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.     

Forest restoration by burning and gap cutting of voluntary set-asides yield distinct immediate effects on saproxylic beetles

Today, the importance of restoring natural forest disturbance regimes and habitat structures for biodiversity is widely recognized. We evaluated the immediate effects of two restoration methods on wood-inhabiting (saproxylic) beetles in boreal forest voluntary set-asides. We used a before-after control-impact experimental set-up in 15 set-asides; each assigned to one of three treatments: (1) restoration burning, (2) gap cutting and (3) no-treatment reference stands. Before treatment, abundance, species richness and assemblage composition of trapped beetles did not differ significantly among treatments. Burning resulted in a significant change in assemblage composition and increased species richness and abundance compared to reference stands. As predicted, saproxylic species known to be fire favoured increased dramatically after burning. The immediate response shows that, initially, fire favoured species are attracted from the surrounding landscape and not produced on site. Gap cutting increased the abundance of cambium consumers but had no significant effect on total species richness or assemblage composition of saproxylic beetles. The stronger effect of burning compared to gap cutting on saproxylic assemblages is probably due to the very specific conditions created by fires that attracts many disturbance-dependent species, but that at the same time disfavour some disturbance-sensitive species. By contrast, gap cutting maintained assemblage composition, increased abundances and is likely to increase species richness in the years to follow, due to elevated level of dead wood. The restoration methods applied in this study may prove particularly useful, partly because of positive effect on saproxylic beetles, but also due to the cost-efficiency of the measures; the voluntary set-asides were already established and the restoration costs fully covered by revenue from the extracted timber.     

Wood-Inhabiting Beetles in Low Stumps,High Stumps and Logs on Boreal Clear-Cuts: Implications for Dead Wood Management

The increasing demand for biofuels from logging residues require serious attention on the importance of dead wood substrates on clear-cuts for the many forestry-intolerant saproxylic (wood-inhabiting) species. In particular, the emerging harvest of low stumps motivates further study of these substrates. On ten clear-cuts we compared the species richness, abundance and species composition of saproxylic beetles hatching from four to nine year old low stumps, high stumps and logs of Norway spruce. By using emergence traps we collected a total of 2,670 saproxylic beetles among 195 species during the summers of 2006, 2007 and 2009. We found that the species assemblages differed significantly between high stumps and logs all three years. The species assemblages of low stumps, on the other hand, were intermediate to those found in logs and high stumps. There were also significant difference in species richness between the three examined years, and we found significant effect of substrate type on richness of predators and fungivores. As shown in previous studies of low stumps on clear-cuts they can sustain large numbers of different saproxylic beetles, including red-listed species. Our study does, in addition to this fact, highlight a possible problem in creating just one type of substrate as a tool for conservation in forestry. Species assemblages in high stumps did not differ significantly from those found in low stumps. Instead logs, which constitute a scarcer substrate type on clear-cuts, provided habitat for a more distinct assemblage of saproxylic species than high stumps. It can therefore be questioned whether high stumps are an optimal tool for nature conservation in clear-cutting forestry. Our results also indicate that low stumps constitute an equally important substrate as high stumps and logs, and we therefore suggest that stump harvesting is done after carefully evaluating measures to provide habitat for saproxylic organisms.     

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.     

Changes in the dominant assembly mechanism drive species loss caused by declining resources

The species–energy hypothesis predicts that more productive areas support higher species richness. Conversely, when resources are reduced, species richness is reduced. Empirical tests of whether extinctions are predominantly caused by environmental constraints or competitive exclusion are lacking. We experimentally reduced dead wood to c. 15% of the initial amount after a major windstorm and examined changes in assembly mechanisms by combining trait‐based and evolutionary species dissimilarities of eight taxonomic groups, differing in their dependence on dead wood (saproxylic/non‐saproxylic). Species richness and assembly mechanisms of non‐saproxylic taxa remained largely unaffected by removal of dead wood. By contrast, extinctions of saproxylic species were caused by reversing the predominant assembly mechanisms (e.g. increasing importance of competitive exclusion for communities assembled through environmental filtering or vice versa). We found no evidence for an intensification of the predominant assembly mechanism (e.g. competitive exclusion becoming stronger in a competitively structured community).     

