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Climate change affects timing of reproduction in many bird species, but few studies have investigated its influence on annual reproductive output. Here, we assess changes in the annual production of young by female breeders in 201 populations of 104 bird species (N = 745,962 clutches) covering all continents between 1970 and 2019. Overall, average offspring production has declined in recent decades, but considerable differences were found among species and populations. A total of 56.7% of populations showed a declining trend in offspring production (significant in 17.4%), whereas 43.3% exhibited an increase (significant in 10.4%). The results show that climatic changes affect offspring production through compounded effects on ecological and life history traits of species. Migratory and larger-bodied species experienced reduced offspring production with increasing temperatures during the chick-rearing period, whereas smaller-bodied, sedentary species tended to produce more offspring. Likewise, multi-brooded species showed increased breeding success with increasing temperatures, whereas rising temperatures were unrelated to reproductive success in single-brooded species. Our study suggests that rapid declines in size of bird populations reported by many studies from different parts of the world are driven only to a small degree by changes in the production of young., This meta-analysis was financed by the grant of the Polish National Science Centre (Narodowe Centrum Nauki) (no. 2017/27/B/NZ8/00465) awarded to Lucyna Hałupka.

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

Landscape conversions induced by human activities can affect dispersal patterns of various bird species and, as a result, affect genetic structure of their populations. Genetic differentiation of bird populations may be enhanced by habitat variation, especially in urban-non-urban systems. The majority of population genetic studies focus on single species, which inflicts limitations for direct comparisons of genetic responses of avian populations to urbanization. Here, we used a set of microsatellite markers to examine genetic diversity, gene flow and population structure in two common parid species, great tits Parus major and blue tits Cyanistes caeruleus occupying three sites in habitats with contrasting urbanization level in central Poland. We found low but significant divergence of urban park population with both suburban and non-urban forest great tit populations, while no differentiation was found between suburban forest and non-urban forest populations. In contrast, no evidence for genetic differentiation was found between blue tit populations from the urban park, suburban forest and non-urban forest sites. We conclude that great tits and blue tits respond to urbanization-related changes in a different way, which may be a result of different rates of migration and/or dispersal, likely higher in blue tits. Some impact may be also induced by interspecific competition. We suggest that changing the focus of urban genetic research from single to multiple species may provide novel insights into how natural populations respond to the processes of urbanization.

Nests of secondary-cavity breeding birds are structures constructed to lay and incubate eggs and raise nestlings to fledging. These nests are characterized by relatively stable conditions, which makes them a suitable habitat for various microorganisms. It has been suggested that bacteria inhabiting nests and nestling skin may affect nestlings both positively and negatively. In this study, nests of Great Tits Parus major were replaced with artificial nests on the fifth day of the nestling phase in two different study sites to create two categories of nests: (i) natural and (ii) artificial with a reduced bacterial load. Four days later, bacterial samples were collected from the nestling skin and from the edge of the nest to assess skin and nest bacterial loads, expressed as colony forming units (CFUs). It was predicted that (i) the previous season occupancy of nest boxes would influence bacterial loads, (ii) the experimental treatment would reduce nest and skin bacterial loads, and (iii) that nest and skin bacterial loads would affect the condition of the nestlings, assessed as the hemoglobin concentration in blood when nestlings were 14 days old. Occupancy in the previous season did not affect the bacterial load. The skin bacterial load was significantly lower in artificial nests, although the nest bacterial load did not differ between natural and artificial nests. Nestlings from artificial nests had higher hemoglobin, and hemoglobin concentration was negatively associated with the nest but not the skin bacterial load. Our results suggest that the bacterial load in nests may negatively affect the physiological condition of avian hosts.

Among environmental factors affecting life - history traits of birds breeding in nest boxes, an influence of microbial communities is relatively poorly understood. In this study, nest boxes used for breeding by great tit (Parus major) and blue tit (Cyanistes caeruleus) were sampled before the start of the breeding season to assess the bacterial loads of the nest box. Samples from the entrance hole and the interior of the nest box were taken at two different study sites: an urban parkland and a natural forest. Nest boxes were sampled to check if their bacterial loads differed between habitats. The second objective of this study was to check whether the occupancy of the nest boxes during the previous season would influence the bacterial load of the nest box. To verify this prediction, two categories of nest boxes were sampled at both study sites: nest boxes occupied by any of the two tit species in the previous season for breeding and nest boxes that had remained empty that year. The bacterial load of the nest box was significantly higher in the forest study area in both the occupied and unoccupied nest boxes. The nest boxes used for breeding in the previous season had significantly higher bacterial loads, but only in the forest area. Our results suggest that the bacterial load of the nest box can vary between habitats and may be positively related to the presence of the nests in the previous breeding season.

