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.     

Effects of repeated investigator handling of Leach's Storm‐Petrel chicks on growth rates and the acute stress response

Disturbance during development may have lasting effects on the growth rates and stress physiology of birds. Although repeated handling by researchers is often necessary, the possible effects of such handling on the development of semi‐altricial young are unclear. We examined the effect of daily handling on growth rates and plasma corticosterone levels of Leach's Storm‐Petrel (Oceanodroma leucorhoa) chicks on Kent Island in the Bay of Fundy, New Brunswick, Canada, during the 2011 nesting season. From post‐hatch day 7 to post‐hatch days 14–36, birds in the experimental group were extracted from burrows and measured (wing, tarsus, and mass) for ～3 min every day, whereas birds in the control group were left undisturbed. After the treatment period, blood was collected from birds in both groups within 3 min of initially reaching into burrows (baseline) and after a 30‐min restraint stress test to assess the effect of early life disturbance on programming of the hypothalamic‐pituitary‐adrenal (HPA) axis. A second acute restraint stress test was conducted three weeks after the end of the treatment period to investigate possible longer term effects of early life disturbance. Growth rates of wings and tarsi were similar for handled chicks (N = 18) and non‐handled control chicks (N = 21), as were baseline and 30‐min acute restraint stress‐induced corticosterone levels. As also reported in previous studies of other altricial and semi‐altricial species, older chicks (42–64 d old) had higher plasma corticosterone levels than younger chicks (21–43 d old) after acute restraint stress tests, reflecting delayed development of the HPA axis. The age‐related increase in HPA axis sensitivity observed prior to fledging could facilitate foraging and predator avoidance behaviors while minimizing exposure to high levels of corticosterone earlier in development. Overall, we found no evidence that repeated disturbance influenced either growth rates or HPA axis programming of Leach's Storm‐Petrel chicks.     

Egg size and asymmetric sibling rivalry in red-winged blackbirds

How big to make an egg is a life history decision that in birds is made coincident with a series of other similar decisions (how many eggs to have, whether to fortify them with maternally derived hormones or immune system boosters, whether to hatch the eggs synchronously or asynchronously). Though within-population variation in egg size in birds has been well studied, its adaptive significance, if any, is unclear. Here we examine within-population variation in egg size in relation to asymmetric sibling rivalry in a 17-year study of red-winged blackbirds (Agelaius phoeniceus), an altricial songbird. Egg mass showed a twofold range of variation, with roughly 80% of the variation occurring across clutches. By commencing incubation before the clutch is complete, mothers create advantaged core and disadvantaged marginal elements within their brood. Previous work on this system has shown that sibling competition is asymmetric, and that core offspring enjoy priority access to food, and as a consequence show higher growth and lower mortality than marginal offspring. Here we examine the effect of initial egg size on nestling growth and survival in relation to these competitive asymmetries. Egg mass was strongly linked to hatchling mass, and remained significantly related to the mass of both core and marginal nestlings; the effect of egg size was stronger for core offspring early in the nestling period, but the disparity between core and marginal nestlings narrowed as they approached fledging age, and slower growing marginals fell victim to brood reduction. The effect of egg mass on survival differed dramatically between core and marginal nestlings. Egg mass was significantly related to the survival of marginal but not core nestlings: below average egg mass was associated primarily with very early mortality. Asymmetric sibling competition is clearly a strong determinant of the consequences of egg size variation.     

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.     

Development of stress response in nestling pied flycatchers

Birds respond to unpredictable events by secreting corticosterone, which induces various responses to cope with stressful situations. However, the evidence is still elusive whether altricial nestlings perceive and respond to external stressors. We investigated the development of adrenocortical stress response to handling-related stressor in nestlings of a small passerine bird, the pied flycatcher (Ficedula hypoleuca). Nestlings were held in isolation from their parents during the experiment to ensure that they indeed respond to handling, not to parental alarm calls. We found that both 9- and 13-day-old nestlings were able to elicit hormonal stress response. Although baseline as well as stress-induced corticosterone levels rose slightly with age, the magnitude of difference between the control and stress-induced levels remained similar in both age groups. However, comparison with adults showed that the stress response of nestlings prior to fledging was still incomplete and significantly lower than in adults. Overall, our results indicate that altricial nestlings do respond to acute stressors, but on the contrary to previous predictions the development of corticosterone stress response during growth period is not gradual and varies remarkably between different passerine species.     

