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.     

Carryover effects and climatic conditions influence the postfledging survival of greater sage‐grouse

Prebreeding survival is an important life history component that affects both parental fitness and population persistence. In birds, prebreeding can be separated into pre‐ and postfledging periods; carryover effects from the prefledging period may influence postfledging survival. We investigated effects of body condition at fledging, and climatic variation, on postfledging survival of radio‐marked greater sage‐grouse (Centrocercus urophasianus) in the Great Basin Desert of the western United States. We hypothesized that body condition would influence postfledging survival as a carryover effect from the prefledging period, and we predicted that climatic variation may mediate this carryover effect or, alternatively, would act directly on survival during the postfledging period. Individual body condition had a strong positive effect on postfledging survival of juvenile females, suggesting carryover effects from the prefledging period. Females in the upper 25th percentile of body condition scores had a postfledging survival probability more than twice that (Φ = 0.51 ± 0.06 SE) of females in the bottom 25th percentile (Φ = 0.21 ± 0.05 SE). A similar effect could not be detected for males. We also found evidence for temperature and precipitation effects on monthly survival rates of both sexes. After controlling for site‐level variation, postfledging survival was nearly twice as great following the coolest and wettest growing season (Φ = 0.77 ± 0.05 SE) compared with the hottest and driest growing season (Φ = 0.39 ± 0.05 SE). We found no relationships between individual body condition and temperature or precipitation, suggesting that carryover effects operated independently of background climatic variation. The temperature and precipitation effects we observed likely produced a direct effect on mortality risk during the postfledging period. Conservation actions that focus on improving prefledging habitat for sage‐grouse may have indirect benefits to survival during postfledging, due to carryover effects between the two life phases.     

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.     

Survival to independence in relation to pre‐fledging development and latitude in songbirds across the globe

Species differ strongly in their life histories, including the probability of survival. Annual adult survival was investigated extensively in the past, whereas juvenile survival, and especially survival to independence, received much less attention. Yet, they are critical for our understanding of population demography and life‐history evolution. We investigated post‐fledging survival to independence (i.e. survival upon leaving the nest until nutritional independence) in 74 species of passerine birds worldwide based on 100 population level estimates extracted from published literature. Our comparative analyses revealed that survival to independence increased with the length of nestling period and relative fledging mass (ratio of fledging mass to adult body mass). At the same time, species with higher nest predation rates had shorter nestling periods and lower relative fledging mass. Thus, we identify an important trade‐off in life history strategies: staying longer in the nest may improve post‐fledging survival due to enhanced flight ability and sensory functions, but at the cost of a longer exposure to nest predators and increased mortality due to nest predation. Additionally, post‐fledging survival to independence did not differ between species from the northern temperate zone vs species from the tropics and southern hemisphere. However, analyses of post‐fledging survival curves suggest that 1) daily survival rates are not constant and improve quickly upon leaving the nest, and 2) species in the tropics and southern hemisphere have higher daily post‐fledging survival rates than northern temperate species. Nevertheless, due to the accumulation of mortality risk during their much longer periods of post‐fledging care, overall survival until independence is comparable across latitudes. Obtaining high‐quality demographic data across latitudes to evaluate the generality of these findings and mechanisms underlying them should be a research priority.     

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.     

Conservation planning for freshwater–marine carryover effects on Chinook salmon survival

Experiences of migratory species in one habitat may affect their survival in the next habitat, in what is known as carryover effects. These effects are especially relevant for understanding how freshwater experience affects survival in anadromous fishes. Here, we study the carryover effects of juvenile salmon passage through a hydropower system (Snake and Columbia rivers, northwestern United States). To reduce the direct effect of hydrosystem passage on juveniles, some fishes are transported through the hydrosystem in barges, while the others are allowed to migrate in‐river. Although hydrosystem survival of transported fishes is greater than that of their run‐of‐river counterparts, their relative juvenile‐to‐adult survival (hereafter survival) can be less. We tested for carryover effects using generalized linear mixed effects models of survival with over 1 million tagged Chinook salmon, Oncorhynchus tshawytscha (Walbaum) (Salmonidae), migrating in 1999–2013. Carryover effects were identified with rear‐type (wild vs. hatchery), passage‐type (run‐of‐river vs. transported), and freshwater and marine covariates. Importantly, the Pacific Decadal Oscillation (PDO) index characterizing cool/warm (i.e., productive/nonproductive) ocean phases had a strong influence on the relative survival of rear‐ and passage‐types. Specifically, transportation benefited wild Chinook salmon more in cool PDO years, while hatchery counterparts benefited more in warm PDO years. Transportation was detrimental for wild Chinook salmon migrating early in the season, but beneficial for later season migrants. Hatchery counterparts benefited from transportation throughout the season. Altogether, wild fish could benefit from transportation approximately 2 weeks earlier during cool PDO years, with still a benefit to hatchery counterparts. Furthermore, we found some support for hypotheses related to higher survival with increased river flow, high predation in the estuary and plume areas, and faster migration and development‐related increased survival with temperature. Thus, pre‐ and within‐season information on local‐ and broad‐scale conditions across habitats can be useful for planning and implementing real‐time conservation programs.     