It has been suggested that extreme weather events may be treated as natural experiments that may unravel the mechanisms by which birds adjust their phenology and breeding parameters to environmental variability. In 2017, a sudden and heavy drop of temperatures for several days affected many European bird populations. This event occurred during the laying–early incubation period in the great tit (Parus major) population in central Poland, causing a large delay in hatching and had sustained reproductive consequences. This cold snap occurring once breeding activity had already started in 2017 was followed by the warm and invariable breeding season of 2018. This natural experiment had an essential influence on great tit reproductive parameters. We found a significant difference in hatching date, number of fledglings, hatching success, and fledging success between 2017 and 2018. In 2017, there were about two fledglings per nest fewer than in 2018. Fledging success was positively associated with hatching delay in 2017, while the relation was negative in 2018. Hatching success differed significantly between both years, being higher in 2018. Mean level of hemoglobin (used as index of body, physiological condition) in the blood of nestling great tits was higher in 2017 than in 2018. We argue that the moment of hatching may be (at least in some years) more tightly related to the moment of maximum food demand of tit nestlings than the traditionally used laying date. Also in extreme years, phenotypic plasticity of hatching delays may be insufficient to adjust the timing of breeding to the upcoming extreme weather events. Further examining its limits may be an important goal for future research.

In seasonal environments, a temporal decline in breeding performance (e.g. clutch size, nestling condition, and fledging success) of altricial bird species is a well-known phenomenon. In this study, we present the effect of laying phenology on the physiological condition of nestling great tits (Parus major) in 14 consecutive breeding seasons. We used blood haemoglobin and baseline glucose concentrations as indicators of nestling physiological condition. Nestling blood haemoglobin reflects food base quality and availability during the breeding period. Blood glucose concentration can be used as a supplementary reverse index of condition, since it is negatively related to environmental quality. It might be indicative of the stress caused by unfavourable extrinsic factors, though, due to potential confounding factors such as adverse weather conditions, low food quality, or feeding interruptions, glucose levels should be used in this ecological context with caution. Great tit nestlings from earlier broods were characterised by higher mean haemoglobin concentrations, indicating a seasonal decline in food quality and availability. The blood glucose concentration displayed an opposite pattern, with nestlings from earlier broods being characterised by lower mean concentrations than those from later broods. However, very little of the variation in blood glucose concentration can be explained by the variation in laying date, which suggests that blood glucose concentration is of little importance in the context of breeding phenology. Our results show that the physiological condition of nestlings of this species decreases as the breeding season progresses, most probably due to environmental factors.

The vegetation structure surrounding nest sites is a crucial component of habitat quality and may have large effects on avian breeding performance. Habitat quality reflects the extent to which the environmental characteristics of an area correspond to the preferred habitat characteristics of the species. The concentration of haemoglobin is considered a simple biochemical indicator of nestling body condition. We present results concerning the effects of variation in habitat characteristics on the concentration of haemoglobin in the blood of 14-day-old nestlings and breeding success of European Pied Flycatchers ( Ficedula hypoleuca) in a mature mixed deciduous forest in central Poland. The haemoglobin concentration of nestlings was higher when there were more native oaks and deciduous, native trees on the territory. Breeding success was reduced by the number of alien oak (the Red Oak, Quercus rubra) and other alien deciduous trees, but increased by the number of native oaks in the territory. This study was conducted on only one site so further research is needed to examine the broader applicability of our results. Our data support the idea that haemoglobin concentration may serve as a simple indicator of body condition in nestlings and is useful in field ecophysiology studies of European Pied Flycatchers.

Female birds are exceptionally consistent in laying rather uniform and individually specific eggs, so that variation in eggs of individual females is low, while variation among females is high. Repeatability is a population-level parameter used in quantitative genetics to quantify the level of intraindividual consistency in relation to interindividual variation in phenotypic traits, in this case, the egg traits. The statistical measure of repeatability is intraclass correlation of particular egg traits clustered in clutches. This study is based on a long-term dataset on repeatabilities of the length, breadth, volume and shape of eggs of Blue Tits Cyanistes caeruleus investigated in two ecologically varying sites (an urban parkland versus a mature deciduous forest) around the city of Łodź, central Poland in 1999–2013. Coefficients of repeatability of egg length, breadth, volume and shape in the study populations of Blue Tits showed some variation between years and study sites, but were generally very high, on average 0.7 or more, and did not significantly differ between the traits within years. The observed differences between repeatabilities do not seem to be strongly associated with year-specific weather indicators for the first half of April. Only coefficients of variation in mean daily temperature influenced repeatabilities of egg length and breadth, whereas effects of mean temperature were non-significant. Repeatability of egg length in the forest and breadth in both sites increased with increasing variation in daily temperature, whereas repeatability of egg length in the urban park site showed a reverse relation with variation in temperature. Year-site-specific mean laying dates, clutch sizes and egg volumes, considered as indicators of breeding conditions, tended to negatively affect the repeatabilities. However, in general, repeatabilities of different egg traits were not consistent in their variation between years and sites, which suggests an important role of stochastic factors in shaping this variation. External factors that are able to force females to diversify traits of eggs in a particular laying sequence do not seem to directly result from the environmental conditions prevailing during a spring period critical for laying eggs in a population in a given year (the first half of April), but seem to be linked to more subtle aspects of variation in the conditions for egg laying, as shown by links with the mean values of basic breeding parameters.

