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We present the main demographic parameters of the Western Siberian pied flycatcher population, nesting in natural habitats, at different stages of population dynamics. The average values, range, boundaries, and direction of changes in the survival of sexually mature individuals, fecundity, and the encounter probability at different ages were analyzed. Using matrix models of population structure and population dynamics, an estimate of the true survival rate of fledglings is given. The range of values of demographic parameters that characterize viable groups of the studied bird species is indicated. [ABSTRACT FROM AUTHOR]

The study of host–parasite co‐evolution is a central topic in evolutionary ecology. However, research is still fragmented and the extent to which parasites influence host life history is debated. One reason for this incomplete picture is the frequent omission of environmental conditions in studies analyzing host–parasite dynamics, which may influence the exposure to or effects of parasitism. To contribute to elucidating the largely unresolved question of how environmental conditions are related to the prevalence and intensity of infestation and their impact on hosts, we took advantage of 25 years of monitoring of a breeding population of pied flycatchers, Ficedula hypoleuca, in a Mediterranean area of central Spain. We investigated the influence of temperature and precipitation during the nestling stage at a local scale on the intensity of blowfly (Protocalliphora azurea) parasitism during the nestling stage. In addition, we explored the mediating effect of extrinsic and intrinsic factors and blowfly parasitism on breeding success (production of fledglings) and offspring quality (nestling mass on day 13). The prevalence and intensity of blowfly parasitism were associated with different intrinsic (host breeding date, brood size) and extrinsic (breeding habitat, mean temperature) factors. Specifically, higher average temperatures during the nestling phase were associated with lower intensities of parasitism, which may be explained by changes in blowflies’ activity or larval developmental success. In contrast, no relationship was found between the prevalence of parasitism and any of the environmental variables evaluated. Hosts that experienced high parasitism intensities in their broods produced more fledglings as temperature increased, suggesting that physiological responses to severe parasitism during nestling development might be enhanced in warmer conditions. The weight of fledglings was, however, unrelated to the interactive effect of parasitism intensity and environmental conditions. Overall, our results highlight the temperature dependence of parasite–host interactions and the importance of considering multiple fitness indicators and climate‐mediated effects to understand their complex implications for avian fitness and population dynamics. [ABSTRACT FROM AUTHOR]

Individual resoluteness during the resumption of feeding of nestlings was examined in tests involving the presentation of an unfamiliar object near the nests of the pied flycatcher (Ficedula hypoleuca) in a natural environment. Over the course of five breeding seasons, 229 individuals were tested, 41 of which were tested 2 to 4 times. Repeat testing of the same birds was conducted at different nests with intervals ranging from 3 to 1103 days (Med = 367). We took into account the number and age of nestlings, nesting periods, the sex and age of the individual and its partner, the presence of molting, the intensity of nestling feeding, the behavior of the partner, as well as the presence of simultaneous bigamy in the male. We revealed a significant repeatability of test results over substantial time intervals (R = 0.21), indicating the individual's stability over time in response (boldness) to changing conditions in the natural environment. [ABSTRACT FROM AUTHOR]

An assessment of the primary sex ratio in the local pied flycatcher population in South Karelia (Russia) is given, and its variations in broods depending on breeding characteristics and characteristics of the parents are analyzed. The proportion of males in 30 clutches examined in 2012 did not differ from the proportion of females and amounted to 50.8% of the total number of eggs laid (n = 199), although it varied from 14.3% to 85.7% in individual nests. A weak positive correlation between the proportion of males in the brood and the timing of breeding and the ornamentation of the feathers that form the white spot on the male's wing (the first tertiary and the 5–6th upper greater coverts of the secondary feathers), and a negative correlation with the length of the wing and the third primary feather of the female were shown. Analysis using generalized linear models showed a significant relationship of the primary sex ratio in the brood only with the length of the female's third primary feather. Possible mechanisms of changes in the sex ratio in broods are discussed. [ABSTRACT FROM AUTHOR]

Preen oil, the secretion of the uropygial gland, may be an important source of body odour in birds. By characterizing the chemical composition of preen oil, we can describe the olfactory phenotypes of birds and investigate whether odours could have a function in sexual signalling or other chemical communication. Here we analysed the preen oil of a wild passerine, the European Pied Flycatcher Ficedula hypoleuca, to find out whether it holds socially relevant information. We sampled both the female and male of breeding pairs during nestling rearing to test for sex differences and within‐pair similarity. We additionally sampled the females during incubation to test for changes across breeding stages and for individual repeatability of chemical profiles. Pair mates had similar chemical profiles in comparison with other breeding adults. Furthermore, we found evidence for sex differences and for changes across breeding stages. Notably, the preen oil of females was more diverse and more volatile than that of males, and the preen oil secreted by females during incubation was more volatile than that secreted during nestling rearing. However, we found no evidence for individual repeatability of chemical profiles across breeding stages in females. Our results point towards a function of preen oil in sexual signalling, although other functions should not be excluded. Our study is a first step towards understanding the role of odours in the social life of an important avian model species used in the study of mate choice and sexual selection. [ABSTRACT FROM AUTHOR]

Anthropogenic materials are often incorporated into bird's nests in marine and terrestrial habitats globally, although we understand very little about the non-random abundance, and variation in the abundance, of anthropogenic materials in the wider environment and in different sections of bird's nests. Here, we address these oversights by examining if the abundance of anthropogenic material in the environment is influenced by local habitat features and if their abundance varies between the outer and inner sections of pied flycatcher (Ficedula hypoleuca) nests in rural woodlands. Locations in the wider environment with farms, houses and barns contained more anthropogenic material than locations without them, whilst the presence of roads, footpaths and hedges was not associated with the abundance of anthropogenic material in the environment. Nestboxes occupied by pied flycatchers had a greater mass of anthropogenic material in the environment surrounding nestboxes than unoccupied nestboxes. Meanwhile, pied flycatchers incorporated more white and to a lesser extent orange anthropogenic material into the inner and outer sections of their nests, whilst not including significant amounts of red, blue and yellow anthropogenic material. Our results suggest that local habitat features influence the abundance of anthropogenic material in the wider environment, which is presumably because such material is more abundant in modified habitats and that anthropogenic materials are non-randomly incorporated into birds’ nests in relation to their colour and also the nest section. Our study therefore increases our understanding of the incorporation of anthropogenic material into birds’ nests.

