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.     

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.     

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.     

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.     

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.     

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).     

Influence of boreal forest succession and dead wood qualities on saproxylic beetles

1 Saproxylic insects, a functional group dominated by beetles, are dependent on dead or moribund trees as habitat elements. 2 Although there are few studies of saproxylic insects from the North American boreal zone, European studies demonstrate that forest harvest can lead to a biologically significant decrease in saproxylic beetle diversity. 3 We studied saproxylic beetles in the North American boreal mixedwood forest using flight intercept traps established on naturally dead and girdled trembling aspen and spruce trees along a successional gradient of undisturbed stands from deciduous to coniferous overstory trees. 4 Composition and diversity of beetle assemblages differed among forest successional types. 5 Snag age class was an important determinant of composition for saproxylic beetle assemblages. 6 Multivariate regression analysis of these data indicated that saproxylic beetles are responding to changes in coarse woody debris, and not to the relative densities of canopy tree species, although these variables are strongly correlated. 7 Coarse woody debris management should be a primary concern in forest management plans seeking to conserve saproxylic organisms and the critical ecosystem functions (i.e. nutrient cycling) in which they participate.     

Ecologie des coléoptères saproxyliques dans les forêts de l’étage montagnard des Alpes du nord françaises

Ecology of the saproxylic beetles in mountain forests of the northern French Alps. Both saproxylic beetles and deadwood stocks were studied in order to detect special deadwood features important for these organisms, and to better understand their ecology. A total of 37 window traps were used in ten stations covering four mountain forests in the French northern Alps. In the same time, the deadwood stocks were inventoried in the ten stations. 181 m³ of deadwood (1219 fragments) were measured and 4268 individuals belonging to 235 saproxylic species were trapped. The deadwood volume ranges from 21.1 to 233.6 m³/ha and the number of beetle species from 27 to 113. A positive relation was observed between the stand age, the whole volume of deadwood and the species richness of saproxylic beetles. However, in some deadwood rich sites the beetle diversity is not as high as expected. This was explained by the negative influence of the management history of the forest. The compositions of saproxylic communities depend also of some characteristics of the deadwood stocks. Xylophagous and zoophagous species prefer “not lying and recently dead softwood” while the mycophagous and saproxylophagous species are strongly dependent on fungi growing on hardwood, respectively under “large surfaces of barks” and in “large woody debris”. Surprisingly, the majority of the saproxylic beetles living on softwood does not depend on the most represented resinous deadwood type: the “decayed and lying softwood”.     

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.     

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.     

Vertical distribution of saproxylic beetles within snag trunks retained in plantation forests

Retention of snags (standing dead trees) is considered to have important effects on saproxylic species conservation in plantation forests because snags would provide vertically stratified deadwood habitats. However, the vertical distribution of saproxylic insects within snag trunks is still unclear. We felled 33 naturally occurring snags of Todo fir Abies sachalinensis in plantation forests and extracted insects from 99 logs sampled from three vertical positions of the snag trunks (basal stem: <2.5 m, lower trunk: 2.5–5 m and upper trunk: >5 m). The mean number of species that emerged from a single log was only 2.69, but we identified 51 morphospecies of saproxylic beetles in total. The total number of species that emerged from the basal stem (34 spp.) was greater than those that emerged from the lower trunk (25 spp.) or the upper trunk (30 spp.). However, rarefaction-extrapolation analysis did not demonstrate a significant difference in species richness among the log positions. Beetle assemblages were separated into two groups by constrained correspondence analysis; one group emerged only from lower and upper trunk logs, while another emerged mainly from basal stem logs. Additionally, vertical position had a significant effect on the distribution of the five main species. Our results show that beetle assemblages within snags in the plantation forests were highly variable, and retaining a sufficient number of high stumps may be important for saproxylic beetle conservation in plantation forests. We propose ‘retention thinning’ as an appropriate method to combine efficient timber production with biodiversity conservation in plantation forests.     

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.     

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.     

