Use of remnant boreal forest habitats by saproxylic beetle assemblages in even-aged managed landscapes

Saproxylic Coleoptera are diverse insects that depend on dead wood in some or all of their life stages. In even-aged boreal forest management, remnant habitats left as strips and patches contain most of the dead wood available in managed landscapes and are expected to act as refuges for mature forest species during the regeneration phase. However, use of remnant habitats by the saproxylic fauna has rarely been investigated. Our objective was to characterize the saproxylic beetle assemblages using clearcuts and forest remnants in western Québec, Canada, and to explore the effects of forest remnant stand characteristics on saproxylic beetle assemblages. We sampled both beetle adults and larvae, using Lindgren funnels and snag dissection, in five habitat locations (clearcuts, forest interiors of large patches, edges of large patches, small patches and cut-block separators) from three distinct landscapes. Adult saproxylic beetles (all feeding guilds combined) had significantly higher species richness and catch rates in small patches compared to forest interiors of large patches; the phloeophagous/xylophagous group had significantly higher species richness only. Small patches, cut-block separators and edges of large patches also had the highest snag density and basal area, increasing habitat for many saproxylic beetles. No significant differences in density of saproxylic larvae were found between habitat patches, but snag dissection nevertheless suggests that snags in forest remnants are used by comparable densities of insects. Saproxylic beetles appear to readily use habitat remnants in even-aged managed landscapes suggesting that forest remnants can insure the local persistence of these species, at least in the timeframe investigated in our study.     

The influence of forest management history on the integrity of the saproxylic beetle fauna in an Australian lowland tropical rainforest

Forest management in temperate and boreal regions is often based on a strong foundation of applied ecological research. Increasingly, this has allowed the needs of saproxylic (dead wood associated) insects to be addressed. However, there has been very little equivalent research in tropical forests, where saproxylic insect faunas are likely to be much richer and where forestry is usually subject to weaker environmental controls. This study compares the saproxylic beetle fauna of old-growth, selectively logged and regrowth rainforest in the Daintree lowlands of northeastern Queensland, Australia. Old-growth levels of abundance, species richness, assemblage composition and guild structure were not maintained in logged and regrowth forest, suggesting that intact assemblages may not survive in the long-term in managed tropical rainforest. However, retaining a continuous supply of commercially overmature trees in the managed stand may prevent a repeat of the widespread extinctions of saproxylic insects witnessed in temperate and boreal forest regions.     

Habitat suitability models of saproxylic red-listed boreal forest species in long-term matrix management: Cost-effective measures for multi-species conservation

Understanding of how a large landscape or network of conservation areas and habitats of red-listed species change in time is an important topic when addressing the temporal interplay between protected areas and matrix. We developed models of habitat suitability indices (HSI) for saproxylic red-listed invertebrate and fungal species, accounting for roughly 70% of all red-listed boreal forest species of the study area in eastern Finland. By using a forestry planning program that incorporates various optimisation methods we analysed trade-offs between timber production and amount of habitats of saproxylic red-listed species within a 60-year period. We also produced production possibility frontiers that show how to increase quality of the matrix with least costs. Moreover, we analysed how habitat suitability criteria used in optimisations affect the area of different habitat quality classes.

Our analysis shows that by adopting HSI models in long-term matrix management, it is possible to increase habitats for several red-listed species without substantial losses in timber production. The increase in habitat area is achieved mainly by decreasing the area that is thinned compared to intensive timber production plan. In the long term, this seems to be a novel cost-effective method to increase the quality of the matrix for red-listed saproxylic species. However, the selected optimisation method and the criteria or specification of the management objective for red-listed forest species have a strong effect on results when HSI models are used in conservation planning. Therefore any practical application must be performed with great care.     

Modelling habitat and spatial distribution of an endangered longhorn beetle - A case study for saproxylic insect conservation

Saproxylic insects are characterised by their exceptional diversity and high proportion of threatened species. No recent studies have demonstrated the validity of habitat suitability analysis for scientifically based habitat management for these species. We studied the habitat requirements of the endangered longhorn beetle Cerambyx cerdo, a species with a supposed keystone function for the saproxylic insect community living on oaks. We used species distribution modelling based on datasets from Central Europe to understand the species-habitat relationships and to find the environmental variables responsible for habitat selection of C. cerdo. Our results show that the most important parameters, insolation, presence of oak sap, bark depth and the distance from the next colonised tree, are able to predict the presence of C. cerdo very well. A spatial validation procedure revealed very similar predictive power, indicating the general validity of our model. Tree-level parameters were shown to have a stronger effect on the occurrence probability than landscape-level predictors. To improve the tree-level conditions (e.g. insolation on the trunk) habitat management in the form of semi-open pasture landscapes is recommended from which many other taxa will also draw considerable benefit. The provision of such conditions over decades is the essential key in the conservation of this longhorn beetle species. The success of the European network of conservation areas “Natura 2000” heavily depends on broad biological knowledge of the designated protected species. The present paper shows that species distribution models can give valuable contributions for conservation in saproxylic insects.     

