Heat stress in a high-latitude seabird: effects of temperature and food supply on bathing and nest attendance of great skuas Catharacta skua

Birds such as great skuas Catharacta skua adapted for successful breeding at high latitudes may experience problems of heat dissipation in mild climates. Great skuas spend time bathing at freshwater sites close to breeding territories and here, we examine impacts of heat stress on bathing, foraging and nest attendance of adults during three breeding seasons with marked variation in the availability of prey (1-group sandeels Ammodytes marinus ). Adults exhibited diurnal variation in bathing activity that matched heat-stress conditions. Moreover more birds bathed on days of higher average heat stress, suggesting that bathing plays a role in thermoregulation. Bathing numbers were lower in years of poor food availability, when adult attendance at territories was low, probably because lower attendance reduced the opportunity for parents to bathe without leaving chicks unattended. Chicks are normally guarded by female parents and fed by males but under conditions of low food availability territorial attendance of breeding pairs was particularly low on days of high heat stress, with chicks regularly left unattended at air temperatures exceeding 14°C. Unattended chicks are at risk of being killed by neighbouring conspecifics and survival of chicks to fledging was low in the two years of low sandeel stocks. Our study indicates that for great skuas, indirect effects of climate change on prey stocks and direct effects on heat stress experienced by adults may be additive.     

Dynamics of House Sparrow Biparental Care: What Contexts Trigger Partial Compensation?

Nest attendance during incubation is characterized by an inequitable division of labor in house sparrows, Passer domesticus, with females spending more time at the nest than males. Previous research has shown that if male contributions are reduced experimentally via testosterone (T) implants, females compensate partially for those reductions, consistent with predictions from most models of negotiated biparental care. In this study, we attempted to identify the cues and contexts generating partial compensation, using data from both unmanipulated parents and pairs with T‐males. Both males and females of this species sometimes leave the nest before their mate returns to relieve them, and we found that these unrelieved departures by unmanipulated individuals occur when partners are on lengthy recesses. Females compensated partially for long male recesses by marginally extending their bouts; most females also slightly reduced their next recess. By contrast, when males left before their mate returned, they left earlier than when they waited for the female. Neither males nor females adjusted their recess lengths after returning to the nest and discovering that their partner was absent. More pronounced changes in nest attendance of unmanipulated parents occurred in the context of ‘visits’, when individuals returned to the nest but then left without relieving their mate. Such visits effectively prolonged the bout of the on‐duty partner and extended the visitor’s recess. Analyses of behavior of T‐males and their mates revealed that T‐males had significantly longer recesses than control males, and that their mates, in turn, had elevated rates of unrelieved departures. T‐males also visited their on‐duty mates more often than control males, whereas female visits to T‐males were rare. Collectively, the predicted changes in female nest attendance associated with lengthy male recesses and male and female visits account reasonably well for the compensatory response of females paired to T‐males. The majority of female compensation was attributable to changes in visit behavior, however, suggesting that much of the negotiation over nest attendance in this species occurs during direct interactions between mates.     

Why do Both Parents Incubate in the Kentish Plover?

Incubation by both parents is a common parental behaviour in many avian species. Biparental incubation is expected if the survival prospects of offspring are greatly raised by shared care, relative to the costs incurred by each parent. We investigated this proposition in the Kentish plover Charadrius alexandrinus, in which both parents incubate the clutch, but one parent (either the male or the female) usually deserts after hatching of the eggs. We carried out a mate‐removal and food supplementation experiment to reveal both the role of the sexes and food abundance in maintaining biparental incubation by removing either the male or the female from the nest for a short period of time. In some nests we provided supplementary food for the parent that remained at the nest to reduce the costs of incubation, whereas other nests were left unsupplemented. Although males spent more time on incubation after their mate had been removed, females’ incubation did not change. Notwithstanding the increased male incubation, total nest attentiveness was lower at uniparental nests than at biparental controls. However, incubation behaviour was not influenced by food supplementation. We conclude that offspring desertion during incubation is apparently costly in the Kentish plover, and this cost cannot be ameliorated with supplementary food.     

