Ostrich chick humoral immune responses and growth rate are predicted by parental immune responses and paternal colouration

One of the most important measures of offspring performance is growth rate, which is often traded off against another important survival trait, immune function. A particular feature of ostrich chicks maintained in farmed environments is that cohorts of chicks vary widely in size. As parents can have a profound effect on the phenotype and fitness of their offspring, we investigated whether chick growth and immune defence were related to variation in levels of immune defence in their genetic parents. As secondary sexual traits of sires could serve as indicators of male quality, and be used in female mating decisions, we also investigated whether chick growth rate and immune defence were related to male plumage and integumentary colouration. We found that offspring growth rates and humoral responses were related to the humoral responses of their parents, suggesting that at least some components of humoral immune capacity are heritable. The white colour of male ostrich feathers was correlated to the humoral response and growth rate of their offspring, suggesting that this visual cue involved in the male courtship display could serve as an important signal to females of male quality, thereby forming the basis of mate choice in this species.     

Sex-specific effects of albumen removal and nest environment manipulation on Barn Swallow nestlings

In avian species, maternal provisioning to the eggs is predicted to be more valuable for the offspring under adverse environmental conditions and intense sibling competition. However, studies manipulating both the amount of maternal pre-hatching resources and the harshness of post-hatching environment have seldom been performed to date. In this experimental study of Barn Swallow (Hirundo rustica) nestlings, we tested the consequences of a reduction in the albumen content of the eggs for fitness-related offspring traits, while performing an unbalanced partial cross-fostering soon after hatching, either increasing or decreasing brood size by one nestling. By molecular sexing of the chicks, we additionally tested for sex-specific sensitivity of individual nestlings to experimental treatments and to sex ratio variation in nestmates. We predicted that chicks hatching from albumen-deprived eggs should suffer more than control chicks from the harsher rearing conditions of enlarged broods. However, although albumen removal depressed chick body mass, chicks hatching from control eggs did not fare better than those hatching from eggs with reduced albumen content in enlarged vs. reduced broods. Albumen removal had sex-specific effects on immunity, with males, but not females, hatching from eggs with reduced albumen content showing a lower T-cell-mediated immune response than controls, suggesting that the two sexes were differentially susceptible to resource deprivation during early ontogeny. In addition, both immune response and chick body mass at age 7 days, when maximum growth rate is attained, declined with an increasing proportion of male nestmates. The effect of brood size manipulation on chick body mass at age 12 days, when peak body mass is attained, was also found to depend on brood sex composition, in that an increase in the proportion of male nestmates depressed offspring body mass in reduced broods, while the reverse was true in enlarged broods. On the whole, these findings suggest that sex differences may exist in environmental sensitivity and patterns of resource allocation among different body functions, and that brood size variation and sex composition may affect offspring fitness-related traits.     

Short- and long-term consequences of prenatal testosterone for immune function: an experimental study in the zebra finch

Hormone-mediated maternal effects play an important role in the formation of a differentiated phenotype. They have been shown to influence a wide array of offspring traits, both early in life and in adulthood. One important offspring trait that is under the influence of maternal androgens is the immune system. In birds, a growing number of studies show that yolk androgens modulate immune function during the chick stage. However, there is a lack of knowledge regarding long-term effects of prenatal androgens on offspring immunity. In this study, we therefore investigated the influence of prenatal testosterone (T) on several measures of immunity in fledgling and adult zebra finches (Taeniopygia guttata). Cell-mediated immune response (towards phytohaemagglutinin, PHA) of fledglings hatching from control eggs was negatively related to brood size, whereas there was no such association for fledglings hatching from eggs with experimentally elevated T levels (T fledglings). Male control fledglings showed reduced mass gain compared to female control fledglings within 24 h after the PHA injection. This pattern was reversed in T fledglings. Total antibody levels in fledglings were not affected by egg treatment. Neither cell-mediated immunity nor total antibody levels in sexually mature zebra finches were influenced by egg treatment. However, there was an immuno-enhancing effect of elevated egg T on both primary and secondary humoral immune responses toward diphtheria and tetanus antigens in ca 5 and 7 month old zebra finches. In addition, the covariation between different immune components differed between T and control offspring, suggesting that egg treatment may have altered the potential trade-offs between different parts of the immune system. Our results suggest that prenatal androgens could be an important factor contributing to individual variation in immune function even in adulthood.     

