Brood Desertion Research Articles

Sex ratio and sex-specific chick mortality in a species with moderate sexual size dimorphism and female brood desertion

An equal sex ratio at the population level is the usual, evolutionarily stable condition. However, at the individual level, it may be adaptive for parents to manipulate the sex of their offspring, especially in species with sexual size dimorphism (SSD) when the costs and benefits of producing sons and daughters can vary. In this study, we investigated the hatching sex ratio (HSR) and fledging sex ratio (FSR) in the Whiskered Tern (Chlidonias hybrida). Despite the fact that SSD exists in Whiskered Terns already at the chick stage, HSR and FSR did not deviate from parity at the population level. We tested the dependence of HSR, FSR and the survival probability of males and females on the individual hatching date, average egg volume (in a clutch) and the number of nestlings. None of these factors influenced HSR. Survival probability was negatively correlated with the number of hatchlings. The proportion of females among the fledglings was positively correlated with the average egg volume per clutch. To better explore the effect of egg volume on the sex ratio, we tested the relationship between exact egg volume and hatchling sex or hatching success at the individual level; but despite the quite large sample size, our analyses failed to reveal any relationship. The sex ratio was equal among chicks that were found dead in a nest, mostly due to starvation, but more female than male chicks disappeared from nests (mostly due to predation), primarily in the first week of life. This indicates that females may be easier to predate, very likely by frogs hunting small chicks.

Eyes on the future: buffering increased costs of incubation by abandoning offspring

AbstractLife history theory states that the resources invested in current reproduction must be traded off against resources needed for survival and future reproduction. Long-lived organisms have a higher residual reproductive value and are therefore expected to be sensitive to reproductive investments that may reduce survival and future reproduction. Individuals within a population may vary in phenotypic quality, experience, access to resources etc. This may affect their optimal reproductive investment level. In this study we manipulated reproductive costs by shortening and extending the incubation period in common eiders Somateria mollissima without altering clutch size. Females whose incubation time was prolonged experimentally, suffered higher mass loss and increased clutch loss/nest desertion. These females were also more prone to abandon their brood after hatching. Both clutch loss and brood abandonment decreased with clutch size in all treatment categories, indicating higher phenotypic quality and/or better access to resources for females producing more eggs. However, although females with prolonged incubation were lighter at hatching, their return rate and breeding performance in the following year were unaffected. These results show that individual quality as expressed through clutch size and body mass is affecting current reproductive investment level as well as future survival and breeding performance. The results also show that individual birds are sensitive to changes in their own condition, and when reproductive effort is approaching a level where survival or future survival may be compromised, they respond by terminating their current reproductive attempt.

Interspecific differences in brood desertion by female diving ducks in relation to duckling age and environmental conditions

Studies on brood desertion in birds have been mainly conducted on species with biparental care, and less often on uniparental species. Females of many duck species remain with their ducklings, unassisted by males, during variable periods of time before the chicks can fledge. In this paper, we examined factors that influence brood desertion by female red-crested pochards ( Netta rufina), common pochards ( Aythya ferina) and white-headed ducks ( Oxyura leucocephala) during a 10-year period in lakes in southern Spain that experienced interannual variations in water levels. At an interspecific level, we found that brood desertion was less frequent in the species that nested earlier (red-crested pochard), while it was more frequent in the species that nested later (white-headed duck). Larger (older) ducklings were deserted more frequently than smaller ones in the three species. However, an interaction between flooding conditions in wetlands and species identity on brood desertion, after accounting for chick age, suggests that there were variations between the ducklings of the three species in their requirements for maternal care, conditioned by environmental conditions. Therefore, our study indicates that environmental conditions may affect parental care. A longer duration of brood attendance by female ducks may buffer against adverse effects of environmental conditions.

Influence of researcher experience and fieldwork intensity on the probability of brood losses in sensitive species: The case of the Red-backed Shrike Lanius collurio

Minimizing disturbance to animals when studying them should be a major consideration in ecological research design, especially when species are known to be sensitive to disturbances. A basic method in the study of bird reproductive ecology involves nest visits, but these can adversely affect birds’ breeding success. Appropriate fieldwork procedures are therefore highly important, especially in the case of timid and sensitive species. This study aimed to assess the impact of the method of searching for nests of Red-backed Shrike Lanius collurio on the probability of these birds abandoning their broods. Our study revealed that observer experience and nest searching intensity did influence the likelihood of brood desertion by this species. Less field experience and a greater nest searching intensity caused the shrikes to abandon their nests more often. We recommend paying scrupulous attention to the behaviour of the birds when searching for nests and when looking for nests after incubation has started.

