Pied Flycatcher Ficedula Hypoleuca Research Articles

Extra-pair matings, context-dependence and offspring quality: a brood manipulation experiment in pied flycatchers

Extra-pair paternity (EPP) in socially monogamous species has been associated with the genetic benefits incurred by females through extra-pair mate choice. There is conflicting evidence in the literature concerning the importance of such benefits possibly due to their context-dependence. To ascertain if there are such context-dependent genetic benefits, we conducted a brood manipulation experiment in pied flycatchers (Ficedula hypoleuca) breeding in central Spain. We found that extra-pair mates (EPM) are frequently not nearest neighbours. Cuckolded males and EPM did not differ from other males in any measured trait, and were not different either when cuckolds and cuckolders were compared within broods. Within-pair offspring (WPO) and extra-pair offspring (EPO) did not differ with respect to heterozygosity and no association of heterozygosity of either offspring or males and females with EPP could be detected. Moreover, the competitive context in the nest induced by brood manipulations affected the growth of nestlings and showed an interaction with paternity differences in nestling mass but in the opposite direction to the prediction of context-dependence, i.e., EPO were lighter than WPO in enlarged broods but not in control broods. Furthermore, EPO had shorter tarsi than WPO in all treatments. Mortality in the nest showed an association with brood manipulation treatment, but not with paternity or its interaction with treatment. Thus, we have not found any evidence of genetic benefits, context-dependent or otherwise, of EPP in our study population. The evidence of poor quality EPO does not support a mixed reproductive strategy of females in our population.

Only Some Ectoparasite Populations are Affected by Nest Re-Use: an Experimental Study on Pied Flycatchers

Summary. Nest re-use in birds is rare but since appropriate cavities may be scarce, cavity-nesting birds may often re-use those that were occupied in previous seasons. Old nest material may contain and/or attract more ectoparasites than fresh material. Therefore it is important to understand the effects of nest re-use on the abundance of different ectoparasite species of different virulence and their impli cation for breeding parameters and nestling condition. We studied the consequences of nest re-use in a population of pied flycatchers Ficedula hypoleuca breeding in central Spain by offering them both nest-boxes with old nest material and cleaned nest-boxes. We monitored breeding activity from the early stages of nest construction until fledging, and then finally removed nests to estimate ectoparasite abun dances. Occupation rates were similar for both treatments. We found that blowfly and flea abundances were significantly higher in old nests than in new nests, but the abundance of mites, the most viru...

Variation in eggshell traits between geographically distant populations of pied flycatchers Ficedula hypoleuca

The expression and impact of maternal effects may vary greatly between populations and environments. However, little is known about large‐scale geographical patterns of variation in maternal deposition to eggs. In birds, as in other oviparous animals, the outermost maternal component of an egg is the shell, which protects the embryo, provides essential mineral resources and allows its interaction with the environment in the form of gas exchange. In this study, we explored variation of eggshell traits (mass, thickness, pore density and pigmentation) across 15 pied flycatcher populations at a large geographic scale. We found significant between‐population variation in all eggshell traits, except in pore density, suggesting spatial variation in their adaptive benefits or in the females’ physiological limitations during egg laying. Between‐ population variation in shell structure was not due to geographic location (latitude and longitude) or habitat type. However, eggshells were thicker in populations that experienced higher ambient temperature during egg laying. This could be a result of maternal resource allocation to the shell being constrained under low temperatures or of an adaptation to reduce egg water loss under high temperatures. We also found that eggshell colour intensity was positively associated with biliverdin pigment concentration, shell thickness and pore density. To conclude, our findings reveal large‐ scale between‐population variation of eggshell traits, although we found little environmental dependency in their expression. Our findings call for further studies that explore other environmental factors (e.g. calcium availability and pollution levels) and social factors like sexual selection intensity that may account for differences in shell structure between populations.

