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Earlier offspring mortality before independence saves resources for kin, which should be more beneficial when food is short. Using 24 years of data on age-specific common tern (Sterna hirundo) chick mortality, best described by the Gompertz function, and estimates of energy consumption per age of mortality, we investigated how energy wasted on nonfledged chicks depends on brood size, hatching order, and annual abundance of herring (Clupea harengus), the main food source. We found mortality directly after hatching (Gompertz baseline mortality) to be high and to increase with decreasing herring abundance. Mortality declined with age at a rate relatively insensitive to herring abundance. The sensitivity of baseline mortality to herring abundance reduced energy wasted on nonfledged chicks when herring was in short supply. Among chicks that did not fledge, last-hatched chicks were less costly than earlier-hatched chicks because of their earlier mortality. However, per hatchling produced, the least energy was wasted on chicks without siblings because their baseline mortality was most sensitive to herring abundance. We suggest that earlier mortality of offspring when food is short facilitates economic adjustment of posthatching parental investment to food abundance but that such economic brood reduction may be constrained by sibling competition.

Brood reduction is a within-brood partial mortality due to sibling rivalry and has been observed in a large number of bird species from diverse taxa. Here, I summarize several hypotheses accounting for the adaptive value of brood reduction and discuss different factors that modulate it. A description of different brood reduction systems is made with focus on Neotropical birds. Finally, an integrative approach analyzing brood reduction in Imperial Shags (Phalacrocorax atriceps) is presented to illustrate causes, consequences, and benefits of this breeding strategy. Fil: Svagelj, Walter Sergio. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Mar del Plata. Instituto de Investigaciones Marinas y Costeras. Universidad Nacional de Mar del Plata. Facultad de Ciencias Exactas y Naturales. Instituto de Investigaciones Marinas y Costeras; Argentina

In animals with uniparental care, the quality of care provided by one sex can deeply impact the reproductive success of both sexes. Studying variation in parental care quality within a species and which factors may affect it can, therefore, shed important light on patterns of mate choice and other reproductive decisions observed in nature. Using Syngnathus typhle, a pipefish species with extensive uniparental male care, with embryos developing inside a brood pouch during a lengthy pregnancy, we assessed how egg size (which correlates positively with female size), male size, and water temperature affect brooding traits that relate to male care quality, all measured on day 18, approximately 1/3, of the brooding period. We found that larger males brooded eggs at lower densities, and their embryos were heavier than those of small males independent of initial egg size. However, large males had lower embryo survival relative to small males. We found no effect of egg size or of paternal size on within-pouch oxygen levels, but oxygen levels were significantly higher in the bottom than the middle section of the pouch. Males that brooded at higher temperatures had lower pouch oxygen levels presumably because of higher embryo developmental rates, as more developed embryos consume more oxygen. Together, our results suggest that small and large males follow distinct paternal strategies: large males positively affect embryo size whereas small males favor embryo survival. As females prefer large mates, offspring size at independence may be more important to female fitness than offspring survival during development., The quality of parental care provided by one parent can affect the reproductive success of both parents and therefore, choosing a good parent can be crucial during mate choice. In the broad-nosed pipefish, large males brood embryos at lower densities, that become larger but have relatively lower survival during brooding than those of small males. As females prefer large mates, our work suggests that offspring size post-birth is more important than number to female fitness.

To compete for parental food deliveries nestling birds have evolved diverse behaviors such as begging displays and sibling aggression. Testosterone has been suggested to be an important mechanism orchestrating such competitive behaviors, but evidence is scarce and often indirect. Siblicidal species provide an interesting case in which a clear dominance hierarchy is established and the dominant chicks lethally attack siblings. We experimentally elevated testosterone in chicks of a facultatively siblicidal species, the black-legged kittiwake, Rissa tridactyla, and showed that testosterone-treated chicks were more aggressive toward their sibling than were control chicks. In such facultatively siblicidal species, chicks normally exhibit intense aggression only when threatened by starvation. Indeed, we found that chicks in relatively poorer condition were more aggressive than were chicks in better condition, even among testosterone-treated chicks, suggesting the action of an additional signal modulating aggression. Relatively larger siblings were also more aggressive than were relatively smaller siblings, confirming the importance of size advantage in determining dominance hierarchies within the brood. In addition, testosterone increased aggression toward a simulated predator, indicating that in kittiwakes testosterone can increase aggression in contexts other than siblicide. Testosterone promoted aggression-mediated dominance, which increased begging although testosterone treatment did not have a significant separate effect on begging. Therefore, testosterone production in the kittiwake and most likely other siblicidal species seems an important fitness mediator already early in life, outside the sexual context and not only manifesting itself in aggressive behavior but also in dominance-mediated effects on food solicitation displays toward parents.

