Post-fledging body mass increase in Common Terns Sterna hirundo: influence of age, sex and year

We studied the inter-year and inter-sex variation of the post-fledging body mass development of Common Terns Sterna hirundo in 2000 and 2001 at the Banter See colony, northern Germany. Here, post-fledglings can be identified and weighed remotely and automatically by a transponder system that makes use of automated balances installed at the colony. Individuals were sexed with PCR amplification methods. After fledging, young generally continued to increase their mass. However, in 2000, the young did not significantly increase their mass during the post-fledging period. In 2001, conditions were more favourable and body mass increased continuously. Further, in 2001, male post-fledglings were significantly heavier than female post-fledglings. Once having left the colony area (on average 18–23 days after fledging in 2000, and 14–16 days after fledging in 2001), post-fledgling body mass had still not reached adult body mass. The longer a juvenile stayed at the colony, the higher was its final body mass, which if acting as a threshold level may control departure time. Neither brood size nor hatching order affected post-fledging mass or period. In the unfavourable year 2000, when many individuals were found dead after fledging, fledging age but not fledging mass was found to be a predictor of post-fledging survival before departure: individuals fledging when older had a lower survival probability. Our results stress the importance of the post-fledging period for body mass increase and survival prior to departure. The variation in post-fledging mass growth between years and between the sexes is discussed with respect to parental effort and a possible selective provisioning of sons over daughters.     

Variation in growth in Sandwich Tern chicks Sterna sandvicensis and the consequences for pre- and post-fledging mortality

Fitness consequences of variation in body mass growth and body condition were studied in a Sandwich Tern Sterna sandvicensis colony on Griend, Dutch Wadden Sea, during 1990–2000. Body mass increment during the linear growth phase predicted nestling survival probabilities accurately. Chicks growing less than 8 g per day had low survival probabilities until fledging, but within a range of 8–11 g per day growth only small effects on chick survival were observed. Effects of slow growth on survival became obvious after about 10 days after hatching. Slow growing chicks reached a much lower fledging mass, whereas slow growth had only small effects on structural size at fledging. Body condition of the chicks was highly variable and had strong effects on survival until fledging. However, body condition during the nestling stage did not influence post-fledging survival. Body condition at fledging had no effects on post-fledging survival and did not affect final mass or body size. It is argued that low fledging mass can be overcome soon after fledging, as parents take their fledglings closer to the foraging areas, thereby avoiding high rates of kleptoparasitism by Black-headed Gulls Larus ridibundus .     

Effects of individual pre-fledging traits and environmental conditions on return patterns in juvenile king penguins

Despite the importance of early life stages in individuals' life history and population dynamics, very few studies have focused on the constraints to which these juvenile traits are subjected. Based on 10 years of automatic monitoring of over 2500 individuals, we present the first study on the effects of environmental conditions and individual pre-fledging traits on the post-fledging return of non-banded king penguins to their natal colony. Juvenile king penguins returned exclusively within one of the three austral summers following their departure. A key finding is that return rates (range 68-87%) were much higher than previously assumed for this species, importantly meaning that juvenile survival is very close to that of adults. Such high figures suggest little juvenile dispersal, and selection occurring mostly prior to fledging in king penguins. Pre-fledging conditions had a strong quadratic impact on juvenile return rates. As expected, cohorts reared under very unfavourable years (as inferred by the breeding success of the colony) exhibited low return rates but surprisingly, so did those fledged under very favourable conditions. Juvenile sojourns away from the colony were shorter under warm conditions and subsequent return rates higher, suggesting a positive effect of climate warming. The longer the post-fledging trip (1, 2 or 3 years), the earlier in the summer birds returned to their natal colony and the longer they stayed before leaving for the winter journey. The presence of juveniles in the colony was more than twice the duration required for moulting purposes, yet none attempted breeding in the year of their first return. Juvenile presence in the colony may be important for acquiring knowledge on the social and physical colonial environment and may play an important part in the learning process of mating behaviour. Further studies are required to investigate its potential implications on other life-history traits such as recruitment age.     

