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In the Galapagos Islands, many endemic landbird populations are declining due to habitat degradation, food availability, introduced species and other factors. Given nestlings typically lack efficient defense mechanisms against parasites, hematophagous ectoparasites such as the larvae of the introduced Avian Vampire Fly, Philornis downsi, can impose high brood mortality and cause threatening population declines in Darwin finches and other landbirds. Here, we assess whether the food compensation hypothesis (i.e., the parents’ potential to compensate for deleterious parasite effects via increased food provisioning) applies to the Green Warbler-Finch. We differentiated nests with low or high infestation levels by P. downsi and quantified food provisioning rates of male and female parents, time females spent brooding nestlings, and nestling growth. Male provisioning rates, total provisioning rates and female brooding time did not significantly vary in relation to infestation levels, nor by the number of nestlings. Opposed to the predictions of the food compensation hypothesis, females showed significantly reduced provisioning rates at high infestation levels. Nestling body mass was significantly lower and there was a reduction of skeletal growth, although not significantly, in highly infested nests. The females’ response to high infestation may be due to parasites directly attacking and weakening brooding females, or else that females actively reduce current reproductive effort in favor of future reproduction. This life-history trade-off may be typical for Darwin finches and many tropical birds with long lifespans and therefore high residual reproductive value. Conservation strategies may not build on the potential for parental food compensation by this species.

After birth, an organism needs to invest both in somatic growth and in the development of efficient immune functions to counter the effects of pathogens, and hence an investment trade-off is predicted. To explore this trade-off, we simultaneously exposed nestling great tits (Parus major) to a common ectoparasite, while stimulating immune function. Using a 2 × 2 experimental design, we first infested half of the nests with hen fleas (Ceratophyllus gallinae) on day 3 post-hatch and later, on day 9-13 post- hatch, and then supplemented half of the nestlings within each nest with an immuno-enhancing amino acid (methio- nine). We then assessed the non-specific immune response by measuring both the inflammatory response to a lipopol- ysaccharide (LPS) and assessing the levels of acute phase proteins (APP). In parasite-infested nestlings, methio- nine had a negative effect on body mass close to fledging. Methionine had an immune-enhancing effect in the absence of ectoparasites only. The inflammatory response to LPS was significantly lower in nestlings infested with fleas and was also lower in nestlings supplemented with methionine. These patterns of immune responses suggest an immuno- suppressive effect of ectoparasites that could neutralise the immune-enhancing effect of methionine. Our study thus suggests that the trade-off between investment in life his- tory traits and immune function is only partly dependent on available resources, but shows that parasites may influence this trade-off in a more complex way, by also inhibiting important physiological functions.

Predation risk is a key driver for the evolution of reproductive strategies and life history traits. In birds, incubation behavior represents one form of parental care where trade-offs between time spent in incubation activities and self-maintenance activities are likely to change in response to predator pressure. This can have strong effects on embryonic development, but is still poorly understood. We investigated the effects of the presence of a nest predator on great tit (Parus major) incubation behavior and the subsequent effects of incubation on nestling morphological traits. We manipulated perceived predation risk using models of short-tailed weasels (Mustela erminea) in combination with great tit alarm calls specific to this predator. Directly after hatching, we swapped whole broods from treated nests with broods from untreated nests to disentangle treatment effects acting during the incubation period from potential carry-over effects on parental care acting on nestlings after hatching. In increased predation risk environments, the number of incubation sessions and recesses, but not their duration, was increased compared to the control group, and the nocturnal incubation session was longer when females were exposed to a predator. Eggs incubated by females under increased predation risk lost more mass over the incubation period compared to the control group. Also, male nestlings hatched from nests exposed to predators were lighter at hatching but were equivalent in weight to their control counterparts at fledging. This suggests that nest predation risk can influence some aspects of incubation rhythm and embryonic development, but has no long-term effects on nestling final body size.

Early-life trade-offs faced by developing offspring can have long-term consequences for their future fitness. Young offspring use begging displays to solicit resources from their parents and have been selected to grow fast to maximize survival. However, growth and begging behaviour are generally traded off against self-maintenance. Oxidative stress, a physiological mediator of life-history trade-offs, may play a major role in this trade-off by constraining, or being costly to, growth and begging behaviour. Yet, despite implications for the evolution of life-history strategies and parent-offspring conflicts, the interplay between growth, begging behaviour and resistance to oxidative stress remains to be investigated. We experimentally challenged wild great tit (Parus major) offspring by infesting nests with a common ectoparasite, the hen flea (Ceratophyllus gallinae), and simultaneously tested for compensating effects of increased vitamin E availability, a common dietary antioxidant. We further quantified the experimental treatment effects on offspring growth, begging intensity and oxidative stress. Flea-infested nestlings of both sexes showed reduced body mass during the first half of the nestling phase, but this effect vanished short before fledging. Begging intensity and oxidative stress of both sexes were unaffected by both experimental treatments. Feeding rates were not affected by the experimental treatments, but parents of flea-infested nests fed nestlings with a higher proportion of caterpillars, the main source of antioxidants. Additionally, female nestlings begged significantly less than males in control nests, whereas both sexes begged at similar rates in vitamin E-supplemented nests. Our study shows that a parasite exposure does not necessarily affect oxidative stress levels or begging intensity, but suggests that parents can compensate for negative effects of parasitism by modifying food composition. Furthermore, our results indicate that the begging capacity of the less competitive sex is constrained by antioxidant availability.

Avian mothers can influence offspring phenotype through the deposition of different compounds into eggs, such as antibodies, hormones and antioxidants. The concentration of carotenoids in yolk is larger than in maternal plasma, suggesting an important role of these compounds for offspring development. Since carotenoids have to be acquired from the diet, they may be available in limiting amounts to the mothers. Here, we investigated the role of egg carotenoids for offspring growth by experimentally increasing the concentration of yolk lutein, the main carotenoid in great tit (Parus major) yolk. We subsequently measured body condition, oxidative stress, immune response, plumage colouration and fledging success. Lutein increased body mass soon after hatching and fledging success, but did not affect tarsus length, oxidative stress, immune response and plumage colouration. The higher content of yolk lutein could have increased body mass by reducing oxidative stress caused by high metabolic rates of rapidly growing embryos or by promoting cell differentiation and proliferation. The positive effect of lutein on fledging success seems to be mediated by its influence on body mass 3 days post-hatch, since these two traits were correlated. The finding that our treatment did not affect traits measured later in the nestling period, except for fledging success, suggests that yolk lutein has short-term effects that are essential to increase survival until fledging. Our study shows the positive effect of yolk lutein on offspring survival in the great tit, and therefore suggests an important role of carotenoid-mediated maternal effects.

