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.     

Ectoparasites and reproductive trade-offs in the barn swallow (Hirundo rustica)

Parents are predicted to trade offspring number and quality against the costs of reproduction. In altricial birds, parasites can mediate these costs because intensity of parasitism may increase with parental effort. In addition, parasites may mediate a trade-off between offspring number and quality because nestlings in large broods may have reduced anti-parasite immune defence. In this study, we experimentally analysed the effect of brood size on infestation by an ectoparasitic mite in nests of barn swallows (Hirundo rustica). Nests with an enlarged brood had larger prevalence and intensity of infestation than those with a reduced brood. Importantly, each nestling in enlarged broods was exposed to a larger number of mites, even when measured on a per nestling basis, than in reduced broods. Nestlings in enlarged broods had smaller body mass and T-cell-mediated immune response compared to reduced broods. T-cell-mediated immune response and feather growth were negatively correlated with per nestling intensity of infestation in enlarged but not in reduced broods. The results suggest that nestlings in enlarged broods have depressed immunity leading to larger per nestling mite infestation. Hence, exposure to parasites of offspring and parents increases with brood size, and parasitism can thus mediate trade-offs between reproduction and number and quality of the progeny in the barn swallow.     

Hatching asynchrony and brood reduction influence immune response in Common Kestrel Falco tinnunculus nestlings

The onset of incubation before the end of laying imposes asynchrony at hatching and, therefore, a size hierarchy in the brood. It has been argued that hatching asynchrony might be a strategy to improve reproductive output in terms of quality or quantity of offspring. However, little is known about the mediating effect of hatching asynchrony on offspring quality when brood reduction occurs. Here, we investigate the relationship between phenotypic quality and hatching asynchrony in Common Kestrel Falco tinnunculus nestlings in Spain. Hatching asynchrony did not increase breeding success or nestling quality. Furthermore, hatching asynchrony and brood reduction had different effects on nestlings’ phytohaematogglutinin (PHA)‐mediated immune response and nestling growth. In asynchronous and reduced broods (in which at least one nestling died), nestlings showed a stronger PHA‐mediated immune response and tended to have a smaller body size compared with nestlings raised in synchronous and reduced broods. When brood reduction occurred in broods hatched synchronously, there was no effect on nestling size, but nestlings had a relatively poor PHA‐mediated immune response compared with nestlings raised in asynchronous and reduced broods. We suggest that resources for growth can be directed to immune function only in asynchronously hatched broods, resulting in improved nestling quality, as suggested by their immune response. We also found that males produced a greater PHA‐mediated immune response than females only in brood‐reduced nests without any effect on nestling size or condition, suggesting that females may trade off immune activities and body condition, size or weight. Overall, our results suggest that hatching pattern and brood reduction may mediate resource allocation to different fitness traits. They also highlight that the resolution of immune‐related trade‐offs when brood reduction occurs may differ between male and female nestlings.     

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.     

Immune function and survival of great tit nestlings in relation to growth conditions

Life history theory predicts a trade-off between number and quality of offspring. Reduced quality with increasing brood size may arise from a decrease in body condition or in immunocompetence that would be important in fighting off virulent parasites by immunologically naive offspring. We tested the effect of rearing conditions on immune function of nestling great tits (Parus major) by reducing or increasing broods by two hatchlings. In the middle of the nestling period (on day 8), nestlings from enlarged broods developed lower T cell responses [as measured from the cutaneous swelling reaction to injection with phytohaemagglutinin (PHA)] and tended to have lower total leukocyte and lymphocyte concentrations in their peripheral blood than nestlings from reduced broods. Brood size manipulation affected the PHA response of nestlings most strongly in small clutches, suggesting that nestling immune function was dependent on their parents’ condition, as estimated by original clutch size. Intra-brood differences in nestling mortality were unrelated to immune parameters, but nestlings in broods without mortality had a stronger PHA response, higher concentration of lymphocytes and higher body mass on day 15 than nestlings in broods with mortality. These results support the prediction that the immune function of altricial birds is affected by rearing conditions, and that growth and immune parameters are related to inter-brood differences in nestling survival. Received: 1 February 1999 / Accepted: 19. July 1999     