Post-fledging survival of individual great tits: the effect of hatching date and fledging mass

Pre-breeeding survival is one of the major sources of individual variation in lifetime reproductive success. However, very little is known about the reasons for differences in survival among individuals during this important phase of the life cycle. Some studies, using local return rates as indices of survival, have shown a relationship between post-fledging survival and fledging date and mass in birds, most of them suggesting directional selection towards heavy masses and early fledging dates. Recent development of capture-recapture models allows the separate estimate of survival and recapture probabilities, as well as the inclusion of individual covariates into the modelling process. We used here these models to explore the relative effects of fledging date and fledging mass on local recruitment of individual great tit Parus major fledglings. Individual capture-recapture histories of 2051 fledglings (cohorts 1992–1999), 184 of which were recaptured as breeding birds during 1993–2000, were used in the analyses. Hatching date, offspring mass at day 15, their squared terms, and interactions between mass and date, were included as covariates into the modelling process. Models with age (fledglings and adults) and time (year) dependence were used. The probability of local recruitment increased with fledging mass in each of the years studied. Fledging date also affected recruitment but, against what is commonly thought, fledgling early is not the best option every year. Either early, intermediate or late fledglings were favoured in different years. This between-year variation in the optimum fledging date offers an alternative explanation to the lack of evolution towards earlier breeding dates, in spite of the advantages of early breeding some years.     

Influence of temperatures during the nestling period on post-fledging survival of great tit Parus major in a Mediterranean habitat

Survival during the first year is one of the most important factors determining fitness in birds. Birds have poor thermoregulatory abilities during the nestling period when the plumage is not fully developed, thus temperature during the nestling period is a serious candidate to affect post-fledging survival, although it has been usually ignored in previous studies. We analysed the relationship between temperatures and post-fledging survival in a great tit Parus major population in Sagunto (Spain) using capture-recapture data from 12 years. Hatching dates, mass at fledging and temperatures during the nestling period (maximum, minimum and mean ambient temperatures as an estimation of the mean and extreme weather conditions that chicks encountered at the nest) were used as individual covariates. Mean post-fledging survival was 0.13±0.01. Adult survival probability was 0.64±0.02. Multi-model inference suggested that post-fledging survival increased with fledging mass, and decreased as temperatures increased. Furthermore, the effect of mass on survival was less important as temperature was higher. We consider that high temperatures affect nestlings' health due to low thermoregulatory abilities of nestlings. Model selection did not support a relationship between hatching dates and survival once mass and temperatures were taken into account. The results suggest the possibility that the effect of date on post-fledging survival found in previous studies was, at least in part, a consequence of the seasonal pattern of temperature variation.     

The post-fledging survival of young Guillemots Uria aalge in relation to hatching date and growth

The post-fledging survival of a total of 1277 young Guillemots Uria aalge ringed in 6 years was assessed using sightings of 267 individuals back at the natal colony and recoveries of 46 ringed birds. In two years there was a significant decline in survival prospects with estimated hatching date, the first time such a trend has been demonstrated in the Alcidae. In these years pairs breeding early had a 2–3 times greater chance of having a young survive to return to the colony than pairs breeding three weeks later. Hatching date had no effect on survival prospects in the other four seasons and in none of the years did chick body condition have a demonstrable effect on post-fledging survival.     

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.     

Quantifying density dependence in a bird population using human disturbance

Although density dependence has long been recognised as vital to population regulation, there have been relatively few studies demonstrating it spatially in wildlife populations, often due to the confounding effects of variation in habitat quality. We report on a study of woodlarks Lullula arborea, a species of European conservation concern, breeding on lowland heath in Dorset, England. We take the novel approach of utilising the birds’ response to human disturbance, which resulted in much of the variation in density but had no direct impact on demographic rates. Within years, in sites with greater density there were smaller mean chick masses, lower post-fledging survival, and higher rates of nestling mortality attributed to starvation. The effects on clutch size and fledging success were confounded by the area of grassland within a site. There was no effect on brood size. Density dependence also operated within sites between years: as density increased there were reductions in mean chick mass and post-fledging survival, while nestling mortality attributed to starvation increased. Density-dependent effects on clutch size were only weakly regulatory, whereas density-dependent starvation and post-fledging mortality rates contributed strongly to differences in overall breeding output. Heavier chicks (when 7 days old) were significantly more likely to fledge and less likely to starve. Broods with heavier chicks were more likely to supply recruits to the breeding population. Nestling mass was not a factor in survival in the immediate post-fledging period, suggesting that density-dependent processes act independently on this stage. We conclude that the number of birds per hectare of suitable habitat is a valid means of expressing density, and that habitat acts as a surrogate for food abundance through which density dependence operates on the woodlark population.     