Development plasticity is a common evolutionary and phenotypic response to poor growth condition, in particular in organisms with determinate growth such as most birds and mammals. Because various body structures can contribute differently to overall fitness, natural selection will adjust the degree of plasticity of each structure to its proportionate contribution to fitness at a given life stage. Two non-mutually exclusive mechanisms can account for plasticity in the growth of offspring to compensate for the effect of parasites. First, if parasite infestation levels fluctuate over the nestling period, parasitized young may show reduced growth until peak parasite infestation, and accelerated growth once the conditions improve (the accelerated growth hypothesis). Secondly, if the period of tissue maturation is not fixed in time, hosts may grow slower than parasite-free hosts but for a longer period of time (the delayed maturation hypothesis). To test whether hosts compensate for the effects of parasites on their development, the load of the blood-sucking louse-fly Crataerina melbae Rondani in the nests of Alpine swifts, Apus melba Linnaeus, was increased or decreased experimentally. Parasite prevalence was 100% in both treatments, but intensity (no. of parasites per nestling) was significantly lower for deparasitized nestlings. In both treatments, parasite intensity increased up to halfway through the rearing period (i.e. 30 days of age) and decreased afterwards. In line with the accelerated growth hypothesis, wings of parasitized nestlings grew at a lower rate than those of deparasitized ones before the peak of parasite infestation, but at a greater rate after the peak. Louse-flies had no significant effect on the growth of body mass. In agreement with the delayed-maturation hypothesis, wings of parasitized nestlings grew for 3 additional days and were of similar size at fledging as in deparasitized birds. In summary, the present study shows in a wild bird population that nestling hosts can compensate for the effect of parasitism on their phenotype. It emphasizes the need to take the dynamics of parasite populations into account in studies of host-parasite relationships, and to investigate the effect of parasites on host development over the entire growing period rather than only at fledging, as employed traditionally.

A major aim of evolutionary biology is to understand why patterns of genomic variation vary among populations and species. Large-scale genomic studies of widespread species are useful for studying how the environment and demographic history shape patterns of genomic divergence, and with the continually decreasing cost of sequencing, such studies are now becoming feasible. Here, we carry out one of the most comprehensive surveys of genomic variation in a wild vertebrate to date; the great tit ( Parus major ) HapMap project. We screened ca 500,000 SNP markers across 647 individuals from 29 populations, spanning almost the entire geographic range of the European great tit subspecies. We found that genome-wide variation was consistent with a recent colonisation across Europe from a single refugium in the Balkans and/or Turkey, with bottlenecks and reduced genetic diversity in island populations. Differentiation across the genome was highly heterogeneous, with clear “islands of differentiation” even among populations which are ostensibly panmictic. These islands of differentiation were consistently found in regions of low recombination, suggesting that background selection can rapidly promote population differentiation among even the most recently colonised populations. We also detected genomic outlier regions that were unique to peripheral great tit populations, most likely as a result of recent directional selection at the range edges of this species. These “unique” outlier regions contained candidate genes for morphology, thermal adaptation and colouration, supporting previous research in this species, and providing avenues for future investigation. Our study suggests that comprehensive screens of genomic variation in wild organisms can provide unique insights into evolution. Author summary Studying patterns of genetic variation is a useful way of determining why populations and species differ in nature. Genetic variation is shaped by natural selection, but also by the present and past size of populations, the amount of migration, and by features of the genome, such as variation in recombination rate, of the organism being studied. Teasing apart the effects of these different processes on genomic diversity is difficult, but one way that this can be achieved is by studying genomic variation across the entire range of a species. We performed a continental-scale analysis of genetic variation in the great tit - a widespread songbird that has been the focus of extensive ecological research. We first used genomic data to reconstruct the historical colonisation of great tits across Europe, and showed that during the last ice age, this species was likely restricted to a single region in Eastern Europe, from which they spread across the continent. We then studied how patterns of variation differ along the genome, and show that recombination rate is a key driver of variation among all populations. Importantly, by comparing many populations we were able to identify genes that have been subject to natural selection in specific geographical regions. We found that natural selection appeared to be strongest in populations on the edges of the great tit’s range acting on traits such as morphology, stress response and colouration. Large-scale genetic analyses such as ours are therefore useful approaches for understanding how evolution operates in the wild.

The use of alternative reproductive tactics (ARTs) is widespread in animals. Males of some species may change tactics depending on age, body condition and social environment. Many bat species are polygynous where a fraction of males only have access to fertile females. For polygynous bats, knowledge of the reproductive success of males using different ARTs is scarce, and it remains unclear how age of males is related to switching decisions between social statuses. We studied a large captive population of Carollia perspicillata, where males are either harem holders, bachelors or peripheral males. Using a multistate procedure, we modelled the age-related switches in reproductive tactics and in survival probability. From the model, we calculated the reproductive success and the frequencies of males displaying different reproductive tactics. As in mammals, the switch between social statuses is often related to age, we predicted that the transition probability of bachelor and peripheral males to harem status would increase with age. We show, however, that social status transition towards a harem holding position was not related to age. Reproductive success changed with age and social status. Harem males had a significantly higher reproductive success than bachelor males except between a short period from 3.8 to 4.4 years of age where success was similar, and a significantly higher reproductive success than peripheral males between 2.6 and 4.4 years of age. Harem males showed a clear decrease in the probability of maintaining social status with age, which suggests that senescence reduces resource holding potential.

Gene flow is usually thought to reduce genetic divergence and impede local adaptation by homogenising gene pools between populations. However, evidence for local adaptation and phenotypic differentiation in highly mobile species, experiencing high levels of gene flow, is emerging. Assessing population genetic structure at different spatial scales is thus a crucial step towards understanding mechanisms underlying intraspecific differentiation and diversification. Here, we studied the population genetic structure of a highly mobile species - the great tit Parus major - at different spatial scales. We analysed 884 individuals from 30 sites across Europe including 10 close-by sites (< 50 km), using 22 microsatellite markers. Overall we found a low but significant genetic differentiation among sites (F-ST = 0.008). Genetic differentiation was higher, and genetic diversity lower, in south-western Europe. These regional differences were statistically best explained by winter temperature. Overall, our results suggest that great tits form a single patchy metapopulation across Europe, in which genetic differentiation is independent of geographical distance and gene flow may be regulated by environmental factors via movements related to winter severity. This might have important implications for the evolutionary trajectories of sub-populations, especially in the context of climate change, and calls for future investigations of local differences in costs and benefits of philopatry at large scales. (C) 2016 The Linnean Society of London, The study was funded by the Swiss National Science Foundation (3100A0-102017 to HR, P2BEP3_152103 to KL and PMPDP3_151361/161858 to ML). We thank L. Gustafsson and J. Forsman who kindly allowed us to use their nest boxes. We gratefully thank E. Bezault, L. Cornetti and two anonymous reviewers for valuable advice on genetic analyses and helpful comments on the manuscript. All samples were collected under licenses of national authorities and financially supported by the Academy of Finland grant (to NP and 265859 to TE), the Netherland Organisation for Scientific Research (NWO-VICI 86503003 to JK and NWO-VICI to MV), the Netherlands Genomics Initiative (Horizon grant to KvO), the Hungarian Scientific Research Fund (OTKA 75618 to JT), the Estonian Ministry of Education and Science (IUT 34-8 to RM), the OSU-OREME, the Spanish Ministry of Education and Science (CGL2010-21933-CO-02 to EB), the Portuguese Foundation for Science and Technology (SFRH/BD/13594/2003 to ACN) and the Agence Nationale de la Recherche (ANR-08-JCJC-0041-01 to BD).