Three-dimensional partitioning of diversity informs state-wide strategies for the conservation of saproxylic beetles

Global conventions on biological diversity force governments to develop region-wide conservation strategies. Such strategies are difficult to design for all taxa because little is known about the important spatial scales. Here we applied additive partitioning of the diversity of saproxylic beetles in Bavarian forests in Southern Germany using a nested hierarchical design of five increasingly broader spatial levels: trap, strata, forest stand, forest site, and ecoregion. We consistently found a significantly higher percentage than expected by chance of between ecoregion diversity and significantly lower α diversity within traps. A significant proportion of β diversity was also found between stands. Analysis of species represented by <0.005% of all specimens in our samples and of species classified as threatened revealed similar results. Critical spatial scales for threatened species encompassed the critical levels of common species. Within habitat substratum guilds, the proportion of β diversity increased from species associated with fresh wood to those associated with rotten wood to those associated with fungi. Our results suggest that the most effective way to ensure saproxylic beetle diversity in a state-wide strategy is to add new conservation sites within different ecoregions and to establish new conservation areas in additional forest stands, rather than to enlarge reserves. Our findings further suggest that monitoring of saproxylic beetle diversity on a broader scale in European temperate forests can be restricted to “monitoring species”, i.e. a subgroup of families easy to identify, and that canopy sampling can be neglected without a substantial loss of information.     

Parasitoids (Hymenoptera, Ichneumonoidea) of Saproxylic beetles are affected by forest successional stage and dead wood characteristics in boreal spruce forest

The habitat requirements and effects of forest management on insects belonging to higher trophic levels are relatively unknown in forest ecosystems. We tested the effect of forest successional stage and dead wood characteristics on the saproxylic parasitoid (Hymenoptera, Ichneumonoidea) assemblage in boreal spruce-dominated forests in northern Sweden. Within each of nine areas, we selected three sites with different management histories: (1) a clear-cut (2) a mature managed forest and (3) an old-growth forest. Parasitoids were collected in 2003 using eclector traps mounted on fresh logs, which were either untreated (control), burned, inoculated with fungi, or naturally shaded, and on artificially-created snags.Both forest type and dead wood characteristics had a significant effect on parasitoid assemblages. Grouped idiobionts and some species, such as Bracon obscurator and Ontsira antica, preferred clear-cuts, while others, such as Cosmophorus regius (Hym., Braconidae) and other koinobionts, were associated with older successional forest stages. No single dead wood substrate was sufficient to support the entire community of parasitoids in any forest type, even when the regular host was present. In particular, snags hosted a different assemblage of species from other types of dead wood, with parasitoids of Tetropium spp. such as Rhimphoctona spp. (Hym., Ichneumonidae) and Helconidea dentator (Hym., Braconidae) being abundant. These results indicate that a diversity of dead wood habitats is necessary to support complete assemblages of beetle-associated parasitoids from early successional stages of dead wood and that parasitoids may be more sensitive to habitat change than their hosts.     

Conservation-oriented forestry and early successional saproxylic beetles: Responses of functional groups to manipulated dead wood substrates

