“Primeval forest relict beetles” of Central Europe: a set of 168 umbrella species for the protection of primeval forest remnants

Identification of forest stands with priority for the conservation of biodiversity is of particular importance in landscapes with a long cultural and agricultural history, such as Central Europe. A group of species with a high indicator value for the naturalness of forest ecosystems are saproxylic insects. Some of these species, especially within the order Coleoptera, have been described as primeval forests relicts. Here, we compiled a list of 168 “primeval forest relict species” of saproxylic beetles based on expert knowledge. These species can serve as focal and umbrella species for forest conservation in Central Europe. They were selected because of their dependence on the continuous presence of primeval forest habitat features, such as over-mature trees, high amounts of dead wood, and dead wood diversity, as well as their absence in managed Central European forests. These primeval forest relict species showed a moderately strong clumping pattern within the phylogeny of beetles, as indicated by phylogenetic signal testing using the D-statistic. When we controlled for phylogenetic relatedness, an ordinal linear model revealed that large body size and preference for dead wood and trees of large diameter are the main characteristics of these species. This list of species can be used to identify forest stands of conservation value throughout Central Europe, to prioritize conservation and to raise public awareness for conservation issues related to primeval forests.     

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.     

Are there saproxylic beetle species characteristic of high dead wood connectivity?

Saproxylic beetles have been shown to be vulnerable to within-forest fragmentation expressed as large distances between single dead wood pieces (low spatial connectivity of dead wood). From samples of a two-year study of saproxylic beetles, species that were characteristic of sites with high dead wood connectivity were identified by Canonical Correspondence Analysis, the method of Dufrêne and Legendre (IndVal) and by considering the species occurring exclusively in sites with high dead wood connectivity. These species differed mainly from the other species by their high habitat specificity. Hence, there arc species-specific responses of saproxylic beetles to the spatial arrangement of dead wood. High dead wood connectivity must be achieved in managed forests to sustain species which are particularly vulnerable to fragmentation.     

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.     

Mimicking natural disturbances of boreal forests: the effects of controlled burning and creating dead wood on beetle diversity

The young successional stages of boreal forests are an important habitat for many saproxylic species. These habitats are formed by disturbances such as forest fires and they are characterized by large volumes of dead wood and sun-exposed conditions. Today, young successional stages of natural origin are very rare in Fennoscandia and there is need for restoration. We constructed a large-scale field experiment in which we studied the effects of two restoration practices on beetle diversity: controlled burning and partial harvesting with creating different volumes of dead wood. We sampled beetles with flight-intercept traps recording a total of 56,031 individuals and 755 species. The species richness and abundance of both saproxylic and non-saproxylic beetles were increased by burning and harvesting but the volume of dead wood created on harvested sites had no short-term effect on species richness or abundance. Rare species, especially saproxylic ones, preferred burned sites and a similar trend was observed among red-listed and pyrophilous species. Burning and harvesting also resulted in different species assemblages and there were some additional differences according to the volume of dead wood. We conclude that fire can be successfully used in restoration of managed boreal forests to increase species diversity and to facilitate the recovery of declined species. However, long-term monitoring is needed to clarify the effects of the restoration practices, in particular those of creating dead wood without using fire.     

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.     

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

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Fast but ephemeral effects of ecological restoration on forest beetle community

Many protected areas have a long history of human intervention before being protected. In protected forests, the past land use has reduced the amount of natural structures, which are crucial substrates for thousands of species. We evaluate the short-term ecological effect of forest restoration (dead wood creation) on conifer-associated saproxylic (dead-wood dependent) beetles. More specifically, we analyze the effect of dead wood creation on the number of beetle species and individuals 1 and 5 years after restoration in spruce and pine forests, using a large-scale monitoring network over Finland. The number of saproxylic beetle species and individuals was larger at restored than at control plots both 1 and 5 years after restoration in both spruce and pine forests. Community composition in restored plots was different from control plots 1 year after restoration, but had returned towards the control plot composition 5 years after restoration, while control plots remained largely unchanged. Both in spruce and pine forests, there were more red-listed and rare saproxylic beetles in restored than in control plots 1 and 5 years after restoration. Our results indicate that restoration has an overall positive influence on saproxylic beetle diversity immediately after dead wood creation, but this effect is rather short-lived. Long term monitoring of restored dead wood is crucial in investigating successional pathways as well as biotic communities in advanced decay stages, and in fully evaluating the ecological effect of dead wood creation as a forest restoration measure.     