Hatching asynchrony and growth trade‐offs within domesticated and wild zebra finch,Taeniopygia guttata,broods

The Australian zebra finch, Taeniopygia guttata, is a widely used model organism, yet few studies have compared domesticated and wild birds with the aim of examining its relevance as an evolutionary model species. Domestic and wild broods hatch over approximately 4 and 2 days, respectively, which is important given that nestlings can fledge after as little as 12 days, although 16–18 days is common. We aimed to evaluate the extent to which the greater hatching asynchrony in domestic stock may effect reproductive success through greater variance in size hierarchies, variance in within‐brood growth rates, and partial brood mortality. Therefore, by simultaneously controlling brood sizes and experimentally manipulating hatching intervals in both domesticated and wild birds, we investigated the consequences of hatching intervals for fledging success and nestling growth patterns, as well as trade‐offs. Fledging success was similarly high in domestic and wild broods of either hatching pattern. Nonetheless, between‐brood analyses revealed that domestic nestlings had significantly higher masses, larger skeletal characters, and longer wings than their wild counterparts, although wild nestlings had comparable wing lengths at the pre‐fledging stage. Moreover, within‐brood analyses revealed only negligible differences between domestic and wild nestlings, and larger effects of hatching order and hatching pattern. Therefore, despite significant differences in the hatching intervals, and the ultimate size achieved by nestlings, the domestication process does not appear to have significantly altered nestling growth trade‐offs. The present study provides reassuring evidence that studies involving domesticated zebra finches, or other domesticated model organisms, may provide reasonable adaptive explanations in behavioural and evolutionary ecology. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 100 , 763–773.     

Ontogeny of Adaptive Antibody Response to a Model Antigen in Captive Altricial Zebra Finches

Based on studies from the poultry literature, all birds are hypothesized to require at least 4 weeks to develop circulating mature B-cell lineages that express functionally different immunoglobulin specificities. However, many altricial passerines fledge at adult size less than four weeks after the start of embryonic development, and therefore may experience a period of susceptibility during the nestling and post-fledging periods. We present the first study, to our knowledge, to detail the age-related changes in adaptive antibody response in an altricial passerine. Using repeated vaccinations with non-infectious keyhole limpet hemocyanin (KLH) antigen, we studied the ontogeny of specific adaptive immune response in altricial zebra finches Taeniopygia guttata. Nestling zebra finches were first injected at 7 days (7d), 14 days (14d), or 21 days post-hatch (21d) with KLH-adjuvant emulsions, and boosted 7 days later. Adults were vaccinated in the same manner. Induced KLH-specific IgY antibodies were measured using ELISA. Comparisons within age groups revealed no significant increase in KLH-specific antibody levels between vaccination and boost in 7d birds, yet significant increases between vaccination and boost were observed in 14d, 21d, and adult groups. There was no significant difference among age groups in KLH antibody response to priming vaccination, yet KLH antibody response post-boost significantly increased with age among groups. Post-boost antibody response in all nestling age groups was significantly lower than in adults, indicating that mature adult secondary antibody response level was not achieved in zebra finches prior to fledging (21 days post-hatch in zebra finches). Findings from this study contribute fundamental knowledge to the fields of developmental immunology and ecological immunology and strengthen the utility of zebra finches as a model organism for future studies of immune ontogeny.     

