Correlation between egg size and clutch size in the Pied Flycatcher Ficedula hypoleuca in cold and warm summers

The correlation between egg size and clutch size was studied in the Pied Flycatcher Ficedula hypoleuca in Finnish Lapland from 1975 to 1994. The hypothesis tested was that the tradeoff phenomenon is masked by warm weather. When data for 20 years were pooled, there was no correlation. However, when years were analysed separately, a negative correlation was observed for five of the years. Over 12 years, the correlation coefficient between egg size and clutch size was not significant, and for 3 years it was significantly positive. The size and magnitude of the correlation were related to temperature during the pre-laying and egg-laying periods. Of the annual variation, 34% was explained by the mean temperature in early summer. It is suggested that trade-off between egg size and clutch size may not be apparent in central (southern) parts of the distribution area of the species where climate is warmer. In the future, global warming may alter reproductive strategies of birds. Long-term data may thus be useful in the analysis of such effects and in testing of theoretical questions in ecology.     

Effects of long-acting testosterone administration on testes in free living pied flycatchers Ficedula hypoleuca

Male pied flycatchers Ficedula hypoleuca given injections of long-acting testosterone, when in full breeding condition, maintained spermatogenesis. In males given vehicle only and in untreated males the testes collapsed at the time of hatching of the brood. The number and secretory activity of the Leydig cells, as indicated by enzymic activities, were markedly lowered after testosterone administration. The results make it unlikely that testosterone is responsible for testicular regression in this species, but it may affect the mechanism regulating the number and activity of the Leydig cells.     

Mate Sampling and Assessment Procedures in Female Pied Flycatchers (Ficedula hypoleuca)

Models of mate sampling strategies predict that choosiness should decrease throughout the breeding season due to increasing costs of delaying mating. Therefore, individuals who start searching mates relatively late, should spend less time on sampling, and sample fewer candidates compared to early individuals. We observed mate searching behavior of female pied flycatchers (Ficedula hypoleuca) by radio-tracking in study areas with 6–12 unpaired males. Contrary to the prediction, the observed numbers of males sampled by the searching females increased with time, i.e. late arriving females visited more males than early arrivers. However, this seems to be due to more active sampling of males in short time by late-arriving females. The observed sampling pattern suggests some kind of comparison tactic, which seems, however, to be very variable among individual females. Mate-assessing females were characterized by a remarkably cryptic behavior, which may be 1) a way of gaining honest information about the male mating status or male/territory quality, or 2) a way of avoiding courtship costs.     

Maternal clutch reduction in the pied flycatcher Ficedula hypoleuca: an undescribed clutch size adjustment mechanism

During eight years of study of a population of pied flycatchers Ficedula hypoleuca breeding in central Spain, we observed throughout incubation 22 cases of intact eggs being placed on the nest rim and clearly outside of the nest bowl and remaining there, with consequent embryo mortality. We assume that the removal of eggs had been performed by the incubating female. There is no evidence that pied flycatcher parents can detect embryo mortality and eject unhatchable eggs. Maternal egg ejection constitutes a direct mechanism of clutch size adjustment during incubation and before hatching of any young of the brood. This adjustment of clutch size happened when the mean and minimum temperature of the day before ejection were especially low. Also, clutch-reducing females were significantly more prone to show moult-breeding overlap than other females, suggesting a lower disposition to invest in reproduction. Clutch-reducing females were also frequently older than four years and had indications of a lower immunocompetence than females not ejecting eggs. Our results support the hypothesis that adverse conditions can elicit parental family size adjustment at every stage of reproduction without the need to depend on mechanisms of sibling competition.     

