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。     

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.     

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.     

The importance of substrate type, shading and scorching for the attractiveness of dead wood to saproxylic beetles

Modern forestry management has reduced the amount of dead wood in forest ecosystems and this has become a serious threat to flora and fauna. Efforts are therefore being made to reverse this trend but one problem is that we still lack detailed knowledge regarding the substrate requirements of many saproxylic species. In a field experiment, conducted in three forest types (forest reserve, mature managed forest and clear-cut), we evaluated the value, from a conservation perspective, of different substrate types (logs, snags and tops) of Norway Spruce, Picea abies, and if the quality of spruce logs as saproxylic habitats can be improved by simple log treatments (scorching and shading). We collected 9982 individuals representing 262 saproxylic beetle species in window traps. Both substrate type and, to a lesser extent, log treatment had a significant effect on the abundance and species richness of saproxylic beetles attracted to the different dead wood substrates. However, more importantly, the composition of the beetle assemblages differed significantly between both substrates and log treatments. Snags, logs and tops all attracted significantly different beetle assemblages and scorched logs differed from untreated control logs. Sixteen red listed species were trapped, with the highest number (11 species) being found on scorched logs. We found strong evidence that some species preferred a specific substrate type, mainly logs, in some cases treated logs (scorched or shaded), but not snags, the substrate commonly provided for conservation purposes on e.g. clear-cuts. This stresses the importance of conducting forestry in such a way that a multitude of both forest habitats and dead wood substrates are available continuously in the forest landscape to maintain biodiversity.     

Dead trees left in clear-cuts benefit saproxylic Coleoptera adapted to natural disturbances in boreal forest

Forest management alters the pattern of forest dynamics from that in natural conditions in the boreal region. In order to examine how certain forestry measures matching natural dynamics affect forest insects, we compared assemblages of saproxylic Coleoptera on dead, standing birch trunks left behind in eight clear-cut areas with corresponding assemblages in seven mature forests in southern and eastern Finland. We used trunk-window traps for sampling. Distinct beetle assemblages were associated with the different habitats. Median numbers of species or specimens caught did not differ between closed forests and clear-cuts, but individual beetle species occurred unevenly among the habitats. Several beetle species associated with open forest habitat, e.g. burned forests or storm-damage areas, including species regarded as threatened in Finland, were found almost exclusively, in clear-cuts. Correspondingly, a number of beetle species occurring frequently in closed forests were not found in clear-cuts. We conclude that dead trunks left in the clear-cut areas may host not only generalist saproxylic species but also many beetle species specialized to warm, sun-exposed environments, and such species may not be able to survive in closed forests. Management measures matching suppressed natural disturbances are found useful in preserving diversity in managed forests.     

Enhancing bark- and wood-boring beetle colonization and survival in vertical deadwood during thinning entries

Snags are used as habitat by several organisms, including bark- and wood-boring beetles, which contribute to snag decomposition and facilitate subsequent snag colonization by other organisms. However, snags seldom occur in young managed forests. This study examines how snag characteristics and spatial arrangement affect bark- and wood-boring beetle colonization and survival in spruce plantations subjected to commercial thinning, thinning with biomass removal and thinning with snag creation where a few clumps of trees were girdled. To this end, we documented the volume of snags, their characteristics and the number of beetle emergence holes in their basal section. Beetle colonization and survival to adulthood in snags was ~33 times greater when expressed per unit area in plantations supplemented with clumps of girdled trees than in other thinning regimes where low densities of beetle emergence holes per hectare were documented. Snag diameter at breast height, snag species and thinning treatments influenced the number of emergence holes per snag. Positive spatial autocorrelation between the abundance of emergence holes per snag was detected but no other spatial effect was noticeable. This work suggests that vertical deadwood is a limiting factor for bark- and wood-boring beetle colonization within plantations and challenges current thinking about the effect of thinning on beetle communities that was developed from window trap studies. These results underline the importance of large diameter snags for beetle conservation within managed forests and demonstrate that tree girdling during thinning entries is a viable method for creating snags to enhance beetle colonization and survival.     

