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.     

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 .     

Energy expenditure, nestling age, and brood size: an experimental study of parental behavior in the great tit Parus major

A brood manipulation experiment on great tits Parus major was performedto study the effects of nestling age and brood size on parentalcare and offspring survival. Daily energy expenditure (DEE)of females feeding nestlings of 6 and 12 days of age was measuredusing the doubly-labeled water technique. Females adjusted theirbrooding behavior to the age of the young. The data are consistentwith the idea that brooding behavior was determined primarilyby the thermoregulatory requirements of the brood. Female DEEdid not differ with nestling age; when differences in body masswere controlled for, it was lower during the brooding periodthan later. In enlarged broods, both parents showed significantlyhigher rates of food provisioning to the brood. Female DEE wasaffected by brood size manipulation, and it did not level offwith brood size. There was no significant effect of nestlingage on the relation between DEE and manipulation. Birds wereable to raise a larger brood than the natural brood size, althoughlarger broods suffered from increased nestling mortality ratesduring the peak demand period of the nestlings. Offspring conditionat fledging was negatively affected by brood size manipulation,but recruitment rate per brood was positively related to broodsize, suggesting that the optimal brood size exceeds the naturalbrood size in this population.     

Great tits, Parus major, lay too many eggs: experimental evidence in mid-boreal habitats

This study shows that great tits lay too large clutches in mid‐boreal habitats. First, breeding success, measured with number of fledglings or proportion of eggs that produce fledglings, in northern Finland (65°N) is much poorer than in central and western Europe. Second, brood size manipulations (ca ±30% of the natural mean) revealed that reduced broods produced equal numbers of and larger‐sized fledglings than control and enlarged broods, giving thus the best fitness value for reduced broods. Third, parents of enlarged broods could not adjust (i.e. increase) their feeding effort to the greater number of nestlings. Fourth, extra feeding (about 1/3 of the theoretical maximal needs of the nestlings) during the nestling period resulted in more numerous and larger‐sized fledglings in comparison to control broods. We suggest that the ultimate explanation for the too large clutches is gene flow from the southern population, which prevents local adaptations in the north. Consequently, the main reason for food limitation during the nestling period is that northern great tits apply “southern” decision rules for timing of breeding, clutch size and foraging behaviour. Thus, they tend to breed too early in comparison to the food abundance peak, lay too large clutches in comparison to the level of resources and, perhaps, forage on a too narrow diet (75% caterpillars). Since the late broods that matched the local food abundance peak did not succeed better than the mismatched earlier ones, the most crucial fault of northern great tits seems to be that they overestimate food abundance during peak demands and lay too large clutches. Another explanation for this could be that northern great tits have adopted a brood reduction strategy. However, the long‐term data reveal that years of high breeding success, which would maintain large clutches in the population, are very rare in the north. Therefore, it is unlikely that a brood reduction strategy per se could explain the phenomenon. Instead, it could work together with the gene flow against local adaptation for clutch adjustment.     

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.     

Responses by Central‐Place Foragers to Manipulations of Brood Size: Parent Flickers Respond to Proximate Cues but do not Increase Work Rate

Manipulations of brood size measure the willingness or ability of parents to invest in offspring and different reproductive roles may lead to differences in feeding effort between the sexes. Parental investment in birds is usually assessed by quantifying feeding rates, but this provides an incomplete picture of parental effort because it fails to account for how parents collect food on the landscape. We studied northern flickers (Colaptes auratus), a woodpecker in which males provide the majority of parental care and used a repeated measures design and short‐term (24 h) brood enlargements (N = 35) and reductions (N = 27) to assess effects of treatment on feeding rates to nestlings and parental foraging behaviour. Parents of enlarged broods did not significantly increase feeding rate, resulting in a decline in nestling mass. Parents of reduced broods decreased their feeding rates by 84%, but increased per capita feeding rates, resulting in nestling mass gain. The variation in feeding rates to enlarged broods was not influenced by feather corticosterone, body condition, feather re‐growth rate or mass change between the incubation and nestling periods. Foraging pattern on the landscape remained the same during the enlarged treatment for both sexes. We conclude that flickers respond to proximate cues in brood demands, but do not increase feeding rates to enlarged broods, at least in the short term. A literature review suggested that this lack of response is atypical for short‐lived species. We hypothesize that parents in species with large home ranges and long nestling periods face energetic limitations that constrain their ability to respond to enlarged broods. We encourage future studies to assess foraging behaviour on the landscape to document important trade‐offs for parents such as predation risk and energy expenditure while feeding offspring.     

