Hematological health state indices predict local survival in a small passerine bird, the great tit (Parus major)

In birds, it has been shown that reproductive effort may impair parental condition, while the relation of different condition indices to subsequent survival is still poorly understood. In this study, we measured body mass and various hematological condition indices in breeding great tits in relation to local survival. Number and quality of nestlings and the occurrence of second broods, potentially reflecting parents' breeding effort, were also considered in analyses. The great tits, both male and female, that returned the following year had had a higher albumin/globulin ratio, lower plasma globulin concentration, and a lower heterophile/lymphocyte ratio during breeding in the preceding year, compared to those who did not return. Surviving males (but not females) also had had a higher level of circulating lymphocytes, compared to nonsurvivors. There was no correlation between breeding effort and survival. We conclude that better immunological state and lower stress in great tits during breeding were positively related to their survival probability.     

The effects of spatial and temporal ecological variation on fatty acid compositions of wild great tits Parus major

In birds, fatty acids (FA) have three main functions; they are structural components of cell membranes, metabolic fuel, and inflammatory molecules. Environmental factors, such as diet and ambient temperature, affect FA composition, thereby function and ultimately fitness. Thus, variation in FA compositions can be the underlying mechanism for varying performance of birds in different habitats. Here we examine variation in plasma FA composition in nestling and adult great tits Parus major, between 1) deciduous and coniferous, and 2) sun‐exposed and shaded habitats. The main results revealed that nestlings had a higher proportion of α‐linolenic acid (α‐LNA) in deciduous habitats and arachidonic acid in coniferous habitats. This reflects a difference in caterpillar availability between habitats with the deciduous habitat being caterpillar‐rich, whereas the coniferous habitats are rich in spiders. In addition, α‐LNA increased with nestling body condition in the coniferous habitat, supporting the importance of caterpillars for fledging success in this species. In line with dietary intake, the proportion of the essential α‐LNA and linoleic acid (LA) increased over the course of the day for all birds. In the deciduous habitat, adult females showed a positive association between LA and body condition. Furthermore, habitat sun‐exposure showed significant interactions with body condition for polyunsaturated FAs in nestlings, and with saturated FA in adult males, which is in accordance with the homeoviscous hypothesis stating that the proportion of saturated FA should decrease with decreasing ambient temperature. Taken together, small‐scale heterogeneity in habitat structure significantly influences FA compositions of great tits. Many of the results can be linked to dietary, and possibly, ambient temperature differences between habitats. These habitat effects on FA compositions can lead to different capacities of individual birds to deal with infections and low temperatures, two stressors that cause major mortality among wild birds.     

Plumage Bacterial Assemblages in a Breeding Wild Passerine: Relationships with Ecological Factors and Body Condition

Microorganisms have been shown to play an important role in shaping the life histories of animals, and it has recently been suggested that feather-degrading bacteria influence the trade-off between parental effort and self-preening behavior in birds. We studied a wild breeding population of great tits (Parus major) to explore habitat-, seasonal-, and sex-related variation in feather-degrading and free-living bacteria inhabiting the birds' yellow ventral feathers and to investigate associations with body condition. The density and species richness of bacterial assemblages was studied using flow cytometry and ribosomal intergenic spacer analysis. The density of studied bacteria declined between the nest-building period and the first brood. The number of bacterial phylotypes per bird was higher in coniferous habitat, while bacterial densities were higher in deciduous habitat. Free-living bacterial density was positively correlated with female mass; conversely, there was a negative correlation between attached bacterial density and female mass during the period of peak reproductive effort. Bacterial species richness was sex dependent, with more diverse bacterial assemblages present on males than females. Thus, this study revealed that bacterial assemblages on the feathers of breeding birds are affected both by life history and ecological factors and are related to body condition.     

Calcium shortage as a constraint on reproduction in great tits Parus major: a field experiment

Several bird species breeding in acidified habitats face serious problems caused by lowered availability of calcium-rich foods. Moreover, recent studies have revealed that calcium shortage causes phenotypic variation in certain reproductive parameters of passerines even in non-acidified habitats. However, to date, no clear evidence has been found of the limiting effect of calcium deficiency on the reproductive output of birds breeding in such habitats. In this paper, we present experimental evidence from a long-term study that calcium shortage may indeed limit reproductive output of passerines that breed in non-acidified habitats. We carried out calcium-supplementation experiments with great tits Parus major breeding in coniferous and deciduous woodlands in Estonia. Calcium-supplemented birds laid more eggs, tended larger broods and raised more fledglings with longer tarsi compared with control birds. Calcium shortage constrained reproduction of tits not only in spring, but also in mid-summer, during second breeding attempts. Moreover, no significant differences were detected between deciduous and coniferous habitats with respect to the effect of calcium supplements on reproductive success.     

