Growth and its relationship to fledging success of African black oystercatcher Haematopus moquini chicks

We investigated the growth of African black oystercatcher Haematopus moquini chicks on Robben Island, South Africa, over three austral summers, 2001-2004. Using a robust regression analysis to determine the growth parameters of chicks of known and unknown age we found that oystercatchers from our study population had a Gompertz growth rate coefficient that was 2% less than predicted for body mass based on the equation for waders. Leg growth lagged initially, then increased and slowed again as the chicks became older, whereas wing growth was slow initially but increased with age. Chicks with small growth rate coefficients for body mass exhibited retarded growth of all body measures except wing length. This enabled these chicks to fledge in a shorter period of time than their slow growth would otherwise allow. The growth rate of body mass was observed to vary greatly between chicks. Fast-growing African black oystercatchers had a shorter pre-fledging period; were larger at fledging and were more likely to fledge successfully. African black oystercatchers display sibling rivalry, and once a dominance relationship is established, the larger chick remains so during the pre-fledging period. Larger siblings fledged earlier and at a heavier mass than the smaller siblings and this may improve their chances of survival. Neither hatching date nor brood size influenced the growth rate coefficients.     

Behavior of adult and young grassland songbirds at fledging

The behavior of adults and young at the time of fledging is one of the least understood aspects of the breeding ecology of birds. Current hypotheses propose that fledging occurs either as a result of parent‐offspring conflict or nestling choice. We used video recordings to monitor the behavior of nestling and adult grassland songbirds at the time of fledging. We observed 525 nestlings from 166 nests of 15 bird species nesting in grasslands of Alberta, Canada, and Wisconsin, USA. Overall, 78% of nestlings used terrestrial locomotion for fledging and 22% used wing‐assisted locomotion. Species varied in propensity for using wing‐assisted locomotion when fledging, with nestling Grasshopper Sparrows (Ammodramus savannarum) and Henslow's Sparrows (Centronyx henslowii) often doing so (47% of fledgings) and nestling Song Sparrows (Melospiza melodia), Common Yellowthroats (Geothlypis trichas), and Chestnut‐collared Longspurs (Calcarius ornatus) rarely doing so (3.5% of fledgings). For 390 fledging events at 127 nests, camera placement allowed adults near nests to be observed. Of these, most young fledged (81.5%) when no adult was present at nests. Of 72 fledging events that occurred when an adult was either at or approaching a nest, 49 (68.1%) involved feeding. Of those 49 fledgings, 30 (62.1%) occurred when one or more nestlings jumped or ran from nests to be fed as an adult approached nests. The low probability of nestlings fledging while an adult was at nests, and the tendency of young to jump or run from nests when adults did approach nests with food minimize opportunities for parents to withhold food to motivate nestlings to fledge. These results suggest that the nestling choice hypothesis best explains fledging by nestlings of ground‐nesting grassland songbirds, and fledging results in families shifting from being place‐based to being mobile and spatially dispersed.     

Post‐fledging survival of altricial birds: ecological determinants and adaptation

For altricial young, fledging is an abrupt step into an unknown environment. Despite increasing numbers of studies addressing the post‐fledging period, our current knowledge of the causes and consequences of post‐fledging survival remains fragmentary. Here, we review the literature on post‐fledging survival of juvenile altricial birds, addressing the following main questions: Is low post‐fledging survival a bottleneck in the altricial reproductive cycle? What is known of proximate and ultimate causal factors such as trophic relations (food and predation), habitat conditions, or abiotic factors acting in the post‐fledging period? We analyzed weekly survival estimates from 123 data series based on studies of 65 species, covering weeks 1–13 post‐fledging. As a general pattern, survival of fledglings was low during the first week post‐fledging (median rate = 0.83), and improved rapidly with time post‐fledging (week 4 median rate = 0.96). For ground‐nesting species, survival immediately after leaving nests was similar to egg‐to‐fledging survival. For species breeding above‐ground, survival during the first week post‐fledging was substantially lower than during both the nestling period and later post‐fledging stages. Thus, the early post‐fledging period is a bottleneck of markedly elevated mortality for most altricial species. Predation was the main proximate cause of mortality. Various factors such as habitat, annual and seasonal variation in the environment, and the physical condition of fledglings have been found to affect post‐fledging survival. Individual survival depended strongly on physical traits such as mass and wing length, which likely influence the ability of fledglings to escape predation. Trophic relationships at various levels are the main ultimate driver of adaptation of traits relevant to survival during the pre‐ and post‐fledging periods. Spatiotemporal dynamics of food resources determine the physical development of juveniles and, in turn, their performance after fledging. However, predators can cause quick and efficient selection for fledgling traits and adult breeding decisions. Parental strategies related to clutch size and timing of breeding, and the age and developmental stage at which young fledge have substantial effects on post‐fledging survival. The intensity and duration of post‐fledging parental investment also influences fledgling survival. Post‐fledging mortality is therefore not a random and inevitable loss. Traits and strategies related to fledging and the post‐fledging stage create large fitness differentials and, therefore, are integral, yet poorly understood, parts of the altricial reproductive strategy.     