Жесткокрылые (Coleoptera) — одна из самых разнообразных групп беспозвоночных в материале из гнезд птиц-дуплогнездников. В ходе исследований на юго-востоке Западной Сибири (Томск) было собрано 50 проб энтомологического материала (беспозвоночные) из гнезд трех видов птиц: Ficedula hypoleuca, Parus major и Phoenicurus phoenicurus. В гнездах двух видов птиц отмечено 40 видов жесткокрылых из 18 семейств. В гнездах Parus major жуки не обнаружены. Среди жуков большая часть таксонов (97,2%) в гнездах представлена пищевыми остатками. В гнездах (и питании) Ficedula hypoleuca в условиях Томска преобладают жужелицы (Carabidae), их доля составила 43,8%. Из нидиколов было обнаружено два вида — Gnathoncus buyssoni (Histeridae) и Dermestes undulatus (Dermestidae). Интересны находки редких видов Otho sphondyloides (Eucnemidae), Ischnodes sibiricus (Elateridae). А вид Aplocnemus nigricornis (Melyridae) впервые указывается для азиатской части Палеарктики, Сибири и Томской области.

Using light-level geolocators, eight European Pied Flycatchers (Ficedula hypoleuca) from two breeding sites in Czechia were tracked. We also gathered all available geolocator tracks on 76 individuals from four European populations and compared the timing of annual cycle events and the African non-breeding sites among all populations. Individuals from both Czech breeding sites had overlapping migration events and non-breeding locations. Four individuals resided in the southwestern edge of Mali, two in Burkina Faso, one in Guinea, and the easternmost one in the Ivory Coast. On average, the birds left the Czech breeding grounds on 8 August and took between one to three stopovers during autumn migration. Birds crossed the Sahara on its western edge on average on 13 September. The mean arrival to the African non-breeding grounds was 47.5 days after departure on 2 October (range 10 September to 10 October). One bird showed intra-tropical movement within West Africa when after a 60-day residency it moved approximately 3° westwards. Estimated locations at the African non-breeding grounds overlapped among tracked birds from five European breeding sites. However, statistically, we could detect longitudinal segregation in two clusters. Birds from the British and Finnish breeding populations shared non-breeding grounds and were located in Africa west of the second cluster of the birds from the Czech and Dutch breeding populations. We show considerable population-specific differences in the timing of annual cycle events. Birds from Dutch breeding sites were the first in all three phases—departure from breeding sites, Sahara crossing and arrival to African non-breeding grounds, followed by the British, Czech, and Finnish birds, respectively. All tracked flycatchers so far fill only the western part of the African non-breeding range. For a complete understanding of the migration pattern in the species, we highlight the need for tracking studies from the eastern part of the range. [ABSTRACT FROM AUTHOR]

The nests of secondary cavity-nesters located in tree cavities may form specific microhabitats of conservation importance due to their limited accessibility and availability. Species-specific nesting materials in nests of different secondary cavity-nesters may furthermore provide very different microhabitats for arthropods. The potential differences in arthropod communities inhabiting nests of different bird species in excavated cavities or nest boxes have, however, rarely been studied despite their relevance for conservation. Here we investigated the diversity and composition of arthropod communities in these different cavity types and bird species' nests in managed boreal forests. We identified morphologically and by DNA-metabarcoding arthropods in nest materials that were collected in and compared between (i) woodpecker-size cavities from seven different combinations of cavity type (nest box or excavated cavity), tree species (aspen or pine) and accumulation history of nest materials (single-season cleaned or uncleaned nest boxes that accumulated nests of passerines or an owl species); and (ii) nests of two different passerine species in small nest boxes. We identified 64 arthropod taxa in ten orders, from which Diptera, Coleoptera, Siphonaptera, and Lepidoptera were the most abundant. Shannon diversity index was similar among the cavity-nest-type combinations, but taxa richness was the highest in the owl nests. The arthropod communities (especially Histeridae beetles) deviated most from the other types of nests in owl and aspen cavity nests with more advanced decomposition of nest material (soil or wet environment related taxa). The differences in arthropod communities between the different nest types point out the importance of the ecological chain "tree cavities—bird nests—arthropod communities". [ABSTRACT FROM AUTHOR]

Parent–offspring conflict over food allocation can be modeled using two theoretical frameworks: passive (scramble competition) and active choice (signaling) resolution models. However, differentiating between these models empirically can be challenging. One possibility involves investigating details of decision-making by feeding parents. Different nestling traits, related to competitive prowess or signaling cryptic condition, may interact additively or non-additively as predictors of parental feeding responses. To explore this, we experimentally created even-sized, small broods of pied flycatchers and manipulated nestling cryptic quality, independently of size, by vitamin E supplementation. We explored how interactions between nestling cryptic condition, size, signals, and spatial location predicted food allocation and prey-testing by parents. Parents created the potential for spatial scramble competition between nestlings by feeding from and to a narrow range of nest locations. Heavier supplemented nestlings grew faster and were more likely to access profitable nest locations. However, the most profitable locations were not more contested, and nestling turnover did not vary in relation to spatial predictability or food supply. Postural begging was only predicted by nestling hunger and body mass, but parents did not favor heavier nestlings. This suggests that size-mediated and spatial competition in experimental broods was mild. Pied flycatcher fathers allocated food in response to nestling position and begging order, while mothers seemingly followed an active choice mechanism involving assessment of more complex traits, including postural intensity interacting with order, position, and treatment, and perhaps other stimuli when performing prey-testings. Differences in time constraints may underlie sex differences in food allocation rules. [ABSTRACT FROM AUTHOR]

The authors have obtained original material on the fauna and population structure of oribatid mites inhabiting nests of the European Pied Flycatcher (Ficedula hypoleuca, Passeriformes, hollow-nesting bird) on the territory of the taiga zone of the European North-East of Russia. Long-term research and the collection of nests were carried out in the green zone of Syktyvkar in 2017–2022. Observations were made for artificial nests (hollows) of a box type with a bottom area of 100 cm2. The material of the tray was collected completely. In 135 studied nests of Pied Flycatchers, 1762 specimens were found and identified for 22 species of oribatid mites from 19 genera and 16 families. In the nests of the Pied Flycatcher, a complex of species was found that is known as an arboricolous species for this region; these are Oribatula (Zygoribatula) propinqua, Oribatula (Z.) exilis, Trichoribates (T.) berlesei, and Ameronothrus oblongus. We suggested that arboricolous species, as well as eurytopic species, can actively inhabit bird nests. Highly numerous in our collections were representatives of the Oribatulidae and Scheloribatidae families; they are Oribatula (Z.) propinqua, Oribatula (Z.) exilis, Oribatula (O.) tibialis, and Scheloribates laevigatus. Epigeic species are dominated by the species number. The fauna of oribatid mites mainly included widespread Holarctic species (54.54%). [ABSTRACT FROM AUTHOR]

