Pied Flycatcher Ficedula Hypoleuca Research Articles

Variation in climate warming along the migration route uncouples arrival and breeding dates

AbstractMigratory species are of special concern in the face of global climate change, since they may be affected by changes in the wintering area, along the migration route and at the breeding grounds. Here we show that migration and breeding times of a trans‐Saharan migrant, the pied flycatcher Ficedula hypoleuca, closely follow local temperatures along the migration route and at the breeding grounds. Because of differences in long‐term temperature trends of short within‐spring periods, the migration period and the time interval between migration and breeding dates of this species have extended in SW Finland. Temperatures in northern parts of Central Europe have risen at the time when the first migrants arrive there, facilitating their migration northward. Temperatures later in the spring have not changed, and the last individuals arrive at the same time as before. The timing of breeding has not advanced because temperatures at the breeding site after arrival have not changed. These results show that the pied flycatchers can speed up their migration in response to rising temperatures along the migration route. Our results strongly indicate that the effects of climate change have to be studied at the appropriate time and geographical scales for each species and population concerned.

Egg colouration and male parental effort in the pied flycatcher Ficedula hypoleuca

A recent hypothesis posits that the bright colours of many avian eggs may act like signals of female genetic quality or condition to males in species with biparental care, inducing them to work harder for their offspring. We measured the colour of blue eggs of pied flycatchers Ficedula hypoleuca in Central Spain with a spectrophotometer on the day of laying, and also quantified the provisioning effort by males and females during the last days of the nestling period. Both chromaticity coordinates in the CIELAB colour space (blue to yellow, green to red) showed significant associations with male provisioning rates, explaining more than 20% of variation in male parental effort. Male provisioning rates were positively correlated with nestling condition, thereby potentially contributing to female fitness. This evidence is only tentative until experimental confirmation, but suggests that males are affected by the colour of their mates’ eggs, a possibility not considered hitherto in the study of sexual selection.

Degradation of song in a species using nesting holes: the Pied Flycatcher Ficedula hypoleuca.

The habitat, but also the nest hole of a hole-nesting species, will degrade the song during transmission. We investigated how the sounds degrade in a sound transmission experiment with the song of the Pied Flycatcher Ficedula hypoleuca (Muscicapidae). Ten different song elements were transmitted to microphones placed inside and outside a nest box. On average, song degradation was much greater inside than outside the nest boxes, especially with respect to excess attenuation and blurring of the song elements. Being inside a nest box therefore strongly reduces a Pied Flycatcher's possibility of detecting and recognizing songs or eavesdropping on singing interactions.

Spatial distribution of breeding Pied Flycatchers Ficedula hypoleuca in respect to their natal sites

Spatial distribution of breeding Pied Flycatchers Ficedula hypoleuca in respect to their natal sites

A trade‐off between two resource‐demanding functions: post‐nuptial moult and immunity during reproduction in male pied flycatchers

Summary Costs of reproduction might be mediated by a resource‐allocation trade‐off between immunity and reproductive effort. Recent studies have suggested that the moult–breeding overlap observed mainly in males may affect this trade‐off. In order to test whether increased investment in immunity may affect the onset of moult or reproductive output in a species with moult–breeding overlap, we immunized breeding male pied flycatchers Ficedula hypoleuca (Pallas) with a non‐pathogenic antigen (sheep red blood cells) in the first week post‐hatching and compared their moult score, parental effort and reproductive success shortly before fledging with that of control males injected with sterile phosphate‐buffered saline. Immunized males provisioning nestlings delayed the onset of post‐nuptial primary moult compared to control males. The experimental activation of the immune defence of males affected their stress levels (heterophile/lymphocyte ratio, heat‐shock proteins) on day 13 after chicks hatched. An activated immune response tended to depress reproductive output, although the experimental design was not efficient enough to find a significant effect because the manipulation was not performed early enough in the nestling period. The results suggest that experimental activation of the males’ immune system decreased the resources necessary for initiating post‐nuptial moult. The trade‐off between immunity and moult onset may imply costs of delayed moult in cases of infection and may interact with reproductive immunosuppression in cases of moult–breeding overlap. Moult, immunity and reproduction are locked in a three‐way interaction which may markedly affect avian life histories.