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.     

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.     

Costs and benefits of parental care in eastern kingbirds

We manipulated brood sizes of eastern kingbirds (Tyrannus tyrannus)to measure the costs and benefits of parental care and to testwhether kingbirds showed evidence of individual optimizationof reproductive effort. We found that the number of feedingtrips (trips/h) increased and that per capita feeding rates(trips/nestling/h) declined as brood size increased. The declinein per capita feeding rates was mostly due to high feeding rateto broods of one: parents made roughly equal number of tripsto feed each nestling in broods of two to five. Nonetheless,nestling mass declined with brood size, probably because largebroods were fed more small prey. Nestling condition (mass adjustedfor structural size) differed only between broods of one andfive. After controlling for effects of brood size, feeding rateshad no supplementary influence on either nestling size or condition,but productivity and feeding rate were positively and significantlyrelated. Adult male condition did not vary with brood size,manipulated brood size, or total feeding rate, but declinedas the pair's per capita feeding rates increased. In addition,males that returned to breed were in better condition beforeleaving for migration than those that failed to return. Femalecondition tended to decline, and the probability of returningto breed dropped when broods were enlarged. However, femalecondition was independent of the probability of returning. Ourresults show that high feeding rates were costly, but that theycarried benefits (greater productivity). Some evidence for individualoptimization of reproductive effort existed: variability innestling and adult female condition were better explained bychanges in brood size than by the actual number of young inthe nest. However, most evidence supported the alternative thatincreased brood size was equally costly for all birds     

Relationships between brood size and parental provisioning performance in chinstrap penguins during the chick guard phase

 Chinstrap penguins (Pygoscelis antarctica) normally lay two eggs, but brood size is often reduced by mortality during incubation or after hatching. We hypothesized that this variation in brood size would affect the parents’ foraging behavior and their chick provisioning performance. We studied patterns of adult foraging trip duration and frequency, food load delivery, and chick growth rates in relation to brood size during the guard phase in four breeding seasons (1991–1994) on Seal Island, Antarctica. Within a given year, parents with two chicks made more frequent foraging trips to sea and may have transported larger food loads to the nest; however, the duration of foraging trips was unrelated to brood size. Overall, parents with two chicks spent ∼15% more time at sea than parents with only one chick. Both the frequency and duration of foraging trips varied between years. Foraging trip duration may partly reflect the birds’ foraging radius, which probably varies with time in response to shifts in krill distribution. Chick growth rate varied betwen years, but was related to brood size only in 1992, when chicks from two-chick broods grew significantly more slowly than chicks from one-chick broods. Food loads transported to chicks, as well as chick growth rates, were highest in 1994, when concurrent hydroacoustic studies indicated that regional krill biomass was severely depressed. This apparent anomaly suggests that the spatial scale of the krill survey may have been too coarse to detect some high-density krill aggregations within the penguins’ foraging range. Received: 26 September 1995 / Accepted: 12 May 1996     

Food Provisioning in Red-Necked Grebes (Podiceps Grisegena) at Common Carp (Cyprinus Carpio) Ponds

Parental feeding patterns were studied in red-necked grebe (Podiceps grisegena) broods throughout the entire period of parental care in a common carp (Cyprinus carpio) fish-pond area in SE Poland in 1993–2002. Fish formed a substantial part of prey provided to the flightless young from their second week of life. Although the numbers of large invertebrates and tadpoles, the alternative prey to fish, did not decrease during the chick rearing period, grebe parents gradually shifted from delivering predominately invertebrates to delivering fish, and the average size of fish fed to chicks increased with brood age. Broods with relatively high fledging success (at least two chicks fledged) had a larger proportion of fish in their diet than broods seriously reduced because of undernourishment. The dive duration of foraging grebe parents did not differ between carp, wild fish and non-fish prey, but carp prey required significantly more time for handling. The percentage of prey rejected by chicks increased over the prefledging period from 2 to 24%. Of the prey rejected, 82% were fish apparently too large for the young to swallow. Fish prevalence in the diet of red-necked grebe chicks at carp ponds contradicts the results of other studies on the feeding habits of the nominative subspecies during breeding season. However, the red-necked grebe is a gape-limited predator and the piscivory of the chicks is limited to small-bodied fish.     