In polygynous mating systems, few males have stable access to sexual mates. With an expected higher copulation rate, harem males may deplete seminal fluids or increase epididymal sperm maturation, generating poor sperm quality. In a first study, we reported a higher sperm quality in sneaker males of Carollia perspicillata. To test whether the lower sperm quality observed in harem males was generated by an elevated copulation rate, we temporarily removed males of both social statuses from the colony. We thus assessed status-related changes of sperm quality resulting from sexual abstinence. Moreover, released from territory and female guarding, harem males were expected to show a reduction in somatic costs. On the basis of sperm competition models, we predicted a higher resource investment in the ejaculate with the reduction of pre-copulatory efforts. In line with our predictions, sperm quality of harem males improved significantly in contrast to sneaker males, whose sperm quality did not change. Without an increase in ejaculate lipid peroxidation, our results also provide evidence that the duration of sexual abstinence was not sufficient to generate sperm oxidative damage through senescence. Harem males did not show a reduction in blood lipid peroxidation or in the ratio of oxidized to reduced glutathione. In line with the maintenance of these somatic costs, harem males did not invest more superoxide dismutase to the ejaculate to maintain sperm quality. Our results suggest that a difference in copulation rate rather than an adaptation to sperm competition provides sneaker males with higher sperm quality in C. perspicillata.

Sperm competition exerts strong selection on males to produce spermatozoa with an optimal morphology that maximizes their fertilization success. Long sperm were first suggested to be favored because they should swim faster. However, studies that investigated the relationship between sperm length and sperm competitive ability or sperm swimming velocity yielded contradictory results. More recently, ratios of the different sections of a spermatozoon (the head, midpiece, and flagellum) were suggested to be more crucial in determining swimming velocity. Additionally, sperm ability to remain and survive in the female storage organs may also influence fertilization success, so that optimal sperm morphology may rather maximize sperm longevity than velocity. In this study, we investigated how sperm morphology is related to sperm velocity and sperm longevity in the house sparrow Passer domesticus. Sperm velocity was found to be correlated with head/flagellum ratio. Sperm with small heads relative to their flagellum showed higher swimming velocity. Additionally, shorter sperm were found to live longer. Finally, we found sperm morphological traits to vary substantially within males and the head/flagellum ratio to be unrelated to total sperm length. We discuss the hypothesis that the substantial within-male variation in sperm morphology reflects a male strategy to produce a diversity of sperm from long, fast-swimming to short, long-living sperm to maximize their fertilization success in a context of sperm competition.

Carotenoid-based colours are widespread in animals and are used as signals in intra- and interspecific communication. In nestling birds, the carotenoids used for feather pigmentation may derive via three pathways: (1) via maternal transfer to egg yolk; (2) via paternal feeds early after hatching when females are mainly brooding; or (3) via feeds from both parents later in nestling life. We analysed the relative importance of the proposed carotenoid sources in a field experiment on great tit nestlings ( Parus major). In a within-brood design we supplemented nestlings with carotenoids shortly after hatching, later on in the nestling life, or with a placebo. We show that the carotenoid-based colour expression of nestlings is modified maximally during the first 6 days after hatching. It reveals that the observed variation in carotenoid-based coloration is based only on mechanisms acting during a short period of time in early nestling life. The experiment further suggests that paternally derived carotenoids are the most important determinants of nestling plumage colour.

The striking variation in colour and maculation of bird eggs has fascinated biologists since centuries, and many hypotheses based on mechanical, physiological or signalling functions have been proposed for its evolution by natural and sexual selection. Protoporphyrin is the main eggshell pigment found in brown maculated eggs, and is assumed to function as a mediator of these selection processes. It is a precursor of heme with pro-oxidant properties, and hence a link between brown maculation and female condition has been proposed and tested in a number of studies, albeit with contrasting results. A variety of different visual methods have been used to quantify outer eggshell pigmentation, which has been assumed to correspond to overall quantity of protoporphyrin in the shell. Yet, this relationship has rarely been tested. The aim of this study was to apply four commonly used methods to assess pigmentation in great tit eggs with protoporphyrin as the predominant eggshell pigment, and to compare the results of these methods. Specifically, we 1) ranked eggshell pigmentation by human naked eye, 2) applied a granularity approach and 3) measured spectrophotometric reflectance of eggshell pigments. Second, we estimated the relationship between outer eggshell pigmentation (i.e. estimated by three different methods above) and true protoporphyrin concentration deposited in the entire shell measured by HPLC. Among-method estimates were significantly correlated for the traits describing pigment ‘darkness’ only. While the model including scores based on human naked eyes explained 16% of the variance of pigment concentration in the entire shell, spectrometry explained 27%, and the granularity approach explained 40%. Thus, the estimation of true pigment concentration in the entire shell from the visible outer side of the shell is most reliable with the granularity approach. It is relevant for studies where the maintenance of the integrity of the eggs is essential.

Scientific interests and conservation needs currently stress the necessity to better understanding bat reproductive biology. In this study, we present the first, safe, inexpensive, and reliable method to obtain sperm from a microbat species (Carollia perspicillata) by electroejaculation. This method revealed to be highly efficient (100% success rate). We obtained ejaculates composed of two characteristically different fractions. We compared three buffers and recommend using an Earle's balanced salt solution as a semen extender. Earle's balanced salt solution provided significant repeatable measure of swimming ability (intraclass correlation coefficient: 0.74, P 0.01) and proportion of motile sperms (intraclass correlation coefficient: 0.08, P = 0.01) and allowed sperm to maintain optimal swimming capacity over time. None of the buffers could dissolve all the coagulated sperm. Although the trypsin buffer freed a larger fraction of spermatozoa in the ejaculate, it impaired swimming ability without improving motility, viability, and stamina. We thus argue that the sperm population analyzed with Earle's balanced salt solution is a representative of the ejaculate. Finally, we found that the mean sperm velocity of C perspicillata (78.8 μm/s) is lower than that predicted by regressing sperm velocity on relative testes mass, a proxy of sperm competition. The question as to whether C perspicillata is an outsider for sperm velocity, or whether bats evolved yet another unique mechanism to cope with sperm competition deserves more investigations.