Clutch Abandoning Parasitised Yellow Warblers Have Increased Circulating Corticosterone With No Effect of Past Corticosterone or Differences in Egg Maculation Characteristics

Parental care can be costly to parents’ fitness. As such, abandonment of the current reproductive attempt may benefit potential future opportunities, maximising lifetime reproductive success. Obligate brood parasitism, a reproductive strategy in which parasites lay their eggs in the nests of other species and rely solely on them to raise the parasitic young, is an ideal system to study brood abandonment. Some parasitised host species have evolved anti-parasitic defences, notably clutch abandonment (egg burial and nest desertion), that may mitigate negative consequences of parasitism. Abandonment of clutches due to parasitism is not unlike abandonment of reproduction in times of stress, suggesting that host responses to parasitism could be triggered at least partly by elevated stress hormones that mediate individual decisions. Yet, the mechanistic basis for clutch abandonment remains unclear. Here, we experimentally parasitised clutches of yellow warblers (Setophaga petechia), a common host of the brown-headed cowbird (Molothrus ater), with model cowbird eggs to examine whether host circulating corticosterone (CORT) differed among females that accepted parasitic eggs or rejected them through clutch abandonment. We also assessed whether feather CORT, a measure of past corticosterone exposure, differed between accepters and abandoners. Finally, we investigated whether egg visual signals, specifically differences in maculation characteristics between model cowbird and host eggs, predicted abandonment of experimentally parasitised clutches. Circulating CORT was higher in females who abandoned their parasitised clutches, but not in those who accepted, relative to controls with no egg addition. Past stress and differences in maculation characteristics did not predict whether individuals accepted or abandoned experimentally parasitised clutches. Moreover, differences in maculation characteristics between the host and model cowbird eggs did not predict CORT levels or nest abandonment. Thus, parasitism with subsequent clutch abandonment may be associated with elevated circulating CORT, but neither past stress nor differences in maculation characteristics influenced abandonment. The combination of these results contributes to our understanding of the roles of corticosterone and egg visual signals in the context of clutch abandonment in brood parasitism specifically, and of parental care more broadly.

Extra-pair paternity in a species with frequent extra-pair courtship feedings, few extra-pair copulations, and male-biased parental care

Patterns of extra pair paternity (EPP) and intraspecific brood parasitism (IBP) were studied in a Whiskered Tern Chlidonias hybrida population characterized by female brood desertion, a high level of extra pair courtship feedings (10%) and a low level of extra pair copulation (EPC; 0.6%). In this study, we used a set of microsatellite loci to analyse parentage in 56 Whiskered Tern families from southern Poland. Depending on the method, we detected that 1.4%–3.6% of chicks were sired by an extra-pair male, and extra-pair chicks were present in 3.6%–8.9% of broods. IBP was observed in 8.9%–14.3% of broods, corresponding to 3.6%–6.4% of the chicks. The low rate of EPP is in agreement with the hypothesis that in species with high male parental investment, females should avoid EPC. The low level of IBP indicates that intraspecific egg dumping is not a common female strategy in Whiskered Tern, similar to many other colonial waterbird species.

Baseline and stress-induced prolactin and corticosterone concentrations in a species with female offspring desertion – The case of Whiskered Tern Chlidonias hybrida

To understand the proximate mechanisms regulating brood desertion, we studied hormonal and behavioural stress responses during the chick-rearing period in adult Whiskered Terns (Chlidonias hybrida), a socially monogamous, semi-precocial species with prolonged post-fledging parental care. In contrast to males, almost all females of this species desert during the chick-rearing and post-fledging periods. Because of the expected link between corticosterone, prolactin and parental investment, we hypothesized that males and females should differ in circulating prolactin and corticosterone concentrations. Baseline hormone concentrations did not differ between males and females. In both sexes, prolactin and corticosterone concentrations decreased and increased in response to acute stress (30 min after capture), respectively. Baseline and stress-induced prolactin concentrations decreased significantly in both sexes with advancing brood age. As expected, males had significantly higher stress-induced prolactin concentrations than females. None of the nine males released after being held in captivity for 24 h deserted, whereas four (29%) of the 14 females kept in captivity for 24 h did so. Altogether, these results suggest that higher prolactin concentrations may be involved in the maintenance of parental care under stress. However, there was no statistically significant difference in stress-induced hormone levels between males, females that deserted and those that returned to the nest after prolonged stress (24 h). Our data indicate that males are probably more resistant to stress as regards the continuation of parental care. The pattern of male and female behavioural and hormonal responses to stress partially predicts their behaviour in terms of natural desertion.

