In search of the best local habitat drivers for saproxylic beetle diversity in temperate deciduous forests

Deadwood-associated species are increasingly targeted in forest biodiversity conservation. In order to improve structural biodiversity indicators and sustainable management guidelines, we need to elucidate ecological and anthropogenic drivers of saproxylic diversity. Herein we aim to disentangle the effects of local habitat attributes which presumably drive saproxylic beetle communities in temperate lowland deciduous forests. We collected data on saproxylic beetles in 104 oak and 49 beech stands in seven French lowland forests and used deadwood, microhabitat and stand features (large trees, openness) as predictor variables to describe local forest conditions. Deadwood diversity and stand openness were consistent key habitat features for species richness and composition in deciduous forests. Large downed deadwood volume was a significant predictor of beetle species richness in oak forests only. In addition, the density of cavity- and fungus-bearing trees had weak but significant effects. We recommend that forest managers favor the local diversification of deadwood types, especially the number of combinations of deadwood positions and tree species, the retention of large downed deadwood and microhabitat-bearing trees in order to maximize the saproxylic beetle diversity at the stand scale in deciduous forests. To improve our understanding of deadwood-biodiversity relationships, further research should be based on targeted surveys on species-microhabitat relationships and should investigate the role of landscape-scale deadwood resources and of historical gaps in continuity of key features availability at the local scale.     

Saproxylic insects and fungi in deciduous forests along a rural–urban gradient

Urbanization is increasing worldwide and is regarded a major threat to biodiversity in forests. As consequences of intensive human use, the vegetation structure of naturally growing urban forests and their amount of deadwood can be reduced. Deadwood is an essential resource for various saproxylic insects and fungi. We assessed the effects of urbanization and forest characteristics on saproxylic insects and fungi. We exposed standardized bundles consisting of each three freshly cut beech and oak branches in 25 forests along a rural–urban gradient in Basel (Switzerland). After an exposure of 8 months, we extracted the saproxylic insects for 10 months using an emergence trap for each bundle. We used drilling chips from each branch to determine fungal operational taxonomic units (OTUs). In all, 193,534 insect individuals emerged from the experimental bundles. Our study showed that the abundance of total saproxylic insects, bark beetles, longhorn beetles, total flies, moths, and ichneumonid wasps decreased with increasing degree of urbanization, but not their species richness. However, the taxonomic composition of all insect groups combined was altered by wood moisture of branches and that of saproxylic beetles was influenced by the degree of urbanization. Unexpectedly, forest size and local forest characteristics had a minor effect on saproxylic insects. ITS (internal transcribed spacer of rDNA) analysis with fungal specific primers revealed a total of 97 fungal OTUs on the bundles. The number of total fungal OTUs decreased with increasing degree of urbanization and was affected by the volume of naturally occurring fine woody debris. The composition of fungal OTUs was altered by the degree of urbanization and pH of the branch wood. As a consequence of the altered compositions of saproxylics, the association between total saproxylic insects and fungi changed along the rural–urban gradient. Our study shows that urbanization can negatively impact saproxylic insects and fungi.     

浙江天目山种植林管理历史及粗死木残体特征影响柳杉腐木甲虫多样性（英文）

不合理的森林管理是导致腐木甲虫多样性丧失的重要原因。在中国亚热带地区, 多样性较高的天然林已被大面积的人工种植林取代, 然而, 这些人工林对腐木甲虫多样性的影响还研究甚少。本研究对浙江天目山自然保护区人工幼龄林（30~40年）、 人工老熟林（80~100年）和半天然混合林（>200年）中柳杉枯立木上的腐木甲虫群落及多样性进行比较。结果表明： 半天然混合林腐木甲虫个体数量（97.4±66.7）显著高于幼龄林（39.9±16.3）和老熟林（21.9±5.9）, 但半天然林混合林(27.9±11.2)与幼龄林(24.1±3.7)腐木甲虫物种数差异并不显著(P>0.05), 而幼龄林的腐木甲虫物种数和个体数量则显著高于老熟林(P<0.05)。腐木甲虫物种数和个体数量与样地粗死木残体体积相关性显著(P<0.05)。典范对应分析和多响应置换过程分析表明腐木甲虫群落特征在不同林型间差异显著(P<0.001)。柳杉枯立木直径、 粗死木残体的直径和数量以及林冠盖度均对腐木甲虫物种组成具有显著影响(P<0.05)。腐木甲虫营养级组成分析也表明, 半天然混合林菌食性甲虫数量显著高于种植林(P<0.001)。结果提示, 提高种植林粗死木残体的数量和质量可以增加腐木甲虫的物种丰富度, 但种植林腐木甲虫多样性可能在随后的演替阶段有所下降, 而且种植林与天然林在腐木甲虫群落组成上差异十分明显。     