Parental attendance and squatting in the Kittiwake Rissa tridactyla during the rearing period

Parental attendance was studied in 1991 in the Cap Sizun Kittiwake colonies (Brittany, France). After a period of continuous guarding lasting on average 22 days, parents left their chicks unattended. Thereafter, parental attendance decreased regularly until fledging. The chick age when first left alone was on average 3 days lower for large broods than for single-chick broods. Moreover, whatever the brood size, chicks from late nests were younger when left unattended. Parental age affected the initiation of first absence. Younger parents reduced their attendance sooner than older parents. About 80% of the nests with chicks were visited by other adults at least once during the absence of the parents, and 50% were visited in the 3 days following the first absence of the breeders. These squatters were mainly failed breeders and prebreeders looking for a future breeding site. The results are discussed in terms of costs and benefits of chick neglect and comparisons were made with data from other studies in North Atlantic and Alaskan colonies. This reflected the flexibility of adult behaviour in relation to brood size and food availability.     

BREEDING BIOLOGY OF THE BEARDED VULTURE IN SOUTHERN AFRICA,PART II: THE NESTLING PERIOD

Brown, C. J. 1990. Breeding biology of the Bearded Vulture in southern Africa, Part I: The nestling period. Ostrich 61: 24–32.

The nestling period of the Bearded Vulture Gypaetus barbatus in southern Africa was 124–128 days. The hatching interval between the normal two-egg. clutch was usually 3–6 days (range 2–9 days Only one nestling per clutch survived to the third day. Tittle sibling aggression and no infanticide took place, but the older nestling dominated the younger which obtained no food. For the first 40 days the nestling was closely brooded. The nest duties were evenly shared by both parents, but females brooded at night. Food was brought to the nest usually once or twice per day by both parents, and was stored behind the nest. During days 41–90 parental attendance steadily decreased. Dunng this stage the female spent more time in the nesting area (57%) and on the nest (91%) than the male. Towards the end of this stage the nestlin started to feed itself but preferred to be fed by a parent. From da 91 to first flight the nestling was left unattended and was visited by its parents only to provide food, which it fed from itself. All pars monitored (40 pair-years) attempted to breed every year. The breeding success (n = 18 pair-years) was 0,89 young fledged per pair per year.     

Prolactin and parental behavior in Adélie penguins: effects of absence from nest, incubation length, and nest failure

Adélie penguin (Pygoscelis adeliae) males and females, nesting in Antarctica, alternate attendance at the nest with absences of many days to forage at sea. We investigated the importance of tactile input from egg and chicks on prolactin levels by observing nest attendance patterns and obtaining blood samples (1) during the first nest exchange of the incubation stage, (2) from birds whose incubation period was artificially increased or decreased by about 10 days, and (3) from birds whose nests had failed. Prolactin levels in females after 8 to 11 days of absence from the breeding colony did not differ from those in incubating males and did not change after females resumed incubation. Moving eggs between nests resulted in nests in which chicks hatched after about 26, 36 (normal), or 46 days. Duration of incubation did not affect prolactin levels in the parents measured during incubation, at the pip stage, hatch stage, or early brood stage. Adults first left their chicks unguarded on about the same calendar date, regardless of chick age. However, chicks from long incubation nests averaged 8 days younger when they were left unguarded than chicks from control or short-incubation nests. In females, there was no effect of nest failure on prolactin levels. In males, prolactin levels were slightly lower after nest failure than in males tending nests. Testosterone was significantly higher in males after nest failure than in males still tending nests. Prolactin is elevated in Adélie penguins as part of the program of cyclical hormonal changes that accompany the lengthy reproductive season and is relatively independent of tactile input. Sustained prolactin secretion is probably required for the maintenance of parental behavior in offshore feeding species that must be absent from the nest for many days at a time.     

Translocation of threatened New Zealand falcons to vineyards increases nest attendance, brooding and feeding rates

Anthropogenic landscapes can be rich in resources, and may in some cases provide potential habitat for species whose natural habitat has declined. We used remote videography to assess whether reintroducing individuals of the threatened New Zealand falcon Falco novaeseelandiae into a highly modified agricultural habitat affected the feeding rates of breeding falcons or related breeding behavior such as nest attendance and brooding rates. Over 2,800 recording hours of footage were used to compare the behavior of falcons living in six natural nests (in unmanaged, hilly terrain between 4 km and 20 km from the nearest vineyard), with that of four breeding falcon pairs that had been transported into vineyards and nested within 500 m of the nearest vineyard. Falcons in vineyard nests had higher feeding rates, higher nest attendance, and higher brooding rates. As chick age increased, parents in vineyard nests fed chicks a greater amount of total prey and larger prey items on average than did parents in hill nests. Parents with larger broods brought in larger prey items and a greater total sum of prey biomass. Nevertheless, chicks in nests containing siblings received less daily biomass per individual than single chicks. Some of these results can be attributed to the supplementary feeding of falcons in vineyards. However, even after removing supplementary food from our analysis, falcons in vineyards still fed larger prey items to chicks than did parents in hill nests, suggesting that the anthropogenic habitat may be a viable source of quality food. Although agricultural regions globally are rarely associated with raptor conservation, these results suggest that translocating New Zealand falcons into vineyards has potential for the conservation of this species.     

