Winter-ground microhabitat use by differently coloured phenotypes affects return rate in a long-distance migratory bird.

Migratory bird populations are declining globally at alarming rates. Non-breeding site conditions affect breeding populations, but generalising non-breeding habitat conditions over large spatial regions cannot address potential fine-scale differences across landscapes or local populations. Plumage characteristics can mediate the effects of environmental conditions on individual fitness. However, whether different phenotypes use distinctive non-breeding sites, and whether they respond to non-breeding site conditions differently remains largely unknown. Stable isotopes (δ13C, δ15N, δ2H) of inert tissues are useful to infer habitat characteristics and geographic origins where those tissues were grown. We collected winter-grown feathers from pied flycatchers (Ficedula hypoleuca) on their breeding grounds over several years from males whose dorsal plumage colouration ranged continuously from brown to black and assessed their stable isotope values as proxies of local habitat conditions. Based on feather δ2H profiles we found that browner males spent their non-breeding season in drier habitats than black males. Assignment to origin analysis shows potential regional non-breeding ground separation between differently coloured males. High within-individual repeatability of both δ13C and δ15N indicate the pied flycatcher males return yearly to similar areas. Blacker males were more likely to return to the breeding grounds after dry years compared with brown males. The opposite was found in wet years. Our study demonstrates that different phenotypes are exposed to different non-breeding site conditions which can differentially affect individual survivorship. This has important ramifications for population dynamics under predicted climate change scenarios where especially brown phenotype pied flycatcher males may be under a risk of decreasing.

Optimised PCR assays for detecting elusive waterfowl from environmental DNA.

For many aquatic and semiaquatic mammal, amphibian and fish species, environmental DNA (eDNA) methods are employed to detect species distribution and to monitor their presence, but eDNA is much less employed for avian species. Here, we developed primers for the detection of true geese and swan species using eDNA and optimised a PCR protocol for eDNA. We selected taiga bean goose (Anser fabalis fabalis) as our focal (sub)species and sampled water from lakes, from which the presence of taiga bean goose was visually confirmed. To test, if taiga bean goose DNA could be detected among DNA of other goose species, we similarly sampled eDNA from a zoo pond housing several Anatidae species. We were able to detect taiga bean goose DNA in all but one of the tested lakes, including the zoo pond. The primers developed are not species-specific, but rather specific to the genus Anser, due to the close relatedness of Anser species, which prevented the development of species-specific primers and the use of, for example, quantitative PCR. We also developed eDNA primers for Branta species and Cygnus species and tested these primers using the same samples. Canada goose (B. canadensis) and barnacle goose (B. leucopsis) DNA were only detected in the zoo pond (in which they were present), as the sampled natural lakes fall outside the range of these species. We detected whooper swan (C. cygnus) DNA in three lakes and the zoo pond (in which the species was present). The eDNA method presented here provides a potential means to monitor elusive goose species and to study the co-occurrence of large waterfowl.

Anthropogenic bottom-up and top-down impacts on boreal breeding waterbirds.

Wetland habitats are changing under multiple anthropogenic pressures. Nutrient leakage and pollution modify physico-chemical state of wetlands and affect the ecosystem through bottom-up processes, while alien predators affect the ecosystems in a top-down manner. Boreal wetlands are important breeding areas for several waterbird species, the abundances of which potentially reflect both bottom-up and top-down ecosystem processes. Here, we use long-term national monitoring data gathered from c. 130 waterbird breeding sites in Finland from the 1980s to the 2020s. We hypothesised that the physico-chemical state of the waters and increasing alien predator abundance both play a role in steering the waterbird population trends. We set out to test this hypothesis by relating population changes of 17 waterbird species to changes in water chemistry and to regional alien predator indices while allowing species-specific effects to vary with foraging niche (dabblers, invertivore divers, piscivorous divers, herbivores), nesting site, female mass and habitat (oligotrophic, eutrophic). We found niche and nesting site-specific, habitat-dependent changes in waterbird numbers. While the associations with higher phosphorus levels and browning water were in overall positive at the oligotrophic lakes, the numbers of invertivore and piscivore diving ducks were most strongly negatively associated with higher phosphorus levels and browning water at the eutrophic lakes. Furthermore, increased pH levels benefitted piscivores. Invertivore diving duck species nesting on the wetlands had declined most on sites with high alien predator indices. Large herbivorous species and species preferring oligotrophic lakes seem to be successful. We conclude that the large-scale breeding waterbird decline in Finland is closely connected to both bottom-up and top-down processes, where negative associations are emphasised especially at eutrophic lakes. Niche-, nest site- and habitat-specific management actions are required to conserve declining waterbird populations. Managing wetlands on catchments level together with alien predator control may provide important approaches to future wetland management.