Is natural hatching asynchrony optimal? An experimental investigation of sibling competition patterns in a facultatively siblicidal seabird

In unpredictable environments, any tactic that enables avian parents to adjust brood size and, thus, energy expenditure to environmental conditions should be favoured. Hatching asynchrony (HA), which occurs whenever incubation commences before clutch completion, may comprise such a tactic. For instance, the sibling rivalry hypothesis states that the hierarchy among chicks, concomitant to HA, should both facilitate the adjustment of brood size to environmental conditions and reduce several components of sibling competition as compared to synchronous hatching, at both brood and individual levels. We thus predicted that brood aggression, begging and feeding rates should decrease and that older chick superiority should increase with HA increasing, leading to higher growth and survival rates. Accordingly, we investigated the effects of an experimental upward and downward manipulation of HA magnitude on behaviour, growth and survival of black-legged kittiwake (Rissa tridactyla) chicks. In line with the sibling rivalry hypothesis, synchronous hatching increased aggression and tended to increase feeding rates by parents at the brood level. Begging rates, however, increased with HA contrary to our expectations. At the individual level, as HA magnitude increased, the younger chick was attacked and begged proportionally more often, experienced a slower growth and a higher mortality than its sibling. Overall, the occurrence of energetic costs triggered by synchronous hatching both for parents and chicks, together with the lower growth rate and increased mortality of the younger chick in highly asynchronous broods suggest that natural HA magnitude may be optimal.     

Food availability and immune capacity in serin (Serinus serinus) nestlings

Parental feeding effort is an important determinant of chick development in birds. Quality and amount of food provided to each nestling, however, might affect the development of the immune system. Food availability around the nest site may account for part of the difference in body condition and immune capacity of nestlings. We tested this idea in the socially monogamous serin (Serinus serinus), investigating growth and immune capacity of nestlings in relation to food availability around the nest site. We used a set of immunological and serological assays, including antibody responses to sheep red blood cells (SRBCs), the T lymphocyte cell-mediated immune response (PHA), granulocyte:lymphocyte ratio (G:L), haematocrit and sedimentation rate. The results suggest that serins raised under poor environmental conditions were less immunocompetent. The immune response to PHA and SRBCs significantly increased with food availability. Haematocrit values were positively, sedimentation rate and the G:L ratio negatively correlated with food availability. Similar results were obtained when examining chick development in relation to the immunological and serological variables. However, there was a negative relationship between parental feeding rates and food availability, as well as between parental feeding rates and chick development. We conclude that body condition and immune capacity of nestlings are strongly correlated with the availability of food around the nest site.     

Opposing selective pressures on hatching asynchrony: egg viability, brood reduction, and nestling growth

At least 19 hypotheses have been proposed to explain the evolutionary significance of avian hatching asynchrony, and hatching patterns have been suggested to be the result of several simultaneous selective pressures. Hatching asynchrony was experimentally modified in the black kite Milvus migrans by manipulating the onset of incubation during the laying period. Delayed onset of incubation reduced egg viability of first-laid eggs, especially when ambient temperature during the laying period was high. Brood reduction (nestling mortality by starvation or siblicide) was more commonly observed in asynchronous nests. The growth rate was slower in synchronous broods, probably due to stronger sibling rivalry in broods with high size symmetry. Last-hatched chicks in synchronous broods fledged at a small size/mass, while in control broods, hatching order affected growth rates, but not final size. Brood reduction, variable growth rates, and the ability to face long periods of food scarcity are probably mechanisms to adjust productivity to stochastic food availability in a highly opportunistic predator. The natural pattern of hatching asynchrony may be the consequence of opposing selective forces. Extreme hatching synchrony is associated with slow growth rates, small final size of last-hatched chicks, and low viability of first-laid eggs, while extreme hatching asynchrony is associated with high mortality rates. Females seem to facultatively manipulate the degree of hatching asynchrony according to those pressures, because hatching asynchrony of control clutches was positively correlated with temperature during laying, and negatively correlated with the rate of rabbit consumption. Received: 25 October 1999 / Revised: 30 May 2000 / Accepted: 25 June 2000     

Effects of elevated yolk testosterone levels on survival, growth and immunity of male and female yellow-legged gull chicks