Substantial amounts of maternal androgens are found in birds' eggs and have been shown to benefit offspring development. Within-clutch patterns of increasing androgen concentrations over the laying sequence are often hypothesized to compensate for the negative effects of hatching asynchrony. However, detrimental effects to offspring fitness of exposure to high yolk androgen levels have also been demonstrated. This suggests that mothers should forego these costs to their offspring when the need for compensation for hatching asynchrony is low or when alternative compensatory strategies, e.g. in terms of increasing egg mass, are available. Here we show that in the south-temperate lesser double-collared sunbird Nectarinia chalybea, a species with hatching asynchrony but also with high survival of last-hatched chicks, mothers do not deposit resources differentially in terms of either yolk androgen concentration or egg mass across the laying sequence. We discuss to what extend this challenges the original explanation of within-clutch variation in these egg parameters and offer some explanation for their between-clutch variation which was related to female body mass.

Many Apodidae, including Streptoprocne biscutata (Sclater, 1866), drop eggs from their nests during incubation. This is interpreted as nest site competition or accident. We provide evidence that egg ejection is deliberate and that this behaviour controls the brood size. Brood sizes were manipulated and nestling growth was measured to test the hypothesis that pairs can regulate brood size during incubation based on current ability to rear nestlings. Natural (control) broods with one, two and three nestlings, and manipulated (experimental) broods reduced to one and increased to two and three young were monitored. Growth rates were measured based on weight, and wing, tail and tarsus lengths of natural and manipulated broods. We compared the slopes of each measure's regression lines of the nestlings of each brood size by t-test. Nestling growth of control nests was similar and relatively little associated with brood size. In broods reduced to one nestling, weight, wing and tail had greater growth rates, and in broods increased to three nestlings growth rates were lower. Weight was most, and tarsus length least influenced by brood size. In general, nestling growth of manipulated nests was inversely proportional to brood size. The results suggest that pairs with larger clutches are in better physical conditions than others. Thus, in experimental broods, pairs are over or under-loaded because feeding activities increase or decrease and these changes affect the growth rate of the nestlings. The present study suggests that egg ejection can control brood size. This behaviour is probably stimulated by physical changes in the adult birds during incubation.

Maternal effects occur when the mother's phenotype influences her offspring's phenotype. In birds, differential allocation in egg yolk components can allow mothers to compensate for the competitive disadvantage of junior chicks. We hypothesize that the parent–older chick conflict peaks at intermediate conditions: parents benefit from the younger chick(s) survival, but its death benefits the older chick in terms of growth and survival. We thus expect maternal compensation to follow a bell‐shaped pattern in relation to environmental conditions. We studied a black‐legged kittiwake (Rissa tridactyla) population where previous results revealed increased allocation of yolk testosterone in younger as compared to older chicks in intermediate conditions, in line with our theoretical framework. We therefore predicted a maternally induced increase in aggressiveness, growth, and survival for younger chicks born in intermediate environmental conditions. Controlling for parental effects and chick sex, we manipulated food availability before egg laying to create a situation with intermediate (Unfed group) and good (Fed group) environmental conditions. Within each feeding treatment, we further created experimental broods where the natural hatching order was reversed to maximize our chances to observe an effect of feeding treatment on the younger chicks' aggressiveness. As predicted, we found that chick aggressiveness was higher in younger chicks born from the Unfed group (i.e., in intermediate environmental conditions), but only when they were put in a senior position, in reversed broods. Predictions on growth and survival were not confirmed. Mothers thus seem to favor the competitiveness of their younger chick in intermediate conditions via egg yolk components, but our study also suggests that hatching asynchrony need to be small for maternal compensation to be efficient. We emphasize the need for further studies investigating other chick behaviors (e.g., begging) and focusing on the relative role of different yolk components in shaping parent–offspring conflict over sibling competition.