You mob my owl, I’ll mob yours: birds play tit-for-tat game

Reciprocity is fundamental to cooperative behaviour and has been verified in theoretical models. However, there is still limited experimental evidence for reciprocity in non-primate species. Our results more decisively clarify that reciprocity with a tit-for-tat enforcement strategy can occur among breeding pied flycatchers Ficedula hypoleuca separate from considerations of byproduct mutualism. Breeding pairs living in close proximity (20–24 m) did exhibit byproduct mutualism and always assisted in mobbing regardless of their neighbours' prior actions. However, breeding pairs with distant neighbours (69–84 m) either assisted or refused to assist in mobbing a predatory owl based on whether or not the distant pair had previously helped them in their own nest defense against the predator. Clearly, these birds are aware of their specific spatial security context, remember their neighbours' prior behaviour, and choose a situation-specific strategic course of action, which could promote their longer-term security, a capacity previously thought unique to primates.

Oxidative damage in relation to a female plumage badge: evidence for signalling costs

Badges of status may be controlled by costs derived from increased aggression from dominant individuals. This cost could be translated into elevated metabolic levels and a concomitant disruption of oxidative balance. Some females in Iberian pied flycatcher Ficedula hypoleuca populations exhibit a white forehead patch similar to that exhibited by all males in this species, functioning in aggressive interactions between females when competing for breeding sites. To test if social stress imposes costs on signalling, we painted white patches on females without natural patches (NP) and compared them with females with natural control (NU). We also over-painted the natural patch in other females (FP) and compared to females with control natural patches (FU). We obtained for the whole sample of females data on reproductive investment, morphology and oxidative damage measured by blood malondialdehydes (MDA), and in a subsample of females variables related to parental care during incubation and the early nestling stage. FP and FU did not differ significantly in any variable which negates an effect of paint itself. However, NP females showed significant higher levels of MDA than NU females when controlling for breeding success for the whole sample, and for female incubation attendance for the parental care subsample. When including the four treatments, there was a significant interaction between the paint treatment and the presence/absence of badges before the experiment when controlling for the significant negative effect of incubation attendance on MDA. Addition of a badge to females without one leads to increased oxidative damage possibly mediated by social control. Badges of status in female pied flycatchers may operate as badges of oxidative status.

Quantitative genetics of basal metabolic rate and body mass in free‐living pied flycatchers

AbstractDespite basal metabolic rate (BMR) being one of the most commonly measured physiological traits and an important indicator of competitive ability, very little is known about its genetic basis and relation to other physiological traits. Here, we present the first attempt to estimate the multivariate basis of BMR using a natural population of pied flycatcher Ficedula hypoleuca breeding in the Tomsk Region, Western Siberia. We show relatively high and significant heritability of whole‐organism BMR, mass‐specific BMR and mass‐independent BMR (h 2 = 0.43, 0.55 and 0.52, respectively), which indicates the potential of these energetic traits to respond to direct selection. In contrast to some previous reports, we found that the genetic correlations between body mass and all three measures of BMR were not significantly different from zero. Independent evolution of body mass and BMR in this species should therefore be possible. Following a previous report, we also estimated the genetic correlations between the different BMR measures and show they are all close to unity, suggesting that they are, from a genetic point of view, a similar trait. Our results are in contrast with previous studies measuring the genetic basis of metabolic rates using aviary‐bred birds and highlight the importance of considering BMR in a natural setting.

Sources of Variation in Enterococci andEnterobacteriaceaeLoads in Nestlings of a Hole-Nesting Passerine

Gut bacterial loads in avian nestlings may be affected by factors such as climate, seasonality and brood size. There is no published information on this subject for wild birds, despite its potential importance for nestling welfare and growth. We studied the associations of these factors with abundances of two common gut bacterial types, enterococci and Enterobacteriaceae, in nestling Pied Flycatchers Ficedula hypoleuca in central Spain. To that end, we obtained faecal samples from 54 broods (102 nestlings) on day 7 and 13 after hatching, for detection and estimation of bacterial abundance. Enterobacteriaceae loads on day 7 were positively correlated with mean temperature during the preceding seven days and negatively correlated with rainfall and hatching date. The negative associations of Enterobacteriaceae loads with rainfall were only found for early broods, at low temperatures. Enterococci loads on day 7 were positively associated with mean temperature. On day 13, Enterobacteriaceae loads were negatively correlated with hatching date, while enterococci loads were positively correlated with mean temperature. Enterobacteriaceae are apparently more sensitive to seasonal changes and climatic variation than enterococci, possibly in relation to variation in diet and nutrition. By contrast, enterococci are only sensitive to thermal variation. The attainment of full thermoregulatory capacity by nestlings reduces climatic effects on Enterobacteriaceae loads.

