Validation of a landscape‐scale planning tool for cavity‐dependent wildlife

Tree cavities provide important habitat for wildlife. Effective landscape‐scale management of cavity‐dependent wildlife requires an understanding of where cavities occur, but tree cavities can be cryptic and difficult to survey. We assessed whether a landscape‐scale map of mature forest habitat availability, derived from aerial photographs, reflected the relative availability of mature trees and tree cavities. We assessed cavities for their suitability for use by wildlife, and whether the map reflected the availability of such cavities. There were significant differences between map categories in several characteristics of mature trees that can be used to predict cavity abundance (i.e. tree form and diameter at breast height). There were significant differences between map categories in the number of potential cavity bearing trees and potential cavities per tree. However, the index of cavity abundance based on observations made from the ground provided an overestimate of true cavity availability. By climbing a sample of mature trees we showed that only 5.1% of potential tree cavities detected from the ground were suitable for wildlife, and these were found in only 12.5% of the trees sampled. We conclude that management tools developed from remotely sensed data can be useful to guide decision‐making in the conservation management of tree cavities but stress that the errors inherent in these data limit the scale at which such tools can be applied. The rarity of tree cavities suitable for wildlife in our study highlights the need to conserve the tree cavity resource across the landscape, but also the importance of increasing the accuracy of management tools for decision‐making at different scales. Mapping mature forest habitat availability at the landscape scale is a useful first step in managing habitat for cavity‐dependent wildlife, but the potential for overestimating actual cavity abundance in a particular area highlights the need for complementary on‐ground surveys.     

Multi‐scale patterns of forest structure and species composition in relation to climate in northeast China

Though a number of studies have focused on the factors that shape the structure and dynamics of temperate forests, little is known about whether these factors vary with spatial scale. In this study we investigated compositional and structural patterns of forests across three spatial scales (plot, local assemblages and regions) in northeast China and asked whether climatic variables shape these patterns. Using a systematic sampling design, we measured diameter at breast height (DBH) and height of trees, and recorded the abundances, percent of cover and heights of shrubs and herbs in 141 plots from 10 nature reserves. We found that summer temperature accounted for most of the variation in species composition, both within and among forest types. DBH, tree height and total basal area all increased significantly with summer temperature while stem density decreased. The DBH frequency distribution depended strongly on temperature (especially winter temperature) and varied among spatial scales, and it tended to be more left‐skewed as temperature increased. Taking together, our results suggest that a warming climate could lead to an increase in tree growth and the changes in size structure of temperate forests in northeast China. In particular, the proportion of large trees will in all likelihood increase while that of smaller trees will decrease. Shifts in forest structure in a warmed world will undoubtedly influence forest management practices, ecosystem dynamics, and species conservation.     

The role of body size in nest-site selection by secondary cavity-nesting birds in a subtropical Chaco forest

Most bird species that nest in tree cavities globally occur in diverse assemblages in little-studied tropical and subtropical forests which have high rates of habitat loss. Conservation of these communities will require an understanding of how species traits, such as body size, influence nest-site selection. We examined patterns of nest-site selection of secondary cavity-nesting birds at the nest patch, tree and cavity scale, and investigated how these patterns are influenced by body size. Using conditional logistic regression, we compared characteristics of 155 nest tree cavities paired with 155 unused tree cavities in quebracho Schinopsis balansae forests in Chaco National Park, Argentina (2016–2018). The odds of a cavity being used for nesting increased with its depth and height above ground, decreased with entrance size, and were greater for dead trees than live. Small-bodied (13–90 g) species used floor diameters in proportion to availability, but medium- (150–200 g) and large-bodied (400–700 g) species selected cavities with larger floors. Model selection indicated that characteristics at the nest patch scale (canopy cover, tree density) had little effect on nest-site selection when cavity-scale variables were included. Cavity floor diameter, entrance size, cavity height and tree diameter (but not cavity depth) increased with body mass, and larger bird species more often used live trees. Two tree species proved to be key for the community: large and medium-sized birds used almost exclusively large live Schinopsis balansae, whereas small birds used live and dead Prosopis spp. in a proportion greater than its availability. Small birds could be differentiated according to species-specific cavity characteristics, but medium and large species overlapped considerably with one another. Although body mass explained much of the overall variation in tree and cavity characteristics between small and medium/large species, several small-bodied species consistently used cavities outside of the expected characteristics for their body size, suggesting that other natural history traits may play important roles in nest-site selection by small-bodied birds. To retain the full suite of secondary cavity-nesters in species-rich tropical and subtropical forests, it is necessary to conserve a diversity of trees and cavities that meet the full range of nesting requirements of these trait-diverse communities.     