Variation in ambient temperature affects various life stages of organisms. It has been suggested that climate change not only implies higher global temperatures but also more unpredictable weather and more frequent extreme weather events. Temperature has a major influence on the optimal laying-incubation-hatching dates of insectivorous passerines, because it poses energetic constraints and affects the timing of food abundance. We have been studying breeding characteristics of great tits Parus major and blue tits Cyanistes caeruleus in two areas, an urban parkland and a deciduous forest, around the city of Łódź since 2002. During the egg-laying period in 2017, both tit species at both study areas faced an unusual cold spell as reflected by a sudden decrease in the mean ambient temperature to ca. 2–3 °C for about 5 days, which caused mean hatching delays of up to 6 days. Since flexibility of behavior plays a major role in adjusting to unpredictable weather conditions, examining its limits may be an important goal for future research.

Partial mortality in broods of altricial passerines results from various reasons, including deficiency of resources, inadequate parental care, diseases and other external factors. In this study we test two hypotheses concerning the phenomenon of partial mortality in nestling altricial passerines: (1) non-adaptive starvation hypothesis based on the assumption that partial brood losses are caused by starvation and undernourishment and has no adaptive value, and (2) adaptive starvation hypothesis assuming that losses from starvation might be elements of an adaptive strategy of reducing surplus nestlings in broods that are over-large. To test the above hypotheses we divided broods of two tit species Great Tit Parus major and Blue Tit Cyanistes caeruleus into two categories: broods with incomplete, less than 100% fledging success and broods with complete 100% fledging success. Subsequently, we compared physiological performance of nestlings between brood categories using blood hemoglobin concentratio...

Birds have specific habitat requirements during the breeding period. The Blue Tit Cyanistes caeruleus is a species evolutionarily associated with forest areas that prefers deciduous and mixed forests, whilst its breeding in urban areas is a relatively recent phenomenon. Long-term data (2002–2015) on reproductive performance (number of hatchlings, hatching success, number of fledglings and fledging success (fledging success as the number of fledglings in relation to the number of hatchlings)) were quantified for two Blue Tit populations in two, floristically and structurally contrasting areas (a mature deciduous forest and an urban parkland) in central Poland. The principal aim of this study is to see whether the habitat type, year or the food availability affect the breeding success of Blue Tits. Forest Blue Tits produced significantly more hatchlings (9.82 ± 2.64 (SD) in the forest vs.9.17 ± 2.16 in the parkland) and fledglings (9.18 ± 2.84 in the forest vs. 8.14 ± 2.68 in the parkland) than urban Blue Tits. The number of fledglings was positively correlated with the number of hatchlings in both study areas. Both forest and urban Blue Tits shared a similar hatching success (85.9 % in the parkland and 85.5 % in the forest), while the fledging success was significantly higher in the forest (83.4 % in the parkland and 86.1 % in the forest). The amount of caterpillar frassfall was also studied (caterpillars are the optimal food for nestlings) at both study areas and it suggested that caterpillars were more abundant in the forest than in the parkland (the maximum amount of frassfall, averaged 0.21 ± 0.11 g frass/m2/day in the urban parkland and 0.59 ± 0.50 g frass/m2/day in the forest in 2003–2015). In the forest area, the mean number of fledglings tended to be related to the amount of frassfall but in the parkland, this relation was non-significant. The long-term dynamics of fledging success in our study sites seems to be mutually independent. Thus low fledging success in the forest site does not mean similarly low fledging success in the urban parkland site and vice versa. We suggest that food availability is one of key drivers of differences in the tits breeding success between both studied habitats.

Plasma biochemistry values are useful tools for ecophysiological research of birds and other vertebrates. They provide data on the physiological and nutritional state of individuals in various life stages. We predicted that blood triglyceride levels correlate with nutritional state of birds and that this trait is temporally variable. Therefore, we studied the effects of year and distinct habitat type (urban parkland vs woodland) on triglyceride concentration in the blood of nestling Blue Tits Cyanistes caeruleus. We ascertained both seasonal and inter-habitat variation within the trait. Specifically, (1) nestlings from trophic rich forest site were characterised with significantly higher mean triglyceride levels, and (2) mean triglyceride content was significantly higher in one out of three breeding seasons. These results suggest that the studied trait may be used as a robust indicator of habitat trophic abundance and quality.

Abstract The blue tit (Cyanistes caeruleus) is one of a few known passerine birds that incorporate fresh fragments of aromatic plants in their nest during the nesting period. In this study, we experimentally tested whether aromatic plants in blue tit nests affect physiological condition of nestlings as indicated by hemoglobin and glucose in the blood. Nestlings in nests supplemented with fresh, aromatic plants had elevated blood hemoglobin, and therefore improved physiological condition, as compared with control nestlings. There was no difference between nestlings in two contrasting habitats (urban parkland and forest). Although mean levels of blood glucose did not differ between treatment and control nestlings, the blood of forest nestlings contained more glucose than the blood of parkland nestlings, which indicates poorer health for forest nestlings. In general, knowledge of the effects of incorporating fresh green, aromatic plant material into avian nests on physiological condition of nestlings is still fragmentary and more experimental studies are needed. Significance statement The blue tit is one of a few known passerine birds that incorporate fresh fragments of aromatic plants in their nest during the nesting period. Little is known about the impact of this material on nestling body condition, and there is a need to understand this behavior and its effects. We tested whether nestlings in nests supplemented with fresh, aromatic plants improved their physiological condition in comparison with control nestlings. The experiment showed that nestlings from supplemented nests improved their physiological condition by exhibiting elevated hemoglobin.

