Long-lasting mobbing of the pied flycatcher increases the risk of nest predation

Mobbing behavior may provide real benefits because mobbing preyindividuals often cause a predator to leave the vicinity. However,mobbing calls of prey can attract acoustically oriented predatorsthus increasing nest predation. Therefore, a real value of mobbingas a type of adaptive behavior may depend on its duration. Inthis experimental study, we tested whether mobbing durationby the pied flycatcher Ficedula hypoleuca, a small hole-nestingpasserine, increases the risk of nest predation. From the topof one nest-box within each of 78 experimental plots, we playedback long calls of pied flycatchers, whereas recordings of shortmobbing calls were played back from the top of another nearbynest-box. The nest-boxes were arranged in pairs, and each ofthem contained a quail Coturnix coturnix egg. Long-call nest-boxeswere depredated by martens Martes martes significantly moreoften than short-call nest-boxes. This predator usually huntsby night and may eavesdrop on the calls of their mobbing preywhile resting nearby during the day. The results of the presentstudy show that long-lasting conspicuous mobbing calls may carrya significant cost for the breeding birds.     

Experimentally reduced male attractiveness increases parental care in the pied flycatcher Ficedula hypoleuca

This study reports effects of experimental manipulations ofreproductive effort and the size of the male's white foreheadpatch (a secondary sexual trait), on provisioning rates, reproductivesuccess, and parental breeding dispersal distance in the piedflycatcher, Ficedula hypoleuca. Parents caring for enlargedbroods resulting from manipulated clutches provisioned nestsat higher rates than parents with reduced broods. Males with a reduced forehead patch fed their nestlings more in relationto males with an unmanipulated forehead patch, and their youngfledging with a longer tarsi. This suggests that males witha reduced attractiveness may perceive their own attractivenessand they devote more time available for parental effort given their poorer prospects in male contest competition and/or femaleattraction for extra-pair copulations. However, their femalesdid not alter their provisioning effort and this runs counterto both the differential allocation and the partner-compensationhypotheses. An artificial decrease in a male secondary sexualtrait led to a wider breeding dispersal distance between successiveyears.     

Descriptive and experimental evidence for timing-mediated polygyny risk in a pied flycatcher Ficedula hypoleuca population

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.     

Adaptive coloration in pied flycatchers (Ficedula hypoleuca)—The devil is in the detail

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.     

Individual color variation and male quality in pied flycatchers (Ficedula hypoleuca): a role of ultraviolet reflectance

Bright coloration of males in many animal species has inspiredresearchers for more than a century. In this field study, weinvestigated whether color variation between individuals isrelated to individual quality in pied flycatcher (Ficedulahypoleuca) males in terms of arrival time at the breeding sites.In addition to traditional visual color scoring, plumage color was measured using spectroradiometric measurements between 320and 700 nm. This range includes the near-ultraviolet wavebandfrom 320 to 400 nm. Males that arrived earlier at breedingsites had higher proportional UV reflectance in the crown andmantle. The proportional UV reflectance in the crown and mantlewas not related to traditionally scored general brownness inmales. However, adult males had a higher proportion of ultravioletin the plumage than yearling males or females. These resultssuggest that in pied flycatcher males, the UV reflectance ofplumage may be positively correlated with individual quality.     

Development of the auditory sensitivity and formation of the acoustically guided defensive behavior in nestlings of the pied flycatcher Ficedula hypoleuca

Age dynamics of generation of the evoked potentials (EP) in the field L of caudal nidopallium (the higher integrative center of the avian auditory system) and development of the auditory-guided defensive behavior were studied in control and visually deprived pied flycatcher Ficedula hypoleuca nestlings. It was shown that the rhythmically organized monofrequency signals with sound frequency 3.5 kHz and higher produced the defensive behavior as the auditory sensitivity to these frequencies matured. After 9 days, the species-specific alarm signal produced more effectively the defensive behavior than the tonal signals. The rhythmically organized sound with filling frequency 0.5 kHz, occupying the less low-frequency diapason than the feeding signal, produced the effect opposite to the alarm signal to increase the nestling mobility. At the initial stage of the defensive behavior development the auditory threshold fell markedly in the frequency diapason corresponding to the frequency diapason of the alarm signal (5–6 kHz), which seemed to facilitate involvement of this diapason signals in the defensive integration. The auditory EP generation thresholds in the whole studied diapason were lower in the visually deprived nestlings than in the normally developing one; however, the ability of the acoustic signals to suppress alimentary reactions fell significantly.     

