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.     

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.     

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.     

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.     

Post-fledging body mass as a determinant of subadult survival in Common Terns Sterna hirundo

Once a bird has fledged it becomes hardly accessible for researchers and consequently knowledge about post-fledging ontogeny is scarce. In this study on juvenile Common Terns (Sterna hirundo) we used an automated transponder-based detection and weighing system at Banter See colony, Northern Germany, which enabled us to investigate body mass growth of post-fledglings and its consequences for their survival until first return to the natal colony when 2 years old. We analysed data from two contrasting breeding seasons, 2000 and 2001, in order to determine inter-year and inter-sex variation of post-fledging parameters assumed to potentially affect subadult survival, such as the period a juvenile is still present at colony surroundings (departure age), its fledging mass and last recorded post-fledging body mass, and hatch date. Using an information-theoretic model selection approach, neither the date of hatching nor the departure age was found to affect survival. The only predictor of survival was last post-fledging body mass whereas fledging mass itself was of minor importance. Although there was weak evidence for an interaction with year, individuals of the cohort 2000, which left the colony area on average 5 g lighter than those reared under the more favourable conditions in 2001, did not exhibit lower return probability. We suggest that under unfavourable conditions selection had eliminated weak individuals prior to fledging or during the post-fledging period. This study underlines the importance of the post-fledging period and its consequences for survival, especially in species with prolonged parental care post-fledging.     

Population decline in tree swallows ( Tachycineta bicolor ) linked to climate change and inclement weather on the breeding ground

Population decline and the threat of extinction are realities currently facing many species. Yet, in most cases, the detailed demographic data necessary to identify causes of population decline are unavailable. Using 43 years (1975−2017) of data from a box-nesting population of tree swallows (Tachycineta bicolor), we identified reduced survival of offspring as a probable demographic cause of population decline. Poor fledging success was associated with increased predation and poor weather conditions during early nestling development. Low juvenile survival and subsequent recruitment was linked to poor weather conditions during the post-fledging period and may also be linked to conditions on the wintering grounds. Regional weather conditions during critical stages of breeding (early nestling and post-fledging) have become progressively worse over the 43-year study period. None of the other factors linked to offspring survival have similarly deteriorated. Overall, our results suggest tree swallows should be added to the growing list of species challenged by climate change, and that other species of aerial insect specialists may face similar impacts of climate change.     

Models of temporal variation in questing activity in individuals of Ixodes recinus (Acari: Ixodidae)

Intensive observations of the questing activity of Ixodes ricinus ticks in the field were made to provide data on the range of durations of periods of continuous questing activity, and of the variation in questing activity between individuals. Continuous periods of questing were observed to extend to a maximum of 28 hours. Substantial variation in questing activity between individuals was observed. Models fitted to the distribution of durations of bouts of questing activity provide insights into the questing ecology of I. ricinus. Results suggest that questing duration may not be solely dependent on the state of hydration of the tick. A function fitted to the frequency distribution of the proportions of active life that individuals spend on questing, provides an empirically-based model that can be used to generate a stochastic expression of variation of questing activity in individuals in a questing population.     

Are numbers enough? Colonizer phenotype and abundance interact to affect population dynamics

1. Ecologists have long recognized that the number of colonizers entering a population can be a major driver of population dynamics, but still struggle to explain why the importance of colonizer supply varies so dramatically. While there are indications that differences in the phenotype among dispersing individuals could also be important to populations, the role of phenotypic variation relative to the number of individuals, and the extent to which they interact, remains unknown. 2. We simultaneously manipulated the phenotype (dispersal duration) and abundance of settlers of a marine bryozoan and measured subsequent population structure in the field. 3. Increases in the number of colonizing individuals increased the subsequent recruitment and biomass of populations, regardless of colonizer phenotype. However, the relationship between colonizer abundance and the subsequent reproductive yield of the population was strongly reduced in populations containing individuals that had long dispersal durations. 4. The interactive effects of colonizer phenotype and abundance on the reproductive yield of populations occurred because longer dispersal durations decreased the proportion of individuals that reproduced. In fact, populations established from a few individuals with short dispersal durations had similar reproductive yield to populations c. 30 times larger established from individuals with long dispersal durations. 5. Interactions between colonizer phenotype and abundance have important implications for predicting population dynamics beyond those previously provided by numerical abundance or recruit phenotype alone.     

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.     

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.     

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.     

