Parental differences in brood provisioning by Hen Harriers Circus cyaneus

Capsule Females varied their provisioning patterns according to brood age and brood size, whereas males did not.

Aims To quantify how parents balance the needs of their offspring for food and protection.

Methods We studied 13 nests from hides and spent on average 101 hours per nest monitoring prey types, provisioning rate and the time spent at the nest by both sexes in relation to brood size and brood age.

Results Males always provided more food than females. Males brought similar amounts of prey items irrespective of brood size and nestling age, whereas females brought more prey and bigger items to larger and older broods. Females spent less time brooding larger broods, particularly early on.

Conclusions Hen Harrier parents share the provisioning burden, with each parent delivering prey as a function of brood care requirements, hunting capability and the behaviour of the other parent.     

Time constraint on food choice in provisioning blue tits, Parus caeruleus: the relationship between feeding rate and prey size

Previous work on food-provisioning behaviour in blue tits suggested that the parents could gather larger prey items only by making longer foraging excursions, for example, by being more selective or by reaching more distant (and less exploited) feeding sites. Here, I show that within-nest, within-day variation in size of prey delivered by the parent could be explained by the time since its last visit. In unmanipulated conditions, size of larvae tended to increase with the time spent away from the nest. A significant positive relationship was more likely at high provisioning rates, suggesting that periods of intense feeding limited the size of prey delivered to the brood. To assess the effect of less intense feeding on prey size, I experimentally increased food availability to the tits. The parents could decide whether to eat the extra food or feed it to the nestlings. In both cases, food supplementation could result in longer time lags between natural feedings. Food-supplemented parents consumed the extra food and fed it to their nestlings, made longer foraging trips and delivered larger natural larvae than controls. In this group, size of larvae was more constant during the observation period and was independent of the time since the parent's last visit. This suggests that, below some value of visit rate, prey size is no longer limited by the duration of the foraging trip. The results support the view that tits continually vary visit rate and prey size. There is some evidence that these adjustments are made by changing food selectivity in response to changes in the state of the brood and of the parents.Copyright 2002 The Association for the Study of Animal Behaviour. Published by Elsevier Science Ltd. All rights reserved .     

Flexibility in the Foraging Behavior of Blue Tits in Response to Short‐Term Manipulations of Brood Size

Previous work suggests that short‐term changes in feeding rate are usually produced by the parent‐offspring interaction. However, few studies have properly tested this assumption. In this study, we attempt to explore the short‐term consequences of daily (within‐pair) brood size manipulations (reduced, original, and enlarged) on feeding behavior (provisioning rates, prey size, and prey type) of Mediterranean blue tits Cyanistes caeruleus. Total provisioning rates were lowest when broods were reduced in size and greatest when broods were enlarged. Mean prey size was also affected by the brood size changes: parents tended to bring larger prey when confronted with low brood demand reinforcing the view that a trade‐off exists between minimizing foraging time and maximizing food quantity. Such differences in feeding frequencies and the load sizes delivered may be explained by changes in the parents’ foraging tactic. Increase of brood size compelled parents to work harder and be less selective in prey choice; we found that stressed birds with a high level of feeding responsibility (hungry nestlings) opted to concentrate on more readily available food items (Tortricids). On the other hand, their immediate reaction when faced with a low level of feeding responsibility was to decrease this prey type in the diet, so that the percentage of other preys (Noctuids) in the diet increased. There was no intersexual difference in the way in which parents responded to the manipulation. In sum, our results revealed a flexibility in foraging strategies of blue tits to cope with changing scenarios, which supports the idea that provisioning behavior is largely governed by nestling demand.     