Synchronous rise and fall of bark beetle and parasitoid populations in windthrow areas

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Saproxylic beetle assemblages related to silvicultural management intensity and stand structures in a beech forest in Southern Germany

Compared to agricultural land and spruce plantations, central European beech-oak forests are often relatively close to natural conditions. However, forest management may alter these conditions. In Steigerwald, southern Germany, a large beech-dominated forest area, three management intensities were applied during the past 30–70 years. Here, we examined the influence of management intensity on saproxylic beetles in >100-year old mature stands at 69 sampling plots in 2004. We sampled beetles using flight-window traps and time standard direct searches. The community structure based on presence/absence data changed remarkably along the gradient from unmanaged to low-intensity to high-intensity management, but these differences were not evident using abundance data from flight interception traps. Saproxylic species richness decreased in intensively managed forests. Elateridae and threatened species richness peaked in unmanaged forests and in forests under low-intensity management. Saproxylic species richness was dependent on certain micro-habitat factors. These factors were (1) the amount of dead wood for Elateridae, overall and threatened saproxylic beetle richness; (2) the amount of flowering plants for Cerambycidae; (3) the richness of wood-inhabiting fungi for Staphylinidae, Melandryidae and overall saproxylic beetle richness; and (4) the frequency of Fomes fomentarius for threatened species. Species richness was better explained by plot factors, such as dead wood or fungi, than by management intensity. These results suggest that the natural variation of dead wood niches (decay stages, snag sizes, tree cavities and wood-inhabiting fungi species) must be maintained to efficiently conserve the whole saproxylic beetle fauna of beech forests. Also, intensive management may alter the specialised saproxylic beetle community even if the initial tree-species composition is maintained, which was the case in our study. For monitoring the ecological sustainability of forest management we must focus on threatened species. If structures alone are sampled then the amount of dead wood is the best indicator for a rich saproxylic beetle fauna.     

Population dynamics in changing environments: the case of an eruptive forest pest species

In recent decades we have seen rapid and co‐occurring changes in landscape structure, species distributions and even climate as consequences of human activity. Such changes affect the dynamics of the interaction between major forest pest species, such as bark beetles (Coleoptera: Curculionidae, Scolytinae), and their host trees. Normally breeding mostly in broken or severely stressed spruce; at high population densities some bark beetle species can colonise and kill healthy trees on scales ranging from single trees in a stand to multi‐annual landscape‐wide outbreaks. In Eurasia, the largest outbreaks are caused by the spruce bark beetle, Ips typographus (Linnaeus), which is common and shares a wide distribution with its main host, Norway spruce (Picea abies Karst.). A large literature is now available, from which this review aims to synthesize research relevant for the population dynamics of I. typographus and co‐occurring species under changing conditions. We find that spruce bark beetle population dynamics tend to be metastable, but that mixed‐species and age‐heterogeneous forests with good site‐matching tend to be less susceptible to large‐scale outbreaks. While large accumulations of logs should be removed and/or debarked before the next swarming period, intensive removal of all coarse dead wood may be counterproductive, as it reduces the diversity of predators that in some areas may play a role in keeping I. typographus populations below the outbreak threshold, and sanitary logging frequently causes edge effects and root damage, reducing the resistance of remaining trees. It is very hard to predict the outcome of interspecific interactions due to invading beetle species or I. typographus establishing outside its current range, as they can be of varying sign and strength and may fluctuate depending on environmental factors and population phase. Most research indicates that beetle outbreaks will increase in frequency and magnitude as temperature, wind speed and precipitation variability increases, and that mitigating forestry practices should be adopted as soon as possible considering the time lags involved.     