Androgen hormones of maternal origin contained in the eggs of avian species are considered to have positive effects on offspring characteristics and performance. However, negative consequences have also been reported, suggesting that mothers may experience a trade-off between beneficial and detrimental effects of egg androgens to offspring fitness. We studied the effects of elevated yolk testosterone (T) concentration on survival, development and phenotype of male and female yellow-legged gull (Larus michahellis) chicks by injecting egg yolks with physiological doses of the hormone. Elevated yolk T resulted in a male-biased post-hatching sex ratio, T-treated clutches producing a greater proportion of males compared to control ones at day 4 post-hatching, likely resulting from a reduction of female embryonic survival, whereas no effect of hormone treatment on hatching success or short-term chick survival was observed. In addition, T depressed post-hatching body mass in both sexes but had no effects on the intensity of the cell-mediated immune response or skeletal growth. No sex differences in egg characteristics or chick phenotype were detected. Time to hatching was not affected by T, but females originating from first laid eggs hatched earlier than males of the same laying order, independently of hormone treatment. However, the implications of sex differences in hatching times are unclear in the study species. Taken together, our results suggest that female yellow-legged gulls may be constrained in transferring androgens to their eggs by negative consequences on the viability of female offspring and growth of chicks of the two sexes.     

Food delivery and sibling competition in experimentally even-aged broods of the cattle egret

Summary The age of nestlings was made even in 12 nests of cattle egrets (Bubulcus ibis) by exchanging young chicks or eggs in a breeding colory in Japan. The growth of chicks in such synchronously hatching broods (SHBs) grew almost as fast as first-hatched chicks in 38 control (asynchronously hatching) broods (AHBs). Four last-hatched chicks in AHBs but no chicks in SHBs died of starvation. The frequency of parental nest-visits with food, that of begging food by chicks, and food mass eaten by chicks were greater in SHBs than in AHBs during the first half of the nestling period and similar thereafter. Dominance rank within each SHB was formed through ritualistic fights among siblings. It was correlated with neither growth rates nor winning ratios in food contests, but in some SHBs the subordinate chicks were attacked more frequently by dominant siblings during food contests than in AHBs. The most subordinate chick in one SHB died as a direct result of such attacks. Sibling aggressions were more frequent in SHBs than in AHBs.     

Inter-sexual differences in T-cell-mediated immunity of black-headed gull chicks (<Emphasis Type="Italic">Larus ridibundus</Emphasis>) depend on the hatching order

Hatching asynchrony in avian species leads to age and size differences between nestlings within a brood, handicapping last-hatched chicks in the sibling rivalry. Starvation due to this competitive disadvantage has been regarded as the primary cause of an increase in mortality with hatching order. However, for gulls it has also been suggested that disease is the cause of mortality for last-hatched chicks, possibly through reduced immunocompetence and thereby an enhanced susceptibility to infection. In addition, the male-biased mortality reported for several gull species may be related to a higher vulnerability to diseases in males compared to females. To determine the potential influence of the immune system on these mortality patterns, we investigated the T-cell-mediated immunity (CMI) of black-headed gull chicks in relation to hatching order and sex. We found a significant decrease in the CMI with hatching order. This result may be causally related to systematic changes in maternal yolk steroids and carotenoids within the laying sequence. For second-laid eggs, male CMI was significantly lower than female CMI. This is possibly linked to higher plasma levels of testosterone in male embryos which might have an immunosuppressive effect. If so, this effect is masked in eggs of either high (first egg) or low (last egg) quality. Chicks with low CMI showed enhanced mortality rates. Thus the differences in immune response are likely to contribute to the observed mortality patterns. However, hatching order significantly affected mortality independently of CMI, suggesting that competitive disadvantage due to hatching asynchrony is also important.Communicated by M. Webster     

Influences of sex,sex composition of brood and hatching order on mass growth in common terns<Emphasis Type="Italic"> Sterna hirundo</Emphasis>

We studied sex-dependent mass growth of chicks of the monomorphic common tern Sterna hirundo which fledged at a colony site in Wilhelmshaven, Germany, between 1997 and 1999. Brood size, brood mates' sex and hatching order (a-, b- and c-chicks) were known for many individuals, which were sexed by molecular techniques. Daily growth rates and age of fledging were independent of sex. However, in all years male chicks reached higher peak mass and fledged up to 5.2% heavier than female chicks. Broods with at least two fledglings showed that besides sex, brood size interacting with sex composition affected chick mass. In mixed broods, brothers had higher peak and pre-fledging mass than sisters they were reared with. Especially in the combination a-daughter and b-son the brothers were heavier. Lowest mass was found in broods with three nest mates of the same sex. A detailed study of 24 three-fledgling broods showed that male c-chicks were heavier than their siblings. The results reveal an advantage for chicks in mixed broods, especially for sons, and more especially if the son was a c-chick. Higher mass and possibly dominance of sons in c-position might be related to higher maternal androgen levels, which are known to increase with each egg laid. The results suggest that even in a monomorphic species, sons might be more expensive to rear, and are discussed with respect to sibling competition, parental effort, survival of sons, as well as to fitness benefits favoring parents producing sons.     