Selection of Nest Trees by Cavity‐nesting Birds in the Neotropical Atlantic Forest

One of the five most important global biodiversity hotspots, the Neotropical Atlantic forest supports a diverse community of birds that nest in tree cavities. Cavity‐nesting birds may be particularly sensitive to forestry and agricultural practices that remove potential nest trees; however, there have been few efforts to determine what constitutes a potential nest tree in Neotropical forests. We aimed to determine the characteristics of trees and cavities used in nesting by excavators (species that excavate their own nest cavity) and secondary cavity‐nesters (species that rely on existing cavities), and to identify the characteristics of trees most likely to contain suitable cavities in the Atlantic forest of Argentina. We used univariate analyses and conditional logistic regression models to compare characteristics of nest trees paired with unused trees found over three breeding seasons (2006–2008). Excavators selected dead or unhealthy trees. Secondary cavity‐nesters primarily selected cavities that were deep and high on the tree, using live and dead cavity‐bearing trees in proportion to their availability. Nonexcavated cavities suitable for birds occurred primarily in live trees. They were most likely to develop in large‐diameter trees, especially grapia Apuleia leiocarpa and trees in co‐dominant or suppressed crown classes. To conserve cavity‐nesting birds of the Atlantic forest, we recommend a combination of policies, economic assistance, environmental education, and technical support for forest managers and small‐scale farmers, to maintain large healthy and unhealthy trees in commercial logging operations and on farms.     

Characteristics of nest cavities of Barrow's Goldeneyes in eastern Canada

ABSTRACT Barrow's Goldeneyes (Bucephala islandica) are secondary cavity nesters found in western North America and, to a lesser extent, in eastern North America. The eastern North American population is concentrated in the province of Québec and totals about 2000 pairs. Characteristics of nest cavities used by Barrow's Goldeneyes have been described in western North America, but no nest cavities have been found in eastern North America. From 2004 to 2008, we searched for nest cavities in the species’ core breeding area in the boreal forests north of the St. Lawrence River. We captured 12 adult females on their breeding grounds and fitted them with transmitters, but none apparently nested so we conducted ground searches in areas near lakes where paired birds were observed. We found 11 cavities, with 10 in dead, decaying trees and one in the dead part of a dying tree. Nine cavities were in white birch (Betula papyrifera) trees. Mean cavity height was 3.5 ± 1.6 (SD) m (range = 1.2–6.6 m) and mean diameter at breast height (DBH) of cavity trees was 37.8 ± 4.7 cm (range = 32.2–47.5 cm). In contrast to the population in western North America, Barrow's Goldeneyes in eastern North America appear to rely on the availability of natural cavities formed in large, decaying trees for nesting. Current forestry regulations in Québec do not promote the retention of either large trees or older forests, reducing the availability of potential nest cavities for Barrow's Goldeneyes and likely threatening their long‐term conservation. Therefore, we recommend that guidelines be developed to promote silvicultural practices aimed at preserving the long‐term availability of large (DBH ≥ 30 cm) decaying trees across the breeding range of Barrow's Goldeneyes.     

西双版纳热带森林树洞丰富度及其分配特点

树洞是森林生态系统的重要结构,对动物多样性的维持起着重要作用。为研究热带森林树洞的数量特征及其分配特点,选取西双版纳20hm2热带森林动态监测样地为研究对象,采用地面观测法调查了样地内所有胸径≥5cm活立木上的树洞。结果表明:(1)该样地森林的树洞密度为108.4个·hm-2,具有树洞的树1725株(占树木总数的6.22%),隶属于208个种;(2)树木个体的树洞数量随径级增加而增加,并在种间变化不均匀;(3)树木出现空心的概率随胸径级的增加而显著增加,并且在树种间存在显著差异;(4)与其他纬度地区森林的树洞密度比较,本研究结果支持了Boyle等(2008)提出的低纬度森林具有较高树洞密度的推测。     

Nest Selection by Cavity-nesting Birds in Subtropical Montane Forests of the Andes: Implications for Sustainable Forest Management