Sex-specific growth of nestlings of the Whiskered Tern Chlidonias hybrida, a species with sexual size dimorphism and female brood desertion

The growth of a nestling during the first few weeks after hatching is crucial for its further life and is sensitive to the conditions experienced during this period. Among species exhibiting Sexual Size Dimorphism (SSD), one of the most important factors influencing growth parameters is the sex of the hatchlings. In this study, we tested whether sex and other factors (hatching date, egg volume, brood size and sex ratio in the brood) were related to sex on growth parameters in Whiskered Tern (Chlidonias hybrida) chicks. In this species, males are 3%–10% larger than females when adult. At hatching, only the total head length was greater in males than females, but values close to fledging were greater for most of the measured parameters (3%—tarsus, 4%—total head length, 13%—body mass). Moreover, the maximum body mass growth rate was higher in males but females entered the rapid growth phase somewhat earlier and achieved the final total head length sooner. Nestling growth was also significantly correlated with brood size and hatching date, though mostly negatively (e.g. lower tarsus and total head length asymptote, lower maximum growth rate and growth rate constant of total head length in bigger and later clutches). The occurrence of SSD during the nestling period and higher maximum growth rate of body mass in males indicates that the costs of raising sons may be higher than of raising daughters. However, we did not find evidence of either sex-biased nestling mortality or skewed sex ratio.

Duration of female parental care and their survival in the little auk Alle alle - are these two traits linked?

Desertion of offspring before its independence by one of the parents is observed in a number of avian species with bi-parental care but reasons for this strategy are not fully understood. This behaviour is particularly intriguing in species where bi-parental care is crucial to raise the brood successfully. Here, we focus on the little auk, Alle alle, a small seabird with intensive bi-parental care, where the female deserts the brood at the end of the chick rearing period. The little auk example is interesting as most hypotheses to explain desertion of the brood by females (e.g. “re-mating hypothesis”, “body condition hypothesis”) have been rejected for this species. Here, we analysed a possible relationship between the duration of female parental care over the chick and her chances to survive to the next breeding season. We performed the study in two breeding colonies on Spitsbergen with different foraging conditions – more favourable in Hornsund and less favourable in Magdalenefjorden. We predicted that in Hornsund females would stay for shorter periods of time with the brood and would have higher survival rates in comparison with birds from Magdalenefjorden. We found that indeed in less favourable conditions of Magdalenefjorden, females stay longer with the brood than in the more favourable conditions of Hornsund. Moreover, female survival was negatively affected by the length of stay in the brood. Nevertheless, duration of female parental care over the chick was not related to their parental efforts, earlier in the chick rearing period, and survival of males and females was similar. Thus, although females brood desertion and winter survival are linked, the relationship is not straightforward.Significance statementWhen bi-parental care is crucial to raise the brood successfully, one parent desertion raises the question of why this happens. We examined this issue in the little auk, a small seabird with females deserting the brood at the end of the chick rearing period. We hypothesised that females deserting the brood save residual energy and, in this way, increases their chance to survive to the next breeding season. We found that duration of female parental care depends on environmental conditions, with longer staying with the brood in less favourable conditions. As expected, female survival decreased with duration of their staying with the brood but it was not related to their parental efforts (i.e. number of chick feedings, duration of foraging flights). In addition, survival of males and females was similar. Thus, although little auk females brood desertion and winter survival are linked, the relationship is not straightforward.