Nestling growth and mortality of Pied Flycatchers Ficedula hypoleuca in relation to weather and breeding effort

The growth pattern and mortality of young Pied Flycatchers Ficedula hypoleuca were studied to focus on the mechanisms and constraints behind the widely studied optimization of clutch size. The clutch sizes were modified, and the growth and survival of chicks from different clutch sizes were monitored along with the prevailing weather during the nestling period to detect the effect of weather on reproductive success. The weather conditions during the feeding period of the nestlings varied within a season as well as between breeding seasons. The prevailing weather markedly affected both the growth rate and the survival of chicks, yet, the effects of weather on growth were not greater in enlarged clutches. The impact of adverse weather was more pronounced in the later phases of nesting, when the food demand of a brood was highest. Brood reduction and total nest losses were more likely during extensive rainfall during the nestling phase and also in the enlarged clutches. Thus, weather is a very important determinant of reproductive success in this species. Weather conditions during the breeding season are unpredictable, however, and therefore brood reduction through sibling competition is a mechanism whereby brood size can be adjusted to the level the parents can rear under the prevailing conditions.     

Patterns of breeding behaviour and plasma levels of hormones in a free-living population of Pied flycatchers, Ficedula hypoleuca

Plasma levels of luteinizing hormone (LH) and steroid hormones were measured by radioimmunoassay in a series of plasma samples collected from a free-living population of Pied flycatchers, Ficedula hypoleuca. Both males and females have similar circulating levels of LH and testosterone when they arrive in the breeding area. However, as territories and pair bonds are established, and as courtship and nest-building ensue, the plasma levels of testosterone in males become elevated over those of females, although a lesser peak also occurs in females. As the egg laying phase and incubation begin, plasma levels of LH and testosterone decline sharply in both sexes. On the other hand polyterritorial males, that establish secondary territories, maintain higher levels of LH and testosterone than males remaining in the home-territory, at least until they return to the home-territory, usually at the middle part of the incubation period. By this time polyterritorial males also have basal levels of these hormones. Curiously, females on home-territories show a small, but significant, rise of circulating LH and testosterone at the time the males return.
In males, plasma levels of corticosterone are elevated from the time of arrival on the breeding grounds until the end of the nest-building period. Thereafter corticosterone levels decline in males staying on the home-territory but remain elevated in polyterritorial males until the latter return to the home-territory. In contrast, circulating levels of corticosterone in females are only elevated for brief period during egg laying.     

Behavioural responses to ectoparasites in pied flycatchers Ficedula hypoleuca: an experimental study

Nests of cavity‐nesting birds usually harbor some species of haematophagous ectoparasites that feed on the incubating adults and nestlings. Given the negative impact of ectoparasites on nestlings there will be selection on hosts to reduce parasite infestations through behavioural means. We have experimentally reduced the abundance of all ectoparasites in nests of pied flycatchers Ficedula hypoleuca to explore both whether there are changes in the frequency and duration of putative anti‐parasite behaviours by tending adults, as well as whether such anti‐parasite behaviours are able to compensate for the deleterious effects that parasites may have on nestlings. Heat treatment of nests substantially decreased the density of ectoparasites, and thereby positively affected nestling growth. The frequency and intensity of female grooming and nest sanitation behaviours during the incubation and nestling periods decreased as a consequence of the experimental reduction of ectoparasite infestation. Although nestlings begged more intensely in infested nests, the experiment had no significant effect on parental provisioning effort. Reduction of parasites resulted in larger nestlings shortly before fledging and increased fledging success. This study shows a clear effect of a complete natural nest ectoparasite fauna on parental behaviour at the nest and nestling growth in a cavity‐nesting bird. Although ectoparasites induce anti‐parasite behaviours in females, these behaviours are not able to fully remove parasite's deleterious effects on nestling growth and survival.     