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.     

Relationship between pre‐ and post‐copulatory traits in Salvator rufescens (Squamata: Teiidae)

Understanding pre‐ and post‐copulatory mechanisms of sexual selection can provide insights into the evolution of male reproductive strategies. The phenotype‐linked fertility hypothesis postulates that male sperm quality and secondary sexual characteristics will positively co‐vary, whereas the sperm competition hypothesis predicts a negative association between those traits. Male reproductive traits often show variation throughout the reproductive period, suggesting that the relationship between pre‐ and post‐copulatory sexual selection may vary temporally. Here, we evaluated the relationship between secondary sexual character and sperm traits and its temporal variation in Salvator rufescens, a south American lizard. We observed a negative relationship between jaw muscle and principal piece length of sperm and a variation in the relationship between pre‐ and post‐copulatory traits throughout the reproductive period. Collectively, our results evidenced a trade‐off between pre‐ and post‐copulatory traits and a strong seasonal flexibility of male reproductive strategies in this lizard species.     

Ecosystem memory of wildfires affects resilience of boreal mixedwood biodiversity after retention harvest

The extent to which past states influence present and future ecosystem characteristics (ecosystem memory (EM)) is challenging to assess because signals of past ecological conditions fade with time. Using data about seven different taxa, we show that ecological gradients initiated by wildfires up to three centuries earlier affect biotic recovery after variable retention harvest in the boreal mixedwood forest. First, we show that fire history over the last 300 years is reflected in pre‐harvest species‐specific stand basal area (BA), with longer times since high severity fire associated with proportionally higher BA of shade‐tolerant softwood species than shade‐intolerant hardwoods. Second, using patterns in the BA of pre‐harvest tree species we link fire history to species composition of pre‐harvest assemblages of bryophytes, herbs, shrubs, regenerated trees, songbirds, spiders and carabid beetles. Finally, we use variance partitioning to compare the importance of species‐specific pre‐ versus post‐harvest BA for explaining the structure of these seven biotic assemblages two, five and ten years after harvest. We detected persistent significant effects of pre‐harvest BA in all post‐harvest biotic assemblages up to ten years after harvest. Pre‐harvest BA was more strongly associated with early post‐harvest understory plant and carabid beetle assemblages than was post‐harvest BA, but the opposite was true for spiders, songbirds and regenerated trees. EM effects were detected two, five and ten years after harvest but temporal patterns varied according to taxa. Thus, EM of fire history can persist at least ten years after variable retention harvest and such effects appear to be stronger for understory plants than for animals. We conclude that management of biological legacies to increase post‐disturbance EM will increase overall resilience and sustainability of these mixedwood forests.     

The influence of sex and body size on nestling survival and recruitment in the house sparrow

Differences in the survival rates of males and females over the period from hatching to recruitment can have important impacts on individual fitness and population demographics. However, whilst the influence of an individual's sex on nestling growth and survival has been well studied, less is known about sex‐specific survival over the period between fledging and recruitment. Here, we analyse nestling survival and recruitment in an isolated, island population of house sparrows (Passer domesticus), using data collected over a 4‐year period. Nestlings that had a greater mass at 1 day old were more likely to fledge. Recruitment was also positively associated with day 11 mass. The positive influence of nestling mass on survival to fledging also increased as brood size increased. There was no difference in the survival of male and female individuals prior to fledging. In contrast, over the period from fledging to recruitment, females had significantly less mortality than males. Recruitment was also positively associated with 11‐day‐old mass. Neither the nestling sex ratio nor the fledging sex ratio deviated from 0.5, but the sex ratio amongst recruits was female biased. Our study shows that sex can influence juvenile survival, but also shows that its effect varies between different life‐history stages; therefore, these stages should be considered separately if we want to understand at what point sex‐specific differences in juvenile survival occur. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 101 , 680–688.     