Urbanization affects the ecological and behavioral traits of various species of animals, including birds.We present results concerning long-term fluctuations in breeding densities of nest-box populations of the Blue Tit Cyanistes caeruleus and the Great Tit Parus major in two, structurally and floristically contrasting types of habitat (an urban parkland and a rich deciduous forest) located 10 km apart, in central Poland. This study was conducted in 1999–2012 in the parkland site and in 2002–2012 in the forest site. We found a strong correlation of year-to-year changes in breeding densities of Great Tits between the parkland site and the forest site and a lack of such a correlation in Blue Tits. Breeding densities of Great Tits were much higher in the parkland than in the forest area every year during the study period. Annual changes in breeding densities were not correlated between the species studied. The North Atlantic Oscillation Index (NAO-winter index) tended to influence the density dynami...

Many avian species in Europe breed earlier as a result of higher temperatures caused by global climate changes. Climate change means not only higher temperatures but also more frequent extreme weather events, sometimes contrasting with the long-term trends. It was suggested that we should look closely at every extreme phenomenon and its consequences for the phenology of organisms. Examining the limits of phenotypic plasticity may be an important goal for future research. Extremely low spring temperatures in 2013 (coldest spring in 40 years) resulted in birds laying unusually late, and it was followed in 2014 by the earliest breeding season on record (warmest spring in 40 years). Here, we present results concerning breeding phenology and double-broodedness in the Great Tit (Parus major) and the Blue Tit (Cyanistes caeruleus) in 2013 and 2014 in an urban parkland and a deciduous forest in central Poland. Great Tits started laying eggs 18.2 days later in 2013 than in 2014 in the parkland, whereas the analogous difference was 21.1 days in the forest. Blue Tits started laying eggs in the parkland 18.5 days later in 2013 than in 2014, while the analogous difference was 21.6 days in the forest. The difference in the proportion of second clutches in Great Tits between 2013 (fewer second clutches) and 2014 (more second clutches) was highly significant in the parkland and in the forest. This rather large extent of breeding plasticity has developed in reaction to challenges of irregular inter-annual variability of climatic conditions. Such a buffer of plasticity may be sufficient for Blue Tits and Great Tits to adjust the timing of breeding to the upcoming climate changes.

This study applies a field experimental approach to an issue of single lead exposures and their effects on specific haematologial indices in nestling songbirds. We used nestling great tits Parus major as a model and showed that higher lead exposure was associated with significant haematocrit decline. In contrast, the assessment of the H/L ratio revealed no significant effect of a single lead exposure., Lead (Pb) is recognized as one of the most toxic trace elements that can induce a wide range of negative health effects in wildlife. Because the investigation of basic environment-sensitive physiological indicators is easy to perform in wild birds, they have been considered as valuable bioindicators of lead contamination. The harmfulness of lead poisoning can depend on the type of exposure, and in most studies of birds, an effect of chronic lead exposition has been considered. In this study, we investigated whether a single exposure to specific doses of lead affected such physiological indices as haematocrit and the heterophil-to-lymphocyte ratio (H/L ratio). For this purpose, we conducted an experiment based on intentional lead supplementation, with the use of two different doses, applied to randomly chosen great tit (Parus major) nestlings from randomly selected broods. A few days after the exposure (when the nestlings were 15 days old), we determined haematocrit and the H/L ratio as potentially sensitive physiological indicators of lead intoxication. We found that the treatment with lead resulted in a significant decline in haematocrit level. In addition, we found that the age when lead exposure occurred can be considered as an important factor influencing haematocrit variation. A decrease in haematocrit was observed on consecutive days of nestling life. In contrast, the H/L ratio did not show any significant response to lead exposure. These results clearly show that the assessment of haematocrit level for nestling great tits can provide a simple and rapid method of indirect evaluation of physiological effects of lead intoxication caused by a single exposure.

Life-history theory suggests that macro- and micro-parasites may generate selection pressures by affecting host health. In the present study, we replaced natural great tit Parus major nests in two, structurally and floristically contrasting sites. Nestlings that were developing in treated nests improved physiological health status in comparison with control nestlings., Most passerines use nests as the exclusive place to lay and incubate eggs and bring nestlings up to fledging. Nests of secondary cavity nesters, like tits, provide a moist, warm and protected habitat for reproduction of blood parasites. Offspring fitness depends on interactions between parental care and environmental constraints. Life-history theory suggests that macro- and micro-parasites may generate selection pressures by affecting host health. In the present study, we replaced natural great tit Parus major nests in two, structurally and floristically contrasting sites (an urban parkland and a rich deciduous forest, located 10 km apart in Łódź, central Poland), with fresh, sterilized, artificial moss-cotton wool nests, twice, on the fifth and tenth day of nestlings life. We then examined haematological condition indicators (haemoglobin and glucose concentrations) of about 14-day-old nestlings. Nestlings that were developing in treated nests improved their health status in comparison with control nestlings. The mean haemoglobin and glucose concentrations (treated and control) also varied between both study areas. Our study confirms that the level of haemoglobin and especially the level of glucose may be treated as reliable indicator of environmental characteristics in great tits.