Stress,immunocompetence and leukocyte profiles of pied flycatchers in relation to brood size manipulation

The two main trade-offs considered determining reproductive patterns in iteroparous organisms are the one between current and future reproduction, and the one between the number and quality of offspring. Recently, it has been suggested that these trade-offs may be mediated by stress-induced reduction in immunocompetence. To test the hypothesis that stress reduces immune function, we investigated the effects of brood size manipulation on stress hormone levels, leukocyte profiles and immune responses against challenge with novel antigens in nestling and parent male pied flycatchers ( Ficedula hypoleuca). In male parents, heterophil (H) and lymphocyte (L) numbers, as well as H/L ratio increased with experimentally enlarged brood size, and corticosterone levels tended to do so, indicating that high parental work load altered their stress level and physiological state. Despite this, we found no effects on humoral immune responsiveness, measured as antibody production against diphtheria-tetanus vaccine. In nestlings, heterophil numbers and H/L ratio increased in enlarged broods, whereas T-cell-mediated immune responsiveness, measured against phytohemagglutinin (PHA), decreased in enlarged broods. The results support the view that growth-stress-induced immunosuppression may be an important physiological pathway mediating the trade-off between the number and viability of offspring. The difference in the observed immune-related responses between nestlings and males may be because we measured different aspects of the immune system (cellular vs humoral). However, it may also be a result of males lowering their own costs by feeding less, (and their mate possibly compensate by feeding more), whereas nestlings cannot escape the costs of increased intra-brood competition.     

Nestlings’ carotenoid feather ornament affects parental allocation strategy and reduces maternal survival

In some birds, feather ornaments are expressed in nestlings well before sexual maturation, possibly in response to parental favouritism towards high‐quality offspring. In species with synchronous hatching, in which nestling ornaments may vary more among than within broods, parents may use this information to adjust their parental allocation to the current brood accordingly. We tested this hypothesis in the rock sparrow, in which a sexually selected yellow feather ornament is also expressed in nestlings. We experimentally enlarged nestlings’ breast patch in a group of broods and sham‐manipulated another group of control broods. Nestlings with enlarged ornament were fed more frequently and defended more actively from a dummy predator than their control counterparts. Mothers from the enlarged group were more likely to lay a second clutch and showed a reduced survival to the next breeding season. These results provide one of the first evidences of differential parental allocation among different broods based directly on nestlings’ ornamentation, and the first, to our knowledge, to show a reduction in maternal survival.     

Sudden weather deterioration but not brood size affects baseline corticosterone levels in nestling Alpine swifts

While evidence is accumulating that stress-induced glucocorticoid responses help organisms to quickly adjust their physiology and behaviour to life-threatening environmental perturbations, the function and the ecological factors inducing variation in baseline glucocorticoid levels remain poorly understood. In this study we investigated the effects of brood size by experimentally manipulating the number of nestlings per brood and the effect of weather condition on baseline corticosterone levels of nestling Alpine swifts (Apus melba). We also examined the potential negative consequences of an elevation of baseline corticosterone on nestling immunity by correlating corticosterone levels with ectoparasite intensity and the antibody production towards a vaccine. Although nestlings reared in enlarged broods were in poorer condition than nestlings reared in reduced broods, they showed similar baseline corticosterone levels. In contrast, nestling baseline corticosterone levels were higher immediately after cold and rainy episodes with strong winds. Neither nestling infestation rate by ectoparastic flies nor nestling antibody production against a vaccine was correlated with baseline corticosterone levels. Thus, our results suggest that altricial Alpine swift nestlings can quickly modulate baseline corticosterone levels in response to unpredictable variations in meteorological perturbation but not to brood size which may be associated with the degree of sibling competition. Apparently, short-term elevations of baseline corticosterone have no negative effects on nestling immunocompetence.     