The importance of a main dish: nestling diet and foraging behaviour in Mediterranean blue tits in relation to prey phenology

Insectivorous birds rely on a short period of food abundance to feed their young; they must time their reproduction to match the timing of Lepidoptera larvae, their main prey. Apart from the net result (i.e. birds are timed or mistimed with respect to the food's peak), an important aspect is the possible influence of other factors, such as the seasonality of the environment or the abundance and diversity of species contributing to the caterpillar peak, on birds' phenology and their ability to cope well with unpredictable food supplies. In a 2-year study, we explored the seasonal variation of nestling diet in Mediterranean blue tits Cyanistes caeruleus and how reproductive parameters (nestling condition, provisioning rates) are affected by the phenology and composition of food. We also examined the influence of the synchrony between offspring needs and caterpillar peak in shaping the composition of the nestlings' diet. We found that the effect of synchrony on nestling condition varied between years which may be partially due to differences in food peak attributes. The adequacy of birds' timing in relation to prey phenology affected foraging decisions; those birds that were not able to correctly adjust their timing were forced to rely on less preferred prey (tortricids). In this sense, we found that relative contribution of tortricids (smaller caterpillars but easier to get) and noctuids (preferred prey but more difficult to find) to the diet influenced nestling condition and parental provisioning effort; parents performed fewer feeding events and reared heavier nestlings as the contribution of noctuids to the diet increased. The relationship between the proportion of caterpillars and nestling mass was curvilinear, whereas that parameter was negatively affected by the percentage of pupae. Our results show how changes in diet composition may contribute to explain the effect of mismatching on birds' breeding performance.     

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.     

Chick parasitism by blowflies affects feeding rates in a Mediterranean population of blue tits

Offspring fitness depends on interactions between parental care and environmental constraints. It has been suggested that in altricial birds parents are able to compensate for the detrimental effects of ectoparasites by improving food provisioning. We tested this prediction in a population of blue tits highly parasitized by blowfly larvae. The frequency of parental feeding visits was significantly higher in parasitized broods than in broods experimentally deparasitized. Despite a strong increase in parental care, chicks of parasitized broods were lighter, smaller, and more anaemic than chicks in deparasitized broods. Parents invest more in feeding parasitized young but cannot fully compensate for the negative effects of parasites, hence young are in poor condition at fledging.     

Importance of feeding ecology to the reproductive success of Blackbirds Turdus merula nesting in rural habitats

The feeding ecology of Blackbirds Turdus merula breeding in contiguous woodland and farmland habitats was studied over three years. The aim of the study was to investigate how reproductive success was influenced by nestling diet and the provisioning rates of parents feeding nestlings. Parental provisioning rates increased with brood size, and consequently individual nestlings were no lighter in larger broods. None of the environmental factors measured had strong effects on parental provisioning rate. The nestling diet was dominated by caterpillars and earthworms, the former occurring in a short period in the middle of the breeding season. The availability of earthworms was higher in woodland and was dependent on rainfall in farmland. Nestling mass and provisioning rates were marginally higher under predominantly earthworm diets. Nestling mass increased with rainfall in farmland only, and was higher in farmland than in woodland or woodland-edge, although it is doubtful whether this result is of any significance for fledgling survival. Overall, Blackbirds were able to provision their nestlings adequately throughout the breeding season across a range of conditions. There was no evidence to suggest that reproductive success was constrained by aspects of feeding ecology within the natural range of brood size.     

Parental care trade‐offs in the inter‐brood phase in Barn Swallows Hirundo rustica