Polymelia is a congenital defect characterized by an excessive number of limbs. Leg malformations are unusual and have rarely been reported in birds. The ultimate mechanism that regulates this type of abnormal development is not yet well understood. Here we report polymelia in a nestling of the European Pied Flycatcher Ficedula hypoleuca. We have ringed more than 12,100 Pied Flycatcher nestlings since 1991 and have never observed this malformation before. To our knowledge, there is no study indicating polymelia in any bird of the order Passeriformes. We observed and studied a Pied Flycatcher nestling that had two extra limbs fused at the pelvis. We used X-rays, Computerized Tomography (CT) scan and Magnetic Resonance Imaging (MRI). We describe the observed malformations and discuss potential causes. The malformation could be due to an embryonic cause, such as a parasitic twin (pygopagus parasitic) or a genetic mutation. [ABSTRACT FROM AUTHOR]

In most animal species, dispersing individuals possess phenotypic attributes that mitigate the costs of colonization and/or increase settlement success in new areas (dispersal syndromes). This phenotypic integration likely affects population dynamics and the direction of selection, but data are lacking for natural populations. Using an approach that combines population dynamics, quantitative genetics, and phenotypic selection analyses, we reveal the existence of dispersal syndromes in a pied flycatcher (Ficedula hypoleuca) population in the Netherlands: immigrants were larger, tended to have darker plumage, bred earlier, and produced larger clutches than local recruits, and some of these traits were genetically correlated. Over time, the phenotypic profile of the population gradually changed: each generation advanced arrival and breeding and exhibited longer wings as a result of direct and indirect selection on these correlated traits. Although phenotypic attributes of immigrants were favored by selection during the early phase of colonization, observed phenotypic changes were similar for immigrants and local recruits. We propose that immigrants facilitated initial population establishment but that temporal changes likely resulted from climate change–induced large-scale selection. This study highlights that newly established populations are of nonrandom composition and that phenotypic architecture affects evolutionary population trajectories. [ABSTRACT FROM AUTHOR]

The authors have obtained original material on the fauna and population structure of oribatid mites inhabiting nests of the European Pied Flycatcher (Ficedula hypoleuca, Passeriformes, hollow-nesting bird) on the territory of the taiga zone of the European North-East of Russia. Long-term research and the collection of nests were carried out in the green zone of Syktyvkar in 2017–2022. Observations were made for artificial nests (hollows) of a box type with a bottom area of 100 cm2. The material of the tray was collected completely. In 135 studied nests of Pied Flycatchers, 1762 specimens were found and identified for 22 species of oribatid mites from 19 genera and 16 families. In the nests of the Pied Flycatcher, a complex of species was found that is known as an arboricolous species for this region; these are Oribatula (Zygoribatula) propinqua, Oribatula (Z.) exilis, Trichoribates (T.) berlesei, and Ameronothrus oblongus. We suggested that arboricolous species, as well as eurytopic species, can actively inhabit bird nests. Highly numerous in our collections were representatives of the Oribatulidae and Scheloribatidae families; they are Oribatula (Z.) propinqua, Oribatula (Z.) exilis, Oribatula (O.) tibialis, and Scheloribates laevigatus. Epigeic species are dominated by the species number. The fauna of oribatid mites mainly included widespread Holarctic species (54.54%).

Simultaneous nesting of six species of secondary hollow-nesting passerine birds in abandoned European Bee-eater Merops apiaster nest-holes has been detected and described. The holes were occupied by Great Tit Parus major, Spotted Flycatcher Muscicapa striata, European Pied Flycatcher Ficedula hypoleuca, Black Redstart Phoenicurus ochruros, Eurasian Tree Sparrow Passer montanus and White Wagtail Motacilla alba, which formed a multi-species settlement in a European Bee-eater colony. [ABSTRACT FROM AUTHOR]

Ecological research is often hampered by the inability to quantify animal diets. Diet composition can be tracked through DNA metabarcoding of fecal samples, but whether (complex) diets can be quantitatively determined with metabarcoding is still debated and needs validation using free‐living animals. This study validates that DNA metabarcoding of feces can retrieve actual ingested taxa, and most importantly, that read numbers retrieved from sequencing can also be used to quantify the relative biomass of dietary taxa. Validation was done with the hole‐nesting insectivorous Pied Flycatcher whose diet was quantified using camera footage. Size‐adjusted counts of food items delivered to nestlings were used as a proxy for provided biomass of prey orders and families, and subsequently, nestling feces were assessed through DNA metabarcoding. To explore potential effects of digestion, gizzard and lower intestine samples of freshly collected birds were subjected to DNA metabarcoding. For metabarcoding with Cytochrome Oxidase subunit I (COI), we modified published invertebrate COI primers LCO1490 and HCO1777, which reduced host reads to 0.03%, and amplified Arachnida DNA without significant changing the recovery of other arthropod taxa. DNA metabarcoding retrieved all commonly camera‐recorded taxa. Overall, and in each replicate year (N = 3), the relative scaled biomass of prey taxa and COI read numbers correlated at R =.85 (95CI:0.68–0.94) at order level and at R =.75 (CI:0.67–0.82) at family level. Similarity in arthropod community composition between gizzard and intestines suggested limited digestive bias. This DNA metabarcoding validation demonstrates that quantitative analyses of arthropod diet is possible. We discuss the ecological applications for insectivorous birds. [ABSTRACT FROM AUTHOR]

Most studies of phenotypic selection in the wild have focussed on morphological and life‐history traits and looked at abiotic (climatic) variation as the main driver of selection. Consequently, our knowledge of the effects of biotic environmental variation on phenotypic selection on sexual traits is scarce. Population density can be considered a proxy for the intensity of intrasexual and intersexual competition and could therefore be a key factor influencing the covariation between individual fitness and the expression of sexual traits. Here, we used an individual‐based data set from a population of pied flycatchers (Ficedula hypoleuca) monitored over 24 years to analyze the effect of breeding density on phenotypic selection on dorsal plumage colouration, a heritable and sexually selected ornament in males of this species. Using the number of recruits as a fitness proxy, our results showed overall stabilizing selection on male dorsal colouration, with intermediate phenotypes being favoured over extremely dark and dull individuals. However, our results did not support the hypothesis that breeding density mediates phenotypic selection on this sexual trait. We discuss the possible role of other biotic factors influencing selection on ornamental plumage. [ABSTRACT FROM AUTHOR]

