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.     

Openness in management: Hands-off vs partial cutting in conservation forests, and the response of beetles

The appropriate management of forest reserves is debated; two major alternatives are succession to ‘wild’ state, or management to produce semi-open stands. For temperate conservation stands, there are no strong experiments replicated at landscape level. In each of 22 forests rich in oaks (Quercus spp.) in Sweden, we set up a closed-canopy wild plot (1 ha), and a cutting plot (1 ha) to produce semi-open conditions, studying them before and after cutting. About 25% of the tree basal area was cut (large trees and dead wood retained) and harvested as bio-fuel, a CO₂ - neutral energy source. We examined the response of beetles and trapped 59,000 individuals (1174 species; 100 red-listed species). For both the guild of herbivorous beetles (222 species) and of saproxylic beetles connected to oak wood (267 species), species richness increased by about 35% in the harvested plots, relative to the wild reference plots. Species composition within the groups changed, though not strongly. Thirteen saproxylic species of 50 analysed, and three herbivores of 12 increased in cutting plots. For red-listed saproxylic beetles, species richness did not change significantly. Regression analyses suggest that more open cutting plots disfavour the red-listed beetles of this forest type. Thus, partial cutting increased species diversity of two beetle groups, probably due to changed microclimate and increase in herbaceous plants, but some red-listed saproxylic beetles may be disfavoured. A hands-off alternative may through storms and other disturbances produce open patches, more dead wood, and favour some species. Combinations of these alternatives, carefully planned at the landscape level, need to be considered.     

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.     

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.     

Diversity of dead wood inhabiting fungi and bryophytes in semi-natural beech forests in Europe

Saproxylic organisms are among the most threatened species in Europe and constitute a major conservation problem because they depend on the most important forestry product - dead wood. Diversity of fungal and bryophyte communities occurring on dead beech trees was analyzed in five European countries (Slovenia, Hungary, The Netherlands, Belgium and Denmark) considering tree level species richness (TLSR), country level species richness (CLSR), frequency distributions of species, occurrence of threatened species and relations between TLSR and decay stage, tree size and countries. Altogether 1009 trees were inventoried in 19 beech dominated forest reserves.The number of fungi in the full dataset was approximately three times larger (456 versus 161 species) and the proportion of low frequent species was higher than among bryophytes. The species richness of bryophytes and fungi was significantly different among countries considering both TLSR and CLSR. In addition the diversity patterns deviated considerably between the two groups of organisms. Slovenian sites appeared to be biodiversity hotspots for bryophytes characterized by high TLSR and CLSR and a high fraction of threatened species. Hungarian sites had somewhat lower bryophyte diversity, while the Atlantic region had deteriorated assemblages. Fungal species richness was very high in Denmark, but the Hungarian and Slovenian sites were richer in threatened and low frequency species. Tree size was better able to explain variation in TLSR in both organism groups than decay stage. TLSR was found to vary significantly between countries but the difference was most considerable in the case of bryophytes.The diversity patterns of both organism groups along the investigated geographical gradient appear to be influenced by both climatic and management related factors (forest history, dead wood availability and continuity, habitat fragmentation). There is no doubt that an increase in the abundance of dead wood in European beech forests will benefit diversity of saproxylic fungi and bryophytes, especially if a continuous presence of large diameter logs are secured within individual stands.     

Conservation of beetles in boreal pine forests: the effects of forest age and naturalness on species assemblages

The effort of boreal forest conservation has emphasised the preservation of old-growth forests while the role of young successional stages in maintaining biodiversity has remained largely unstudied. We compared the richness of beetle species and composition of species assemblages between managed and seminatural forests in five stages of forest succession. The sites were in boreal sub-xeric pine-dominated forests in eastern Finland. Seminatural study sites, especially the recently burned sites, were important habitats for threatened and near-threatened species. We propose that young stages of natural succession should be included in the network of protected forest areas. On the other hand, the composition of saproxylic species assemblages in seminatural forests differed from the assemblages in managed forests, indicating also the need to improve the forest management guidelines so that they better address the requirements of species protection. Regeneration methods applied should resemble or mimic the natural disturbances more closely.     