Pair coordination is related to later brood desertion in a provisioning songbird

Raising a family is not free of conflict for parents, as each parent benefits when its partner provides more care for the offspring. Resolving this conflict requires cooperation between the parents. One way to achieve such cooperation might be to coordinate parental provisioning by synchronizing (i.e. returning to the nest at the same time) or alternating (i.e. taking turns) offspring provisioning at the nest. Empirical studies in birds indicate that pair coordination of the nest visits is common; however, it is unknown whether this behaviour is directly related to different outcomes of sexual conflict, such as brood desertion. We used the rock sparrow, Petronia petronia, a species with high levels of sexual conflict, to explore whether alternation and synchrony of the nest visits were related to later brood desertion. Pairs with no desertion alternated and synchronized their nest visits more than pairs in which one sex deserted. This difference in coordination was not simply a by-product of differences in provisioning by the parents. Synchrony of the visits also increased with offspring age in the pairs with no desertion. We provide evidence, for the first time to our knowledge, that the degree of parental coordination is strongly associated with the ultimate consequence of sexual conflict, brood desertion, supporting the idea that coordination in parental behaviour might promote conflict resolution.

Parental feeding responses to experimental short-term partner removal in a species with male and female brood desertion

As parental care is costly, it can be expected that there will be a sexual conflict between parents over the individual levels of parental investment because each parent has limited resources to invest in a reproductive event. Theoretical models of parental investment predict that when one parent reduces its parental effort, the other parent should adjust its effort facultatively to compensate for the decrease in the mate’s contribution. Here, we tested for facultative adjustments in care in the rock sparrow, Petronia petronia. In this species, both sexes can desert the brood, creating the potential for strong sexual conflict over parental care to occur. To that end, we examined how rock sparrow parents adjust their level of care in response to an experimental mate removal for a limited time period, mimicking the starting phase of the desertion process. We compared male and female provisioning rates before and after an experimental mate removal. Males behaved according to the prediction of compensatory adjustment, as they allocated more care to the offspring and returned faster to the nest after female removal, whereas females showed no response. Our study shows sexual differences in response to mate removal in a species with biparental care and strong sexual conflict over parental care, and suggests that males may use the female absence to determine their actual parental effort.Significance statementStrong differences in parental care, with one sex providing more care than the other, are widespread across the animal kingdom. Theoretical models of parental investment predict that when one parent reduces its care, the mate should adjust its care facultatively to compensate. Here, we tested for facultative adjustments in care in the rock sparrow when their mates were experimentally removed for a limited time period. We show that only male behavior was consistent with theoretical predictions as they increased their frequency of provisioning significantly after mate removal, whereas females showed no response. Our study suggests that members of a pair do not make independent decisions, but negotiate to modify their effort in direct response to the prior effort of their mate.

Evaluating the Efficacy of Brood Flush Counts: A Case Study in Two Quail Species

Abstract. Taking brood flush counts is a common sampling method that has been used for decades to estimate brood and chick survival in many gallinaceous bird species. However, brood survival estimates based upon flush counts may be biased because of low detection probabilities, occurrence of brood amalgamations, brood abandonment, and brooding adult mortality. Given that brood flush counts are still commonly used to estimate brood survival, and in some cases extrapolated to provide an estimate of chick survival, it is important to evaluate biases associated with this method. Therefore, we evaluated the use of brood flush counts to estimate brood survival of 2 gallinaceous birds: Northern Bobwhite (Colinus virginianus) and Scaled Quail (Callipepla squamata). To gain a better understanding of some of the mechanisms that may bias brood survival estimates, we radio-tagged Northern Bobwhite and Scaled Quail chicks (8–12 days old) and juveniles (4–6 weeks old) in 67 broods. We used radio-tagged chicks and juven...

An experimental test on time constraint and sexual conflict over parental care.

Because parental care is costly, a sexual conflict between parents over parental investment is expected to arise. Parental care behavior is an adaptive decision, involving trade-offs between remating, and consequently desertion of the brood, and continuing parental effort. If the main advantage of desertion is remating, then this will be a time constraint, because the deserting individual will require a certain minimum period of time to breed again in the same breeding season. So, a short breeding season should force certain individuals to desert the first brood to have enough time to successfully complete their second breeding attempt. The rock sparrow, Petronia petronia, is an unusual species in which brood desertion can occur in both sexes and the breeding season is quite short so it is a good species to investigate the role of time constraint on brood desertion. For 3 years, I investigated the brood desertion modality of the rock sparrow. Then, for 2 years, I removed a group of experimental nest boxes during the autumn. Later, I re-installed the experimental nest boxes after the start of the breeding season (2 weeks after the first egg was laid), mimicking a shortening of the breeding season for the (experimental) pairs that used experimental nest boxes. I found that in the experimental pairs, the percentage of deserting individuals was significantly higher than in the control groups, and the deserting individuals were older females. This experiment adds to our knowledge of timing of reproduction effects on individual decisions to desert by showing that a short and delayed breeding season may have different effects on males and females. To my knowledge, this is the first experimental study that demonstrates a direct link between time constraint and brood desertion.