Large-scale genotypic identification reveals density-dependent natal dispersal patterns in an elusive bird of prey.

BACKGROUND: Natal dispersal, the distance between site of birth and site of first breeding, has a fundamental role in population dynamics and species' responses to environmental changes. Population density is considered a key driver of natal dispersal. However, few studies have been able to examine densities at both the natal and the settlement site, which is critical for understanding the role of density in dispersal. Additionally, the role of density on natal dispersal remains poorly understood in long-lived and slowly reproducing species, due to their prolonged dispersal periods and often elusive nature. We studied the natal dispersal of the white-tailed eagle (Haliaeetus albicilla) in response to local breeder densities. We investigated the effects of the number of active territories around the natal site on (a) natal dispersal distance and (b) the difference between natal and settlement site breeder density. We were interested in whether eagles showed tendencies of conspecific attraction (positive density-dependence) or intraspecific competition (negative density-dependence) and how this related to settlement site breeder density. METHODS: We used a combination of long-term visual and genotypic identification to match individuals from their breeding site to their natal nest. We identified natal dispersal events for 355 individuals hatched between 1984 and 2015 in the Baltic Sea coast and Arctic areas of Finland. Of those, 251 were identified by their genotype. RESULTS: Individuals born in high-density areas dispersed shorter distances than those born in low-density areas, but settled at lower density breeding sites in comparison to their natal site. Eagles born in low natal area densities dispersed farther but settled in higher density breeding sites compared to their natal site. CONCLUSIONS: We show that eagles might be attracted by conspecifics (positive density-dependence) to identify high-quality habitats or find mates, but do not settle in the most densely populated areas. This indicates that natal dispersal is affected by an interplay of conspecific attraction and intraspecific competition, which has implications for population dynamics of white-tailed eagles, but also other top predators. Furthermore, our study demonstrates the value of long-term collection of both nestling and (non-invasive) adult DNA samples, and thereafter using genotype matching to identify individuals in long-lived and elusive species.

A gradual migratory divide determines not only the direction of migration but also migration strategy of a social migrant bird.

Migratory divides separate populations of migratory animals, facilitating the evolution of intraspecific differences in migration strategies. Migration strategies are expected to be different for birds using different flyways and environments, but the knowledge regarding the impact of the flyway on individual migration strategies is scarce. By using satellite tracking and neckband resightings, we reveal the existence and structure of a gradual migratory divide between two European flyway populations of greylag geese Anser anser. Birds breeding at the far end of the Gulf of Bothnia in the Baltic Sea coast use the Western Flyway, those breeding in the Gulf of Finland the Central Flyway and those breeding between these extremes scatter to the two flyways. By using Gaussian process modelling, we show that migration strategies differed between the flyways. The birds using the Western Flyway migrated earlier in autumn, performed longer annual migration and made a clear stopover during migration, whereas the birds using the Central Flyway flew directly to their wintering sites. The gradual migratory divide that also divides migration strategies provides insights into migratory divides on birds with learned migration. Distinct migration strategies in different flyways provide exciting possibilities to further study the factors driving migration strategies.

Early-life diet specificity is associated with long-lasting differences in apparent survival in a generalist predator.