Extra-pair copulation (EPC) occurred in most socially monogamous bird species. The mechanisms leading to the frequent occurrence of extra-pair offspring (EPO, EPY) in socially monogamous couples, as well as the 'function' of EPC, are the subjects of strong debates and raise many unanswered questions. We studied the relationship between extra-pair paternity (EPP) and the different characteristics of males and females in the European pied flycatcher in Western Siberia (Russia). The analysis was based on the genotyping of 232 males, 250 females, 1485 nestlings (250 nests). The European pied flycatchers were predominantly socially and genetically monogamous, but about 20% of birds could be involved in EPP. Loss of paternity tended to be more frequent in one-year-old males. EPCs could be multiple: one individual may have up to three extra-pair partners. The EPP rate was independent of the breeding time. The extra-pair mates of an individual were mainly its near neighbours. The EPC status of an individual was unrelated to most of its morpho-physiological traits. The occurrence of EPP was almost twice as high in females nesting in good quality territories. The fitness of within-pair offspring, EPO, paternal half-sibs of EPO and maternal half-sibs of EPO did not differ statistically significantly. Assuming very low heritability of extra-pair mating, we argued that EPCs could be incidental side effects (by-product) of selection. We believe that the evolution and maintenance of extra-pair mating are the episelective processes in the case of the European pied flycatcher. [ABSTRACT FROM AUTHOR]

Songbird populations are in decline all over the world, and our understanding of the causal mechanisms remains surprisingly limited. It is important to identify the extent of individual variations in migratory behaviour to better understand species' ability to respond to environmental change. We describe the annual migratory behaviour of British breeding European Pied Flycatchers Ficedula hypoleuca by using light‐recording geolocators. During both autumn and spring migrations, individuals used previously unknown staging areas on the Iberian Peninsula and northern West Africa. Furthermore, partial sex‐specific segregation in the location of non‐breeding areas was observed within West Africa, with females located west of males. We also found sex‐based phenological differences, with females staying longer in non‐breeding areas and undertaking the spring Sahara Desert crossing later than males. Irrespective of sex, multiple use of the two predominant staging regions was identified during both migrations, with 63% of individuals stopping more than once in these regions. We also identified instances of migratory behaviours rarely documented in individually tracked songbirds. These include making daytime landfall during the Sahara crossing, and a case of a temporary retreat migration, with an individual aborting a spring Sahara crossing before making a second successful crossing 14 days later. Together, our results show variability in migratory behaviour both between sexes and between individuals. For Pied Flycatchers, such flexible migratory behaviour may increase their resilience to environmental change. [ABSTRACT FROM AUTHOR]

Abstract Understanding the origin and persistence of phenotypic variation within and among populations is a major goal in evolutionary biology. However, the eagerness to find unadulterated explanatory models in combination with difficulties in publishing replicated studies may lead to severe underestimations of the complexity of selection patterns acting in nature. One striking example is variation in plumage coloration in birds, where the default adaptive explanation often is that brightly colored individuals signal superior quality across environmental conditions and therefore always should be favored by directional mate choice. Here, we review studies on the proximate determination and adaptive function of coloration traits in male pied flycatchers (Ficedula hypoleuca). From numerous studies, we can conclude that the dark male color phenotype is adapted to a typical northern climate and functions as a dominance signal in male–male competition over nesting sites, and that the browner phenotypes are favored by relaxed intraspecific competition with more dominant male collared flycatchers (Ficedula albicollis) in areas where the two species co‐occur. However, the role of avoidance of hybridization in driving character displacement in plumage between these two species may not be as important as initially thought. The direction of female choice on male coloration in pied flycatchers is not simply as opposite in direction in sympatry and allopatry as traditionally expected, but varies also in relation to additional contexts such as climate variation. While some of the heterogeneity in the observed relationships between coloration and fitness probably indicate type 1 errors, we strongly argue that environmental heterogeneity and context‐dependent selection play important roles in explaining plumage color variation in this species, which probably also is the case in many other species studied in less detail.

Abstract Ecological research is often hampered by the inability to quantify animal diets. Diet composition can be tracked through DNA metabarcoding of fecal samples, but whether (complex) diets can be quantitatively determined with metabarcoding is still debated and needs validation using free‐living animals. This study validates that DNA metabarcoding of feces can retrieve actual ingested taxa, and most importantly, that read numbers retrieved from sequencing can also be used to quantify the relative biomass of dietary taxa. Validation was done with the hole‐nesting insectivorous Pied Flycatcher whose diet was quantified using camera footage. Size‐adjusted counts of food items delivered to nestlings were used as a proxy for provided biomass of prey orders and families, and subsequently, nestling feces were assessed through DNA metabarcoding. To explore potential effects of digestion, gizzard and lower intestine samples of freshly collected birds were subjected to DNA metabarcoding. For metabarcoding with Cytochrome Oxidase subunit I (COI), we modified published invertebrate COI primers LCO1490 and HCO1777, which reduced host reads to 0.03%, and amplified Arachnida DNA without significant changing the recovery of other arthropod taxa. DNA metabarcoding retrieved all commonly camera‐recorded taxa. Overall, and in each replicate year (N = 3), the relative scaled biomass of prey taxa and COI read numbers correlated at R = .85 (95CI:0.68–0.94) at order level and at R = .75 (CI:0.67–0.82) at family level. Similarity in arthropod community composition between gizzard and intestines suggested limited digestive bias. This DNA metabarcoding validation demonstrates that quantitative analyses of arthropod diet is possible. We discuss the ecological applications for insectivorous birds.

Long-term and short-term changes in ambient temperature can cause stress in birds, leading to changes in the level of hematological parameters. The H:L ratio (heterophil-to-lymphocyte ratio) is a hematological index that allows for the assessment of the stress induced by environmental changes, including weather conditions. In this paper, we examined the influence of temperatures and the sum of precipitation on the health of nestling pied flycatchers (Ficedula hypoleuca) by using the H:L ratio reflecting the body's response to stress. All examined temperature indicators influenced the H:L ratio, yet the average value of daily minimum temperature during the first 12 days of nestling life had the strongest influence, maximum temperature had the weakest effect, while precipitation had no significant influence. Our research indicates that even a small increase in temperature caused a stress reaction in nestling pied flycatchers, which was reflected by an increase in the H:L ratio. The increase in the stress index (H:L ratio) was probably a result of poor weather conditions (precipitation, low temperature), which prevented the adult birds from actively foraging and properly feeding the nestlings. [ABSTRACT FROM AUTHOR]