Moult of three Palaearctic migrants in their West African winter quarters

Some theories about moult strategies of Palaearctic passerine migrants assume that birds adapt timing of moult to environmental conditions such as rainfall on their African wintering grounds. Species wintering in the northern tropics should limit moult to the period shortly after their arrival at the end of the rainy season. Passerine migrants wintering in West Africa should also moult more rapidly compared to related species or conspecific populations that moult elsewhere. We investigated the moult of melodious warblers Hippolais polyglotta, willow warblers Phylloscopus trochilus and pied flycatchers Ficedula hypoleuca wintering in Comoe National Park, Ivory Coast, between October 1994 and April 1998. In contrast to previous studies we did not restrict our analyses to moult of flight feathers but also included moult of body feathers. The results differed partially from the general assumptions of previous authors. Melodious warblers moulted twice: a complete moult shortly after their arrival, and a moult of body feathers and in some cases some tertials and secondaries in spring. Willow warblers moulting flight feathers were found between December and March with the majority moulting in January and February. Primary moult was not faster compared to populations moulting in central Africa and South Africa. Body feather moult varied strongly among individuals with birds in heavy moult between December and April. Pied flycatchers moulted body feathers and tertials between January and April. Birds with growing feathers were found throughout the whole period including the entire dry season. Moult strategies are thus not readily related to a few environmental factors in general and our results show that factors other than mere resource availability during certain times on the wintering grounds are likely to govern the timing of moult.

Cross-fostering of Pied Flycatchers (Ficedula Hypoleuca) to Heterospecific Hosts In the Wild: a Study of Sexual Imprinting

Learning from parents during early development may crucially influence future mate choice decisions of birds. Such sexual imprinting is thought to be important to many fields of evolutionary ecology, including sexual selection, hybridization, speciation, and interspecific brood parasitism. Most results have been obtained from controlled experiments on captive birds. Hence, there is a need to study sexual imprinting and the development of species recognition under more natural circumstances. I have cross-fostered a migratory bird, the pied flycatcher Ficedula hypoleuca, to nests of two resident species, the larger great tit Parus major, and the smaller blue tit P. caeruleus, by adding a single egg to the clutch, or by swapping whole clutches. No cross-fostered bird recruited to the local breeding population from the mixed brood treatment but sample size was small. However, for the whole brood treatment, cross-fostering had no apparent effect, compared to controls, on local recruitment, natal dispersal, mating success, breeding success, or sexual display responses to intruders. Hence, there were no signs that the cross-fostered birds were sexually imprinted on the host. The results are discussed in relation to sexual imprinting, natal dispersal, interspecific brood parasitism, and conservation of endangered birds.

Observer error in measurements of nestling wing length in small passerines

Observer error in measurements of nestling wing length in small passerines Wing lengths of nestlings of Great Tit Parus major, Blue Tit P. caeruleus and Pied Flycatcher Ficedula hypoleuca measured by one observer (WK) were compared with the measurements taken concurrently by one of 30 other observers. In total 1321 pairs of measurements were analysed. The differences between the measurers were found to: (1) depend on wing length; (2) vary between distinct wing-length classes and species; (3) be bigger in the case of inexperienced measurers (4) be small, only exceeding ± 1 mm in 7% cases and ± 2 mm in 0.5% cases. Such small measurement errors did not significantly bias the wing-length-based age estimation. The average differences between the age estimates derived from wing length taken by WK and other experienced measurers ranged from -0.3 to +0.3 day for various measurers, species and wing-length classes when 1-3 day-old nestlings (1-5 day-old in Blue Tit) were excluded. For the latter nestlings as well as for inexperienced measurers that range was -0.4-. +0.8 day.

Heterogeneous Maturation of Retina of the Pied Flycatcher Ficedula hypoleuca

A method is developed to determine the moment of establishment of functional maturity of cones by the appearance of their lipid droplet. Using this method, the process of differentiation of photoreceptor layer cells is traced and general regularities are elucidated of growth and maturation of the bifoveal retina of chicks of the pied flycatcher Ficedula hypoleuca. Development of retina occurs heterochronously; the first colored lipid droplets determining mature photoreceptors are photooptically revealed locally in the temporal retina area at the 5th day; at the 7th day, the lipid droplets appear within the limits of the central fovea; then the maturation wave moves from the temporal and central fovea towards periphery of retina. By the 14th day the entire area of retina turns out to be filled with mature photoreceptors. It is suggested that the predominant early maturation of photoreceptor cells of the temporal area is due to a necessity of the forestalling development of the retina parts providing the short-focus binocular vision that is the most important for establishment of chick alimentary behavior and start of hunting on their own after flight from the nest.

