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.     

Effect of age,haemosporidian infection and body condition on pair composition and reproductive success in Great Tits Parus major

Several factors shape lifetime reproductive success, including genetic background, body condition, environmental conditions and ecological interactions such as parasitism. Adults often show higher reproductive success than their young conspecifics, especially in long-lived bird species, and this may be explained by the cumulative effects of an increase in reproductive experience and the selection of high-quality individuals from one year to the next. To test whether this pattern also exists in short-lived bird species, we used 13 years of monitoring data from two Great Tit Parus major populations. The effects of male and female age on several reproductive parameters were analysed in 419 pairs of Great Tits, while accounting for body condition and infection by haemosporidian parasites. Reproductive success was mainly affected by the age-class of males. Pairs containing a sub-adult male fledged one-third fewer chicks than pairs containing an adult male. The difference was not caused by variation in male fertility but could have been caused by better parental care provided by adult birds. In addition to lower reproductive success, first-year males also had reduced access to mating compared with adult males, suggesting an avoidance of sub-adult males by females. Nestling body condition was positively correlated with parental body condition, and the body condition of male and female members of breeding pairs was positively correlated. Finally, the number of fledged chicks was mainly affected by the infection status of males. These results temper our previously published results showing an effect of infection on Great Tit reproduction regardless of their sex. In our previous study, and as in most cases, the status of the partner was not taken into account and we show here that this is essential because it can lead to a biased interpretation of the results.     

Environmental drivers of roosting body mass variation in boreal Great Tits Parus major

Small wintering passerines adaptively modulate daily body mass acquisition as part of their energy management policy. However, whether birds optimize overnight mass loss or body mass at dawn remains poorly understood. We studied environmental correlates of individual variation in body mass at dusk, overnight mass loss and body mass at dawn in a wild population of Great Tits Parus major wintering in northern Fennoscandia. Body mass at dusk, overnight mass loss and body mass at dawn were independent of prevailing conditions despite extremely low night ambient temperatures. Body mass at dusk was higher in males than in females, and decreased throughout winter and when snowfall was higher in the previous month. Overnight mass loss increased with precipitation during the previous week and tended to be higher in mid‐winter, when nights were longest. However, birds reduced overnight mass loss with higher temperatures in the previous week and higher precipitation in the previous 2 weeks. Dawn body mass was strongly correlated with dusk body mass and overnight mass loss, and showed only mild associations with weather variables once dusk mass was accounted for. Body mass in roosting boreal Great Tits seems to be constrained by recent snowfall as the winter progresses, but otherwise appears to be mostly unaffected by previous and current temperatures, suggesting a regular use of facultative hypothermia.     

Body temperature responses of Great Tits Parus major to handling in the cold

Animals typically respond to stressful stimuli such as handling by increasing core body temperature. However, small birds in cold environments have been found to decrease body temperature (T_b) when handled over longer periods, although there are no data extending beyond the actual handling event in such birds. We therefore measured both the initial T_b decrease during ringing and standardized T_b sampling, and subsequent recovery of T_b after this handling protocol in wild Great Tits Parus major roosting in nestboxes in winter. Birds reduced their T_b by 2.3 °C during c. 4 min of handling. When birds were returned to their nestboxes after handling, T_b decreased by a further 1.9 °C over c. 2 min, reaching a T_b of 34.6 °C before taking 20 min to rewarm to 2.5 °C above their initial T_b. The T_b reduction during handling could be a consequence of increased heat loss rate from disrupted plumage insulation, whereas T_b reduction after handling might reflect reduced heat production. These are important factors to consider when handling small birds in the cold.     

Nesting outcomes under anthropogenic change – effects of changing climate and nestbox provision on the reproduction of Great Tits Parus major

Global anthropogenic changes are significantly impacting the ecology and evolution of many species. Among temperate taxa, changes to reproductive phenology as a result of warming springs are apparent. However, how such responses to abiotic change interact with biotic impacts resulting from human management interventions are less clear. Here we examine the response of a range of breeding metrics (laying date, clutch size, hatching and fledging success) to interactions between climatic variables and changes in conspecific density (and hence intraspecific competition) resulting from changes in nestbox provision. Using a 37‐year dataset on the Great Tit Parus major we found little evidence for interacting effects of these two drivers. Instead we found that either climatic or competitive effects were the key influence on different metrics. Annual mean laying date substantially advanced with a warming climate, whereas clutch size, hatching success and fledging success were significantly inversely associated with conspecific density. Annual variance in clutch size and hatching success increased weakly with measures of conspecific density, but there was no association between either climatic or density measures and the annual variance in laying date or fledging success. Increasing conspecific densities therefore resulted in years with lower, but more variable, reproductive rates. These results highlight the importance of adaptive provisioning of nestboxes to enhance reproductive output of hole‐nesting birds regardless of climate‐induced phenological change. Such management is likely to be appropriate for assisting with sustaining populations responding to a changing climate.     