The post‐fledging period in a tropical bird: patterns of parental care and survival

How environmental conditions affect the timing and extent of parental care is a fundamental question in comparative studies of life histories. The post‐fledging period is deemed critical for offspring fitness, yet few studies have examined this period, particularly in tropical birds. Tropical birds are predicted to have extended parental care during the post‐fledging period and this period may be key to understanding geographic variation in avian reproductive strategies. We studied a neotropical passerine, the western slaty‐antshrike Thamnophilus atrinucha, and predicted greater care and higher survival during the post‐fledging period compared to earlier stages. Furthermore, we predicted that duration of post‐fledging parental care and survival would be at the upper end of the distribution for Northern Hemisphere passerines. Correspondingly, we observed that provisioning continued for 6–12 weeks after fledging. In addition, provisioning rate was greater after fledging and offspring survival from fledging to independence was 75%, greater than all estimates from north‐temperate passerines. Intervals between nesting attempts were longer when the first brood produced successful fledglings compared to nests where offspring died either in the nest or upon fledging. Parents delayed initiating second nests after the first successful brood until fledglings were near independence. Our results indicate that parents provide greater care after fledging and this extended care likely increased offspring survival. Moreover, our findings of extended post‐fledging parental care and higher post‐fledging survival compared to Northern Hemisphere species have implications for understanding latitudinal variation in reproductive effort and parental investment strategies.     

Laying date,body mass and tick infestation of nestling tropical Roseate Terns Sterna dougallii predict fledging success,first‐year survival and age at first return to the natal colony

Both intrinsic and extrinsic factors recorded at individual nests can predict offspring fitness and survival but few studies have examined these effects in the tropics. We recorded nestling survival, post‐fledging survival and age at first return of Roseate Terns breeding at Aride Island, Seychelles, over a 12‐year period (1998–2009). Nest data recorded at the egg, nestling and fledging stages were collected during six breeding seasons (1998, 2001–2005) and a capture‐mark‐recapture dataset of six cohorts of fledglings was obtained from 2001–2009. Logistic regression models were used to assess the predictive effect of reproductive variables on fledging success, while multistate capture‐mark‐recapture models were used to estimate post‐fledging survival and return–recruitment probabilities to the natal site. Nestling survival probability increased with earliness of laying and was negatively affected by tick infestation during the growth period (0–23 days). Fledging probability was also positively related to chick body condition, whereas other pre‐fledging reproductive parameters such as clutch size and egg size were not influential. A multistate modelling of age‐specific survival and return–recruitment (transition) rates found that first‐year survival differed between cohorts and was also negatively affected by tick infestation. Annual survival stabilized from age 2 onwards at 0.83 ± 0.02. Transition rates were positively related to body condition at fledging, with heavier individuals returning for the first time to the natal colony at a younger age compared with lighter individuals. These results highlight the importance of local conditions encountered by tropical seabirds during the breeding season in shaping demographic parameters.     

Timing of fledging is influenced by glucocorticoid physiology in Laysan Albatross chicks

Fledging is a major life transition for birds, when juveniles move from the safety of a nest into an environment where they must find food and avoid predators. The timing of fledging within a season can have significant effects on future survival and breeding success. Proximate triggers of fledging are unknown: though wing development is likely a primary factor, other physiological changes, such as elevated plasma corticosterone (CORT), may affect fledging behavior. Laysan Albatross (Phoebastria immutabilis) chicks have an extended post−hatching period during which they reach 150% of adult mass. However, approaching fledging, chicks fast for days to weeks and lose mass while still putting energy into feather growth. We evaluated chick morphology and physiology to elucidate proximate triggers of fledging. As in some other species, CORT increased as chicks fasted and lost body mass. At the same time, corticosteroid binding globulin (CBG) declined, thus amplifying free CORT prior to fledging. Once chicks reached a morphological threshold, free CORT levels predicted how long they stayed at the colony: chicks with higher free CORT fledged sooner. To perturb the relationship between body condition, endocrine physiology, and fledging behavior, we supplementally fed chicks for the month before fledging. Fed birds had a slower decrease in body mass, slower decrease in CBG, slower increase in free CORT, and stayed at the colony longer after reaching a morphological threshold. Our study suggests that as chicks lose mass, free CORT acts as a signal of energetic or nutritional state to adjust the timing of fledging.     