Early exposure to a bacterial endotoxin advances the onset of moult in the European starling

In animals, events occurring early in life can have profound effects on subsequent life‐history events. Early developmental stresses often produce negative long‐lasting impacts, although positive effects of mild stressors have also been documented. Most studies of birds have investigated the effects of events occurring at early developmental stages on the timing of migration or reproduction, but little is known on the long‐term effects of these early events on moulting and plumage quality. We exposed European starling Sturnus vulgaris nestlings to an immune challenge to assess the effects of a developmental stress on the timing of the first (post‐juvenile) and second (post‐breeding) complete annual moult, the length of the flight feathers, and the length and colouration of ornamental throat feathers. The nestlings were transferred to indoor aviaries before fledgling and kept in captivity until the end of post‐breeding moult. Individuals treated with Escherichia coli lypopolysaccharide (LPS) started both moult cycles earlier compared to control siblings. Moult duration was unaffected by the immune challenge, but an advanced moult onset resulted in a longer moult duration. Moreover, female (but not male) throat feather colouration of LPS‐injected individuals showed a reduced UV chroma. We argue that an early activation of the immune system caused by LPS may allow nestlings to better cope with post‐fledging stresses and lead to an earlier moult onset. The effect of early LPS exposure was remarkably persistent, as it was still visible more than one year after the treatment, and highlighted the importance of early developmental stresses in shaping subsequent major life‐history traits, including the timing of moult in birds.     

Environmental influences on the evolution of growth and developmental rates in passerines

Abstract.— The reasons why growth and developmental rates vary widely among species have remained unclear. Previous examinations of possible environmental influences on growth rates of birds yielded few correlations, leading to suggestions that young may be growing at maximum rates allowed within physiological constraints. However, estimations of growth rates can be confounded by variation in relative developmental stage at fledging. Here, we re-estimate growth rates to control for developmental stage. We used these data to examine the potential covariation of growth and development with environmental variation across a sample of 115 North American passerines. Contrary to previous results, we found that growth rates of altricial nestlings were strongly positively correlated to daily nest predation rates, even after controlling for adult body mass and phylogeny. In addition, nestlings of species under stronger predation pressure remained in the nest for a shorter period, and they left the nest at lower body mass relative to adult body mass. Thus, nestlings both grew faster and left the nest at an earlier developmental stage in species with higher risk of predation. Growth patterns were also related to food, clutch size, and latitude. These results support a view that growth and developmental rates of altricial nestlings are strongly influenced by the environmental conditions experienced by species, and they generally lend support to an adaptive view of interspecific variation in growth and developmental rates.     

Timing of incorporation of docosahexaenoic acid into brain and muscle phospholipids during precocial and altricial modes of avian development

We investigated the possibilities that the proportion of docosahexaenoic acid (DHA) in phospholipids of brain and skeletal muscle at hatch, and the ontogenetic timing of the DHA accretion spurt in these tissues, might serve as indices of neonatal functional maturity that discriminate between precocial and altricial avian developmental modes. Comparison of the fatty acid profiles of the initial and residual yolks of two free-living altricial species, the swallow (Hirundo rustica) and the sparrow (Passer domesticus), reveals that, in contrast to precocial birds, there is no preferential uptake of DHA from the yolk during embryonic development. At hatch, the proportions of DHA in brain phospholipid (wt.% of fatty acids) of the swallow and sparrow, at 8.1% and 5.0%, respectively, are far lower than the values (16.9-19.6%) reported for non-altricial species. This reflects a marked difference in the timing of the brain DHA accretion spurt, which occurs during the first half of the embryonic period of precocial birds, but is largely delayed until after hatching in the altricial species. By the time of fledging, the proportion of DHA in the swallow brain phospholipid has increased to 14.3%. For non-altricial birds, the brain DHA concentration at hatch shows little interspecies variation, despite major differences in yolk DHA content. The proportions of DHA in leg muscle phospholipid of the newly hatched swallow and sparrow, at 2.9% and 2.5%, respectively, are far lower than the value (6.7%) for the precocial chicken. Again, this relates to differences in developmental timing, with muscle DHA accretion occurring in the first half of the chicken's embryonic period, whereas, in the swallow, this increase is delayed until after hatching. By the time of fledging in the swallow, DHA forms 9.3% of muscle phospholipid fatty acids, equivalent to the level attained in chicken muscle at the mid-embryo stage. The results indicate a clear distinction between altricial and non-altricial avian species in the timing of tissue DHA accretion during development, presumably reflecting differences in neonatal functional maturity.     