Beneficial Effects of Cloacal Bacteria on Growth and Fledging Size in Nestling Pied Flycatchers (Ficedula Hypoleuca) in Spain

AbstractEffects of bacteria on avian hosts in the wild have received little attention until recently. Whereas the pathogenic effects of bacteria are well known, positive effects of symbiotic bacteria are more rarely considered. Nestling growth has important repercussions for offspring fitness in avian populations and may be affected by microbial colonization of the gut. Enterococcus faecalis is a common opportunistic pathogen, whereas E. faecium has been used as a growth promoter because it interacts competitively with pathogenic bacteria, E. faecalis included. We followed the growth in tarsus length and mass of 18 Pied Flycatcher (Ficedula hypoleuca) broods. Chicks were weighed and measured on days 4, 8, 10, and 13 after hatching. On day 13, wing length was also measured and cloacal swabs were taken of two chicks in each brood for detection of enterococci. In all, the methods used allowed us to detect six species of bacteria among a possibly much richer community. Most chicks had E. faecalis, whereas E. faecium was less prevalent. There was a negative association between scores for E. faecalis and for the rest of the species pooled. The presence of E. faecalis showed no detectable association with nestling mass or size at any age, whereas the presence of the other species showed significantly positive associations with mass and size on day 13, but not before. Presence of E. faecium on its own was positively associated with nestling mass and size shortly before fledging. E. faecium may act as a growth promoter in the wild through its competitive interactions with facultative pathogenic bacteria. The presence of some microbes are critically important in avian growth and development.

Seasonal trade-offs in cell-mediated immunosenescence in ruffs (Philomachus pugnax).

The immune system is an energetically expensive self-maintenance complex that, given the risks of parasitism, cannot be carelessly compromised. Life-history theory posits that trade-offs between fitness components, such as self-maintenance and reproduction, vary between genders and age classes depending on their expected residual lifetime reproductive success, and seasonally as energetic requirements change. Using ruff (Philomachus pugnax), a bird with two genetically distinct male morphs, we demonstrate here a decrease in male immunocompetence during the breeding season, greater variance in immune response among males than females, immunosenescence in both sexes and male morphs, and a seasonal shift in the age range required to detect senescence. Using a phytohaemagglutinin delayed hypersensitivity assay, we assessed cell-mediated immunity (CMI) of males of typical breeding age during the breeding and nonbreeding seasons, and of a larger sample that included females and birds of a greater age range during the non-breeding period. CMI was higher for breeding-aged males in May than in November, but the increase was not related to age or male morph. In November, mean CMI did not differ between the sexes, but the variance was higher for males than for females, and there were no differences in mean or variance between the two male morphs. For both sexes and male morphs, CMI was lower for young birds than for birds of typical breeding ages, and it declined again for older birds. In males, senescence was detected in the non-breeding season only when very old birds were included. These results, generally consistent with expectations from life-history theory, indicate that the immune system can be involved in multifarious trade-offs within a yearly cycle and along an individual's lifetime, and that specific predictions about means and variances in immune response should be considered in future immunoecological research.

Do birds behave in order to avoid disclosing their nest site?

Capsule Three similarly sized passerine species with various breeding strategies behaved differently in the presence of models of mammalian and avian predators. Aims We tested whether solitary breeders behave in the presence of a predator in order to avoid disclosing their nest site in: hole-nesting Pied Flycatcher Ficedula hypoleuca, shrub-nesting Blackcap Sylvia atricapilla and ground-nesting Meadow Pipit Anthus pratensis. Methods The behaviour of parents in response to two types of predator model (visually oriented Corvid birds and olfactory oriented Mustelid mammals) placed sequentially at 40, 5 and 1 m distance from the nest was recorded from a shelter. Results The hypothesis was supported in open-nesting species (Meadow Pipit and Blackcap), as parents did not approach the nest in the presence of a predator. In the hole-nesting Pied Flycatcher the parents disclosed the nest site in most cases (by entering the nest). The intensity of alarm calling increased with decreasing distance of a predator from the nest in all species except Meadow Pipit in the presence of Raven Corvus corax models. The intensity of attacking changed only in Meadow Pipit with decreasing distance of Stoat Mustela erminea from the nest. Conclusion The results showed that anti-predator behaviour was species-specific, depending on type of predator, habitat and nest inaccessibility.