In seasonal environments with limited time and energy resources, double‐brooded birds face trade‐offs in the timing of their two reproductive attempts and in the effort allocated to the first and the second broods. In the Barn Swallow Hirundo rustica a long care period for the first brood enhances the survival of first‐brood chicks, but also delays the start of the second brood, which in turn reduces the survival prospects of second‐brood chicks. Probably as a response to this trade‐off, double‐brooded Barn Swallows reduce the period of post‐fledging care for first‐brood fledglings. By radiotracking whole families, we investigated the determinants of this behaviour and its consequences for the survival of the first‐brood fledglings. The end of the females’ investment in post‐fledging care of the first brood was related to the beginning of egg synthesis for the second clutch. With the start of egg synthesis, females significantly reduced provisioning rates to the first‐brood fledglings to less than one‐fifth of the previous rates, while the proportion of time they spent foraging remained high. Assuming that the females’ foraging success was constant, we conclude that their energy income was allocated to egg production rather than fledgling provision. Males did not compensate for the females’ reduced feeding rates. Thus the start of egg production for the second clutch had a marked effect on the quantity of food received by first‐brood fledglings. In parallel with the changes in parental behaviour and provisioning rates, we observed a marked drop in the daily survival rate of first‐brood chicks. These results support the hypothesis that females face a strong trade‐off in the allocation of energy to subsequent broods. Energy allocation to a second clutch involves a cost in terms of reduced provisioning, and as a result the survival of first‐brood chicks is compromised. This is probably outweighed by the improved success of an early second brood.     

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.     

Reproductive effort and T-lymphocyte cell-mediated immunocompetence in female pied flycatchers Ficedula hypoleuca

The physiological mechanism underlying the cost of reproduction may consist of immunodepression caused by increased parental effort. Here, we report effects of experimental manipulation of clutch size on T-lymphocyte cell-mediated immune response in female pied flycatchers, Ficedula hypoleuca. Parents with reduced broods provisioned at lower rates than those caring for control and enlarged broods three days after hatching. Parents caring for enlarged broods provisioned nests at higher rates 13 days after chick hatching than those feeding control and reduced broods. Females with enlarged broods weighed less than females with control or reduced broods. No effect of experimental treatment on nestling mass and size was found. The response to the injection of phytohaemagglutinin in the wing-web of females decreased with increasing brood size and with increasing provisioning rate when the chicks were three days old, when controlling for the negative effect of female mass on response. The T-lymphocyte cell-mediated response decreased from the reduced to the control, and from this to the enlarged group, when controlling for female mass. This effect of experimental manipulation of clutch size was significant and consistent with a trade-off between maternal effort and immunocompetence.     

Facultative Adjustment of Brood Sex Ratio in Response to Indirect Manipulation of Behaviour

Sex allocation theory states that parents should adjust their offspring sex ratio according to the expected fitness returns from sons and daughters. Several recent studies indicate that such adaptive manipulation of offspring sex ratio is achievable, and that it may be influenced by e.g. morphological characters. Here we manipulate behaviour through interspecific cross-fostering of great tits ( Parus major ) and blue tits ( Cyanistes caeruleus ), and investigate its effect on the offspring sex ratio of adults that were themselves cross-fostered as chicks. The experience of being raised by a different species has previously been shown to result in aberrant species assortative behaviour and song, and a lowered dominance status during winter. Brood sex ratios of conspecifically breeding pairs with and without cross-fostered members were compared. Broods with at least one cross-fostered parent contained significantly more males than did control broods. Sex of cross-fostered parents did not influence the brood sex ratio. We conclude that female great tits and blue tits seem to be able to adjust the sex ratio of their broods, and that changes in their own or their partners' behaviour may elicit such adjustments.     

Brood size and body condition in the House Sparrow Passer domesticus: the influence of brooding behaviour

In many bird species, females undergo a marked decline in body condition during the first days of the nestling period. This decline may be because brooding young chicks reduces the time available for foraging. Alternatively, it might be viewed as an adaptive way to reduce flight costs when the food demand of the brood is highest. To test these hypotheses we modified the brooding commitment of House Sparrows Passer domesticus by manipulating brood size to see if changes in time spent brooding affects adult body condition. During the nestling period, females provided on average three times as much brooding as males. Reduced broods received 14% more brooding than large broods and time spent brooding declined with brood size and chick age according to an exponential decay function. Male body condition was unaffected by brood size and remained stable throughout the reproductive period. Body condition of females with enlarged broods decreased gradually during the nestling period, whereas that of females tending reduced broods dropped abruptly and significantly upon hatching. This resulted in females with reduced broods having lower body condition during the first half of the nestling period than those with enlarged broods. The sharp drop in body condition of females with reduced broods coincided with the period that brooding was most intensive. Indeed, female body condition at the end of the nestling period was negatively correlated with the proportion of time they spent brooding during the first half of the nestling period. Thus, the probable lower homeothermic capacities of reduced broods implies a higher brooding commitment for female House Sparrows that, in turn, may reduce their opportunity to forage and consequently also their body condition.