Nesting and post-fledging predation risk influence diel patterns of songbird fledging

Among stages of avian ontogeny, the act of nest departure or fledging is an abrupt transition into a new environment and a major leap toward independence for offspring. In altricial birds, the timing (i.e. time of day) of fledging is notable in that many species tend to fledge early in the morning. Past studies have proposed nest predation as a key factor driving birds to fledge earlier in the morning (the ‘survival hypothesis’), whereby offspring avoid peak times of nest predation that occur later in the day. A natural extension of this hypothesis is the predation of offspring post-fledging, whereby offspring are also timing their fledging with future survival prospects outside of the nest. However, few studies have investigated fledging behaviour in the context of both nesting and post-fledging predation. To help fill this knowledge gap, we investigated factors driving the timing and duration of fledging across six songbird species in the context of offspring predation: daily nest mortality, post-fledging mortality and diel patterns of nest predation risk. We found that > 60% of songbirds fledged early in the morning, whereas the peaks in nest predation risk occurred several hours post-fledging. Furthermore, species under greater risk of nest predation fledged earlier in the day and in closer succession to their siblings. Parameters of post-fledging mortality were poor predictors of fledging timing, but individuals from broods of species under higher risk of post-fledging mortality fledged in closer succession to their siblings. These results provide evidence in support of the survival hypothesis, and suggest that songbirds fledge in the morning to avoid peak times of nest predation risk that occur later in the day (~ 8 h after civil dawn). Such results corroborate past research highlighting predation on dependent offspring as a key factor driving variation in life histories across animal taxa; however, estimates of post-fledging mortality suggest that nest predation alone does not fully explain variation in fledging behaviour among species. Future research is therefore needed to investigate the contribution of other factors, such as energetics, parent–offspring conflict and diel patterns of post-fledging survival, which may help to mediate diel patterns of fledging within and among songbird species.     

To what extent do environmental factors affect the long-distance nocturnal post-fledging movements of the Reed Warbler?

We studied the effects of weather and the lunar cycle on long-distance nocturnal pre-migratory flights of Reed Warblers (Acrocephalus scirpaceus). Noturnal tape luring was used to capture the birds, and the study was carried out in a habitat atypical of this species on the Courish Spit (southeastern Baltic) between1999 and 2002. A total of 443 juvenile Reed Warblers were captured during 120 nights of trapping. Based on data on the moult and body condition of the birds, it was possible to identify 163 individuals as being on post-fledging movements. More than half of the birds (54.0%) were captured at the end of night during the nautical and civil twilight period; the remaining individuals were caught during the astronomical twilight period and darkest periods of night. Many birds performed pre-migratory flights in the middle of the night, the period during which a large part of the moon was visible. We suggest that the increased visibility under the full moon may provide conditions in which Reed Warblers increase the distance or intensity of nocturnal post-fledging movements. During the entire course of the night, the birds generally preferred to fly under rainless conditions, limited cloud cover and/or in still air. The birds performed flights under winds stronger than 2 m s⁻¹ when the wind was blowing along the axis of the spit. We also suggest that the birds generally fly along the spit. We found a weak but significant relationship between the numbers of Reed Warblers captured during post-fledging movements and the individual weather parameters and their interaction. Our data suggest that endogenous stimuli rather than weather parameters or lunar cycle phase determine the decision of Reed Warblers to undertake pre-migratory long-distance flights at night.     