Climate change, breeding date and nestling diet: how temperature differentially affects seasonal changes in pied flycatcher diet depending on habitat variation

1. Climate warming has led to shifts in the seasonal timing of species. These shifts can differ across trophic levels, and as a result, predator phenology can get out of synchrony with prey phenology. This can have major consequences for predators such as population declines owing to low reproductive success. However, such trophic interactions are likely to differ between habitats, resulting in differential susceptibility of populations to increases in spring temperatures. A mismatch between breeding phenology and food abundance might be mitigated by dietary changes, but few studies have investigated this phenomenon. Here, we present data on nestling diets of nine different populations of pied flycatchers Ficedula hypoleuca, across their breeding range. This species has been shown to adjust its breeding phenology to local climate change, but sometimes insufficiently relative to the phenology of their presumed major prey: Lepidoptera larvae. In spring, such larvae have a pronounced peak in oak habitats, but to a much lesser extent in coniferous and other deciduous habitats. 2. We found strong seasonal declines in the proportions of caterpillars in the diet only for oak habitats, and not for the other forest types. The seasonal decline in oak habitats was most strongly observed in warmer years, indicating that potential mismatches were stronger in warmer years. However, in coniferous and other habitats, no such effect of spring temperature was found. 3. Chicks reached somewhat higher weights in broods provided with higher proportions of caterpillars, supporting the notion that caterpillars are an important food source and that the temporal match with the caterpillar peak may represent an important component of reproductive success. 4. We suggest that pied flycatchers breeding in oak habitats have greater need to adjust timing of breeding to rising spring temperatures, because of the strong seasonality in their food. Such between-habitat differences can have important consequences for population dynamics and should be taken into account in studies on phenotypic plasticity and adaptation to climate change.

Bacterial degradability of an intrafeather unmelanized ornament: a role for feather-degrading bacteria in sexual selection?

The impact of feather-degrading bacilli on feathers depends on the presence or absence of melanin. In vitro studies have demonstrated that unmelanized (white) feathers are more degradable by bacteria than melanized (dark) ones. However, no previous study has looked at the possible effect of feather-degrading bacilli on the occurrence of patterns of unmelanized patches on otherwise melanized feathers. The pied flycatcher Ficedula hypoleuca Pallas, 1764 is a sexually dimorphic passerine with white wing bands consisting of unmelanized patches on dark flight feathers. These patches are considered to be a sexually selected trait in Ficedula flycatchers, especially in males, where the patches are more conspicuous (larger and possibly whiter) than in females. Using in vitro tests of feather bacterial degradation, we compared the degradability of unmelanized and melanized areas of the same feather for 127 primaries collected from the same number of individuals in a population breeding in central Spain (58 males and 69 females). In addition, we also looked for sex differences in feather degradability. Based on honest signalling theory and on the fact that there is stronger sexual selection for males to signal feather quality than in females, we predicted that unmelanized areas should be more degradable by bacteria than melanized ones within the same feather, and that these unmelanized areas should also be more degradable in males than in females. We confirmed both predictions. Microstructural differences between cross-section dimensions of unmelanized and melanized barbs, but not differences in the density of barbs within unmelanized and melanized areas of feathers in males and females, could partly explain differences in degradability. This is the first study to show differences in bacterial degradability among markings on the same feather and among unmelanized feather patches between males and females as predicted by sexual selection theory.

Parental infanticide in birds through early eviction from the nest: rare or under‐reported?