Female birds of several species have control over the production of daughters and sons. However, most studies failed to find a relationship between egg size and sex. This is intriguing as adjustment of egg size would constitute a powerful tool for the female to meet different resource demands of the sexes, particularly in size dimorphic species. Our results show that, within clutches of black-headed gulls (Larus ridibundus) the proportion of males was positively associated with egg mass. This applied for all three laying positions, independently of the absolute egg mass. There was a significant relationship between the distribution of the sexes over the laying sequence and the egg mass change. When egg mass decreased over the sequence, first-laid eggs were male biased and last-laid eggs female biased, and vice versa. The potential adaptive value of this allocation strategy is evaluated with regard to male sensitivity to egg quality and competitive differences between the sexes.

The amount of time a bird allocates to incubation is likely to be limited by energetic constraints. If food is abundant, energetic constraints may be reduced and the time spent incubating (incubation attendance) may increase. Moreover, the onset of incubation in relation to clutch completion may be advanced, resulting in a higher degree of hatching asynchrony. We measured the effect of experimentally increased food availability on incubation attendance and an estimate of hatching asynchrony in the Australian reed warbler Acrocephalus australis. Supplementary food was provided every other day, from a few days before the start of egg laying until just prior to hatching. Incubation attendance was measured with temperature loggers at nests receiving supplementary food and control nests. Hatching asynchrony was inferred from mass and size differences between siblings shortly after hatching. We found that 1) food supplementation resulted in an increase in incubation attendance, when comparing both nests receiving supplementary food to control nests as well as feeding to non-feeding days in nests receiving supplementary food, and 2) food supplementation resulted in a greater hatching asynchrony, without affecting clutch size, average egg volume or the likelihood of eggs hatching. This suggests that food availability acts in a proximate way to modify the extent of incubation attendance and hatching asynchrony. We discuss the adaptive significance of increased incubation attendance and a shift in the degree of hatching asynchrony in relation to food availability.

All parental hosts of heterospecific brood parasites must pay the cost of rearing non-kin. Previous research on nest parasitism by brown-headed cowbirds (Molothrus ater) concluded that competitive superiority of the typically more intensively begging and larger cowbird chick leads to preferential feeding by foster parents and causes a reduction in the hosts' own brood. The larger size of cowbird nestlings can be the result of at least two causes: (1) cowbirds preferentially parasitize species with smaller nestlings and lower growth rates; and/or (2) cowbirds hatch earlier than hosts. I estimated the cost of cowbird parasitism for each of 29 species by calculating the difference between hosts' published brood sizes in nonparasitized and parasitized nests and using clutch size to standardize values. In this analysis, greater incubation length and lower adult mass, surrogate measures of the hatching asynchrony and size difference between parasite and hosts, were both related to greater costs of cowbird parasitism without bias owing to phylogeny. To establish causality, I manipulated clutch contents of eastern phoebes (Sayornis phoebe) and examined whether earlier hatching by a single cowbird or phoebe egg reduces the size of the rest of the original host brood. As predicted, greater hatching asynchrony increased the proportion of the original phoebe brood that was lost. This measure of the cost of parasitism was partially owing to increased hatching failure of the original eggs in asynchronous broods but was not at all related to the size differences of older and younger conspecific nestmates. However, proportional brood loss owing to an earlier hatching conspecific was consistently smaller than brood loss owing to asynchronous cowbirds in both naturally and experimentally parasitized phoebe nests. These results imply that although hatching asynchrony is an important cause of the reduction of host broods in parasitized clutches, competitive features of cowbird nestlings remain necessary to explain the full extent of hosts' reproductive costs caused by interspecific brood parasitism. Copyright 2003.