New conservation-oriented forestry aims to maintain intact populations of forest organisms by improving the conservation value of managed forests and providing protected areas. We tested the conservation value of treatments of dead wood for assemblages of early successional saproxylic beetles. In nine areas in northern Sweden, we selected one clear-cut, one mature managed forest and one reserve. In 2001-2002, we placed three blocks of spruce logs, each containing control, burned and shaded logs and a high stump (“snag”) at each site. Saproxylic beetles emerging from the dead wood were collected using emergence traps and beetles flying close to it were collected using flight-intercept traps. After one year of exposure, assemblage composition was examined, with respect to nutritionally-defined functional groups, red-listed species and fire-favoured species. Experimental snags were most complementary to control logs, supporting different assemblages of cambium consumers and fungivores and supporting more red-listed individuals. Burned logs supported depauperate assemblages, particularly with respect to cambium consumers, while shading of logs affected assemblages of fungivores, but only on clear-cuts. Despite containing less dead wood, managed forests provided valuable habitat, supporting similar assemblages of saproxylic beetles to reserves. Most functional groups were less abundant on clear-cuts than in older forests, but fire-favoured species were more common on clear-cuts, suggesting that clear-cuts may support assemblages of species associated with natural disturbances, if suitable substrates are available. Utilization of logs by saproxylic beetles changes over time, so long-term monitoring of our experimental logs will determine their lifetime conservation value.     

Factors and scales potentially important for saproxylic beetles in temperate mixed oak forest

The influence of environmental factors on species richness and species composition may be manifested at different spatial levels. Exploring these relationships is important to understand at which spatial scales certain species and organism groups become sensitive to fragmentation and changes in habitat quality. At different spatial scales we evaluated the potential influence of 45 factors (multiple regression, PCA) on saproxylic oak beetles in 21 smaller broadleaved Swedish forests of conservation importance (woodland key habitats, WKH). Local amount of dead wood in forests is often assumed to be important, but two landscape variables, area of oak dominated woodland key habitats within 1 km of sites and regional amount of dead oak wood, were the main (and strong) predictors of variation in local species richness of oak beetles. The result was similar for red-listed beetles associated with oak. Species composition of the beetles was also best predicted by area of oak woodland key habitat within 1 km, with canopy closure as the second predictor. Despite suitable local quality of the woodland key habitats, the density of such habitat patches may in many areas be too low for long-term protection of saproxylic beetles associated with broadleaved temperate forests. Landscapes with many clustered woodland key habitats rich in oak should have high priority for conservation of saproxylic oak beetles.     

Learning from a “benign neglect strategy” in a national park: Response of saproxylic beetles to dead wood accumulation

Increasing demands for firewood owing to rising energy costs have accelerated discussions about the amount of dead wood needed for conservation. A sharp increase in dead wood caused by bark beetles in a German national park provides lessons for management of commercial and protected forests. We investigated the effects of dead wood due to bark beetle infestation as well as tree senility on abundance and richness of saproxylic species of beetles. Increasing amounts of spruce dead wood and opening of the canopy by bark beetles had positive effects on the abundance of host-generalist, conifer-specialist, and red-listed saproxylic beetles. Broadleaf specialists were positively associated with the amount of broadleaf dead wood and negatively associated with canopy openness. Gradient analysis of beetle assemblages revealed two major environmental axes: canopy openness and amount of dead wood. We found a threshold for community divergence at a canopy openness of 23% (confidence interval CI: 11-49) and at an amount of dead wood of 64 m³ ha⁻¹ (CI: 35-160). Critically endangered species served as indicators of dense and open forests, but only when the amount of dead wood was high. Our results suggest that, to maintain saproxylic beetle assemblages, the amount of dead wood in commercial montane forests (at present ≈15 m³ ha⁻¹) needs to be tripled, with a focus on broadleaf wood in dense stands and spruce wood in open stands. For large protected areas in Europe, our data suggest that bark beetle infestation and senescence without active forest management improves habitat conditions for saproxylic beetles.     

The effect of forest clearcutting in Norway on the community of saproxylic beetles on aspen

In a study of the effect of forest clear-cutting on the saproxylic beetle fauna in aspen in Norwegian boreonemoral forest, we find that both sun exposure and substrate are important structuring factors for the community of saproxylic beetles. Even though the species number and abundance is rather similar across the gradient of sun exposure, there is a major turnover in species composition in decaying aspen from mature forest to clear-cuttings. Our results confirm that aspen is an important element for beetle biodiversity both because of a generally species-rich community and a high number of rare and threatened (red-listed) beetles. There is a higher probability of presence of red-listed beetle species in snags than in logs. Sun exposure increases the probability of presence of red-listed species, but this was largely an effect of one dominating species. We conclude that retention of trees when clear-cutting is an important means for safeguarding the fauna of saproxylic beetles in aspen.     