Numerous studies from different locations in Europe show that nest-box populations of tits in urban areas lay earlier and produce fewer eggs than do tits in rural areas. We collected data on laying dates and clutch size in two great tit Parus major populations nesting in oak deciduous forest and urban parkland areas, only about 10 km apart, in central Poland over 11 years. The abundance of caterpillars, the optimal food of breeding tits, at both breeding areas was also quantified. We analysed long-term trends in the timing of egg laying and clutch size. We focused on the effects of year, habitat type, insect availability and weather conditions shortly before egg laying on the patterns of variation in laying date and clutch size. In general, our study supports earlier generalisations on urban and rural tit populations. In particular, our finding that great tits initiated breeding consistently earlier in the urban site than in the forest area supports earlier studies on urbanised birds. The mean laying date was strongly and negatively associated with air temperature between 15 March and 15 April in both habitats. Mean clutch size was lower in the parkland than in the forest population, at least partly in response to the greater abundance of caterpillars in the forest. A novel result of this study is that clutch size differed between habitats to a varying degree in different years, in association with a stronger response to caterpillar abundance in the forest than in the park. Clutch size tended to decline with the progress of the breeding season within years in the parkland site, especially in “early years”, but not in the forest habitat.

Sequentially laid eggs of small songbirds are built from the current income of resources and therefore can be subject to different environmental influences on the short-term and long-term time scales. In this paper we study variation in egg length, breadth, volume and shape in Pied Flycatchers Ficedula hypoleuca in Łodź, central Poland, during 2002–2012, in relation to climatic and habitat-related factors. We found a significant decreasing trend in egg length, breadth and volume across the study years and a slight tendency for eggs to be a little more elongated in a forest site than in an urban parkland site. Even though mean monthly temperature in May is known to have increased since the 1970s in the area of Łodź, the average ambient temperature during the time of formation of eggs in particular clutches tended to decrease, with increasing variability at the same time, in 2002–2012. Egg size traits showed a positive association with average temperature and a negative relation with variability i...

Most passerines use nests as the exclusive place to lay and incubate eggs and bring nestlings up to fledging. Nests of secondary cavity nesters, like tits, provide a moist, warm and protected habitat for reproduction of blood parasites. Offspring fitness depends on interactions between parental care and environmental constraints. Life-history theory suggests that macro- and micro-parasites may generate selection pressures by affecting host health. In the present study, we replaced natural great tit Parus major nests in two, structurally and floristically contrasting sites (an urban parkland and a rich deciduous forest, located 10 km apart in Łodź, central Poland), with fresh, sterilized, artificial moss-cotton wool nests, twice, on the fifth and tenth day of nestlings life. We then examined haematological condition indicators (haemoglobin and glucose concentrations) of about 14-day-old nestlings. Nestlings that were developing in treated nests improved their health status in comparison with control nestlings. The mean haemoglobin and glucose concentrations (treated and control) also varied between both study areas. Our study confirms that the level of haemoglobin and especially the level of glucose may be treated as reliable indicator of environmental characteristics in great tits.

Egg size is a basic species-specific life-history trait in birds which influences offspring performance through the nestling stage and, therefore, fitness. Shells of snails are the principal source of calcium for female parids and some other passerines during egg formation. Shortages in the availability of snail shells may lead to laying smaller eggs with thinner shells. In urban green spaces that require protection against pests, the numbers of snails may become greatly reduced in response to molluscicide treatments. The use of a molluscicide treatment against pest slugs in the Botanic Garden, Łodź, Poland, in 2013 and 2014 provided a unique quasi-experimental setting to study the effect of molluscicide on both non-target snails and Blue Tit Cyanistes caeruleus eggs. Using a forest study site where no treatment was applied as a control area, we tested for effects of the molluscicide treatment in the Botanic Garden on the density of snails and on the egg volume of Blue Tits. We found that after the molluscicide treatment a dramatic drop in snail numbers occurred, which likely restricted the availability of snail shells for egg laying Blue Tits in the Botanic Garden. Correspondingly, egg sizes significantly decreased in the Garden, while they stayed constant over time in the forest. Eggs in bigger clutches were more strongly affected by the molluscicide treatment than eggs in smaller clutches. Our quasi-experimental study suggests that Blue Tits are sensitive to poor availability of snail shells. This is manifested by laying smaller eggs, which require less calcium to be incorporated in the eggshell, in response to a dramatic drop in the availability of snail shells. This result is expected to raise awareness of potential indirect and inadvertent effects of park pest management on biota other than molluscan pests.

In this study, we examined heavy-metal concentrations in feathers of nestling great tits Parus major and blue tits Cyanistes caeruleus at two different sites (urban parkland vs. deciduous forest) located in the Łódź agglomeration in relation to interyear variation. We found that tit species did not differ significantly in lead and cadmium concentrations. Zinc concentration was significantly higher in blue tits. We also found that lead and cadmium levels in blue tit nestlings and the level of lead in great tit nestlings were higher in the parkland site than in the woodland site. We explain habitat variation in heavy-metal concentrations in feathers of nestlings by different levels of contamination at study sites. For both tit species, significant variation in heavy-metal amounts accumulated by nestlings was found between years with the lowest value in a year with the lowest value of rainfall. We suggest that the interyear variation may be accounted for by differences in rainfall, thus influencing quantities of trace elements bioavailable in the environment.