Landscape context affects the success of habitat restoration: large‐scale colonization patterns of saproxylic and fire‐associated species in boreal forests

Aim Restoration of habitats may be used as a conservation tool when ecosystems have lost their natural structure, dynamics or functioning over large areas. Controlled and planned use of fire could be an effective way to restore habitats of many threatened species in boreal forests where fire suppression has been effective. We asked whether the large‐scale landscape context affects the occurrence of rare and threatened species in forest habitats that have been burned to restore their fire‐related structures. Location Boreal forests in southern Finland. Methods We designed a large‐scale field experiment that included nine Pinus sylvestris forests (5–10 ha each) in southern Finland. Sites were located in two regions: (1) in eastern region with shorter management history and (2) in western region where intensive forestry has continued longer. We evaluated whether restoration of dead/burned wood is beneficial for rare and conservation‐dependent species and measured the recovery of pyrophilous and red‐listed insects (beetles and flatbugs) in burned forests, using standardized sampling effort. Altogether, 956 individuals of 29 red‐listed and pyrophilous species were sampled. Results Rare species colonized areas quickly, but there was a clear difference in species richness between the regions. The eastern forests harboured higher species richness after restoration. In these sites, the average species richness was 13.7 species per site, whereas in western forests it was 5.0 species per site. Similar pattern was also observed in subgroups: the corresponding numbers for pyrophilous species were 9.7 vs. 3.8, for red‐listed 8.7 vs. 2.3 and for red‐listed pyrophiles 4.7 vs. 1.2. Main conclusions Introducing fire back to boreal forests can aid in the recovery of rare species, but the landscape context considerably affects the success of restoring species. If restored habitats are located in landscapes that have lost their natural properties long ago, the success of restoration seems to be more challenging than in landscapes where habitats have been modified more recently.     

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.     

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.     

Some of the rarest European saproxylic beetles are common in the wilderness of Northern Mongolia

Natural landscapes characterized by heavy disturbance regimes were displaced in Europe by managed cultural landscapes over the past centuries. The associated loss of biological legacies, such as dead or dying trees, has exposed numerous saproxylic species to high risks of extinction. In contrast, extensive wilderness forests in Northern Mongolia have been sustained owing to significant cultural differences. Here we used saproxylic beetle abundance data gathered during two sampling campaigns in the Mongolian taiga to address whether (1) the saproxylic beetle fauna of the Mongolian taiga is comparable to that of European boreal forests, (2) fires are a natural disturbance regime, indicated by the occurrence of many pyrophilous species, and (3) species rare in Europe are also rare in the Northern Mongolian wilderness. Of 191 saproxylic beetle species identified, 150 (79 %) were also found in Europe. The high number of pyrophilous beetle species (20) indicated that natural species communities are well adapted to this disturbance regime. The species rarity in Germany was significantly positively correlated with the species rarity in Finland, but the species rarity in these two countries was negatively correlated with that in the Mongolian wilderness. Our results indicated that wilderness areas with natural disturbances provide biological legacies important for rare species. Therefore, exploitation of the unique, remaining natural landscapes of the Palaearctic wilderness areas should be stopped. Moreover, we urge conservationists to expand controlled burning for restoration at relict sites of rare boreal species also outside Fennoscandia.     

Patterns of saproxylic beetle succession in loblolly pine

• 1 Patterns of insect succession in dead wood remain unclear, particularly beyond the first several years of decay. In the present study, saproxylic beetles were sampled from loblolly pine (Pinus taeda L.) logs aged between 1 month and 9 years old using both emergence traps attached to logs in the field and rearing bags in the laboratory.
• 2 Species richness peaked within the first year as a result of a diverse assemblage of bark beetles, wood‐borers and predators associated with young logs. After the phloem phase, there were no significant differences in species richness among decay classes.
• 3 Beetle communities differed significantly among decay classes, with 25 and seven species being significantly associated with young and old logs, respectively.

     

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.     

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.     

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.     

