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.     

Diet of Starling Sturnus vulgaris nestlings on farmland: the importance of Tipulidae larvae

Capsule?Tipulidae larvae are a key resource for adult Starlings provisioning their young on lowland farmland.

Aims?(i) To describe Starling nestling diet on lowland farmland. (ii) To examine the effects of brood size variation and nestling age on Starling parental food provisioning.

Methods?Over 4 years, we observed parental food provisioning at 42 nests across 4 sites on lowland farmland in Oxfordshire, UK using nestbox video recorders.

Results?Tipulidae larvae were the most frequent prey item recorded in provisioning loads (52%), the next most frequent were winged insects (11%), then Coleoptera larvae (6%), Lepidoptera larvae (5%) and Lumbricidae (5%), with 21% of prey unidentified. Estimates of prey lengths, dry mass and total energy content also confirmed that Tipulidae larvae were the key prey source. Generalized linear mixed-effects models showed that larger broods received fewer Tipulidae larvae per nestling per day and that the proportion of Tipulidae larvae in the diet did not vary with brood size or nestling age.

Conclusion?Our results support the idea that Tipulidae larvae are the primary dietary item for nestling Starlings on lowland farmland.     

Great Spotted Cuckoo Nestlings but not Magpie Nestlings Starve in Experimental Age‐Matched Broods

Nestlings of non‐evicting avian brood‐parasites have to compete for food with foster parents' own nestlings. The outcome of these competitive contests is determined mainly by body size differences between parasitic and host nestlings. As part of the coevolutionary arms race between brood parasites and their hosts at the nestling stage, it has been reported that some host foster parents discriminate against parasitic chicks and are reluctant to feed them. Here, by experimentally creating size‐matched broods of different composition (only magpie Pica pica chicks, only great spotted cuckoo Clamator glandarius chicks or mixed broods), we show that great spotted cuckoo chicks starved in 20.2 per cent (17 of 84) of the parasitized magpie nests even in absence of size asymmetries, while in none (0 of 72) of the nests a magpie chick starved. As far as we know, this is the first record of non‐evictor brood parasitic nestlings starving without being smaller than their host nestmates in a frequently used host species. Nest composition had no effect on chick starvation. The cuckoo nestling starved even in two of the nests occupied by only one cuckoo chick. Our results could be explained by (1) magpies being reluctant to feed cuckoo chicks; (2) parasitic chicks receiving lower‐quality food items or cuckoo nestlings being sensitive to some particular component of the diet (e.g. cereal grains); and (3) the existence of cuckoo chick discrimination ability by magpie foster parents.     

Winter diet of Common Buzzards Buteo buteo on a Scottish grouse moor

Capsule: The winter diet of Common Buzzards Buteo buteo on a Red Grouse Lagopus lagopus scotica moor was dominated by small mammals, whilst grouse were a minor prey item.

Aims: To assess winter diet of Common Buzzards from pellets collected at roost sites on and around a managed Red Grouse moor, and to explore temporal, spatial and age-related variation in diet composition.

Methods: Forty-four winter roost sites were located during two winters using a combination of observations from vantage points and individual Common Buzzards equipped with either radio or satellite transmitters. Pellets were collected between October and March each winter and analysed to assess dietary composition.

Results: Small mammals were the main prey in both years, comprising 60–67% of items and occurring in 88–92% of pellets. Diet varied between years, with more lagomorphs and birds (passerines, corvids and pigeons) but fewer Red Grouse eaten when grouse abundance declined. Grouse formed 1.1% and 0.6% of prey items, and occurred in 3% and 2% of pellets from each winter, respectively.

Conclusion: Common Buzzards rely on small mammal prey during winter. When available, Red Grouse are a minor dietary component, the amount of which reflects their abundance in the environment. The opportunism of Common Buzzards can result in temporal variation in winter diet.     