The major aim of this study was to examine the influence of the European Pine Marten ( Martes martes) predation on the heights of nests of Blue Tits ( Cyanistes caeruleus) and Great Tits ( Parus major) in nestboxes of different security levels. To this end we performed a comparison of nest heights before and after introducing an anti-predator device to two size-types of wooden nestboxes set up in a deciduous forest. Blue Tit and Great Tit nest sizes were influenced by the application of plastic entrance tubes as anti-predator devices that elongated a distance between the entrance and the interior of the nestbox. Nests were taller in nestboxes equipped with anti-predator devices regardless of nestbox size. Nests located in smaller nestboxes were destroyed by Martens more frequently than nests built in bigger nest boxes. Most nest functions in cavity-nesting birds, such as filling the excessive cavity space, maintenance of proper humidity, thermal, light and sanitary conditions for eggs and nestlings, improve with nest sizes. Because, in contrast, the risk of nest predation by martens increases with nest size (height within the nestbox), the nest predation constitute a major factor that constrains cavity-nesting birds in constructing tall nests.

Enduring changes in the surrounding environment cause long-term stress in birds, which can affect the activity of the immune system and lead to changes in heterophil-to-lymphocyte ratios (H:L). The aim of this paper is to investigate if the H:L ratios of male and female Great Tits Parus major during the first breeding attempt differ between two sites of dissimilar habitats – an urban parkland site and a deciduous forest site. In the final model we found a difference between sites, while separate submodels for the study sites showed that females had higher H:L ratio than males in the urban parkland site, with the opposite result being found in the forest site. We suppose that the pattern of variation in H:L ratios we found may result from greater parental expenditures of females as compared to males. Such difference in the H:L ratio between habitats could be the reason for the substantial difference in trophic conditions between the forest and park areas during the breeding of tits.

The aim of this study was to determine the possibility of using feathers of blue tit nestlings to assess the level of endogenous accumulation of lead. For this purpose we conducted an experiment with lead application to randomly chosen nestlings from eight randomly drawn broods. Five days after the exposure, feathers of lead-treated nestlings had significantly higher lead concentrations than control nestlings. This result suggests that feathers can be used as reliable non-destructive bioindicators to assess the level of heavy metals originating from contaminated food, which is of great significance for comparative studies on ecological consequences of pollution.

Physiological functions of growing nestlings are thought to be traded-off in relation to rearing conditions, with the resulting physiological state of fledglings having important long-lasting consequences for their fitness. By manipulating brood size up and down, and, separately, by supplying additional food (mealworms — larvae of Tenebrio molitor) we tested if alterations of the rearing conditions would influence nestling performance in Blue Tits Cyanistes caeruleus and Great Tits Parus major. Brood size manipulation affected body mass, heterophil-to-lymphocyte ratio (H:L) and fledging probability in both species and the level of triglycerides in nestling Great Tits. Extra food supply influenced only fledging probability, with no other effect on indicators of nestling performance. An effect on nestling body mass and a lack of effect on cell-mediated immune response in the brood-size experiment suggest that nestlings in enlarged broods sacrificed growth to maintain immunity. In general, effects ...

The Great Tit Parus major and the Blue Tit Cyanistes caeruleus are the only Western Palearctic Parids that maintain numerous urban populations as well as forest populations. Because of their evolutionary history both these species are best adapted to different types of deciduous and mixed forests. Ecological conditions in cities are different from those dominating in forests, especially in such aspects as: habitat fragmentation, tree species composition, microclimate, human activity, predators and food conditions. The tits breeding in cities start laying eggs earlier in the season, lay smaller clutches and fledge fewer fledglings of lower quality. Yet urban populations are often relatively stable in numbers. This may result from the fact that survival of winter is higher in cities due to increased availability of food and milder weather.

High metabolic rates of birds demand an efficient oxygen transport system; this is ultimately based on the oxygen carrying capacity of haemoglobin. Therefore patterns of variation in blood haemoglobin content of wild birds are an important aspect of functional ecology. In this paper we report results concerning variation in haemoglobin con- centration in the blood of adult Blue Tits Cyanistes caeruleus and Great Tits Parus major during the nestling rearing peri- od (days 7-15 of nestlings life) of first broods in central Poland in 2003-2009. The most important findings of this study are: (i) average haemoglobin concentration in blood differs between Blue Tits and Great Tits, with higher values in Blue Tits; (ii) males differ from females in both these species, with higher values in females, and (iii) there is also significant variation among years, with parallel tendencies for both species. We explain the patterns of haemoglobin content vari- ation in adult tits by differences in metabolic demands for oxygen transported by blood. The demands are higher for the smaller-bodied species (Blue Tits), heavier working sex (females) and in years with worse physical and trophic con- ditions, though only non-significant relationships with weather conditions (temperature and rainfall) or food availabil- ity (measured by frassfall) were found during our long-term study.