Fledgling survival increases with development time and adult survival across north and south temperate zones

Slow life histories are characterized by high adult survival and few offspring, which are thought to allow increased investment per offspring to increase juvenile survival. Consistent with this pattern, south temperate zone birds are commonly longer‐lived and have fewer young than north temperate zone species. However, comparative analyses of juvenile survival, including during the first few weeks of the post‐fledging period when most juvenile mortality occurs, are largely lacking. We combined our measurements of fledgling survival for eight passerines in South Africa with estimates from published studies of 57 north and south temperate zone songbird species to test three predictions: (1) fledgling survival increases with length of development time in the nest; (2) fledgling survival increases with adult survival and reduced brood size controlled for development time; and (3) south temperate zone species, with their higher adult survival and smaller brood sizes, exhibit higher fledgling survival than north temperate zone species controlled for development time. We found that fledgling survival was higher among south temperate zone species and generally increased with development time and adult survival within and between latitudinal regions. Clutch size did not explain additional variation, but was confounded with adult survival. Given the importance of age‐specific mortality to life history evolution, understanding the causes of these geographical patterns of mortality is important.     

Preparation for flight: pre‐fledging exercise time is correlated with growth and fledging age in burrow‐nesting seabirds

Chicks of many burrow‐nesting seabirds are known to repeatedly emerge from their nests (these trips being termed ‘excursions’) and exercise their wings prior to fledging, but this behavior is poorly documented in the literature, and thus the relationship between growth and exercise remains unclear. Here, we used infrared video cameras placed in front of streaked shearwater Calonectris leucomelas nests during the chick‐emergence period to examine correlations between chick excursions and parameters known to be important for juvenile survival after fledging. In addition, we also attached acceleration‐temperature loggers to several chicks in order to evaluate the relationship between excursion time and time spent exercising the wing muscles (i.e. flapping). Chicks that undertook longer excursions exhibited more rapid increases in wing length and larger body masses at fledging, and also fledged earlier. Correlations between fitness‐related parameters and excursion time indicate that excursions during the emergence period might offer insights into the various relationships between growth and behavior and/or the mechanisms underlying offspring survival following fledging.     

The direct regeneration hypothesis in northern forests

Question: Can the direct regeneration hypothesis (DRH) be used to predict post‐disturbance regeneration after fire, wind disturbance, and clearcutting in northern forests? Do life‐history traits such as regeneration strategy and shade tolerance influence post‐disturbance regeneration success of tree species? Location: Northern forests in North America. Methods: A meta‐analysis was conducted by collecting published data on pre‐ and post‐disturbance stand compositional characteristics in the northern forests. For each tree species, compositional difference (CD) was calculated as the difference between basal area proportions of the post‐ and pre‐disturbance stands, but for post‐disturbance stands <25 years of age, post‐disturbance proportions were calculated based on relative stem density. Results: Species response to disturbances was best explained by regeneration strategy, while disturbance type had no effect on CD. The proportion of broadleaf trees with either strong or weak vegetative reproduction ability increased after all disturbances. Serotinous species had CD values not significantly different from zero after fire, while CD for semi‐serotinous species was negative. The post‐disturbance proportions of non‐serotinous conifers decreased after all forms of disturbance. Conclusions: All disturbances promote broadleaf trees, regardless of regeneration strategy (suckering, sprouting, or seeding). The DRH is supported for conifers with serotinous cones after fire. Fire causes local extinction of non‐serotinous conifers, while wind and clearcutting only decrease the proportion of non‐serotinous conifers because of partial survival of seed sources and advanced regeneration. This study suggests that increasing stand‐replacing disturbances associated with global climate change will promote broadleaf trees in northern forests.     

Increase of feather quality during moult: a possible implication of feather deformities in the evolution of partial moult in the great tit Parus major