Increasing temperature may compensate for lower amounts of dead wood in driving richness of saproxylic beetles

Global warming and land‐use change are expected to be additive threats to global diversity, to which insects contribute the highest proportion. Insects are strongly influenced by temperature but also require specific habitat resources, and thus interaction between the two factors is likely. We selected saproxylic beetles as a model group because their life cycle depends on dead wood, which is highly threatened by land use. We tested the extent to which higher temperatures compensate for the negative effects of low amounts of dead wood on saproxylic beetle species richness (Temperature–Dead wood compensation hypothesis) on both a macroclimate and a topoclimate scale (north‐ and south‐facing slopes). We analyzed 1404 flight‐interception trap catches across Europe to test for interaction effects of temperature and dead‐wood amount on species richness. To experimentally test our findings from the activity trap data, we additionally reared beetles from 80 bundles of dead wood initially exposed at high and low elevations. At the topoclimate scale, we analyzed trap catches and reared beetles from dead wood exposed in 20 forest stands on south‐facing and north‐facing slopes in one region. On the macroscale, both temperature and dead‐wood amount positively affected total and threatened species richness independently, but their interaction was significantly negative, indicating compensation. On both scales and irrespective of the method, species richness decreased with temperature decline. Our observation that increasing temperature compensates for lower amounts of dead wood has two important implications. First, managers of production forests should adapt their dead‐wood enrichment strategy to site‐specific temperature conditions. Second, an increase in temperature will compensate at least partially for poor habitat conditions in production forests. Such a perspective contrasts the general assumption of reinforcing impacts of global warming and habitat loss on biodiversity, but it is corroborated by recent range expansions of threatened beetle species.     

Biodiversity conservation in old-growth boreal forest: black spruce and balsam fir snags harbour distinct assemblages of saproxylic beetles

Biodiversity conservation of forest ecosystems strongly relies on effective dead wood management. However, the responses of saproxylic communities to variations in dead wood characteristics remains poorly documented, a lack of knowledge that may impede the development of efficient management strategies. We established the relationship between saproxylic beetles—at the species and community levels—and attributes of black spruce and balsam fir in old-growth boreal forests. The relationship was first evaluated for individual snag bole segments, and then for forest stands. A total of 168 bole sections were collected in summer 2006 along a compositional gradient ranging from black spruce-dominated stands to balsam fir-dominated ones, in a boreal forest dominated by >90-year-old stands. A total of 16,804 beetles belonging to 47 species emerged from bole segments, with 21% of the species being found exclusively in black spruce snags and 36% exclusively in balsam fir snags. Black spruce and balsam fir snags thus contributed differently to forest biodiversity by being inhabited by different saproxylic communities. Wood density was an important attribute in the host-use patterns for several species of saproxylic beetles, but no relationship was found between snag availability within stands and abundance of beetles strongly linked to either black spruce or balsam fir. Our study outlines the relative contribution of tree compositional diversity to saproxylic species, while highlighting the contribution of black spruce and balsam fir to animal diversity in old-growth boreal forests.     

Effects of different tree harvesting intensities on saproxylic beetle diversity in coniferous and broad-leaved mixed forest in the Changbai Mountain,Northeast, China

1. Saproxylic beetles have gained increasing attention due to their role in the decomposition of rotting wood in forests. Studying the response of saproxylic beetles to tree harvesting is important for developing harvesting strategies that consider conservation of saproxylic beetle diversity.
2. We report results from a case study in which we designed four treatment stands to test the effects of forest harvesting intensity on saproxylic beetle diversity, harvest intensities of 0% untreated control (CK), 17.2% light harvest (LT), 34.7% moderate harvest (MT) and 51.9% high harvest (HT). Flight intercept traps were used to collect specimens of saproxylic beetles in each stand at 2, 3 and 4 years post-harvest.
3. The richness and abundance of saproxylic beetles were higher in MT and HT than in CK. Twelve of the 15 indicator species were significantly associated with MT or HT, whereas only three species were significantly associated with CK.
4. We found that moderate and high intensity harvesting affected the composition and increased beetle abundance and richness of saproxylic beetles, and light intensity harvesting had no effect on the beetle community in years 2, 3 and 4 post-harvest. There was no difference in the beetle community composition between moderate and high harvest stands.