Wild passerines, especially tits, utilize snail shells as the main source of calcium necessary for laying females to construct egg shells. This research found that the two study areas, representing two habitat types — a mature decid- uous forest and a human-disturbed parkland — are inhabited by different snail assemblages: both species richness and density are much higher in the parkland than in the forest. This means that less calcium is available to female tits in the forest than in the parkland, which could result in calcium limitation in the former habitat. Egg size traits, i.e. volume, length and breadth, in the Blue Tit show a consistent long-term pattern of variation that reflects the pattern of calcium availability: egg trait values are higher in the parkland than in the woodland. No habitat-related variation in egg size traits was found in the Great Tits. We suggest that the lack of a relation between Great Tit egg characteristics and snail availability results from the higher ecological plasticity of this species in comparison with the Blue Tit, including its abil- ity to exploit alternative sources of calcium.

The physiological condition of nestling altricial birds depends on the quantity and quality of food delivered to them by parents. One indicator of the condition of Great Tit Parus major nestlings is the haemoglobin concentration in their blood. The present study demonstrates the influence of weather conditions (temperature and rainfall) on nestling haemoglobin concentrations during two consecutive breeding seasons in two different habitat types (parkland vs. woodland) in the city of Łodź in Central Poland. This influence probably results from the effects of weather on the trophic base of the Tits. Dry, hot weather strongly affected bush and herbal foliage later in the breeding season (mid-June to mid-July) in 2006, presumably by interfering with the development of herbivorous arthropod populations. This in turn caused food shortages for second broods of Great Tits, which resulted in nestlings having low haemoglobin levels. In the following year, temperature was on average lower, and rainfall was regular but not very heavy. These conditions enabled the development of arthropod assemblages, and the trophic base for birds was much richer. Haemoglobin concentrations in the blood of nestlings from second broods were significantly higher than those of first broods and, unexpectedly, second-brood nestlings in 2007 were on average in better physiological state than first-brood nestlings in 2006 in both habitats. The relationship between haemoglobin concentration, brood category and year was very similar to that for nestling body mass. However, it was independent of both body mass and brood size. In some years and under certain conditions, second broods can be more successful than first broods.

Producing high quality offspring of good physiological performance, able to survive to independence and, then, to reproductive maturity is a major component of life history strategies. The ability of nestling altricial birds to develop a good physiological condition depends to a large extent on the amount and quality of food provided by par- ents, as well as other aspects of parental care. We hypothesized that experimental changes to the original brood size should affect both parental Blue Tits and their offspring, resulting in corresponding changes in the body condition of the nestlings. Over two breeding seasons, using two habitat sites, we conducted an experiment with two manipulative treatments applied to broods of three-day-old nestlings — the reduction or enlargement of broods by three nestlings, and one non-manipulative control treatment. Our aim was to test whether the experiment would affect a number of different measures of nestling condition: blood concentrations of hemoglobin and glucose, heterophil-to-lymphocyte ratio and morphometric condition index, all being analyzed when the nestlings were 13 days old. We found no effect in the case of hemoglobin, despite the fact that it had previously been shown to be sensitive to large-scale differences in trophic conditions between habitats and years and to the experimental removal of nest parasites. All the remaining variables, i.e. heterophil-to-lymphocyte ratio, glucose concentration and morphometric condition index, responded to the experimental treatments, showing different but reasonable patterns of variation. We suggest that an experimental increase in brood size definitely hinders the development of nestling physiological condition, but even an experimen- tal reduction of broods can affect some physiological indicators (glucose), probably because of readjustments in the feeding rate.

Nutrition during the time spent at the nest is critical for the development of the body condition in altricial birds, including nestling blue tits (Cyanistes caeruleus). Trophic conditions tend to be variable in time and space, which should influence the physiological condition of growing birds. In this paper, we consider haemoglobin concentration as a general indicator of chick physiological condition. Its variation is both consistent within broods and indicative/prognostic of survival from the moment of hatching to fledging. We show that haemoglobin concentration of nestling blue tits is on average higher in years and habitats characterised by rich abundance of caterpillars than under poor caterpillar availability. Haemoglobin seems to quite exactly reflect even relatively subtle differences in trophic conditions.

Theory suggests that macro- and micro-parasites may be important factors of selection for life-histories. They generate selection pressures by detrimental effects on host health. Nests of secondary cavity nesters provide a convenient habitat for an assemblage of parasites exploiting nestlings. In this study, natural blue tit Parus caeruleus nests (26) were replaced with clean artificial nests, twice during the nestling stage. This treatment caused an increase of 7–10.5 g/l in hemoglobin level of 12-day-old nestlings in comparison with control nestlings. Nestlings that developed in parasite–pathogen-free nests improved their health status. The experimental sterilization did not affect a morphometric index of condition. Potential effects on condition indices might be masked by trophic conditions.

Environmental factors affecting trophic conditions act as stressors on nestling altricial birds. Access of parental birds to a sufficient supply of food in a limited period of the nestling stage differ in time and space, depending on nesting habitat, prey density and weather conditions. Heterophil-to-lymphocyte ratio (H/L) is considered as a reliable indicator of prolonged stress reaction in birds. In this study we examine if variation in H/L shows consistent spatio-temporal patterns in nestlings of two parids, blue tit Cyanistes caeruleus and great tit Parus major. We found that blue tit nestlings had on average higher H/L than great tit nestlings, which corresponds with the ecological sensitivity of these species. In both species H/L was higher in a poor parkland habitat than in a high quality forest habitat. In nestling blue tits, higher H/L values occurred in years characterized by more extreme weather conditions and worse caterpillar availability. Such consistent patterns of variation in the H/L ratio of nestling blue tits and great tits suggest that, when age-dependent effects are controlled, the ratio can be used as an indicator of physiological stress that is generated by food-related stressors differing in space and time. In particular, elevated H/L ratios are indicative of human-induced changes in the structure of breeding habitats.