Early-life conditions can have long-term fitness consequences. However, it is still unclear what optimal rearing conditions are, especially for long-lived carnivores. A more diverse diet ('balanced diet') might optimize nutrient availability and allow young to make experiences with a larger diversity of prey, whereas a narrow diet breadth ('specialized diet') might result in overall higher energy net gain. A diet that is dominated by a specific prey type (i.e. fish, 'prey type hypothesis') might be beneficial or detrimental, depending for example, on its toxicity or contaminant load. Generalist predators such as the white-tailed eagle Haliaeetus albicilla provide an interesting possibility to examine the relationship between early life diet and long-term offspring survival. In the Åland Islands, an archipelago in the Baltic Sea, white-tailed eagles live in various coastal habitats and feed on highly variable proportions of birds and fish. We use data from 21,116 prey individuals that were collected from 120 territories during the annual surveys, to examine how early-life diet is associated with apparent annual survival of 574 ringed and molecular-sexed eaglets. We supplement this analysis by assessing the relationships between diet, reproductive performance and nestling physical condition, to consider whether they are confounding with possible long-term associations. We find that early-life diet is associated with long-term fitness: Nestlings that are fed a diverse diet are in lower physical condition but have higher survival rates. Eagles that are fed more fish as nestlings have lower survival as breeding-age adults, but territories associated with fish-rich diets have higher breeding success. Our results show that young carnivores benefit from a high diversity of prey in their natal territory, either through a nutritional or learning benefit, explaining the higher survival rates. The strong relationship between early-life diet and adult survival suggests that early life shapes adult foraging decisions and that eating fish is associated with high costs. This could be due to high levels of contaminants or high competition for fish-rich territories. Long-lasting consequences of early-life diet are likely not only limited to individual-level consequences but have the potential to drive eco-evolutionary dynamics in this population.

Species-specific song responses emerge as a by-product of tuning to the local dialect.

Oscine birds preferentially respond to certain sounds over others from an early age, which focuses subsequent learning onto sexually relevant songs.1,2,3 Songs vary both across species and, due to cultural evolution, among populations of the same species. As a result, early song responses are expected to be shaped by selection both to avoid the fitness costs of cross-species learning4 and to promote learning of population-typical songs.5 These sources of selection are not mutually exclusive but can result in distinct geographic patterns of song responses in juvenile birds: if the risks of interspecific mating are the main driver of early song discrimination, then discrimination should be strongest where closely related species co-occur.4 In contrast, if early discrimination primarily facilitates learning local songs, then it should be tuned to songs typical of the local dialect.5,6,7 Here, we experimentally assess the drivers of song discrimination in nestling pied flycatchers (Ficedula hypoleuca). We first demonstrate that early discrimination against the songs of the closely related collared flycatcher (F. albicollis) is not strongly affected by co-occurrence. Second, across six European populations, we show that nestlings' early song responses are tuned to their local song dialect and that responses to the songs of collared flycatchers are similarly weak as to those of other conspecific dialects. Taken together, these findings provide clear experimental support for the hypothesis that cultural evolution, in conjunction with associated learning predispositions, drives the emergence of pre-mating reproductive barriers.

Trophic Dynamics of Mercury in the Baltic Archipelago Sea Food Web: The Impact of Ecological and Ecophysiological Traits.

We investigated trophic dynamics of Hg in the polluted Baltic Archipelago Sea using established trophic magnification (TMFs) and biomagnification factors (BMFs) on a comprehensive set of bird, fish, and invertebrate species. As different ecological and ecophysiological species traits may affect trophic dynamics, we explored the effect of food chain (benthic, pelagic, benthopelagic) and thermoregulatory strategy on trophic total Hg (THg) dynamics, using different approaches to accommodate benthopelagic species and normalize for trophic position (TP). We observed TMFs and most BMFs greater than 1, indicating overall THg biomagnification. We found significantly higher pelagic TMFs (3.58-4.02) compared to benthic ones (2.11-2.34) when the homeotherm bird species were excluded from models, but not when included. This difference between the benthic and pelagic TMFs remained regardless of how the TP of benthopelagic species was modeled, or whether TMFs were normalized for TP or not. TP-corrected BMFs showed a larger range (0.44-508) compared to BMFs representing predator-prey concentration ratios (0.05-82.2). Overall, the present study shows the importance of including and evaluating the effect of ecological and ecophysiological traits when investigating trophic contaminant dynamics.