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。     

Reviewing the strength of evidence of biodiversity indicators for forest ecosystems in Europe

With a growing number of forest biodiversity indicators being applied in forest policy documents and even more being suggested by the scientific community, there is a need to evaluate, review and critically assess the strength of evidence for individual indicators, their interrelationships and potential overlaps and gaps. Biodiversity indicators proposed for forest ecosystems in Europe were reviewed with the overarching aim of providing advice on strategic selection and combination of indictors. The objectives were to (1) establish interrelationships between indicators and their indicandum (i.e. the indicated aspect of biodiversity); (2) assess the strength of scientific evidence for individual indicators; and (3) identify a set of indicators with confirmed validity for further scientific testing and inclusion in long-term reporting and decision-making regarding forest biodiversity. Ten indicator groups and 83 individual indicators were identified with application from stand scale up to landscape scale in 142 eligible scientific papers. In 62 of the 142 studies no statistical correlations between indicator(s) and indicandum were performed and 42 (out of the 62) did not even present a clear indicandum. In the remaining 80 studies, 412 correlations between indicator and indicandum were identified. However, only six correlations were assessed as being supported by strong evidence, i.e. three or more studies found statistical correlation between the indicator and indicandum, and no studies reported contradictory results. For the species richness relationships, there was strong evidence for positive correlations between deadwood volume and wood-living fungal species richness; deadwood volume and saproxylic beetle species richness; deadwood diversity and saproxylic beetle species richness; age of canopy trees and epiphytic lichen species richness. There was strong evidence for a negative correlation between tree canopy cover and spider species richness. Concerning species composition-related correlation, there was strong evidence that the species composition of epiphytic lichens changed with the age of canopy trees. These results imply that the validity of most indicators on which monitoring and conservation planning are based are weakly scientifically supported and that further validation of current biodiversity indicators for forest ecosystems is needed.     

A decade of change in the saproxylic beetle fauna of eucalypt logs in the Warra long-term log-decay experiment,Tasmania. 2. Log-size effects,succession, and the functional significance of rare species

The first decade of sequential and cyclical sampling of the saproxylic beetles of twelve freshly felled Eucalyptus obliqua logs at Warra, Tasmania has allowed comparisons between larger-diameter mature and smaller-diameter regrowth log-classes and between successive sampling cycles and years; and consideration of the interplay between these two aspects. The two log-classes support different assemblages, with the mature log-class hosting consistently more species, more unique species, and proportionally more obligately saproxylic species. Assemblages change seasonally and year-to-year, demonstrating succession. While changes in the assemblages of mature and regrowth log-classes follow similar trajectories, they remain distinct at every point in time. These differences remain apparent when considering sub-sets of the assemblages based on the rarity of the species involved, their flightedness, saproxylicity and larval feeding guild. This study suggests a need to incorporate the conservation of coarse woody debris derived from mature trees into production forestry practices. There is a particular need to devise silvicultural and/or planning systems that cater for the retention and long-term recruitment of mature trees, since these are the only source of the larger-diameter logs that were identified in this study as having particular ecological value. Through continuing the Warra long-term log-decay experiment over the next century or more, a more complete picture of the saproxylic beetle fauna will progressively emerge, together with a better understanding of the management requirements of the fauna.     

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.     

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.     

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.     

八大公山亚热带森林木质残体中大型无脊椎动物群落特征

木质残体可为大型无脊椎动物提供重要栖息地、食物等资源, 并影响其生物多样性。目前针对不同树种、径级及分解阶段的木质残体如何调控土壤大型无脊椎动物群落结构尚不清楚, 相关研究在亚热带森林地区尤为稀缺。为此, 本文选取湖南省八大公山国家级自然保护区柳杉(Cryptomeria fortunei)、亮叶水青冈(Fagus lucida)及檫木(Sassafras tzumu) 3种树种为研究对象, 每种树种分别选取两类径级(直径分别为10 ± 2 cm、4 ± 2 cm)不同分解阶段的木质残体, 对其中的大型无脊椎动物进行调查。调查于2020年10-11月完成。结果显示: 共捕获大型无脊椎动物2,558只, 隶属4门10纲23目, 不同树种的优势类群、常见类群及稀有类群均存在差异。亮叶水青冈木质残体中大型无脊椎动物个体密度显著高于柳杉和檫木。亮叶水青冈和檫木大径级木质残体中大型无脊椎动物Shannon-Wiener多样性指数显著高于小径级, 3个树种大径级木质残体中大型无脊椎动物的类群数、特有类群数均大于小径级。木质残体中大型无脊椎动物的Shannon-Wiener多样性指数、Simpson优势度指数及Pielou均匀度指数与木材密度显著负相关, 表明随着分解的进行木质残体中大型无脊椎动物群落呈明显变化趋势。木质残体的理化性质(相对含水率、全氮、全碳及碳氮比)和土壤温度、湿度与木质残体中大型无脊椎动物群落特征具有相关性。研究初步表明, 大型无脊椎动物群落特征在所选树种、径级及分解阶段木质残体中具有差异, 在亚热带森林中同时保留不同树种、不同大小径级的木质残体或有利于增加大型无脊椎动物多样性。