Feather development under environmental stress: lead exposure effects on growth patterns in Great Tits Parus major

Capsule Regrowth rate of tail feathers is more strongly affected compared to feather length and symmetry.

Aims To assess the value of avian feathers as bioindicators.

Methods The origin and persistence of fluctuating asymmetry (FA) in homologous pairs of regrowing feathers was studied in captive birds under different levels of environmental stress, respresented by exposure to lead (Pb). Homologous feathers of individually housed birds were plucked synchronously or with a delay of seven days. We measured growth rate, regeneration time, final size and FA of regrown feathers and related them to Pb stress.

Results Asymmetry decreased as feathers reached their final length. This was not due to compensatory growth but rather a consequence of the programmed growth trajectory of single feathers. Tail feathers grown under higher Pb pollution showed increased regeneration times, decreased growth rates and shorter lengths, but no changes in development times nor in FA. For differences between both (i) original and induced feathers and (ii) control and Pb treatment, effect sizes of parameters related to feather development (growth bar width, growth rate, regeneration and development time) were consistently larger than those related to the resulting phenotype (feather length and FA).

Conclusions Growth bar widths in particular provide an applicable, sensitive and reliable indicator of adverse conditions such as Pb pollution and experimental conditions. In general, phenotypic characteristics that retrospectively allow estimation of growth rates may be more suitable for monitoring environmental stress than sizes or asymmetries of full-grown traits.     

Increasing the perceived predation risk changes parental care in female but not in male Great Tits Parus major

In birds, little is known about how the presence of predators alters parental food distribution decisions among nestlings. We found that experimentally increasing perceived predation risk changed parental care in female but not in male Great Tits Parus major. Females fed the lightest and average nestlings at similar rates under control conditions when predation risk was not manipulated but ignored the lightest nestling under increased perceived predation risk. Moreover, females reduced the duration of nest visits greatly after encountering a model predator, suggesting that the perception of predators may facilitate brood reduction mechanisms.     

Provision of nestboxes raises the breeding density of Great Tits Parus major equally in coniferous and deciduous woodland

Nestbox provision is a technique used to increase nest-site availability for secondary cavity-nesting birds. However, little is known about the demographic consequences of nestbox provision in different habitat types. To assess how nestbox provision affects the density of hole-nesting birds simultaneously in two contrasting habitats, we compared the breeding density of Great Tits along transects without nestboxes with that in transects where nestboxes were provided. Although the initial density of breeders was considerably higher in the deciduous habitat than in the coniferous habitat, provision of nestboxes increased density by a similar number of additional pairs in each habitat type. Thus, the provision of nestboxes in managed coniferous forests may be as effective in increasing the breeding opportunities of cavity nesters as in deciduous stands. Moreover, previous research showed that pairs in deciduous habitat with nestboxes have consistently lower breeding success than those in coniferous habitat with nestboxes. It is possible that the addition of nestboxes in the preferred habitat increased density to such an extent that density-dependent effects became apparent.     

Great tits (Parus major) reduce body mass in response to wing area reduction: a field experiment

Flight performance is crucial in determining whether a smallbird will survive an attack by a predator. Given the importanceof body mass in determining flight performance, it has beensuggested that birds should strategically regulate body massas a response to predation risk. However, all experiments upto now have been carried out with captive birds, comparing experimental to control birds. Here we present data from thefirst experiment in the field using a within-individuals experimentaldesign. The wing area of wild great tits, Parus major, wasreduced by reversibly taping primaries five to seven. Thisallowed for the same individual to alternatively act as controlor experimental bird. Great tits reduced body mass (but not pectoral muscle width) during episodes of wing area reduction,lending support to the view that the reduction in body massexperienced by birds during molt is a strategy rather thanthe result of energetic stress. Theoretical models establishingthe different trade-offs that determine optimal body mass should therefore take into account this important life-history episode.     