We collected 328 freshly laid Great Tit (Parus major) eggs from 38 clutches in 1999 to determine the relationship of whole egg weight, wet yolk weight, wet albumen weight, dry shell weight, and the occurrence of laying gaps with mean ambient temperature in the three days preceding laying, while controlling for laying date and position in the laying sequence. We also reanalyzed existing data on whole egg weight in 1978 and 1979 in relation to temperature, controlling for the same other variables. Egg weight was correlated with temperature in 1978 and 1979, but not in 1999. None of the egg components was related to temperature preceding laying, apart from a nonsignificant tendency for albumen weights to be lower at higher temperature. Egg weight increased with laying date in 1978 and 1979 after controlling for other variables. In 1999, albumen and shell weight increased seasonally, but shell weight only did so to the extent expected on the basis of the seasonal increase in whole egg weight. Yolk weight did not increase significantly seasonally, but neither did yolk weight relative to whole egg weight decrease significantly. Whole egg weight and the weights of all components varied between clutches. Most weights also varied through the laying sequence. The probability of a laying gap occurring varied between clutches, and increased with position in the laying sequence and decreased with temperature and laying date. Eggs preceding laying gaps were lighter and contained less albumen but had higher shell weights than other eggs. Eggs following laying gaps also had heavier shells relative to the size of the egg than other eggs.

1. Females of egg-laying vertebrates may adjust the development of their offspring to prevailing environmental conditions by regulating the deposition of hormones into their eggs. Within- and amng-clutch variation in levels of steroid hormones were studied in the egg yolks of the Black-Headed Gull (Larus ridibundus, Linnaeus) in relation to environmental conditions at the nest site, This species breeds in colonies of different densities and in different habitats, and the chicks hatch asynchronously.2. Egg yolks contained very high levels of androstenedione, substantial levels of testosterone and moderate levels of 5alpha-dihydrotestosterone. Oestrogen (17-oestradiol) was not detected.3. Androgen levels increased strongly with laying order, irrespective of egg or yolk mass. This may compensate for the disadvantages of the later hatching chicks. These results have implications for adaptive hypotheses that were proposed for asynchronous incubation.4. Eggs of lighter clutches contained more androgens, perhaps to compensate for a lower nutritional quality of these eggs.5. Birds breeding in the periphery of a colony, being relatively more aggressive and having relatively large territories, laid eggs that contained more androgens than those of birds breeding in the centre. These high yolk androgen levels may facilitate growth and motor development of the chicks, which may be especially important for chicks developing at the periphery of a colony. Reduced levels may be adaptive for birds breeding in the centre, where risk of infectious diseases is high, since steroids may be immunosuppressive.6. Corrected for nest distance, clutches of birds in high vegetation, where predation risk is less severe and therefore competition for nest sites perhaps high, contained relatively high levels of androgens. It is suggested that the level of yolk androgens reflects the hormonal condition of the female, that in turn is influenced by her characteristics such as her age and aggressiveness, and the level of social stimulation.

Offspring are often produced in excess as insurance against stochastic events or unpredictable resources. This strategy may result in high early-life mortality, yet age-specific mortality before offspring independence and its associated costs have rarely been quantified. In this study, we modelled age-specific survival from hatching to fledging using 24 years of data on hatching order (HO), growth and age of mortality of more than 15 000 common tern ( Sterna hirundo ) chicks. We found that mortality peaked directly after hatching, after which it declined rapidly. Mortality hazard was best described with the Gompertz function, and was higher with later HO, mainly due to differences in baseline mortality hazard, rather than age-dependent mortality. Based on allometric mass–metabolism relationships and detailed growth curves of starving chicks, we estimated that the average metabolizable energy intake of non-fledged chicks was only 8.7% of the metabolizable energy intake of successful chicks during the nestling phase. Although 54% of hatchlings did not fledge, our estimates suggest them to have consumed only 9.3% of the total energy consumption of all hatched chicks in the population before fledging. We suggest that rapid mortality of excess offspring is part of an adaptive brood reduction strategy to the benefit of the parents.

