death, predation might have resulted in total nest failures.     

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.     

Effects of habitat fragmentation on provisioning rates, diet and breeding success in two species of tit (great tit and blue tit)

The aim of this study was to examine the effects of forest fragmentation on the ability of parent birds to provide their young with an adequate food supply. To examine whether prey population densities of the great tit (Parus major L.) and the blue tit (P. caeruleus L.) vary between study areas in different forest size classes we compared provisioning rates and chick diet and related these parameters to breeding success. We filmed 217 nests over two breeding seasons and collected data on frass fall as a general estimate of caterpillar availability. Nests which were attended by none or one parent only during filming (n = 46) were excluded from the analyses. In both years and for both species feeding rates were highest in the smallest fragments and lowest in the large forest. There was also a suggestion that differences in feeding rates between areas vary between years. We found no consistent tendency for prey size to change with forest size, although both species brought slightly smaller prey items to the nest in the smallest forest fragments and feeding rates correlated negatively with prey size. Caterpillars were the main item fed to nestlings, in both species. We found no evidence to suggest that either frass fall or the proportion of caterpillars in the diet varied with forest size. There was also no correlation between mean frass fall and the total number of caterpillars brought to the nests, in either species. Breeding success, as measured by clutch size, brood size, fledging weight and fledging success, did not differ between the small fragments and the large forest, in either species. There was also no relationship between provisioning rate (as concerns volume of prey fed to nestlings and the quality of chick diet) and breeding success parameters. In conclusion, this study does not suggest suboptimal foraging or breeding conditions in small fragments compared to a nearby large forest, for either species. Received: 23 June 1997 / Accepted: 29 December 1997     

Reproductive responses of temperate and boreal Tengmalm's Owl Aegolius funereus populations to spatial and temporal variation in prey availability

Environmental variation across space and time can strongly influence life‐history strategies in vertebrates. It has been shown that the reproductive success of birds of prey is closely related to food availability. However, relatively little is known about intraspecific differences in reproductive success of birds in relation to varying ecological conditions across environmental gradients. We investigated the reproductive performance of Tengmalm's Owls Aegolius funereus in a temperate (Czech Republic, 50°N) and a boreal (Finland, 63°N) population in relation to long‐term variations in the abundance of their main prey (small rodents). Prey densities at the northern site were much higher, but there were also large inter‐annual fluctuations and years with steep summer declines of vole densities. Northern owls laid larger clutches but offspring production per nest was similar at both study sites. This resulted from higher nestling mortality in the northern population, especially in nests established later in the season. Despite much greater nesting losses due to predation by Pine Martens Martes martes, productivity at the population level was about four times greater at the temperate site, mainly due to the much higher breeding densities compared with Finland. Tengmalm's Owls at the temperate study site may benefit from relatively stable prey abundance, a more diverse prey community that offers alternative prey during vole scarcity, longer nights in summer that allow more time for foraging, and a lower level of interspecific competition with other vole‐specialized predators.     

Patch time allocation and patch sampling by foraging great and blue tits

The rate at which parents deliver energy to their brood is an important factor in avian reproduction because poor condition caused by malnutrition may reduce the offspring's survival to breeding. Models of central place foraging predict that nesting parents should optimize their prey delivery rate by minimizing travelling distances and by selecting patches where the gain per unit cost is high. I investigated the allocation of searching time amongst food patches in the home ranges of breeding great tits, Parus major, and blue tits P. caeruleus, by radiotracking. The density of locations in individual trees was positively correlated with prey biomass within trees and negatively with the distance of the trees from the nest. These two factors explained 52% of the variance in the allocation of the birds' search time. In rich patches, food was reduced considerably within 20 m of the nests, and the birds' travelling distances increased significantly during the nestling period. In parallel to foraging selectively in rich resources near the nest, the birds continually sampled the trees in their territory. The average surplus search time due to resource exploration was 1.52 times (range 1.25-1.99) the expected search time if the birds had exclusively used the most profitable patch. Despite considerable effort in patch sampling, the overall search time per unit prey was 30% better than expected by an equal use of trees. The results suggest that foraging tit parents come close to the maximum rate of prey delivery possible in a given patch distribution. Copyright 2000 The Association for the Study of Animal Behaviour.     

Breeding Phenology of Birds: Mechanisms Underlying Seasonal Declines in the Risk of Nest Predation

Seasonal declines in avian clutch size are well documented, but seasonal variation in other reproductive parameters has received less attention. For example, the probability of complete brood mortality typically explains much of the variation in reproductive success and often varies seasonally, but we know little about the underlying cause of that variation. This oversight is surprising given that nest predation influences many other life-history traits and varies throughout the breeding season in many songbirds. To determine the underlying causes of observed seasonal decreases in risk of nest predation, we modeled nest predation of Dusky Flycatchers (Empidonax oberholseri) in northern California as a function of foliage phenology, energetic demand, developmental stage, conspecific nest density, food availability for nest predators, and nest predator abundance. Seasonal variation in the risk of nest predation was not associated with seasonal changes in energetic demand, conspecific nest density, or predator abundance. Instead, seasonal variation in the risk of nest predation was associated with foliage density (early, but not late, in the breeding season) and seasonal changes in food available to nest predators. Supplemental food provided to nest predators resulted in a numerical response by nest predators, increasing the risk of nest predation at nests that were near supplemental feeders. Our results suggest that seasonal changes in foliage density and factors associated with changes in food availability for nest predators are important drivers of temporal patterns in risk of avian nest predation.     