Life-history theory predicts a trade-off for allocation of limited resources to reproduction and self-maintenance; however, many of the underlying physiological mechanisms remain elusive. There is growing evidence for oxidative stress to play an essential role in this trade-off because some by-products from the immune system and from normal metabolism generate reactive oxygen species that can cause oxidative damage. We manipulated reproductive effort of male and female great tits shortly before reproduction by clipping feathers of either the male or female parent of pairs of known age, given that parental effort may differ between the sexes and change over the lifetime of an individual. We quantified the effect of the treatment on morphological, physiological, behavioral, and reproductive traits. We found that feather clipping led to a decrease in parental body mass and to a reduced clutch size. Nestlings raised by clipped fathers showed reduced body mass although feeding rate was equally high between clipped and control individuals. In contrast to our predictions, we found that the feather clipping did not affect oxidative status. However, independently of the treatment, adult males had higher antioxidant capacity than females and older males showed higher oxidative damage compared with yearlings. Thus, our results suggest that the self-maintenance was prioritized over reproduction. It suggests that males are more susceptible to increased workload than females and thus more likely to reduce allocation of resources to reproduction.

The evolution of conspicuous begging displays has been suggested as the outcome of a conflict where offspring attempt to manipulate food allocation beyond the parental optimum. One resolution for the conflict arises via costs of begging, and oxidative stress has been proposed as a major mechanism for causing begging-induced costs. Although begging can be a physically demanding activity, the evidence for causing oxidative stress is scarce. Great tit (Parus major) parents provide food at the nest mostly from 2 different locations, which in consequence relaxes nestling competition. Here, we manipulated nestling competition by forcing parents to feed from a single location and supplemented half of the nestlings in each brood with vitamin E to test if this major antioxidant can alleviate a potential oxidative cost of begging. The design increases the cost of begging without altering parental feeding rates. Begging intensity was significantly higher when parents fed from a single location. Body mass and antioxidant capacity were not affected by the increase in begging, but oxidative damage was lower in females of the increased begging group compared with those in the control group, independent of vitamin E supplementation. The results suggest that oxidative stress is rather a minor cost of begging. Vitamin E–supplemented nestlings had a higher probability to fledge, which underlines the important role of vitamin E during development, although this might not be due to its role as an antioxidant.

The activation of immune defences counteracts pathogens but mounting an immune response is costly and can negatively impact life history traits. Immune activation releases highly reactive species that kill pathogens but can also cause oxidative damage to host tissues and these negative effects may therefore constrain further investment in immune responses. To offset these toxic effects animals rely on a complex system of antioxidants. Here we tested if vitamin E a dietary antioxidant can reduce oxidative damage induced by an immune challenge and thus enhance the immune response. In a 2 x 2 experimental design we supplemented great tit nestlings with either vitamin E or a placebo and then injected them with either a bacterial lipopolysaccharide (LPS) or a buffer solution (PBS) as a control. LPS treated nestlings mounted an inflammatory response and increased antioxidant capacity without any change in ROM (reactive oxygen metabolites) an index of early oxidative damage. These results suggest that the likely transient increase in reactive species of the LPS injection was counteracted by a rise in endogenous antioxidant defences that was independent of supplementary dietary antioxidants. Indeed vitamin E supplementation neither affected oxidative status nor enhanced the immune response suggesting that in our experimental condition great tit nestlings were not limited in vitamin E and in antioxidants in general. Overall our results show that birds can mount an effective antioxidant response to face an immune challenge and can therefore avoid stress caused by a transient increase in reactive species generated by immune activation. (C) 2014 Elsevier Inc. All rights reserved.

Aggressive behaviours have an important impact on the social organization of animals and on the social status of individuals especially in gregarious species. Agonistic interactions between territory holders are essential to set and reinforce territorial borders. Additionally agonistic displays are used to demonstrate ownership of a territorial site and may indicate social status of the signaller. Between neighbouring territory owners dynamic borders require frequent interactions. In daily repeated aggressive encounters ritualization can help to avoid the need for serious fights and their costly consequences. In the bat Carollia perspicillata a gregarious frugivore with resource defence polygyny males defend territories at valuable roosting sites for females. Working with a captive bat colony of about 400 individuals housed under seminatural conditions in a tropical zoo we found that males defended territories aggressively by the use of a succession of displays forming a ritualized structure. Simultaneously males used three different vocalization types during aggressive displays namely down sweeps warbles and aggressive trills. A statistical analysis of 58 aggressive trills from five adult males showed that they contained sufficient variation to encode an individual signature. Using a habituation dishabituation paradigm playback experiment we found that males could discriminate between vocalizing males based on aggressive calls alone. Such discrimination is probably useful for distinguishing between neighbouring territory owners and more unfamiliar intruders and thus allows for an economical response. (C) 2014 The Association for the Study of Animal Behaviour. Published by Elsevier Ltd. All rights reserved.

Early-life stressful conditions can shape individual phenotypes and ultimately influence fitness. Oxidative stress is a pervasive threat that affects many fitness-related traits and can modulate life-history trade-offs. Yet, the extent to which exposure to oxidative stress during early life can have long-lasting effects on key fitness-related traits remains to be elucidated, particularly in natural populations of vertebrates. Using a wild population of great tits Parus major, we experimentally dosed 11-day-old birds with paraquat, a pro-oxidant molecule, aiming at increasing oxidative stress. One year later, we recaptured 39 of them as adult recruiting breeders and quantified effects of the paraquat exposure on their resistance to oxidative stress, carotenoid-based plumage coloration and male sperm performance. Despite the absence of a short-term effect of paraquat on oxidative stress measured two days later, the pre-fledging exposure to paraquat induced a reduction in individual oxidative damage measured at adulthood. Paraquat-dosed individuals also had brighter plumage, but no effect was observed on male sperm performance. For the first time in a natural population of vertebrates, we experimentally show that an early-life acute exposure to a pro-oxidant has long-lasting effects on individual resistance to oxidative stress at adulthood. Our results are in line with the environmental matching and the hormesis hypotheses but may also reflect selective disappearance of individuals with lower resistance to oxidative stress.