Ground beetles (Carabidae) and edge effect in oak-hornbeam forest and grassland transects

We evaluated ground beetle diversity in relation to forest edge between an oak-hornbeam forest and adjacent herbaceous grassland. To test our hypothesis that the diversity of ground beetles was higher in the forest edge than the interior, pitfall trap samples were taken along two forest-grassland transects in northern Hungary. The diversity of ground beetles was significantly higher at the forest edge and in the grassland than in the forest interior. Ground beetle assemblages in the forest interior, forest edge and grassland could be separated from each other by ordination. Indicator species analysis detected five groups of species: habitat generalists, grassland-associated species, forest generalists, forest specialists, and edge-associated species. Rank correlation indicated leaf litter, herb, canopy cover, and prey abundance as the most important factors influencing carabid diversity. The high diversity of the forest edge resulted from the presence of edge-associated species and of species characteristic of adjacent habitats. Forest edges seem to play an important role in maintaining diversity. Serving as source habitats, edges also contribute to the recolonisation by ground beetles after habitat destruction or other disturbance in the adjacent habitats.     

Sampling saproxylic beetles: lessons from a 10-year monitoring study

We sampled saproxylic beetles using trunk window traps in two birch-dominated forests in Finland during 1990-1999. The sampling scheme, 10 traps attached to living fruiting bodies of Fomes fomentarius growing on dead birches, remained unchanged in both forests throughout the study period. Beetles belonging to 32 selected families were identified every year, whereas all species were identified during the last 4 years. Total number of identified individuals was 40,294 and number of species 583, of which 258 were saproxylic (dependent on dead wood). Species richness of rare and threatened saproxylic beetles in the samples varied a lot between the years and did not fluctuate synchronously between the forests. Variation between years was smaller when all saproxylic species were pooled together and some abundant species fluctuated synchronously in the two forests. Similarity indices and DCA-ordination did not generally suggest decreasing similarity between samples with increasing temporal isolation. Incidence-based similarities of common saproxylic species within and between forests and years were high (means 0.7-0.8), whereas those of rare saproxylics were roughly 50% smaller and much more variable. More than 75% of the common saproxylic species found during the entire 10-year period were detected already after 3 years of sampling but accumulation of rare and threatened species was much slower. Our results suggest that: (1) occurrence of rare and threatened species in samples is much less predictable than that of common species and, e.g. reserve selection based on rarities should be made cautiously; (2) estimation of total number of threatened species in a forest is very difficult, because such species accumulate slowly in the samples; (3) samples from different years can be comparable in certain cases.     

The effects of forest patch size and matrix type on changes in carabid beetle assemblages in an urbanized landscape

In this study we compared ground beetles (Carabidae) from a range of different forest fragments along an urbanization gradient in Brussels, Belgium. We address the following questions: (i) How does the degree of urbanization in the surrounding habitat affect forest beetles, and does it interact with the effects of patch size and distance to forest edge? (ii) Do these factors have a different effect at the level of individual species, habitat affinity groups or total community? During 2002 we sampled 13 forest plots in 10 forest patches, ranging in size from 5.27 to 4383 ha. The beetles were captured using transects of pitfall traps from the edge to a distance of 100 m into each woodland and identified to species level. Effects of urbanization, forest size and forest edge were evaluated on total species number, abundance and habitat affinity groups and ten abundant, widespread model carabid species. Overall, the effects of urbanization, forest size and edge effects slightly influenced total species richness and abundance but appeared to have a major effect on ground beetle assemblages through species specific responses. More urbanized sites had significantly fewer forest specialists and more generalist species. Large forest fragments were favoured by forest specialist species while generalist species and species frequently associated with forest (forest generalists) dominated the smaller forests. Forest edges mainly harboured generalist species while forest specialist species were more frequent into the forests if the forest patches were large enough, otherwise they disappeared due to the destruction or impoverishment of their habitat. Our results show the importance of differentiating between habitat affinity, especially habitat generalists versus specialists, the latter having a higher value in nature conservation, and merely the quantity of species represented in human-dominated areas.     