There is a need to study the effects of urbanization on wildlife in order to understand the ecological implications of increasing urbanization and find out how to reduce its threats to biodiversity. The blue tit evolved as a forest species and prefers deciduous and mixed forests, whereas its nesting in urban habitats is a more recent phenomenon. Our long-term study of blue tit populations has been conducted in two habitats: an urban parkland (frequently visited by people) and a deciduous forest outside of the city. Using linear mixed modeling, we revealed that a relationship of blue tit breeding success (and the number of fledglings) with thermal conditions in May differed between the urban parkland and the forest. While the relationship was positive in the forest, it was negative in the parkland. In addition, breeding success in the parkland increased with increasing number of rainy days in May. We argue that the main possible reason for such patterns is human activity in the parkland, which interferes with tit parental care, especially the regular feeding of nestlings, whereas it is evidently associated with weather conditions. Human disturbance in the forest is likely to be negligible.

Physiological tools can be used to identify the sources and consequences of stressors on animals. Understanding the influences of variation in habitat quality and anthropogenic disturbance on organism condition and health may improve future management and conservation. We present results concerning variation in haemoglobin and glucose concentrations in the blood of about 14-day-old nestling Pied FlycatchersFicedula hypoleuca in central Poland over a 4-year period, 2011–2014, in a deciduous forest. The most important findings of the study are: (1) the concentration of haemoglobin and glucose of the nestlings from the same brood tended to be consistently similar, with much variation occurring among broods; (2) repeatability of haemoglobin concentration was higher than repeatability of glucose concentration; (3) mean levels of haemoglobin and glucose varied among years; (4) haemoglobin and glucose concentrations were negatively correlated; and (5) there was a positive relationship between haemoglobin levels and breeding success. All procedures were approved by the Local Ethical Committee and the State Office for Environment Protection. We thank A. Jaksa, D. Man´kowska, M. Janiszewska and J. Białek for their help and consent in conducting research in the areas under their administration. The study was founded by a grant from the Polish Ministry of Science and Higher Education No. N N304 045136 and University of Ło´dz´ (No. 506/829). We are obliged to P. Procter for linguistic consultation. We thank both reviewers for their valuable and constructive comments.

The impact of climatic changes on life cycles by re-scheduling the timing of reproduction is an important topic in studies of biodiversity. Global warming causes and will probably cause in the future not only raising temperatures but also an increasing frequency of extreme weather events. In 2013, the winter in central and north Europe ended late, with low temperatures and long-retained snow cover--this extreme weather phenomenon acted in opposition to the increasing temperature trend. In 2013, thermal conditions measured by the warmth sum in the period 15 March–15 April, a critical time for early breeding passerines, went far beyond the range of the warmth sums for at least 40 preceding years. Regardless of what was the reason for the extreme early spring 2013 and assuming that there is a potential for more atypical years because of climate change, we should look closely at every extreme phenomenon and its consequences for the phenology of organisms. In this paper, we report that the prolonged occurrence of winter conditions during the time that is crucial for Blue Tit (Cyanistes caeruleus) and Great Tit (Parus major) reproduction caused a substantial delay in the onset of egg laying in comparison with typical springs.

Integration of landscape ecology and conservation physiology has been recommended as a potentially useful way to investigate consequences of human-induced changes in habitats for animal populations. A central goal of this paper was to examine if a simple physiological parameter displays any consistent patterns of spatio-temporal variation. Blood glucose concentration in birds reflects their high metabolic demands and may be influenced by a number of environmental factors. Therefore we present results concerning variation in glucose concentration in the blood of c. 14-day-old nestling blue tits (Cyanistes caeruleus) in central Poland in an 8-year period, 2005–2012, in two landscapes: an urban parkland and a deciduous forest. The most important findings of the study were: (1) mean levels of blood glucose varied markedly among years, most probably due to variable weather conditions, (2) glucose concentrations were significantly higher in the parkland study site than in the forest site, (3) heavier nestlings had lower glucose levels, and (4) high glucose levels were negatively correlated with fledging and breeding success. Thus we have confirmed that a consistent spatio-temporal pattern really exists.

Dates of egg laying, clutch size, the numbers of hatchlings and fledglings, and egg dimensions were studied in Annaba, Algeria in 2004-2005. The onset of laying is related to the end of the wet period and shows some inter-year variation. 38-45% of breeding pairs produce two broods per year. Mean clutch size declines with the progress of the breeding season. Mean values vary between the first (4.7 eggs) and second clutch (3.5 eggs) but not between years. Hatching and fledging success is relatively low in comparison with other Western Palearctic populations, probably because of environmental constraints operating at the southern edge of the House Martin breeding area. The relatively small size of Algerian House Martin eggs fits the general geographic pattern of variation.