Ectoparasites are a ubiquitous environmental component of breeding birds, and it has repeatedly been shown that hematophagous ectoparasites such as fleas and mites reduce the quality and number of offspring of bird hosts, thereby lowering the value of a current brood. Selection acting on the hosts will favor physiological and behavioral responses that will reduce the parasites' impact. However, the results of the few bird studies that addressed the question of whether parasitism leads to a higher rate of food provisioning are equivocal, and the begging response to infestation has rarely been quantified. A change in begging activity and parental rate of food provisioning could be predicted in either direction: parents could reduce their investment in the brood in order to invest more in future broods, or they could increase their investment in order to compensate for the parasites' effect on the current brood. Since the nestlings are weakened by the ectoparasites they may beg less, but on the other hand they may beg more in order to obtain more food. In this study we show experimentally that (1) hen fleas (Ceratophyllus gallinae) reduce the body mass and size of great tit (Parus major) nestlings, (2) nestlings of parasitized broods more than double their begging rate, (3) the male parents increase the frequency of feeding trips by over 50%, (4) the females do not adjust feeding rate to the lowered nutritional state of nestlings, and (5) food competition among siblings of parasitized broods is increased. Ultimately the difference in the parental feeding response may be understood as the result of a sex-related difference in the trade-off of investing in current versus future broods.

Reproduction is known to increase the basal metabolic rate and generate oxidative stress, a possible proximate cost of reproduction. Carotenoids have been shown to be in vitro antioxidant molecules and, in a number of instances, to contribute in vivo to the antioxidant protection of the organism against the deleterious effects of free radicals and oxidative stress. These compounds are also involved in the up- and downregulation of the immune system. Thus, carotenoids may improve a male’s health status and condition during breeding and enhance his attractiveness through, for example, a higher investment into mating activities. The differential allocation hypothesis predicts that females should invest more in reproduction when mated to more attractive partners. Therefore, a supplementary dose of carotenoids during breeding should increase male attractiveness and translate into higher reproductive success via a higher reproductive effort by their mate. We tested this hypothesis in great tits by supplementing males with carotenoids during their female’s fertile period. We subsequently transferred entire clutches into unmanipulated foster nests. Thus, any effect of our carotenoid supplementation to males on their reproductive success must be due to female differential reproductive investment. Offspring sired by carotenoid-supplemented males were found to grow bigger and heavier and to fledge more successfully. Carotenoid-supplemented males also lost less paternity. Our results illustrate the fitness benefits males can accrue from carotenoids and underline the selective pressure imposed on males to optimize carotenoid acquisition. Key words: carotenoids, differential allocation hypothesis, extrapair paternity, great tit, Parus major. [Behav Ecol]

Oxidative stress was recently demonstrated to affect several fitness-related traits and is now well recognized to shape animal life-history evolution. However, very little is known about how much resistance to oxidative stress is determined by genetic and environmental effects and hence about its potential for evolution, especially in wild populations. In addition, our knowledge of phenotypic sexual dimorphism and cross-sex genetic correlations in resistance to oxidative stress remains extremely limited despite important evolutionary implications. In free-living great tits (Parus major), we quantified heritability, common environmental effect, sexual dimorphism and cross-sex genetic correlation in offspring resistance to oxidative stress by performing a split-nest cross-fostering experiment where 155 broods were split, and all siblings (n = 791) translocated and raised in two other nests. Resistance to oxidative stress was measured as both oxidative damage to lipids and erythrocyte resistance to a controlled free-radical attack. Both measurements of oxidative stress showed low additive genetic variances, high common environmental effects and phenotypic sexual dimorphism with males showing a higher resistance to oxidative stress. Cross-sex genetic correlations were not different from unity, and we found no substantial heritability in resistance to oxidative stress at adult age measured on 39 individuals that recruited the subsequent year. Our study shows that individual ability to resist to oxidative stress is primarily influenced by the common environment and has a low heritability with a consequent low potential for evolution, at least at an early stage of life.

© 2014 The Authors. Nests are structures built to support and protect eggs and/or offspring from predators, parasites, and adverse weather conditions. Nests are mainly constructed prior to egg laying, meaning that parent birds must make decisions about nest site choice and nest building behavior before the start of egg-laying. Parent birds should be selected to choose nest sites and to build optimally sized nests, yet our current understanding of clutch size-nest size relationships is limited to small-scale studies performed over short time periods. Here, we quantified the relationship between clutch size and nest size, using an exhaustive database of 116 slope estimates based on 17,472 nests of 21 species of hole and non-hole-nesting birds. There was a significant, positive relationship between clutch size and the base area of the nest box or the nest, and this relationship did not differ significantly between open nesting and hole-nesting species. The slope of the relationship showed significant intraspecific and interspecific heterogeneity among four species of secondary hole-nesting species, but also among all 116 slope estimates. The estimated relationship between clutch size and nest box base area in study sites with more than a single size of nest box was not significantly different from the relationship using studies with only a single size of nest box. The slope of the relationship between clutch size and nest base area in different species of birds was significantly negatively related to minimum base area, and less so to maximum base area in a given study. These findings are consistent with the hypothesis that bird species have a general reaction norm reflecting the relationship between nest size and clutch size. Further, they suggest that scientists may influence the clutch size decisions of hole-nesting birds through the provisioning of nest boxes of varying sizes.

Summary Oxidative stress is the imbalance between the production of reactive species and antioxidants, which causes damage to lipids, proteins and DNA. Antioxidants, like vitamins and carotenoids, can limit oxidative damage and can therefore regulate the trade-off between growth, which is a period of high reactive species production, and self-maintenance. However the role of carotenoids as antioxidant in vivo has been debated, and it has been suggested that carotenoid-based signals indicate the availability of non-pigmentary antioxidants (e.g. vitamins) that protect carotenoids from oxidation, known as the "protection hypothesis". To evaluate the importance of vitamins versus carotenoids as antioxidants during growth and to test the protection hypothesis, we supplemented nestling great tits Parus major 3, 5 and 7 days after hatching with a single dose of carotenoids and/or vitamins in a 2x2 full-factorial design. We subsequently measured body condition, antioxidant capacity, oxidative damage, fledging success and plumage reflectance. Vitamins enhanced antioxidant capacity, but did not affect oxidative damage. Vitamin-treated nestlings had higher growth rates and higher probability to fledge. In contrast, carotenoids did not affect any of these traits. Furthermore carotenoid-based colouration increased over the breeding season in nestlings that received vitamins only. This study shows that vitamins are limiting for growth rate and fledging success, and it suggests that vitamins could regulate the trade-off between growth and self-maintenance in favour of the former. Moreover our results are consistent with the idea that carotenoids are minor antioxidants in birds, but they do not support the "protection hypothesis".