Haemoglobin concentration and body condition of nestling Great Tits Parus major : a comparison of first and second broods in two contrasting seasons

The physiological condition of nestling altricial birds depends on the quantity and quality of food delivered to them by parents. One indicator of the condition of Great Tit Parus major nestlings is the haemoglobin concentration in their blood. The present study demonstrates the influence of weather conditions (temperature and rainfall) on nestling haemoglobin concentrations during two consecutive breeding seasons in two different habitat types (parkland vs. woodland) in the city of Łódź in Central Poland. This influence probably results from the effects of weather on the trophic base of the Tits. Dry, hot weather strongly affected bush and herbal foliage later in the breeding season (mid-June to mid-July) in 2006, presumably by interfering with the development of herbivorous arthropod populations. This in turn caused food shortages for second broods of Great Tits, which resulted in nestlings having low haemoglobin levels. In the following year, temperature was on average lower, and rainfall was regular but not very heavy. These conditions enabled the development of arthropod assemblages, and the trophic base for birds was much richer. Haemoglobin concentrations in the blood of nestlings from second broods were significantly higher than those of first broods and, unexpectedly, second-brood nestlings in 2007 were on average in better physiological state than first-brood nestlings in 2006 in both habitats. The relationship between haemoglobin concentration, brood category and year was very similar to that for nestling body mass. However, it was independent of both body mass and brood size. In some years and under certain conditions, second broods can be more successful than first broods.     

Genomic dissection of variation in clutch size and egg mass in a wild great tit (Parus major) population

Clutch size and egg mass are life history traits that have been extensively studied in wild bird populations, as life history theory predicts a negative trade‐off between them, either at the phenotypic or at the genetic level. Here, we analyse the genomic architecture of these heritable traits in a wild great tit (Parus major) population, using three marker‐based approaches – chromosome partitioning, quantitative trait locus (QTL) mapping and a genome‐wide association study (GWAS). The variance explained by each great tit chromosome scales with predicted chromosome size, no location in the genome contains genome‐wide significant QTL, and no individual SNPs are associated with a large proportion of phenotypic variation, all of which may suggest that variation in both traits is due to many loci of small effect, located across the genome. There is no evidence that any regions of the genome contribute significantly to both traits, which combined with a small, nonsignificant negative genetic covariance between the traits, suggests the absence of genetic constraints on the independent evolution of these traits. Our findings support the hypothesis that variation in life history traits in natural populations is likely to be determined by many loci of small effect spread throughout the genome, which are subject to continued input of variation by mutation and migration, although we cannot exclude the possibility of an additional input of major effect genes influencing either trait.     

Relationships among oxidative status,breeding conditions and life‐history traits in free‐living Great Tits Parus major and Common Starlings Sturnus vulgaris

The interest shown by ecologists in antioxidants and oxidative stress as potential modulators of life‐history trade‐offs has expanded greatly in recent years. However, we still know very little about natural variation in oxidative damage and antioxidant capacity in free‐living animals. In this study, we describe the natural variation in three components of oxidative balance in nestlings and breeding females in free‐living Great Tits Parus major and Common Starlings Sturnus vulgaris in central Italy, and relate these to breeding conditions and life‐history traits. Our results suggest that there are associations among oxidative physiology, reproductive activity, growth pattern and season in wild birds, but that the nature of these associations might be species‐specific rather than general across species.     

Daily rhythms in activity and mRNA abundance of enzymes involved in glucose and lipid metabolism in liver of rainbow trout,Oncorhynchus mykiss. Influence of light and food availability

ABSTRACT

We have investigated the magnitude of diurnal variation in back squat and bench press performance using the MuscleLab force velocity transducer. Thirty resistance-trained males (mean ± SD: age 21.7 ± 1.4 years; body mass 80.5 ± 4.5 kg; height 1.79 ± 0.06 m) underwent two sessions at different times of day: morning (M, 07:30 h) and evening (E, 17:30 h). Each session included a period when rectal temperature (T_rec) was measured at rest, a 5-min standardized 150 W warm-up on a cycle ergometer, then defined programme of bench press (at 20, 40 and 60 kg) and back squat (at 30, 50 and 70 kg) exercises. A linear encoder was attached to an Olympic bar used for the exercises and average force (AF), peak velocity (PV) and time-to-peak velocity (tPV) were measured (MuscleLab software; MuscleLab Technology, Langesund, Norway) during the concentric phase of the movements. Values for T_rec at rest were higher in the evening compared to morning values (0.48°C, P < 0.0005). Daily variations were apparent for both bench press and back squat performance for AF (1.9 and 2.5%), PV (8.3 and 12.7%) and tPV (?16.6 and ?9.8%; where a negative number indicates a decrease in the variable from morning to evening). There was a main effect for load where AF and tPV increased and PV decreased from the lightest load to the heaviest for both bench press and back squat (47.1 and 80.2%; 31.7 and 57.7%; ?42.1 and ?73.9%; P < 0.0005 where a negative number indicates a decrease in the variable with increasing load). An interaction was found only for tPV, such that the tPV occurs earlier in the evening than the morning at the highest loads (60 and 70 kg) for both bench press and back squat, respectively (mean difference of 0.32 and 0.62 s). In summary, diurnal variation in back squat and bench press was shown; and the tPV in complex multi-joint movements occurs earlier during the concentric phase of exercise when back squat or bench press is performed in the evening compared to the morning. This difference can be detected using a low cost, portable and widely available commercial instrument and enables translation of past laboratory/tightly controlled experimental research in to main-stream coaching practice.     