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.     

The effects of windthrow on forest insect communities: a literature review

This paper reviews the effects that windstorm-induced drastic changes (micro-climate, soil, vegetation, and ground structural heterogeneity) have on forest insect communities. In the current context of shady and CWD-deprived managed forests, windthrow gaps act as regional biodiversity hotspots by maintaining habitat continuity in a mosaic landscape, and by facilitating the breeding and population growth of clearing specialists and saproxylic species. Windthrow gaps are dead-wood islands where forest protection and habitat conservation goals may stand against each other. Besides the quantitative effect of dead wood on bark beetle outbreaks and saproxylic diversity, the latter is favoured by key dead-wood micro-habitats such as large logs, snags and sun-exposed coarse woody debris. The role of natural enemies and sanitation operations in regulating pest outbreaks is discussed. Heterogeneous openings provide many micro-habitats favouring flower-visiting insects, phytophages on saplings, on fallen tree crowns, and on diverse understory flora, as well as ground insects on specific micro-sites.     

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.     

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.     

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.     

Carabid beetle (Coleoptera: Carabidae) diversity in the black spruce succession of eastern Canada

This study provides the first assessment of carabid beetle diversity in a natural forest context that encompasses a complete black spruce (Picea mariana) natural succession. Boreal forest conservation has been based on several assumptions about forest age that only consider species richness without accounting for species composition. It has also been guided by studies of incomplete chronologies that do not include naturally burned or old-growth stages. Twenty-one forest stands of different ages following fire - from recently burned to old-growth stages (0-340 years of age) - were sampled, revealing a strong relationship between age of forest and diversity of Carabidae. Over time, species richness followed a parabolic U-shaped pattern both with observed (Obs) and predicted richness (ACE). Chronological clustering identified four groups of species in the succession: the ‘burned’ group characterized 0-2-year-old forests, ‘regenerating’ (21-58 year), ‘mature’ (70-170 year) and ‘old-growth’ (177-340 year). The time spans corresponding to each of these assemblages lengthen with age of forest at an exponential rate. Ward’s and K-means (clustering without constraint) provided support for the four assemblages but showed variation between individual successions, particularly for the ‘regenerating’ assemblage, identified as the most heterogeneous. The IndVal method identified characteristic species in every stage of the succession, particularly Sericoda spp. in the burned stage and Dromius piceus and Platynus mannerheimii associated with old-growth stands. The results obtained here show that diversity of Carabidae varies in primeval conditions according to age of forest and such variation should be taken into account when conservation issues are involved.     

Polypore diversity in managed and old-growth boreal Picea abies forests in southern Finland

Polypore communities were compared between mature managed, overmature managed, and old-growth spruce-dominated forests in southern Finland. A total of 85 polypore species, with 6000 records, were found in 16 sample plots, each 4 ha in size. Old-growth stands had on average 80% more species than mature stands, and 38% more species than overmature managed stands. Variation in polypore species richness was best explained by diversity of dead wood and the number and volume of dead trees. The best predictor for the number of threatened polypore species was the number of cut stumps. Threatened species were practically confined to old-growth forests and to stands in which the amount of dead wood exceeded 20 m³/ha. This figure appears to represent a stand-level threshold value for the amount of dead wood, below which the persistence of threatened species becomes unlikely. Our results suggest that a significant increase in the amount of dead wood (e.g. by leaving large retention trees and even by killing trees) is needed in managed forests before they become suitable habitats for threatened polypores.     