Handicapped males and extrapair paternity in pied flycatchers: a study using microsatellite markers

We report an attempt to induce extrapair copulations and fertilizations in a species with a low intensity of sperm competition, the pied flycatcher Ficedula hypoleuca. Shortly after pair formation males were made less attractive to females by removing certain wing and tail feathers. Earlier research has shown that this manipulation reduces a male's pairing success. The idea was to test whether females mated to such males (N= 9) were more likely to obtain extrapair fertilizations than females mated to unmanipulated controls (N= 9). Paternity testing was carried out on all 98 young in the 18 broods, using a set of six microsatellite markers isolated from the species. Extrapair fertilizations were revealed in only three (17%) broods; two broods of handicapped males and one of a control male. A total of seven (7%) offspring were not genetically related to their putative father, a level which agrees well with results of other studies of this and other populations. We conclude that there was no evidence to suggest that the fertilization pattern was altered by the experimental manipulation. One reason for the lack of response could be that female mate choice in this species is based on male phenotypic, and not genotypic, quality.     

Properties of food loads delivered to nestlings in the pied flycatcher: effects of clutch size manipulation, year, and sex

Experimental manipulation of the number of altricial offspring is supposed to modify parental expenditure in birds. In addition to the observed increase in parental feeding rate, it is also possible that the choice of prey or the size of load may change with the changing demand for food. Sexual differences in the provisioning response are also expected, on the basis of earlier studies. We examined the effect of brood size manipulation on choice of prey brought to nestlings and load size in the pied flycatcher. The composition and size of loads differed between years, possibly depending on varying availability of different prey types. Males responded to brood size enlargement by gathering heavier loads, whereas females showed no response. The alteration of load size in males was not explained by a larger number of prey items or mean prey size, but was a combination of these components. It is likely that males also increased their work rate in response to increased food demand at the nest. The absence of response in females might be because they are unable to increase work rate any further, or because food delivery rate in females can not be optimized by changing load properties. Received: 18 December 1997 / Accepted: 1 March 1998     

Do female pied flycatchers seek extrapair copulations with familiar males? A test of the incomplete knowledge hypothesis

In birds with biparental care, great variation exists in thefrequency of extrapair paternity. Several hypotheses have beenproposed to account for this variation. We tested the incompleteknowledge hypothesis, which states that females are constrainedin their knowledge of male quality and that this influencestheir willingness to engage in extrapair copulations (EPC).By selective removal and release of female pied flycatchersFicedula hypoleuca, we created a situation where females finallysettled with a social mate close to the site where a formersocial mate was breeding. According to the incomplete knowledgehypothesis, this would lower the threshold for females to seekextrapair copulations in cases where their former social matewas of higher quality than the one finally chosen. The hypothesiswas not supported because manipulation of female settlementdid not increase frequency of extrapair paternity, not evenin cases where the female nested close to the previous mateand the current mate apparently was of lower quality becausehe was younger and more dull colored. However, we found that when extrapair paternity did occur, the cuckolder tended tobe a familiar male (i.e., the female's initial social mate).     