Different effects of coarse woody material on the species diversity of three saproxylic beetle families (Cerambycidae, Melandryidae, and Curculionidae)

The relationships between coarse woody material (CWM) and the diversity of three saproxylic beetle families (Cerambycidae, Melandryidae, and Curculionidae) were investigated. These three families responded differently toward the quality and quantity of CWM. The species richness of cerambycid beetles increased in forests with more CWM in the early stage of decomposition. The richness of saproxylic curculionids increased in stands with more tree species. No clear trend was evident for melandryid diversity. The CWM differed in quality and quantity among three forest types [middle-aged larch (Larix kaempferi [Lamb.] Carrière) plantation, secondary forest, and old-growth forest] and between two forest-management practices (thinning and long rotation). Forest type and management practices affected CWM and the number of tree species, and CWM and the number of tree species influenced the diversity of saproxylic families differently. Because each decomposition stage is considered to be important for saproxylic beetles at the species level, a constant supply of CWM is necessary to maintain a suitable balance of CWM through these decomposition stages in forests to maintain the diversity of saproxylic beetles.     

Are bark beetle outbreaks less synchronous than forest Lepidoptera outbreaks?

Comparisons of intraspecific spatial synchrony across multiple epidemic insect species can be useful for generating hypotheses about major determinants of population patterns at larger scales. The present study compares patterns of spatial synchrony in outbreaks of six epidemic bark beetle species in North America and Europe. Spatial synchrony among populations of the Eurasian spruce bark beetle Ips typographus was significantly higher than for the other bark beetle species. The spatial synchrony observed in epidemic bark beetles was also compared with previously published patterns of synchrony in outbreaks of defoliating forest Lepidoptera, revealing a marked difference between these two major insect groups. The bark beetles exhibited a generally lower degree of spatial synchrony than the Lepidoptera, possibly because bark beetles are synchronized by different weather variables that are acting on a smaller scale than those affecting the Lepidoptera, or because inherent differences in their dynamics leads to more cyclic oscillations and more synchronous spatial dynamics in the Lepidoptera.     

Influence of tree hollow characteristics and forest structure on saproxylic beetle diversity in tree hollows in managed forests in a regional comparison

Tree hollows are among the rarest habitats in today''s Central European managed forests but are considered key structures for high biodiversity in forests. To analyze and compare the effects of tree hollow characteristics and forest structure on diversity of saproxylic beetles in tree hollows in differently structured managed forests, we examined between 41 and 50 tree hollows in beech trees in each of three state forest management districts in Germany. During the two‐year study, we collected 283 saproxylic beetle species (5880 individuals; 22% threatened species), using emergence traps. At small spatial scales, the size of hollow entrance and the number of surrounding microhabitat structures positively influenced beetle diversity, while the stage of wood mould decomposition had a negative influence, across all three forest districts. We utilized forest inventory data to analyze the effects of forest structure in radii of 50–500 m around tree hollows on saproxylic beetle diversity in the hollows. At these larger spatial scales, the three forest management districts differed remarkably regarding the parameters that influenced saproxylic beetle diversity in tree hollows. In Ebrach, characterized by mostly deciduous trees, the amount of dead wood positively influenced beetle diversity. In the mostly coniferous Fichtelberg forest district, with highly isolated tree hollows, in contrast, only the proportion of beech trees around the focal tree hollows showed a positive influence on beetle diversity. In Kelheim, characterized by mixed forest stands, there were no significant relationships between forest structure and beetle diversity in tree hollows. In this study, the same local tree hollow parameters influenced saproxylic beetle diversity in all three study regions, while parameters of forest structure at larger spatial scales differed in their importance, depending on tree‐species composition.     

Horizontal and vertical distribution of saproxylic beetles (Col., Buprestidae, Cerambycidae, Scolytinae) across sections of forest edges

Abstract:  We evaluated the preferred home ranges of three saproxylic beetle taxa along transects from the open field into the forest interior, and from the forest floor up to the canopy. By means of trap sets on metal scaffolds, vertical and horizontal strata were sampled across two types of forest edges: soft-edge ecotones with a gradual transition from the field into the forest and hard edges with an abrupt transition. The forest edges consisted of different strata such as herbaceous fringe, shrub belt, unmanaged forest and managed forest. The thermophilic buprestids were mainly caught in the open land (herbaceous fringe and agricultural land) and in the upper forest mantle. In general, the cerambycids were most abundant in the open land and the lower forest mantle, but a few species favoured the forest interior. The bark beetles (Scolytinae) were equally distributed in all habitats. These distribution patterns of the taxa were observed in terms of both species numbers and abundances. Each species with at least five collected specimens was assigned to one of the three habitat types: open land, forest mantle and forest interior. Of 74 ranked species, only 16% were prevalent in the forest interior and are thus considered to be true forest species. The other 84% of the species were attributed to open land or the forest mantle and are, therefore, forest edge species. Soft forest edges generally supported a higher species richness than hard edges, particularly as regards Cerambycidae and Scolytinae. In terms of Shannon diversity, soft edges tended to be more diverse in buprestids and cerambycids. Overall, the forest interior showed the least species richness and diversity. Therefore, for the conservation of saproxylic beetles, not only the amount and quality of dead wood is important, but also the presence and design of forest boundary structures.     