Here we investigate the change in feather quality during partial post‐juvenile and complete post‐breeding moult in great tit Parus major by measuring the change in the number of fault bars and feather holes on wing and tail feathers. Feathers grown during ontogeny usually are of lower quality than feathers grown following subsequent moults at independence. This is reflected by higher number of fault bars and feather holes on juveniles compared to adults. Fault bars are significantly more common on tail and proximal wing feathers than on the distal remiges, indicating a mechanism of adaptive allocation of stress induced abnormalities during ontogeny into the aerodynamically less important flight feathers. On the contrary, feather holes produced probably by chewing lice have a more uniform distribution on wing and tail feathers, which may reflect the inability of birds to control their distribution, or the weak natural selection imposed by them. The adaptive value of the differential allocation of fault bar between groups of feathers seems to be supported by the significantly higher recapture probability of those juvenile great tits which have fewer fault bars at fledging on the aerodynamically most important primaries, but not on other groups of flight feathers. The selection imposed by feather holes seems to be smaller, since except for the positive association between hatching date, brood size and the number of feather holes at fledging, great tits' survival was not affected by the number of feather holes. During post‐juvenile moult, the intensity of fault bars drops significantly through the replacement of tail feathers and tertials, resulting in disproportional reduction of the total number of fault bars on flight feathers related to the number of feathers replaced. The reduction in the number of fault bars during post‐juvenile moult associated with their adaptive allocation to proximal wing feathers and rectrices may explain the evolution of partial post‐juvenile moult in the great tit, since the quality of flight feathers can be increased significantly at a relatively small cost. Our results may explain the widespread phenomenon of partial post‐juvenile moult of flight feathers among Palearctic passerines. During the next complete post‐breeding moult, the total number of fault bars on flight feathers has remained unchanged, indicating the effectiveness of partial post‐juvenile moult in reducing the number of adaptively allocated fault bars. The number of feather holes has also decreased on groups of feathers replaced during partial post‐juvenile moult, but the reduction is proportional with the number of feathers moulted. In line with this observation, the number of feather holes is further reduced during post‐breeding moult on primaries and secondaries, resulting in an increase in feather quality of adult great tits.     

An experimental test of predator–parasite interaction in a passerine bird

Experimental studies incorporating multiple trophic levels are scarce but of increasing interest for understanding ecological communities. Here we investigated interactive effects of perceived predation risk and parasite pressure on life‐history traits in a hole‐nesting bird, and the effects of predation risk on parasite success. In a 3 × 2 experimental design we increased perceived predation risk for breeding great tits Parus major via simulations of either nest‐predators (woodpeckers) or post‐fledging predators (sparrowhawks) close to nests, and used a non‐predatory species (song thrush) as a control. Concurrently, half of the nests in each treatment were either infested with ectoparasites, or kept parasite‐free. Regarding the predation risk – parasite interaction, exposure to nest‐predators tended to lower wing and sternum growth rates of nestlings in the absence, but not the presence, of parasites. In the presence of parasites, exposure to a post‐fledging, but not to a nest‐predator, led to significantly reduced wing growth. Mass and tarsus length were not affected by predator exposure, but ectoparasites had slight positive effects on mass gain. In the last third of the nestling period, overall nestling size was significantly smaller when exposed to a post‐fledging predator than to a nest‐predator, but neither differed from the control. Parental feeding rates were not affected by the treatments, but parents became less selective towards food items under either predation risk. Hen‐flea population sizes (adult or larvae) in nests were not affected by predation risk treatment of hosts. In summary, we found some evidence for an interactive effect of predation risk and parasite pressure on nestling growth. The complexity of the interaction, combined with certain inconsistencies of the effects and potential statistical artifacts, prevent however a straightforward interpretation of the results. The insights from the study are useful for designing additional experiments to further investigate the complexity of predator–parasite interactions in wild populations.     

Facultative and non‐facultative sex ratio adjustments in a dimorphic bird species

If parental allocation to each offspring sex has the same cost/benefit ratio, Fisher's hypothesis predicts a sex ratio biased towards the cheaper sex. However, in dimorphic birds there is little evidence for this, especially at hatching. We investigated the pre‐fledgling 1) sex ratio, 2) body condition and 3) sex‐differential mortality in a population of the glossy ibis Plegadis falcinellus, in southern Spain between 2001 and 2011. We defined two age groups for the period between hatching and fledging. We also compared pre‐fledgling with the autumn sex ratio. Metabolic rates were estimated by the doubly labeled water (DLW) technique to establish that sons (the bigger sex) were 18% more energy demanding than daughters, and to compute the predicted Fisher's sex ratio (0.465). As population size increased between years, body condition decreased in both sexes, and mortality increased more for daughters than sons prior to fledging. At the same time, the proportion of males among chicks close to fledging increased (average sex ratio: 0.606) while the proportion close to hatching decreased (average sex ratio: 0.434, in line with Fisher's prediction). Furthermore, the proportions of males at fledging and the following autumn were negatively correlated across years. We suggest that, as population density increased and conditions worsened the larger sex had relatively higher survival. These differences in survival produce a shift from a facultative female‐biased sex ratio at hatching into a non‐facultative male‐biased sex ratio of fledglings. Additionally, the excess of males at fledging was counterbalanced by sex‐related dispersal during the autumn. Overall, glossy ibis sex ratio is a product of a combination of facultative and non‐facultative adjustments triggered by environmental conditions, driven by rapid population growth, and mediated by highly interrelated life‐history traits such as body condition, mortality, and dispersal.     