Assemblages of saproxylic beetles on large downed trunks of oak

Old living oaks (Quercus robur) are known as a very species‐rich habitat for saproxylic beetles, but it is less clear to what extent such veteran trees differ from an even rarer feature: downed trunks of large oaks. In this study, we set out to sample this habitat, using window traps, with two aims: (1) to describe the variation of assemblages among downed trunks of different type and (2) to compare beetles on downed oaks with data from veteran standing trees. The results showed that trunk volume and sun exposure better explained assemblages as well as species numbers on downed trunks than did decay stage. Furthermore, species classified as facultative saproxylic species showed weak or no differentiation among downed trunks. Species with different feeding habits showed no apparent differentiation among downed trunks. Furthermore, species composition on dead, downed oak trunks differed sharply from that of living, veteran oaks. Wood or bark feeders were more common on veterans than downed trunks, but there was no difference for those species feeding on fungi or those feeding on insects and their remains. In conclusion, for a successful conservation of the saproxylic beetle fauna it is important to keep downed oak trunks, and particularly large ones, in forest and pastures as they constitute a saproxylic habitat that differs from that of living trees.     

The saproxylic beetle assemblage associated with different host trees in Southwest China

Dead wood is a habitat for many insects and other small animals, some of which may be rare or endangered and in need of effective protection. In this paper, saproxylic beetle assemblages associated with different host trees in the subtropical forests in southwestern China were investigated. A total of 277 species (1 439 specimens) in 36 beetle families were collected from 117 dead wood samples, of which 101 samples were identified and respectively belonged to 12 tree genera. The number of saproxylic beetle species varied greatly among logs of different tree genera, with the highest diversity on logs of Juglans. Generally, broad‐leaved trees had a higher richness and abundance of saproxylic species than coniferous trees. Cluster analysis revealed that assemblages from broad‐leaved tree genera were generally similar (except for Betula) and assemblages from coniferous trees formed another distinct cluster. The subsequent indicator analysis proposed that there are different characteristic species for different cluster groups of host tree genera. In our study, log diameter has no positive influence on beetle species density. Conversely, comparisons of individual‐based rarefaction curves suggested that beetle species richness was highest in the small diameter class both in coniferous and broad‐leaved tree genera. With increased wood decay, proportion of habitat specialists (saproxylic beetles living on one tree genus) decreased, whereas proportion of habitat generalists (living on more than three tree genera) increased. The beetle species density was found to be higher in early stages, and decreased in later stages as well. A negative influence of altitude on saproxylic beetle species richness and abundance was detected. It was indicated that different tree genera and altitudes possibly display cross effects in modulating the altitudinal distribution and host preference of the beetles.     

Diameter of downed woody debris does matter for saproxylic beetle assemblages in temperate oak and pine forests

Deadwood is widely recognized to be an important issue for biodiversity conservation in forest ecosystems. Establishing guidelines for its management requires a better understanding of relationships between woody debris characteristics and associated species assemblages. Although deadwood diameter has been identified as an important factor predicting occurrence of many saproxylic species, the boundary between small and large diameter has not yet been precisely defined. In commercial forests, it is also of critical importance to know which diameter is large enough to host the beetle species associated with large logs in order to ensure cost-effectiveness of biodiversity conservation measures. We investigated the differences in saproxylic beetle assemblages among four different diameter classes of downed woody oak and maritime pine debris, in France. Beetles were sampled using in situ emergence traps. The diameter of deadwood pieces ranged from 1 to 40 cm. No patterns of nestedness associated with the gradient of diameter size were identified for either tree species. More indicator saproxylic species were observed in large logs and branches than in small logs. A clear distinction appeared in assemblage composition around the 5-cm diameter threshold whereas no similar pattern occurred around the 10 cm value, i.e. the classical threshold used in forestry to distinguish fine woody debris from coarse woody debris. For both tree species, the mean body length of beetles increased with the diameter of deadwood suggesting that the quantity of available resources per piece may constitute a limiting factor for large beetle species. This study confirms that not only large deadwood pieces are relevant for saproxylic biodiversity conservation but also the smallest pieces. Therefore, forest managers would be well advised to maintain a high diversity of deadwoods to maintain saproxylic biodiversity.     