Parental provisioning behaviour in Pied Flycatchers Ficedula hypoleuca is well adjusted to local conditions in a mosaic of deciduous and coniferous habitat

Capsule Pied Flycatchers are better able than Great Tits to adjust their feeding behaviour to varying conditions in the same area.

Aims Great Tits breeding in a mosaic of deciduous and coniferous forests in the northern temperate region exhibit consistently lower breeding success in their preferred deciduous habitat than in coniferous habitat. This was explained by the unexpectedly poor nestling feeding conditions in deciduous forests of this region. We studied whether the same paradox applies to Pied Flycatchers that occupy the same habitats in the same area.

Methods Parental provisioning behaviour was studied using video‐recording and experimental manipulation. Caterpillar abundance and basic breeding parameters were measured in different habitat types.

Results Parental provisioning frequency and the proportion of caterpillars in nestling diet was lower, while food objects were on average larger, in coniferous compared with deciduous habitat. However, the total volume of caterpillars and adult Lepidoptera delivered to nestlings did not differ between habitats. In contrast to Great Tits, offspring body parameters in Pied Flycatchers did not differ between habitat types.

Conclusions These results demonstrate how the relative suitability of particular habitat types varies between species and is dependent upon geographical location.     

Provision of nestboxes raises the breeding density of Great Tits Parus major equally in coniferous and deciduous woodland

Nestbox provision is a technique used to increase nest-site availability for secondary cavity-nesting birds. However, little is known about the demographic consequences of nestbox provision in different habitat types. To assess how nestbox provision affects the density of hole-nesting birds simultaneously in two contrasting habitats, we compared the breeding density of Great Tits along transects without nestboxes with that in transects where nestboxes were provided. Although the initial density of breeders was considerably higher in the deciduous habitat than in the coniferous habitat, provision of nestboxes increased density by a similar number of additional pairs in each habitat type. Thus, the provision of nestboxes in managed coniferous forests may be as effective in increasing the breeding opportunities of cavity nesters as in deciduous stands. Moreover, previous research showed that pairs in deciduous habitat with nestboxes have consistently lower breeding success than those in coniferous habitat with nestboxes. It is possible that the addition of nestboxes in the preferred habitat increased density to such an extent that density-dependent effects became apparent.     

Providing nest boxes for hole-nesting birds – Does habitat matter?

Nest boxes are a popular management tool to increase nest site availability for hole-nesting birds, but biological consequences of this technique in different habitats are poorly studied. In our study area in southwestern Estonia, nest boxes for small passerines were set up in deciduous and coniferous woods. Great tits Parus major preferred the food-rich deciduous habitat for breeding, as judged by higher nest-box occupation, earlier egg-laying and larger clutches and eggs. However, in coniferous habitat more and heavier young fledged per nest, and the return rate of both fledglings and adults was higher. We propose two mutually non-exclusive explanations, both related to the maladaptive outcome of the provision of nest boxes: (i) in the preferred habitat, nest boxes caused a supra-optimal breeding density leading to an ecological trap; (ii) boxes drastically improved the non-preferred habitat, but birds were unable to exploit the breeding habitat fully. One should be careful in providing large numbers of artificial nest sites in preferred habitats. Sometimes it would be more preferable to improve less favourable habitats by removing critical constraints.     

Body Condition Indices Predict Reproductive Success but Not Survival in a Sedentary,Tropical Bird

Body condition may predict individual fitness because those in better condition have more resources to allocate towards improving their fitness. However, the hypothesis that condition indices are meaningful proxies for fitness has been questioned. Here, we ask if intraspecific variation in condition indices predicts annual reproductive success and survival. We monitored a population of Neochmia phaeton (crimson finch), a sedentary, tropical passerine, for reproductive success and survival over four breeding seasons, and sampled them for commonly used condition indices: mass adjusted for body size, muscle and fat scores, packed cell volume, hemoglobin concentration, total plasma protein, and heterophil to lymphocyte ratio. Our study population is well suited for this research because individuals forage in common areas and do not hold territories such that variation in condition between individuals is not confounded by differences in habitat quality. Furthermore, we controlled for factors that are known to impact condition indices in our study population (e.g., breeding stage) such that we assessed individual condition relative to others in the same context. Condition indices that reflect energy reserves predicted both the probability of an individual fledging young and the number of young produced that survived to independence, but only during some years. Those that were relatively heavy for their body size produced about three times more independent young compared to light individuals. That energy reserves are a meaningful predictor of reproductive success in a sedentary passerine supports the idea that energy reserves are at least sometimes predictors of fitness. However, hematological indices failed to predict reproductive success and none of the indices predicted survival. Therefore, some but not all condition indices may be informative, but because we found that most indices did not predict any component of fitness, we question the ubiquitous interpretation of condition indices as surrogates for individual quality and fitness.     

