Flexible responses to stage‐specific offspring threats

When caring for their young, parents must compensate for threats to offspring survival in a manner that maximizes their lifetime reproductive success. In birds, parents respond to offspring threats by altering reproductive strategies throughout the breeding attempt. Because altered reproductive strategies are costly, when threats to offspring are limited, parents should exhibit a limited response. However, it is unclear if response to offspring threat is the result of an integrated set of correlated changes throughout the breeding attempt or if responses are a flexible set of dissociable changes that are stage‐specific. We test these hypotheses in a system where house wrens (Troglodytes aedon) compete for nesting cavities with Carolina chickadees (Poecile carolinensis) by usurping and destroying their nests during the early stage of the breeding attempt (the egg stage). Due to the specificity of the house wren threat, we can test whether parental responses to an offspring threat show flexibility and stage specificity or if parental strategies are an integrated and persistent response. We monitored nests in a natural population to compare life history traits of chickadees nesting in boxes that were in the presence of house wrens to chickadees nesting in boxes that did not overlap with house wrens. Carolina chickadees that nested near house wrens laid significantly smaller clutch sizes (early change in reproductive strategy) but did not alter nestling provisioning or nestling stage length (late change in reproductive strategy), suggesting that chickadees respond in a flexible and stage‐specific manner to the threat of house wrens. By responding only when a threat is highest, parents minimize the cost of antithreat responses. Our study suggests that parents can respond in subtle and nuanced ways to offspring threats in the environment and specifically alter reproductive behaviors at the appropriate stage.     

Nest cavity orientation in black‐capped chickadees Poecile atricapillus: do the acoustic properties of cavities influence sound reception in the nest and extra‐pair matings?

Birds that nest in cavities may regulate nest microclimate by orienting their nest entrance relative to the sun or prevailing winds. Alternatively, birds may orient their nest entrance relative to conspecific individuals around them, especially if the acoustic properties of cavities permit nesting birds to better hear individuals in front of their nest. We measured the cavity entrance orientation of 132 nests and 234 excavations in a colour‐banded population of black‐capped chickadees Poecile atricapillus for which the reproductive behaviour of nesting females was known. Most chickadees excavated cavities in rotten birch Betula papyrifera, aspen Populus tremuloides and maple Acer saccharum. Nest cavities showed random compass orientation around 360° demonstrating that chickadees do not orient their cavities relative to the sun or prevailing winds. We also presented chickadees with nest boxes arranged in groups of four, oriented at 90° intervals around the same tree. Nests constructed in these nest box quartets also showed random compass orientation. To test the acoustic properties of nest cavities, we conducted a sound transmission experiment using a microphone mounted inside a chickadee nest. Re‐recorded songs demonstrate that chickadee nest cavities have directional acoustic properties; songs recorded with the cavity entrance oriented towards the loudspeaker were louder than songs recorded with the cavity entrance oriented away from the loudspeaker. Thus, female chickadees, who roost inside their nest cavity in the early morning during their fertile period, should be better able to hear males singing the dawn chorus in front of their nest cavity. Using GIS analyses we tested for angular‐angular correlation between actual nest cavity orientation and the azimuth from the nest tree to the territories and nest cavities of nearby males. In general, nest cavity entrances showed no angular‐angular correlation with neighbourhood territory features. However, among birds who followed a mixed reproductive strategy and nested in the soft wood of birch and aspen trees, nest cavity entrances were oriented towards their extra‐pair partners. We conclude that nest cavity orientation in birds may be influenced by both ecological and social factors.     