Sampling saproxylic beetle assemblages in dead wood logs: comparing window and eclector traps to traditional bark sieving and a refinement

The use of saproxylic beetle community as a metric to evaluate nature conservation measures in forests requires efficient methods. We first compare traditional bark sieving to a potential improvement (extracting beetles from whole bark with Tullgren funnels) to determine the most efficient. Secondly we compare this most efficient bark sampling to eclector and window traps. At the species, family, and functional group levels, we consider species richness, abundance and practical aspects. Traditional bark sieving missed >50% of the individual beetles compared to whole bark sampling so we recommend the latter. Window traps caught large numbers of mobile saproxylic beetles, but a high proportion of non-saproxylics results in high sorting cost; bark sampling and eclector traps had a high proportion of saproxylics and obligate saproxylics. Compared to bark sampling, eclector traps are non-destructive, and monitor the whole saproxylic assemblage (i.e. also beetles inside the wood). Overall, window traps are useful because they capture saproxylic beetles attracted to dead wood and sample the local species pool, whereas eclector traps capture the saproxylics that actually emerge from a particular piece of dead wood, and thus are suited to detailed studies. Overall, we suggest that a combination of these two best methods is highly complementary.     

Linking taxonomical and functional biodiversity of saproxylic fungi and beetles in broad‐leaved forests in southern Italy with varying management histories

Abstract

The fundamental ecological significance of deadwood decomposition in forests has been highlighted in several reviews, some conclusions regarding silviculture being drawn. Old‐growth forests are natural centres of biodiversity. Saproxylic fungi and beetles, which are vital components of these ecosystems, occupy a variety of spatial and trophic niches. Fungal and beetle diversity on coarse woody debris (CWD) was analysed in 36 forest sites in the Cilento and Vallo di Diano National Park, Italy. The data were analysed by DCA and Spearman’s rank correlation. The results provide empirical evidence of the existence of a pattern of joint colonization of the woody substrate by fungi and beetles, which includes an assemblage of reciprocal trophic roles within fungal/beetle communities. These organisms act together to form a dynamic taxonomical and functional ecosystem component within the complex set of processes involved in wood decay. The variables most predictive of correlations between management‐related structural attributes and fungal/beetle species richness and their trophic roles for old‐growth forest are: number of logs, number of decay classes and CWD total volume. Deadwood spatio‐temporal continuity should be the main objective of forest planning to stop the loss of saproxylic fungal and insect biodiversity.     

Are biodiversity patterns of saproxylic beetles shaped by habitat limitation or dispersal limitation? A case study in unfragmented montane forests

Understanding the processes that shape biodiversity patterns is essential for ecosystem management and conservation. Local environmental conditions are often good predictors of species distribution and variations in habitat quality usually positively correlate to species richness. However, beside habitat limitation, species presence-absence may be constrained by dispersal limitation. We tested the relative importance of both limitations on saproxylic beetle diversity, using forest continuity as a surrogate for dispersal limitation and stand maturity as a surrogate for habitat limitation. Forest continuity relies on the maintenance of a forest cover over time, while stand maturity results in the presence of old-growth habitat features. Forty montane beech-fir forests in the French pre-Alps were sampled, under a balanced sampling design in which forest continuity and stand maturity were crossed. A total of 307 saproxylic beetle species were captured using flight-interception traps and Winkler–Berlese extractors. We explored the response of low- versus high-dispersal species groups to forest continuity and stand maturity. Saproxylic beetle diversity increased significantly with stand maturity and was mostly influenced by variables related to deadwood diversity at the stand scale and suitable habitat availability at the landscape scale. Surprisingly, no evidence of dispersal limitation was found, as diversity patterns were not influenced by forest continuity and associated variables, even for low-dispersal species. Our study demonstrates that in an unfragmented forest landscape, saproxylic beetles are able to colonize recent forests, as long as local deadwood resources are sufficiently diversified (e.g. tree species, position, diameter and/or decay stage).     