Properties of food loads delivered to nestlings in the pied flycatcher: effects of clutch size manipulation, year, and sex

Experimental manipulation of the number of altricial offspring is supposed to modify parental expenditure in birds. In addition to the observed increase in parental feeding rate, it is also possible that the choice of prey or the size of load may change with the changing demand for food. Sexual differences in the provisioning response are also expected, on the basis of earlier studies. We examined the effect of brood size manipulation on choice of prey brought to nestlings and load size in the pied flycatcher. The composition and size of loads differed between years, possibly depending on varying availability of different prey types. Males responded to brood size enlargement by gathering heavier loads, whereas females showed no response. The alteration of load size in males was not explained by a larger number of prey items or mean prey size, but was a combination of these components. It is likely that males also increased their work rate in response to increased food demand at the nest. The absence of response in females might be because they are unable to increase work rate any further, or because food delivery rate in females can not be optimized by changing load properties. Received: 18 December 1997 / Accepted: 1 March 1998     

Parental Self-Feeding Effects on Parental Care Levels and Time Allocation in Palestine Sunbirds

The trade-off between parents feeding themselves and their young is an important life history problem that can be considered in terms of optimal behavioral strategies. Recent studies on birds have tested how parents allocate the food between themselves and their young. Until now the effect of food consumption by parent birds on their food delivery to their young as well as other parental activities has rarely been studied. I have previously shown that parent Palestine sunbirds (Nectarinia osea) will consume nectar and liquidized arthropods from artificial feeders. However, they will only feed their young with whole arthropods. This provided a unique opportunity to experimentally manipulate the food eaten by parents independent of that fed to their offspring. Here, I hypothesized that parents invest in their current young according to the quality of food that they themselves consume. Breeding pairs with two or three nestlings were provided with feeders containing water (control), sucrose solution (0.75 mol) or liquidized mealworms mixed with sucrose solution (0.75 mol). As food quality in feeders increased (from water up to liquidized mealworms mixed with sucrose solution): 1) Parents (especially females) increased their food delivery of whole arthropod prey to their young. 2) Only males increased their nest guarding effort. Nestling food intake and growth rate increased with increasing food quality of parents and decreasing brood size. These results imply that increasing the nutrient content of foods consumed by parent sunbirds allow them to increase the rate at which other foods are delivered to their young and to increase the time spent on other parental care activities.     

Cost of reproduction and allocation of food between parent and young in the swift (Apus apus)

We manipulated brood sizes to promote different levels of parentaleffort in the common swift (Apus apus). This provided a powerfulmethod for testing hypotheses regarding parental investmentdecisions concerning optimal allocation strategies between parentsand young. Data were analyzed on a visit-by-visit basis regardingchanges in parental and chick body mass, the mass of prey delivered,and the estimated mass of parental self-feeding. Our resultswere consistent with current theory in that food delivery increasedwith brood size, whereas the food received per chick, and hencemean chick body mass, decreased with brood size. Parental bodymass decreased with brood size and increasing parental effortbut recovered quickly during lower levels of chick feeding immediatelybefore fledging, suggesting some short-term cost of reproduction.Parents feeding at the highest level experienced criticallylow body mass and responded by a temporary cessation of chickfeeding. On any one foraging trip, total mass of prey captureddid not differ between brood sizes, but load mass deliveredto the young was negatively related to the amount of estimatedparental self-feeding. Allocation decisions of parents feedingthemselves and their young matched differential allocation theories,but estimated provisioning efficiency of parents at differentbody masses did not suggest any adaptive advantage from parentalmass loss.     

Biparental care: short-term manipulation of partner contribution and brood size in the starling, Sturnus vulgaris

In species with biparental care, the evolutionarily stable investmentstrategy depends, among other factors, on the reproductive valueof the brood and on the contribution of one's partner. A shortfallin work rate by one parent should be partially compensated forby its partner, while both parents should invest more effortwhen there are more young. We simultaneously tested these predictionsby attaching weights to one member of pairs of European starlings(Sturnus vulgaris), thus reducing one partner's nest visitationrate, and by daily manipulation of brood sizes (between threeand seven chicks). Both sexes compensated to an equivalent degreefor their partner's lower work rate, while at the same timeadjusting flexibly to the daily brood size changes by increasingvisit rates to larger broods. However, this ability to compensatewas limited in the larger brood sizes, perhaps due to an upperlimit on provisioning rate. Weighted birds and their partnersshowed differences in prey types delivered, relative to controls,taking a lower proportion of leatherjackets. With regard tothe proximate cues affecting parental provisioning rate, beggingnoise levels (automatically recorded by computer from microphonesin the nest-box) increased with brood size, but average lengthof begging bout did not. As expected, visit rates per chickdecreased as brood size increased, and this was reflected inlower weight gains per day by chicks in larger broods. Theseresults agree in general with games theory models but revealimportant departures from our previous work with respect to(1) parents reaching an apparent ceiling to nest visitationrate at high brood sizes and (2) the ability to track thesebrood demands with short-term adjustments.     