The breeding biology of the Scimitarbilled Woodhoopoe

Steyn, P. 1999. The breeding biology of the Scimitarbilled Woodhoopoe. Ostrich 70 (3&4): 173–178.

The breeding biology of the Scimitarbilled Woodhoopoe Rhinopomastus cyanomelas was studied at two sites in Zimbabwe over a 13-year period from 1964–1977. The pairs were resident, remained together throughout the year, and inspected their nest sites occasionally during the non-breeding season. The breeding season extended from August to December with a marked September/October peak. Eggs were laid at daily intervals. Clutch size averaged 2.9 (range 2–4). Incubation began either with the penultimate or last laid egg. During the 13–14 day incubation period the female left the nest only occasionally each day and was reliant on the male for food. This pattern continued for four days after the chicks hatch. Thereafter she started to forage and gradually increased her contribution to chick provisioning until she overtook that of the male. With one exception, he never fed the chicks directly and delivered the food to the female. The nestlings were brooded overnight for the first two weeks. The anti-predator response of the young included a malodorous brown exudation from the preen gland and unpleasant liquid excreta. The nestling period was 21–24 days and the young left the vicinity of the nest with their parents and did not return to roost in it. Twelve breeding cycles were monitored and 76% of eggs laid (n=37) produced fledged young. Second broods were raised in the same nest on two occasions after successful rearing of the first, presumably by the same pair, but the birds were not individually marked. There was no evidence of helpers at the nest.     

Feeding frequency influences crèching age in the Dalmatian pelican, Pelecanus crispus

Crèching behaviour is common in colonial seabirds; nevertheless, the factors inducing chicks to aggregate remain relatively poorly understood. It has been proposed that brood size, laying date and nest attendance are important factors in the formation of a crèche. Moreover, in most species of pelicans, chicks join crèches following the development of homoeothermy and coincident with the end of the brooding behaviour. We studied effects of feeding rate, nest attendance, brood size, laying date and homoeothermy on the age at which chicks entered the crèche at a colony of Dalmatian pelicans (Pelecanus crispus), in Srebarna, Bulgaria. Single chicks were fed more frequently than chicks from two-chick broods. Unlike American white pelicans (Pelecanus erythrorhynchos), Dalmatian pelicans maintained brooding behaviour a further 9 days after chicks had developed thermoregulation abilities. In contrast to nests with two chicks, nests with only one chick were never left unattended by the parents before the chick reached the crèching stage. Laying date, nest attendance and brood size did not affect the age that the chick entered the crèche. The age the chick entered the crèche was not correlated with the age of homoeothermy acquisition, but chicks significantly joined the crèche at younger ages when the mean number of feeds per chick per day during the rearing period in the nest was higher. This result suggests an implication of growth rate in the crèching age. Joining the crèche earlier can provide benefits that could have strong implications for the chicks’ future reproductive lives.     

THE TIMING OF BIRDS' BREEDING SEASONS

Examination of survival rdtes of nestlings and fledglings of some species show that there is a strong tendency for those young which are hatched earliest in the season to have the greatest chance of surviving to breed. Since natural selection so strongly favours parents who leave many surviving young, the question arises as to why other birds breed later than the date at which they could most successfully raise their young.
It is suggested that the food supply for the breeding females immediately prior to the breeding season may limit their ability to form eggs and the females may thus not be able to lay at the time which would result in young being in the nest at the best time for raising them, but as soon after this time as the female is able to produce her eggs. Not all species are likely to be prevented, by food shortage, from breeding at the best time for raising young and the groups of birds most likely to be affected are discussed.     

BREEDING BIOLOGY OF THE BEARDED VULTURE IN SOUTHERN AFRICA,PART I: THE PRELAYING AND INCUBATION PERIODS

Brown, C. J. 1990. Breeding biolo of the Bearded Vulture in southern Africa, Part I: The pre-laying and incubation periods. Ostrich 61: 24–32.

