The impact of increased food availability on reproduction in a long-distance migratory songbird: implications for environmental change?
ABSTRACT: Many populations of migratory songbirds are declining or shifting in distribution. This is likely due to environmental changes that alter factors such as food availability that may have an impact on survival and/or breeding success. We tested the impact of experimentally supplemented food on the breeding success over three years of northern wheatears (Oenanthe oenanthe), a species in decline over much of Europe. The number of offspring fledged over the season was higher for food-supplemented birds than for control birds. The mechanisms for this effect were that food supplementation advanced breeding date, which, together with increased resources, allowed further breeding attempts. While food supplementation did not increase the clutch size, hatching success or number of chicks fledged per breeding attempt, it did increase chick size in one year of the study. The increased breeding success was greater for males than females; males could attempt to rear simultaneous broods with multiple females as well as attempting second broods, whereas females could only increase their breeding effort via second broods. Multiple brooding is rare in the study population, but this study demonstrates the potential for changes in food availability to affect wheatear breeding productivity, primarily via phenotypic flexibility in the number of breeding attempts. Our results have implications for our understanding of how wheatears may respond to natural changes in food availability due to climate changes or changes in habitat management.
Project description:In migrating animals protandry is the phenomenon whereby males of a species arrive at the breeding grounds earlier than females. In the present study we investigated the proximate causes of protandry in a migratory songbird, the northern wheatear Oenanthe oenanthe. Previous experiments with caged birds revealed that males and females show differentiated photoperiod-induced migratory habits. However, it remained open whether protandry would still occur without photoperiodic cues. In this study we kept captive first-year birds under constant photoperiod and environmental conditions in a "common garden" experiment. Male northern wheatears started their spring migratory activity earlier than females, even in the absence of environmental cues. This indicates that protandry in the northern wheatear has an endogenous basis with an innate earlier spring departure of males than females.
Project description:Quantifying among-individual variation in life-history strategies, and associated variation in reproductive performance and resulting demographic structure, is key to understanding and predicting population dynamics and life-history evolution. Partial migration, where populations comprise a mixture of resident and seasonally migrant individuals, constitutes a dimension of life-history variation that could be associated with substantial variation in reproductive performance. However, such variation has rarely been quantified due to the challenge of measuring reproduction and migration across a sufficient number of seasonally mobile males and females. We used intensive winter (non-breeding season) resightings of colour-ringed adult European shags (Phalacrocorax aristotelis) from a known breeding colony to identify resident and migrant individuals. We tested whether two aspects of annual reproductive performance, brood hatch date and breeding success, differed between resident and migrant males, females and breeding pairs observed across three consecutive winters and breeding seasons. The sex ratios of observed resident and migrant shags did not significantly differ from each other or from 1:1, suggesting that both sexes are partially migratory and that migration was not sex-biased across surveyed areas. Individual resident males and females hatched their broods 6 days earlier and fledged 0.2 more chicks per year than migrant males and females on average. Resident individuals of both sexes therefore had higher breeding success than migrants. Hatch date and breeding success also varied with a pair's joint migratory strategy such that resident-resident pairs hatched their broods 12 days earlier than migrant-migrant pairs, and fledged 0.7 more chicks per year on average. However, there was no evidence of assortative pairing with respect to migratory strategy: observed frequencies of migrant-migrant and resident-resident pairs did not differ from those expected given random pairing. These data demonstrate substantial variation in two key aspects of reproductive performance associated with the migratory strategies of males, females and breeding pairs within a partially migratory population. These patterns could reflect direct and/or indirect mechanisms, but imply that individual variation in migratory strategy and variation in pairing among residents and migrants could influence selection on migration and drive complex population and evolutionary dynamics.
Project description:Adverse weather conditions during parental care may have direct consequences for offspring production, but longer-term effects on juvenile and parental survival are less well known. We used long-term data on reproductive output, recruitment, and parental survival in northern wheatears (Oenanthe oenanthe) to investigate the effects of rainfall during parental care on fledging success, recruitment success (juvenile survival), and parental survival, and how these effects related to nestling age, breeding time, habitat quality, and parental nest visitation rates. While accounting for effects of temperature, fledging success was negatively related to rainfall (days > 10 mm) in the second half of the nestling period, with the magnitude of this effect being greater for breeding attempts early in the season. Recruitment success was, however, more sensitive to the number of rain days in the first half of the nestling period. Rainfall effects on parental survival differed between the sexes; males were more sensitive to rain during the nestling period than females. We demonstrate a probable mechanism driving the rainfall effects on reproductive output: Parental nest visitation rates decline with increasing amounts of daily rainfall, with this effect becoming stronger after consecutive rain days. Our study shows that rain during the nestling stage not only relates to fledging success but also has longer-term effects on recruitment and subsequent parental survival. Thus, if we want to understand or predict population responses to future climate change, we need to consider the potential impacts of changing rainfall patterns in addition to temperature, and how these will affect target species' vital rates.