Radio‐transmitters do not affect seasonal productivity of female Golden‐winged Warblers

Investigating the potential effects of handling and marking techniques on study animals is important for correct interpretation of research results and to effect progress in data‐collection methods. Few investigators have compared the reproductive output of radio‐tagged and non‐radio‐tagged songbirds, and no one to date has examined the possible effect of radio‐tagging adult songbirds on the survival of their fledglings. In 2011 and 2012, we compared several parameters of reproductive output of two groups of female Golden‐winged Warblers (Vermivora chrysoptera) breeding in Minnesota, including 45 females with radio‐transmitters and 73 females we did not capture, handle, or mark. We found no difference between groups in clutch sizes, hatching success, brood sizes, length of incubation and nestling stages, fledging success, number of fledglings, or survival of fledglings to independence. Thus, radio‐tags had no measurable impact on the productivity of female Golden‐winged Warblers. Our results build upon previous studies where investigators have reported no effects of radio‐tagging on the breeding parameters of songbirds by also demonstrating no effect of radio‐tagging through the post‐fledging period and, therefore, the entire breeding season.     

Positive correlations between pre‐ and post‐copulatory sexual traits in warblers

Theoretical models predict that investment in pre‐copulatory and post‐copulatory sexually selected traits should trade‐off. At the macroevolutionary scale, the majority of studies to date have focused on male weaponry as the target of pre‐copulatory sexual selection, but the trade‐off should equally apply to traits used to attract females, such as bird song and plumage. We studied the Old World leaf warblers (Phylloscopidae), a group of socially monogamous songbirds that experience relatively high levels of sperm competition. We examined the relationships between song duration and number of elements in the song with sperm length across 21 species, and between the same song variables and combined testes mass in a subset of these species (n = 10). Across species, these song variables and testes mass/sperm length were generally positively correlated, albeit not statistically significantly so or with borderline significance. In contrast to theory, we found no evidence for negative associations between pre‐ and post‐copulatory traits. We argue that this is a consequence of males of some species investing more into overall fertilization success (i.e. the sum of pre‐ and post‐copulatory sexual selection) than males of other species, and high fertilization success is achieved through investment into both mate attraction and sperm competition.     

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.     

Sex,growth rate,rank order after brood reduction,and hatching date affect first‐year survival of long‐lived Herring Gulls

Among most species of birds, survival from hatching throughout the first year of life is generally lower than subsequent survival rates. Survival of young birds during their first year may depend on a combination of selection, learning, unpredictable resources, and environmental events (i.e., post‐fledging factors). However, knowledge about post‐fledging development in long‐lived species is usually limited due to a lengthy immature stage when individuals are generally unobservable. Therefore, pre‐fledging characteristics are often used to predict the survival of young birds. We assessed effects of nestling growth rates, hatching date, hatching asynchrony, brood size and rank order after brood reduction, and sex on first‐year survival of 137 fledglings using a mark‐resighting analysis. We found that the survival probability (Φ_1yr = 0.39) of first‐year Herring Gulls (Larus argentatus) in our study colony located at the outer port of Zeebrugge (Belgium) was lower than that of older individuals (Φ_>1yr = 0.75). All 10 models best supported by our data included nestling growth rate, suggesting that variability in first‐year survival may be linked primarily to individual variation in growth. First‐year survival was negatively correlated with hatching date and rank order after brood reduction. Hence, carry‐over effects of breeding season events such as timing of breeding, early development, and social status had an influence on survival of Herring Gulls after fledging. Furthermore, we found sex‐biased mortality in first‐year Herring Gulls, with females (Φ_1yr = 0.45) surviving better than males (Φ_1yr = 0.38). Although adult survival is generally regarded as the key parameter driving population trajectories in long‐lived species, juvenile survival has recently been acknowledged as an important source of variability in population growth rates. Thus, increasing our knowledge of factors affecting age‐specific survival rates is necessary to improve our understanding of population dynamics and ultimately life‐history variation.