Parental provisioning behaviour in a flock-living passerine, the Vinous-throated Parrotbill Paradoxornis webbianus

The amount of food delivered by parents to their chicks is affected by various life history traits as well as environmental and social factors, and this investment ultimately determines the current and future fitness of parents and their offspring. We studied parental provisioning behaviour in the Vinous-throated Parrotbill Paradoxornis webbianus, a species with an unusual social system that is characterised by flock-living, weak territoriality and variable nesting dispersion. Parental provisioning rate had a positive influence on chick mass gain, suggesting that provisioning rate is an effective measure of parental investment in this species. Males and females fed nestlings at approximately the same rate, and no other carers were observed at nests. Parents coordinated provisioning rates so that they mostly fed chicks synchronously. However, the extent to which parents coordinated provisioning was associated with their social environment, synchrony being positively related to local breeding density and negatively to nearest neighbour distance. The rate at which parents provisioned nestlings showed the same relationships with social measures, being greatest at higher density and when neighbours were closer. Visit rate was also related to chick age, but not to brood size, brood sex ratio, extra pair paternity, laying date, temperature, parents’ body characters, time of day or year. We conclude that a breeding pairs’ social environment plays an important role in determining parental investment, probably through its effects on the opportunities that parents have for foraging with conspecifics.     

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.     

Parental Sex Differences in Food Allocation to Junior Brood Members as Mediated by Prey Size

Individual offspring within a brood may receive different amounts of provisioning from the male and female parents. Some hypotheses suggest that this bias is the result of an active and adaptive choice by parents. An alternative hypothesis is that feeding biases arise as a result of a constraint of fitting large prey items into small gapes. In an experiment with pied flycatchers, Ficedula hypoleuca , we tested for sex-biased allocation to junior nestlings in asynchronous broods and whether this could be explained by active parental choice or by passive allocation according to prey size and gape size. In both control broods and broods with experimentally increased degree of asynchrony, prey types did not differ between parents but females brought smaller prey than males at younger but not older nestling stages. At younger but not older nestling stages, the majority of feeds to junior nestlings were from females, and the smaller nestlings consumed smaller prey than older siblings. However, there was no evidence of active preference of small nestlings by females as parents did not differ in the tendency to bypass a begging senior nestling in order to feed a junior nestling. Provisioning rates by females were lower than those by males when nestlings were young and we suggest that foraging time constraints caused by the need to brood offspring result in females bringing smaller prey than males. In turn, the larger prey brought by males was more often transferred to larger offspring after the smaller ones failed to swallow it. In such cases, 'preferential' feeding of small nestlings by females may simply be a passive side effect of foraging constraints and gape-size limitations.     

Parent-offspring interactions in food provisioning of Manx shearwaters: implications for nestling obesity