The importance of forest type,tree species and wood posture to saproxylic wasp (Hymenoptera) communities in the southeastern United States

Although the forests of the southeastern United States are among the most productive and diverse in North America, information needed to develop conservation guidelines for the saproxylic (i.e., dependent on dead wood) fauna endemic to the region is lacking. Particularly little is known about the habitat associations and requirements of saproxylic parasitoids even though these organisms may be even more vulnerable than their hosts. We sampled parasitoids emerging from dead wood taken from two forest types (an upland pine-dominated forest and a lowland hardwood-dominated forest), three tree species (Liquidambar styraciflua L., Pinus taeda L., and Quercus nigra L.) and two wood postures (standing dead trees (i.e., snags) and fallen logs) in South Carolina. Parasitoid abundance did not differ between forest types or among tree species, but did differ between wood postures, being higher in snags than logs. This difference may have been due to the logs being in contact with the ground or surrounding vegetation and therefore less accessible to parasitoids. Parasitoid abundance and density decreased with height on both snags and logs. Species richness did not differ between forest types, among tree species or between wood postures. According to analysis of similarities, parasitoid communities did not differ between forest types, but did differ among tree species. The wasp communities associated with the different tree species and posture combinations were distinct. In addition, communities associated with the upper boles and crowns of snags were distinct from those occurring lower on snags. These results emphasize the importance of maintaining tree diversity in managed forests as well as retaining or creating entire snags at the time of harvest.     

Restoration of beech forest for saproxylic beetles—effects of habitat fragmentation and substrate density on species diversity and distribution

The influence of spatial location and density of beech snags on species diversity and distribution patterns of saproxylic beetles was studied in a 2,400 ha forest landscape in southern Sweden. Complete snag surveys were combined with a beetle survey using small window traps directly attached to the beech snags. The density of beech snags ≥30 cm dbh varied between one and seven snags per ha within the study area, corresponding to 1.1–5.1 m³/ha. A total of 2,610 specimens of 180 saproxylic beetles species were trapped, of which 19 species were red-listed. Within the study area, the number of red-listed and formerly red-listed species was highest around traps in old-growth stands, intermediate in managed stands contiguous with old-growth and lowest in managed stands isolated from old-growth by a two km-wide zone without beech forest. Logistic regressions revealed negative relationships between distance to old-growth forest and occurrence of eleven species, among them six red-listed or formerly red-listed species. The number of non red-listed species was not correlated with isolation from old-growth forest. The number of red-listed species also increased with snag density within 200–300 m around the traps. Our results suggest that red-listed species generally have a lower dispersal capacity than other saproxylic beetles. We conclude that retention of dead wood close to existing populations is more beneficial for red-listed species than an even distribution of snags across the forest landscape.     

Effects of landscape composition and substrate availability on saproxylic beetles in boreal forests: a study using experimental logs for monitoring assemblages

Intensive forestry practises in the Swedish landscape have led to the loss and fragmentation of stable old‐growth habitats. We investigated relationships between landscape composition at multiple scales and the composition of saproxylic beetle assemblages in nine clear‐cut, mature managed and old‐growth spruce‐dominated forest stands in the central boreal zone of Sweden. We set out fresh spruce and birch logs and created spruce snags in 2001–2002 to experimentally test the effects of coarse woody debris (CWD) type and forest management on the composition of early and late successional, and red‐listed saproxylic beetle assemblages. We examined effects of CWD availability at 100 m, and landscape composition at 1 and 10 km on saproxylic beetle abundances. Additionally, we tested whether assemblage similarity decreased with increasing distance between sites. We collected beetles from the experimental logs using eclector and window traps in four periods during 2003. CWD was measured and landscape composition data was obtained from maps of remotely sensed data. The composition of saproxylic beetles differed among different CWD substrates and between clear‐cuts and the older stand types, however differences between mature managed and old‐growth forests were significant only for red‐listed species. Assemblage similarities for red‐listed species on clear‐cuts were more different at greater distances apart, indicating that they have more localised distributions. CWD availability within 100 m of the study sites was rarely important in determining the abundance of species, suggesting that early successional saproxylic beetles can disperse further than this distance. At a larger scale, a large area of suitable stand types within both 1 and 10 km resulted in greater abundances in the study sites for several common and habitat‐specific species. The availability of suitable habitat at scales of 1–10 km is thus likely to be important in the survival of many saproxylic species in forestry‐fragmented areas.     