Development of sustainable forestry has been hampered in tropical countries by a scarcity of research on the ecological effects of logging. We focused on cavity-nesting birds, a group known to be sensitive to logging. Cavities used for nesting were not a random subset of all available suitable cavities. Birds selected cavities that were relatively high above the ground, had smaller entrances, and were excavated by woodpeckers. The use of tree species was also not random: Calycophyllum multiflorum , Blepharocalyx gigantea , and Podocarpus parlatorei were disproportionately important. Cavity nests were also more likely to be found in areas with trees with high mean diameter at breast height. This study emphasizes the need to maintain some unlogged forest patches within logging areas and retain certain species of trees. This study has implications for forest management in Argentina, where a new law mandates the sustainable use of forest resources and where many landowners are interested in forest certification.     

Can we effectively stop the expansion of trees on wetlands? Results of a birch removal experiment

The main aim of this study was to determine the effectiveness of removing invasive downy birch in non-forest peatland applying a single cut. We tested whether the removal of birch was positively related to the height of cutting, season of the year and age of the trees. Our study showed that birch trees cut at the higher tested height exhibited a lower survival rate than those cut below or just above the ground level. Furthermore, the winter cut produced more non-sprouting stumps than those recorded after the cut performed in autumn. In our opinion, winter cutting at breast height is the best management strategy for downy birch control. However, we suppose that the effective elimination of birch by applying a single cut is not possible, and the treatment should be repeated in subsequent years or boosted by additional treatments.     

The effects of elk on aspen in the winter range in Rocky Mountain National Park

National parks face problems in managing wild ungulates and their forage resources, including small park sizes and incomplete protection of winter ranges, absence of major predators, and influences from exterior management activities Our study focuses on the effects of elk Cervus elaphus browsing on aspen Populus tremuloides in the elk winter range of Rocky Mountain National Park Elk can prevent successful regeneration of aspen by suckers, and increase the mortality of established trees Here we quantify the effect of elk on aspen in the elk winter range of the park particularly since the adoption of a policy of “natural regulation” in 1968 We collected a stratified-random sample of aspen cohorts in the elk winter range, and also inside two long-term exclosures In each cohort we tallied live and dead trees by diameter, recorded the height and percentage of stem surface covered by bark-stripping, and estimated the date of regeneration of the cohort using increment cores We also tallied live and dead aspen suckers (young trees < 2 5 in tall) by height and the number of live and dead branches on each sucker Aspen outside the exclosures are declining as evidenced by 1) high mortality among established trees, including entire stands that are dead, 2) low density of live suckers, insufficient to regenerate the stand, 3) suckers that are excessively branched and seldom surviving to reach tree height, and 4) almost no cohort regeneration since adoption of natural regulation Exclosures and early photos of the elk winter range suggest these are anomalous conditions Over the last century, aspen cohorts regenerated only when there were fewer than ca 600 elk on the park's elk winter range, far fewer than the present estimated 1600 head Fire suppression, natural succession, fluctuations in beaver populations, climatic fluctuations, and other factors are of no or lesser importance than elk in the regeneration and survival of aspen cohorts on the elk winter range The decline of aspen is a concern when management is focused on the ecosystem scale rather than simply the scale of wildlife and their primary forage resources     

Cavity construction by reintroduced populations of Red‐cockaded Woodpeckers (Dryobates borealis)

Although species reintroduction attempts are now common, monitoring of reintroduction attempts rarely extends beyond initial population establishment. This short timespan likely fails to document long‐term population stability, subtle changes in behavior, and the potentially larger effects that some reintroduced species may have on other species. The Red‐cockaded Woodpecker (RCW; Dryobates borealis) is an important habitat specialist and ecosystem engineer that excavates cavities in living trees. Excavation of natural RCW cavities can take years to complete, but they also persist for many years and are used by many other species. We quantified characteristics of cavity trees excavated by RCWs (n = 44) in two populations that were reintroduced to unoccupied areas more than 10 years earlier. We measured features associated with heartwood rot and used generalized linear mixed effects regression to determine whether these features differed for trees selected for cavity excavation compared with random neighboring trees. We also assessed population trends for cavity‐nesting species that commonly used RCW cavities on one of the sites. Height of first live limb was the only factor distinguishing natural RCW cavity trees from control trees. Four of six cavity‐nesting species monitored increased significantly following RCW establishment. The increases may relate to the many natural and artificial cavities created during the reintroduction effort. Future reintroductions of the RCW should lead to successful natural cavity excavation if sufficiently large trees with smaller live crowns are present. Future efforts may also benefit the broader community of cavity‐nesting birds.     