Distribution and reproductive success of the Pied Flycatcher Ficedula hypoleuca in relation to forest patch size and vegetation characteristics; the effect of scale

We studied habitat choice, distribution and reproductive success in the Pied Flycatcher Ficedula hypoleuca in relation to the spatial structure of a fragmented forest area. Two hierarchical levels of forest habitat structure were used: (1) forest patch size (macrohabitat) and (2) vegetative structure within forest patches (microhabitat). In spring, both males and females settled preferentially in large and medium-sized forest stands (>1 ha) where breeding density was also higher than in small stands (<1 ha). Stands <5 ha were occupied later, and the proportion of unpaired males was higher there than in large stands (>5 ha). The known age distributions of breeding birds and breeding success were independent of forest patch size. Nest predation rate was not associated with stand size or nest distance from the forest edge. The preference of breeding birds for large forest patches was presumably related to the higher amount of resources (nest sites, mates, food) a large patch can offer for reproduction compared with a small patch. At the microhabitat level, territories of old males were characterized by relatively more deciduous trees that contained more invertebrate food than coniferous tree-dominated territories of yearling males. The reproductive success of old males, as measured by the number of fledged young per male, was higher than that of yearling males. This suggests that the larger body size and blacker plumage of old males possibly contributed to male dominance and that old males excluded younger males from preferred habitats through territorial behaviour. Our results suggest that habitat choice of the Pied Flycatcher was affected by both micro- and macrohabitat. On the basis of settlement pattern, density and reproductive success, the distribution of Pied Flycatchers across forest stands of different sizes followed the ideal-free distribution model, whereas at the microhabitat level, age-related unequal distribution of males followed the ideal-despotic model of Fretwell and Lucas.     

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 colorwas 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.     

Environmental and genetic variation in the haematocrit of fledgling pied flycatchers Ficedula hypoleuca

We exposed females of a highly placentotrophic viviparous scincid lizard (Pseudemoia pagenstecheri) to various environmental factors during pregnancy, and quantified the effects of these treatments on their offspring. The clear result was that the phenotypes of neonatal lizards can be substantially modified by the environment that their mother experiences during gestation. Restricting prey availability to the females reduced the size of their offspring. Limiting the females' basking opportunities delayed their seasonal timing of parturition, and modified body proportions (tail length relative to snout-vent length) of the neonates. More surprisingly, female lizards that were regularly exposed to the scent of sympatric lizard-eating snakes gave birth to offspring that were heavier, had unusually long tails relative to body length, and were highly sensitive to the odour of those snakes (as measured by tongue-flick responses). The neonates' antipredator responses were also modified by the experimental treatment to which their mother was exposed. The modifications in body mass, tail length and response to snake scent plausibly reduce the offspring's vulnerability to predatory snakes, and hence may constitute adaptive maternal manipulations of the neonatal phenotype. Received: 6 July 1998 / Accepted: 5 December 1998     

Predation by sparrowhawks Accipiter nisus on male and female pied flycatchers Ficedula hypoleuca in relation to their breeding behaviour and foraging

Bright plumage, song display, and aggressive resource defence in males may cause higher predation on males than on females during the breeding season. However, in birds, higher predation on females is sometimes observed. Parental investment may be high in females (egg-laying, incubation and feeding of offspring), which might lead to a high risk of predation. We studied predation by sparrowhawks Accipiter nisus in relation to behaviour in pied flycatchers Ficedula hypoleuca where breeding males are more conspicuous than females in plumage and behaviour. Male pied flycatchers generally occupied more exposed perches than females. Females were more mobile and foraged more than males, especially prior to and during incubation. During the incubation and nestling stages, when predation on the sexes could be directly compared, sparrowhawks took about the same number of male and female pied flycatchers. During incubation, however, females spent about 77% of the day in the nest and were 4.7 times more vulnerable than males per unit of time available (i.e. outside the nest). A comparison with the chaffinch Fringilla coelebs , where hawks took more females than males, indicates that timing of breeding, foraging behaviour and parental roles of males and females affect predation risk.     