Robinia pseudoacacia as a surrogate for native tree species for saproxylic beetles inhabiting the riparian mixed forests of northern Italy

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Tree killing by Ips typographus (Coleoptera: Scolytidae) at stand edges with and without colonized felled spruce trees

Abstract 1 To maintain biodiversity in managed spruce forests in Sweden more wind‐felled trees must be retained. However, there is concern among forest owners that this may result in higher tree mortality caused by the spruce bark beetle, Ips typographus (L.) (Col. Scolytidae). 2 To simulate wind‐felled trees, living spruce trees were cut at spruce stand edges bordering fresh clear‐cuttings. Treatments comprised edges with zero, one or five cut trees colonized by I. typographus. Edges with naturally wind‐felled trees colonized by I. typographus were also included in the analyses. 3 During the two following summers, the number of trees killed by I. typographus did not differ between edges with and without felled trees, or between edges with one or five felled trees. 4 Within edges with felled trees, there were more killed trees close to the felled trees than at other parts of the edges. Thus, felled trees provided focal points for attacks within edges. 5 It is concluded that small numbers of wind‐felled trees colonized by I. typographus may be left near spruce stand edges without increasing the risk of beetle‐induced tree mortality.     

Beetle responses to artificial gaps in an oceanic island forest: implications for invasive tree management to conserve endemic species diversity

Natural forests are often replaced by invasive alien trees on isolated oceanic islands. Adequate eradication of invasive trees should be conducted with the goal of biodiversity conservation, because islands support many endemic organisms that depend on native forests. An invasive alien tree, Bischofia javanica Blume (Euphorbiaceae), has invaded and replaced natural forests on the oceanic Ogasawara (Bonin) Islands, Japan, in the northwestern Pacific Ocean. To determine how the removal of B. javanica trees affects insect diversity, we examined flying beetles captured using Malaise traps in B. javanica forests on Hahajima. The abundance, species density, and species composition of wood-boring beetles (Coleoptera: Cerambycidae, Elateridae, Mordellidae, and Scolytidae) were compared between closed-canopy sites and gaps created by girdling B. javanica trees in alien forests during two seasons (June–July and October–November 2005). Of the collected beetles, 75.8, 87.5, 90.0, and 0.0% of cerambycid, elaterid, mordellid, and scolytid beetle species, respectively, were endemic to the Ogasawara Islands. More cerambycid, elaterid, and mordellid individuals were captured in June–July than in October–November; the number of scolytid individuals did not differ between seasons. More cerambycid, elaterid, and scolytid individuals were captured in artificial gaps than on the closed-canopy forest floor. Although fewer mordellid individuals were captured in gaps, more endemic mordellids were captured in gaps. More cerambycid and scolytid species were captured in artificial gaps than in closed-canopy areas. The positive responses of beetles to artificial gaps suggest that the removal of B. javanica increases beetle diversity and the abundance of endemic beetles.     

Field response of non‐target beetles to Ips sexdentatus aggregation pheromone and pine volatiles

Although mass trapping cannot be a definitive control measure, it is one of the few ones available to contain the destruction of millions of cubic metres of conifer forests perpetrated every year worldwide by bark beetles. However, using bark beetle aggregation pheromones during both monitoring and control programs may negatively affect other saproxylic insects. The aim of this study was to describe the response of both Ips sexdentatus and its saproxylic beetle associates, especially predators, to traps baited with a commercial blend of I. sexdentatus aggregation pheromone. Furthermore, the usefulness of adding pine volatiles, such as (?)‐α‐pinene and ethanol, to the pheromone was discussed. The commercial blend proved to be attractive to I. sexdentatus adults, both when used alone and together with pine volatiles. Pheromone attractiveness, however, was lessened by the addition of the volatiles. The pheromone blend proved to be attractive to Thanasimus formicarius, as well as to other predator species. Overall, although during our study, traps baited only with (?)‐α‐pinene and ethanol attracted some predator specimens, I. sexdentatus pheromone traps were more attractive. Our study confirms that calendar differences in flight activity between the bark beetle and its predators are substantial; therefore, they should be taken into account when planning control measures. According to our data, the commercial blend of I. sexdentatus pheromone seems to be the most effective, among the baits used, in catching I. sexdentatus adults, while reducing the impact on T. formicarius.     