Rove beetles and ground beetles (Coleoptera: Staphylinidae, Carabidae) as indicators of harvest and regeneration practices in western Canadian foothills forests

Staphylinid beetle assemblages from coniferous foothills forest in west-central Alberta, Canada were studied via pitfall trapping to examine the effects of stand age and possible edge effects. Sites included a chronosequence of stands from 1 to 27 years post-harvest, and four types of mature forest that had not been disturbed by fire for at least 80 years. In all, 19 sites were sampled between 1989 and 1991. A total of 98 species were identified, nine of which are reported for the first time in Alberta. Staphylinids were more abundant in mature forest stands but assemblages were more diverse in regenerating stands. Thirty-four rove beetle species showed significant indicator value for particular stands or groups of stands, including mature forest, young forest, and open ground specialists. After harvesting, the catch rate of many forest species decreased dramatically, and open ground species were more commonly collected. Populations of some forest species remained active on logged sites for one or 2 years before disappearing. As stands regenerated, they were colonized by species characteristic of young stands, but true forest species were found only in older unharvested stands. The beetle assemblages from regenerating stands became more similar to those from mature stands as they aged, but still differed considerably from them 27 years after harvesting. Transects across forest-clearcut edges revealed a significant beetle response to habitat edges. Staphylinids assemblages were compared to the ground beetle (Carabidae) assemblage sampled via the same pitfall trapping regime. Mature forest specialists are threatened by fragmentation and loss of habitat. In order to conserve these beetle assemblages, forest managers should retain adequate patches of older successional stages on working landscapes.     

Saproxylic beetles in natural and man-made deciduous high stumps retained for conservation

Intensive forest management in Scandinavia has decreased the amount of dead wood required by saproxylic (wood-living) organisms. To reduce this problem, some dead wood is now retained during forest operations, often in the form of man-made high stumps (ca. 4 m high). Most often these stumps are cut with a harvester, although the stumps in this study were made with explosives. The aims of this study were to determine whether such stumps of aspen (Populus tremula) and birch (Betula spp.) could be used by red-listed saproxylic beetles (Coleoptera), and to examine how the fauna of man-made high stumps differs from that of natural stumps. We also studied how tree species, sun-exposure, stage of decay and trunk diameter influenced the fauna. In 169 samples of bark from high stumps 116 saproxylic species were found, of which 21 were red-listed. Many species, including red-listed ones, were more associated with man-made stumps than with natural stumps. However, in total, more species were found in the natural than in the man-made stumps. This is probably because man-made stumps provide a more homogeneous type of wood substrate than natural ones. Among the other variables the difference between aspen and birch was the most important. We conclude that man-made high stumps are valuable habitats for many saproxylic beetle species.     

Effects of climate change on insect defoliator population processes in Canada's boreal forest: Some plausible scenarios

Insect populations have a substantial impact on Canada's forest. They are a dominating disturbance factor and during outbreaks they can cause tree mortality over vast areas of forest. If the predicted climate changes take effect, the damage patterns caused by insects may be drastically altered, especially for the many insects whose occurrence in time and space is severely limited by climatic factors. This possibility substantially increases the uncertainties associated with the long-term planning of pest control requirements, with hazard rating models, with depletion forecasts, and with projections for the sustainability of future timber supplies. Moreover, because insect damage affects the rates of various processes in nutrient and biogeochemical cycling, potential changes in damage patterns can affect ecosystem resilience. This paper presents a number of plausible scenarios that describe how some key processes in the boreal forest's insect defoliator outbreak systems may respond to climate change. The spruce budworm,Choristoneura fumiferana Clem. (Lepidoptera: Tortricidae), is used as an illustrative case study throughout. The potential importance of phonological synchrony in the dynamical interactions between species is emphasised. It is argued that natural selection may be a particularly important process in the response of insects to climate change and that climate change may already be influencing some insect lifecycles. The importance of threshold effects, rare but extreme events, and transient dynamics is emphasised, and the inadequacy of ‘equilibrium’ models for forest:pest systems noted. We conclude by discussing approaches to developing forecasts of how one of the boreal forest's insect defoliator-based disturbance regimes, as a whole, might respond to climate change.     

Assessing the extinction vulnerability of wood-inhabiting fungal species in fragmented northern Swedish boreal forests

Fragmentation of old-growth forests and greatly reduced amounts of coarse dead wood in managed forests threat the persistence of many saproxylic species in boreal Fennoscandia. Individual old-growth forest remnants may lose species over time as they pay off their extinction debt. We tested this by comparing the observed site occupancy of individual wood-inhabiting fungal species in isolated old-growth stands (i.e. woodland key habitats; WKHs) with statistical predictions of their occupancy assuming potential extinction debt had already been paid off. The occupancy of species was analysed in two sets of WKHs differing in time since isolation (i.e. recent and old isolates).Few species occurred more frequently than expected in WKHs. However, patterns across species and across all WKHs masked important differences among species in their risk of facing future extinction. The site occupancy decreased significantly between recent and old isolates for a group of annual, red-listed specialist fungal species, suggesting that an extinction debt in WKHs may exist among specific species confined to coarse dead wood and old-growth forest habitat. Generalist species that also occur in the surrounding matrix showed no negative trends, or actually increased in site occupancy, making future extinctions less likely. Thus, continuing loss of threatened species are likely if not preservation of WKHs are combined with other conservation efforts in managed forest landscapes. Natural forest landscapes may serve as important references when aiming to identify species in risk of future extinction but more detailed knowledge about the biology of the most vulnerable species is also required.     