Procellariiform seabirds such as the Manx shearwater Puffinus puffinus, rear only one chick at a time but may breed many times in their lives; parents should thus limit food delivery to the chick in keeping with the balance between current and future reproductive output. Yet procellariiform chicks accumulate large quantities of lipid, which may provide a buffer against pronounced and unpredictable variation in food provisioning, resulting in part from an inability of parents to regulate food supply to the nest. We switched chicks between nests to examine the roles of parents and offspring in controlling food delivery. The serial autocorrelation in age-specific body masses for unmanipulated chicks decreased from 0.61 (P< 0.01) to 0.35 (NS) over a period of 15 days and remained nonsignificant thereafter. By contrast, the serial autocorrelation for switched chicks increased from 0.64 (P< 0.01) to 0.83 (P< 0.001) and the serial cross-correlation rose from 0.23 (NS) to 0.50 (P< 0.05). These results supported both chick determination and parental determination models of food provisioning, indicating that chicks conveyed information about their nutritional status, which parents acted upon by adjusting their rate of food delivery. We discuss these results in relation to the optimization of nestling lipid reserves and parental foraging effort. We suggest that information conveyed by the chick's begging intensity serves to reduce the provisioning rate to well-fed chicks, but parents cannot or do not increase food provisioning to poorly fed chicks. Such adjustment of food provisioning does not refute the hypothesis that nestling obesity provides a buffer against highly variable food delivery. Copyright 1999 The Association for the Study of Animal Behaviour.     

Intensity of mouth coloration in Blackcap Sylvia atricapilla nestlings affects food distribution among siblings but not provisioning of the whole brood

Among various begging stimuli, mouth coloration has received increasing attention in recent years, and previous research has demonstrated that mouths of nestling Canaries Serinus canaria get redder with the extent of food deprivation and that parents preferentially feed nestlings of redder gapes. This study assesses whether the intensity of red mouth colour in nestling Blackcaps Sylvia atricapilla is a signal in parent–offspring communication. This is one of the few species with a naturally red gape in which the function of mouth redness has been tested. Three predictions were experimentally tested: (1) reddening the gape of a single nestling within a brood increases its provisioning in relation to other siblings; (2) reddening the gapes of all nestlings within a brood increases parental feeding rate; and (3) food deprivation increases nestling mouth redness. The effect of nestling quality on mouth redness was also assessed. The intensity of gape coloration affected food distribution, but in a way opposite to that expected: an increase in mouth redness of the nestling caused reduced feeding by parents. However, reddening the gapes of all nestlings had no effect on provisioning of the whole brood, suggesting that Blackcap parents use different cues for provisioning particular nestlings and the whole brood. Intensity of mouth redness in Blackcap nestlings was not affected either by food deprivation or by nestling quality in terms of mass and rank in the nest.     

Parental provisioning in a variable environment: Evaluation of three foraging currencies and a state variable model

We estimated the reproductive success of black terns (Chlidonias niger) based on three optimal foraging currencies (maximizing the net rate of energy intake, daily delivery rate, and efficiency, respectively) and a state variable model. There was a broad range of capture intervals (the time required for the parent to capture a single prey) when the flight speeds predicted by the three currencies were so high that they resulted in daily provisioning costs which parents could not fully recover through self-feeding. Whenever the efficiency currency produced higher estimates of reproductive success, parents lost comparatively less weight than when they foraged as rate-maximizers. If parents did not experience any weight loss, the net rate and efficiency currencies made equivalent fitness projections. However, both of these currencies provided lower fitness returns than daily delivery rate at longer capture intervals. There were a number of capture intervals when estimates of reproductive success from the state variable model and at least one of the foraging currencies were equal. Provisioning behaviour under the state variable model was much more flexible and parents were therefore able to reduce their self-feeding rate on days when food was particularly scarce, thereby increasing the total delivery to the nest. This resulted in higher fitness returns than was possible under the foraging currencies. Our results suggest that efficiency-maximizing is more likely to provide fitness returns that are equivalent to the state variable model in comparison with the rate-maximizing alternatives. Furthermore, only the efficiency currency and the state variable model made predictions of flight speed that were similar to speeds measured in black tern parents provisioning young at natural nests.     