In southern Africa the Bearded Vulture Gpaetus barbatus lays its eggs in mid-winter. between the second half of May and the first week of July. Pairs became more active in their nesting areas about six weeks before laying and usually roosted there at night. Courtship flights were less frequent and demonstrative than in Eurasian birds and took place mainly in the late afternoons. During the pre-laying period most nest visits (77%) were to bring nesting material, 92% by the male. All nesting material was arranged by the female. Copulation was always preceded by allopreening, and occurred most frequently in the mornings. No copulation or courtship display took place after the first egg had been laid. Of 18 clutches, 16 (89%) contained two eggs and the remainder one egg. The laying interval was usually 3–5 days (range 2–9 days). Incubation started with the first egg and was evenly shared by both parents during the day, but only the female incubated at night, individual pairs maintained distinctive nest attendance and foraging period timetables, which allowed sufficient time for self-foraging by both parentes. No food was brought into the nest during the pre-laying and incubation periods, but in some pairs food was cached in nearby potholes in cliffs. The incubation period was 56–57 days.     

OBSERVATIONS ON THE POST-FLEDGING DEPENDENCE PERIOD OF CAPE VULTURES

Robertson, A. S. 1985. Observations on the post-fledging dependence period of Cape Vultures. Ostrich 56: 58–66.

Cape Vultures were observed during their post-fledging dependence period at a colony in the Cape Province, South Africa. Information is presented on the length of the period, behaviour of juveniles and of parents at the nest, survival of juveniles, aggressive interactions between parents and juveniles and retention of the nest site following breeding. At the colony, juveniles initiate contact with their parents, which supply food to their own offspring at the natal site only. Parental aggression was observed over an average period of five months after juveniles had left the nest (range 32–218 days); at two nest sites, the period overlapped with the next season's incubation period, although no transfer of food was observed during this period-of overlap.     

Nest Attendance does not Predict Offspring Desertion by Eurasian Penduline Tit Parents

Parental care is costly, and in many organisms, the male or the female parent benefits from reducing its own care which may be compensated for by its mate. One of the parents may even face all costs of parental care if its mate deserts and leaves him/her to care for the offspring alone. Theoretical models have generated contrasting predictions as to how parents negotiate a resolution of this sexual conflict over care, although empirical tests are largely lacking. We investigated pre‐desertion behaviour (nest attendance) of a highly polygamous passerine, the Eurasian penduline tit ( Remiz pendulinus) that exhibits intense sexual conflict over care culminating in clutch desertion by the male, by the female or by both parents. We conjectured that nest attendance of parents should predict desertion, so that the lack of care provided by the deserting parent may be compensated for. By analysing over 200 000 video frames (2.50 ± 1.36 d per pair, 302 ± 170 min/d) of 20 pairs, we show that nest desertion is not a gradual process and cannot be predicted by nest attendance. These results are consistent with the argument that the predictable desertion may not be evolutionarily stable, and suggest that male and female penduline tits do not negotiate clutch desertion.     

Reed warblers guard against cuckoos and cuckoldry

Previous studies have shown that reed warblers, Acrocephalus scirpaceus, are more likely to reject a cuckoo, Cuculus canorus, egg if they have seen a cuckoo at their nest. This suggests that they would benefit from watching out for cuckoos. We tested whether presentations of a cuckoo mount near the nest (to simulate nest inspection) led to increased nest attendance by the warblers. Cuckoo presentations at completed nests before laying, when males guarded their females closely, led to desertion at 40% of nests before any eggs were laid (there were no desertions after presentations of a jay,Garrulus glandarius , a nest predator). In the remaining cases, there was no effect of the cuckoo on nest attendance before laying began, but a marked increase in male nest attendance (compared with jay and no-presentation controls) on the days the first and second eggs were laid. Cuckoo presentations at the one-egg stage led to the same increase in male nest attendance as did the prelaying presentations. Increased male nest attendance at the one-two-egg stage was not at the expense of mate guarding, because this declined anyway when laying began, and it did not lead to increased paternity loss compared with controls. Overall, 15% of broods had one or two extrapair young (6% of all young extrapair). We conclude that male reed warblers do increase nest guarding in response to cuckoos, but only after their females have begun egg laying, when there are less likely to be costs in lost paternity. Females did not increase nest guarding, perhaps because they need to spend more time foraging during the egg-laying period. Our results suggest that cuckoos should be secretive not only when they lay but also when they monitor host nests beforehand. Copyright 2003 The Association for the Study of Animal Behaviour. Published by Elsevier Science Ltd. All rights reserved.      