The pattern of Dunlin Calidris alpina distribution and abundance in relation to habitat variation in the Flow Country of northern Scotland

Capsule Folivorous caterpillars constituted the majority of nestlings’ food in a primeval forest. Blue Tit broods only partially matched the caterpillar peak, and the mismatch did not affect food composition or nesting success.

Aims To describe factors influencing the timing of reproduction in Blue Tits under primeval conditions (Bia?owie?a National Park, Poland) and to check whether they schedule breeding so as to synchronize broods with a seasonal caterpillar peak.

Methods We gathered information on phenology of leaf development, seasonal availability of folivorous caterpillars (frass collection), timing of Blue Tit breeding, composition of its nestling food, and nest fate over a three-year period.

Results Caterpillars constituted c. 74% of nestling diet, but only 17–65% of broods matched the caterpillar peak in any season. Neither total nest loss, nor frequency of brood reduction depended on the level of mismatch. Caterpillar availability was probably adequate every year, regardless of the amount of mismatch, and no selective advantage of precise matching was detectable. Phenological events at all trophic levels occurred earlier in warmer springs. Egg-laying coincided with tree bud burst and appearance of caterpillars, but was not critically dependent on their timing.

Conclusion The observations are consistent with the view that Blue Tits under primeval conditions in Bia?owie?a National Park, Poland, breed as early as possible, rather than synchronizing their breeding with the caterpillar peak later in the season.     

Habitat selection and foraging success of Hen Harriers Circus cyaneus in west Scotland

The foraging behaviour of Hen Harriers breeding in west Scotland was studied in a variety of afforested and moorland habitats. Time budget information from sample areas, stratified by habitat type and controlled for nest distance, was used to investigate habitat selection. Preferences were then related to measures of foraging performance in each habitat. Hen Harriers foraged preferentially over young first rotation coniferous forests, and selected heathland and grassland habitats ahead of closed canopy woodland. Harrier preference for the various habitats was positively correlated with the frequency with which they struck at, and captured, prey. Prey strike success was much lower than that reported for moorland managed for Red Grouse Lagopus lagopus scoticus. The results explain many of the reported changes in local Hen Harrier populations. The extent of young first rotation forestry is currently in decline. It is predicted that this will lead to a reduction in numbers of Hen Harriers breeding in west Scotland, and an increase in the proportion of the British harrier population occupying moorland managed for Red Grouse.     

Corticosterone levels in host and parasite nestlings: Is brood parasitism a hormonal stressor?

Parasite chicks from non-evictor species usually try to monopolize host parental care, thereby increasing considerably the level of food competition in the nest. Here, we propose that brood parasitism is an important stressor for host and parasite nestlings and explore this hypothesis in the non-evictor great spotted cuckoo (Clamator glandarius) and its main hosts, the same-sized black-billed magpie (Pica pica) and the larger carrion crow (Corvus corone). We experimentally created 3-nestling broods of different brood compositions (only cuckoo chicks, only host chicks, or cuckoo and host chicks together) and measured baseline corticosterone levels of nestlings along their developmental period (early, middle and late). We found that brood parasitism increased corticosterone levels in magpie nestlings in the mid and late nestling period compared to those raised in unparasitized nests. Interestingly, carrion crow nestlings from parasitized nests only increased their corticosterone levels in the mid nestling period, when the competition for food with the cuckoo nestling was highest. Our results suggest that brood parasitism could be a potential physiological stressor for host nestlings, especially during the developmental stages where food requirements are highest. Conversely, cuckoo nestlings could be physiologically adapted to high competition levels since they did not show significant differences in corticosterone levels in relation to brood composition.     