Enzyme histochemical studies in testes and seminal vesicles of the Pied flycatcher (Ficedula hypoleuca) during the breeding season

Enzymatic conditions in testes and seminal vesicles of the Pied flycatcher during the breeding season were studied by histochemical techniques. The following enzyme activities were studied: glucose-6-phosphatase dehydrogenase (G-6-PDH), NADPH₂-diaphorase, glutamic dehydrogenase (GDH), NADH₂-diaphorase, adenosine triphosphatase (ATPase) and acid phosphatase. Phospholipids and PAS-positive substances were also studied. Towards the end of the nest-building period, a time when the males showed maximal sexual activities, some enzyme activity changes were observed. There was an increase of the GDH activity in the sperm bundles, an increase of the ATPase activity in the interstitium and tunica propria of the tubuli seminiferi. The phospholipid content increased in all spermatogenetic cells and the PAS-positive material increased in the Sertoli cells and spermatozoa. The enzymic activities indicated a steroid production in the Leydig cells.
The epithelium of the seminal vesicles did not show any variations in enzyme activities as long as the Leydig cells persisted. Most enzymes showed a very high activity during this time, indicating a secretory function of the epithelium. The amount of phospholipids increased at the end of the nest-building period. Parallel with the degeneration of the epithelium the enzymic activities, as well as the phospholipid content, were lowered markedly.     

Growth and mortality of nestling great tits (Parus major) and pied flycatchers (Ficedula hypoleuca) in a heavy metal pollution gradient

We studied nestling growth, growth abnormalities, mortality and breeding success of two hole-nesting passerines, the great tit (Parus major) and the pied fly-catcher (Ficedula hypoleuca), at 14 study sites around a copper smelter complex in Harjavalta, south-west Finland in 1991–1993. The main pollutants in the area are heavy metals and sulphuric oxides. Exposure of birds to heavy metals was shown by measuring their faecal concentrations. Copper, nickel and lead contents of nestling faeces were high near the factory and decreased with distance away from the pollution source. F. hypoleuca nestlings suffered high mortality very close to the factory complex, but did relatively well at all other sites. Breeding success of P. major was below background levels up to 3–4 km from the pollution source and nestlings grew poorly close to the factory. Growth abnormalities of legs and wings in F. hypoleuca nestlings were significantly more common near the factory than farther away. In contrast, F. hypoleuca nestlings grew equally well at all distances. The poor breeding success of F. hypoleuca close to the factory complex is probably related to the high amount of heavy metals in its diet, and low availability of calcium-rich food items may enhance this effect. We suggest that the poor breeding success of P. major is related to habitat changes that have taken place around the factory. The different responses of these two bird species are probably due to their different diet. Our results show convincingly that species-specific differences in response should be carefully considered when planning schemes for air pollution monitoring.     

Divergent character displacement in the song of two allospecies: the Pied Flycatcher Ficedula hypoleuca, and the Collared Flycatcher Ficedula albicollis

Songs of the three flycatcher forms, Ficedula hypoleuca hypoleuca, Ficedula albicollis albicollis and Ficedula albicollis semitorquata , were distinguishable from one another using a multivariate analysis. F. albicollis semitorquata , which has an intermediate song character, is considered to be a primary allopatric form while the nominate subspecies of albicollis is assumed to be dispersed into sympatry with F. h. hypoleuca . The differences in song between the nominate albicollis and F. hypoleuca are accentuated in their zone of sympatry and are mostly due to divergence in the albicollis song. The song is primarily an epigamic vocalization in the two species and conforms to the pattern characteristic of divergent character displacement associated with reproductive behaviour. Two alarm calls are quite similar in the two species and are the main vocalizations associated with territorial combat. Their interspecific territorial behaviour has apparently not been affected in areas of sympatry.     