Multiple Lines of Egg Defense in a Neotropical Arachnid with Temporary Brood Desertion

AbstractEgg predation is the one of the main costs of brood desertion in many ectothermic animals. When stressful environmental conditions constrain parental activities to only some periods of the day, the combination of physical or chemical defenses may attenuate the costs related to egg loss during periods of temporary parental absence. Females of the harvestman Neosadocus maximus periodically abandon their clutches to shelter or forage. They also cover their eggs with a hygroscopic mucus coat and seem to lose fewer eggs to predation than other syntopic harvestmen whose eggs lack the mucus coat. Using two species of N. maximus egg predators, we demonstrate that eggs whose mucus coat was experimentally removed suffered higher predation rate than eggs whose mucus coat was left intact. We argue that this mucus provides physical protection against egg predators, especially small arthropods. A similar mucus coat has independently evolved in other two clades of Neotropical harvestman in which males care for the eggs and typically leave their clutches unattended for several hours a day. We propose that the presence of multiple lines of egg defense may have evolved as a way of lowering the costs imposed by intra‐ and interspecific egg predation during periods of temporary brood desertion.

The influence of brood loss on nest abandonment decisions in largemouth bass Micropterus salmoides.

Largemouth bass Micropterus salmoides broods were experimentally reduced in size to test whether brood size (BS) and simulated brood depredation affect the decision by a male to continue providing care for its brood or to abandon that brood prematurely before its offspring reach independence. The highest ranked of the generalized linear models predicting brood abandonment was based on the number of offspring remaining in a nest following brood devaluation, indicating that parental male fish reassess the value of a brood following perturbation. Paternal M. salmoides were more likely to abandon their broods if initial BS was small before devaluation, and if there was a greater decrease in BS, indicating a threshold for both the amount of brood loss and remaining BS. Larger, older males were also less likely to abandon their brood than smaller, younger conspecifics. These results have broad implications for determining drivers of parental care trade-offs and how individuals assess the value of a brood.

Behavioural and hormonal stress responses during chick rearing do not predict brood desertion by female in a small Arctic seabird

We examined behavioural and hormonal stress responses in a small seabird (little auk, Alle alle), which exhibits a transition from biparental to male-only care towards the end of the nesting period, in order to understand the mechanisms underlying this parental strategy. We hypothesized that the male staying with the chick should be less sensitive to stressors. As such the male might offer the offspring more efficient protection during the fledging period than the female. We tested this hypothesis by observing male and female behaviour in a neophobia test. We also measured the birds' baseline and stress-induced levels of corticosterone and prolactin using the standardized capture-and-restraint protocol. Both sexes respond rapidly to foreign objects, delaying the entry time to the nest with food, consuming the food load, and/or temporarily abandoning feeding. However, we did not find any differences between the sexes in the frequency of each behaviour or in the time of the first reaction to the experimental treatment. Level of both corticosterone and prolactin increased after the experimental treatment. However, we did not find sex differences in baseline and stress-induced hormone levels. The results indicate that the males are as much sensitive to the stress situation as the females. Thus, the pattern of male and female behavioural and hormonal responses to stress does not predict their behaviour at the final breeding stage.

Seasonal Variation in Offspring Sex Ratio in the Snowy Plover

The Snowy Plover (Charadrius nivosus) is unique in being a determinate layer of an odd modal clutch size and in having a variable mating system in which female brood desertion occurs regularly. These traits make determining Snowy Plover offspring sex ratios important not only for long-term population stability, as the species is of conservation concern, but also for application to sex allocation theory. In this study, we determined Snowy Plover offspring sex ratios, examined differential costs of producing male and female offspring, and evaluated sex ratio variation in relation to maternal condition, habitat condition, and time during the nesting season on saline lakes of the Southern High Plains of Texas. Examination of 245 chicks from 118 clutches during 1999–2000 and 2008–2009 showed that male offspring were more costly to produce than female offspring; however, offspring sex ratio did not differ from parity, but was slightly male-biased in most years. The probability of producing a male offspring was greater both earlier and later in the breeding season than in the middle. As the availability of saline lake surface water and the subsequent availability of food vary unpredictably throughout the breeding season, depending on precipitation events, we suggest that sex ratio adjustment in unpredictable environments may not be straightforward and may follow nonlinear models and/or vary annually The effects such changes in sex ratios may have on population growth and stability remain unknown.