Avian parental infanticide is apparently a rare phenomenon judging from the scant reports in the literature. However, unexplained cases of single nestling disappearance or of apparent fall from the nest are rather common and are routinely attributed to parental eviction of dead nestlings, single chick predation or dislodging of nestlings through errors by nestlings or parents. During long‐term studies of pied flycatchers Ficedula hypoleuca in central Spain 29 cases of eviction from the nest cup of mostly hatchlings or small, heterothermic nestlings were recorded, leading invariably and quickly to death. Parental errors and failed attempts to predate single chicks are unlikely in these cases. Although the incidence of recorded evictions was mostly below 5% of nests, it reached higher levels in years and populations where reproductive success was low. Some cases were preceded by prior egg eviction. In these studies, there were unexplained disappearances affecting 3.7% of eggs laid which amounted to 20% of all losses. During studies of three other passerine cavity nesting species, similar cases were observed. The rarity of reports of parentally evicted nestlings is apparently based on the difficulty of directly observing parents evicting live nestlings but may also be due to observer bias as ejections of dead nestlings, erroneous dislodgings or single‐chick predation events are also rarely observed but are normally given as reasons for single‐chick losses. Field researchers should attempt to determine the real causes of partial brood loss before assuming that parsimony excludes parental infanticide through eviction.

Dynamics and reproduction of a nest-box breeding population of Pied Flycatcher Ficedula hypoleuca in a subalpine birch forest in Swedish Lapland during a period of 46 years

A nest-box breeding population of Pied Flycatcher Ficedula hypoleuca in subalpine birch forest at Ammarnäs in northern Sweden was studied in 1965–2010. The population showed on average a significantly decreasing trend during the initial 25 years, after which it remained on a static level. The yearly variation of the population size was significantly correlated with breeding result in preceding years, which in turn was affected by e.g. nest predation, adverse climatic factors, clutch size, and incidence of defective egg shell formation. It is also suggested that the population decline during the initial 10–15 study years followed from over-establishment of the breeding habitat. The average breeding result was 2.3 fledglings per pair, which is far below the number needed for the population to be self-reproducing. The occurrence of unusually high rates of nest predation and defective egg shells seems to be passing phenomena, predicted to approach zero after about 30 and 60 years, respectively, after the start of the study. But even with these factors eliminated and with other conditions remaining unchanged, the Ammarnäs population will continue to be a sink population.

Correlations between long distance migrants in year-to-year fluctuations of arrival timing

This study investigated the year-to-year fluctuations in arrival timing in the UK of 23 species of summer migrants. Median arrival dates, estimated from BirdTrack survey data from 2002–10, and Cheshire and Wirral first-arrival dates were analysed using Pearson correlations. Highly statistically significant correlations were found for pairings within a group comprising some warblers and the Pied Flycatcher Ficedula hypoleuca, in particular for pairings of species which have similar average arrival dates. In contrast, no significant correlations were observed for pairings involving the earlier-arriving Willow Warbler Phylloscopus trochilus and later-arriving Spotted Flycatcher Muscicapa striata. No significant correlations with any other warbler species were apparent for a third species, Lesser Whitethroat Sylvia curruca, which, unlike the other species, follows an easterly migration path through the Middle East before travelling northwest across Greece, northern Italy and central Europe. It is suggested that these correlations in arrival timing may arise from the within-year similarities in local weather conditions encountered en route by species with similar spring migration strategies. Significant correlations were also identified between some other pairings, for example those of Common Cuckoo Cuculus canorus, Barn Swallow Hirundo rustica, House Martin Delichon urbicum and Tree Pipit Anthus trivialis with various other species, but were generally lacking for some other species.

Prevalence ofSalmonellaandYersiniain Free-Living Pied Flycatchers (Ficedula hypoleuca) in Central Spain

Salmonella and Yersinia are important enteropathogens in poultry and can affect birds of all ages, including embryos. These food-borne zoonotic enteropathogens are of great economic and medical concern worldwide and are intensely studied in poultry. Information regarding the prevalence of these bacteria in wild birds is scarce and biased toward avian species ecologically linked to humans, which have often been incriminated as both reservoirs and disseminators of these enteropathogens. The prevalence of Salmonella and Yersinia recovered from both the feces and eggs in a population of female pied flycatchers (Ficedula hypoleuca) breeding in nest-boxes in central Spain was evaluated. Salmonella arizonae was recovered from the feces of one female but was not recovered from eggs. Yersinia was not detected in either the feces or eggs. These results may suggest that Salmonella and Yersinia may be uncommon in this population studied and may indicate that these birds are unlikely reservoirs of Salmonella and Yersinia.