贵州花溪大学城破碎化林地鸟类多样性与嵌套分布格局

为揭示城镇化进程中生境破碎化对鸟类多样性及分布格局的影响, 本研究于2017-2019年每年的4-8月使用样线法对贵州花溪大学城26块破碎化林地(面积介于0.3-290.4 ha)中的鸟类群落进行了10次调查。共记录到鸟类78种, 隶属于11目37科。其中, 东洋界物种数占56.4%, 古北界物种数占32.1%, 广布种占11.5%; 有中国特有种1种。剔除高空飞行、非森林鸟类及偶然出现物种后, 不同斑块中的鸟类物种数介于12-55之间, 平均每个斑块有23.2 ± 10.5种。线性回归分析显示, 鸟类物种丰富度与林地斑块的面积有显著相关性, 斑块面积越小, 鸟类物种丰富度越低; 斑块隔离度对物种丰富度没有显著影响。基于物种多度分布矩阵的WNODF (weighted nestedness metric based on overlap and decreasing fill)嵌套分析显示, 不同斑块中鸟类群落呈现出反嵌套结构。小斑块中鸟类物种丰富度较低可能与植物丰富度较低、食物资源稀缺和繁育条件不足有关, 但短距离的隔离对鸟类迁入或扩散影响有限。环境过滤效应、种间竞争或优先效应可能导致不同斑块间存在较大的物种组成差异, 从而导致反嵌套格局。因此, 本研究建议在城市规划建设中应注重维持栖息地的完整性, 对不同面积大小的破碎化斑块都应加以保护。     

A comparison of life‐history and parental care in temperate and tropical wrens

Comparing closely related species that live in different environments is a powerful way to understand selective pressures that influence life‐history evolution. We examined a suite of life‐history traits and parental care in neotropical buff‐breasted wrens Cantorchilus leucotis and north‐temperate Carolina wrens Thryothorus ludovicianus (Family Troglodytidae), to test hypotheses about life‐history evolution. As expected, buff‐breasted wrens exhibited smaller clutch sizes and higher annual adult survival than Carolina wrens. We found minimal support for the nest predation hypothesis, as nest survival and age‐corrected provisioning rates to whole broods were similar between species, and number of breeding attempts and breeding season length were greater in temperate wrens. Critical predictions of the food limitation hypothesis were not supported; in particular age‐corrected provisioning rates per nestling were higher in the tropical than temperate species. The adult survival and offspring quality hypothesis garnered the most support, as buff‐breasted wrens exhibited greater age‐corrected provisioning rates per nestling, a longer nestling period, longer re‐nesting intervals following nest success, and lower annual fecundity than Carolina wrens. Despite similarly prolonged breeding seasons, reproductive strategies differ between species with buff‐breasted wrens investing considerably in single broods to optimize first‐year survival and Carolina wrens investing in multiple small broods to optimize annual fecundity.     

Patterns of extra‐pair paternity in mountain chickadees

Extra‐pair paternity (EPP) is common in chickadees and often attributed to the good genes hypothesis. Females generally seek dominant males, who are typically larger, older and sing at higher rates than subordinate males, as extra‐pair sires. In other songbird species, habitat quality and urbanization have been found to influence EPP. Mountain chickadees commonly inhabit suburban habitat, and previous research on our population has shown urbanization may provide benefits to these adaptable songbirds. Here, we ask how individual condition and urbanization influence rates of EPP in mountain chickadees. Over three breeding seasons, we monitored mountain chickadee nests in urban and rural habitat, and determined parentage by genotyping adults and nestlings at six microsatellite loci. Extra‐pair paternity is common in mountain chickadees, with extra‐pair offspring (EPO) in 43.2% of nests and accounting for 17.9% of offspring. We found tenuous support for the good genes hypothesis with females tending to engage in EPCs with older males. However, we did not find an influence of male or female condition on the proportion of EPO in a nest. In addition, we did not find a significant effect of habitat on EPP rates, suggesting the impacts of urbanization on mountain chickadee reproduction may not extend to altering extra‐pair behaviour.     

Urban living alters moult dynamics in a passerine

Urbanization and habitat fragmentation can alter the timing of life history events, potentially leading to phenological mismatches, carryover effects, and fitness costs. Whereas urbanization and fragmentation are known to alter important aspects of breeding in many bird species, little is known about the effects of urbanization and habitat fragmentation on moult. To investigate the effects of urbanization and fragmentation on the annual moult, we compared the moult dynamics (onset, duration, and intensity) of urban, fragmented forest, and contiguous forest populations of the Carolina chickadee, a North American resident passerine that moults once per year immediately following the breeding season. Over three years, moult dynamics were similar in contiguous and fragmented forest populations, but wing moult started significantly earlier, and onset of moult varied less among years, in urban chickadees than in forest chickadees (fragmented and contiguous habitats pooled). Duration of wing moult did not differ between urban and forest populations, but urban birds moulted significantly fewer feathers simultaneously during peak moult, suggesting that individual feathers grew more rapidly. Our results show that urban living alters critical aspects of moult dynamics in a widespread songbird. Given the importance of moult dynamics for fitness during subsequent life history stages, and the large number of songbird species inhabiting urban areas, these results reveal previously unrecognized and potentially costly carryover effects of urban living on songbirds.     