Nestling immunocompetence and testosterone covary with brood size in a songbird

The social and ecological conditions that individuals experience during early development have marked effects on their developmental trajectory. In songbirds, brood size is a key environmental factor affecting development, and experimental increases in brood size have been shown to have negative effects on growth, condition and fitness. Possible causes of decreased growth in chicks from enlarged broods are nutritional stress, crowding and increased social competition, i.e. environmental factors known to affect adult steroid levels (especially of testosterone and corticosteroids) in mammals and birds. Little, however, is known about environmental effects on steroid synthesis in nestlings. We addressed this question by following the development of zebra finch (Taeniopygia guttata) chicks that were cross-fostered and raised in different brood sizes. In line with previous findings, nestling growth and cell-mediated immunocompetence were negatively affected by brood size. Moreover, nestling testosterone levels covaried with treatment: plasma testosterone increased with experimental brood size. This result provides experimental evidence that levels of circulating testosterone in nestlings can be influenced by their physiological response to environmental conditions.     

Cost of reproduction, T-lymphocyte mediated immunocompetence and health status in female and nestling barn swallows Hirundo rustica

We investigate the trade-off between reproductive effort, health status and T-lymphocyte acquired immunity in female and nestling barn swallows Hirundo rustica using a brood size manipulation experiment. Maternal and total feeding effort increased with experimental brood size. Parents did not fully compensate for the increased food demand of the enlarged broods and as a consequence the per capita feeding rate of nestlings decreased with increasing experimental brood size. Body mass and a measure of T-cell mediated immunity in 12 days old nestlings also decreased with increasing experimental brood size. Different leucocyte concentrations and the heterophile/lymphocyte ratio – an index of stress – of nestlings did not change in relation to experimental brood size, suggesting that within brood competition did not affect stress to nestlings. The brood size manipulation had a significant effect on maternal T-cell mediated immunity, measured by the phytohemagglutinin skin test, but not on maternal body mass, haematocrit or differential or total white blood cell counts. Our results seem to support the prediction that under mild work stress females respond first by reducing the energetically expensive acquired immunity. Different leucocyte types and the heterophile/lymphocyte ratio appear less sensitive to parental workload.     

Forest fragmentation is associated with primary brood sex ratio in the treecreeper (Certhia familiaris)

We studied the primary brood sex ratio of an old-growth forest passerine, the Eurasian treecreeper (Certhia familiaris), along a gradient of forest fragmentation. We found evidence that male nestlings were more costly to produce, since they suffered twofold higher nestling mortality and were larger in body size than females. Furthermore, the proportion of males in the brood was positively associated with the provisioning rate and the amount of food delivered to the nestlings. During the first broods, a high edge density and a high proportion of pine forests around the nests were related to a decreased production of males. The densities of spiders, the main food of the treecreeper, were 38% higher on spruce trunks than on pine trunks. This suggests that pine-dominated territories with female-biased broods may have contained less food during the first broods. The observation was further supported by the fact that the feeding frequencies were lower in territories with high proportions of pines. In the second broods, territories with a high forest patch density produced female-biased broods, whereas high-quality territories with a large amount of deciduous trees and mixed forests produced male-biased broods. Our results suggest that habitat quality as measured by habitat characteristics is associated with sex allocation in free-living birds.     

ENVIRONMENTAL INFLUENCES ON GROWTH AND SURVIVAL OF NESTLING HOUSE MARTINS DELICHON URBICA

Growth of nestling House Martins was studied in relation to (a) conditions in the external environment and (b) aspects of their breeding biology. The dependence of growth performance on (1) hatchling weights, (2) relative difference in hatchling weights within broods, (3) brood size,(4) season, (5) earliness of breeding in relation to other pairs in the colony, (6) timing of breeding in relation to the median breeding week of the colony and (7) aerial food abundance, was investigated by step-down multiple regression analysis. Up to the stage of the peak brood weight, early laying, small brood sizes and high hatchling weights were associated with higher nestling growth rates. Large relative differences in hatchling weights however tended to depress mean brood weights and increase weight differences (= size hierarchies) within broods. These differences in hatchling weights were considered to contribute significantly to 23% of all nestling deaths, because small, late hatching nestlings suffered very high mortality even when food was abundant. The nestlings which died showed a progressive reduction on growth rates and all succumbed before the 11th nestling day. Because these differences in hatchling weights can be linked to the food supply during laying rather than immediately prior to their death, it is considered that the mortality of these nestlings can ultimately be attributed to the low quality of eggs from which they hatched. There was a tendency for pre-hatching factors to diminish in importance throughout growth, while post-hatching factors increased in importance and, with one exception, were responsible for explaining all the significant variance in the growth characteristics of fledglings. The exception was that differences in wing-lengths in broods could be linked with weight differences at hatching. Food shortages lowered average brood weights prior to fledging. Because pairs breeding during the median breeding week had lighter young, it was inferred that competition for food during this peak of breeding activity had the effect of lowering nestling growth performance, although the overall effect was considered to be small. Early breeding pairs tended to have larger broods, and these large broods showed a lowered growth performance. However, early breeding pairs had relatively smaller weight and wing-length differences, in broods of a given size, than occurred in broods of late breeders. It was therefore concluded that early breeding pairs had some attribute which tended to minimize certain disadvantages of large broods. This effect appeared to be linked to the pair, rather than to season or food supply.     