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Conserving species relies upon acquiring an understanding of their use of habitat, yet our understanding of the use of habitat by co-existing species of different guilds at microgeographic scales remains poor. In particular, the use of habitat by woodland species is of conservation concern because of widespread declines in woodland biodiversity. Woodland bird declines have been ascribed, in part, to high deer densities because their browsing reduces the availability of nesting sites and food. We quantify the microgeographic use of habitat by Roe Deer (Capreolus capreolus) and of Great Tits (Parus major), Blue Tits (Cyanistes caeruleus) and Pied Flycatchers (Ficedula hypoleuca) in a heterogenous woodland landscape. We examined the use of habitat at microgeographic scales by the deer and the three bird species in relation to whether the local habitat was flat or wet or had a path, fence or wall within a 25-m radius of 206 randomly selected locations. We first examined if the occupancy rates of nestboxes in those locations were correlated with the number of Roe Deer lays and second, examined if the use of habitat by the Roe Deer and the bird species were associated with each of the habitat features that we quantified. We begin by showing that the use of habitat by Roe Deer is incongruent with the use of habitat by Great Tits, Blue Tits and Pied Flycatchers during the breeding season. Also, whilst all three bird species showed no, or weak, habitat preferences, the Roe Deer preferred daytime lay sites that were in flat areas of wet woodland close to paths, whilst there were no significant effects of the presence of fences and walls. These findings show that the Roe Deer and the three bird species differ in the use of habitat within a heterogenous woodland landscape, meaning that their use of habitat did not overlap at microgeographic scales. Meanwhile, the deer showed preferences for flat areas of wet woodland, whilst none of the bird species exhibited such preferences, and we discuss the implications of our findings for the management of woodlands.

The origin of natural selection is linked to environmental heterogeneity, which influences variation in relative fitness among phenotypes. However, individuals in wild populations are exposed to a plethora of biotic and abiotic environmental factors. Surprisingly, the relative influence of multiple environmental conditions on the relative fitness of phenotypes has rarely been tested in wild populations. Identifying the main selection agent(s) is crucial when the target phenotype is tightly linked to reproduction and when temporal variation in selection is expected to affect evolutionary responses. By using individual‐based data from a 29‐year study of a short‐lived migratory songbird, the pied flycatcher (Ficedula hypoleuca), we studied the relative influence of 28 temperature‐ and precipitation‐based factors at local and global scales on selection on breeding time (egg laying) at the phenotypic level. Selection, estimated using the number of recruits as a proxy for fitness, penalized late breeders. Minimum temperatures in April and May were the environmental drivers that best explained selection on laying date. In particular, there was negative directional selection on laying date mediated by minimum temperature in April, being strongest in cold years. In addition, nonlinear selection on laying date was influenced by minimum temperatures in May, with selection on laying date changing from null to negative as the breeding season progressed. The intensity of selection on late breeders increased when minimum temperatures in May were highest. Our results illustrate the complex influence of environmental factors on selection on laying date in wild bird populations. Despite minimum temperature in April being the only variable that changed over time, its increase did not induce a shift in laying date in the population. In this songbird population, stabilizing selection has led to a three‐decade stasis in breeding time. We suggest that variation in the effects of multiple climatic variables on selection may constrain phenotypic change. [ABSTRACT FROM AUTHOR]

The age-related variability of the frequency and structural–temporal parameters of the advertising song of Ficedula hypoleuca males in natural habitats and the urban environment were analysed. In the natural habitats, the range of male song frequencies, the maximal frequency, the number of figures, and the length of the song decreased from the third year of life. Similar age-related song changes in urban birds are observed earlier, from the second year of life. Urban birds have shown that the higher minimum song frequency increases with age, adapting the song to noise pollution. [ABSTRACT FROM AUTHOR]

Age-related variation in reproductive performance is central for the understanding of population dynamics and evolutionary processes. Our understanding of age trajectories in vital rates has long been limited by the lack of distinction between patterns occurring within- and among-individuals, and by the lack of comparative studies of age trajectories among traits. Thus, it is poorly understood how sets of demographic traits change within individuals according to their age. Based on 40 years of monitoring, we investigated age-related variation in five reproductive traits in female pied flycatchers (Ficedula hypoleuca) including laying date, clutch size, brood size, nest success (probability that a nest produces at least one chick) and egg success of successful nests (proportion of eggs resulting in a chick). We disentangled within- from among-individual processes and assessed the relative contribution of within-individual age-specific changes and selective appearance and disappearance. Finally, we compared the aging pattern among these five reproductive traits. We found strong evidence for age-specific performance including both early-life improvement and late-life decline in all reproductive traits but the egg success. Furthermore, the aging patterns varied substantially among reproductive traits both for the age of peak performance and for the rates of early-life improvement and late-life decline. The results show that age trajectories observed at the population level (cross-sectional analysis) may substantially differ from those occurring at the individual level and illustrate the complexity of variation in aging patterns across traits. [ABSTRACT FROM AUTHOR]

The dynamics of the species richness, breeding density, and structure of the community of birds occupying nest-boxes upon a 50-fold reduction of atmospheric emissions from the Middle Ural Copper Smelter was analyzed based on long-term continuous observations (1989–2020). The total density in the deciduous forest was lower in the heavily polluted zone (impact zone) than in the background zone, while there were no differences in the coniferous forest between the zones. The total breeding density of the community of hole-nesters increased in the impact zone over the 30 years (especially, in the deciduous forest in 2010–2020) after the significant reduction of industrial emissions. The breeding density of the two most abundant species varied in different directions: it increased for pied flycatcher and decreased for common redstart. This led to a change of dominants in the impact zone: redstart dominating until 2012 was replaced by pied flycatcher. The differences in the dynamics of species density are probably determined by the beginning of vegetation recovery near the smelter, which is favorable for pied flycatcher but not for redstart. [ABSTRACT FROM AUTHOR]

Abstract Background Nest construction is a key element of avian reproductive behaviour and the result is often a complex structure that is used for incubation of eggs, which represents an extended phenotype. It is known that nest construction is a plastic behaviour but the extent to which plasticity is observed in a single species with a wide geographical distribution is largely unknown. This study sought to better understand variation in nest size and composition across a very wide geographical area. The hypothesis suggested that location would affect size but not composition of nests of the European Pied Flycatcher (Ficedula hypoleuca). Methods Nests and reproductive data were collected from seventeen study sites, spread over 6° of latitude and 3.3° of longitude on the island of Great Britain. Dimensions of nests were measured before they were deconstructed to determine the masses and types materials used in the outer nest and the cup lining. Results Geographical variation was observed in base thickness of nests but not many other dimensions. Nests varied in composition but were mainly made of leaf, moss, bark, grass, root and fern. Moss was used more to the north and east of the study area compared with more leaf mass towards the south and west. The species of leaf and bark used in the nests varied between geographical locations. Additionally, the use of leaves or bark from a particular tree species did not reflect the incidence of the tree species in the immediate territory. Conclusions This study showed that nest composition was affected by geographical location over a wide area. Variation between nests at each location was high and so it was concluded that differences in nest composition reflect individual selection of materials but evidence is such that it remains unclear whether this is deliberate to fulfil a specific role in the nest, or simply opportunistic with birds simply picking up materials with the appropriate characteristics as they find them outside their nestbox.