Arrival date, age and breeding success in white stork Ciconia ciconia

Early arrival to breeding grounds is a life history trait in birds that can result in fitness benefits. We studied the relationship between arrival date and breeding success of individuals in a central Iberian population of white stork Ciconia ciconia , between 1999 and 2005, and the ways in which other potential factors, such as age or sex, affect this relationship. Our results showed that age was the factor most closely related to arrival date and breeding success. Older individuals returned earlier to the breeding grounds, achieved larger clutch sizes and produced more chicks than younger birds. After controlling statistically for age effect, breeding probability (laid eggs or not) and laying date were still significantly explained by arrival date. A higher probability of failure to reproduce (no eggs laid) was found in birds arriving later than in those arriving early. However, clutch size and nestling success (number of nestlings in the nest 40 days after hatching) were not correlated with arrival date. Food availability in the study area throughout the breeding cycle, due to a nearby rubbish dump, could be the factor mitigating differences in clutch size and nestling success related to individual arrival date.     

Horned larks on the Tibetan Plateau adjust the breeding strategy according to the seasonal changes in the risk of nest predation and food availability

Songbirds in seasonal environments often adjust their breeding strategy according to spatial or temporal changes in breeding conditions. Here we investigate how horned larks Eremophila alpestris, a multi‐brooded songbird on the Tibetan Plateau, responded to the changing risk of nest predation and food availability across breeding attempts. We showed that both nest concealment and food supply increased with plant growth, and horned larks adjusted their breeding strategies accordingly. First they selected nest‐sites where predator density was low, which enhanced nest survival. Second, clutch size increased with improving breeding conditions. They did not adopt an ‘egg‐size’ strategy as egg size did not change with laying sequence or breeding attempt. Instead, they adopted the ‘brood survival (feeding later‐hatched nestlings more)’ and ‘brood reduction (feeding early‐hatched nestlings more)’ strategies during early and later attempts. Moreover, nestlings’ growth varied with breeding attempt: more energy was invested into the growth of body mass during the first attempt but more energy was expended on the growth of linear structures during later attempts. This difference in energy allocation reflected changing food availability. We suggest that temporal changes of environmental factors are also the important force driving the evolution of avian breeding strategies.     

Within-season sequential density dependence regulates breeding success in mallards Anas platyrhynchos

Density dependence in vital rates is a key issue in population ecology but remains largely unexplored experimentally. We studied breeding success, lake use, and prey availability in wild mallards Anas platyrhynchos on small nemoral lakes in a replicated, two-year cross-over experiment in which pair density was increased. The number of wild mallards that settled on lakes prior to introductions of extra pairs did not differ between control and introduction years. Introductions led to a lake-level reduction in the number of broods observed. However, the number of stage 2+ (almost fledged) ducklings did not differ between treatments, nor did lake utilization by nonbreeding adults, broods and ducklings. Prey resource availability differed greatly among lakes, but it did not correlate with breeding success. Partialling out the possible effect of food competition from wild adult nonbreeding mallards did not change this conclusion. Our study demonstrates sequential density dependence in breeding success; introductions caused a decrease in brood number, but despite fewer broods a similar number of nearly fledged ducklings were produced. We suggest that predation and/or lake change of broods soon after hatching created these patterns. We conclude that using a single and late measure of breeding success such as fledged birds can mask regulatory processes. Implications of density dependence and its relation to individual reproductive success are understood better if breeding success is decomposed into nest success, duckling survival and fledgling survival.     

Anthropogenically-Mediated Density Dependence in a Declining Farmland Bird

Land management intrinsically influences the distribution of animals and can consequently alter the potential for density-dependent processes to act within populations. For declining species, high densities of breeding territories are typically considered to represent productive populations. However, as density-dependent effects of food limitation or predator pressure may occur (especially when species are dependent upon separate nesting and foraging habitats), high territory density may limit per-capita productivity. Here, we use a declining but widespread European farmland bird, the yellowhammer Emberiza citrinella L., as a model system to test whether higher territory densities result in lower fledging success, parental provisioning rates or nestling growth rates compared to lower densities. Organic landscapes held higher territory densities, but nests on organic farms fledged fewer nestlings, translating to a 5 times higher rate of population shrinkage on organic farms compared to conventional. In addition, when parental provisioning behaviour was not restricted by predation risk (i.e., at times of low corvid activity), nestling provisioning rates were higher at lower territory densities, resulting in a much greater increase in nestling mass in low density areas, suggesting that food limitation occurred at high densities. These findings in turn suggest an ecological trap, whereby preferred nesting habitat does not provide sufficient food for rearing nestlings at high population density, creating a population sink. Habitat management for farmland birds should focus not simply on creating a high nesting density, but also on ensuring heterogeneous habitats to provide food resources in close proximity to nesting birds, even if this occurs through potentially restricting overall nest density but increasing population-level breeding success.     