EVOLUTION OF OBLIGATE SIBLICIDE IN BOOBIES. 2: FOOD LIMITATION AND PARENT–OFFSPRING CONFLICT

Proximate limitation on parental food delivery has long been invoked to explain the evolution of single-chick broods of pelagic seabirds such as masked boobies (Sula dactylatra). A second possible proximate limit on brood size is siblicide driven by genetic parent–offspring conflict (POC) over brood size, if siblicidal offspring can reduce brood size to one even if the parents' optimal brood size is greater than one. I tested these two hypotheses by experimentally suppressing obligate siblicide in masked booby broods and comparing breeding parameters of these broods with unmanipulated single-chick control broods. Per capita mortality rate of experimental nestlings was higher than that of controls, but this deficit was more than made up by larger brood size. Parents of experimental broods brought more food to offspring, had higher fledging success, and apparently incurred no additional major short-term cost of reproduction, relative to parents of control broods, thus refuting the food limitation hypothesis. Estimates of inclusive fitness of chicks in experimental broods were higher than were those of control nestlings, a result inconsistent with the POC hypothesis that the siblicidal offspring's optimal brood size is one while the parents' optimum is greater than one. This discrepency between natural brood size and apparent brood size optima might be resolved in several ways: experimental artifacts may give misleading estimates of optimal brood size; experimental and control offspring may have different reproductive values at the time of fledging; nestling masked boobies may face a special frequency-dependent case of POC in which the high risk of sharing a nest with a siblicidal sibling makes invasion of other behavioral genotypes difficult even when offspring and parent inclusive fitnesses are higher from a nonsiblicidal brood of two than from a brood of one.     

NESTLING SURVIVAL AND NESTLING WEIGHTS IN THE HOUSE SPARROW AND TREE SPARROW PASSER SPP. AT OXFORD

Nestling survival and nestling weights in P. domesticus and P. montanus were studied in 1961 and 1963–64 at Oxford. This paper concludes a study of factors influencing the reproductive rate. Taking all losses into account, P. domesticus reared an average of 1^.6 nestlings per brood (45%) and P. montanus 2.7 nestlings per brood (59%). About a third of all broods of both species failed completely to survive the nestling period. In P. domesticus these failures were most numerous in the middle part of the breeding season and are attributed to nutritional deficiencies derived from unsuitable food provided as a consequence of a seasonal food shortage, but in P. montanus complete brood failures occurred mostly in the second half of the nestling period and are attributed to predation. 43 broods of P. domesticus and one brood of P. montanus were weighed daily. Those of P. domesticus were classified as (1) successful broods—in these some nestlings died in the larger brood-sizes, apparently through starvation; (2) long-lived unsuccessful broods—in these the nestlings died at intervals and failure was attributed to nutritional deficiencies; and (3) short-lived unsuccessful broods. A slight decrease in the weights of nestlings in successful broods at the end of the nestling period is attributed to the utilization of insulating fat facilitated by the completion of the feather covering. Nestlings of both species left the nest at 88–89% of the adult weight. Taking all “successful” broods together, the percentage survival rates on nestling day 13¹/₂ (day of hatching = day ¹/₂) in P. domesticus were 81–82% in b/2–3, 70% in b/4 and 56% in b/5 (a situation paralleled in this respect by P. hispaniolensis), but in P. montanus they were c. 82% in all brood-sizes. Hence, in P. domesticus b/4 probably gave rise to the largest number of nestlings reared per brood, while in P. montanus most nestlings were produced by the largest brood-size. Weighings of many broods on day 13¹/₂ showed two trends in the weight of the nestlings: (1) in both species the weight of the nestling decreased as the number of survivors from each initial brood-size decreased; (2) between successive initial brood-sizes the weight of the nestling of P. domesticus decreased with increasing brood-size but in P. montanus there was no change. The losses in the larger broods of P. domesticus occurred mostly in the first half of the nestling period—apparently in association with the asynchronous hatching of the eggs and as a consequence of the limitation on the feeding frequency of the adults. Nestling survival was lowest in the larger broods in the middle of the breeding season and contrasted with the mid-season increase in mean clutch-size. It is suggested that in the study area there was a (possibly unnatural) shortage of food suitable for nestlings in the middle of the season. It is suggested that in P. domesticus the unexpectedly low feeding frequencies of the adults with large broods, apparently causing their low survival rates, may be an adaptation evolved to obtain the maximum amount of food in the presence of other adults which would be attracted to a food source by higher rates of activity. The breeding success calculated from data derived from the whole of this study was 35% for P. domesticus and 49% for P. montanus (2.9 and 3.9 nestlings per breeding pair per year respectively). It is suggested that the population of P. domesticus was much closer to a critical limiting factor, e.g. food supply, than that of P. montanus. This may account for the striking differences between the two species in their nestling survival rates and their nestling weights in relation to brood-size; in particular, the success of the larger broods of P. montanus may have been a temporary phenomenon.     