Chick-a-dee call variation in Carolina chickadees and recruiting flockmates to food

The "chick-a-dee" call of many Paridae species (titmice, tits,and chickadees) is structurally complex and functions in socialcohesion. Studies with different Parid species suggest thatvariation in the note composition of calls relates to a widevariety of contexts. An earlier study with Carolina chickadees(Poecile carolinensis), the focal species of the present study,found that receivers responded differently to playback callsdiffering in note composition in feeding contexts. Here, weaddressed whether signalers actually produce calls differingin note composition in feeding contexts and whether those callsmight serve a recruitment function. In a first study, we foundthat the first chickadee to take seed from a feeding stationproduced calls with a greater number of D notes before the secondchickadee arrived to take seed, compared with after the secondchickadee arrived to take seed. This suggests that calls witha large number of D notes might serve a general recruitmentfunction. We tested this idea in a second study, using playbacksof calls containing a large number of D notes or a small numberof D notes at different sites. We found that the latency fora first chickadee to come into a site and take seed was shorterfor playback variants containing a large number of D notes.Thus, in Carolina chickadees, chick-a-dee calls containing alarge number of D notes may function to recruit other flockmembers to a discovered food source.     

Greater hippocampal neuronal recruitment in food-storing than in non-food-storing birds

Previous research has shown heightened recruitment of new neurons to the chickadee hippocampus in the fall. The present study was conducted to determine whether heightened fall recruitment is associated with the seasonal onset of food-storing by comparing neurogenesis in chickadees and a non-food-storing species, the house sparrow. Chickadees and house sparrows were captured in the wild in fall and spring and received multiple injections of the cell birth marker bromodeoxyuridine (BrdU). Birds were held in captivity and the level of hippocampal neuron recruitment was assessed after 6 weeks. Chickadees showed significantly more hippocampal neuronal recruitment than house sparrows. We found no seasonal differences in hippocampal neuronal recruitment in either species. In chickadees and in house sparrows, one-third of new cells labeled for BrdU also expressed the mature neuronal protein, NeuN. In a region adjacent to the hippocampus, the hyperpallium apicale, we observed no significant differences in neuronal recruitment between species or between seasons. Hippocampal volume and total neuron number both were greater in spring than in fall in chickadees, but no seasonal differences were observed in house sparrows. Enhanced neuronal recruitment in the hippocampus of food-storing chickadees suggests a degree of neurogenic specialization that may be associated with the spatial memory requirements of food-storing behavior.     

Genetic effects of habitat fragmentation in Viola pubescens (Violaceae), a perennial herb with chasmogamous and cleistogamous flowers

The reproductive biology of a plant species is important in the response of populations to habitat fragmentation, especially if plant-pollinator interactions are disrupted. The genetic effects of forest fragmentation were examined in the common understorey herb Viola pubescens, a species that produces self-pollinated cleistogamous (CL) flowers and potentially outcrossing chasmogamous (CH) flowers. Using allozymes, we measured genetic variation in different sized populations. These were located in woodlots of various sizes (0.5-40.5 ha) and distances from one another (0.3-46 km) within the agricultural landscape of central Ohio in the Midwestern United States. Changes in forest cover of each woodlot within the past 180 years were determined from historical sources and aerial photographs. Woodlot and population sizes were significantly and positively correlated with measures of genetic variation (A, P, HO and HE), with variation highest in populations in the largest woodlot population and lowest in the smallest woodlot population. Most large woodlots resulted from fluctuations in forest cover over the past 60 years, while smaller fragments remained the same size. Overall, populations in Crawford County were genetically differentiated from one another (theta = 0.34), but there was no relationship between genetic and geographical distance. Preliminary evidence for a single year indicated a high rate of outcrossing in most populations. Despite the CH/CL reproductive advantage and apparent outcrossing, populations of V. pubescens in small woodlots remain susceptible to potentially detrimental effects of fragmentation such as genetic drift and reduced levels of genetic variation.     