In migratory species, the timing of arrival at the breeding grounds is a life-history trait with major fitness consequences. The optimal arrival date varies from year-to-year, and animals use cues to adjust their arrival dates to match this annual variation. However, which cues they use to time their arrival and whether these cues actually predict the annual optimal arrival date is largely unknown. Here, we integrate causal and evolutionary analysis by identifying the environmental variables used by a migratory songbird to time its arrival dates and testing whether these environmental variables also predicted the optimal time to arrive. We used 11 years of male arrival data of a pied flycatcher population. Specifically, we tested whether temperature and normalized difference vegetation index (NDVI) values from their breeding grounds in the Netherlands and from their wintering grounds in Ivory Coast explained the variation in arrival date, and whether these variables correlated with the position of the annual fitness peak at the breeding grounds. We found that temperature and NDVI, both from the wintering and the breeding grounds, explained the annual variation in arrival date, but did not correlate with the optimal arrival date. We explore three alternative explanations for this lack of correlation. Firstly, the date of the fitness peak may have been incorrectly estimated because a potentially important component of fitness (i.e., migration date dependent mortality en route or directly upon arrival) could not be measured. Secondly, we focused on male timing but the fitness landscape is also likely to be shaped by female timing. Finally, the correlation has recently disappeared because climate change disrupted the predictive value of the cues that the birds use to time their migration. In the latter case, birds may adapt by altering their sensitivity to temperature and NDVI.

Extra-pair copulation (EPC) occurred in most socially monogamous bird species. The mechanisms leading to the frequent occurrence of extra-pair offspring (EPO, EPY) in socially monogamous couples, as well as the ‘function’ of EPC, are the subjects of strong debates and raise many unanswered questions. We studied the relationship between extra-pair paternity (EPP) and the different characteristics of males and females in the European pied flycatcher in Western Siberia (Russia). The analysis was based on the genotyping of 232 males, 250 females, 1485 nestlings (250 nests). The European pied flycatchers were predominantly socially and genetically monogamous, but about 20% of birds could be involved in EPP. Loss of paternity tended to be more frequent in one-year-old males. EPCs could be multiple: one individual may have up to three extra-pair partners. The EPP rate was independent of the breeding time. The extra-pair mates of an individual were mainly its near neighbours. The EPC status of an individual was unrelated to most of its morpho-physiological traits. The occurrence of EPP was almost twice as high in females nesting in good quality territories. The fitness of within-pair offspring, EPO, paternal half-sibs of EPO and maternal half-sibs of EPO did not differ statistically significantly. Assuming very low heritability of extra-pair mating, we argued that EPCs could be incidental side effects (by-product) of selection. We believe that the evolution and maintenance of extra-pair mating are the episelective processes in the case of the European pied flycatcher.

Understanding the origin and persistence of phenotypic variation within and among populations is a major goal in evolutionary biology. However, the eagerness to find unadulterated explanatory models in combination with difficulties in publishing replicated studies may lead to severe underestimations of the complexity of selection patterns acting in nature. One striking example is variation in plumage coloration in birds, where the default adaptive explanation often is that brightly colored individuals signal superior quality across environmental conditions and therefore always should be favored by directional mate choice. Here, we review studies on the proximate determination and adaptive function of coloration traits in male pied flycatchers (Ficedula hypoleuca). From numerous studies, we can conclude that the dark male color phenotype is adapted to a typical northern climate and functions as a dominance signal in male–male competition over nesting sites, and that the browner phenotypes are favored by relaxed intraspecific competition with more dominant male collared flycatchers (Ficedula albicollis) in areas where the two species co‐occur. However, the role of avoidance of hybridization in driving character displacement in plumage between these two species may not be as important as initially thought. The direction of female choice on male coloration in pied flycatchers is not simply as opposite in direction in sympatry and allopatry as traditionally expected, but varies also in relation to additional contexts such as climate variation. While some of the heterogeneity in the observed relationships between coloration and fitness probably indicate type 1 errors, we strongly argue that environmental heterogeneity and context‐dependent selection play important roles in explaining plumage color variation in this species, which probably also is the case in many other species studied in less detail. [ABSTRACT FROM AUTHOR]

Males and females take part in extra-pair copulations in most socially monogamous bird species. The mechanisms leading to the frequent occurrence of extra-pair offspring in socially monogamous couples are strongly debated and unresolved, and they are often difficult to distinguish from one another. Most hypotheses explaining the evolution of extra-pair reproduction suggest selective and adaptive scenarios for their origination and persistence. Is extra-pair paternity a heritable trait? We evaluated the heritability of extra-pair paternity in the pied flycatcher (Ficedula hypoleuca) nesting in Western Siberia. Estimated heritability was low: depending on the model used, the point estimate of the heritability (mode) varied from 0.005 to 0.11, and the bounds of the 95% confidence interval are [0–0.16] in the widest range. Thus, it seems that extra-pair mating behaviour in the pied flycatchers is a plastic phenotypic mating tactic with a small or no genetic component. Our data can help to understand the evolution of extra-pair mating behaviour in socially monogamous species.

The structure of the diet of passerine bird nestlings was analyzed in order to identify possible ways to reduce food competition between these species during the breeding period. We assumed that analysis of the effects of obtaining diets of heterospecific birds by nestlings in nature can help better understand their species-specific adaptations to living conditions, and we considered the case of joint nesting of the coal tit (Periparus ater (L. 1758)) (CT) and pied flycatcher (Ficedula hypoleuca (Pallas 1764)) (PF) in Moscow oblast as a key link. A PF male attracted a female to an artificial nest box, which has been already occupied by a CT pair. The females of both species incubated a mixed clutch sitting side by side. The CT nestlings hatched earlier than the PF nestlings, and the parents of both species began to feed them. The CT nestlings started to die one by one on the 4th day of joint feeding, and the last nestling died on the 11th day. This case encouraged us to compare the diets of nestlings from 22 broods of CTs and PFs in the same region. To compare the ratios of nestlings, we collected the portions of food that were delivered to nestlings by their parents and analyzed video recordings from the nests. The diet of PF nestlings was significantly more diverse than in CT nestlings (Berger–Parker reverse indices (1/d) were 3.01 and 2.11, respectively). Caterpillars and spiders (objects with a high content of carotenoids and taurine important for the development of nestlings) dominated in the CT diet. In PF feed, these objects occupied a much smaller portion. In addition to the above, compared with coal tits, PFs brought the following to nestlings: a significant number of objects with rough, highly chitinized covers (Coleoptera, Homoptera, etc.); objects with a sharp taste, such as true bugs (Heteroptera), ladybirds (Coleoptera, Coccinellidae), and millipedes (Diplopoda, Julidae); insects containing cantharidin toxin (soldier beetles, Coleoptera, Cantharidae). The nestlings developed successfully in PF nests to which parents brought such feed objects. The dimensional ranges of objects in the CT and PF diets completely overlapped. According to our opinion, the rougher, highly chitinized food and some poisonous insects (Cantharidae) in the PF diet make fundamentally important differences in the nutrition of PFs and CTs. Most likely, PFs have resistance to cantharidin and some other toxins that are contained in insects and therefore have a wider trophic niche. [ABSTRACT FROM AUTHOR]