Saproxylic beetle diversity in a managed boreal forest: importance of stand characteristics and forestry conservation measures

Saproxylic beetles constitute a significant proportion of boreal forest biodiversity. However, the long history of timber production in Fennoscandia has significantly reduced the availability of dead wood and is considered a threat to the conservation of saproxylic beetle assemblages. Therefore, since the mid‐1990s dead wood retention in harvested stands has formed an integral part of silvicultural practices. However, the contribution of this biodiversity‐orientated management approach to conserving saproxylic beetle assemblages in boreal forest landscapes that include production forestry remains largely untested. We examined differences in resident saproxylic beetle assemblages among stands under different management in a boreal forest landscape in Central Sweden, and in particular stands managed according to new conservation‐orientated practices. We also investigated the relationship between beetle diversity and forest stand characteristics. Bark of coarse woody debris (CWD) was sieved for beetles in old managed stands, unmanaged nature reserves, and set‐aside areas, and clear‐cut stands harvested according to certification guidelines [new forestry (NF) clear‐cuts]. All stand types contributed significantly to the total diversity of beetles found. While stand size, position, and distance to nearest reserve were unimportant, both the quality and the quantity of CWD in stands contributed significantly to explaining beetle abundance and species richness. This extends the previous findings for red‐listed invertebrates, and shows that heterogeneous substrate quality and a range of management practices are necessary to maintain saproxylic beetle diversity in boreal forest landscapes that include production forestry. The unique abiotic conditions in combination with the abundant and varied CWD associated with NF clear‐cuts form an important component of forest stand heterogeneity for saproxylic beetles. It is thus essential that sufficient, diverse, CWD is retained in managed boreal landscapes to ensure the conservation of boreal saproxylic beetle assemblages.     

Investigating the abundance and flight period of bark beetles (Coleoptera: Curculionidae: Scolytinae) over elevational gradients in Sitka spruce forests

1. A warming climate, as predicted under current climate change projections, is likely to influence the population dynamics of many forest insect species. Numerous bark beetle species in both Europe and North America have already responded to a warming climate by significantly expanding their geographical ranges.
2. The aim of the current study was to investigate how populations of bark beetles within stands of Sitka spruce, a widely planted non-native commercial plantation tree species in the U.K., were likely to respond to a warming climate. Experimental plots were established in stands of Sitka spruce over elevational gradients in two commercial forest plantations, and the abundance and emergence times of key bark beetle species were assessed over a 3-year period using flight interception traps. The air temperature difference between the lowest and highest experimental plot in each forest was consistently >1°C throughout the 3-year period.
3. In general, the abundance of the most dominant bark beetle species (e.g. Trypodendron, Dryocoetes, Hylastes spp.) was higher, and emergence times tended to be earlier in the year at the lower elevation plots, where temperatures were higher, although not all bark beetle species responded in the same manner.
4. The results of the study indicated that, under the projected future climate warming scenarios, monoculture Sitka spruce stands at low elevations may potentially be more vulnerable to significant outbreak events from existing or invasive bark beetle species. Hence, consideration of establishing more resilient forests of Sitka spruce by diversifying the species composition and structure of Sitka spruce stands is discussed.

     

New Insights into the Consequences of Post-Windthrow Salvage Logging Revealed by Functional Structure of Saproxylic Beetles Assemblages

Windstorms, bark beetle outbreaks and fires are important natural disturbances in coniferous forests worldwide. Wind-thrown trees promote biodiversity and restoration within production forests, but also cause large economic losses due to bark beetle infestation and accelerated fungal decomposition. Such damaged trees are often removed by salvage logging, which leads to decreased biodiversity and thus increasingly evokes discussions between economists and ecologists about appropriate strategies. To reveal the reasons behind species loss after salvage logging, we used a functional approach based on four habitat-related ecological traits and focused on saproxylic beetles. We predicted that salvage logging would decrease functional diversity (measured as effect sizes of mean pairwise distances using null models) as well as mean values of beetle body size, wood diameter niche and canopy cover niche, but would increase decay stage niche. As expected, salvage logging caused a decrease in species richness, but led to an increase in functional diversity by altering the species composition from habitat-filtered assemblages toward random assemblages. Even though salvage logging removes tree trunks, the most negative effects were found for small and heliophilous species and for species specialized on wood of small diameter. Our results suggested that salvage logging disrupts the natural assembly process on windthrown trees and that negative ecological impacts are caused more by microclimate alteration of the dead-wood objects than by loss of resource amount. These insights underline the power of functional approaches to detect ecosystem responses to anthropogenic disturbance and form a basis for management decisions in conservation. To mitigate negative effects on saproxylic beetle diversity after windthrows, we recommend preserving single windthrown trees or at least their tops with exposed branches during salvage logging. Such an extension of the green-tree retention approach to windthrown trees will preserve natural succession and associated communities of disturbed spruce forests.