Do parents play a role in the timing and process of fledging by nestling Mountain Bluebirds?

What causes young birds to leave nests remains unclear for almost all altricial species. For many years, the assumption was that parents often controlled the time of fledging by coaxing young from nests, e.g., by holding food within view, but out of reach, of nestlings. This assumption, though, was based solely on scattered anecdotal reports of such behavior. We used continuous video‐recording of nests to assess the role of parents, if any, in the timing and process of fledging of cavity‐nesting Mountain Bluebirds (Sialis currucoides). We placed perches ~50 cm in front of nest‐box entrances to give parents ample opportunity to display food to nestlings. We found no evidence that parents routinely initiated the fledging process. On the day of fledging, parents did not perch on supplemental perches with food more often, or for longer periods of time, than on the day before fledging. Also, after going to nest‐box entrances, parents never held food away from a nestling reaching for the food. Parents were usually absent (16 of 19 cases) when the first nestling fledged. In the remaining three cases, a parent perched with food in view of a nestling for 8, 15 and 65 s, respectively, just before that nestling fledged. Although these might have appeared to be attempts at coaxing, in each case, the parent was encountering, for the first time, a nestling partially emerging from the nest entrance. Parents may simply have hesitated to approach nests because the nestling's position prevented parents from delivering food in the normal manner. Finally, the rate at which parents fed nestlings on the day of fledging did not differ from the rate the day before, suggesting that parents do not try to use hunger to induce fledging. Our results are consistent with previous research suggesting that, in Mountain Bluebirds, it is a nestling that initiates fledging, typically when it reaches some threshold state of development.     

The evolution of crocodilian nesting ecology and behavior

Crocodilians comprise an ancient and successful lineage of archosaurs that repeatedly raises questions on how they survived a mass extinction and remained relatively unchanged for ~100 million years. Was their success due to the change‐resistant retention of a specific set of traits over time (phylogenetic conservatism) or due to flexible, generalist capabilities (e.g., catholic diets, phenotypic plasticity in behavior), or some combination of these? We examined the evolution of reproductive ecology and behavior of crocodilians within a phylogenetic perspective, using 14 traits for all 24 species to determine whether these traits were phylogenetically constrained versus (ecologically) convergent. Our analysis revealed that the ancestral crocodilian was a mound nester that exhibited both nest attendance and defense. Nesting mode exhibited 4–5 transformations from mound to hole nesting, a convergence of which habitat may have been a driving factor. Hole nesters were more likely to nest communally, but this association may be biased by scale. Although there were exceptions, mound nesters typically nested during the wet season and hole nesters during the dry season; this trait was relatively conserved, however. About two‐thirds of species timed their nesting with the wet season, while the other third timed their hatching with the onset of the wet season. Nest attendance and defense were nearly ubiquitous and thus exhibited phylogenetic conservatism, but attendance lodging was diverse among species, showing multiple reversals between water and burrows. Collectively, our analysis reveals that reproductive trait evolution in crocodilians reflects phylogenetic constraint (nest attendance, nest defense), ecological convergence (seasonal timing of nesting, nest attendance lodging), or both (mode of nesting). Some traits (e.g., communal nesting and mode of nesting) were autocorrelated. Our analysis provides a framework for addressing hypotheses raised for why there has been trait convergence in reproductive ecology and behavior in crocodilians and why some traits remained phylogenetically conserved.     

THE BREEDING OF A PARADISE FLYCATCHER

1. At Amani, Tanganyika Territory, the Paradise Flycatchers use the softest moss for their nests and make them durable by oversewing the rim with cobweb. Sites are on the forest edge nearly always over water. Theree hours of observation were made at three nests in spells of about eight hours at a time.
2. On the whole the males and females shared the care of the eggs and the young about equally, but there were wide differences in this respect from day to day
3. The eggs were covered for over 90% of the observed time, and a high proportion of spells "on" were terminated by thc mate's arriving to take over. Duration of individual spells "on" varied up to two hours, but the favourite duration (nearly half of all the spells) was about 30–40 minutes. This applied both to spells "on" terminated by the initiative of the sitting bird and to those terminated by the arrival of the mate. The possibility suggests itself that an internal rhythm was operating, irrespective of whether a bird was on or off the nest. Nevertheless, out of 39 occasions (all short) when the eggs were uncovered through the sitter's departure without relief, the same bird returned on 18, which suggests that when off the nest both parents "keep an eye on it" and react to the situation "nest uncovered".
4. Brooding of the young amounted at first to nearly the same high percentage of time as the brooding o f the rggs, but was in much shorter spells. It stopped abruptly about the fifth day. others followed a parent.
5. Each nestling of a brood of two in a nest received more food (largely butterflies) than each in two broods of three, 2.6-5.4 feeds per hour compared with 14-2.5 and 1.3-1.8.
6. Some of the young that wcre seen to fly left the nest in the absence of the parents, All left between dawn and noon. Nestling period about 11 days irrespective of the amount of food received.     