Project description:Warmer springs may cause animals to become mistimed if advances of spring timing, including available resources and of timing of breeding occur at different speed. We used thermal sums (cumulative sum of degree days) during spring to describe the thermal progression (timing) of spring and investigate its relationship to breeding phenology and demography of a long-distant migrant bird, the northern wheatear (Oenanthe oenanthe L.). We first compare 20-year trends in spring timing, breeding time, selection for breeding time, and annual demographic rates. We then explicitly test whether annual variation in selection for breeding time and demographic rates associates with the degree of phenological matching between breeding time and thermal progression of spring. Both thermal progression of spring and breeding time of wheatears advanced in time during the study period. But despite breeding on average 7 days earlier with respect to date, wheatears bred about 4 days later with respect to thermal spring progression. Over the same time period, selection for breeding time changed from distinct within-season advantage of breeding early to no or very weak advantage. Furthermore, demographic rates (nest success, fledgling production, recruitment, adult survival) and nestling weight declined markedly by 16%-79%. Those temporal trends suggest that a reduced degree of phenological matching may affect within-season fitness advantage of early breeding and population demographic rates. In contrast, when we investigate links based on annual variation, we find no significant relationship between either demographic rates or fitness advantage of early breeding with annual variation in the degree of phenological matching. Our results show that corresponding temporal trends in phenological matching, selection for breeding time and demographic rates are inconclusive evidence for demographic effects of changed phenological matching. Instead, we suggest that the trends in selection for breeding time and demographic rates are due to a general deterioration of the breeding environment.
Project description:Environmental factors can shape reproductive investment strategies and influence the variance in male mating success. Environmental effects on extrapair paternity have traditionally been ascribed to aspects of the social environment, such as breeding density and synchrony. However, social factors are often confounded with habitat quality and are challenging to disentangle. We used both natural variation in habitat quality and a food supplementation experiment to separate the effects of food availability-one key aspect of habitat quality-on extrapair paternity (EPP) and reproductive success in the black-throated blue warbler, Setophaga caerulescens. High natural food availability was associated with higher within-pair paternity (WPP) and fledging two broods late in the breeding season, but lower EPP. Food-supplemented males had higher WPP leading to higher reproductive success relative to controls, and when in low-quality habitat, food-supplemented males were more likely to fledge two broods but less likely to gain EPP. Our results demonstrate that food availability affects trade-offs in reproductive activities. When food constraints are reduced, males invest in WPP at the expense of EPP. These findings imply that environmental change could alter how individuals allocate their resources and affect the selective environment that drives variation in male mating success.
Project description:In seasonal environments, a temporal decline in breeding performance (e.g. clutch size, nestling condition, and fledging success) of altricial bird species is a well-known phenomenon. In this study, we present the effect of laying phenology on the physiological condition of nestling great tits (Parus major) in 14 consecutive breeding seasons. We used blood haemoglobin and baseline glucose concentrations as indicators of nestling physiological condition. Nestling blood haemoglobin reflects food base quality and availability during the breeding period. Blood glucose concentration can be used as a supplementary reverse index of condition, since it is negatively related to environmental quality. It might be indicative of the stress caused by unfavourable extrinsic factors, though, due to potential confounding factors such as adverse weather conditions, low food quality, or feeding interruptions, glucose levels should be used in this ecological context with caution. Great tit nestlings from earlier broods were characterised by higher mean haemoglobin concentrations, indicating a seasonal decline in food quality and availability. The blood glucose concentration displayed an opposite pattern, with nestlings from earlier broods being characterised by lower mean concentrations than those from later broods. However, very little of the variation in blood glucose concentration can be explained by the variation in laying date, which suggests that blood glucose concentration is of little importance in the context of breeding phenology. Our results show that the physiological condition of nestlings of this species decreases as the breeding season progresses, most probably due to environmental factors.