We tested the hypothesis that mother birds counterbalance the negative effects of hatching asynchrony for later–hatched chicks by increasing the yolk androgen concentrations in consecutive eggs of their clutch. In doing so, they may adaptively tune each offspring9s competitive ability and, thus, growth and survival. However, evidence in support of this hypothesis is contradictory. The yolk concentrations of maternal androgens in the eggs of black–headed gulls increase significantly with the laying order of the eggs in a clutch. We experimentally tested the functional consequences of this increase on chick development under natural conditions by injecting eggs with either an oil or androgen solution. We created experimental clutches in which androgen levels either stayed constant or increased with laying order while controlling for differences in egg quality by using only first–laid eggs. We then compared development, growth and survival between these broods. Androgen treatment enhanced embryonic development because androgen–treated eggs hatched half a day earlier than controls, while their size at hatching was similar to oil–treated controls. Androgen treatment did not increase chick survival, but it enhanced growth. Androgen–treated, third–hatched chicks had a higher body mass and longer legs than thirdndash;hatched chicks that hatched from oil–treated eggs. At the same time, growth of first chicks (which were all oil treated) was reduced by the presence of two androgen–treated siblings, suggesting that yolk androgens enhance the competitive ability of later–hatched chicks. Our results support the hypothesis that transfer of different amounts of androgens to the eggs of a clutch is a mechanism by which mothers maximize their reproductive output.

We investigated the fledging probability of oystercatcher, Haematopus ostralegus, chicks as a function of hatching order, brood size, territory quality and food availability. Sibling dominance was related to the hatching order in both low- ('leapfrogs') and high-quality ('residents') territories. Differences in hatchling mass might have aided the establishment of a dominance hierarchy, since breeders produced small late eggs and hatchlings. These mass differences were most pronounced in leapfrogs, and in large broods in years with lower food availability ('poor' years). Late hatchlings fledged less often and with lower body masses compared to early hatchlings in all situations. Leapfrogs produced smaller broods and hatched their broods more asynchronously in poor years than leapfrogs breeding in years with more available food ('good' years) and residents breeding in both poor and good years. Large brood sizes resulted in lower survival of hatchlings in poor years. These results favour the 'brood reduction' hypothesis. However, contrary to the expectations of this hypothesis, hatching order also affected fledging success in residents. Moreover, large brood size resulted in higher survival of hatchlings in good years, particularly in residents. Thus, although large broods experienced losses due to sibling competition in some years, they nevertheless consistently produced more fledglings per brood in all years, both as leapfrogs and residents. We believe this effect is due to parental quality correlating with initial brood size. Most leapfrogs, at best, fledged one chick successfully each year, losing chicks due to starvation. Nevertheless, leapfrog broods were reduced in size after hatching significantly less quickly than resident broods. These results suggest that breeders lay and hatch insurance eggs to compensate for unpredictable losses due to the high predation rates on both nests (ca 50%) and chicks (ca 90%), in accordance with the 'nest failure' hypothesis.

Food availability is an important limiting factor for avian reproduction. In altricial birds, food limitation is assumed to be more severe during the nestling stage than during laying or incubation, but this has yet to be adequately tested. Using food-supplementation experiments over a 5-year period, we determined the degree and timing of food limitation for burrowing owls (Athene cunicularia) breeding in Canada. Burrowing owls are an endangered species and food limitation during the nestling stage could influence reproductive performance of this species at the northern extent of their range. Supplemented pairs fledged on average 47% more owlets than unfed pairs, except during a year when natural food was not limiting (i.e., a prey irruption year). The difference in fledgling production resulted from high nestling mortality in unfed broods, with 96% of all nestling deaths being attributed to food shortage. Supplemental feeding during the nestling period also increased fledgling structural size. Pairs fed from the start of laying produced the same number of hatchlings as pairs that received no supplemental food before hatch. Furthermore, pairs supplemented from egg laying to fledging and pairs supplemented during the nestling period alone had the same patterns of nestling survival, equal numbers of fledglings, and similar fledgling mass and structural size. Our results provide empirical support for the hypothesis that the nestling period is the most food-limited phase of the breeding cycle. The experimental design we introduce here could be used with other altricial species to examine how the timing of food limitation differs among birds with a variety of life-history strategies. For burrowing owls, and other species with similar life histories, long-term, large-scale, and appropriately timed habitat management increasing prey abundance or availability is critical for conservation. Our results provide empirical support for the hypothesis that the nestling period is the most food-limited phase of the breeding cycle. For burrowing owls, and other species with similar life histories, long-term, large-scale, and appropriately timed habitat management increasing prey abundance or availability is critical for conservation.