     

Ten years after: Release cutting around old oaks still affects oak vitality and saproxylic beetles in a Norway spruce stand

1. It is often suggested to release oaks (Quercus robur) from competition, to ensure their survival and boost their conservational value. However, few studies have explored how long-lasting this effect is and how it affects saproxylic beetles.
2. Ten years after cutting, we investigated effects of different release cutting levels (high, medium, and no release) around 140-year-old retained oaks in a commercial forest with Norway spruce (Picea abies).
3. We evaluated oak vitality using crown and dead wood measurements. Saproxylic beetles were caught in window traps, identified to species level, and grouped according to their association with oak and/or Norway spruce.
4. Released oaks had more light, higher temperatures, greater crown growth, and less dead wood in the crown compared to the no-release control.
5. After 10 years we still found a higher abundance of oak-associated beetles and higher overall species richness of saproxylic beetles in the released oaks. Beetle species composition significantly differed between released oaks and control.
6. We suggest avoiding planting trees beneath retained oaks when regenerating conifer forests and to proceed with conservation management during subsequent thinning by removing regeneration under the oak crowns. We emphasize the benefits of monitoring the retained oaks to maintain tree vitality, habitat quality, and insolation.

     

Colonisation of ephemeral forest habitats by specialised species: beetles and bugs associated with recently dead aspen wood

The most appropriate strategy for preserving fragmented populations depends on a species’ ability to colonise distant habitat patches. Insects associated with early decay stages of dead wood are expected to have a high capacity to colonise new habitat patches. To study the dispersal ranges of beetles (Coleoptera) and flat bugs (Hemiptera: Aradidae) dependent on recently dead aspen (Populus tremula) wood in Finland, we set out 58 piles of recently cut aspen logs at various distances up to 1.6 km from forests that contained a high density of old aspen trees. We captured insects by trunk window-traps, and counted beetles’ exit holes. Habitat connectivity was measured in terms of the amount of suitable aspen-wood in the surrounding environment, with the closest dead wood items up-weighted by a negative-exponential function. The log-piles attracted many saproxylic insects including four red-listed aspen-specialist species. The exposure of log-piles to the sun, and high levels of habitat connectivity increased the species richness of aspen-specialists, whereas bark peeling by moose decreased richness. The spatial scale at which species richness had its strongest response to habitat was 93 m. Among individual species there was a wide variability in spatial scale of response. This study supports the view that conservation efforts in boreal forests should be concentrated on sites where colonisation by target species is most likely. Restoration of habitat by re-locating logs may be useful at localities with a rich and specialised fauna but which have too low rate of formation of dead wood by natural processes.     

Micro and macro-habitat associations in saproxylic beetles: implications for biodiversity management

Restoration of habitats is critically important in preventing full realization of the extinction debt owed as a result of anthropogenic habitat destruction. Although much emphasis has been placed on macrohabitats, suitable microhabitats are also vital for the survival of most species. The aim of this large-scale field experiment was to evaluate the relative importance of manipulated microhabitats, i.e., dead wood substrates of spruce (snags, and logs that were burned, inoculated with wood fungi or shaded) and macrohabitats, i.e., stand types (clear-cuts, mature managed forests, and forest reserves) for species richness, abundance and assemblage composition of all saproxylic and red-listed saproxylic beetles. Beetles were collected in emergence traps in 30 forest stands in 2001, 2003, 2004 and 2006. More individuals emerged from snags and untreated logs than from burned and shaded logs, but species richness did not differ among substrates. Assemblage composition differed among substrates for both all saproxylics and red-listed saproxylic species, mainly attributed to different assemblage composition on snags. This suggests that the practise of leaving snags for conservation purposes should be complemented with log supplementation. Clear-cuts supported fewer species and different assemblages from mature managed forests and reserves. Neither abundance, nor species richness or assemblage composition differed between reserves and mature managed forests. This suggests that managed stands subjected to selective cutting, not clear-felling, maintain sufficient old growth characteristics and continuity to maintain more or less intact assemblages of saproxylic beetles. Thus, alternative management methods, e.g., continuity forestry should be considered for some of these stands to maintain continuity and conservation values. Furthermore, the significantly higher estimated abundance per ha of red-listed beetles in reserves underlines the importance of reserves for maintaining viable populations of rare red-listed species and as source areas for saproxylic species in boreal forest landscapes.     