Large‐scale effect of climate change on breeding parameters of pied flycatchers in Western Europe

Variations in laying date and clutch size of pied flycatchers Ficedula hypoleuca across populations throughout western Europe are examined in relation to climatic fluctuations, measured by the winter North Atlantic Oscillation (NAO) index. Across breeding sites, the winter‐NAO index affected laying date such that females lay earlier after warmer and moister winters (positive values of winter NAO‐index). Female pied flycatchers breed progressively earlier because presumably the whole breeding season is being shifted, as a direct result of the positive values of winter NAO‐index. Moreover, clutch size of pied flycatchers across populations was negatively related to winter NAO‐index during the last 50 yr. These analyses controlled for potentially confounding variables such as latitude, longitude, elevation and habitat of each study site. The present study conclude that pied flycatchers across western Europe are breeding earlier and laying smaller clutch sizes and that the most likely cause is a long‐term increase in spring temperature. On the other hand, this study shows that climate change may not act uniformly between breeding populations in Western Europe. From those results, this study concludes that northern pied flycatcher populations are more sensitive to climate change than southern populations breeding in montane habitats.

Impact of busy roads on breeding success in pied flycatchers Ficedula hypoleuca.

The impact of dense traffic on the breeding success in pied flycatchers (Ficedula hypoleuca) was studied using roadside nest-boxes. Nest site selection and breeding success of flycatchers were observed in relation to the distance from the road. The number of occupied territories was no higher closer to the road than it was deeper inside the forest. The distance to the road had no effect on the laying date, clutch size, or brood size. However, nests closer to the road were more likely to fail at the chick stage. The number of broods that were closer to roads and were lost completely was significantly higher than those further away. As a consequence, the number of fledglings per breeding attempt decreased closer to roads. The nestlings typically died as older chicks when the parent birds were providing them with maximum quantities of food. A possible reason for the declining reproductive success was the traffic-related mortality of parent birds, as weather conditions were not especially adverse for raising healthy offspring.

Does supplementary calcium reduce the cost of reproduction in the Pied Flycatcher Ficedula hypoleuca?

Studies in acidified as well as in naturally base‐poor areas have recently revealed that availability of extra calcium‐rich food items is an important component of habitat quality affecting breeding performance in several bird species. However, these mostly short‐term studies have provided equivocal results concerning the exact consequences of calcium shortage on different species in different regions. We studied the effect of calcium availability on reproduction of the Pied Flycatcher Ficedula hypoleuca breeding in pine forests in Estonia, NE Europe, over a period of 4 years. Experimental pairs were provided with supplementary calcium‐rich material when breeding, while control pairs were left unsupplemented. Experimental females laid larger eggs and their nestlings had longer tarsi than those of controls. Moreover, the mass and condition of females tending larger than average clutches were increased by calcium‐supplementation. Our results provide the first experimental evidence that calcium availability may affect the overall cost of reproduction in free‐living passerines. We compared these results with similar data for the Great Tit Parus major, collected from the same area during the same study period. Great Tits responded to low calcium availability mainly by restrained reproductive behaviour and reduced breeding success, while Pied Flycatchers invested significantly more in current reproductive effort despite the increased cost of reproduction. Thus, the effects of calcium deficiency on birds seem to be species‐specific or population‐specific. This partly explains discrepancies between the results of earlier studies.

Mate choice in Pied Flycatchers Ficedula hypoleuca: can females use song to find high‐quality males and territories?

High‐quality male Pied Flycatchers Ficedula hypoleuca (defined by brighter plumage, better condition and more experience) have previously been shown to have larger syllable repertoires and greater song versatility than males of inferior quality. Thus, by preferring more complex songs, females could choose a high‐quality male. Females may also use song as a cue to find a high‐quality territory since early arriving males may obtain the best territories and these males have more complex songs than late‐arriving males. We found that males with more complex songs had a greater chance of becoming paired and stayed unpaired for a shorter period than males with less elaborate songs. When controlling for arrival order, however, only strophe versatility was still correlated with pairing order. Males defending popular territories had more complex and longer songs and were also in better body condition than males in less popular territories. A multiple logistic regression showed that song length was important in explaining whether a male defended a popular nestbox or not. Thus, male arrival time seems to be important in deciding the quality of a male's territory, which in turn explains female choice. However, song quality seems to add important information. Thus, females could find both high‐quality males and high‐quality territories by using song cues during mate choice.

Different responses to cold weather in two pied flycatcher populations

We compared how breeding parameters differ according to prevailing weather conditions between a marginal, subarctic (69°N) and temperate (61°N) population of the pied flycatcherFicedula hypoleuca,a small migratory insectivorous passerine. We predicted that the effects of weather on breeding performance (clutch size, hatching success, nestling growth, fledging success) would be greater at northern latitudes, where the weather conditions are more extreme and unpredictable. We found that the breeding parameters, except clutch size, were not, however, inferior in the north. Northern birds, unlike the southern ones, responded to colder conditions by laying smaller clutches and maintaining a larger energy reserve (indicated by higher female body mass and higher levels of subcutaneous fat). If a cold spell occurred during the nestling period, southern flycatchers had 5–10% lower fledging success than the northern ones. Our results indicate that in the north, the breeding individuals coped with cold and variable weather better than the individuals in the southern population. This could be adaptive, because at high latitudes there is a higher probability of cold weather at the time of breeding.