Handicapped females receive more feedings during incubation from their mates: support for the female nutrition hypothesis

The female nutrition hypothesis posits that provisioning intensity of incubating females by their mates may depend on female needs and ensure proper incubation and a corresponding high hatching and breeding success of breeding pairs. Here, we have handicapped female pied flycatchers Ficedula hypoleuca at the beginning of incubation by clipping two primaries on each wing and filmed nests during incubation and later nestling provisioning to estimate male involvement in incubation feeding at the nest and in offspring care. Incubation feeding was more frequent at late nests. Correcting for this seasonal effect, incubation feeding was significantly affected by treatment and twice as high at experimental as at control nests. There was no effect of the experiment on female incubation attendance. The handicap did not result in any effect on hatching and breeding success, nestling growth and male or female provisioning and mass at the end of the nestling period. Males adjust their incubation feeding activity at the nest to female energetic requirements during incubation.

A rare case of double‐brooding in a Pied Flycatcher Ficedula hypoleuca

Capsule A report of double‐brooding in a Pied Flycatcher Ficedula hypoleuca in Central Poland.

A 64-year study of a Pied Flycatcher Ficedula hypoleuca population

A nestbox breeding population of Pied Flycatchers Ficedula hypoleuca was monitored during 64 years in a mixed coniferous-deciduous forest area near Örebro, South Central Sweden. The population showed a continuous average decline of c. 0.5 pairs per year, from c. 50% to c. 20% occupancy. There was a strong negative correlation between flycatcher numbers and those of other hole-nesters, but it is doubtful whether any causal relationship existed. Mean laying date was 24 May (17 May—3 June) and showed a negative correlation with mean May temperatures, yet no significant trend over the six decades. Mean laying date did, however, show an increased variation during the last 20 years. Mean clutch size varied between 5.76 and 7.08, with a mean of 6.34. It varied more during the last 30 years but without any significant relationship with mean laying date. A mean of 5.96 young fledged from broods that produced at least one fledgling vs 5.34 for all broods; the lower figure was mainly a consequence of nest predation by Pine Marten Martes martes, particularly after the early 1980s.

Do Male Pied Flycatchers (Ficedula hypoleuca) Adjust Their Feeding Effort According to Egg Colour?

The colour of blue-green bird eggs has been hypothesized to signal female quality to attending males, who may adjust their level of investment in the brood accordingly. The hypothesis has gained support in studies of Spanish pied flycatchers Ficedula hypoleuca. We performed a cross-fostering experiment in a Norwegian population of pied flycatchers to provide an independent test of the sexually selected egg colour hypothesis in this species. Egg colour was not significantly correlated with estimates of female quality (clutch size, average egg volume, first egg laying date and feeding rate). There was a significant decrease in chroma and increase in brightness and egg volume during the laying sequence, with some marked differences between six-egg and seven-egg clutches that might reflect differences in female quality. However, we found no significant influence of egg colour (foster or original clutch) on male feeding rate or offspring viability (hatching success, average mass and fledging success) and no significant difference in feeding rate for males with six-egg and seven-egg foster or original clutches. We conclude that Norwegian male pied flycatchers do not use egg colour as a cue to female quality, thus questioning the generality of previous support for the sexually selected egg colour hypothesis from this species.