Temperature characteristics of winter roost-sites for birds and mammals: tree cavities and anthropogenic alternatives

The microclimate of potential roost-sites is likely to be a crucial determinant in the optimal roost-site selection of endotherms, in particular during the winter season of temperate zones. Available roost-sites for birds and mammals in European high trunk orchards are mainly tree cavities, wood stacks and artificial nest boxes. However, little is known about the microclimatic patterns inside cavities and thermal advantages of using these winter roost-sites. Here, we simultaneously investigate the thermal patterns of winter roost-sites in relation to winter ambient temperature and their insulation capacity. While tree cavities and wood stacks strongly buffered the daily cycle of temperature changes, nest boxes showed low buffering capacity. The buffering effect of tree cavities was stronger at extreme ambient temperatures compared to temperatures around zero. Heat sources inside roosts amplified Δ T (i.e., the difference between inside and outside temperatures), particularly in the closed roosts of nest boxes and tree cavities, and less in the open wood stacks with stronger circulation of air. Positive Δ T due to the installation of a heat source increased in cold ambient temperatures. These results suggest that orchard habitats in winter show a spatiotemporal mosaic of sites providing different thermal benefits varying over time and in relation to ambient temperatures. At cold temperatures tree cavities provide significantly higher thermal benefits than nest boxes or wood stacks. Thus, in winter ecology of hole-using endotherms, the availability of tree cavities may be an important characteristic of winter habitat quality.     

Hibernation by tree-roosting bats

In summer, long-eared bats (Nyctophilus spp.) roost under bark and in tree cavities, where they appear to benefit from diurnal heating of roosts. In contrast, hibernation is thought to require a cool stable temperature, suggesting they should prefer thermally insulated tree cavities during winter. To test this prediction, we quantified the winter thermoregulatory physiology and ecology of hibernating tree-roosting bats, Nyctophilus geoffroyi and N. gouldi in the field. Surprisingly, bats in winter continued to roost under exfoliating bark (65%) on the northern, sunny side of trees and in shallow tree cavities (35%). Despite passive re-warming of torpid bats by 10-20 degrees C per day, torpor bouts lasted up to 15 days, although shorter bouts were also common. Arousals occurred more frequently and subsequent activity lasted longer on warmer nights, suggesting occasional winter foraging. We show that, because periodic arousals coincide with maximum roost temperatures, when costs of rewarming and normothermic thermoregulation are minimal, exposure to a daily temperature cycle could largely reduce energy expenditure during hibernation. Our study provides further evidence that models of torpor patterns and energy expenditure from hibernators in cold temperate climates are not directly applicable in milder climates, where prolonged torpor can be interspersed with more frequent arousals and occasional foraging.     

Behavioral thermoregulation of wild Japanese macaques: comparisons between two subpopulations

We studied the behavioral thermoregulation of Japanese macaques in two troops that live in the coniferous (1,000-1,200 m in elevation) and coastal forests (0-200 m in elevation) of Yakushima. Frequency of sunbathing, huddling, and microhabitat selection during inactivity was compared. The difference in mean annual air temperature between the forests was more than 7 degrees C. In both forests, when the weather was clear, macaques spent more time being inactive in the sunshine in winter than in autumn. In winter, they huddled more often when it was clear than when cloudy. Microhabitat selection to stay in the sunshine during winter differed between the two forests. In winter, macaques spent more time inactive in open habitats in the coniferous forest and in the trees in the coastal forest than in autumn, respectively. This difference is related to the lower crown height in the coastal forest and the large open habitats (logged area) available only in the coniferous forest. In winter, skin temperature measured by temperature-sensitive transmitters was 1.32-1.71 degrees C higher when sunbathing, and 0.83-4.75 degrees C higher when huddling than staying in the shade without huddling. In winter, the proportion with which they stayed in the sunshine or huddled in winter did not differ between the two forests, in spite of the difference in air temperature. This suggests that Japanese macaques respond to seasonal changes in air temperature, not the absolute temperature, and that they acclimatize themselves to thermal conditions that require behavioral thermoregulation only during the season when thermoregulation is most costly.     