In social systems with alternative reproductive tactics, sneakers face a higher level of sperm competition than harem males and hence are predicted to allocate more resources to ejaculates. Antioxidants can protect sperm against oxidative stress, and thus, their allocation to the ejaculate may depend on mating tactic. In this study on the frugivorous bat Carollia perspicillata, we assessed, for harem and sneaker males, four sperm mobility traits, blood and ejaculate markers of the redox balance and the ejaculate to blood ratios of the redox markers. Under higher sperm competition, sneaker males should allocate proportionally more antioxidant resources to the protection of sperm than harem males. In contrast, harem males should favour pre-copulatory functions, which comprise the protection of blood. We found significantly higher sperm velocity and sperm survival in sneakers. There was no correlation between sperm mobility and sperm enzymatic antioxidant activity or ejaculate levels of lipid peroxidation (oxidative damage). Ejaculate levels of lipid peroxidation and sperm survival showed a significantly positive correlation, which could be attributed to the role of reactive oxygen species for sperm capacitation. Harem and sneaker males showed similar levels of redox balance markers in ejaculate and blood. However, harem males showed a higher ratio of oxidized over reduced glutathione in blood, which may indicate higher cellular stress due to higher metabolism. Overall, our findings suggest that sneakers of C. perspicillata compensate for a higher level of sperm competition by higher sperm mobility. In social systems with alternative reproductive tactics, sneakers face higher level of sperm competition than harem males and hence are predicted to allocate more resources to ejaculates. Antioxidants can protect sperm against oxidative stress, and thus, their allocation to the ejaculate may depend on mating tactic. In this study on the frugivorous bat Carollia perspicillata, we found sperm swimming significantly faster and longer in sneaker males compared to harem males. However, traits other than the investigated antioxidant may favour higher sperm mobility. Measured redox pattern in blood of harem males may indicate higher cellular stress due to higher metabolism. Our results provide support to the current sperm competition models at the intraspecific level, which is still debated for internal fertilizers. This study contributes to better understanding the trade-offs and adaptations resulting from alternative reproductive tactics in mammals.

In many bird species offspring hatch over hours or days which leads to an age and size hierarchy within broods. The function of hatching asynchrony is much debated and it has been suggested that the induced size hierarchy among offspring may be an adaptive maternal mechanism for maximizing reproductive output under variable environmental conditions. The best known hypothesis to explain the adaptive value of hatching asynchrony the `brood reduction' hypothesis holds that a size hierarchy among offspring allows for a quick adaptive adjustment of brood size to unpredictable feeding conditions and thus benefits parents. To test the consequences of hatching asynchrony on offspring growth and food provisioning we experimentally manipulated the onset of incubation of eggs within broods of great tits Parus major to induce either synchronous or asynchronous hatching and then manipulated brood size after hatching to simulate favourable control or harsh conditions. We did not find a difference in offspring mortality between asynchronous and synchronous broods under any of these conditions. In harsh conditions nestling mass and size were reduced in asynchronous broods compared to synchronous ones. The opposite pattern occurred under control conditions. Although our results showed that induced hatching asynchrony alters offspring phenotype they do not provide strong support for the brood reduction hypothesis as a mechanism to explain hatching asynchrony. (C) 2013 The Association for the Study of Animal Behaviour. Published by Elsevier Ltd. All rights reserved.

Marie Vaugoyeau [et al.], The increase in size of human populations in urban and agricultural areas has resulted in considerable habitat conversion globally. Such anthropogenic areas have specific environmental characteristics, which influence the physiology, life history, and population dynamics of plants and animals. For example, the date of bud burst is advanced in urban compared to nearby natural areas. In some birds, breeding success is determined by synchrony between timing of breeding and peak food abundance. Pertinently, caterpillars are an important food source for the nestlings of many bird species, and their abundance is influenced by environmental factors such as temperature and date of bud burst. Higher temperatures and advanced date of bud burst in urban areas could advance peak caterpillar abundance and thus affect breeding phenology of birds. In order to test whether laying date advance and clutch sizes decrease with the intensity of urbanization, we analyzed the timing of breeding and clutch size in relation to intensity of urbanization as a measure of human impact in 199 nest box plots across Europe, North Africa, and the Middle East (i.e., the Western Palearctic) for four species of hole-nesters: blue tits (Cyanistes caeruleus), great tits (Parus major), collared flycatchers (Ficedula albicollis), and pied flycatchers (Ficedula hypoleuca). Meanwhile, we estimated the intensity of urbanization as the density of buildings surrounding study plots measured on orthophotographs. For the four study species, the intensity of urbanization was not correlated with laying date. Clutch size in blue and great tits does not seem affected by the intensity of urbanization, while in collared and pied flycatchers it decreased with increasing intensity of urbanization. This is the first large-scale study showing a species-specific major correlation between intensity of urbanization and the ecology of breeding. The underlying mechanisms for the relationships between life history and urbanization remain to be determined. We propose that effects of food abundance or quality, temperature, noise, pollution, or disturbance by humans may on their own or in combination affect laying date and/or clutch size., E. Barba acknowledges funding of project CGL2013-48001-C2-1-P (Spanish Ministry of Science and Innovation).

Host range is a key element of a parasite's ecology and evolution and can vary greatly depending on spatial scale. Generalist parasites frequently show local population structure in relation to alternative sympatric hosts (i.e. host races) and may thus be specialists at local scales. Here, we investigated local population specialization of a common avian nest-based parasite, the hen flea Ceratophyllus gallinae (Schrank), exploiting two abundant host species that share the same breeding sites, the great tit Parus major (Linnaeus) and the collared flycatcher Ficedula albicollis (Temminck). We performed a cross-infestation experiment of fleas between the two host species in two distinct study areas during a single breeding season and recorded the reproductive success of both hosts and parasites. In the following year, hosts were monitored again to assess the long-term impact of cross-infestation. Our results partly support the local specialization hypothesis: in great tit nests, tit fleas caused higher damage to their hosts than flycatcher fleas, and in collared flycatcher nests, flycatcher fleas had a faster larval development rates than tit fleas. However, these results were significant in only one of the two studied areas, suggesting that the location and history of the host population can modulate the specialization process. Caution is therefore called for when interpreting single location studies. More generally, our results emphasize the need to explicitly account for host diversity in order to understand the population ecology and evolutionary trajectory of generalist parasites.