American Black Duck Nesting Ecology in North Carolina

North Carolina, USA, is the southernmost extent of the American black duck's (Anas rubripes) breeding range; however, little is known about their nesting ecology in this region. We located and monitored 140 nesting black ducks over 2 years (2017–2018) to quantify preferred nesting habitat and assess nesting productivity within coastal North Carolina. We located nests in brackish marshes (75%) and man-made dredge spoil islands (25%) at a density of 1 nest/22 ha. Black ducks selected high marsh and nested an average of 21.81 m from open water at a mean elevation of 1.36 m. In preferred nesting habitat, visual obstruction readings were 0.50 m with a maximum mean vegetation height of 0.81 m and land cover consisted largely of grasses (84.6%). Apparent nest success rates varied from 31% (2017) to 63% (2018) across years. The majority (72.2%) of variability in nest success was best predicted by nest location (mainland marsh, natural island, or spoil island), vegetation density, maximum vegetation height, and year. Management for breeding black ducks in coastal North Carolina should focus on promoting selected nesting habitat and reducing nest predators. Prescribed burns, used to set back succession on spoil islands and in brackish marshes should be conducted in the winter or in the early growing season not to exceed the twenty-fifth quantile date of black duck nest initiation (2 Apr). © 2021 The Wildlife Society.     

Experimental test of nest-site limitation in mature mixed forests of central British Columbia,Canada

Nest-site availability limits cavity-using populations in many harvested forests; however, little is known about the extent of nest-site limitation in mature forests with a full complement of excavator species and intact processes of cavity creation and loss. To examine the role of nest-site availability in limiting cavity-using populations in mature mixed conifer forests in central British Columbia, Canada, we conducted an 11-year before-after control-impact experiment in which we increased nest-site availability via nest box addition. Our 7 sites (3 treatments, 4 controls) had low cavity densities (<2/ha) prior to treatment and cavity occupation rates were also low (<10%/yr), which is a relationship often cited in the literature as evidence of non-limitation in cavity-nesting populations. Following nest box addition at our treatment sites, which tripled the availability of cavities, total density of bird and mammal nests more than tripled. Density of mountain chickadee (Poecile gambeli) nests increased 9-fold on treatment sites and returned to pre-treatment levels following box removal, suggesting that chickadee populations were limited by cavity availability at our study sites. Density of red squirrel (Tamiasciurus hudsonicus) and northern flying squirrel (Glaucomys sabrinus) nests and roosts also increased significantly at treatment sites following box addition and declined following box removal. We noted little change in chickadee or squirrel nest density at control sites monitored concurrently. Squirrels preferred large-sized over small-sized boxes, and significantly enlarged the entrance areas of small boxes by chewing, suggesting that there may have been a shortage of suitable nest and roost sites for them in our study area. We contend that low cavity occupancy rates may not accurately reflect nest-site availability for cavity nesters in mature forests, and that cavity size may influence the true availability of cavities on the landscape. © 2011 The Wildlife Society.     

Partial host fidelity in nest selection by the shiny cowbird (Molothrus bonariensis), a highly generalist avian brood parasite

Obligate avian brood parasites can be host specialists or host generalists. In turn, individual females within generalist brood parasites may themselves be host specialists or generalists. The shiny cowbird Molothrus bonariensis is an extreme generalist, but little is known about individual female host fidelity. We examined variation in mitochondrial control region sequences from cowbird chicks found in nests of four common Argentinean hosts. Haplotype frequency distributions differed among cowbird chicks from nests of these hosts, primarily because eggs laid in nests of house wrens Troglodytes aedon differed genetically from those laid in nests of the other three hosts (chalk-browed mockingbird Mimus saturninus, brown-and-yellow marshbird Pseudoleistes virescens, and rufous-collared sparrow Zonotrichia capensis). These differences in a maternally inherited marker indicate the presence of a nonrandom laying behaviour in the females of this otherwise generalist brood parasite, which may be guided by choice for nest type, as house wrens nest in cavities whereas the other three species are open cup nesters.     