An energetic correlate between colony size and foraging effort in seabirds, an example of the Adélie penguin Pygoscelis adeliae

Central-place foraging seabirds alter the availability of their prey around colonies, forming a "halo" of reduced prey access that ultimately constrains population size. This has been indicated indirectly by an inverse correlation between colony size and reproductive success, numbers of conspecifics at other colonies within foraging range, foraging effort (i.e. trip duration), diet quality and colony growth rate. Although ultimately mediated by density dependence relative to food through intraspecific exploitative or interference competition, the proximate mechanism involved has yet to be elucidated. Herein, we show that Adélie penguin Pygoscelis adeliae colony size positively correlates to foraging trip duration and metabolic rate, that the metabolic rate while foraging may be approaching an energetic ceiling for birds at the largest colonies, and that total energy expended increases with trip duration although uncompensated by increased mass gain. We propose that a competition-induced reduction in prey availability results in higher energy expenditure for birds foraging in the halo around large colonies, and that to escape the halo a bird must increase its foraging distance. Ultimately, the total energetic cost of a trip determines the maximum successful trip distance, as on longer trips food acquired is used more for self maintenance than for chick provisioning. When the net cost of foraging trips becomes too high, with chicks receiving insufficient food, chick survival suffers and subsequent colony growth is limited. Though the existence of energetic studies of the same species at multiple colonies is rare, because foraging metabolic rate increases with colony size in at least two other seabird species, we suggest that an energetic constraint to colony size may generally apply to other seabirds.     

Foraging Trade‐offs between Prey Size,Delivery Rate and Prey Type: How Does Niche Breadth and Early Learning of the Foraging Niche Affect Food Delivery?

Optimal foraging theory suggests that avian parents should prefer the most energetically efficient (largest) prey items when delivering food to offspring at a central place. However, during periods of high demand, selectivity of prey may decline, leading to the delivery of smaller and/or less nutritious items. We compared foraging trade‐offs between great tits (Parus major) which had a wider feeding niche than blue tits (Cyanistes caeruleus). We also compared the foraging efficiency of cross‐fostered young, which had learned the spatial foraging niche and prey size of the foreign species, to that of control conspecifics. Mean delivery rates did not differ between control and cross‐fostered parents of either species but as delivery rates increased, prey size declined for both species and both treatment groups. However, across the range of increasing delivery rates, parents were able to increase the total biomass of prey delivered. Cross‐fostering did not alter the proportion of different prey taxa in the diet, but cross‐fostered birds shifted the size of the prey taken to that of their foster species. Consistent with their broader feeding niche, great tits, but not blue tits, incorporated more unpalatable items (flies) as delivery rates increased. Although great tits foraged less efficiently in the blue tit niche, paradoxically, blue tits seem to deliver more prey biomass when foraging in the great tit niche.     

Prey preparation by adult Great Tits Parus major feeding nestlings

Some birds prepare food items before giving them to their nestlings. We studied the relationships between the degree of prey preparation and prey size, nestling age, brood size and time of season. We estimated the degree of preparation of 513 animal prey items, taken by using neck collars, brought to nestling Great Tits Parus major. Prey preparation increased with prey size and decreased as the nestlings grew older, as brood size increased and as the season progressed. Other factors, such as nutrient concentration (through removal of low-quality or deleterious parts) or palatability (considering scaly moth forewings unpalatable), seem also to be important in determining prey preparation. Our results suggest that the degree of prey preparation is a compromise between the benefits gained by the nestlings (ingestion and digestion of prey is facilitated) and the costs to the parents (mainly time allocated to prey preparation).     

Does resource availability affect offspring begging and parental provisioning in a partially begging species?

In species where parents repeatedly provide their offspring with food, the offspring often communicate their need to the parents. Burying beetles, which breed on a wide size range of carcasses of small vertebrates, are interesting model systems to test theories on begging, because the larvae show partial begging, that is, they obtain food through both signalling to their parents (begging) and feeding directly from the carcass. We manipulated resource availability inNicrophorus vespilloides by providing parents with mouse carcasses spanning a wide size range, and allowing them to rear the larvae that hatched, so that both the amount of resources and the number of siblings varied. Time spent begging by each larva was strongly influenced by the time parents spent near the larvae. Brood size had a nonlinear effect on larval begging, with begging increasing with brood size for relatively smaller broods and decreasing again for larger broods. Carcass size and number of parents present had no effect on begging. Time spent provisioning the larvae by the parents was strongly associated with the time spent begging by each larva. Parents spent more time provisioning under biparental care than under uniparental care, while brood size and carcass size had no significant effect. These findings suggest that the larvae adjust their begging to the behaviour of their parents and the number of siblings, but not to the amount of resources. Furthermore, parents adjust the time spent provisioning to the average amount of begging by each larva in the brood, and not to the availability of resources.     