Nest Site Selection by Western Gray Squirrels at Their Northern Range Terminus

ABSTRACT The western gray squirrel (Sciurus griseus) in Washington, USA, is limited to 3 disjunct areas and is a state threatened species. Information is lacking for the North Cascades population, which is the northernmost population for the species. Squirrels in this population exist without oaks (Quercus spp.) that provide forage and cavities for maternal nests elsewhere in their range. During May 2003 to August 2005, we studied selection of nest sites and nest trees by 18 radiocollared squirrels in Okanogan County, Washington. Without oak cavities, females reared their young in dreys. General nest-tree characteristics were similar to characteristics of western gray squirrel nest trees in Southeastern Cascades: relatively tall ponderosa pines (Pinus ponderosa) ≥ 40 cm diameter at breast height. Results from conditional logistic models determined that the odds of a squirrel selecting a tree for nesting increased with greater diameter at breast height and with infection by dwarf mistletoe (Arceuthobium spp.). Nest sites with high selection probability by squirrels had greater basal area and number of tree species than available unselected sites. Retention of forest patches that include a mix of conifer species or conifer and deciduous trees and moderate to high basal area could promote nesting opportunities, connectivity for arboreal travel, as well as abundance and diversity of hypogeous fungi. Experiments to test the efficacy of retaining untreated patches of varying size (including trees infected with mistletoe) on nesting by western gray squirrels within stands managed for fire suppression and forest health would provide important information about the effects of forest fuel management on arboreal wildlife.     

Site occupancy and cavity use by the northern flying squirrel in the boreal forest

Tree cavities are used as shelter and breeding nests by numerous avian and mammalian species. In cold environments, tree cavities are often proposed as the best winter nest choice because of the superior protection they offer from precipitation, wind, cold temperatures, and predators. As such, they represent a critical resource, which has the potential to limit population size of non-excavating species. We assessed factors affecting site occupancy in the boreal forest by northern flying squirrels, a secondary user of tree cavities, and to identify which nest type is preferred during the colder days of the autumn–winter period. We trapped flying squirrels twice in 59 aspen-dominated stands in the autumn period using low- (1.5 m above ground-level) and high-mounted (4 m) traps to determine site occupancy. A total of 85 individuals were captured on 2,880 trap-nights. During the winter period, we radio-tracked 26 individuals to 87 diurnal nests in 220 locations. None of the habitat variables considered (cavity availability, woody debris, and lateral cover) explained site occupancy. Detectability decreased with precipitation, and was lower using high traps than low traps. Both females and males used tree cavities (26%), external nests (39%), and ground nests (35%). In cold weather, females preferred ground nests, whereas males preferred external nests. Our results do not support the hypothesis that tree cavities represent a limiting factor to northern flying squirrels in cold environments. Instead, this species seems to be a generalist and is opportunistic, using a variety of nest types. Nevertheless, practices ensuring the persistence of large diameter live cavity trees, providing better insulative properties, are likely to increase the relative use of tree cavities as nest sites by northern flying squirrels. © 2011 The Wildlife Society.     

Variables associated with the occurrence of Ips beetles,red turpentine beetle and wood borers in live and dead ponderosa pines with post‐fire injury

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Water availability as driver of birch mortality in Hustai National Park,Mongolia

Trees in the forest-steppe ecotones face stress due to reduced water availability as a consequence of more extreme seasonal fluctuations in precipitation and temperature. Together with browsing pressure this can hinder tree growth, tree regeneration and competition between trees and grasses. We studied the impact of both stress factors on the mortality of birch trees in two forest sites at Hustai National Park, Mongolia, by applying tree-ring research to determine growth-limiting factors and assessing browsing pressure on young and adult birch. We expected warm and dry summer conditions as main growth limiting factor. Moreover, we expected a positive relation between deer density and tree mortality with browsing mainly affecting smaller trees with a low diameter at breast height (DBH). We found that the growth in both birch populations is mainly driven by winter precipitation and – to a lesser extent – negatively affected by high summer temperature. This suggests that water availability as defined by soil moisture, especially at the beginning of the growing season is crucial for birch growth in our study area. For mortality we found significant differences between both populations, but no significant relationship with deer density. In plots with high mortality rates mean tree height of the remaining living trees decreased. These results suggest that under expected climate change with declining annual precipitation rates, the birch forest of Hustai National Park is converting into a steppe ecosystem, like described for other forest ecosystems in this ecotone.     