Experimental studies incorporating multiple trophic levels are scarce but of increasing interest for understanding ecological communities. Here we investigated interactive effects of perceived predation risk and parasite pressure on life-history traits in a hole-nesting bird, and the effects of predation risk on parasite success. In a 3 × 2 experimental design we increased perceived predation risk for breeding great tits Parus major via simulations of either nest-predators (woodpeckers) or post-fledging predators (sparrowhawks) close to nests, and used a non-predatory species (song thrush) as a control. Concurrently, half of the nests in each treatment were either infested with ectoparasites, or kept parasite-free. Regarding the predation risk – parasite interaction, exposure to nest-predators tended to lower wing and sternum growth rates of nestlings in the absence, but not the presence, of parasites. In the presence of parasites, exposure to a post-fledging, but not to a nest-predator, led to significantly reduced wing growth. Mass and tarsus length were not affected by predator exposure, but ectoparasites had slight positive effects on mass gain. In the last third of the nestling period, overall nestling size was significantly smaller when exposed to a post-fledging predator than to a nest-predator, but neither differed from the control. Parental feeding rates were not affected by the treatments, but parents became less selective towards food items under either predation risk. Hen-flea population sizes (adult or larvae) in nests were not affected by predation risk treatment of hosts. In summary, we found some evidence for an interactive effect of predation risk and parasite pressure on nestling growth. The complexity of the interaction, combined with certain inconsistencies of the effects and potential statistical artifacts, prevent however a straightforward interpretation of the results. The insights from the study are useful for designing additional experiments to further investigate the complexity of predator–parasite interactions in wild populations.

Positive correlations between heterozygosity and fitness traits are frequently observed, and it has been hypothesized, but rarely tested experimentally, that parasites play a key role in mediating the heterozygosity-fitness association. We evaluated this hypothesis in a wild great tit (Parus major) population by testing the prediction that the heterozygosity-fitness association would appear in broods experimentally infested with a common ectoparasite, but not in parasite-free broods. We simultaneously assessed the effects of parental and offspring heterozygosity on nestling growth and found that body mass of nestlings close to independence, which is a strong predictor of post-fledging survival, increased significantly with nestling levels of heterozygosity in experimentally infested nests, but not in parasite-free nests. Heterozygosity level of the fathers also showed a significant positive correlation with offspring body mass under an experimental parasite load, whereas there was no correlation with the mothers' level of heterozygosity. Thus, our results indicate a key role for parasites as mediators of the heterozygosity-fitness correlations.

Exposure of mothers to risk of predation can induce phenotypic changes in offspring as shown in several species. We previously found that cross-fostered great tit (Parus major) chicks of females exposed to increased predation risk were smaller and lighter, but had faster wing growth than control cross-fostered chicks, possibly improving predator-escaping abilities. Here we examined the possible role of maternal steroids deposited in eggs as an underlying mechanism. We collected eggs from female great tits under either experimentally increased predation risk (PRED) or control treatments (CON) and analyzed the concentration of testosterone, androstenedione, and progesterone in the yolks. PRED eggs contained lower levels of testosterone than CON eggs, but levels of androstenedione and progesterone did not differ. The smaller size and mass of chicks found in the previous study may thus be explained by the lower testosterone concentrations, since yolk testosterone is known to boost growth and development. Alternatively, testosterone may act as a modulator of differential investment into morphological traits, rather than a simple growth enhancer, explaining lower body mass in conjunction with the accelerated wing growth. This could possibly occur concurrently with other hormones such as corticosterone. (C) 2012 Elsevier Inc. All rights reserved.

Social structures such as families emerge as outcomes of behavioural interactions among individuals, and can evolve over time if families with particular types of social structures tend to leave more individuals in subsequent generations. The social behaviour of interacting individuals is typically analysed as a series of multiple dyadic (pair-wise) interactions, rather than a network of interactions among multiple individuals. However, in species where parents feed dependant young, interactions within families nearly always involve more than two individuals simultaneously. Such social networks of interactions at least partly reflect conflicts of interest over the provision of costly parental investment. Consequently, variation in family network structure reflects variation in how conflicts of interest are resolved among family members. Despite its importance in understanding the evolution of emergent properties of social organization such as family life and cooperation, nothing is currently known about how selection acts on the structure of social networks. Here, we show that the social network structure of broods of begging nestling great tits Parus major predicts fitness in families. Although selection at the level of the individual favours large nestlings, selection at the level of the kin-group primarily favours families that resolve conflicts most effectively.

Oxidative stress is considered to act as a universal physiological constraint in life-history evolution of animals. This should be of interest for extra-pair paternity behaviour, and we tested here the prediction that offspring arising from extra-pair matings of female great tits show higher resistance to oxidative stress than within-pair offspring. Resistance to oxidative stress, measured as the whole blood resistance to a controlled free-radical attack, was significantly higher for extra-pair offspring as predicted although these were not heavier or in better body condition than within-pair offspring. Since resistance to oxidative stress has been suggested to enhance survival and reproductive rates, extra-pair offspring with superior resistance to oxidative stress, be it through maternal effects or paternal inheritance, may achieve higher fitness and thus provide significant indirect fitness benefits to their mothers. In addition, because oxidative stress affects colour signals and sperm traits, females may also gain fitness benefits by producing sons that are more attractive (sexy-sons hypothesis) and have sperm of superior quality (sexy-sperm hypothesis). Heritability of resistance to oxidative stress as well as maternal effects may both act as proximate mechanisms for the observed result. Disentangling these two mechanisms would require an experimental approach. Future long-term studies should also aim at experimentally testing whether higher resistance to oxidative stress of EP nestlings indeed translates into fitness benefits to females.

Organisms are often simultaneously prey for predators and hosts for parasites. The two challenges require different defence mechanisms which may be traded off if based on the same resources. We investigated how predation risk perceived by mothers affects via a maternal effect the survival of parasites feeding on offspring. In an experiment on great tits Parus major we manipulated the perceived predator density for females before and during ovulation by simulating the presence of either birds of prey (sparrowhawks) or controls (song thrushes) in their breeding habitat. To test exclusively for the maternal effect as a consequence of the higher perceived predation risk the offspring were raised by foster parents without treatment. We then let groups of hen fleas the main ectoparasite of the great tit feed on nestlings from both treatment groups for a fixed period of time and thereafter measured flea survival under standardized laboratory conditions. Fleas feeding on offspring of mothers experiencing increased predation risk before egg laying survived significantly longer than those feeding on control nestlings. Higher survival may be associated with maternal stressors transferred via the egg to the host offspring or may result from the resolution of an investment trade off whereby maternal resources can be used to prepare the offspring to deal with either increased predation risk or parasites. This finding emphasizes the need for understanding multitrophic interactions arising from the interplay between parasite and predator induced maternal effects and their consequences for parasite population dynamics the composition of ecological communities and host parasite coevolution. (C) 2011 The Association for the Study of Animal Behaviour. Published by Elsevier Ltd. All rights reserved.