Similarity in the begging calls of nestling Red‐winged Blackbirds

ABSTRACT Although individually distinct begging calls may permit parents to recognize their offspring, birds nesting in dense breeding colonies where fledglings intermingle might benefit from additional adaptations. For example, if the calls of all nestlings in a brood were similar, parents would need to recognize only one brood call instead of the identity calls of each nestling. We recorded nestling Red‐winged Blackbirds (Agelaius phoeniceus) to determine whether their calls function to identify individuals (identity call hypothesis) or broods (brood call hypothesis). We used spectrogram cross‐correlation and dynamic time warping as well as call duration, peak frequency, and frequency range to estimate the similarity of begging calls of nestling Red‐winged Blackbirds. We recorded individual nestlings on day 5 and on day 9 of the nestling period to determine whether calls of individuals were more similar than calls of different nestlings, and whether calls of broodmates were more similar than calls of nestlings from different broods. We found that calls of 8‐d‐old individuals were more similar than calls of different nestlings, but the calls of broodmates were not more similar than those of nestlings from different broods. These results were consistent with the identity call hypothesis. We then compared begging calls of pairs of nestlings recorded separately and together on day 9. We found that the calls of 8‐d‐old nestlings recorded together were more similar than when they were recorded separately. In addition, using playback of begging calls from normal broods and artificial “broods” constructed from the calls of single nestlings, we found that females returned with food sooner in response to the calls of single nestlings (with enhanced call similarity) than to those of normal broods. Our results suggest that similar begging calls may be beneficial for both nestlings and parents, with broodmates fed at higher rates when their calls are more similar and, after fledging, parents needing to recognize only one brood call instead of the identity calls of each fledgling.     

The direct and indirect effects of predation by Hen Harriers Circus cyaneus on trends in breeding birds on a Scottish grouse moor

Two phases of an experimental reduction in management at Langholm Moor in southwest Scotland provided an insight into factors that determined trends in the abundance of moorland birds. In 1992 the historical control of breeding Hen Harriers Circus cyaneus stopped and in 2000 grouse moor management was discontinued. Golden Plover Pluvialis apricaria , Lapwing Vanellus vanellus , Curlew Numenius arquata and Red Grouse Lagopus lagopus scoticus declined, whilst Carrion Crow Corvus corone and Snipe Gallinago gallinago increased. Hen Harriers increased from two to 20 breeding females, then declined back to two. Lapwing abundance was positively associated with that of Hen Harriers, and moorland passerines (Skylark Alauda arvensis , Meadow Pipit Anthus pratensis and Stonechat Saxicola torquata ) were negatively associated. Golden Plover, Lapwing, Curlew, Red Grouse, Skylark and Hen Harrier were more abundant when the moor was managed for grouse, whilst Carrion Crow, a common predator of clutches of ground-nesting birds, increased during the second half when management ceased. Increased Crow numbers, together with an increase in Red Foxes Vulpes vulpes , probably contributed to the observed bird declines.     