The process of fledging in the Mountain Bluebird

Fledging is a critical event in the avian breeding cycle, but remains unstudied in almost all species. As a result, little is known about factors that cause nestlings to leave nests. We documented fledging behavior in a box‐nesting population of Mountain Bluebirds (Sialia currucoides) using radio‐frequency identification. We attached a passive integrated transponder (PIT tag) to the leg of each nestling in 40 nests. An antenna checked for the presence of a transponder signal (i.e., a nestling) at nest‐box entrances every 2 s. The time of last detection of a nestling was taken as the time that nestling fledged. We found that fledging began when the oldest nestlings were 15–22 d old. Broods that were ahead in development, as measured by primary feather length, fledged at relatively younger ages. All nestlings fledged on the same day at 33 nests (83%) and over 2 d at remaining nests. When all nestlings fledged on the same day, fledging usually began in the morning and median time between the first and last fledging was 55 min (range = 2.3 min–10.6 h). When young fledged over 2 d, fledging always began >8 h after sunrise and usually just one nestling fledged the first day, suggesting that this fledging may have been accidental. Clutches in our population often hatch asynchronously, which sets up a hierarchy within broods in developmental state, size, and competitive ability. In such situations, fledging may be initiated by one of the most‐developed and hence most‐competitive nestlings in a brood, presumably when it reaches a certain threshold state of development. Alternatively, fledging may begin when a less‐developed, less‐competitive, and probably hungrier nestling leaves the nest, presumably to gain better access to food. We used the proportion of time that a nestling was able to occupy the nest‐box entrance late in the nestling stage, waiting to intercept parents with food, as an index of nestling competitive ability. Assuming that the number of nest entrance detections reliably indicates nestling competitive ability, we found that the most‐competitive nestling fledged first at over half of all nests, supporting the notion that fledging usually begins when oldest nestlings reach a threshold state of development.     

A developmental constraint on the fledging time of birds

We examined the hypothesis that the rate of bone growth limits the minimum fledging time of birds. Previous observations in California gulls indicate that linear growth of wing bones may be the rate limiting factor in wing development. If bone growth is rate limiting, then birds with relatively long bones for their size could be expected to have longer fledging periods than birds with relatively short bones. We tested this by comparing the length of wing bones, relative to body mass, to the relative length of fledging periods among 25 families. The results support the hypothesis. A strong correlation exists between relative fledging period and relative bone length. Species which have relatively long bones for their body size tend to take longer to fly. In contrast, parameters that influence flight style and performance, such as size of the pectoralis muscle and wing loading, show little or no correlation with fledging time. The analysis also indicates that, when altricial and precocial species are considered together, bone length is more highly correlated with fledging time than is body mass or rate of increase in body mass during growth. These observations suggest that linear growth of bones does limit the growth of avian wings and that it is one of the factors that influences the fledging time of birds.     

Breeding biology of the thin-billed prion<Emphasis Type="Italic"> Pachyptila belcheri</Emphasis> at New Island,Falkland Islands: egg desertion,breeding success and chick provisioning in the poor season 2002/2003

We present data on breeding success, chick growth and chick feeding in thin-billed prions, Pachyptila belcheri, at New Island, Falkland Islands, in the breeding season 2002/2003. As in many populations of seabirds in the region, the overall breeding success was very low. This was mainly caused by low rate of observed burrow occupancy (60%) and hatching (57%) of thin-billed prions, while chick survival was closer to normal. Sixty-eight percent of the chicks survived to fledging. In total, a chick was successfully reared in 23% of the nests or from 38% of recorded eggs. The failed eggs were found to be incubated for 30 days, on average. The time of egg desertion coincided with the time of desertion of other seabirds at New Island, with a period of high sea-surface temperatures and low catch rates by the commercial fisheries. We describe chick growth and use repeated weighings, corrected for metabolic loss, to estimate meal sizes. Chicks were fed a mean 39.2 g in 77% of the nights. We discuss possible reasons for the observed extremely low hatching success, and compare with the breeding success of other seabirds at the Falkland Islands.     