Nestling testosterone controls begging behaviour in the pied flycatcher, Ficedula hypoleuca

Begging signals and endogenous testosterone (T) levels of young birds have been shown to be positively correlated. If T is causally involved in controlling the level of begging effort, an endocrine control mechanism could explain the evolution of begging as a costly signal reflecting need. We tested experimentally whether elevated circulating T levels enhanced begging behaviour in nestling pied flycatchers, Ficedula hypoleuca. A pilot study confirmed that nestling T levels could be elevated within a natural physiological range using an oral dose of T. After T-dosing, nestling begging behaviour was measured as: i) the duration of begging displays and ii) the maximum height of begging stretches. Our results show that nestling T levels were elevated at 90 min post dosing and that at this time point both measures of begging behaviour were performed more intensely by T-dosed nestlings than controls. Nestling begging displays in response to dosing varied between individuals, which in part was explained either by the date in the breeding season or nestling mass. The results of this study confirm the causal nature of T in controlling nestling begging signals and suggest that it may be part of the mechanism that controls begging behaviour in nestling birds.     

Behavioral responses to conspecific mobbing calls are predator‐specific in great tits (Parus major)

When facing a predator, animals need to perform an appropriate antipredator behavior such as escaping or mobbing to prevent predation. Many bird species exhibit distinct mobbing behaviors and vocalizations once a predator has been detected. In some species, mobbing calls transmit information about predator type, size, and threat, which can be assessed by conspecifics. We recently found that great tits (Parus major) produce longer D calls with more elements and longer intervals between elements when confronted with a sparrowhawk, a high‐threat predator, in comparison to calls produced in front of a less‐threatening tawny owl. In the present study, we conducted a playback experiment to investigate if these differences in mobbing calls elicit different behavioral responses in adult great tits. We found tits to have a longer latency time and to keep a greater distance to the speaker when sparrowhawk mobbing calls were broadcast. This suggests that tits are capable of decoding information about predator threat in conspecific mobbing calls. We further found a tendency for males to approach faster and closer than females, which indicates that males are willing to take higher risks in a mobbing context than females.     

Black-and-white plumage in male pied flycatchers (Ficedula hypoleuca) reduces the risk of predation from sparrowhawks (Accipiter nisus) during the breeding season

Bright colors in birds might signal that they are undesirableas prey (aposematic), an idea that has been difficult to test.When stuffed pied flycatchers Ficedula hypoleuca are exposedto migrating sparrowhawks Accipiter nisus in spring or in autumn,the hawks attack cryptic females more often than bright males.To achieve better statistical control and to assess whethermale plumage also reduces predation risk in the breeding seasonand in the nesting habitat of the pied flycatcher, I placedpairs of male and female flycatcher mounts in similar positionsnear 22 nests of sparrowhawks. The hawks attacked mainly femalemounts, verifying that the preference is real. The sparrowhawkscaught at least 19 live pied flycatchers; 12 young, 5 adultmales, 1 adult female, and 1 female or young. Hawks that caughtan adult male seemed to prefer attacking female mounts. I discussthree interpretations of these results, suggesting that black-and-whitemale flycatchers may benefit from being a novel and aberrantprey, at least early in the breeding season.     

Evidence for the signaling function of egg color in the pied flycatcher Ficedula hypoleuca

A recent hypothesis proposes that the bright colors, especiallyblue and green, of many avian eggs may function as signals offemale or offspring phenotypic quality or condition to malesin species with biparental care, inducing them to allocate moreeffort to their offspring. The pigment determining blue andgreen egg colors is an antioxidant whose availability for eggshellcoloring may be limited. To test the signaling function on aspecies with blue eggs, the pied flycatcher Ficedula hypoleuca,we measured egg color with a spectrophotometer on the day oflaying and obtained two principal components from their reflectancespectra that together explained 99% of variation and representedshell lightness, and hue and saturation, respectively. We alsomeasured female immunocompetence during the nestling periodthrough the response to phytohemagglutinin as a measure of cell-mediatedimmunity and the response to a tetanus vaccination as a measureof humoral immunity. The total amount of immunoglobulins inblood of females and of nestlings before fledging was also estimated.Mean within-clutch egg darkness was positively associated withboth measures of female immunocompetence, while better femalecondition was associated with colors tending away from intermediateand toward short wavelengths. Ageing female laid lighter eggs.The mean within-brood level of nestling IgY was also associatedwith mean within-clutch egg colors tending away from intermediateand toward short wavelengths. Mean egg darkness decreased linearlyduring the laying sequence, suggesting pigment limitation. Maleswere observed frequently visiting nests during the laying period,allowing them to observe eggs before the start of incubation.These results support the signaling hypothesis for explainingbright colors of avian eggs.     