喀斯特森林生态系统的小生境及其土壤异质性

通过对贵州南部茂兰喀斯特森林生态系统中小生境的土壤进行调查及采样分析,探讨喀斯特微地貌的土壤多样性及土壤性质的变化。结果表明:在小生境地表微形态和微地貌空间变异的影响下,石坑、石沟、石缝、石洞和土面的土壤分布及其性质出现明显的水平空间变异。石洞土壤砂粒含量、>5 mm团聚体数量、容重明显高于石坑、石沟、石缝和土面的土壤,而<1 mm团聚体数量、通气孔隙度及腐殖酸、全氮、全磷、有效氮的含量则相反;石坑土壤<1 mm团聚体数量、通气孔隙度及有机碳、腐殖酸、全氮、有效磷、有效钾、有效铜、有效锌的含量显著高于石洞、石沟、石缝和土面的土壤,而黏粒含量、>5 mm团聚体数量、容重则明显低于土面、石沟和石缝的土壤;石沟土壤的黏粒含量显著低于土面和石缝的土壤,而<1 mm团聚体数量及有机碳、腐殖酸、全氮的含量则相反;石缝土壤有机碳、有效钾的含量明显低于土面、石沟的土壤。初步将喀斯特微地貌的土壤类型分为石洞型、石缝型、土面-石沟型和石坑型,其中土面-石沟型土壤分布最多,其次为石缝型。     

Effects of anthropogenic land use on forest birds and butterflies in Subic Bay, Philippines

Despite the loss of 83% of native forests in the Philippines, little is known on the effects of this massive habitat loss and degradation on its forest biotas. This is a cause for concern because of the threat posed to the country’s large number of endemic taxa. To investigate the impacts of anthropogenic disturbance, forest birds and butterflies were surveyed in closed and open canopy forests, as well as suburban, rural and urban areas within the Subic Bay Watershed Reserve and Olongapo City in western Luzon. Measures of forest species richness and population densities for both taxa were similar in the two forest types, but showed different patterns in the other habitats. Indirect gradient analysis showed that forest bird species were positively correlated with vegetation variables (i.e., canopy cover, tree density, height to inversion and ground cover), while forest butterflies were not strongly correlated to any of the measured habitat variables. Community composition of birds in forests was distinct from those in modified habitats, while butterfly communities were more similar. A simulation showed that canopy cover of 60% or higher was required by 24 of the 26 bird species that were sensitive to canopy loss. Endemicity and nesting strata were the significant predictors of vulnerability to habitat disturbance for birds, while endemicity and larval hostplant specificity were significant for butterflies. Both taxa were negatively affected by anthropogenic disturbance but may respond to different components in the habitat (i.e., structure and resources), and thus cannot be used as surrogates of each other. Conservation of forests with contiguous canopy cover should be prioritized, and more ecological research is needed to improve the knowledge on the effects of disturbance on Philippine biodiversity.     

Habitat and survival of capercaillie Tetrao urogallus nests and broods in the Bavarian alps

The capercaillie Tetrao urogallus has become endangered in central Europe, most probably as a result of habitat changes. Breeding ecology was studied by following radio-tagged hens through egg-laying, incubation, and chick-rearing between 1988 and 1992. Loss of chicks rather than nests was the main determinant of reproductive success. All adult hens incubated, two-thirds of the nests hatched, and one-fifth of the chicks survived until autumn. Both nests and broods were mostly found in habitats with rich ground vegetation. Nest cover was probably a major factor for hatching success. Brood home ranges averaged 148 ha during the period from hatching to late summer. Broods preferred old forest stands with rich ground vegetation and high invertebrate abundance, which occur under moderate canopy cover. Bilberry Vaccinium myrtillus was an essential feature for brood habitat at forest stand, home range, and landscape scale, and should therefore be a major goal for capercaillie habitat management in central Europe.