Post‐independence mortality of juveniles is driven by anthropogenic hazards for two passerines in an urban landscape

Urban environments impose novel selection pressures with varying impacts across species and life history stages. The post‐fledging stage for migratory passerines, defined as the period of time from when hatch‐year birds fledge until their first migration, is a poorly understood component of annual productivity that potentially limits population growth. We studied two migratory passerines with positive and negative population responses to urbanization, respectively: gray catbird Dumetella carolinensis and wood thrush Hylocichla mustelina. Our goals were to estimate post‐fledging survival rates for urban bird populations and determine which features of the urban landscape impact mortality risk during the post‐fledging stage. From 2012–2014, we tracked 127 fledglings (60 gray catbirds and 67 wood thrushes). Over 55 d after fledging, cumulative survival of gray catbirds (0.32 [95% CI: 0.22–0.47]) was approximately half that of wood thrushes (0.63 [95% CI: 0.52–0.75]). Thus, survival rates during the post‐fledging stage, taken in isolation, do not explain differential trajectories of gray catbird and wood thrush populations in urban environments. Most mortality (86%) for both species was due to predation. However, after reaching independence from parental care, 6 birds (9.4% of mortalities) died of anthropogenic causes (e.g. building, car strikes). Crossing roads significantly increased mortality risk, but increasing daily movement distance decreased mortality risk. Our results raise the question of whether anthropogenic sources of mortality are compensatory or additive to natural mortality; we emphasize the need to monitor fledgling survival beyond the parental‐dependence stage in order to fully understand the impacts of anthropogenic hazards on juvenile birds.     

Influence of hatching order on growth rate and resting metabolism of kestrel nestlings

Hatching asynchrony in altricial birds may result in a competitive disadvantage for the youngest nestlings compared to older siblings. We studied the effects of a size hierarchy on the growth rate of Eurasian kestrels Falco tinnunculus chicks in nests with and without access to supplemented food in western Finland. Body mass stopped increasing on the 19th day after hatching while body size, estimated by a combination of bone and feather lengths continued to increase at least until fledging at 26 days. Body condition, reflecting muscle and fat, did not change markedly during the growth period from the 12th day to fledging. Body temperature and resting metabolism were usually lower in nestlings 12 days old than in nestlings at fledging. Growth of body mass, size and condition, and resting metabolism were delayed in last-hatched nestlings aged 19 days. Just before fledging, last-hatched nestlings attained a similar body mass and size, and had a similar resting metabolism to those of older siblings. At fledging, only in nests without access to supplemented food was the body condition of last-hatched chicks lower than that of its siblings, but in nests with access to supplemented food no such difference was detected. Our results highlight that the level of lipids in the last-hatched nestling can be affected by the food restriction imposed by hatching order.     

Why renew fresh feathers? Advantages and conditions for the evolution of complete post‐juvenile moult

Juveniles of several passerine species renew all of their fresh juvenile feathers immediately after fledging (complete post‐juvenile moult), in contrast to the majority, which perform a partial post‐juvenile moult. To understand the adaptive roles of this phenomenon we compared the quality of juvenile plumage in species that perform a complete post‐juvenile moult with that of species which perform a partial post‐juvenile moult; we similarly compared juveniles and adults in each of these groups. The quality of feathers was measured by mass of primaries, colour, and length. In species which perform a complete post‐juvenile moult the plumage quality of second‐year individuals, in their first breeding season, is similar to the plumage quality of adults, unlike those species that perform a partial post‐juvenile moult. In species which perform complete post‐juvenile moult, the quality of the feathers grown in the nest is lower than the quality of adult post‐breeding feathers. In contrast, in species which perform partial post‐juvenile moult the quality of the feathers grown in the nest is similar to that of adult post‐breeding feathers. We found that a complete post‐juvenile moult strategy is much more common 1) in residents and short‐distance migrants than in long‐distance migrants, 2) in southern latitudes, 3) in species with medium body mass and 4) in omnivores and granivores. Our results indicate two adaptive roles of the complete post‐juvenile moult strategy: 1) achieving high quality plumage in the first year which may increase individual survival probability and fitness and 2) allocating fewer resources to nestling plumage and more to nestling development, which enables the nestlings to leave the nest earlier, thus reducing the probability of encountering nest predators. We suggest that the complete post‐juvenile moult, immediately after fledging, is an optimal strategy in favourable habitats and under low time constraints, as in some tropical ecosystems.     