Is hatching asynchrony beneficial for the brood?

Many hypotheses have been proposed to explain why female birdsstart to incubate before clutch completion (IBCC). Some of thosesuggest that the resulting hatching asynchrony (HA) is adaptivebecause it increases the size hierarchy among offspring andin turn reduces nestling competition and energy demands duringthe peak feeding period. Others argue that IBCC is a good strategyin unpredictable environments. When food conditions deteriorate,the large size hierarchy quickly results in the death of thelast hatched nestlings, allowing the remaining ones to surviveand fledge in better condition. In comparison, under favorableconditions, all nestlings can fledge independent of hatchingorder. To test these hypotheses, we performed a brood size manipulationexperiment (as a simulation of good and bad years) in collaredflycatchers Ficedula albicollis and examined the effect of sizehierarchy on offspring and brood performance. We found thatchicks with an initial size disadvantage experienced reducedbody mass growth and had shorter feathers at fledging in bothreduced and enlarged broods. In enlarged broods, they also fledgedwith a smaller skeletal size. Although broods on average orparents could possibly still benefit from HA when food is scarce,this was not seen in the current study. Parental survival wasnot related to the size hierarchy in the broods, and the averagebody mass growth of the nestlings was slower in broods witha high initial size variance. We therefore conclude that HAand the resulting size hierarchy are probably detrimental forthe growth of nestlings in both good and bad years, at leastin species where nestling mortality does not occur early inlife.     

Age‐related prenatal maternal effects and postnatal breeding experience have different influences on nestling development in an altricial passerine

Reproductive success and nestling performance are related to the age of parents across several vertebrate taxa. However, because breeding experience and prenatal maternal investment in reproduction often covary, the source of these age‐related differences can be difficult to determine. In this study, we evaluated the influence of prenatal maternal effects and postnatal breeding experience on the performance of nestling tree swallows Tachycineta bicolor by conducting a carefully controlled partial cross‐fostering experiment. We swapped half‐broods of nestlings between the nest of a young first‐time breeding female and the nest of a female known to have previously raised and fledged young. Our manipulation did not influence the within‐brood nestling hierarchies, and controlled for the effects of egg laying order. We found that nestlings of older females were heavier just prior to fledging regardless of the breeding experience of the attending female. In addition, fledglings raised by experienced females grew their flight feathers faster, and had greater probability of fledging. Our study demonstrates that prenatal investment in reproduction by older females can have long‐term consequences on nestling mass, and suggests limited potential for compensatory mass gains prior to fledging. Because our analyses controlled for feeding rates, our results also suggest that foraging quantity and quality are not the only benefits nestlings gain by being raised by an experienced female.     

Nestling telomere shortening,but not telomere length,reflects developmental stress and predicts survival in wild birds

Developmental stressors often have long-term fitness consequences, but linking offspring traits to fitness prospects has remained a challenge. Telomere length predicts mortality in adult birds, and may provide a link between developmental conditions and fitness prospects. Here, we examine the effects of manipulated brood size on growth, telomere dynamics and post-fledging survival in free-living jackdaws. Nestlings in enlarged broods achieved lower mass and lost 21% more telomere repeats relative to nestlings in reduced broods, showing that developmental stress accelerates telomere shortening. Adult telomere length was positively correlated with their telomere length as nestling (r = 0.83). Thus, an advantage of long telomeres in nestlings is carried through to adulthood. Nestling telomere shortening predicted post-fledging survival and recruitment independent of manipulation and fledgling mass. This effect was strong, with a threefold difference in recruitment probability over the telomere shortening range. By contrast, absolute telomere length was neither affected by brood size manipulation nor related to survival. We conclude that telomere loss, but not absolute telomere length, links developmental conditions to subsequent survival and suggest that telomere shortening may provide a key to unravelling the physiological causes of developmental effects on fitness.     