Contrasting Responses of Saproxylic Insects to Focal Habitat Resources: The Example of Longhorn Beetles and Hoverflies in Belgian Deciduous Forests

Although both saproxylic longhorn beetles and hoverflies benefit from the presence of woody substrates for reproduction, they differ in their requirements for floral resources and for microbiotopes of overmature and senescent trees. This led us to expect contrasting responses between the two species groups in relation to these essential resources. We examined this prediction in 22 mature oak- and beech-dominated stands of southern Belgium by relating their species assemblages to local vegetation structure and composition, altitude and landscape composition. Stands were organised in pairs as a function of their overall dead wood supply. Free-hanging window traps, stump emergence traps and Malaise traps produced 30 longhorn beetle species (1637 individuals) and 106 hoverfly species (3020 individuals). Paired-comparisons controlling for annual variation in captures showed that, unlike saproxylic hoverflies, stands with dead wood hosted more species and individuals of longhorn beetles. Accordingly, the two species groups were found to be independent on ordination axes, responding to different sets of environmental conditions. While stands dominated by oaks with a high snag volume were highly favoured by longhorn beetles, saproxylic and threatened syrphids were limited to open-stands with large trees and a well-developed, species rich herb layer providing the floral resources required for their reproduction. Our results suggest that, when defining criteria to identify or restore important habitats for saproxylic insect conservation, variables related to different aspects of dead wood supply should not be the only criteria taken into account.     

Evaluation of window flight traps for effectiveness at monitoring dead wood-associated beetles: the effect of ethanol lure under contrasting environmental conditions

1 Subsequent to the diversity of saproxylic beetles being proposed as a management tool in forestry, more explicit knowledge about the efficiency and selective properties of beetle sampling methods is needed.
2 We compared saproxylic beetle assemblages caught by alcohol-baited or unbaited window traps in different forest contexts. Considering that trap attractiveness depends on kairomone concentrations, we appraised whether the trap efficiency was influenced by trap environment (openness and local supply of fresh dead wood).
3 Saproxylic beetles were sampled using 48 cross-vane window flight traps, arranged in paired designs (alcohol-baited/unbaited), in eight ancient and eight recent gaps (open stands), and eight closed-canopy control stands in an upland beech forest in the French Pyrenees.
4 Baited traps were more efficient than unbaited traps in terms of abundance and richness in our deciduous forests. The ethanol lure did not have any repellent effect on the individual response of saproxylic taxa.
5 The influence of local environmental conditions on trap attractiveness was observed. Openness had a significant moderate effect on species richness. Trap attractiveness was slightly reduced in the alcohol-saturated environment of recent gaps probably due to a disruption by local fresh dead-wood concentrations of the kairomonal response of saproxylic beetles to baited traps ('alcohol disruption').
6 Because the ethanol lure enhanced the probability of species detection, it may be useful in early-warning surveillance, monitoring and control of wood borers, despite slight influences of local conditions on baited trap efficiency.     

Role of urban green spaces for saproxylic beetle conservation: a case study of tenebrionids in Rome,Italy

Forested urban areas provide many important ecosystem services and their preservation is considered of paramount importance. Although urban forest are known to host a high diversity of saproxylic beetles (i.e. those associated with dead wood), contributions dealing with the role of urban green spaces for their conservation are lacking. We investigated the importance of urban green spaces for saproxylic and non-saproxylic tenebrionid beetles in urban Rome. Based on species vulnerability scores we calculated two indices of area prioritisation, the Biodiversity Conservation Concern (BCC) and the Biodiversity Conservation Weight (BCW) for saproxylic and non-saproxylic species. Site area and forest surface correlated positively with saproxylic richness, whereas site isolation correlated negatively with non-saproxylic richness. BCC and BCW values for saproxylic species were positively correlated with distance from the city centre. For non-saproxylic species, BCW values were negatively correlated with distance from adjacent areas. These results suggest that saproxylic beetles require large areas covered by forest, but are not strongly influenced by isolation, which is important for non-saproxylic species. Non-saproxylic tenebrionids have limited dispersal capabilities, which explains their sensitivity to isolation, but are generally eurytopic species frequently found even in the city centre. By contrast, most saproxylic species are able to fly, but are mainly found in peripheral areas with large and relatively well preserved forest fragments. Maintaining and possibly enhancing connectivity among green spaces is important for the conservation of non-saproxylic species, whereas preserving large forest surfaces, especially in peripheral areas, is needed for the conservation of saproxylic species.     