Provisioning behaviour at the nest in single-parent versus biparental nests and male versus female parents in the common redstart (<Emphasis Type="Italic">Phoenicurus phoenicurus</Emphasis>)

We analysed video-sequences of undisturbed parental provisioning behaviour on 12 nests of common redstart (Phoenicurus phoenicurus). In 4 of the 12 nests, chicks were fed by a single parent only. We compared provisioning rate of chicks, time spent on the nest and food allocation rules between nests with uniparental and biparental care and between male and female parents in biparental nests. In nests with a single parent, the frequency of feeding visits per parent was higher than in biparental nests. As a result, the rate of food provisioning of chicks was similar in uniparental and biparental nests. The food allocation rules did not differ between uniparental and biparental nests. In biparental nests, male and female provisioning behaviour was similar though with two exceptions: males had a strong preference for feeding chicks in front positions in the nest and females spent a longer time on the nest after feeding. We conclude that single common redstart parents are able to compensate fully for the absence of the other parent through increased provisioning efforts, and that in biparental nests, males and females contribute equally to the provisioning of the young.     

Azure‐winged Magpies Cyanopica cyanus trade off reproductive success and parental care by establishing a size hierarchy among nestlings

It is common in birds that the sizes of nestlings vary greatly when multiple young are produced in one nest. However, the methods used by parents to establish size hierarchy among nestlings and their effect on parental provisioning pattern may differ between species. In the Azure‐winged Magpie Cyanopica cyanus, we explored how and why parents controlled the sizes of nestlings. Asynchronous hatching was the main cause of size hierarchy within the brood, although the laying of larger eggs later in the laying sequence reduced this effect. Parents with asynchronous broods produced more eggs and fledged more nestlings than those with synchronous broods but their brood provisioning rates, food delivery per feeding bout and feeding efficiency did not differ. We performed a cross‐fostering experiment to synchronize some asynchronous broods. Provisioning rates of asynchronous broods were lower than those of synchronized broods, but the daily growth rates and fledging body mass of their nestlings were not different. Our findings indicate that parents of asynchronous broods can achieve higher reproductive success than those of synchronous broods based on the same parental care, and the same reproductive success as those of synchronized broods based on less parental care. It appears that parent birds can better trade off reproductive success and parental care by establishing a size hierarchy among nestlings.     

The manipulation of food resources reveals sex-specific trade-offs between parental self-feeding and offspring care

Parent Palestine sunbirds (Nectarinia osea) feed on flower nectar that is not fed to their nestlings. This phenomenon provided a unique opportunity to manipulate self-feeding rates of parent birds independently of the rate at which they feed arthropod prey to their offspring. Based on provisioning models, we predicted that parents would invest more in their young as the energy content of their own food increased. From our earlier work, we also predicted that the levels of sex-specific activities of males and females would differ as the energy content of their food increased. Sunbird pairs with two or three nestlings were provided with feeders containing a low-, medium- or high-concentration sucrose solution. As the sugar concentration increased, the females delivered arthropods at a greater rate to their nestlings, removed proportionally more faecal sacs and spent longer at the nest, while the males increased their mobbing effort. Nestling food intake and body mass, but not tarsus length or bill size, were larger in small broods than in large broods, and increased with increasing feeder sugar concentration. These results imply that increasing the energy content of food consumed by parent sunbirds allows them to increase the rate at which other foods are delivered to their young and to increase other parental care activities as well. The results also add credence to the idea that behavioural decisions reflect life-history trade-offs between parental self-feeding and investment in current young.