Sex ratio of Parus major and P. caeruleus broods depends on parental condition and habitat quality

Brood sex ratio was studied in 88 families of Parus caeruleus (blue tit) and 95 families of P. major (great tit) in deciduous and mixed forest habitats differing in food availability. As a food specialist, the blue tit is expected to be more sensitive to the nutritional differences between the habitats than a food generalist such as the great tit. A shift of brood sex ratio towards males was detected for great tits in the high quality habitat, but there was no significant impact of parental condition or the number of nestlings. In contrast, brood sex ratio of blue tits was not affected by habitat quality. In blue tits, male condition correlated positively with a male-biased sex ratio. Habitat quality, however, affected the body mass differences of male and female blue tit siblings, and nestlings developed differently. The low quality habitat had a negative effect on the sexual dimorphism of siblings in male-biased broods, and the condition of offspring was bad. Nevertheless, sexual dimorphism cannot explain the differences between great and blue tits with respect to the correlation of sex ratio and individual condition.     

Personality and urbanization: behavioural traits and <Emphasis Type="Italic">DRD4</Emphasis> SNP830 polymorphisms in great tits in Barcelona city

Most examples of adaptation to the urban environment relate to plasticity processes rather than to natural selection. Personality, however, defined as consistent individual differences in behaviour related to exploration, caution, and neophobia, is a good behavioural candidate character to study natural selection in relation to the urban habitat due to its heritable variation. The aim of this paper was to analyse variation in personality by comparing urban and forest great tits Parus major using standard tests of exploratory behaviour and boldness. We studied personality in 130 wild great tits captured in Barcelona city and nearby forests and found that urban birds were more explorative and bolder towards a novel object than forest birds. Genotype frequencies of the DRD4 SNP830 polymorphism, a gene region often associated with personality variation, varied significantly between forest and urban birds. Behavioural scores, however, were not correlated with this polymorphism in our population. Exploration scores correlated to boldness for forest birds but not for urban birds. Our findings suggest that the novel selection pressures of the urban environment favour the decoupling of behavioural traits that commonly form behavioural syndromes in the wild.     

Habitat quality as a predictor of spatial variation in blue tit reproductive performance: a multi-plot analysis in a heterogeneous landscape

Vertebrate studies have rarely investigated the influence of spatial variation in habitat richness on both short-term (breeding) and long-term (offspring recruitment) reproductive performance using simultaneously multi-patch, multi-habitat type and multi-year approaches at landscape level. Here we present results of such an approach using the influence of two oak tree (Quercus ilex, Q. humilis) species on reproductive performance in Corsican blue tits (Parus caeruleus ogliastrae) as a model system. We found that blue tits breeding in rich broad-leaved deciduous patches consistently laid eggs earlier in the season, and produced larger clutches and more fledglings of higher quality, than those breeding in poor evergreen patches. Also, parents, especially males, were in better physical condition in the broad-leaved deciduous than in the evergreen patches. Surprisingly, estimates of long-term effects of reproduction, such as recruitment rates of locally born offspring, did not differ between the two habitat types. Our results suggest that short-term breeding performance and phenotypic quality of both chicks and parents do not necessarily provide reliable information about contributions to following generations at a scale larger than that of the local study plot. Differences in reproductive performance between the two oak habitat types could not be attributed to density-dependent effects, differences in levels of nest predation, or differences in age structure of the birds. We suggest that habitats that are optimal for breeding are not necessarily optimal for survival after the breeding season.     