Influence of macroclimate and local conservation measures on taxonomic,functional, and phylogenetic diversities of saproxylic beetles and wood-inhabiting fungi

Wood-inhabiting fungi and saproxylic beetles are threatened by habitat degradation. Our understanding of the importance of macroclimate and local factors determining their taxonomic diversity has increased, but determinants of functional and phylogenetic diversity are poorly understood. We investigated assemblages of wood-inhabiting fungi and saproxylic beetles along a 1000 m elevational gradient of a temperate low mountain range. We (i) tested the relative importance of macroclimate (i.e. elevation) and local variables (microclimate, i.e. canopy closure, amount and diversity of dead wood) in determining observed and rarefied diversities and (ii) explored whether determinants of observed functional and phylogenetic diversities match those of taxonomic diversity. For both taxa, the determinants of observed phylogenetic and functional diversities largely matched those of taxonomic diversity. The diversity of wood-inhabiting fungi was predominantly determined by local variables, whereas that of saproxylic beetles was determined by both local variables and elevation. Taxonomic and phylogenetic diversities of saproxylic beetles decreased with increasing elevation, but standardized functional richness and entropy of both groups increased with increasing elevation. Diversities of wood-inhabiting fungi increased with canopy closure, while diversities of saproxylic beetles decreased with increasing canopy closure. Microclimate and dead-wood amount and diversity affected the observed and rarefied diversity of both saproxylic taxa, which justifies conservation actions that focus on attributes of dead wood and canopy cover. The contrasting responses of fungi and beetles highlight the need for amounts of diverse dead wood in the various microclimates to preserve functional and phylogenetic diversities of saproxylic organisms.     

Boxes mimicking tree hollows can help conservation of saproxylic beetles

Old hollow trees have declined in Europe and many saproxylic (i.e. wood-dwelling) invertebrates living on them are threatened. The aim of this study was to investigate to what extent artificial habitats can be exploited by saproxylic beetles. To mimic the conditions in tree hollows, we constructed wooden boxes filled with different combinations of substrates like oak saw dust, oak leaves, a dead hen (Gallus domesticus), chicken dung, lucerne flour or potatoes and placed them on tree trunks. To investigate the importance of distance from dispersal sources, we placed boxes at different distances (0–1,800 m) from three species-rich sites with high densities of hollow oaks. Over 3 years, 3,423 specimens of 105 saproxylic beetle species were caught in 47 boxes. Among beetles found in hollow oaks that were either tree-hollow species, bird nest species, or wood rot species, 70% were also found in the boxes. A dead hen added to the artificial wood mould gave a higher number of beetle specimens. The number of species associated with tree hollows in oak decreased with distance from sites with hollow oaks. In conclusion, the prospects for using artificial environments for boosting substrate availability, or to fill spatial and temporal gaps therein, for saproxylic beetles are good.     

Spatial and temporal scales relevant for conservation of dead-wood associated species: current status and perspectives

Dead wood is a key substrate for forest biodiversity, hosting a rich and often threatened biodiversity of wood-living species. However, the relationship between the occurrence of dead wood and associated species is modified by several environmental factors. Here we review the present state of knowledge on how dead wood on different spatial and temporal scales affects saproxylic biodiversity. We searched for peer-reviewed studies on saproxylic species that compared dead wood distribution on at least two spatial or temporal scales. We scanned close to 300 articles, of which 34 fit our criteria. 20 studies were directed towards the current amount of dead wood at different scale levels and how this relates to the abundance or occurrence of saproxylic species, embracing scales from 10 m to 10 km. 14 studies compared time-lagged effects of dead wood, covering time-lags from 25 years to more than 200 years. The reviewed articles focused mainly on European forest and addressed invertebrates (mostly beetles), alone or in combination with fungi (27 articles), fungi (six articles), or lichens (one article). Although the significance of dead wood for forest biodiversity is firmly established, the reviewed studies show that we still have limited knowledge of the relationship between saproxylic biodiversity and spatial and temporal scales. Based on the reviewed studies, we conclude that there is large variation in response to spatial and temporal dead wood patterns between different taxa and sub-groups. Still, several of the reviewed papers indicate that time-lagged effects deserve more attention, especially on a landscape scale and for specialized or red-listed species. Further work is required before firm management recommendations can be suggested.