Comparing food limitation among three stages of nesting: supplementation experiments with the burrowing owl

Food availability is an important limiting factor for avian reproduction. In altricial birds, food limitation is assumed to be more severe during the nestling stage than during laying or incubation, but this has yet to be adequately tested. Using food‐supplementation experiments over a 5‐year period, we determined the degree and timing of food limitation for burrowing owls (Athene cunicularia) breeding in Canada. Burrowing owls are an endangered species and food limitation during the nestling stage could influence reproductive performance of this species at the northern extent of their range. Supplemented pairs fledged on average 47% more owlets than unfed pairs, except during a year when natural food was not limiting (i.e., a prey irruption year). The difference in fledgling production resulted from high nestling mortality in unfed broods, with 96% of all nestling deaths being attributed to food shortage. Supplemental feeding during the nestling period also increased fledgling structural size. Pairs fed from the start of laying produced the same number of hatchlings as pairs that received no supplemental food before hatch. Furthermore, pairs supplemented from egg laying to fledging and pairs supplemented during the nestling period alone had the same patterns of nestling survival, equal numbers of fledglings, and similar fledgling mass and structural size. Our results provide empirical support for the hypothesis that the nestling period is the most food‐limited phase of the breeding cycle. The experimental design we introduce here could be used with other altricial species to examine how the timing of food limitation differs among birds with a variety of life‐history strategies. For burrowing owls, and other species with similar life histories, long‐term, large‐scale, and appropriately timed habitat management increasing prey abundance or availability is critical for conservation.     

Parental Alarm Calls of the White‐Crowned Sparrow Fail to Stimulate Corticosterone Production in Nest‐Bound Offspring

Alarm calling by parents is widespread among animals and has strong implications for parent and offspring fitness, yet it is virtually unknown whether parental alarm calls can initiate a corticosterone response in offspring. We investigated whether parental alarm calls of the white‐crowned sparrow, Zonotrichia leucophrys, activated the corticosterone response of their nest‐bound young, as such a response might prepare older nestlings for premature fledging and increase their survival when contacted by a predator at the nest. We conducted an experiment in which nestlings were either exposed to parent alarm calls (treatment) or experienced a period without parental alarm calls (control) immediately prior to blood sampling. We then sampled nestlings to measure corticosterone levels within 4 min of first contact (baseline corticosterone) and 60 min later (handling‐induced corticosterone). Young nestlings (i.e. 3–4 d post‐hatch) did not exhibit a corticosterone response to parental alarm calls or to handling, as mean corticosterone levels were similar in the control and treatment groups for both baseline and 60‐min post‐baseline samples. Against our predictions, there was no difference in mean levels of baseline corticosterone between control and treatment groups in older nestlings (i.e. 7?8 d post‐hatch) that were capable of surviving out of the nest. However, we did find a significant increase in mean levels of corticosterone after handling in both groups, which indicated that older nestlings were able to mount a functional corticosterone response when confronted with a potential predator. Why older nestlings did not initiate a corticosterone response after exposure to parental alarm calls is unclear but may have occurred because the costs of mounting such a response outweighed the benefits, perhaps because of growth or developmental costs.     

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.     

Do parents play a role in the timing and process of fledging by nestling Mountain Bluebirds?