Summary 1. Predation risk is known to influence behaviour and development, especially in breeding animals. Mothers may be selected to transfer information about the intensity of such risk to their offspring through maternal effects and thus influence their development. 2. Here, we test for this maternal effect via manipulation of perceived predation risk by exposing great tit females before and during ovulation to stuffed models and sounds of either a predator bird (sparrowhawk –Accipiter nisus) or of a non-predatory control (song thrush –Turdus philomelos) in their environment. Offspring of exposed mothers were then raised by foster parents subjected to no treatment in order to separate maternal effects from effects during post-hatching parental care. 3. Nestlings of mothers under increased predator density were smaller than those of control mothers yet showed higher growth rates of the wings. Additionally, first-year recruits from the predator treatment had longer wings at maturity. 4. This maternal effect may be a passive consequence of higher circulating stress hormone levels in mothers. The accelerated wing growth during the nestling stage may be a result of compensatory growth. 5. Alternatively, the accelerated wing growth, coupled with the longer wings at maturity, suggest that the maternal response to the environmental risk may be adaptive since lower weight and bigger wings are a selective advantage for predator evasion. In this case the maternal effect probably influences the distribution of resources to different growth functions in offspring. 6. We show for the first time through an environmental rather than a direct hormonal manipulation, that predation risk may elicit adaptive maternal effects in birds.

Oxidative stress has been suggested as a proximate cost of reproduction and hence as a major constraint in the evolution of life histories, and it is therefore thought that antioxidants alleviate the effects of reproductive effort on oxidative stress. Furthermore, carotenoid-based ornaments have been proposed to mirror male ability to resist oxidative stress. Using a full-factorial experimental design in a natural population of great tits Parus major, we manipulated brood size and supplemented the male parent with either carotenoids or a placebo. We then assessed antioxidant capacity via a measure of whole blood resistance to a free radical attack during the nestling rearing period. Males of enlarged broods showed impaired antioxidant capacity 5 days after the brood size manipulation. However, 13 days after manipulation, they had their antioxidant capacity restored, an effect that may be due to the development of compensatory antioxidant mechanisms or due to reduced investment in the current reproduction in favor of future survival and reproduction. Carotenoid supplementation did not affect male antioxidant capacity nor was the interaction with the brood manipulation significant. Males with stronger carotenoid-based plumage colors did not show higher antioxidant capacity 5 days after the brood size manipulation, but after 13 days, the relationship was highly significant. This study on a natural population shows that larger brood size can induce a transient decrease in antioxidant capacity. It also supports the hypothesis that carotenoid-based plumage may signal male ability to resist oxidative stress, particularly during the energetically demanding nestling rearing period. Key words: antioxidant capacity, brood size manipulation, carotenoid-based coloration, carotenoids, oxidative stress, sexual selection. [Behav Ecol]

Sperm cells are highly vulnerable to free radicals, and sperm quality and male fertility are critically affected by oxidative stress. Recently, sexual ornaments, particularly carotenoid-based colourful traits, have been proposed to depend on a male's capacity to resist oxidative stress, and thus to signal sperm quality. We conducted an experimental test of this hypothesis on great tits Parus major, in which adults are sexually dichromatic in carotenoid-based breast plumage. We report the first evidence that ornaments and sperm quality may be linked through oxidative stress. When experimentally subjected to oxidative stress resulting from increased workload, less colourful males suffered a greater reduction in sperm motility and swimming ability, and increased levels of sperm lipid peroxidation compared to more colourful males. Moreover, the level of sperm lipid peroxidation was negatively correlated with sperm quality. Finally, carotenoid supplementation increased sperm quality of less colourful males, suggesting that pale males are deficient in carotenoid antioxidants.

Early developmental conditions have major implications for an individual’s fitness. In species where offspring are born simultaneously, the level of sibling competition for food access is intense. In birds, high sibling competition may subject nestlings to decreased growth rate as a result of limited food and increased levels of oxidative stress through high metabolic activity induced by begging behaviors. We manipulated the level of sibling competition in a natural population of great tits and assessed the consequences for nestling body condition and resistance to oxidative stress. In a full factorial design, we both augmented brood size to increase sibling competition and supplemented the male parents with physiological doses of carotenoids thereby doubling the natural carotenoid intake, aiming at increasing the males’ investment in current reproduction and thereby decreasing sibling competition. Nestling body mass was reduced by the brood enlargement and enhanced by the carotenoid supplementation of fathers. Nestling resistance to oxidative stress, measured as total antioxidant defenses in whole blood, was not influenced by the treatments. Because nestlings experience high metabolic activities, an absence of an effect of sibling competition on free radicals production seems unlikely. Nestling body mass decreased and resistance to oxidative stress tended to increase with initial brood size, and hence these correlational effects suggest a trade-off between morphological growth and development of the antioxidant system. However, the result of the experimental treatment did not support this trade-off hypothesis. Alternatively, it suggests that nestling developed compensatory mechanisms that were not detected by our antioxidant capacity measure. Key words: antioxidant capacity, body condition, brood size manipulation, carotenoid supplementation, Parus major, sibling competition. [Behav Ecol]

During egg formation, female birds deposit antibodies against parasites and pathogens they were exposed to before egg laying into the yolk. In captive bird species, it has been shown that these maternal immunoglobulins (maternal yolk IgGs) can protect newly hatched offspring against infection. However, direct evidence for such benefits in wild birds is hitherto lacking. We investigated (1) if nestling Great Tits Parus major originating from eggs with naturally high levels of maternal yolk IgG are less susceptible to a common, nest-based ectoparasite, (2) if maternal yolk IgGs influence nestling development and in particular, their own immune defence, and (3) if there is a negative correlation between levels of maternal yolk IgG in host eggs and the reproductive success of ectoparasitic fleas feeding on the nestlings. Counter to expectations, we found no indication that maternally transferred yolk IgGs have direct beneficial effects on nestling development, nestling immune response or nestling resistance or tolerance to fleas. Furthermore, we found no negative correlation between host yolk IgG levels and parasite fecundity. Thus, whereas previous work has unequivocally shown that prenatal maternal effects play a crucial role in shaping the parasite resistance of nestling birds, our study indicates that other egg components, such as hormones, carotenoids or other immuno-active substances, which bird females can adjust more quickly than yolk IgG, might mediate these effects.