In many species, embryos are exposed to maternal hormones in utero, in the egg, or in the seed. In birds, mothers deposit substantial testosterone into their eggs, which enhances competitive ability of offspring. These maternal testosterone concentrations vary systematically within clutches in different patterns and may enable mothers to adaptively fine-tune competitive hierarchies within broods. We performed a comparative analysis to investigate this hypothesis using a broad set of avian species. We expected species with small size differences among siblings (arising from small hatching asynchrony or slow growth rates) to aim for survival of the whole brood in good years and therefore compensate last-hatching eggs with relatively more testosterone. We expected species with large size differences among siblings (large hatching asynchrony or fast growth rates) to produce surplus young as insurance against failed offspring and to facilitate elimination of redundant surplus young by bestowing last-hatching eggs with relatively less testosterone. As predicted, we found that maternal testosterone compensation to last-hatching eggs is stronger when size differences among siblings become smaller. Maternal testosterone compensation to last-hatching eggs also correlated negatively with hatching asynchrony and growth rates. These findings provide evidence for correlated evolution of several maternal effects that together support different maternal reproductive strategies.

From an evolutionary and adaptive perspective, the occurrence and extent of parental care in animals (investment in offspring beyond the genetic contribution) is expected to depend on an optimal (fitness-maximizing) balance of its costs and benefits. Males of the sex-role reversed broad-nosed pipefish, Syngnathus typhle, which care for eggs and embryos and may perform brood reduction, are particularly interesting in this respect. In this thesis, I explore paternal care in the broad-nosed pipefish with special focus on the causes and consequences of brood reduction. Firstly, I demonstrate that brooding males absorb nutrients from reduced embryos in their brood pouch, presumably benefiting their own nutrition. This indicates that brood reduction in this species is similar to filial cannibalism (eating of own young), found in many other fish species with paternal care. Furthermore, as the embryos did not absorb nutrients originating from siblings in the same pouch, there is no support for ‘nurse eggs’ in this pipefish species (paper I). When comparing brooding males with non-brooding males, brooding males were on average in better condition, as measured by hepatosomatic index (HSI). This is contrary to the expectation that the metabolic and other costs of brooding males should deplete their energy reserves. However, given the results above (paper I), the higher HSI in brooding males may to some extent stem from nutrients acquired through brood reduction (paper V). This uptake is, however, unlikely to fully compensate for the costs of paternal care since another experiment (paper IV) showed that males brooding full broods had lower survival, despite a larger relative brood reduction, compared to males with half-sized broods (paper V). Furthermore, the more eggs a male received, the more eggs were reduced, which supports the indication that the cost of brooding is related to brood size (paper II and IV). This clearly indicates that embryos are competing over paternal care and that paternal care is a costly and limiting resource. Costs of paternal care in terms of reduced energy reserves (HSI) and lower survival (paper IV) suggests that males in poor condition (low HSI) should have a lower optimal brood size and thus show a larger relative brood reduction, as found in paper III. Also, males with low HSI received fewer eggs at mating and were more likely to suffer mortality from an infection (paper III). When comparing monogynously and polygynously mated males, only monogynous males showed significant brood reduction, whereas polygynous males did not (paper II). This may be caused by either increased paternal provision or less competition due to less overlapping needs within half-sib than full-sib broods. Also, broods from larger females survived better than broods from smaller females (paper III and V). This may show that large eggs have certain qualities which allow them to survive better, or that males provide better care to eggs from larger females, reflecting the sexual conflict of embryo competition and cryptic male choice that may be operating in this species. In summary, I have found that several factors influence the process of brood reduction (filial cannibalism) in the broad-nosed pipefish. These factors include the number of female mating partners, brood size, egg size and male condition. Consequently, these factors are important in parental conflicts and parent-offspring conflicts over the level of parental care and brood reduction.

Crested penguins (genus Eudyptes) have a peculiar hatching pattern, with the first-laid egg (A-egg) hatching after the second-laid egg (B-egg) and chicks from A-eggs typically having a much lower survival probability. Maternal yolk androgens have been suggested to contribute to the competitive superiority of the B-chick in southern rockhopper penguins Eudyptes chrysocome, given their important role in mediating sibling competition in other species. We therefore increased the yolk androgen levels in freshly-laid eggs and examined the consequences for sibling competition - via effects on embryonic developmental times, chick growth and early survival. We placed one androgen-treated egg and one control egg into each foster nest, matching them for mass, laying date and laying order. The androgen treatment did not significantly affect embryonic developmental times or chick measurements at hatching. However, elevated yolk androgen levels benefitted chick growth in interaction with the number of siblings in a brood. Chicks from androgen-treated eggs had faster growth in the presence of a sibling than chicks from control eggs. Under these circumstances they also had a higher survival probability. Thus maternal androgens appear to reinforce the observed hatching pattern, facilitating brood reduction. This contrasts to most previous studies in other species where yolk androgens have been shown to compensate for the negative consequences of delayed hatching within the brood hierarchy.