What causes young birds to leave nests remains unclear for almost all altricial species. For many years, the assumption was that parents often controlled the time of fledging by coaxing young from nests, e.g., by holding food within view, but out of reach, of nestlings. This assumption, though, was based solely on scattered anecdotal reports of such behavior. We used continuous video‐recording of nests to assess the role of parents, if any, in the timing and process of fledging of cavity‐nesting Mountain Bluebirds (Sialis currucoides). We placed perches ~50 cm in front of nest‐box entrances to give parents ample opportunity to display food to nestlings. We found no evidence that parents routinely initiated the fledging process. On the day of fledging, parents did not perch on supplemental perches with food more often, or for longer periods of time, than on the day before fledging. Also, after going to nest‐box entrances, parents never held food away from a nestling reaching for the food. Parents were usually absent (16 of 19 cases) when the first nestling fledged. In the remaining three cases, a parent perched with food in view of a nestling for 8, 15 and 65 s, respectively, just before that nestling fledged. Although these might have appeared to be attempts at coaxing, in each case, the parent was encountering, for the first time, a nestling partially emerging from the nest entrance. Parents may simply have hesitated to approach nests because the nestling's position prevented parents from delivering food in the normal manner. Finally, the rate at which parents fed nestlings on the day of fledging did not differ from the rate the day before, suggesting that parents do not try to use hunger to induce fledging. Our results are consistent with previous research suggesting that, in Mountain Bluebirds, it is a nestling that initiates fledging, typically when it reaches some threshold state of development.     

Vitamin E improves growth of collared flycatcher Ficedula albicollis young: a supplementation experiment

In altricial birds, the quantity and quality of food provided by parents is a crucial determinant of nestling performance. Vitamin E is an important micronutrient with various physiological functions, including a positive role in the antioxidant system. Sufficient intake of vitamin E has been shown to condition normal avian development in poultry, yet, our knowledge of the role of vitamin E in free‐living birds is limited. Thus, we experimentally examined the effects of vitamin E on nestling development in the collared flycatcher Ficedula albicollis. We supplemented nestlings with vitamin E and evaluated their growth and survival till fledging. Increased availability of vitamin E did not affect body mass, wing length or survival, but improved tarsus growth. The effect of supplementation on tarsus length changed over season and with initial body mass. Supplemented nestlings that were smaller at hatching and those that hatched later in the season grew longer tarsi compared to the control. Our results suggest that 1) vitamin E may be limiting for the development of collared flycatcher nestlings, 2) seasonal changes of vitamin E availability may affect breeding success of collared flycatchers, and 3) increased income of vitamin E may improve growth of nestlings with bad start in life.     

Contrasting patterns of post‐fledging movements of two sympatric warbler species with different life‐history strategies

Broad‐scale movements of migrant songbirds during the post‐fledging period are hypothesized to aid in the development of navigational abilities, to allow individuals to prospect for future breeding territories (combined as regional exploration), or as representing the commencement of migration. Using an automated radio telemetry array, we compared broad‐scale post‐fledging movements of hatch‐year individuals from two closely related species: blackpoll warblers Setophaga striata and myrtle warblers Setophaga coronata coronata. These two species have contrasting migratory strategies (long‐distance vs short‐distance), and we studied populations from two different islands in Nova Scotia that have different geographical landscape features. Locally‐hatched individuals affixed with VHF radios in August were tracked throughout the Gulf of Maine region for up to 2.5 months after tagging. Departure date and direction, daily probability of initiating a flight, daily displacement, total displacement and net displacement were assessed to see if there was support for the commencement of migration or regional exploration hypotheses. We observed differences between both species and islands. Compared to blackpolls, myrtles departed later, had more variable timings and directions of departure, made fewer regional‐scale flights, were more directional in their movements, and had higher net displacement. Total displacement and daily flight distances were similar between species. Variability of departure behaviour of myrtles was observed on the island farther from the mainland and both species made longer flights from that island. These results are consistent with the hypothesis that hatch‐year blackpoll movements are a form of regional exploration and hatch‐year myrtle movements represent the initial stages of migration. Species differences may be related to migratory strategy (long‐distance vs short‐distance), where the need to acquire information during post‐fledging for navigational purposes is higher for blackpolls than myrtles. Island differences suggest that habitat quality and ecological barriers influence broad‐scale movements, and myrtles are more facultative in their behaviour than blackpolls.     

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

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