Effect of deadwood management on saproxylic beetle richness in the floodplain forests of northern Italy: some measures for deadwood sustainable use

Saproxylic beetles may act as bio-indicators of high-quality mature woodlands, and their conservation is strongly linked to the quality and quantity of deadwood in a biotope. We tested the effect of deadwood accumulation and habitat variables on saproxylic species richness by investigating six sampling sites under different deadwood management practices that belong to both alluvial and riparian mixed forests of the Po plain, Italy. We sampled 43 obligate saproxylic species. The main factor predicting saproxylic species richness was the amount of deadwood measured by both log diameter and volume. We found a threshold of 0.22 m diameter (confidence interval CI 0.18–0.37 m) and 32.04 m³/ha volume (CI 16.09–64.09 m³/ha) below which saproxylic beetle richness would be significantly reduced and a threshold of 35 m³/ha dead wood volume (CI 33–40 m³/ha) over which species richness increases by <5 %. The other deadwood and environmental components influenced saproxylic beetle richness to a lesser extent; some of them, however, should still be considered for proper management. Forest structure variables describing forest density such as large trees and basal areas have a negative effect on species richness. According to the results of our study, stumps and advanced decaying class are positively correlated, while small logs are negatively correlated to species richness. Thus, in extensively managed forests, the regular cutting of trees should be implemented to create artificial stumps, in order to assure a continuity of deadwood and, in the meantime, increase the number and width of openings in the forest. Moreover, prolonging rotation times can assure the presence of deadwood at intermediate/later stages of decay.     

Tree selection by the endangered beetle <Emphasis Type="Italic">Rosalia alpina</Emphasis> in a lapsed pollard beech forest

Abandoned pollard beech forests are particular habitats that may require special conservation efforts to preserve the endangered beetle Rosalia alpina, a model species whose protection may perpetuate the habitat of many other saproxylic species. Forest use can determine the tree selection and population size of R. alpina, variables previously not researched in pollard forests. Selected tree traits and population size, indicated by presence of adult specimens and recent exit holes, were determined in a pollard beech forest before the implementation of habitat manipulations targeted to preserve the species. Multivariate analysis showed tree condition (living, snag, fallen) and clearing size and aspect as influential variables, with the former explaining more variance than the latter. R. alpina positively selected pollard snags and trees located in big and dry clearings, avoiding those in small clearings or shaded areas. Snags showed most adults (90 %) and recent emergence holes (84 %). Distance to the nearest occupied tree, trunk diameter, bark thickness and presence of sap leaks had no effect on the occurrence of R. alpina. Population size was estimated in 0.1 individuals day^?1 ha^?1, and 38 % of available habitat trees showed exit holes, values lower than those observed in other European countries. These results show the need to start re-pollardings in the forest to extend pollard standing life and enhance exposure to sunlight. Pinpointing pollard snags and clearing the surrounding vegetation are recommended as the first measures to be taken in order to favor R. alpina populations on pollard stands with closed canopies.     

Effects of forest continuity on flying saproxylic beetle assemblages in small woodlots embedded in agricultural landscapes

Ancient forests are of considerable interest for strategies for biodiversity conservation. However, in European forest landscapes fragmented and harvested for a long time forest continuity might be no longer a key driver for flying organisms such as saproxylic beetles. In a study based on paired samples (n = 60 stands, p = 180 traps) of ancient and recent forests, we investigated the effects of forest continuity on saproxylic beetle assemblages in two French regions. Mean species richness was significantly related with deadwood volume in ancient forests, but not in recent forests. This loss of relationship between assemblages and their environment suggests that dispersal limitation is at work, at least for some species. Forest continuity had a significant effect on mean species richness and on the mean number of common species, but not on rare species. Forest continuity had a significant effect on assemblage composition in one out of the four cases tested. In both regions, we identified species associated with either recent or ancient forests. Finally, mean body size of species was significantly smaller in recent forests compared with ancient ones, as was their tree diameter preference, despite a higher volume of large deadwood in recent forests. These results lend support to using forest continuity as a criterion to identify sites of conservation importance, even in highly fragmented landscapes.     

Influence of snag characteristics on saproxylic beetle assemblages in a south Swedish beech forest

Effects of snag characteristics on saproxylic beetles were studied in an area of managed beech forest in southern Sweden. A snag survey was combined with a beetle survey using 30 small window traps directly attached to beech snags. The total number of species was lower in the snags which were most decayed than in the three younger decay classes, while the number of red-listed species remained the same regardless of the stage of decay. The number of fresh wood species declined and the number of rot hole species increased with increasing snag decay. The diversity of fungicolous and decayed wood species peaked at the intermediate stage of decay in the snags. CCA ordination confirmed that the stage of decay in the snags was most important for species composition, followed by sun exposure. There were no general differences in species density and composition between managed and unmanaged stands. Our study suggests that most species are able to find suitable habitat within a radius of a few kilometres and that the total amount of habitat in an unfragmented forest area is more important for species diversity than the spatial distribution of this habitat. Our data also shows that species diversity increases with habitat diversity. Snags formed from giant beech trees seem to be particularly important for rare species living in rot holes. We conclude that for a high species diversity there is a requirement for snags in different stages of decay, size and degree of sun exposure.     