Counting Flocks of Roosting Birds by Photography

A BTO nest record cards analysis. The early breeding season of the Dipper appears to be an adaptation to a food supply that is most abundant in early spring; and the consistent reuse of the same nest for replacement and true second clutches enables the birds to save time and utilise more efficiently the temporary abundance of food.     

Helper contributions in the cooperatively breeding laughing kookaburra: feeding young is no laughing matter

I studied the contributions of individuals to incubation and nestling feeding in a population of cooperatively breeding laughing kookaburras, Dacelo novaeguineae. In most cooperatively breeding birds where nest success is limited by nestling starvation, related helpers increase the overall level of provisioning to the nest, thus boosting the production of nondescendent kin. However, although partial brood loss is the largest cause of lost productivity in kookaburra nests, additional helpers failed to increase overall provisioning. Instead, all group members, but especially helpers, reduced their feeding contributions as group size increased. Breeders and helpers reduced the size of prey delivered, and helpers also reduced the number of feeding visits. An important benefit of helping in kookaburras may be to allow all group members to reduce their effort. Within groups, contributions to care depended on status, sex, group size and the brood size. Breeding males delivered the most food. Breeding females provisioned less than their partner, but their effort was comparable to that of male helpers. Female helpers contributed the least food. Incubation effort followed similar patterns. The relatedness of helpers to the brood had no impact on their provisioning. Across all group sizes, helpers generally brought larger items to the nest than breeders. Copyright 2000 The Association for the Study of Animal Behaviour.     

Brood size manipulations in Great Horned Owls Bubo virginianus: are predators food limited at the peak of prey cycles?

The reproduction of raptors strongly depends on food resources. It is unclear whether predators experience superabundant food during cyclic peaks of prey populations. In order to test this hypothesis, four pairs of Great Horned Owls Bubo virginianus with two young were subjected to brood size manipulations during high densities of cyclic Snowshoe Hare Lepus americanus populations in southwestern Yukon, Canada. Broods older than 35 days were temporarily enlarged by one, and then by two, young. No effects were observed when one owlet was added, but the addition of two young resulted in significant weight losses in manipulated broods. Females with enlarged broods moved farther from their nest sites at night, presumably reflecting increased hunting effort, and also spent less time near the nest during the day. Food additions to enlarged broods returned the parental behaviour to normal. We conclude that these large predators did not experience superabundant food at this stage of the breeding season during a peak in cyclic prey.     

Growth and decline of a penguin colony and the influence on nesting density and reproductive success

Colonial breeding is characteristic of seabirds but nesting at high density has both advantages and disadvantages and may reduce survival and fecundity. African penguins (Spheniscus demersus) initiated breeding at Robben Island, South Africa in 1983. The breeding population on the island increased in the late 1990s and early 2000s before decreasing rapidly until 2010. Before the number breeding peaked, local nest density in the areas where the colony was initiated plateaued, suggesting that preferred nests sites were mostly occupied, and the area used by breeding birds expanded. However, it did not contract again as the population decreased, so that nesting density varied substantially. Breeding success was related positively to the prey available to the breeding birds and negatively to local nest density, particularly during the chick-rearing period, suggesting a density-dependence operating through social interactions in the colony, possibly exacerbated by poor prey availability when the breeding population was large. Although nest density at Robben Island was not high, nesting burrows, which probably reduce the incidence of aggressive encounters in the colony, are scarce and our results suggest that habitat alteration has modified the strength of density-dependent relationships for African penguins. Gaining a better understanding of how density dependence affects fecundity and population growth rates in colonial breeders is important for informing conservation management of the African penguin and other threatened taxa.     

Food or nesting place? Identifying factors limiting Wryneck populations

In recent decades, farmland bird populations have declined strongly as a consequence of agriculture intensification. Birds may have lost breeding sites, food supply or other crucial resources, with the role of multiple factors often remaining unclear. The ant-eating and cavity-breeding Wryneck (Jynx torquilla) may be limited by the availability of cavities, the number of ants or their accessibility. By comparing occupied and unoccupied breeding territories, we investigated the relative role of these factors in the decline of Wrynecks. We compared the characteristics of known Wryneck breeding territories (availability of breeding cavities, food abundance and ground vegetation structure) with randomly selected, fictitious territories (n = 154) in Western Switzerland. We also studied environmental factors that may affect ant nest density. The probability of territory occupancy strongly increased with both nestbox availability and ant abundance. In addition, this probability peaked around 50% of bare ground cover. Habitat types that harbour low ant abundance such as cropland and grassland were avoided. Ant nest density decreased with increasing amounts of bare ground, and it was particularly high in vineyards. Our results showed that breeding cavities, food availability and its accessibility all limit Wryneck distribution. The maintenance and restoration of ant rich grassland, interspersed with patches of bare ground and with hollow trees or dedicated nestboxes in the surroundings, are essential to preserve Wryneck populations. Such a habitat structure could be achieved even in intensively farmed habitats, such as in vineyards or fruit tree plantations.