The effects of force‐fledging and premature fledging on the survival of nestling songbirds

Despite the broad consensus that force‐fledging of nestling songbirds lowers their probability of survival and therefore should be generally avoided by researchers, that presumption has not been tested. We used radiotelemetry to monitor the survival of fledglings of Ovenbirds Seiurus aurocapilla and Golden‐winged Warblers Vermivora chrysoptera that we unintentionally force‐fledged (i.e. nestlings left the nest in response to our research activities at typical fledging age), that fledged prematurely (i.e. nestlings left the nest earlier than typical fledging age), and that fledged independently of our activities. Force‐fledged Ovenbirds experienced significantly higher survival than those that fledged independent of our activities, and prematurely fledged Ovenbirds had a similarly high survival to those that force‐fledged at typical fledging age. We observed a similar, though not statistically significant, pattern in Golden‐winged Warbler fledgling survival. Our results suggest that investigator‐induced force‐fledging of nestlings, even when deemed premature, does not necessarily result in reduced fledgling survival in these species. Instead, our results suggest that a propensity or ability to fledge in response to disturbance may be a predictor of a higher probability of fledgling survival.     

Post‐fledging survival of Little Owls Athene noctua in relation to nestling food supply

Among the range of determinants of post‐fledging survival in altricial birds, the energy supply to the growing juveniles is likely to play a central role. However, the exact mechanisms shaping post‐fledging survival are poorly understood. Using a food supplementation experiment, we determined the effect of variation in food supply on the survival of juvenile Little Owls Athene noctua from hatching to 2 months post‐fledging. Experimental broods were food‐supplemented for 36 days during the nestling and the early post‐fledging period. The fate of 307 juveniles (95 of them provided with extra food) was determined by nest monitoring and radiotelemetry. In unsupplemented birds, the rates of survival measured at 5‐day intervals were lowest during the nestling stage, remained low during the early post‐fledging stage and steadily increased after about 2 weeks post‐fledging. Food supplementation substantially increased nestling survival, but we detected no direct treatment effect on post‐fledging survival. Instead, we found a strong indirect effect of food supplementation, in that fledglings of good physical condition had markedly higher chances of surviving the post‐fledging period compared with those in poor condition. Experimental food supplementation increased survival over the first 3 months from 45% to 64.6%. This suggests that energy reserves built up during the nestling stage influence post‐fledging survival and ultimately parental reproductive output. The low nestling and post‐fledging survival shows that the early life‐history stages constitute a crucial bottleneck of reproductive ecology in Little Owls. The strong treatment effects on the number of independent offspring indicate that natural variation in food supply is an important determinant of spatio‐temporal patterns in Little Owl demography.     

Variation in nestling body condition and wing development predict cause‐specific mortality in fledgling dickcissels

Phenotypic traits developed in one life‐history stage can carryover and affect survival in subsequent stages. For songbirds, carryover effects from the pre‐ to post‐fledging period may be crucial for survival but are poorly understood. We assessed whether juvenile body condition and wing development at fledging influenced survival during the post‐fledging period in the dickcissel Spiza americana. We found pre‐ to post‐fledging carryover effects on fledgling survival for both traits during the ‘early part’ – first four days – of the post‐fledging period. Survival benefits of each trait depended on cause‐specific sources of mortality; individuals in better body condition were less likely to die from exposure to adverse environmental conditions, whereas those with more advanced wing development were less likely to be preyed upon. Fledglings with more advanced wing development were comparatively more active and mobile earlier in the post‐fledging period, suggesting they were better able to avoid predators. Our results provide some of the first evidence linking development of juvenile phenotypic traits to survival against specific sources of post‐fledging mortality in songbirds. Further investigation into pre‐ to post‐fledging carryover effects may yield important insights into avian life‐history evolution.     

Accelerated growth rates in late‐hatched Rhinoceros Auklet Cerorhinca monocerata chicks depend on food conditions and growth stage: an experimental approach

In some bird species, the survival of chicks hatching later in the season is lower than those hatched earlier due to increased risk of predation and a seasonal decline in feeding conditions. To reduce these risks, it might be advantageous for late‐hatched chicks to grow faster and hence fledge at younger age. In this experimental study, the growth rates of early‐ and late‐hatched Rhinoceros Auklet Cerorhinca monocerata chicks were compared under average and poor food supplies in captivity. Controlling for potentially confounding effects of chick mass at 10 days old, chick age and nest‐chamber temperature, late‐hatched chicks had higher wing growth rate than early‐hatched chicks before attaining the minimum wing length required for fledgling under both average and poor food supplies. After attaining the minimum wing length, however, late‐hatched chicks had a lower fledging mass, indicating a potential cost that could diminish the early advantage of fast wing growth.     