Brood reduction in the American white pelican (Pelecanus erythrohynchos)

Summary American white pelicans (Pelecanus erythrorhynchos) breeding in colonies at East Shoal Lake, Manitoba, Canada exhibited a mean hatching asynchrony of 2.5 days in 2-egg clutches. This resulted in a size difference between chicks which facilitated sibling dominance, harassment and lack of food for the subordinate chick. Only one young survived per nest. In marked broods, the secondhatched chick survived in 20% of successful nests. Manipulated clutch sizes (1, 2 and 3 eggs or chicks per nest) revealed that the presence of a second chick contributes significantly to the reproductive success of the parents. Results support the hypothesis that the second egg functions as a form of insurance against early loss of the first egg or chick. The parents, by establishing hatching asynchrony, by nonintervention in sibling aggression, and by selectively feeding the dominant chick, maximize their chance of rearing the most viable young.     

Seasonal decline of offspring quality in the European starling Sturnus vulgaris: an immune challenge experiment

In seasonally fluctuating environments, timing of reproduction is a crucial determinant of fitness. Studies of birds show that late breeding attempts generally result in offspring of lower reproductive value, with lower recruitment and long-term survival prospects. Several proximate mechanisms, including a seasonal decline of immune system functioning, may lead to a seasonal decline of offspring fitness. We investigated seasonal variation in offspring quality by subjecting first- and second-brood chicks of a sexually size dimorphic species, the European starling Sturnus vulgaris, to an immune challenge with a bacterial endotoxin (LPS), and evaluated their growth and physiological response in terms of total plasma antioxidant capacity (TAC), concentration of reactive oxygen metabolites and hematocrit. LPS challenge did not affect chick growth or oxidative status. However, hematocrit of second-brood chicks was higher in LPS chicks compared to controls. Body mass halfway through the rearing period (days?8–9 post-hatching), TAC and hematocrit were lower among second- vs. first-brood chicks. Interestingly, sexual dimorphism in body mass at days?8–9 post-hatching markedly differed between broods, first-brood males being 4.7% and second-brood males 22.7% heavier than their sisters, respectively. Pre-fledging mortality occurred among second-brood chicks only and was strongly female-biased. Our findings suggest that starling chicks, even if in poor conditions, are little affected by a bacterial challenge, at least in the short-term. Moreover, our study indicates that sex differences in body size, possibly mediated by sex-specific maternal investment in egg size, may heavily impact on pre-fledging survival in a different way in the course of the breeding season, resulting in sex-specific seasonal decline of offspring fitness. Finally, we suggest that levels of circulating antioxidants should be regarded among the proximate causes of the association between timing of fledging and long-term survival in avian species.     

Is asynchronous hatching adaptive in herring gulls (Larus argentatus)?

Hatching asynchrony commonly induces a size hierarchy among siblings and the resultant competition for food between siblings can often lead to starvation of the smallest chicks within a brood. We created herring gull (Larus argentatus) broods with varying degrees of hatching synchrony by manipulating the timing of incubation while maintaining the originally laid eggs. The degree of hatching asynchrony affected sibling size hierarchy at the time of hatching of the last-hatched ”c-chick.” In unmanipulated broods, there was no disadvantage of being a c-chick. However, when asynchrony was experimentally increased, we found reduced survival of the c-chick only in the exaggerated asynchronous experimental group. The effects were observable only during the first 10 days of chick life. We recorded no cases of the chicks dying of starvation. Furthermore, behavioral observations indicated that there was no sibling competition, and no selective feeding of larger sibs in the study colony. We propose that the observed lower survival rates of c-chicks in exaggerated asynchronous broods resulted from their lesser motor abilities, affecting their chances of escaping predators. Fledging success for the whole colony was generally high and almost half of all pairs fledged all three chicks, which is indicative of a good feeding environment. We argue that normal hatching asynchrony is a favorable solution in a good feeding environment, but that increased asynchrony reduces breeding success. We do not view asynchrony in the herring gull as an adaptation for brood reduction and propose instead that it may come about because there has been selection for incubation to start before clutch completion. Received: 14 April 1999 / Received in revised form: 20 October 1999 / Accepted: 23 January 2000     