Early-life reproduction is associated with increased mortality risk but enhanced lifetime fitness in pre-industrial humans

The physiology of reproductive senescence in women is well understood, but the drivers of variation in senescence rates are less so. Evolutionary theory predicts that early-life investment in reproduction should be favoured by selection at the cost of reduced survival and faster reproductive senescence. We tested this hypothesis using data collected from preindustrial Finnish church records. Reproductive success increased up to age 25 and was relatively stable until a decline from age 41. Women with higher early-life fecundity (ELF; producing more children before age 25) subsequently had higher mortality risk, but high ELF was not associated with accelerated senescence in annual breeding success. However, women with higher ELF experienced faster senescence in offspring survival. Despite these apparent costs, ELF was under positive selection: individuals with higher ELF had higher lifetime reproductive success. These results are consistent with previous observations in both humans and wild vertebrates that more births and earlier onset of reproduction are associated with reduced survival, and with evolutionary theory predicting trade-offs between early reproduction and later-life survival. The results are particularly significant given recent increases in maternal ages in many societies and the potential consequences for offspring health and fitness.     

Adaptive Plasticity in Wild Field Cricket’s Acoustic Signaling

Phenotypic plasticity can be adaptive when phenotypes are closely matched to changes in the environment. In crickets, rhythmic fluctuations in the biotic and abiotic environment regularly result in diel rhythms in density of sexually active individuals. Given that density strongly influences the intensity of sexual selection, we asked whether crickets exhibit plasticity in signaling behavior that aligns with these rhythmic fluctuations in the socio-sexual environment. We quantified the acoustic mate signaling behavior of wild-caught males of two cricket species, Gryllus veletis and G. pennsylvanicus. Crickets exhibited phenotypically plastic mate signaling behavior, with most males signaling more often and more attractively during the times of day when mating activity is highest in the wild. Most male G. pennsylvanicus chirped more often and louder, with shorter interpulse durations, pulse periods, chirp durations, and interchirp durations, and at slightly higher carrier frequencies during the time of the day that mating activity is highest in the wild. Similarly, most male G. veletis chirped more often, with more pulses per chirp, longer interpulse durations, pulse periods, and chirp durations, shorter interchirp durations, and at lower carrier frequencies during the time of peak mating activity in the wild. Among-male variation in signaling plasticity was high, with some males signaling in an apparently maladaptive manner. Body size explained some of the among-male variation in G. pennsylvanicus plasticity but not G. veletis plasticity. Overall, our findings suggest that crickets exhibit phenotypically plastic mate attraction signals that closely match the fluctuating socio-sexual context they experience.     

Variation in flight duration among individual Tetraopes beetles: Implications for studies of insect flight

Milkweed beetles, Tetraopes tetraophalmus (Forster) (Cerambycidae), were flight tested three times weekly throughout their lives. Flight durations peaked early in life and then declined rapidly with age. Significant variation existed (1) between individuals, with some flying for long periods of time, others for only a few seconds, and (2) within individuals, with some flying for long periods on some test days and very briefly or not at all on other days. Long and short fliers were indistinguishable on the basis of size, sex, or lifespan. The data show that studies of insect flight will underestimate the number of long fliers in a population by as much as 50% or more unless individuals are flight tested more than once.     

EVOLUTIONARY ENDOCRINOLOGY OF JUVENILE HORMONE ESTERASE: FUNCTIONAL RELATIONSHIP WITH WING POLYMORPHISM IN THE CRICKET,GRYLLUS FIRMUS

The existence, nature, and physiological consequences of genetic variation for juvenile hormone esterase (JHE) activity was studied in the wing-polymorphic cricket, Gryllus firmus. Hemolymph (blood) JHE activity was sixfold lower in nascent short-winged (SW) females, relative to nascent long-winged (LW) females during the last juvenile stadium (stage). Morph-associated genetic variation for JHE activity had two causes, variation in loci: (1) regulating whole-organism enzyme activity; and (2) controlling the degree to which JHE is secreted into the blood Reduced JHE activity in nascent SW-selected individuals was associated with reduced in vivo juvenile hormone catabolism. This suggests that variation in JHE activity during juvenile development may have important physiological consequences with respect to the regulation of blood levels of juvenile hormone and consequent specification of wing morph. This is the first definitive demonstration of genetic variation for hormonal metabolism in any insect and a genetic association between hormone metabolism and the subsequent expression of morphological variation (wing morph). However, we have not yet firmly established whether these associations represent causal relationships In contrast to the clear association between JHE activity and wing morph development, we observed no evidence indicating that variation in JHE activity plays any direct or indirect role in causing the dramatic differences in ovarian growth between adult wing morphs. Variation in JHE activity also does not appear to be important in coordinating the development of wing morph with the subsequent expression of reproductive differences between adult morphs. Finally genetic variation for the developmental profiles of JHE activity during juvenile and adult stages are remarkably similar in three Gryllus species. This suggests that genetic correlations between JHE activities during different periods of development, which underlie these activity profiles, have been conserved since the divergence of the three Gryllus species.     