Effects of water discharge on fledging time,growth, and survival of piping plovers on the Missouri River

River flow management and modification is a global issue, and its effects on river-dependent organisms are pervasive. Flow modification can directly affect avian species through mortality or habitat loss, but less is known about indirect and sublethal effects of flow modification on reproductive output in these species. Young birds are more vulnerable to predation between hatching and fledging than after flight is achieved, but tradeoffs must be made to balance growth and survival. Predation pressure appears to be a significant factor affecting the time to fledging in altricial birds, but less is known about this threat for precocial birds. Birds reaching fledging earlier should have greater rates of survival to migration because their predator escape repertoire includes flight at an earlier age. We evaluated the effect of varying outflows from the Gavins Point Dam on the growth, age at fledging, and survival of piping plover (Charadrius melodus) chicks on the Missouri River (2006–2009). The study was characterized by 2 relatively high flow years (2006 and 2009) and 2 relatively low flow years (2007 and 2008). We used success rate in recapturing chicks in capture–mark–recapture models as an index for fledging. We attempted to recapture all chicks (n = 1,099) by hand every 3–4 days throughout the season to acquire morphological measurements. Models indicated that as flows from the dam increased, age at fledging increased. We also found that increasing flows were associated with decreasing daily survival rates (β_flow = −2.401, 95% CI: −4.351 to −0.452). Flow was also negatively related to chick mass gain, but we found less evidence for an effect on wing-chord length. Increased flows covered wet-substrate foraging habitat, and likely affected plover reproductive output directly through chick survival and indirectly through decreased growth and increased fledging times. © The Wildlife Society, 2013     

Mathematical modeling of appendicular bone growth in glaucous‐winged gulls

Development of locomotor activity is crucial in tetrapods. In birds, this development leads to different functions for hindlimbs and forelimbs. The emergence of walking and flying as very different complex behavior patterns only weeks after hatching provides an interesting case study in animal development. We measured the diaphyseal lengths and midshaft diameters of three wing bones (humerus, ulna, and carpometacarpus) and three leg bones (femur, tibiotarsus, and tarsometatarsus) of 79 juvenile (ages 0–42 days) and 13 adult glaucous‐winged gulls (Larus glaucescens), a semiprecocial species. From a suite of nine alternative mathematical models, we used information‐theoretic criteria to determine the best model(s) for length and diameter of each bone as a function of age; that is, we determined the model(s) that obtained the best tradeoff between the minimized sum of squared residuals and the number of parameters used to fit the model. The Janoschek and Holling III models best described bone growth, with at least one of these models yielding an R² ≥ 0.94 for every dimension except tarsometatarsus diameter (R² = 0.87). We used the best growth models to construct accurate allometric comparisons of the bones. Early maximal absolute growth rates characterize the humerus, femur, and tarsometatarsus, bones that assume adult‐type support functions relatively early during juvenile development. Leg bone lengths exhibit more rapid but less sustained relative growth than wing bone lengths. Wing bone diameters are initially smaller than leg bone diameters, although this relationship is reversed by fledging. Wing bones and the femur approach adult length by fledging but continue to increase in diameter past fledging; the tibiotarsus and tarsometatarsus approach both adult length and diameter by fledging. In short, the pattern of bone growth in this semiprecocial species reflects the changing behavioral needs of the developing organism. J. Morphol., 2009. © 2008 Wiley‐Liss, Inc.     

Buffering against food availability? The physical environment has little influence on breeding performance of fairy prions (Pachyptila turtur)