Maternally transferred thyroid hormones and life-history variation in birds.

In vertebrates, thyroid hormones (THs) play an important role in the regulation of growth, development, metabolism, photoperiodic responses and migration. Maternally transferred THs are important for normal early phase embryonic development when embryos are not able to produce endogenous THs. Previous studies have shown that variation in maternal THs within the physiological range can influence offspring phenotype. Given the essential functions of maternal THs in development and metabolism, THs may be a mediator of life-history variation across species. We tested the hypothesis that differences in life histories are associated with differences in maternal TH transfer across species. Using birds as a model, we specifically tested whether maternally transferred yolk THs covary with migratory status, developmental mode and traits related to pace-of-life (e.g. basal metabolic rate, maximum life span). We collected un-incubated eggs (n = 1-21 eggs per species, median = 7) from 34 wild and captive bird species across 17 families and six orders to measure yolk THs [both triiodothyronine (T3) and thyroxine (T4)], compiled life-history trait data from the literature and used Bayesian phylogenetic mixed models to test our hypotheses. Our models indicated that both concentrations and total amounts of the two main forms of THs (T3 and T4) were higher in the eggs of migratory species compared to resident species, and total amounts were higher in the eggs of precocial species, which have longer prenatal developmental periods, than in those of altricial species. However, maternal yolk THs did not show clear associations with pace-of-life-related traits, such as fecundity, basal metabolic rate or maximum life span. We quantified interspecific variation in maternal yolk THs in birds, and our findings suggest higher maternal TH transfer is associated with the precocial mode of development and migratory status. Whether maternal THs represent a part of the mechanism underlying the evolution of precocial development and migration or a consequence of such life histories is currently unclear. We therefore encourage further studies to explore the physiological mechanisms and evolutionary processes underlying these patterns.

Within-individual repeatability in telomere length: A meta-analysis in nonmammalian vertebrates.

Telomere length is increasingly used as a biomarker of long-term somatic state and future survival prospects. While most studies have overlooked this aspect, biological interpretations based on a given telomere length will benefit from considering the level of within-individual repeatability of telomere length through time. Therefore, we conducted a meta-analysis on 74 longitudinal studies in nonmammalian vertebrates, with the aim to establish the current pattern of within-individual repeatability in telomere length and to identify the methodological (e.g., qPCR/TRF) and biological factors (e.g., age class, phylogeny) that may affect it. While the median within-individual repeatability of telomere length was moderate to high (R = 0.55; 95% CI: 0.05-0.95; N = 82), marked heterogeneity between studies was evident. Measurement method affected the repeatability estimate strongly, with TRF-based studies exhibiting high repeatability (R = 0.80; 95% CI: 0.34-0.96; N = 25), while repeatability of qPCR-based studies was markedly lower and more variable (R = 0.46; 95% CI: 0.04-0.82; N = 57). While phylogeny explained some variance in repeatability, phylogenetic signal was not significant (λ = 0.32; 95% CI: 0.00-0.83). None of the biological factors investigated here significantly explained variation in the repeatability of telomere length, being potentially obscured by methodological differences. Our meta-analysis highlights the high variability in within-individual repeatability estimates between studies and the need to put more effort into separating technical and biological explanations. This is important to better understand to what extent biological factors can affect the repeatability of telomere length and thus the interpretation of telomere length data.

Population differences in the length and early-life dynamics of telomeres among European pied flycatchers.

Telomere length and shortening rate are increasingly being used as biomarkers for long-term costs in ecological and evolutionary studies because of their relationships with survival and fitness. Both early-life conditions and growth, and later-life stressors can create variation in telomere shortening rate. Studies on between-population telomere length and dynamics are scarce, despite the expectation that populations exposed to varying environmental constraints would present divergent telomere length patterns. The pied flycatcher (Ficedula hypoleuca) is a passerine bird breeding across Eurasia (from Spain to western Siberia) and migrating through the Iberian Peninsula to spend the nonbreeding period in sub-Saharan Africa. Thus, different populations show marked differences in migration distance. We studied the large-scale variation of telomere length and early-life dynamics in the pied flycatcher by comparing six European populations across a north-south gradient (Finland, Estonia, England and Spain) predicting a negative effect of migration distance on adult telomere length, and of nestling growth on nestling telomere dynamics. There were clear population differences in telomere length, with English birds from midlatitudes having the longest telomeres. Telomere length did not thus show consistent latitudinal variation and was not linearly linked to differences in migration distance. Early-life telomere shortening rate tended to vary between populations. Fast growth was associated with shorter telomeres in the early life, but faster nestling growth affected telomeres more negatively in northern than southern populations. While the sources of between-population differences in telomere-related biology remain to be more intensively studied, our study illustrates the need to expand telomere studies at the between-population level.