Seasonal variability in habitat structure may have shaped acoustic signals and repertoires in the black-capped and boreal chickadees

Many songbird species have evolved multiple vocalizations, or repertoires, that function to communicate various biological signals. More diverse repertoires may have evolved in response to the effects of seasonal variation in habitat structure on signal transmission. Such changes in habitat necessarily occur for migrating species, but they also occur for resident species that occupy deciduous forests. The black-capped chickadee (Poecile atricapillus) possesses a chick-a-dee call and a fee-bee song, but the closely related boreal chickadee (P. borealis) lacks a song. Consistent with the habitat variability hypothesis, the black-capped chickadee possesses a larger repertoire and primarily occupies deciduous forests, whereas the songless boreal chickadee occurs more often in coniferous forests. We explored the ecological basis of this hypothesis by recording audio playbacks of two species in two habitat types during two seasons. Specifically, we played both songs and calls of the black-capped chickadee and calls of the boreal chickadee in deciduous and coniferous habitats, prior to and after leaf-out. We measured attenuation and degradation in re-recorded vocalizations. For black-capped chickadees, the song was less degraded than the call in post-leaf, deciduous forests. The boreal chickadee call attenuated more quickly in all treatments, but maintained its acoustic structure better than both black-capped chickadee vocalizations in coniferous forests. Our results support the hypothesis that variable habitats provided a seasonal transmission benefit for both song and call in the black-capped chickadee, but that the transmission benefit of song is lost in the less variant coniferous forests, which may underlie the absence of a song in the boreal chickadee.     

Manipulation of laying effort reveals habitat-specific variation in egg production constraints in Great Tits (Parus major)

It has been suggested that a bird's clutch size is not limited by the amount of resources available at the time of laying but that differences in the availability of food for nestlings is the ultimate underlying factor determining spatio-temporal variations in clutch size. However, habitat-related variations in egg production ability has yet to be investigated explicitly. We studied the breeding of Great Tits Parus major in deciduous and coniferous forests in the same area. The sizes of both the clutches and the eggs were, on average, larger in the former habitat than in the latter. A number of females were induced to lay more eggs than usual by removing four eggs from designated experimental clutches early in the laying period. These manipulated females laid approximately one egg more than control females, with the number of additional eggs laid not differing between the habitats. However, in both study years the relative size of the extra eggs – relative to the mean size of earlier laid eggs of the same clutch – was smaller in the coniferous habitat than in the deciduous habitat, while there was no habitat-related difference in the relative size of the last-laid eggs of control clutches. This result indicates that some form of proximate limitation during egg-laying period can contribute to the relatively small clutches and eggs in the coniferous habitat. Our results emphasize the need to take egg production costs into account when attempting to account for spatial variation in the reproductive behaviour of birds.     

Geographic variation in breeding parameters of the Pied Flycatcher Ficedula hypoleuca

Variation in laying date, clutch-size and number of fledglings of the Pied Flycatcher Ficedula hypoleuca was studied in relation to altitude, habitat and latitude, using data from 99 breeding areas from 35̀ N to 70̀ N. Laying date significantly increased with altitude and showed a significant quadratic relationship with latitude, with the earliest values at about 50̀ N. This quadratic relationship shows the same pattern as the relationship between arrival time of adults on the breeding areas and latitude. Clutch-size variation was significantly related to altitude and habitat, small clutches being laid at high altitude and in coniferous forests. Number of fledglings was significantly affected by habitat type, with lower values in the coniferous than in the deciduous forest. Mean clutch-size and mean number of fledglings showed the same quadratic relationship with latitude. In contrast to some earlier studies, an increased clutch-size with latitude was found, mainly because of new data from areas in the southern part of the species' distribution.     

Nonrandom dispersal drives phenotypic divergence within a bird population

Gene flow through dispersal has traditionally been thought to function as a force opposing evolutionary differentiation. However, directional gene flow may actually reinforce divergence of populations in close proximity. This study documents the phenotypic differentiation over more than two decades in body size (tarsus length) at a very short spatial scale (1.1 km) within a population of pied flycatchers Ficedula hypoleuca inhabiting deciduous and coniferous habitats. Unlike females, males breeding in the deciduous forest were consistently larger than those from the managed coniferous forest. This assortment by size is likely explained by preset habitat preferences leading to dominance of the largest males and exclusion of the smallest ones toward the nonpreferred coniferous forest coupled with directional dispersal. Movements of males between forests were nonrandom with respect to body size and flow rate, which might function to maintain the phenotypic variation in this heritable trait at such a small spatial scale. However, a deeply rooted preference for the deciduous habitat might not be in line with its quality due to the increased levels of breeding density of hole‐nesting competitors therein. These results illustrate how eco‐evolutionary scenarios can develop under directional gene flow over surprisingly small spatial scales. Our findings come on top of recent studies concerning new ways in which dispersal and gene flow can influence microevolution.     