Decline of a montane Mediterranean pied flycatcher Ficedula hypoleuca population in relation to climate

Certain populations of long‐distance migratory birds are suffering declines, which may be attributed to effects of climate change. In this article, we have analysed a long‐term (1991–2015) data set on a pied flycatcher Ficedula hypoleuca population breeding in nest‐boxes in a Mediterranean montane oak forest, exploring the trends in population size due to changes in nestling recruitment, female survival and female immigration. We have related these changes in population parameters to local climate, winter NAO index and to breeding density. During the last 25 yr the population has declined by half, mainly in association with a decrease in nestling mass and structural size which had repercussions on the probability of nestling recruitment to the population. Lower local nestling recruitment in certain years was linked to lower female immigration rate in the same years. On the other hand, the local survival of females remained stable throughout the study period. Laying date and breeding success were negatively affected by local temperatures while breeding, recruitment rate likewise by minimum temperature prior to breeding in April. As minimum April temperatures have increased across the study period, this may have affected recruitment and immigration rates negatively. On the other hand, tarsus length and body mass of nestlings were positively associated with winter NAO index, pointing to more global climatic links. Moreover, there was also a negative temporal trend in body mass of adults, implying increasingly difficult conditions for breeding. Declining recruit production in the study area could be attributed to a mismatch between the timing of arrival and breeding in the population, and the peak of food availability in this area.     

Climatic effects on timing of spring migration and breeding in a long-distance migrant, the pied flycatcher Ficedula hypoleuca

Climate change has advanced the breeding dates of many bird species, but for few species we know whether this advancement is sufficient to track the advancement of the underlying levels of the food chain. For the long-distance migratory pied flycatcher Ficedula hypoleuca the advancement in breeding time has been insufficient to maintain the synchrony with their main food sources. The timing of arrival in the breeding areas from their African wintering grounds is likely to constrain the advancement of breeding date. We hypothesise that this is because in Africa they cannot predict the advancement of spring in their breeding habitat. However, long-distance migrants may advance their arrival time by migrating faster when circumstances en route are favourable. In this study we show that both arrival and breeding date depend on temperatures at their main North African staging grounds, as well as on temperature at the breeding grounds. Male arrival and average laying date were not correlated, but the positive effect of temperature in North Africa on breeding dates suggests that breeding date is indeed constrained by arrival of females. Long-distance migrants thus are able to adjust arrival and hence breeding by faster spring migration, but the degree of adjustment is probably limited as timing schedules in spring are tight. Furthermore, as climate change is affecting temperatures differently along the migratory flyway and the breeding areas, it is unlikely that arrival dates are advanced at the same rate as the timing of breeding should advance, given the advancement of the underlying levels of the food chain.     

Polygyny in the pied flycatcher,Ficedula hypoleuca: a test of the deception hypothesis

The deception hypothesis has been proposed as an explanation for polygyny in the pied flycatcher. According to this hypothesis, already-mated males hide their mating status with polyterritorial behaviour and thereby increase their chances of obtaining a second mate. In a study area at Oslo, Norway, secondary females raised 84% as many fledglings as did concurrent monogamous and primary females. The unmated males sang most of the time near their nest site, whereas the already-mated males frequently disrupted singing for longer periods in their secondary territories to visit their primary nest; such visits to the primary nest occurred both before and after the time of their second mating. The behaviour of the males suggests that deception of females is not an evolutionarily stable strategy, as an observant female would soon discern the male's status. Another difficulty with the deception hypothesis is that secondary females laid larger clutches than primary females. The number of young fledged from secondary nests was not dependent on the distance to the primary nest. The deception hypothesis was not supported by the data, and the reduced reproductive success of secondary females may be explained by the cost of searching for a mate.