Major population splits coincide with episodes of rapid climate change in a forest-dependent bird.

Climate change influences population demography by altering patterns of gene flow and reproductive isolation. Direct mutation rates offer the possibility for accurate dating on the within-species level but are currently only available for a handful of vertebrate species. Here, we use the first directly estimated mutation rate in birds to study the evolutionary history of pied flycatchers (Ficedula hypoleuca). Using a combination of demographic inference and species distribution modelling, we show that all major population splits in this forest-dependent system occurred during periods of increased climate instability and rapid global temperature change. We show that the divergent Spanish subspecies originated during the Eemian-Weichselian transition 115-104 thousand years ago (kya), and not during the last glacial maximum (26.5-19 kya), as previously suggested. The magnitude and rates of climate change during the glacial-interglacial transitions that preceded population splits in pied flycatchers were similar to, or exceeded, those predicted to occur in the course of the current, human-induced climate crisis. As such, our results provide a timely reminder of the strong impact that episodes of climate instability and rapid temperature changes can have on species' evolutionary trajectories, with important implications for the natural world in the Anthropocene.

Birds of three worlds: moult migration to high Arctic expands a boreal-temperate flyway to a third biome.

BACKGROUND: Knowledge on migration patterns and flyways is a key for understanding the dynamics of migratory populations and evolution of migratory behaviour. Bird migration is usually considered to be movements between breeding and wintering areas, while less attention has been paid to other long-distance movements such as moult migration. METHODS: We use high-resolution satellite-tracking data from 58 taiga bean geese Anser fabalis fabalis from the years 2019-2020, to study their moult migration during breeding season. We show the moulting sites, estimate the migratory connectivity between the breeding and the moulting sites, and estimate the utilization distributions during moult. We reveal migration routes and compare the length and timing of migration between moult migrants and successful breeders. RESULTS: All satellite-tracked non-breeding and unsuccessfully breeding taiga bean geese migrated annually to the island of Novaya Zemlya in the high Arctic for wing moult, meaning that a large part of the population gathers at the moulting sites outside the breeding range annually for approximately three months. Migratory connectivity between breeding and moulting sites was very low (rm = - 0.001, 95% CI - 0.1562-0.2897), indicating that individuals from different breeding grounds mix with each other on the moulting sites. Moult migrants began fall migration later in autumn than successful breeders, and their overall annual migration distance was over twofold compared to the successful breeders. CONCLUSIONS: Regular moult migration makes the Arctic an equally relevant habitat for the taiga bean goose population as their boreal breeding and temperate wintering grounds, and links ecological communities in these biomes. Moult migration plays an important role in the movement patterns and spatio-temporal distribution of the population. Low migratory connectivity between breeding and moulting sites can potentially contribute to the gene flow within the population. Moult migration to the high Arctic exposes the population to the rapid impacts of global warming to Arctic ecosystems. Additionally, Novaya Zemlya holds radioactive contaminants from various sources, which might still pose a threat to moult migrants. Generally, these results show that moult migration may essentially contribute to the way we should consider bird migration and migratory flyways.

Spatial and dietary sources of elevated mercury exposure in white-tailed eagle nestlings in an Arctic freshwater environment.