The breeding performance of seabirds is constrained by the availability of marine‐based prey, which may be influenced by competition with other seabirds, and environmental conditions. Fairy prion (Pachyptila turtur) populations have declined substantially in New Zealand since the introduction of mammalian predators; remaining small populations provide an opportunity to examine the effect of environmental factors on reproductive success in the absence of competition. Using 11 years of nest monitoring data and eight years of chick measurements we investigated (i) the link between physical environmental factors, breeding success rates and chick fledging size, and (ii) the relationship between chick fledging size and likelihood of natal‐colony recapture. Despite annual variations in the Southern Oscillation Index (SOI) and sea surface temperatures (SST), breeding success and chick fledging weights remained similar throughout the entire period, possibly due to the use of behavioural buffering mechanisms by parents. Fledgling wing lengths differed among years, and were predicted by early winter and early spring values of the SOI and late spring and summer SSTs. In years of high SOI and high SST, fairy prions fledged with longer wings, possibly due to increased availability of the euphausiid Nyctiphanes australis, their main prey. River flows and rainfall were unrelated to breeding success or chick measurements: the influence of freshwater on the marine system at this locality acts at smaller timescales than those studied here. Using the 2004 and 2005 cohorts, there was no link between fledging size and likelihood of recapture, except for the 2005 cohort which had unusually low 6‐year recapture rates, and shorter‐than‐average wing lengths. A combination of low sea temperatures and an El Niño event in 2005 may have led to a reduction in euphausiid availability, ultimately affecting post‐fledging survival.     

Cooperative breeding shapes post‐fledging survival in an Afrotropical forest bird

For avian group living to be evolutionary stable, multiple fitness benefits are expected. Yet, the difficulty of tracking fledglings, and thus estimating their survival rates, limits our knowledge on how such benefits may manifest postfledging. We radio‐tagged breeding females of the Afrotropical cooperatively breeding Placid greenbul (Phyllastrephus placidus) during nesting. Tracking these females after fledging permitted us to locate juvenile birds, their parents, and any helpers present and to build individual fledgling resighting datasets without incurring mortality costs or causing premature fledging due to handling or transmitter effects. A Bayesian framework was used to infer age‐specific mortality rates in relation to group size, fledging date, maternal condition, and nestling condition. Postfledging survival was positively related to group size, with fledglings raised in groups with four helpers showing nearly 30% higher survival until independence compared with pair‐only offspring, independent of fledging date, maternal condition or nestling condition. Our results demonstrate the importance of studying the early dependency period just after fledging when assessing presumed benefits of cooperative breeding. While studying small, mobile organisms after they leave the nest remains highly challenging, we argue that the telemetric approach proposed here may be a broadly applicable method to obtain unbiased estimates of postfledging survival.     

Facultative adjustment of pre‐fledging mass recession by nestling chimney swifts Chaetura pelagica

In species susceptible to mass‐dependent flight costs, mass recession prior to fledging may ensure that fledglings have appropriate wing loading. Our objectives were to determine if mass recession by chimney swift Chaetura pelagica nestlings is intrinsically controlled or facultatively adjusted by nestlings, and if mass recession is driven by changes in parental (i.e. reduced provisioning rates) or nestling (i.e. reduced begging) behavior. Nestling swifts (n = 50 in 17 broods) were divided into three treatment groups: controls, half‐weighted, or weighted. Half‐weighted and weighted nestlings had 0.6–0.7 or 1.2–1.3‐g lead weights, respectively, glued to body feathers on their backs during the period from 16 to 26 d post‐hatching. Weighted nestlings lost more mass than control and half‐weighted nestlings. After accounting for the added weights, control nestlings also had a higher wing loading than weighted nestlings. Video recordings revealed that provisioning rates of adult swifts did not vary throughout the nestling period, but the percent time nestlings spent begging increased slightly with age. Differences in mass recession among nestlings in different treatment groups resulted in convergence toward similar wing loading values likely optimal for flight efficiency. Mechanism(s) involved in this process remain unclear because provisioning rates were similar (from day 12 to 26 post‐hatching) whereas percent begging time by nestlings tended to increase with nestling age. However, weighted nestlings may have lost more mass than control nestlings by soliciting less food from adults than siblings, being more active, losing more water due to tissue maturation, or through some combination of two or more of these factors.     

Effect of food availability on nestling growth and fledging success in manipulated pallid swift broods

Growth rate and fledging success were assessed in natural and manipulated broods of the pallid swift Apus pallidus. Daily measurements of chick mass, wing length, and insect abundance allowed us to examine the short-term variation of chick growth in relation to food availability.
The number of fledged nestlings increased with brood size. Wing length and body mass were slightly but significantly smaller in larger broods, and the nestlings of enlarged broods needed longer to fledge. We discuss how these differences could influence survival after fledging.
Hatching asynchrony caused a significant difference in growth among siblings, and the difference between the oldest and youngest chick was greater in larger broods.
Chick growth was independent of daily food availability. We suggest that this was due to an increased effort of the parents at their expense, when food availability was poor.
The ability of this species to raise an additional chick is in line with most findings on birds, but partially in contrast with results for the common swift in which, at least during poor seasons, the additional nestling caused an increased mortality and lowered the reproductive success.