Intraseasonal effects of clutch manipulation on parental provisioning and residual reproductive value of eastern phoebes (Sayornis phoebe)

Summary First clutches of double-brooded eastern phoebes Sayornis phoebe were manipulated (up two eggs, down 2 eggs or no change) to test for intraseasonal reproductive tradeoffs and to test whether size of first brood influenced food delivery rates to nestlings and nestling quality in second broods.Considering all nests from both broods, rate of feeding nestlings increased linearly with brood size but nestling mass per nest decreased with increasing brood size. High nestling weights in small broods may have resulted from parents delivering better quality food, but we did not test this.Among treatment groups in first broods, nestlings from decreased broods weighed more than those in control or increased broods. Treatment did not influence the likelihood that second nests would be attempted after successful first nests nor did it alter the interval between nests. Nestlings of parents that renested weighed more than those of parents that did not, regardless of treatment, suggesting that post-fledging care may preclude renesting. Mass of individual females did not change between broods, regardless of brood size. Clutch sizes of second attempts were not affected by manipulations of first broods but increasing first broods reduced the number of nestlings parents were able to raise to day 11 in their second broods. However, manipulation of first broods did not affect mean nestling mass per nest of nestlings that survived to day 11.In phoebes, parents of small first broods are able to raise nestlings in better condition. We predict that in harsh years, parents of small first broods would be more likely to renest. Parents of enlarged first broods sacrificed quality of offspring in second broods, which seems a reasonable strategy if nestlings from second broods have lower reproductive value.     

Brood size and begging intensity in nestling birds

Theoretical models suggest that sibling competition should selectfor conspicuous begging signals. If so, begging intensity mightbe expected to increase with the number of competitiors. Thepurpose of our study was to examine the relationship betweenbegging intensity and brood size using nestling tree swallows(Tachycineta bicolor) as our model. Over 2 years, we videotapedbegging behavior in unmanipulated broods of different sizes.We found that begging intensity increased with brood size. Theaverage weight of nestlings in each brood did not vary withbrood size, but feeding rate per nestling decreased with broodsize, suggesting that nestlings in larger broods begged moreintensively, possibly because they were hungrier. We also conductedan experiment to examine the effect of nest mates on beggingin different-sized broods. We found that nestlings with similarweights, previous competitive environments, and food deprivationbegged more intensively in large broods than in small broods.Overall, our study indicates that begging intensity increaseswith brood size in tree swallows. This relationship may resultfrom interactions among brood mates rather than from lower feeding rates to individual nestlings in larger broods.     

Status of the Hen Harrier Circus cyaneus in the UK and Isle of Man in 2016

Capsule: The fifth UK and the Isle of Man survey of Hen Harrier in 2016 showed varying trends by country and region.

Aim: To estimate the size of the breeding Hen Harrier population (with associated 95% confidence intervals) in the UK and Isle of Man, constituent countries and Scottish regions, in 2016 and calculate population change over the five surveys to date.

Methods: Complete surveys were made of all 10-km squares likely to be occupied by breeding Hen Harriers in England, Wales, Northern Ireland and the Isle of Man, using standard methods developed for previous national surveys. In Scotland, self-selected 10-km squares were surveyed by volunteers and a stratified random selection of the remaining 10-km squares within the known breeding range was surveyed.

Results: The UK and Isle of Man Hen Harrier population was estimated at 575 territorial pairs (95% confidence limits, 477–694), a non-significant decline of 13% since 2010 but a significant decline of 24% since 2004. Scotland held the majority (80%) of the population with 460 (359–573) territorial pairs. Elsewhere, 46 territorial pairs were recorded in Northern Ireland, 35 in Wales, 30 in the Isle of Man and four in England. Significant decreases were recorded in the number of pairs in Scotland using grouse moor (?57%) and young forest (?54%).

Conclusion: The combined breeding population of Hen Harriers in the UK and Isle of Man has shown a non-significant decline between 2010 and 2016. There were notable decreases in England, Northern Ireland and Wales since 2010.     

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.     