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.     

Prudent investment in rearing nestlings in bull-headed shrikes

I investigated seasonal changes in the relationships between brood size, body mass of nestlings and body mass of parents of the bull-headed shrike, Lanius bucephalus, in Ishikari, northern Japan. When the broods were 12days old, the body mass of the heaviest nestling in a brood did not differ among brood sizes, or throughout the season. However, the body mass of the lightest nestlings in a brood was different among brood sizes. The body mass of the lightest nestling in five- and six-nestling broods decreased throughout the season. The lightest nestling in four-nestling broods, and the lightest and the second lightest nestlings in five-nestling broods, were significantly lighter than the heaviest nestling in broods of this size. It is likely that pairs with six nestlings at 12days old can feed at least five of these nestlings enough to ensure their survival . The standardized body mass of parents (SBM), which was defined as the body mass divided by the length of the tarsus, did not differ among brood sizes, or throughout the season. It is possible that the relationship between the constancy of the SBM and the seasonal decline in the body mass of nestlings indicates that bull-headed shrikes have a limit to their parental efforts.     

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.     

Genetic and environmental components of growth in nestling blue tits (Parus caeruleus)

We investigated the effect of brood‐size mediated food availability on the genetic and environmental components of nestling growth in the blue tit (Parus caeruleus), using a cross‐fostering technique. We found genetic variation for body size at most nestling ages, and for duration of mass increase, but not of tarsus growth. Hence, nestling growth in our study population seems to have the potential to evolve further. Furthermore, significant genotype–environment interactions indicated heritable variation in reaction norms of growth rates and growth periods, i.e. that our study population had a heritable plasticity in the growth response to environmental conditions. The decreasing phenotypic variance with nestling age indicated compensatory growth in all body traits. Furthermore, the period of weight increase was longer for nestlings growing up in enlarged broods, while there was no difference to reduced broods in the period of tarsus growth. At fledging, birds in enlarged broods had shorter tarsi and lower weights than birds in reduced broods, but there was no difference in wing length or body condition between the two experimental groups. The observed flexibility in nestling growth suggests that growing nestlings are able to respond adaptively to food constraint by protecting the growth of ecologically important traits.     

Haemoglobin concentration and body condition of nestling Great Tits Parus major : a comparison of first and second broods in two contrasting seasons

The physiological condition of nestling altricial birds depends on the quantity and quality of food delivered to them by parents. One indicator of the condition of Great Tit Parus major nestlings is the haemoglobin concentration in their blood. The present study demonstrates the influence of weather conditions (temperature and rainfall) on nestling haemoglobin concentrations during two consecutive breeding seasons in two different habitat types (parkland vs. woodland) in the city of Łódź in Central Poland. This influence probably results from the effects of weather on the trophic base of the Tits. Dry, hot weather strongly affected bush and herbal foliage later in the breeding season (mid-June to mid-July) in 2006, presumably by interfering with the development of herbivorous arthropod populations. This in turn caused food shortages for second broods of Great Tits, which resulted in nestlings having low haemoglobin levels. In the following year, temperature was on average lower, and rainfall was regular but not very heavy. These conditions enabled the development of arthropod assemblages, and the trophic base for birds was much richer. Haemoglobin concentrations in the blood of nestlings from second broods were significantly higher than those of first broods and, unexpectedly, second-brood nestlings in 2007 were on average in better physiological state than first-brood nestlings in 2006 in both habitats. The relationship between haemoglobin concentration, brood category and year was very similar to that for nestling body mass. However, it was independent of both body mass and brood size. In some years and under certain conditions, second broods can be more successful than first broods.