Research into the effect of environmental variables on reproductive success of tropical raptors is often constrained by the lack of information on breeding biology. We provide the first detailed information of the breeding biology and nestling development of the Grasshopper Buzzard Butastur rufipennis, an Afrotropical migratory raptor threatened by extensive land transformation in its breeding range. Breeding coincided with the transition from the dry to the wet season. The mean incubation period was 30 d and mean fledging period 36 d. The breeding period was characterised by rapid establishment of territories and short post-fledging dependency periods related to the onset of migration shortly after breeding. Growth rate of body mass, tarsus, wing and primary feathers were similar between sexes, which showed significant but moderate body mass dimorphism at fledging (♂85–90% of ♀). Second hatchlings were smaller in body mass and structural dimensions compared to similarly-aged first hatchlings and singles of the same sex. Survival of second hatchlings was related to body mass ratio with first hatchlings, whereas brood reduction occurred through food competition and siblicide. We provide ageing formulae and photographic records to facilitate further studies of Grasshopper Buzzard nestling development and reproductive success in the region.Keywords: ageing, Butastur rufipennis, hatch order, reproduction, siblicideOSTRICH 2012, 83(3): 137–146

No environment is truly constant in time. As a result, animals have evolved multiple adjustments to cope with such fluctuations. However, the allocation of ffort to costly activities that imply long-term commitments, such as breeding, may be extremely challenging when future resources change constantly and unpredictably, a context that has received little investigation. To fill this gap, we studied the breeding response by a wetland-dependent raptor, the black kite Milvus migrans, to within and between-years fluctuations in resource availability (inunda- tion levels). The breeding performance of the population was decomposed into reproductive components expressed in a sequence of successive tasks along the breeding cycle (e.g. timing of laying, clutch size, hatching success, brood reduction). Variation in each component was related to resource levels observed at different key dates of the season in order to test whether and when population-level reproduction was adjusted to available resources. Along a 22-year time-series, inundation levels fluctuated unpredictably within and among years, and mostly affected the later components of kites’ reproduction, such as hatching success and the incidence of brood reduction, which were the main determinants of the population yearly breeding output. Results were consistent with multiple adjustments to cope with uncertainty. As the season progressed and resources became easier to assess, a bet-hedging waiting strategy based on a conservatively small, invariant and asynchronous clutch gave way to real-time resource-track- ing mechanisms mediated by progressive adjustments to current prey availability, so that population-level breeding rates were determined and tuned to resources rather late in the season. Such adjustments were the likely outcome of the interaction between parental tactics and environmental constraints. Behavioural flexibility, such as dietary opportunism, probably promoted further resistance to resource oscillations. Given that all ecosystems show some degree of unpredictability, resource-tracking adjustments, such as the ones depicted here, are likely to be commonplace in most communities.

We document the occurrence of fault bars in a population of nestling Ospreys Pandion haliaetus under natural conditions. Ospreys had an average of 9.9 fault bars on their rectrices, however variation was large. Fault bar formation declined linearly with age and increased symmetrically from outer to inner rectrices. Fault bar incidence is consistent in all plumage groups and those groups most essential for flight are least affected. We also examine fault bar occurrence in relation to two hypotheses: food shortage and handling effect. The former is investigated indirectly by comparing the number and intensity of rectrix fault bars in nestlings to their feeding rank and to their brood size. Neither variable has a significant effect on fault bar severity. The possibility of weather-mediated food shortage is examined, but no strong effect of weather is detected. The role of handling is assessed by comparing the number and intensity of fault bars in nestlings experiencing different numbers of nest visits. Nestlings visited repeatedly had more fault bars providing support for the handling hypothesis. Analysis of the timing of fault bar formation with respect to nest visits is suggestive as further evidence for the role of handling.