Forest community survey and the structural characteristics of forests in China

Most of the world’s terrestrial biome types can be found in China. To systematically investigate species composition and structure of China’s forest communities, we launched a long‐term project consisting forest vegetation surveys across China’s mountains in the mid 1990s. Over the study period, we have conducted vegetation surveys for 65 mountains and collected vegetation data from about 1500 forest plots, using consistent sampling protocols. In this paper we first introduce the aims, protocols, and major research themes of the project, and then describe the major characteristics of forest communities and their geographic patterns and climatic controls. As latitude increased, diameter at breast height (DBH) and height of trees increased, while individual density of trees and woody species richness decreased. Total basal area (TBA) of trees and species richness of herbs did not vary with latitude. Contemporary climate seems to drive these patterns: temperature was the leading factor for DBH, precipitation was most important for tree height and individual density, actual evapotranspiration (a surrogate of productivity) determined woody (trees and shrubs) species richness, and rainfall was the major controller of the herb species richness. The species–abundance relationship showed that species dominance (measured by the number of individuals per species) declined significantly from boreal forests to evergreen broadleaf forests from north to south. Our results are in line with the idea that productivity drives woody species richness. Similarly, we find that biomass (measured as TBA) is invariant along the environmental gradients. However, individual density varies dramatically, in contrast to the assumptions underlying the metabolic theory of ecology.     

How do Araucaria angustifolia trees grow in overstocked stands?

The objective of this study was to evaluate growth along the stem of Araucaria angustifolia (Bertol.) Kuntze trees competing in overstocked stands, in order to identify periods when growth and trunk shape are differentiated during the trees' lifespan. The research was carried out in a planted forest of Araucaria angustifolia established in 1946 in the Açungui National Forest in Campo Largo, Paraná, Brazil, when these trees were 65 years old. One thinning was recorded, at some time between 1970 and 1980. Forty-six trees were selected and divided into three development classes (DC) at 65 years of age; these classes considered diameter at 1.30 m (breast height, dbh) with a range of 20 cm (from 10 cm to 70 cm). In addition to dbh, total tree height, and crown height and diameter were measured in the field. From each tree, 14 disks were removed to analyze growth rings and confirm the age of the stand. Some trees in the smallest DC (10 ≤ dbh < 30 cm) were the product of natural regeneration (younger trees that grew after the initial planting). In 63% of the trees, at least one growth ring was missing at breast height. Missing rings at breast height were more common in trees with smaller dbh and crown diameters. The need for more growing space was observed at different periods during the studied lifespan of the trees from three DCs. It resulted in changes in stems shape from conical to cylindrical. Different growth patterns could be observed during the lives of some trees as they outgrew their competition.     

Decaying trees improve nesting opportunities for cavity‐nesting birds in temperate and boreal forests: A meta‐analysis and implications for retention forestry

Many studies have dealt with the habitat requirements of cavity‐nesting birds, but there is no meta‐analysis on the subject and individual study results remain vague or contradictory. We conducted a meta‐analysis to increase the available evidence for nest‐site selection of cavity‐nesting birds. Literature was searched in Web of Science and Google Scholar and included studies that provide data on the habitat requirements of cavity‐nesting birds in temperate and boreal forests of varying naturalness. To compare nest and non‐nest‐tree characteristics, the following data were collected from the literature: diameter at breast height (DBH) and its standard deviation (SD), sample size of trees with and without active nest, amount of nest and available trees described as dead or with a broken crown, and amount of nest and available trees that were lacking these characteristics. Further collected data included bird species nesting in the cavities and nest‐building type (nonexcavator/excavator), forest type (coniferous/deciduous/mixed), biome (temperate/boreal), and naturalness (managed/natural). From these data, three effect sizes were calculated that describe potential nest trees in terms of DBH, vital status (dead/alive), and crown status (broken/intact). These tree characteristics can be easily recognized by foresters. The results show that on average large‐diameter trees, dead trees, and trees with broken crowns were selected for nesting. The magnitude of this effect varied depending primarily on bird species and the explanatory variables forest type and naturalness. Biome had lowest influence (indicated by ΔAIC). We conclude that diameter at breast height, vitality, and crown status can be used as tree characteristics for the selection of trees that should be retained in selectively harvested forests.