Early survival and growth in populations of grayling with recent common ancestors—field experiments

Among eight populations of grayling Thymallus thymallus that shared common ancestors 8–28 generations earlier, mean egg mortality ranged between 1·2 and 59·8%; mortality during yolk sac absorption was low (0middot;0–8middot;0%) and mean survival to swim-up was high (90–97%), with two exceptions (20% and 50%). Survival probabilities differed significantly among populations, even after statistically adjusting for size at swim-up and water depth, water velocity, gravel size and temperature. Development time from fertilization to first observed swim-up was significantly different among populations, ranging between 27 to 40 days, or 264 to 280 degree-days. Mean specific growth rates during the first external feeding period varied between 1·2 and 2·0% day^-1 among populations. Population-specific growth rates were independent of temperature. Rates of divergence were of the same order as in other salmonid studies showing rapid evolution. The phenotypic differences may be due partly to genetic differentiation among the populations.     

Long-term evaluation of body mass at weaning and postweaning survival rates of Weddell seals in Erebus Bay, Antarctica

Variability in juvenile survival rate is expected to be an important component of the dynamics of long-lived animal populations. Across a range of species, individual variation in juvenile body mass has been shown to be an important cause of variation in fates of juveniles. Our goal in this paper was to estimate age-specific apparent survival rates for Weddell seals ( Leptonychotes weddellii ) in Erebus Bay, Antarctica, and to investigate hypotheses about relationships between body mass at weaning and apparent survival rate for juveniles. Mark–resighting models found average apparent juvenile survival rate (survival from weaning to age 3) was similar between males and females, and revealed positive relationships between body mass at weaning and apparent juvenile survival rate. The effects of body mass on juvenile survival rate differed between the sexes and the relationship between body mass and survival rate was stronger in males. These results indicate that the magnitude of energy transferred from mother to pup during lactation likely has important consequences on offspring survival rate and maternal fitness.     

Sources of (co)variation in alternative siring routes available to male great tits (Parus major)

Males of socially monogamous species can increase their siring success via within‐pair and extra‐pair fertilizations. In this study, we focused on the different sources of (co)variation between these siring routes, and asked how each contributes to total siring success. We quantified the fertilization routes to siring success, as well as behaviors that have been hypothesized to affect siring success, over a five‐year period for a wild population of great tits Parus major. We considered siring success and its fertilization routes as “interactive phenotypes” arising from phenotypic contributions of both members of the social pair. We show that siring success is strongly affected by the fecundity of the social (female) partner. We also demonstrate that a strong positive correlation between extra‐pair fertilization success and paternity loss likely constrains the evolution of these two routes. Moreover, we show that more explorative and aggressive males had less extra‐pair fertilizations, whereas more explorative females laid larger clutches. This study thus demonstrates that (co)variation in siring routes is caused by multiple factors not necessarily related to characteristics of males. We thereby highlight the importance of acknowledging the multilevel structure of male fertilization routes when studying the evolution of male mating strategies.     

Opposing effects on glutathione and reactive oxygen metabolites of sex,habitat, and spring date,but no effect of increased breeding density in great tits (Parus major)

Oxidative stress (i.e., more oxidants than antioxidants) has been proposed as a proximate currency in life‐history trade‐offs, which if studied in an ecological setting allow a more realistic perspective on the origin and evolution of trade‐offs. Therefore, the aim here was to investigate the impact of ecological and individual factors for variation in markers of oxidative stress using both experimental and correlational data. Total glutathione (tGSH), oxidized glutathione (GSSG), plasma antioxidant capacity (OXY), and plasma‐reactive oxygen metabolites (ROM) were measured in more than 700 breeding great tits (Parus major). The main results revealed a pronounced sex difference, with females having lower ROM and OXY, but higher tGSH compared with males. In addition, birds breeding in the evergreen areas had higher tGSH compared with those in the deciduous habitat, but the experimentally manipulated breeding density had no significant effect on any of the redox markers. Independent of the sex differences, the larger the reproductive investment the lower the ROM of both males and females. Taken together, the extracellular markers – ROM and OXY – revealed similar results and were highly correlated. Interestingly, the direction of their effects was in the opposite direction to the endogenously synthesized tGSH and GSSG. This highlights the need to combine extracellular markers with endogenously synthesized antioxidants to understand its implications for the origin and evolution of trade‐offs in an ecological setting.