Providing chicks with extra food lowers male but not female provisioning in the House Sparrow Passer domesticus

We assessed whether adult House Sparrows Passer domesticus adjusted their provisioning in response to an experimental increase in the nutritional condition of their nestlings. When we supplemented chicks directly with additional food, male parents, but not female parents, reduced their provisioning. The results for males, but not females, run contrary to a previous experiment in this species. In addition, female provisioning was positively associated with both brood size and the age of the brood. In contrast, whereas male provisioning was positively associated with brood size, males did not increase provisioning as their chicks grew older. Males, but not females, exhibited repeatability in their provisioning. Food supplementation had a larger positive effect upon nestling survival in smaller broods than in larger broods. Overall, there appear to be fundamental differences between males and females in how decisions regarding the level of parental investment in the current brood are made.     

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.     

Supplementary feeding increases nestling feather corticosterone early in the breeding season in house sparrows

Several studies on birds have proposed that a lack of invertebrate prey in urbanized areas could be the main cause for generally lower levels of breeding success compared to rural habitats. Previous work on house sparrows Passer domesticus found that supplemental feeding in urbanized areas increased breeding success but did not contribute to population growth. Here, we hypothesize that supplementary feeding allows house sparrows to achieve higher breeding success but at the cost of lower nestling quality. As abundant food supplies may permit both high‐ and low‐quality nestlings to survive, we also predict that within‐brood variation in proxies of nestling quality would be larger for supplemental food broods than for unfed broods. As proxies of nestling quality, we considered feather corticosterone (CORT_f), body condition (scaled mass index, SMI), and tarsus‐based fluctuating asymmetry (FA). Our hypothesis was only partially supported as we did not find an overall effect of food supplementation on FA or SMI. Rather, food supplementation affected nestling phenotype only early in the breeding season in terms of elevated CORT_f levels and a tendency for more variable within‐brood CORT_f and FA. Early food supplemented nests therefore seemed to include at least some nestlings that faced increased stressors during development, possibly due to harsher environmental (e.g., related to food and temperature) conditions early in the breeding season that would increase sibling competition, especially in larger broods. The fact that CORT_f was positively, rather than inversely, related to nestling SMI further suggests that factors influencing CORT_f and SMI are likely operating over different periods or, alternatively, that nestlings in good nutritional condition also invest in high‐quality feathers.     

Habitat and diet of young grouse broods: resource partitioning between Capercaillie (Tetrao urogallus) and Black Grouse (Tetrao tetrix) in boreal forests

We compared habitat use and diets of young Capercaillie and Black Grouse broods in a boreal forest in southeast Norway. We used pointing dogs to search for broods (N = 83) in mature “natural” forest types and examined the crop content of 66 chicks 1–9 weeks old. We also measured the abundance of insects in the habitats where broods were found. Although overlapping substantially in both habitat and diets, there were notable differences: Capercaillie broods were more frequently recorded in bilberry-dominated forest types, whereas Black Grouse preferentially used pine bog forest, a more open habitat with little bilberry. Capercaillie chicks ate proportionally more insects, particularly lepidopteran larvae, and insects dominated their diet for a longer period of time (until age 28–29 days) than in Black Grouse (14–15 days). After reaching their peaks, the quantity of insects in the crops declined rapidly especially in Capercaillie, and in one of 2 years this occurred at a time when insects, including larvae, were still abundant in the habitats. Among plant foods, both species ate large amounts of Bilberry (Vaccinium myrtillus) and Bog Whortleberry (V. uliginosum). The main difference between species was a large proportion of both over-wintered and new, not yet ripe, berries of Cranberry (Oxycoccus quadripetalus) in Black Grouse, and a higher proportion of the forb Melampyrum sylvaticum in Capercaillie. The difference in diets reflected their differential use of habitats; the Vaccinium-preferred habitats of Capercaillie were richer in insects, particularly larvae, than the pine bog habitat preferred by Black Grouse. Because insects, especially larvae, comprised a larger proportion of the diet of Capercaillie chicks and chicks of this species need more food to sustain their rapid growth, Capercaillie is likely to be more sensitive to variation in insect food than Black Grouse. Also, by reducing the abundance of bilberry, the main host plant of larvae chick food, clearcutting forestry has negative effects on the brood habitat quality of both species.     

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.