Predator burden and past investment affect brood abandonment decisions in a parental care‐providing teleost

Summary Reproductive success of parental care‐providing species is contingent upon the level of parental investment into a brood. When the expected fitness contribution of a brood is outweighed by the costs of parental care (e.g. decreasing parent somatic condition), an individual may reduce its level of care, or may altogether abandon a brood in trade‐off for potential future reproductive success. Leading parental care hypotheses have established parental stress, oxidative stress, nutrition and androgen condition, as well as reproductive value and the threat of brood depredation, as drivers of investment into parental care. The concomitant effects of parent physiological condition and the threat of brood depredation have yet to be considered in their direct effect on the reproductive success of a parental care‐providing individual. Using largemouth bass (Micropterus salmoides) as a model parental care‐providing species, we investigated the relative influence of these factors, together, on a direct fitness measure: the decision by a parent to abandon its brood. By employing a novel combination of multivariate and information‐theoretic modelling, our findings indicate that reproductive success of parental largemouth bass is contingent primarily on the threat of depredation to a brood (i.e. the density of brood predators adjacent to nest locations), past investment into a brood and reproductive value of the brood. Parent physiological condition had limited influence on the decision by largemouth bass to abandon parental care. Modelling indicated that parental circulating androgen concentration and antioxidant capacity may play a minimal role in driving brood abandonment; nutrition and stress condition of the parent did not differ between reproductive outcomes. Findings here suggest that a holistic approach should be implemented when studying parental care and that a definite experimental endpoint, e.g., brood abandonment or reproductive success, be utilized as a direct metric of the cost to fitness of parental care decisions.

Mating strategies, parental investment and mutual ornamentation in Iberian Rock Sparrows (Petronia petronia)

Relatively few bird species show complex social mating systems whose preponderance in a population is likely to affect the patterns of parental care observed there. In turn, parental investment is likely to be related to the expression of certain ornaments, which may reveal information on the bearer’s individual quality. Here we address both issues in a species characterised by several forms of parental care (both biparental and uniparental care) and in which both sexes possess a yellow breast patch, the rock sparrow (Petronia petronia). In our population, males contributed more to the care of the young in comparison with other populations. Social monogamy was the most frequent mating pattern and the percentage of cases of female (or male) brood desertion was lower with respect to that reported in previous studies, suggesting a flexible behaviour of this species to deal with different social environments. Birds did not pair assortatively with respect to the size of the yellow breast patch and we found no significant relationship between this trait and the frequency with which parents provisioned their chicks. However, we observed a positive relationship between male yellow patch size and nestling tarsus length, which suggests that more ornamented males are better parents. Males, but not females, differentially allocated parental investment in response to female ornamentation, although the benefits that males may gain from choosing more attractive females remain unidentified. Our results on paternal care investment along with previous studies on this species, reinforcing the view that the rock sparrow constitutes a good model to study sexual conflict over parental care under different social environments.

When and why does my mother leave me? The question of brood desertion in the Dovekie (Alle alle)

ABSTRACT. Many avian species exhibit a transition from biparental to uniparental care, but the reasons for this pattern are largely unknown. We examined the transition from biparental to paternal-only care in a small Arctic seabird, the Dovekie (Alle alle). We monitored colony attendance of males and females throughout the chick-rearing period in 2008–2010, investigating the sex ratio of breeders and following individually marked parents of both sexes in the colony at Magdalenefjorden (northwestern Spitsbergen). To understand reasons for the female's behavior, we also examined the birds' parental efforts and changes in chick body mass. We found that females fed the chicks less than males but provisioned the chick until very late in the chick-rearing period. Brood desertion occurred in 95% of nests and took place 3 days, 2 days, or 1 day prior to fledging (in 24%, 39%, and 69% of nests, respectively), on the 26th day of the chick's life, on average. Chicks experienced body-mass recession beginning, on aver...