The effect of body condition on the timing and success of breeding in Little Penguins Eudyptula minor

We studied the effect of parental body condition on the breeding biology of Little Penguins Eudyptula minor . Daily attendance patterns and body mass were recorded using an Automated Penguin Monitoring System, which collected arrival and departure masses for approximately 200 breeding birds over the 2000 and 2001 breeding seasons. Breeding success varied between the two years; 2000 was a year of average breeding success (fledging 1.07 chicks per pair), and 2001 a year of poor breeding success (fledging 0.53 chicks per pair). In both years, adult body condition (body mass divided by flipper length) increased significantly prior to laying. The laying period began over a month later in 2000 than in 2001, and birds in 2000 exhibited significantly better body condition at laying. However, the mean laying dates in 2000 were less variable than in the 2001 breeding season. Body condition appeared to influence both the time of breeding and breeding synchrony in Little Penguins. Breeding success was correlated with adult body condition at incubation in 2000 but not in 2001, indicating that success was not solely influenced by adult body condition at incubation.     

Are there trade-offs between pre- and post-fledging survival in black brent geese?

1.?The growth period is an important determinant of fitness later in life through its effects on first-year survival and future reproduction. Choices by adult females about where to rear their offspring strongly affect growth rates and offspring fitness in geese. 2.?Individual female black brent (Branta bernicla nigricans) tend to raise their broods in the same areas each year, and these areas are consistently ranked with respect to growth rates of goslings. Therefore, some females consistently rear their broods on areas resulting in lower post-fledging fitness. 3.?We explore the potential that growth rates of offspring (and associated fitness consequences) are traded off against other vital rates influencing fitness of either adult females or goslings. Growth of goslings primarily influences fitness after fledging, so one hypothesis is that survival before fledging, which is influenced by predation, is traded off against growth rates and post-fledging survival. 4.?We estimated pre-fledging and post-fledging survival for goslings reared on areas used by broods from the Tutakoke River black brent colony. We examined recaptures, recoveries by hunters and resightings of brent marked as goslings with webtags and standard leg rings. These data were analyzed using capture-mark-recapture models in program mark to derive separate estimates of pre- and post-fledging survival for 18 cohorts (1987-2004) of black brent goslings across seven brood rearing areas (BRAs). 5.?Estimates of pre-fledging survival probability varied from 0·00?±?0·00 (mean?±?95% confidence interval) to 0·92?±?0·1; and estimates of post-fledging survival probability varied from 0·00?±?0·00 to 1·00?±?0·08. Substantial variation existed both among BRAs and years but post-fledging survival declined substantially during the study. 6.?Pre- and post-fledging survival were positively correlated, exhibiting a quadratic relationship (?(post-fledging survival) =?1·00 (±0·47)x-0·83 (±0·480)x(2) , where x?=?pre-fledging survival). Therefore, we did not find a trade-off between pre- and post-fledging survival in black brent goslings across BRAs, suggesting that factors other than foraging conditions and predation on goslings must influence selection of BRAs.     

Fitness consequences of timing of breeding in birds: date effects in the course of a reproductive episode

In seasonal environments, avian reproductive performance almost generally declines in the course of the season. Quantifying the associated fitness consequences of timing of breeding, i.e. of date‐related factors, is important for understanding the evolution of temporal patterns in avian life‐histories and for predicting consequences of climate change. The seasonal decline can also be caused by an effect of parental quality: individuals with high phenotypic quality may breed early. The results of existing experimental studies investigating whether date or quality effects cause the seasonal decline are inconsistent, indicating that both mechanisms might be involved. However, it remains unclear to what extent the confounding effect of quality occurs and what the fitness consequences of timing per se over a whole breeding episode are. In a cross‐fostering experiment using the barn swallows’ second broods we evaluated the causes for the seasonal decline in reproductive performance for three distinct periods of a reproductive attempt, the early nestling period, the late nestling period and the post‐fledging period, and we assessed the overall fitness consequences of timing per se. A seasonal decline in juvenile feather growth rate was mainly due to date effects in the late nestling period, although we determined quality effects during early nestling development. Date effects on survival were present in the post‐fledging period, but not in the nestling period. The decline in feather length due to date effects in the nestling period accounted for 9% of the seasonal decline in post‐fledging survival, whereas date effects arising only in the post‐fledging period caused 91% of the decline. These results suggest that date effects increase in the course of a reproductive episode. Thus, the benefits of an early timing of breeding can be quantified only when considering also the post‐fledging period. We suggest that the timing of breeding evolved through a trade‐off between date‐related benefits and quality‐related costs of early breeding.     