Males and females take part in extra-pair copulations in most socially monogamous bird species. The mechanisms leading to the frequent occurrence of extra-pair offspring in socially monogamous couples are strongly debated and unresolved, and they are often difficult to distinguish from one another. Most hypotheses explaining the evolution of extra-pair reproduction suggest selective and adaptive scenarios for their origination and persistence. Is extra-pair paternity a heritable trait? We evaluated the heritability of extra-pair paternity in the pied flycatcher (Ficedula hypoleuca) nesting in Western Siberia. Estimated heritability was low: depending on the model used, the point estimate of the heritability (mode) varied from 0.005 to 0.11, and the bounds of the 95% confidence interval are [0-0.16] in the widest range. Thus, it seems that extra-pair mating behaviour in the pied flycatchers is a plastic phenotypic mating tactic with a small or no genetic component. Our data can help to understand the evolution of extra-pair mating behaviour in socially monogamous species. [ABSTRACT FROM AUTHOR]

In polygynous species with biparental care, mates are often acquired in succession. Most research has focussed on the cost of polygyny in secondary females, but primary females may also suffer from reduced paternal care. The likelihood of sharing a male may be higher for early laying females, which could counteract the fitness benefits of breeding early. In this study, we use 12 years of data on pied flycatchers Ficedula hypoleuca, to show that the likelihood of becoming a primary female of a polygynous male declines over the season. Moreover, we provide experimental evidence that early breeding elevates polygyny risk, through an experimental manipulation that introduced early breeding females to a population with later breeding phenology. We found that, independently of breeding date, primary females slightly more often experienced complete brood failures than monogamous females, but did not differ in number of fledged offspring among successful broods or number of locally returning recruits. However, apparent survival in subsequent years was substantially lower in primary females, indicating that they may compensate for reduced male care at the expense of future reproduction. Our study reveals that polygyny risk indeed increases with early breeding and entails a local survival cost for primary females. However, this cost is likely largely outweighed by fitness benefits of early breeding in most years. Hence it is unlikely that the increased polygyny risk of early breeding counteracts the fitness benefits, but it may reduce selection for breeding extremely early. [ABSTRACT FROM AUTHOR]

Parasitoid wasps may act as hyperparasites and sometimes regulate the populations of their hosts by a top-down dynamic. Nasonia vitripennis (Walker, 1836) is a generalist gregarious parasitoid that parasitizes several host flies, including the blowfly Protocalliphora Hough, 1899 (Diptera, Calliphoridae), which in turn parasitizes bird nestlings. Nonetheless, the ecological factors underlying N. vitripennis prevalence and parasitoidism intensity on its hosts in natural populations are poorly understood. We have studied the prevalence of N. vitripennis in Protocalliphora azurea (Fallén, 1817) puparia parasitizing wild populations of pied flycatcher (Ficedula hypoleuca) and blue tit (Cyanistes caeruleus) birds in two Mediterranean areas in central and southern Spain. We found some evidence that the prevalence of N. vitripennis was higher in moist habitats in southern Spain. A host-dependent effect was found, since the greater the number of P. azurea puparia, the greater the probability and rate of parasitoidism by the wasp. Our results also suggest that N. vitripennis parasitizes more P. azurea puparia in blue tit nests than in pied flycatcher nests as a consequence of a higher load of these flies in the former. Based on the high prevalence of N. vitripennis in P. azurea puparia in nature, we propose that this wasp may regulate blowfly populations, with possible positive effects on the reproduction of both bird species. [ABSTRACT FROM AUTHOR]

Abstract Background Social learning allows animals to eavesdrop on ecologically relevant knowledge of competitors in their environment. This is especially important when selecting a habitat if individuals have relatively little personal information on habitat quality. It is known that birds can use both conspecific and heterospecific information for social learning, but little is known about the relative importance of each information type. If provided with the choice between them, we expected that animals should copy the behaviour of conspecifics, as these confer the best information for that species. We tested this hypothesis in the field for Pied Flycatchers Ficedula hypoleuca arriving at their breeding grounds to select a nest box for breeding. We assigned arbitrary symbols to nest boxes of breeding pied flycatchers (conspecifics) and blue and great tits, Cyanistes caeruleus and Parus major (heterospecifics), in 2014 and 2016 in two areas with different densities of tits and flycatchers. After ca 50% of flycatchers had returned and a flycatcher symbol was assigned to their nest box, we gave the later arriving flycatchers the choice between empty nest boxes with either a conspecific (flycatcher) or a heterospecific (tit) symbol. Results As expected, Pied Flycatchers copied the perceived nest box choice of conspecifics, but only in areas that were dominated by flycatchers. Against our initial expectation, flycatchers copied the perceived choice of heterospecifics in the area heavily dominated by tits, even though conspecific minority information was present. Conclusions Our results confirm that the relative density of conspecifics and heterospecifics modulates the propensity to copy or reject novel behavioural traits. By contrasting conspecific and heterospecific ecology in the same study design we were able to draw more general conclusions about the role of fluctuating densities on social information use.