Estimation of heritability and repeatability of resting metabolic rate in birds by the example of free-living pied flycatchers Ficedula hypoleuca (Aves: Passeriformes)

Repeatability of a trait estimated over long time intervals and its heritability make it possible to assess whether it can serve as an individual characteristic and be subject to selection. Heritability and repeatability of energetic traits are still poorly studied in birds. The most important physiological characteristic of the homoiotherms is the minimal level of energy expenditure, referred to as the resting metabolic rate (RMR), which, in the absence of the expenditure for biomass growth, does not exceed the basal metabolic rate (BMR). We have estimated the BMR repeatability in adult free-living pied flycatchers in Moscow-region (55°44′ N, 36°51′ E; 1992–2008) and Tomsk (56°20′ N, 84°56′ E; 2008–2009) populations over intervals of 40 days to 3 years. In the Moscow-region pied flycatcher population, the BMR repeatability recorded during the same period of the reproductive season was τ = 0.34 ± 0.10 for a 1-year interval (n = 80), τ = 0.60 ± 0.15 for a 2-year interval (n = 19), and τ = 0.85 ± 0.13 for a 3-year interval (n = 6). In the Tomsk population, the BMR repeatability for the 1-year interval was τ = 0.49 ± 0.11 (n = 50). The repeatability of a trait is a measure of its constancy in time and specifies the upper limit of its heritability. RMR heritability was assessed in cross-fostering experiments in the Moscow-region free-living pied flycatcher population in 2003–2005. The RMRs of chicks and BMRs of their biological fathers displayed a positive correlation, whereas no correlation was found between the RMRs of chicks and their foster fathers. The RMR heritability value was h2 = 0.43 ± 0.17 (n = 210). The obtained estimates for repeatability and heritability of fundamental energetic traits in pied fly-catcher are rather high for a physiological trait and suggest the existence of evolutionary stable diversity of the avian population in the BMR.

Vocal Plasticity - are Pied Flycatchers, Ficedula Hypoleuca, Open-Ended Learners?

In some songbird species, large song repertoires are advantageous in female attraction, whereas song sharing with neighbours may give an advantage in male–male competition. Open-ended learners, with the ability to memorize new song elements throughout their lives, may learn from territorial neighbours and thus benefit from increasing both repertoire size and song sharing. A distinction needs to be made between true adult song learning, i.e. memorization of novel song elements, and vocal plasticity resulting in changes in the use of previously memorized elements, such as the use of hidden repertoires or increased production of previously rare syllable types. We assessed the ability of adult pied flycatcher Ficedula hypoleuca males to learn previously unheard song elements and to change their song production in response to playback of unfamiliar, conspecific song, emulating a singing neighbour. After a 1-week playback treatment, three out of 20 subjects had learned foreign song elements, providing evidence from the wild that pied flycatchers are true open-ended learners. However, the syllable sharing with the playback stimulus repertoires had not changed, and the males’ repertoires had decreased rather than increased. Hence, we did not find support for increased syllable sharing with neighbours or increased repertoire size as functions of adult song learning in pied flycatchers. Because pied flycatcher song seems to serve mainly for mate attraction, copying of attractive syllable types is a possible alternative function of adult song learning in this species.

Nest Construction Costs Affect Nestling Growth: A Field Experiment in a Cavity-Nesting Passerine

Nest construction effort is seldom taken into account in avian life history studies, although activity duration and the amount of nest material collected may affect the condition of parent birds and their subsequent capacity to care for dependent offspring. In the present work, we reduced the effort needed to construct the nest in a cavity nester, the Pied Flycatcher Ficedula hypoleuca, where females are the main builders. To this end, we placed stored complete nests in certain nest-boxes as soon as nest building was initiated, whereas stored complete nests were placed in the control nests-boxes as soon as nest building had ceased. Thus, no nest-box contained the original nest. Pairs collected more nest material and for longer periods in the control nests, although the experimental pairs still added some nest material after the manipulation. Nest-boxes were filmed for 1 to 3 h during incubation and on days 4 and 11 of the nestlings' life. Female attendance during incubation was negatively affected by nest mass, this parameter explaining almost half the variation in attendance, which suggests an effect of nest mass on incubation efficiency. The female provisioning effort on day 4 was significantly higher in the experimental nests, whereas other parental care variables were affected by date, weather conditions and brood size or brood mass. Mean tarsus length at fledging was significantly longer in the exper- imental nests. The savings by females in nest construction effort were expressed in higher provisioning at early nestling stages and in improved nestling growth.