Cuckoo growth performance in parasitized and unused hosts: not only host size matters

The quality and quantity of food delivered to young are among the major determinants of fitness. A parental provisioning capacity is known to increase with body size. Therefore, brood parasitism provides an opportunity to test the effects of varying provisioning abilities of different-sized hosts on parasitic chick growth and fledging success. Knowledge of growth patterns of common cuckoo, Cuculus canorus, chicks in nests of common hosts is very poor. Moreover, no study to date has focused on any currently unused hosts (i.e., suitable cuckoo host species in which parasitism is currently rare or absent). Here, I compare the growth performance of cuckoo chicks in nests of a common host (the reed warbler, Acrocephalus scirpaceus) and two unparasitized hosts (the song thrush, Turdus philomelos, and the blackbird, Turdus merula). Parasitic chicks were sole occupants of the observed nests, thus eliminating the confounding effect of competition with host chicks. Experiments revealed striking differences in parasitic chick growth in the two closely related Turdus hosts. Cuckoo chicks cross-fostered to song thrush nests grew much quicker and attained much higher mass at fledging than those in nests of their common reed warbler host. Alternatively, parasitic chicks in blackbird nests grew poorly and did not survive until fledging. I discuss these observations with respect to host selection by parasitic cuckoos.     

Provisioning rules and chick competition in asynchronously hatching common terns (Sterna hirundo)

Interactions between nestling birds and their parents are models for examining parent–offspring communication and sibling competition. Most studies have focused on species where young are restricted to a nest. However, offspring of many species are mobile and fed by parents for an extended period post-hatch. These chicks mobility may provide an opportunity to examine the role of signalling and physical competition on parental feeding decisions. We examined parental provisioning rules in relation to offspring behaviour and hatching order (i.e., competitive ability) in a species with mobile young, the common tern. We determined that about 95% of feedings were directed to the first chick to reach the parent when it landed with food. We developed a probabilistic model to predict the likelihood of a chick reaching the parent first, and thus receiving food. Our model showed that begging intensity, feeding history, and the interaction between begging intensity and relative proximity to the parent best predicted which chick would arrive first. Increased begging was associated with arriving first significantly more when a chick was relatively further from the parent than when it was closer than its siblings. Independently of these factors, larger, earlier-hatched chicks were more likely to be fed than smaller, later-hatched chicks. Additional analyses showed that parents landed closer to more intensively begging chicks, however, increased begging did not explain the advantage of earlier-hatched chicks because begging intensity did not vary with hatching order. Instead, earlier-hatched chicks were more likely to outrun later-hatched siblings and reach the parent first.     

Causes of couspecific nest parasitism in the northern masked weaver

Summary Conspecific nest parasitism is a common reproductive strategy in the northern masked weaver (Ploceus taeniopterus). Parasites appear to be females with nests of their own who lay an additional egg in another female's nest as a way to enhance their reproductive success. Brood size in the northern masked weaver is practically constrained to three: starvation in four-chick broods is very common. The constraint on brood size is probably imposed by the extreme hatching asynchrony characteristic of this species: the last egg in a four-egg clutch hatches 72 h after the first egg, and when a chick starves, it is almost always the youngest chick. Late-hatching chicks also grow more slowly than do early-hatching chicks. If a female were to lay a fourth egg in her own nest, there is little chance that it would succeed. However; if she places it in another female's nest before that host lays her third egg, then it may have a greater chance of success.     