Cost-effectiveness of silvicultural measures to increase substrate availability for red-listed wood-living organisms in Norway spruce forests

It is important that measures to maintain biodiversity are taken in a way that is cost-effective for the landowner. We analyzed the cost-effectiveness of silvicultural measures that aim at increasing the substrate availability for red-listed (species that are threatened, near threatened or where species probably are threatened but data is deficient) saproxylic (wood-inhabiting) organisms. We modelled stands of Norway spruce (Picea abies) in three regions of Sweden by using computer simulations and a database with substrate requirements of saproxylic beetles and cryptogams on the Swedish Red-List. Conclusions concerning cost-effectiveness of silvicultural measures depend on the extinction thresholds of the species they are intended to conserve; measures that generate only small amounts of coarse woody debris (CWD) may provide too little substrate to be useful for species with high extinction thresholds. In northern Sweden, forestland is relatively inexpensive, so a cost-effective strategy to increase the amount of spruce CWD was to set aside more forests as reserves. In central and southern Sweden, more emphasis should instead be given to increasing the amount of CWD in the managed forest. The regulations by the Forest Stewardship Council (FSC) could be made more cost-effective by prescribing creation of more high stumps and retention of larger amounts of naturally dying trees. Large-sized CWD, CWD from slow-growing trees, and CWD in late decay stages are substrate types that were particularly rare in managed forest in relation to unmanaged forests. Manual soil scarification and retention of living trees are measures that can increase the proportion of these underrepresented CWD types.     

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.     

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.     

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.     

The importance of secondary forest blocks for terrestrial mammals within an Acacia/secondary forest matrix in Sarawak, Malaysia

With the rapid conversion of tropical forests in Asia to plantation forests for timber or biofuel production there is a need to determine if these forests serve any viable role in wildlife conservation. We used infra-red trip cameras to survey for large terrestrial mammals within an Acacia/secondary forest matrix being created for pulpwood production in Sarawak, Malaysia. We detected at least 27 species of mammals within the matrix and 18 species were detected in both Acacia and secondary forest. Using occupancy modeling to determine important covariates for seven mammal species within the project area, six of these species were sensitive to forest type, and most were sensitive to the amount of secondary forest within 1 km of the sample point. For four species (sun bear, Helarctos melayanus; common porcupine, Hystrix brachyura; mousedeer, Tragulus spp.; and pig-tailed macaque, Macaca nemestrina), the mean distance of detection from large secondary forest blocks was significantly closer than expected from the distribution of sample points. Most species used Acacia forest less frequently than secondary forest, possibly for transit or foraging, with the exception of bearded pigs (Sus barbatus), sambar (Rucervus unicolor), and civets which were more common in Acacia stands. The amount of secondary forest preserved within forest plantations seems to be the best measure of conservation potential for these industrial forests. Forest plantations can provide a conservation value if managed properly and those retaining significant amounts of secondary forest should be eligible for a base level of certification as they comprise a significant portion of the landscape in this region.     

Wood-inhabiting macrofungi in Danish beech-forests - conflicting diversity patterns and their implications in a conservation perspective

Macrofungal diversity was investigated on 281 decaying beech trees distributed across 14 forests in Denmark, based on sporocarp inventories. Two aspects of diversity were considered, i.e., species richness per fallen tree and the incidence of red-listed species occurrence per tree. For both diversity measures the effects of both tree and site variables were tested. In total, 319 fungal species were identified, including 28 red-listed. Decay stage and wood volume were identified as key variables influencing species richness as well as red-listed species incidence. Red-listed species, however, showed a preference for more decayed trees than non red-listed species. Further, red-listed species incidence was found to be significantly higher on broken trees, compared to fallen trees with a distinct root-plate, indicating tree death cause to be important for some red-listed species. The relations between diversity measures and site variables were conflicting. Species richness per tree decreased with increasing maximum tree age and dead wood continuity, possibly a consequence of competitive exclusion of unspecialised opportunistic species in old-grown stands. For red-listed species the opposite trend was evident, and it is concluded that forest history may have fundamental effects on the community structure of wood-inhabiting fungi. Accordingly, simple species richness may be a misleading conservation measure if the aim is to conserve the most threatened aspects of forest biodiversity.     

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.