Gender and viability selection on morphology in fledgling pied flycatchers.

Until recently, analyses of gender-dependent differences in viability selection and the ontogeny of sexual size dimorphism have been plagued by difficulties in determining the sex of nestling birds on the basis of morphology. Recently, this problem was overcome using molecular sex identification to report for the first time body-size-mediated antagonistic selection on the viability of male and female collared flycatchers. We used molecular sex identification to analyse natural selection on fledgling viability, sexual size dimorphism and effects of parasites in relation to gender in a Mediterranean population of the related pied flycatcher Ficedula hypoleuca. There was directional positive selection on fledgling weight but no selection on tarsus length. Fledgling weight was the most important determinant of fledgling survival, with heavier fledglings having increased viability. Although selective trends were of the same sign for both sexes, only among female fledglings were selection differentials and gradients statistically significant. Therefore, similar trends in selection were revealed in analyses of a data set where sex was ignored and in separate analyses using same-sex sibship trait means. Mite nest ectoparasites negatively affected fledgling weight, and the effects were stronger in female than male fledglings. There was no effect of parasitism on the tarsus length in males, as previously reported in retrospective analyses performed without knowledge of sex until recruitment. Overall, selection on fledgling viability on the basis of morphological traits and hatching date was not confounded by an individual's gender.

Impaired flight ability during incubation in the pied flycatcher

During the breeding season, many female passerine birds increase in body mass before egg laying, maintain a relatively high body mass during incubation, and then drop back to the original level during the chick‐rearing period. The post‐hatching reduction in body mass, which can be as large as 10–20%, has been suggested to represent an adaptive mass loss to reduce wing loading, thereby increasing parental flight efficiency when chicks have hatched and have to be fed. Here we present the first study of changes in flight ability from incubation to chick rearing in birds. Wild female pied flycatchers Ficedula hypoleuca flew more slowly during incubation than during chick rearing; a 7% reduction in body mass after the chicks had hatched was associated with a 10% increase in vertical take‐off speed. Furthermore, the flight muscle size of the females tracked the reduction in wing load, suggesting that muscle size was adaptively reduced when no longer needed. Since incubation‐feeding by males reduces the time females have to spend outside the nest foraging, our results suggest that in addition to increasing female nutritional status and reducing incubation time, incubation‐feeding will also reduce predation risk during the period when females face impaired flight ability.

Predicting Life-Cycle Adaptation of Migratory Birds to Global Climate Change Coppack T. & C. Both 2002. Predicting life-cycle adaption of migratory birds to global climate change. In: Both C. & T. Piersma (eds) The avian calendar: exploring biological hurdles in the annual cycle. Proc. 3rd Conf. European Ora. Union, Groningen, August 2001. Ardea 90(3) special issue: 369–378.

Analyses of long-term data indicate that human-caused climatic changes are affecting bird phenology in directions consistent with theoretical predictions. Here, we report on recent trends in the timing of spring arrival and egg laying found within a western European Pied Flycatcher Ficedula hypoleuca population. Mean egg laying date has advanced over the past 20 years in this population. The advancement in egg laying date was stronger than the advancement of spring arrival, suggesting that Pied Flycatchers are changing these stages of their annual cycle at different rates. It could be shown that selection for earlier breeding had increased. Hence, the observed adjustment in laying date did not match the advancement of spring. Our findings raise general questions about the adaptability of migratory birds to rapid environmental changes. Adaptive advancement of reproduction in response to increasing spring temperatures and to the concomitant advancement of food supply could be held back, because annual breeding and migration cycles are controlled primarily by endogenous rhythms and photoperiodic cues which do not relate to temperature. Migrants may have several options for arriving earlier on the breeding grounds, including an increase in migration speed, earlier departure from the wintering area or a shortening of migration distance. Changes in migratory behaviour could be accomplished either by phenotypic plasticity or by selection on different genotypes. Although descriptive field data provide compelling evidence for changes in, and possible constraints on, the timing of breeding and migration, their explanatory power in predicting the limits of adaptation remains restricted. We review recent experimental approaches, which explicitly test the relative roles of genetic versus environmental factors in the adaptation of life-cycle timing to global environmental changes.