Modeling Systematic Change in Stopover Duration Does Not Improve Bias in Trends Estimated from Migration Counts

The use of counts of unmarked migrating animals to monitor long term population trends assumes independence of daily counts and a constant rate of detection. However, migratory stopovers often last days or weeks, violating the assumption of count independence. Further, a systematic change in stopover duration will result in a change in the probability of detecting individuals once, but also in the probability of detecting individuals on more than one sampling occasion. We tested how variation in stopover duration influenced accuracy and precision of population trends by simulating migration count data with known constant rate of population change and by allowing daily probability of survival (an index of stopover duration) to remain constant, or to vary randomly, cyclically, or increase linearly over time by various levels. Using simulated datasets with a systematic increase in stopover duration, we also tested whether any resulting bias in population trend could be reduced by modeling the underlying source of variation in detection, or by subsampling data to every three or five days to reduce the incidence of recounting. Mean bias in population trend did not differ significantly from zero when stopover duration remained constant or varied randomly over time, but bias and the detection of false trends increased significantly with a systematic increase in stopover duration. Importantly, an increase in stopover duration over time resulted in a compounding effect on counts due to the increased probability of detection and of recounting on subsequent sampling occasions. Under this scenario, bias in population trend could not be modeled using a covariate for stopover duration alone. Rather, to improve inference drawn about long term population change using counts of unmarked migrants, analyses must include a covariate for stopover duration, as well as incorporate sampling modifications (e.g., subsampling) to reduce the probability that individuals will be detected on more than one occasion.     

Changes in the proportion of young birds in the hunting bag of Eurasian wigeon: long-term decline,but no association with climate

The proportion of first-year birds in annual wing samples provided by hunters has been used as a measure of breeding success in waterfowl. The proportion of first-year birds in the wing samples of Eurasian wigeon (Mareca penelope) from Denmark and the UK shows a long-term decline, probably reflecting a decrease in breeding success. However, previous studies report conflicting results in the relationship between variation in the proportion of first-year birds and variation in climatic conditions. We used wing data of hunter-shot Eurasian wigeon from Finland to study whether the proportion of first-year birds shows a similar long-term decline and whether between-year variation in the proportion of young is associated with variation in climatic conditions. We found a long-term decline in the proportion of first-year birds. The proportion of young also varied considerably between years, but this variation was not associated with weather or the climatic variables considered for the breeding and wintering periods. More research is needed concerning factors that affect long-term changes and annual variation in the proportion of young in the hunting bag and on the suitability of this index to measure productivity in ducks.     

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.     

Nestling telomere shortening,but not telomere length,reflects developmental stress and predicts survival in wild birds

Developmental stressors often have long-term fitness consequences, but linking offspring traits to fitness prospects has remained a challenge. Telomere length predicts mortality in adult birds, and may provide a link between developmental conditions and fitness prospects. Here, we examine the effects of manipulated brood size on growth, telomere dynamics and post-fledging survival in free-living jackdaws. Nestlings in enlarged broods achieved lower mass and lost 21% more telomere repeats relative to nestlings in reduced broods, showing that developmental stress accelerates telomere shortening. Adult telomere length was positively correlated with their telomere length as nestling (r = 0.83). Thus, an advantage of long telomeres in nestlings is carried through to adulthood. Nestling telomere shortening predicted post-fledging survival and recruitment independent of manipulation and fledgling mass. This effect was strong, with a threefold difference in recruitment probability over the telomere shortening range. By contrast, absolute telomere length was neither affected by brood size manipulation nor related to survival. We conclude that telomere loss, but not absolute telomere length, links developmental conditions to subsequent survival and suggest that telomere shortening may provide a key to unravelling the physiological causes of developmental effects on fitness.     

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.