Saproxylic beetles in artificially created high-stumps of spruce and birch within and outside hotspot areas

Artificially created high-stumps (snags) are created regularly during forest felling operations in Swedish coniferous production forests. The saproxylic beetle fauna in high-stumps of spruce and birch, on 20 clearcuts in Southern Sweden, were sampled by bark sieving. Ten of the clearcuts were located in hotspot areas with documented occurrence of many red-listed saproxylic beetle species. The other ten clearcuts were located in a typical production forest landscape (i.e. the matrix). Our aim was to investigate whether the benefit of creating high-stumps differs if the clearcuts is located in a hotspot area or in the matrix. In total 4,179 saproxylic beetles were found, belonging to 66 species, 9 of which were red-listed. Birch high-stumps hosted more species, on average, than spruce high-stumps. In an ordination analysis, tree species had the strongest explanatory effect among the environmental variables. No difference in beetle fauna could be found between the hotspot and matrix clearcuts, for neither birch nor spruce, according to all parameters: species numbers, species composition and red-listed species. The study does not indicate that conservation efforts in coniferous production forests should be concentrated to hotspot areas.     

Temporal variation in saproxylic beetle assemblages in a Mediterranean ecosystem

One of the main challenges in biological conservation has been to understand species distribution across space and time. Over the last decades, many diversity and conservation surveys have been conducted that have revealed that habitat heterogeneity acts as a major factor that determines saproxylic assemblages. However, temporal dynamics have been poorly studied, especially in Mediterranean forests. We analyzed saproxylic beetle distribution at inter and intra-annual scales in a “dehesa” ecosystem, which is a traditional Iberian agrosilvopastoral ecosystem that is characterized by the presence of old and scattered trees that dominate the landscape. Significant differences in effective numbers of families/species and species richness were found at the inter-annual scale, but this was not the case for composition. Temperature and relative humidity did not explain these changes which were mainly due to the presence of rare species. At the intra-annual scale, significant differences in the effective numbers of families/species, species richness and composition between seasons were found, and diversity partitioning revealed that season contributed significantly to gamma-diversity. Saproxylic beetle assemblages exhibited a marked seasonality in richness but not in abundance, with two peaks of activity, the highest between May and June, and the second between September and October. This pattern is mainly driven by the seasonality of the climate in the Mediterranean region, which influences ecosystem dynamics and imposes a marked seasonality on insect assemblages. An extended sampling period over different seasons allowed an overview of saproxylic dynamics, and revealed which families/species were restricted to particular seasons. Recognizing that seasons act as a driver in modelling saproxylic beetle assemblages might be a valuable tool in monitoring and for conservation strategies in Mediterranean forests.     

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.     

Deadwood as a surrogate for forest biodiversity: Meta-analysis of correlations between deadwood volume and species richness of saproxylic organisms

Deadwood is an important component of forest ecosystems on which many forest dwelling species depend. Deadwood volume is therefore widely used as an indicator of forest biodiversity, notably throughout Europe. However, using deadwood as an indicator has mostly been based on boreal references, and published references for temperate forests are scarce. As a result, the magnitude of the relationship between deadwood volume and species richness remains unclear for saproxylic species. We used meta-analysis to study the correlation between deadwood volume and the species richness of saproxylic beetles and fungi relative to several predictors at the forest stand level: biome, type of deadwood (log, snag, and stump) and decay class (fresh vs. decayed). We showed that the correlation between deadwood volume and species richness of saproxylic organisms was significant but moderate (r = 0.31), and that it varied only slightly between logs and snags or between decay stages. However, we found a strong biome effect: deadwood volume and species richness were more correlated in boreal forests than in temperate forests. This could be attributed both to differences in the history of forest management between biomes and to varying landscape patterns. Finally, we conclude that total deadwood volume is probably not a sufficient indicator of saproxylic biodiversity, and those additional variables (notably at the landscape level) such as type of deadwood or decay class should be integrated in deadwood monitoring. In addition, further forest research is needed to better assess the quantitative relationship between deadwood and saproxylic biodiversity, and in order to build indicators adapted to different biome contexts.