Tests of association between the humoral immune response of red-winged blackbirds (<Emphasis Type="Italic">Agelaius phoeniceus</Emphasis>) and male plumage,testosterone, or reproductive success

We explored the potential relationships between male traits, reproductive success, testosterone levels in the breeding season, and humoral immunocompetence in male red-winged blackbirds (Agelaius phoeniceus). Males responded to a single injection of diphtheria-tetanus vaccine by producing antibodies to both antigens. The primary responses to each antigen were positively correlated with each other, but the relationship was clearly non-linear. No male responded strongly to diphtheria without also responding strongly to tetanus, but many males had strong responses to tetanus and weak responses to diphtheria. Response to tetanus was positively associated with hormone levels, whereas the relationship between testosterone and the response to diphtheria was weakly and nonsignificantly negative. We found no convincing relationships between immune responses and male morphological traits (epaulet size, body blackness, and body size), male success in male-male competition (territory size and tenure), or male reproductive success (number of mates, average clutch size, proportion of offspring sired, or proportion of young fledged). These results do not support testosterone acting as a constraint on immunity. Testosterone could be mediating a condition-dependent trade-off between sexual ornaments and the immune system, but the evidence for this was weak, as neither any male sexual trait nor any measure of performance was associated with immune responses. We could not eliminate the possibility that male sexual traits advertise other aspects of immunity that are not dependent upon condition or hormones.     

Sibling aggression, hatching asynchrony, and nestling mortality in the black kite (Milvus migrans)

In siblicidal species, hatching asynchrony could act to reduce sibling rivalry or promote the death of last-hatched chicks. The pattern of hatching asynchrony was experimentally altered in the black kite Milvus migrans. Hatching asynchrony in control broods was intermediate between those of experimentally synchronised and asynchronised broods. Sibling aggression and wounds on the chicks were more commonly observed early in the nestling period and in synchronous nests. Serious injuries were observed on last-hatched chicks in asynchronous nests, as were observations of intimidated or crushed chicks. Sibling aggression was related to food abundance, but some chicks died at an early age in nests with abundant food (cainism). Cainism was more commonly found in asynchronous nests. For species with facultative siblicide, moderate hatching asynchrony could be a compromise between reducing sibling rivalry and avoiding large size differences between sibs that would result in cainism. Female black kites preferentially fed the smallest chicks and exhibited behaviours to reduce sibling aggression, contrary to observations in other siblicidal species. In a highly opportunistic forager such as the black kite, a strategy may exist to protract the life of all the chicks in the brood, waiting for unpredictable situations of food overabundance. This would induce the appearance of a parent-offspring conflict over brood reduction, reflected in the existence of a possible anticipated response by some of the chicks (cainism) and in the appearance of special behaviours by the parents to selectively feed smaller chicks or reduce sibling aggression. In this facultatively siblicidal species, cainism does not seem to be the final stage of an evolutionary trend favouring the raising of high-quality chicks, but a manifestation of a parent-offspring conflict over brood size. Received: 9 March 1998 / Accepted after revision: 8 August 1998     

Adjustment of parental effort in the puffin; the roles of adult body condition and chick size

 We examined the adjustment of parental effort of puffins by switching 20-day-old chicks randomly between parents of known body condition. Among unmanipulated birds mass gain (5–20 days) and mass of 20-day-old chicks was positively correlated with the body condition of parents at day 6. During the first 5 days after chick switching 28% (n = 55) of the parents deserted their foster chick. Parents which deserted their foster chick originally had a chick of their own that was smaller than that of those which did not desert their foster chick. Whether parents deserted their foster chick was also negatively related to the size of the foster chick. The mass of the foster chick was more important than the size of the parents' own chick in determining the desertion rate of chicks. The mass gain of the foster chick during the first 5 days after switching was positively related to the body condition of foster parents and also positively related to the mass of the foster parents' own chick, but negatively related to the size of the foster chick. The results suggest that puffins adjust their parental effort according to both their own body condition and the size of the chick. The latter may indicate the chick's prospect of survival and recruitment to the population. Received: 20 January 1996 / Accepted after revision: 27 October 1996     

The role of helpers in feeding chicks in cooperatively breeding green (red-billed) woodhoopoes

Summary Observations were made of ten green (red-billed) woodhoopoe Phoeniculus purpureus flocks during the breeding season in order to quantify the relationship between flock size and the amount of food delivered to chicks. The study period was kept short specifically to minimize the effects of environmental stochasticity. Neither woodhoopoe feeding visit rates nor the total amount of food brought to chicks increased with flock size. Although nonbreeders did not increase the net rate of food provisioning to chicks, they reduced parental input in chick rearing, and hence energy expenditure by the breeding pair. However, over an 8-year study period, which includes data for 144 flock years, this did not result in increased breeding frequency or enhanced survival of breeders. There is thus no evidence that helpers' feeding contributions to young per se influence the indirect fitness of helpers.