The natural quantity of the pied flycatcher in deciduous, mixed, and coniferous forests and reproductive parameters in deciduous and mixed stands of subtaiga forests in the southeast of Western Siberia has been studied comparatively based on materials collected in the period from 1980 to 2017. The natural quantity of the species at the nesting period is greater in mixed and deciduous forests when compared to coniferous forests. Nesting density, clutch size, volume of eggs, success, and productivity of reproduction in nest boxes are higher in deciduous forests. The data on the natural quantity and breeding of pied flycatchers in the study area are compared with the similar indicators in the European part of the range. [ABSTRACT FROM AUTHOR]

Organisms need to time their annual-cycle stages, like breeding and migration, to occur at the right time of the year. Climate change has shifted the timing of annual-cycle stages at different rates, thereby tightening or lifting time constraints of these annual-cycle stages, a rarely studied consequence of climate change. The degree to which these constraints are affected by climate change depends on whether consecutive stages are causally linked (scenario I) or whether the timing of each stage is independent of other stages (scenario II). Under scenario I, a change in timing in one stage has knock-on timing effects on subsequent stages, whereas under scenario II, a shift in the timing of one stage affects the degree of overlap with previous and subsequent stages. To test this, we combined field manipulations, captivity measurements and geolocation data. We advanced and delayed hatching dates in pied flycatchers (Ficedula hypoleuca) and measured how the timing of subsequent stages (male moult and migration) were affected. There was no causal effect of manipulated hatching dates on the onset of moult and departure to Africa. Thus, advancing hatching dates reduced the male moult-breeding overlap with no effect on the moult-migration interval. Interestingly, the wintering location of delayed males was more westwards, suggesting that delaying the termination of breeding carries over to winter location. Because we found no causal linkage of the timing of annualcycle stages, climate change could shift these stages at different rates, with the risk that the time available for some becomes so short that this will have major fitness consequences. [ABSTRACT FROM AUTHOR]

Reproduction is energetically expensive and to obtain sufficient energy, animals can either alter their metabolic system over time to increase energy intake (increased-intake hypothesis) or reallocate energy from maintenance processes (compensation hypothesis). The first hypothesis predicts a positive relationship between basal metabolic rate (BMR) and energy expenditure (DEE) because of the higher energy demands of the metabolic system at rest. The second hypothesis predicts a trade-off between different body functions, with a reduction of the BMR as a way to compensate for increased daytime energetic expenditure.We experimentally manipulated the workload of wild pied flycatchers by adding or removing chicks when chicks were 2 and 11 days old. We then measured the feeding frequency (FF), DEE and BMR at day 11, allowing us to assess both short- and long-term effects of increased workload. The manipulation at day 2 caused an increase in FF when broods were enlarged, but no response in DEE or BMR, while the manipulation at day 11 caused an increase in FF, no change in DEE and a decrease in BMR in birds with more chicks. Our results suggest that pied flycatchers adjust their workload but that this does not lead to a higher BMR at night (no support for the increasedintake hypothesis). In the short term, we found that birds reallocate energy with a consequent reduction of BMR (evidence for the compensation hypothesis). Birds thus resort to short-term strategies to increase energy expenditure, which could explain why energy expenditure and hard work are not always correlated in birds. [ABSTRACT FROM AUTHOR]

• It is unclear if wild birds randomly or non-randomly collect nest materials. • We provided pied flycatchers with wool, feathers and deer hair close to their nests. • Nest building females collected the deer hair, whilst avoiding the wool and feathers. • Females thus showed strong preferences for some nest materials but not others. • Birds select nest material in a non-random manner from the wider environment. Nest building is a taxonomically widespread behaviour that consists of the construction of a suitable receptacle with collected materials for the incubation of eggs and sometimes for the raising of offspring. The use of specific nest materials has important fitness consequences for avian parents and offspring because they help to determine the thermal, parasitic and bacterial environment within nests and may also influence parental investment via intraspecific signalling. However, we presently know very little about the process by which nest materials are selected from the wider environment and specifically, it is unclear whether wild birds randomly or non-randomly select nest materials in relation to their local availability. Here, we report an experiment in which we provided experimental pairs of pied flycatchers (Ficedula hypoleuca) with wool, feathers and deer hair – commonly used nest materials found in their woodland habitats – close to their nests during the nest building period whilst control pairs were not provided with any materials. We found that females at experimental nests showed very clear preferences for deer hair, whilst almost completely avoiding the wool and feathers, thereby demonstrating that females exhibited very strong preferences for certain nest materials but not others. We therefore conclude that birds select nest materials in a non-random manner and do not simply use the materials most commonly available to them. [ABSTRACT FROM AUTHOR]

Prolonged stress can have long-lasting effects on an individual physiology and growth. However, the impact of chronically elevated glucocorticoids on the expression of early antipredator responses is still poorly documented. In this study I simulated the effect of repeated acute stress on offspring phenotype in free-living pied flycatchers (Ficedula hypoleuca) by administering adrenocorticotropic hormone (ACTH) to nestlings for 6 days. The results showed that frequent induction of stress responses by ACTH injections, independently of parental care, adversely affected offspring final body size, wing length and baseline corticosterone levels. Nestling behavioural activity did not differ between ACTHand saline-treated groups during exposure to control sounds while behavioural activity during exposure to alarm calls was reduced in manipulated offspring only. I conclude that prolonged physiological stress may have short-term benefits to nest-bound offspring, such as more effective antipredator behaviour, but at the expense of negative effects on body size and developmental speed. [ABSTRACT FROM AUTHOR]

Few studies on song learning have been conducted in the field. Male pied flycatchers (Ficedula hypoleuca) seem to learn song syllables in the breeding grounds. Adult males arrive first to the breeding grounds and sing more complex songs until they have paired. Early arriving juveniles may have more opportunities to learn new syllables from adult neighbours and develop more complex songs than later arriving juveniles. However, results suggest that arrival time is not important in song complexity development, but that males develop greater song complexity (repertoire size and song versatility) regardless as they age. [ABSTRACT FROM AUTHOR]

Singing is a key element of songbirds’ behavioral repertoire, particularly for males, which sing during the breeding season to defend resources against other males and to attract females. Different song traits may convey honest information about males’ qualities or conditions, which may be used by females to select their mates. Traits under strong sexual selection have an important component of additive genetic variation (i.e., the main genetic inheritance from parents), and so relatively high heritability; therefore, it can be expected that song traits also do. Although the act of singing is an innate behavior, and thus, genetically determined, songbirds need to learn their songs and therefore the genetic contribution to song traits may be reduced by the effect of environmental factors. We tested this hypothesis in seven song traits recorded in the long-distance migratory bird, the pied flycatcher (Ficedula hypoleuca). From a 23-year database (1992–2015), we obtained songs for 28 father–son pairs, and for each song trait we applied parent–offspring regressions to estimate heritability. The type of syllables sung are learned from tutors, and here we also determined the cultural contribution of fathers to the song repertoires of their sons, by quantifying the percentage of syllables that sons shared with their fathers, and compared this with what sons shared with other males in the population (e.g., neighbors). The heritabilities of song traits were highly variable (ranging from −0.22 to 0.56), but most of these were around zero and none of them were significant. These results indicate that the seven song traits are most likely determined by environmental factors. Sons shared more syllables with their fathers than with neighbors (21% vs. 3%), suggesting that fathers are important song tutors during the nestling period. We conclude that there is a cultural inheritance from fathers to their sons’ syllable repertoires, but there is no strong evidence for a genetic contribution of fathers to the seven song traits studied.