Human-induced mercury (Hg) contamination is of global concern and its effects on wildlife remain of high concern, especially in environmental hotspots such as inland aquatic ecosystems. Mercury biomagnifies through the food web resulting in high exposure in apex predators, such as the white-tailed eagle (Haliaeetus albicilla), making them excellent sentinel species for environmental Hg contamination. An expanding population of white-tailed eagles is inhabiting a sparsely populated inland area in Lapland, northern Finland, mainly around two large reservoirs flooded 50 years ago. As previous preliminary work revealed elevated Hg levels in this population, we measured Hg exposure along with dietary proxies (δ13C and δ15N) in body feathers collected from white-tailed eagle nestlings in this area between 2007 and 2018. Mercury concentrations were investigated in relation to territory characteristics, proximity to the reservoirs and dietary ecology as potential driving factors of Hg contamination. Mercury concentrations in the nestlings (4.97-31.02 µg g-1 dw) were elevated, compared to earlier reported values in nestlings from the Finnish Baltic coast, and exceeded normal background levels (≤5.00 µg g-1) while remaining below the tentative threshold of elevated risk for Hg exposure mediated health effect (>40.00 µg g-1). The main drivers of Hg contamination were trophic position (proxied by δ15N), the dietary proportion of the predatory fish pike (Esox lucius), and the vicinity to the Porttipahta reservoir. We also identified a potential evolutionary trap, as increased intake of the preferred prey, pike, increases exposure. All in all, we present results for poorly understood freshwater lake environments and show that more efforts should be dedicated to further unravel potentially complex pathways of Hg exposure to wildlife.

Identifying the paths of climate effects on population dynamics: dynamic and multilevel structural equation model around the annual cycle.

How environmental factors influence population dynamics in long-distance migrants is complicated by the spatiotemporal diversity of the environment the individuals experience during the annual cycle. The effects of weather on several different aspects of life history have been well studied, but a better understanding is needed on how weather affects population dynamics through the different associated traits. We utilise 77 years of data from pied flycatcher (Ficedula hypoleuca), to identify the most relevant climate signals associated with population growth rate. The strongest signals on population growth were observed from climate during periods when the birds were not present in the focal location. The population decline was associated with increasing precipitation in the African non-breeding quarters in the autumn (near the arrival of migrants) and with increasing winter temperature along the migration route (before migration). The number of fledglings was associated positively with increasing winter temperature in non-breeding area and negatively with increasing winter temperature in Europe. These possible carry-over effects did not arise via timing of breeding or clutch size but the exact mechanism remains to be revealed in future studies. High population density and low fledgling production were the intrinsic factors reducing the breeding population. We conclude that weather during all seasons has the potential to affect the reproductive success or population growth rate of this species. Our results show how weather can influence the population dynamics of a migratory species through multiple pathways, even at times of the annual cycle when the birds are in a different location than the climate signal.

Interplays between pre- and post-natal environments affect early-life mortality, body mass and telomere dynamics in the wild.

Early-life conditions are crucial determinants of phenotype and fitness. The effects of pre- and post-natal conditions on fitness prospects have been widely studied but their interactive effects have received less attention. In birds, asynchronous hatching creates challenging developmental conditions for the last-hatched chicks, but differential allocation in last-laid eggs might help to compensate this initial handicap. The relative importance and potential interaction between pre- and post-hatching developmental conditions for different fitness components remains mostly unknown. We manipulated hatching order in wild pied flycatchers (Ficedula hypoleuca), creating three groups: natural asynchrony (last-laid eggs hatching last), reversed asynchrony (last-laid eggs hatching first) and hatching synchrony (all eggs hatching at once). We examined the effects of these manipulations on early-life survival, growth and telomere length, a potential cellular biomarker of fitness prospects. Mortality was mostly affected by hatching order, with last-hatched chicks being more likely to die. Early-life telomere dynamics and growth were influenced by the interplays between laying and hatching order. Last-laid but first-hatched chicks were heavier but had shorter telomeres 5 days after hatching than their siblings, indicating rapid early growth with potential adverse consequences on telomere length. Synchronous chicks did not suffer any apparent cost of hatching synchronously. Impaired phenotypes only occurred when reversing the natural hatching order (i.e. developmental mismatch), suggesting that maternal investment in last-laid eggs might indeed counterbalance the initial handicap of last-hatched chicks. Our experimental study thus highlights that potential interplays between pre- and post-natal environments are likely to shape fitness prospects in the wild.