Conspecific olfactory preferences and interspecific divergence in odor cues in a chickadee hybrid zone

Understanding how mating cues promote reproductive isolation upon secondary contact is important in describing the speciation process in animals. Divergent chemical cues have been shown to act in reproductive isolation across many animal taxa. However, such cues have been overlooked in avian speciation, particularly in passerines, in favor of more traditional signals such as song and plumage. Here, we aim to test the potential for odor to act as a mate choice cue, and therefore contribute to premating reproductive isolation between the black‐capped (Poecile atricapillus) and Carolina chickadee (P. carolinensis) in eastern Pennsylvania hybrid zone populations. Using gas chromatography–mass spectrometry, we document significant species differences in uropygial gland oil chemistry, especially in the ratio of ester to nonester compounds. We also show significant preferences for conspecific over heterospecific odor cues in wild chickadees using a Y‐maze design. Our results suggest that odor may be an overlooked but important mating cue in these chickadees, potentially promoting premating reproductive isolation. We further discuss several promising avenues for future research in songbird olfactory communication and speciation.     

Forest size and avian diversity in New Jersey woodlots with some land use implications

Summary The effect of forest size on avian diversity was studied in oak forest patches in rural New Jersey. Number of bird species continued to increase significantly in old oak woods up through 40 ha. This was due to the progressive addition of carnivorous species with increasing minimum forest size requirements. One large woodlot had more species than the same area subdivided into smaller woodlots. To maintain maximum regional diversity more than three large forests are required. Primary land use priority should be to protect large forests. Second priority should be to maintain a high density of small woodlots.     

Variation in Note Composition of Chick‐a‐dee Calls is Associated with Signaler Flight in Carolina Chickadees,Poecile carolinensis

Chick‐a‐dee calls are used in a wide range of social contexts in Poecile (chickadee) species. These calls comprise a number of distinct note types. Earlier naturalistic observational studies suggested that the ‘C’ note type was used frequently in calls of Carolina chickadees (P. carolinensis) in the context of flight. We conducted three field studies with Carolina chickadees to test in more experimentally manipulative ways whether Chick‐a‐dee calls with more ‘C’ notes were associated with flight. The three studies differed in how they elicited flight behavior from chickadees, as well as in the likely arousal levels experienced by the birds. First, we captured chickadees and released them, recording any calls they produced in flight or when later perched after escape. Second, we approached chickadees that were foraging near the ground in field settings and recorded any calls they produced when perched compared to when they were in flight. Third, from a distant blind, we recorded chickadees flying to and from feeding stations, in which the closest perching substrate/cover was at least 2 m away from the feeding station, thus requiring flight. In all three studies, calls contained more ‘C’ notes when birds were in flight compared to when they were perched. This work expands our understanding of variation in note composition of chick‐a‐dee calls beyond the contexts of food and predator detection to the context of movement. Studies are now needed to test whether such variation in chick‐a‐dee calls brings about group cohesion.     

Edge effects on ground beetles at the woodlot–field interface are short‐range and asymmetrical

• 1 Boundaries between woodlots and agricultural habitats are numerous in temperate agricultural landscapes and influence ecological processes in both woodlots and agricultural habitats.
• 2 We aimed to determine how far the species assemblage of ground beetles in woodlot and open habitats was influenced by the presence of the woodlot–field boundary.
• 3 We studied the distribution of ground beetles on both sides of the boundaries of four woodlots along transects of pitfall traps (n = 140). The depth of edge influence (i.e. the distance from the boundary at which the presence of the boundary has no more significant influence) on the species assemblage of ground beetles in each woodlot and in each agricultural habitat was determined with nonlinear canonical analysis of principal coordinates, an ordination method that is followed by nonlinear regression of the principal coordinates on distance from the boundary.
• 4 The depth of edge influence on the species assemblages of ground beetles was asymmetrical relative to the boundary: it was generally higher and had higher variability in open habitats (14.4 ± 12.3 m) than in woodlots (4.9 ± 2.3 m). Species assemblages of ground beetles in edges were a mix between both adjacent species assemblages. Edge effects in woodlots were deeper in the woodlots exhibiting a deeper penetration of open habitat species. Symmetrically, edge effects in open habitat were deeper in the open habitats with a deeper diffusion of forest species into the open habitat.
• 5 Forest ground beetles were not threatened by edge effects. Rather, edge effects are likely to benefit agriculture, mostly through the dispersal of predatory forest species into agricultural fields.