No effect of habitat fragmentation on post-fledging, first-year and adult survival in the middle spotted woodpecker

Despite its relevance for the dynamics of populations, the ecological mechanisms underlying juvenile and adult survival are poorly known in most bird species. This study focuses on the effect of habitat fragmentation on early post-fledging, first-year and adult survival of the middle spotted woodpecker Dendrocopus medius by combining data of radio-tagged and ringed birds. Among juveniles, most deaths occurred during the first three weeks after fledging (survival rate: 0.359±0.077) and were mainly caused by predation. After independence, birds faced another critical period during their first autumn-winter that lowered first-year survival further (0.255±0.044), whereas adult mortality was considerably lower (annual survival rate: 0.786±0.074). We did not find any significant effect of habitat fragmentation (measured as patch size and connectivity) on juvenile or adult survival. Sex ratio at fledging did not differ significantly from parity (proportion of females: 0.513) and was not correlated to patch size. Regardless of age, survival did not differ between the sexes, suggesting that a female-biased mortality was not the mechanism behind the presence of unpaired territorial males in this population. Lighter nestlings underwent significantly higher post-fledging mortality, indicating that conditions in the nest may substantially affect survival later in life.     

Individual condition,but not fledging phenology,carries over to affect post-fledging survival in a Neotropical migratory songbird

Migratory animals face severe time and energy constraints during their annual cycle. These constraints may be exacerbated in young animals by conditions experienced during development that can affect both phenotype and phenology. For young migratory songbirds, the period between fledging and autumn migration, the post-fledging period, is believed to represent a time of intense selective pressure. However, there has yet to be a study that has assessed post-fledging survival for the entirety of the post-fledging period, probably due to the challenge of following juveniles as they move broadly across the landscape (tens to hundreds of kilometres). To overcome this challenge, we used an automated radiotelemetry array spanning 60 000 km² in southern Ontario, Canada, and miniature digital radiotelemetry tags to track 216 juvenile Barn Swallows Hirundo rustica continuously from fledging to migration. We hypothesized that young that fledged in better condition and earlier in the breeding season would have higher survival relative to birds fledging in poorer condition, because they have more energy to deal with resource constraints, and that early-fledging birds would depart on migration earlier than late-fledging birds because there is probably a fixed period of time required post-fledging to prepare for migration. We found that average cumulative apparent survival was 42% and that condition in the nest was a strong positive predictor of post-fledging apparent survival. We also found that birds that fledged earlier in the season departed on migration earlier in the autumn relative to late-fledging birds. Contrary to our prediction, average apparent survival was equal for early- and late-fledging birds. Our results suggest that factors during development that promote better nestling condition are critical for predicting future apparent survival prior to migration. Differences in annual apparent survival between early- and late-fledging songbirds, as commonly observed, may be driven by events occurring at later stages of the annual cycle.     

THE SELECTION OF TITS PARUS SPP. BY SPARROWHAWKS ACCIPITER NISUS

Selection of the Wytham Wood population of tits by Sparrowhawks during the tit post-fledging period was studied over three years. Within three weeks of the median date of tit fledging, juvenile Blue Tits were selected more frequently than juvenile Great Tits but thereafter juvenile Great Tits were selected more frequently; it is suggested that the former was due to greater vulnerability of Blue Tits in the two to three weeks after fledging, and the latter due to changes in availability due to immigration/emigration of juvenile tits. When juvenile tits killed by hawks were compared on the basis of brood and physical characteristics with their cohorts in the entire population and in those surviving to the following year, only one factor, fledging date, was found to have affected selection by hawks. It is believed that this was because hawks selected late fledged young still in family parties in preference to early fledged young which had already become independent of their parents and were foraging in the better cover of the forest canopy. When adults killed by hawks were compared with all adults available for selection on the basis of species, sex, timing of nesting and whether or not they had been born in the wood, the only selection trend found was that male Great Tits were taken more often than females due, possibly, to greater risk of exposure for males while foraging or during territorial behaviour. When compared with availability, juveniles were selected more frequently than adults in only one year, possibly a result of yearly differences in the number of prey available per hawk, the rate of non-predator related juvenile mortality or unknown hawk hunting strategies. The overall finding of relative non-selectivity by hawks was attributed to the surprise factor associated with the hunting methods used by Sparrowhawks in woodland.     