Project description:Early arrival at the breeding site positively affects the breeding success of migratory birds. During migration, birds spend most of their time at stopovers. Therefore, determining which factors shape stopover duration is essential to our understanding of avian migration. Because the main purpose of stopover is to accumulate fat as fuel for the next flight bout, fuel reserves at arrival and the accumulation of fuel are both expected to affect stopover departure decisions. Here, we determined whether fuel reserves and fuel accumulation predict a bird's motivation to depart, as quantified by nocturnal migratory restlessness (Zugunruhe), using northern wheatears (Oenanthe oenanthe) that were captured and temporarily contained at spring stopover. We found that fuel reserves at capture were positively correlated with Zugunruhe, and negatively correlated with fuel accumulation. This indicates that fat birds were motivated to depart, whereas lean birds were set on staying and accumulating fuel. Moreover, the change in fuel reserves was positively correlated with the concurrent change in Zugunruhe, providing the first empirical evidence for a direct link between fuel accumulation and Zugunruhe during stopover. Our study indicates that, together with innate rhythms and weather, the size and accumulation of fuel reserves shape stopover duration, and hence overall migration time.
Project description:Migratory flight is physiologically highly demanding and has been shown to negatively affect multiple parameters of constitutive immune function (CIF), an animal's first line of physiological defence against infections. In between migratory flights, most birds make stopovers, periods during which they accumulate fuel for the next flight(s). Stopovers are also commonly thought of as periods of rest and recovery, but what this encompasses is largely undefined. Here, we show that during stopover, northern wheatears Oenanthe oenanthe, a long-distance migratory bird, can rapidly increase constitutive innate immune function. We caught and temporarily caged birds under ad libitum food conditions at a stopover site in autumn. Within 2 days, most birds significantly increased complement activity and their ability to kill microbes. Changes in immune function were not related to the birds' food intake or extent of fuel accumulation. Our study suggests that stopovers may not only be important to refuel but also to restore immune function. Additionally, the increase in CIF could help migrating birds to deal with novel pathogens they may encounter at stopover sites.
Project description:Several studies on birds have proposed that a lack of invertebrate prey in urbanized areas could be the main cause for generally lower levels of breeding success compared to rural habitats. Previous work on house sparrows Passer domesticus found that supplemental feeding in urbanized areas increased breeding success but did not contribute to population growth. Here, we hypothesize that supplementary feeding allows house sparrows to achieve higher breeding success but at the cost of lower nestling quality. As abundant food supplies may permit both high- and low-quality nestlings to survive, we also predict that within-brood variation in proxies of nestling quality would be larger for supplemental food broods than for unfed broods. As proxies of nestling quality, we considered feather corticosterone (CORT f), body condition (scaled mass index, SMI), and tarsus-based fluctuating asymmetry (FA). Our hypothesis was only partially supported as we did not find an overall effect of food supplementation on FA or SMI. Rather, food supplementation affected nestling phenotype only early in the breeding season in terms of elevated CORT f levels and a tendency for more variable within-brood CORT f and FA. Early food supplemented nests therefore seemed to include at least some nestlings that faced increased stressors during development, possibly due to harsher environmental (e.g., related to food and temperature) conditions early in the breeding season that would increase sibling competition, especially in larger broods. The fact that CORT f was positively, rather than inversely, related to nestling SMI further suggests that factors influencing CORT f and SMI are likely operating over different periods or, alternatively, that nestlings in good nutritional condition also invest in high-quality feathers.
Project description:We have sequenced a partial transcriptome of the Northern Wheatear (Oenanthe oenanthe), a species with one of the longest migrations on Earth. The transcriptome was constructed de novo using RNA-Seq sequence data from the pooled mRNA of six different tissues: brain, muscle, intestine, liver, adipose tissue and skin. The samples came from nine captive-bred wheatears collected at three different stages of the endogenous autumn migratory period: (1) lean birds prior the onset of migration, (2) during the fattening stage and (3) individuals at their migratory body mass plateau, when they have almost doubled their lean body mass. The sample structure used to build up the transcriptome of the Northern Wheatears concerning tissue composition and time guarantees the future survey of the regulatory genes involved in the development of the migratory phenotype. Through the pre-migratory period, birds accomplish outstanding physical and behavioural changes that involve all organ systems. Nevertheless, the molecular mechanisms through which birds synchronize and control hyperphagia, fattening, restlessness increase, immunity boosting and tuning the muscles for such endurance flight are still largely unknown. The use of RNA-Seq has emerged as a powerful tool to analyse complex traits on a broad scale, and we believe it can help to characterize the migratory phenotype of wheatears at an unprecedented level. The primary challenge to conduct quantitative transcriptomic studies in non-model species is the availability of a reference transcriptome, which we have constructed and described in this paper. The cDNA was sequenced by pyrosequencing using the Genome Sequencer Roche GS FLX System; with single paired-end reads of about 400 bp. We estimate the total number of genes at 15,640, of which 67% could be annotated using Turkey and Zebra Finch genomes, or protein sequence information from SwissProt and NCBI databases. With our study, we have made a first step towards understanding the migratory phenotype regarding gene expression of a species that has become a model to study birds long-distance migrations.