     

Landscape-dependent distribution of northern forest birds in winter

We evaluated the effects of landscape structure, along a broad gradient of deforestation (forest cover 8–88%, 500-m radius), on the spatial distribution of forest birds exposed to winter climatic conditions, in Quebec, Canada. Concurrently, we conducted an experiment to determine if these effects would persist if an unlimited source of energy, provided by food-supplementation, became available. We analyzed these effects at the population level, using count data of black-capped chickadees Poecile atricapillus , but also at the community level, referring to species richness. In one of the two years of the study, before food-supplementation began (November), "forest integrity" (a composite of forest cover and edge density) was positively associated with chickadee abundance and species richness. Each year, forest integrity was also positively associated with chickadee abundance and species richness in landscapes that were supplemented (December–February). However, in control landscapes, during the food-supplementation period, chickadee abundance and species richness tended to decrease with an increase in forest integrity. We argue that the more forested control landscapes facilitated winter emigration of juveniles and transient birds. Conversely, our results further suggest that, in the highly deforested and fragmented control landscapes, birds became "gap-locked" when rigorous winter climatic conditions exacerbated already existing movement constraints.     

上海闵行区园林鸟类群落嵌套结构

城市中的园林绿地呈现斑块状分布,其栖息地特征与岛屿栖息地相似。2008年11月至2009年10月,对上海市闵行区内的7块城市绿地进行调查,记录雀形目鸟类的分布情况,并运用Nestedness temperature calculator软件,检验其群落结构是否符合嵌套结构。运用Arc GIS软件分析该地区的卫星图片,收集7块样地的面积、绿地盖度、水源距离和人为干扰程度等数据,结合实地调查所得到的数据,分析这一嵌套结构的形成原因和影响因素。结果显示：上海市闵行区城市绿地中的雀形目鸟类分布是显著的嵌套结构,园林面积、绿地面积和水源情况都对其嵌套结构有显著影响。但是与真正岛屿上存在的群落分布嵌套结构不同,人为干扰程度对这一结构也有非常明显的影响。基于上述结果可以看出,影响上海市园林鸟类的群落嵌套结构的主要原因是栖息地的结构和人为干扰程度。因此,建议在规划和建设城市公园和绿地时,应该偏重于面积较大,植被盖度和丰富度高,结构合理的园林,并且尽量减少人为干扰。     

Frequency sensitivity in the auditory periphery of male and female black-capped chickadees (Poecile atricapillus)

The black-capped chickadee is a songbird that has been used extensively as a model of animal communication in field and laboratory settings. Although many studies have focused on the complex call and song systems of the black-capped chickadee, relatively fewer studies have focused on chickadee audition. However, we do know from behavioral and molecular work that chickadees (and auditory processing areas in their brains) discriminate between artificially generated tones, between conspecific and heterospecific vocalizations, and among different types of conspecific vocalizations. In this paper we investigate peripheral auditory processing of frequency in the black-capped chickadee and the potential influence of sex on frequency sensitivity using a technique called auditory evoked potentials. We found that male and female black-capped chickadees did not differ in any measure of frequency sensitivity. Both sexes had the greatest sensitivity to frequencies between 2 and 4 kHz. This range of frequencies is well represented in black-capped chickadee song, partially supporting the idea that sender and receiver coevolve. Finally, we suggest that the call and song system of North American parids make them an ideal taxonomic group for comparative work exploring the relationship between call systems and the evolution of auditory processing.     

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.