Structural and functional aspects of variation of the breeding plumage ornamentation in the male Pied Flycatcher Ficedula hypoleuca (Aves: passeriformes)

Inter- and intraindividual variation of the breeding plumage ornamentation in male Pied Flycatcher Ficedula hypoleuca from the Moscow province population (55 degrees 44' N 36 degrees 51' E) was studied using long-term data (1996-2005). Eumelanin prevailing colour type (CT) scored according to the seven-step Dorst scale (Dorst, 1936), age, and reproductive status during the current breeding season were analyzed as potential sources of variation of the white ornamentation on the forehead, the wings, and the tail in males. Male CT strongly varied, from almost black to cryptic brownish (2-7, average 4.4, N = 1403). On average CT became I1degree darker with age and tended to be stable after the age of two years. Initial and final CT strongly correlated (r = 0.76). Wing ornamentation and forehead patch (FP) were associated with both initial and final CT, whereas correlation between tail ornamentation and CT was observed only in aged males. Ornamentation was most variable in males with cryptic (pale) CT. Such males had, on average, less intense forehead and wing ornamentation than conspicuous males, compensating this difference by having more intense ornamentation next year. These data suggest the weakening of morphogenetic relations between prevailing colour and ornamentation in males with cryptic phenotype. The return rate (rate of conspicuous males returning to their previous nesting territory) was not related to the success of their previous breeding attempts. Among pale males, the return rate was higher in breeders than in non-breeders. The relation between FP shape and reproductive status was pronounced only in pale males. Among pale males, previous breeders and non-breeders displayed no difference in the degree of prevailing colour darkening next year. In conspicuous males, previous breeding led to the weakening of dorsal eumelanin pigmentation of the new plumage, which suggests the existence of tradeoffs between reproductive efforts and the development of rich eumelanin coloration. Among pale males, previous breeders displayed a higher rate of FP widening next year than previous non-breeders. Diffrerent patterns of relations between male ornamentation and reproductive experience suggest that the females can use plural clues to evaluate the quality of their potential mates.     

Egg shell quality,clutch size and hatching success of the great tit (Parus major) and the pied flycatcher (Ficedula hypoleuca) in an air pollution gradient

Egg shell thickness, egg volume, clutch size and hatching success of Parus major and Ficedula hypoleuca were studied at 14 study sites around a copper smelter complex in Harjavalta, south-west Finland, in 1991–1993. In 1991–1992 unhatched eggs were collected to measure shell quality. F. hypoleuca was more susceptible to pollutants than P. major, the response of which was weaker in all aspects studied. Egg shells of F. hypoleuca were about 17% thinner and eggs were about 8% smaller in volume near the factory than at a distance of 10 km. The clutch size of F. hypoleuca was significantly smaller and hatching success markedly lower at a study site next to the factory complex than at all other sites. In P. major, variation in shell thickness and egg volume was not significantly related to the distance from the pollution source. Clutch size and hatching success of P. major did not significantly differ among study sites, although the trend in hatching success was in the same direction as in F. hypoleuca. Clutches of both species contained less shell material and both species had more nests without eggs near the factory than further away. The surface structure of the eggshells was studied by scanning electron microscope. Especially in F. hypoleuca, the egg shell surface was more rough and porous near the factory. The roles of Ca and heavy metals in shell thinning are discussed.     

Use of materials in nest construction by Pied Flycatchers Ficedula hypoleuca reflects localized habitat and geographical location

Capsule: Pied Flycatchers Ficedula hypoleuca use different materials to construct their nests according to local availability and geographical location.

Aims: This study tested the hypotheses that Pied Flycatcher nests were constructed from the leaves of different tree species in proportion to their occurrence within the breeding territories and that nest composition varied between geographical locations.

Methods: In Lancashire, Pied Flycatcher nests were collected from nestboxes built in locations dominated by different tree species and were deconstructed to determine which materials were used.

Results: The leaves found in nests generally reflected their availability within the locality of the territories rather than showing evidence of selective use of some leaf species. However, the use of moss was dominated by the use of one species in all but two nests. Nests from Lancashire were significantly different in composition when compared with published data for nests from north Wales and central Spain.

Conclusion: Pied Flycatchers exhibit plasticity in nest construction behaviour because they were opportunistic in their choice of most leaves as nesting materials, although they may be selective in their choice of moss.