Growth, fledging success and post-fledging survival of juvenile Oystercatchers Haematopus ostralegus

We studied the consequences of differences in growth rate on the subsequent survival of Oystercatcher Haematopus ostralegus chicks. Fledging success increased sharply with growth rate, from zero in chicks growing at less than 6 g per day to about 85% in chicks growing at more than 10 g per day. The age at which chicks fledged varied from 27 to 52 days. Chicks which fledged at an early age displayed a much faster growth rate than later fledging chicks. Although slow growth resulted in a considerable prolongation of the period before fledging, slow-growing chicks fledged at a smaller size and with a lower body-weight than fast-growing chicks. After fledging, all chicks remained almost completely dependent on their parents up to an age of 3 months and often longer.
Almost 40% of the fledglings eventually returned to the breeding area. This figure probably reflects post-fledging survival. Age and size at fledging had no effect on a chick's probability of return. Body-weight at fledging had a small positive correlation with the return probability, but this was not statistically significant. We conclude that although slow growth severely reduces a chick's chance of fledging, it probably does not result in irreversible damage causing an increased risk of mortality during the first years after fledging. Apparently, any possible disadvantage associated with small size or low body-weight could be compensated for after fledging.     

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.     

Post-fledging interactions between the Common Cuckoo Cuculus canorus and its cavity-nesting Common Redstart Phoenicurus phoenicurus host

Brood parasite–host interactions during the incubation and nestling stages have been well studied, but the post-fledging period remains virtually unknown. Using radiotracking, we provide the first detailed data on post-fledging interactions between the Common Cuckoo Cuculus canorus and its only regular cavity-nesting host, the Common Redstart Phoenicurus phoenicurus. Cuckoos raised alone (‘solitary’) fledged at higher mass, with higher wing and tarsus length and started to fly at a younger age than Cuckoos raised alongside young Redstarts (‘mixed’). However, a further 23 fledging and post-fledging parameters measured at five pre-determined times (fledging, first-flight, predation, starvation, independence) did not differ between solitary and mixed Cuckoos. In addition, none of the parameters measured during the post-fledging period (growth, dispersal distances, number of flights) differed between solitary and mixed Cuckoos. Redstart fledglings from non-parasitized broods (‘solitary’) showed generally similar fledging and post-fledging parameters to fledglings reared alongside a Cuckoo (‘mixed’). Surprisingly, there were no significant differences in post-fledging predation rate, starvation or overall survival rates between mixed and solitary Cuckoos or mixed and solitary Redstarts. Thus, during the post-fledging period, mixed Cuckoo fledglings successfully compensated for the poorer performance experienced during the nestling stage whereas mixed and solitary Redstarts did not differ in any measured parameters. This suggests that the regular occurrence of mixed broods in this host–parasite system – which is unique among the many Cuckoo hosts – is evolutionarily stable for both hosts and parasites.     

Early life events carry over to influence pre-migratory condition in a free-living songbird

Conditions experienced during development can have long-term consequences for individual success. In migratory songbirds, the proximate mechanisms linking early life events and survival are not well understood because tracking individuals across stages of the annual cycle can be extremely challenging. In this paper, we first use a 13 year dataset to demonstrate a positive relationship between 1(st) year survival and nestling mass in migratory Savannah sparrows (Passerculus sandwichensis). We also use a brood manipulation experiment to show that nestlings from smaller broods have higher mass in the nest relative to individuals from larger broods. Having established these relationships, we then use three years of field data involving multiple captures of individuals throughout the pre-migratory period and a multi-level path model to examine the hypothesis that conditions during development limit survival during migration by affecting an individual's ability to accumulate sufficient lean tissue and fat mass prior to migration. We found a positive relationship between fat mass during the pre-migratory period (Sept-Oct) and nestling mass and a negative indirect relationship between pre-migratory fat mass and fledging date. Our results provide the first evidence that conditions during development limit survival during migration through their effect on fat stores. These results are particularly important given recent evidence showing that body condition of songbirds at fledging is affected by climate change and anthropogenic changes to landscape structure.