Climate change and perishable food hoards of an avian predator: Is the freezer still working?

Changing climate can modify predator-prey interactions and induce declines or local extinctions of species due to reductions in food availability. Species hoarding perishable food for overwinter survival, like predators, are predicted to be particularly susceptible to increasing temperatures. We analysed the influence of autumn and winter weather, and abundance of main prey (voles), on the food-hoarding behaviour of a generalist predator, the Eurasian pygmy owl (Glaucidium passerinum), across 16 years in Finland. Fewer freeze-thaw events in early autumn delayed the initiation of food hoarding. Pygmy owls consumed more hoarded food with more frequent freeze-thaw events and deeper snow cover in autumn and in winter, and lower precipitation in winter. In autumn, the rotting of food hoards increased with precipitation. Hoards already present in early autumn were much more likely to rot than the ones initiated in late autumn. Rotten food hoards were used more in years of low food abundance than in years of high food abundance. Having rotten food hoards in autumn resulted in a lower future recapture probability of female owls. These results indicate that pygmy owls might be partly able to adapt to climate change by delaying food hoarding, but changes in the snow cover, precipitation and frequency of freeze-thaw events might impair their foraging and ultimately decrease local overwinter survival. Long-term trends and future predictions, therefore, suggest that impacts of climate change on wintering food-hoarding species could be substantial, because their 'freezers' may no longer work properly. Altered usability and poorer quality of hoarded food may further modify the foraging needs of food-hoarding predators and thus their overall predation pressure on prey species. This raises concerns about the impacts of climate change on boreal food webs, in which ecological interactions have evolved under cold winter conditions.

Age and sex differences in numerical responses, dietary shifts, and total responses of a generalist predator to population dynamics of main prey.

Fluctuations in the abundance of main prey species might shape animal communities, by inducing numerical responses and dietary shifts in predators. Whether numerical responses and dietary shifts differ among individuals of different age and sex has so far gained little attention. These differences could affect how much predators consume main and alternative prey, thus causing variation in predation pressure on main and alternative prey species. We studied the effect of fluctuating main prey abundance (voles) in autumn on the age and sex composition of a food-hoarding population of Eurasian pygmy owls Glaucidium passerinum (327 individuals), and on the species composition of their food stores in western Finland during 2003-2017 (629 food stores). Numbers of yearlings (< 1-year old) of both sexes and adult (+ 1-year old) females increased with increasing vole abundance. During low vole abundance, adult owls stored more small birds and less small mammals than yearlings. Females stored more small mammals than males and showed a tendency to store less birds. The amount of consumed birds (the most important alternative prey), and in particular of crested, willow, great, and blue tits, increased with low vole densities. Our results show that numerical, functional, and total responses of pygmy owls, and probably also other vertebrate predators, to the availability of the main prey in winter are shaped by the age and sex composition of the predator population, which both show large spatio-temporal variation in boreal forests.

Impact of continuous predator threat on telomere dynamics in parent and nestling pied flycatchers.

In addition to direct mortality, predators can have indirect effects on prey populations by affecting prey behaviour or physiology. For example, predator presence can increase stress hormone levels, which can have physiological costs. Stress exposure accelerates the shortening of telomeres (i.e. the protective caps of chromosomes) and shorter telomeres have been linked to increased mortality risk. However, the effect of perceived predation risk on telomeres is not known. We investigated the effects of continuous predator threat (nesting Eurasian pygmy owl Glaucidium passerinum) on telomere dynamics of both adult and partially cross-fostered nestling pied flycatchers (Ficedula hypoleuca) in the wild. Females nesting at owl-inhabited sites showed impaired telomere maintenance between incubation and chick rearing compared to controls, and both males and females ended up with shorter telomeres at owl-inhabited sites in the end of chick rearing. On the contrary, both original and cross-fostered chicks reared in owl sites had consistently longer telomeres during growth than chicks reared at control sites. Thus, predation risk may cause a long-term cost in terms of telomeres for parents but not for their offspring. Predators may therefore affect telomere dynamics of their preys, which could have implications for their ageing rate and consequently for population dynamics.