Aspects of the breeding ecology and behaviors of the Bar-tailed Lark (Ammomanes cinctura) from Ha’il region in north of Saudi Arabia

The Bar-tailed Lark (Ammomanes cinctura) breeds in desert and semi-desert areas of the Saharo-Sindian region, from north-west Africa through the arid plains of the Arabian Peninsula to the Sind. Despite having a wide distribution, little information is available on the breeding ecology of this species. This study was conducted in a desert in the north of Saudi Arabia, where the daytime ambient temperature may exceed 40 °C. In contrast, the night ambient temperature may reach less than 10 °C in late spring and early summer. The objectives of this study were to collect some baseline data on some aspects of the breeding ecology of this species and to record the nest attendance behavior. The study found that Bar-tailed Larks preferred to nest under shrub trunks, which may camouflage both nests and incubating parents against predators and protect eggs, nestlings and incubating parents from hostile weather conditions. Moreover, nest attendance was high, as Bar-tailed Lark parents incubated their eggs 95.97 ± 2.62% over the entire day, and they seemed to maintain the eggs at temperatures around 23–33 °C. In addition, they incubated more at night than during the daytime. Temperatures under the shrubs at night fell below 21 °C, thus parents increased the nest attendance to warm the eggs and prevent the embryos from exposure to lethal temperatures.     

Corticosterone Promotes Scramble Competition Over Sibling Negotiation in Barn Owl Nestlings (Tyto alba)

In species with parental care, siblings compete for access to food resources. Typically, they vocally signal their level of need to each other and to parents, and jostle for the position in the nest where parents deliver food. Although food shortage and social interactions are stressful, little is known about the effect of stress on the way siblings resolve the conflict over how food is shared among them. Because glucocorticoid hormones mediate physiological and behavioral responses to stressors, we tested whether corticosterone, the main glucocorticoid in birds, modulates physical and vocal signaling used by barn owl siblings (Tyto alba) to compete for food. Although corticosterone-implanted (cort-) nestlings and placebo-nestlings were similarly successful to monopolize food, they employed different behavioral strategies. Compared to placebo-nestlings, cort-individuals reduced the rate of vocally communicating with their siblings (but not with their parents) but were positioned closer to the nest-box entrance where parents predictably deliver food. Therefore, corticosterone induced nestlings to increase their effort in physical competition for the best nest position at the expense of investment in sib?Csib communication without modifying vocal begging signals directed to parents. This suggests that in the barn owl stress alters nestlings?? behavior and corticosterone could mediate the trade-off between scramble competition and vocal sib?Csib communication. We conclude that stressful environments may prevent the evolution of sib?Csib communication as a way to resolve family conflicts peacefully.     

How different provisioning strategies result in equal rates of food delivery: an experimental study of blue tits Parus caeruleus

Food provisioning in birds requires a considerable amount of time and usually has to be traded-off against other parental and non-parental activities. I investigated experimentally the rate at which blue tit Parus caeruleus parents deliver food to their brood after a change in food availability. The main argument behind this study is that parents enjoying an additional food source may use less time for self-feeding and therefore use more time for food provisioning. This could increase the rate at which food is brought to the nest. However, a prey choice model that takes the energetic needs of the parent into account allows for the possibility that the food-supplemented parents would deliver the same amount of food by increasing prey size (through an increase in prey selectivity) and reducing visit rate. The field data indicate that the parents changed provisioning strategy when food-supplemented: they fed the chicks natural food less frequently, but brought larger larvae. On the whole, delivery rate of natural food was the same or lower than in controls. The results suggest that food-supplemented parents used the time saved to increase their degree of food selectivity. When the gains from an increased delivery rate are not worth the increased costs (mainly resulting from an increased visiting rate), the parent with low energetic need may increase selectivity to provide the same amount of food to the brood as the unmanipulated parent, but at a lower cost.