Long-term consequences of early ontogeny in free-living Great Tits Parus major

Environmental factors during early development may have profound effects on subsequent life-history traits in many bird species. In wild birds, sex-specific effects of early ontogeny on natal dispersal and future reproduction are not well understood. The objective of this work was to determine whether hatching date and pre-fledging mass and condition of free-living Great Tits Parus major have any subsequent effect on individuals’ natal dispersal and reproductive performance at first breeding. Both males and females dispersed longer distances in coniferous than in deciduous forests, while dispersal was condition-dependent only in males (heavier as nestlings dispersed farther). In females, mass and condition at pre-fledging stage correlated significantly with clutch size, but not with subsequent reproductive performance as measured by fledging success or offspring quality. In contrast, heavier males as nestlings had higher future fledging success and heavier offspring in their broods compared with those in worse condition as nestlings. The hatching date of female as well as male parents was the only parental parameter related to the number of eggs hatched at first breeding. These results indicate that pre-fledging mass and condition predict subsequent fitness components in this bird species. We suggest that sex-specific relationships between a disperser’s condition and its selectivity with respect to breeding habitat and subsequent performance need to be considered in future models of life-history evolution.     

Female genetic heterogeneity affects the reproduction of great tits (Parus major L., 1758) in low-quality woodlands

Genetic heterogeneity is considered an important parameter for individual fitness and reproductive success. In 1999 and 2000, we studied the population genetics of great tit ( Parus major L., 1758) in southwestern Germany from two different forest types (deciduous and mixed-coniferous), which may significantly differ in prey diversity and/or food supply. Adults of 99 families were genotyped at four enzyme and eight microsatellite loci, in order to estimate individual heterozygosity. In the mixed-coniferous forest, a significant positive correlation between the genetic heterogeneity of females and early egg-laying date and clutch size was detected. Early egg-laying date and increased clutch size are conditions that positively affect the number of fledglings. This effect of individual heterozygosity was not observed in the deciduous woodland. Maternal genetic heterogeneity, however, did not correlate with fledgling condition, and individual heterozygosity of fathers had no impact on breeding success in either habitat. The positive effect of female genetic heterogeneity on brood size of great tits in mixed-coniferous forests is attributed to early egg-laying date, i.e. a maternal effect, rather than to a specific mating strategy that optimizes fitness through an increased brood size and the quality of offspring.     

四川小寨子沟自然保护区大熊猫生境植物群落结构特征

通过沿海拔梯度的系统调查,对小寨子沟自然保护区大熊猫生境的植物群落种类组成和结构进行了初步研究。结果表明:(1)小寨子沟大熊猫生境的植被类型有2种:阔叶林和针叶林;(2)小寨子沟自然保护区内分布有青川箭竹、缺苞箭竹、华西箭竹、冷箭竹等多种箭竹,这些作为大熊猫食料的箭竹广布在各种植被类型中,尤其是针阔混交林和亚高山针叶林;(3)大熊猫对生境的群落结构有较广的适宜性,阔叶林、针阔混交林、亚高山针叶林均可作为它的适宜生境;(4)小寨子沟自然保护区内的植被类型是随海拔的升高而自然过渡的,呈现出受人类活动影响较小的自然状态。     

Winter bird communities in the Åland archipelago: an island biogeographic point of view

The distribution of wintering land birds was examined in the archipelago of Åland (60°N, 20°E) along two ecological dimensions: (1) the island continuum from small skerries ((0.5 ha) to the mainland of Åland (97000 ha), and (2) the habitat range of the mainland of Åland. In the species-area relationship, both the exponential and power function models fitted well. The increase of the number of species with island size was interpreted as a consequence of increasing habitat diversity on larger islands. The smallest islands supported only few seed-eating species utilizing the islands in a fine-grained fashion; none of these birds belonged to the breeding fauna of the islands. With increasing island size and habitat diversity, insectivores and species tied to the human culture were added. On the mainland of Åland, more than half of the birds were observed near human habitations: bird communities wintering in pine forests were richer than in deciduous forests. Proportion of wintering species was greatest in the breeding communities of pine forests. The proportion of wintering species was presumably regulated by the degree of structural change in the habitat between summer and winter, this change being most drastic in deciduous habitats The species observed could be broadly classified into three feeding categories: opportunists ., omnivores and scavengers, dependent on the diverse food items provided by man; food specialists , species éating seeds and berries, ranging widely in the islands and habitats; and insectivorous habitat specialists , inhabiting coniferous habitats, and islands with coniferous (pine) forests. Opportunists, in particular, gel increasing wintering opportunities by the impact of man.