Effects of Clearfell Harvesting on Lucanid Beetles (Coleoptera: Lucanidae) in Wet and Dry Sclerophyll Forests in Tasmania

The effects of commercial forestry harvest and regeneration practices (clearfelling and slash-burning) on the lucanid fauna of the wet sclerophyll forests of southern Tasmania and the dry sclerophyll forests of eastern Tasmania were examined using pitfall catches. Lucanids are saproxylic beetles, dependent on dead, moribund and decaying wood. Samples taken from old-growth forest and from a chronosequence of sites regenerating after logging, in each forest type, were used to compare the species richness and abundance of the lucanid assemblages. In both forest types, species richness and abundance was highest in the youngest regeneration sites (1–3 year), reflecting the species richness of the original and adjacent unlogged forest, lowest in the older (20–25 year) sites, and variable in the old-growth sites. TWINSPAN cluster analysis showed no clear distinction between regeneration and old-growth forest. The post-harvest slash and stump residue provided an important refugium and initial habitat, but our research indicates that some species may not maintain populations in the long term. Our results suggest that most species of lucanids will find a continuous supply of suitable habitat only in old-growth forests; and such species may become less common as clearfell harvesting leads to a replacement of heterogeneous old-growth forest with single-aged monospecific stands. Continuity of supply of wood in all decay stages, the maintenance of sufficient source areas, and biological connectivity between old-growth stands to enable dispersal, are all likely to be essential to maintain lucanid beetle community integrity. If similar principles apply to other saproxylic species of invertebrate, then clearfelling and slash-burning may cause a gradual extinction of an important element of the forest biota.     

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.     

Species richness of phytophagous beetles in the tropical tree Brosimum utile (Moraceae): the effects of sampling strategy and the problem of tourists

Abstract.  1. Phytophagous beetles on six mature living trees and two dead trees of Brosimum utile (Moraceae) were surveyed during 1 year in a tropical wet forest in Panama. The dead trees were surveyed both as suspended in the canopy and after falling down to the understorey.
2. Canopy access was provided by a construction crane and sampling was performed by beating and hand-collecting. The same amount of time was spent on each tree in order to standardise sampling effort. A list of all species associated with the tree is presented.
3. A total of 3009 individuals representing 364 species were collected. Tourists were excluded from the analyses by recording host associations directly and by probability assessments of host associations based on abundance categories. A total of 2603 individuals and 244 species were associated with the tree. The proportion of tourists in the trees increased with sample size.
4. A single mature living tree had on average 58.5 ± 6.5 species. The local species richness of B. utile was estimated as 2.5 times higher than in a single mature tree; however, a substantial increase in species richness was attained when dead wood habitats were included. Saproxylic species made up 82% of the total material.
5. The investigated habitat types of B. utile constituted distinct, complementary species assemblages. Similarity between saproxylic species of dead suspended wood and dead understorey wood of the same tree was 0.2 (Morisita–Horn index), confirming a prominent vertical stratification among this guild.     

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

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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.     

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.     

Inter‐assemblage facilitation: the functional diversity of cavity‐producing beetles drives the size diversity of cavity‐nesting bees

Inter‐specific interactions are important drivers and maintainers of biodiversity. Compared to trophic and competitive interactions, the role of non‐trophic facilitation among species has received less attention. Cavity‐nesting bees nest in old beetle borings in dead wood, with restricted diameters corresponding to the body size of the bee species. The aim of this study was to test the hypothesis that the functional diversity of cavity‐producing wood boring beetles ‐ in terms of cavity diameters ‐ drives the size diversity of cavity‐nesting bees. The invertebrate communities were sampled in 30 sites, located in forested landscapes along an elevational gradient. We regressed the species richness and abundance of cavity nesting bees against the species richness and abundance of wood boring beetles, non‐wood boring beetles and elevation. The proportion of cavity nesting bees in bee species assemblage was regressed against the species richness and abundance of wood boring beetles. We also tested the relationships between the size diversity of cavity nesting bees and wood boring beetles. The species richness and abundance of cavity nesting bees increased with the species richness and abundance of wood boring beetles. No such relationship was found for non‐wood boring beetles. The abundance of wood boring beetles was also related to an increased proportion of cavity nesting bee individuals. Moreover, the size diversity of cavity‐nesting bees increased with the functional diversity of wood boring beetles. Specifically, the mean and dispersion of bee body sizes increased with the functional dispersion of large wood boring beetles. The positive relationships between cavity producing bees and cavity nesting bees suggest that non‐trophic facilitative interactions between species assemblages play important roles in organizing bee species assemblages. Considering a community‐wide approach may therefore be required if we are to successfully understand and conserve wild bee species assemblages in forested landscapes.