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Conservation of biodiversity requires in-depth knowledge of trait-environment interactions to understand the influence the environment has on species assemblages. Saproxylic beetles exhibit a wide range of traits and functions in the forest ecosystems. Understanding their responses to surrounding environment thus improves our capacity to identify habitats that should be restored or protected. We investigated potential interactions between ecological traits in saproxylic beetles (feeding guilds and habitat preferences) and environmental variables (deadwood, type and age of surrounding forest). We sampled beetles from 78 plots containing newly created high stumps of Scots pine and Silver birch in boreal forest landscapes in Sweden for three consecutive years. Using a model based approach, our aim was to explore potential interactions between ecological traits and the surrounding environment at close and distant scale (20 m and 500 m radius). We found that broadleaf-preferring beetle species are positively associated with the local broadleaf-originated deadwood and broadleaf-rich forests in the surrounding landscapes. Conifer-preferring species are positively associated with the local amount of coniferous deadwood and young and old forests in the surrounding landscape. Fungivorous and predatory beetles are positively associated with old forests in the surrounding landscapes. Our results indicate that both local amounts of deadwood and types of forests in the landscape are important in shaping saproxylic beetle communities. We particularly highlight the need to increase deadwood amounts of various qualities in the landscape, exempt older forests from production and to increase broadleaf-rich habitats in order to meet different beetle species' habitat requirements. Trait responses among saproxylic beetles provide insights into the significance of broadleaf forest and dead wood as essential attributes in boreal forest restoration, which helps conservation planning and management in forest landscapes. • Trait – environment interactions of saproxylic beetles are investigated. • Broadleaf-preferring beetles thrive with local broadleaved deadwood. • Broadleaf-rich forests contribute to the abundance of broadleaf-preferring beetles. • Fungivores and predators positively interact with old forests in the landscape. • Deadwood increase and a 'broadleafication' of boreal forests need to be implemented. [ABSTRACT FROM AUTHOR]

Management of forest for wood production has altered ecosystem structures and processes and led to habitat loss and species extinctions, worldwide. Deadwood is a key resource supporting forest biodiversity, and commonly declines following forest management. However, different forest management methods affect dead wood differently. For example, uneven-aged silviculture maintains an age-stratified forest with ongoing dead wood production, while even-aged silviculture breaks forest continuity, leading to long periods without large trees. We asked how deadwood-dependent beetles respond to different silvicultural practices and if their responses depend on deadwood volume, and beetles preference for decay stages of deadwood. We compared beetle assemblages in five boreal forest types with different management strategies: clearcutting and thinning (both representing even-aged silviculture), selective felling (representing uneven-aged silviculture), reference and old growth forest (both uneven-aged controls without a recent history [~50 years] of management, but the latter with high conservation values). We collected beetles using window traps and by sieving the bark from experimental logs (bolts). Beetle assemblages on clear-cuts differed from all other stand types, regardless of trapping method or decay stage preference. Thinning differed from reference stands, indicating incomplete recovery after clear-cutting, while selective felling differed only from clear-cuts. In contrast to our predictions, early and late successional species responded similarly to different silvicultural practices. However, there were indications of marginal assemblage differences both between thinned stands and selective felling and between thinned and old growth stands (p = 0.10). The stand volume of early decay stage wood influenced assemblage composition of early, but not late successional species. Uneven-aged silviculture maintained species assemblages similar to those of the reference and old growth stands and might therefore be a better management option when considering biodiversity conservation. [ABSTRACT FROM AUTHOR]

Uneven-aged silviculture may better maintain species assemblages associated with old-growth forests than clear felling in part due to habitat heterogeneity created by maintaining standing retention strips adjacent to harvest trails. Retention strips and harvest trails created at the time of tree removal will likely have different microclimate and may harbor different assemblages. In some cases, the resultant stand heterogeneity associated with uneven-aged silviculture may be similar to natural small-scale disturbances. For beetles, increased light and temperature as well as potential access to young vegetation and deadwood substrates present in harvset trails may harbor beetle assemblages similar to those found in natural gaps. We sampled saproxylic beetles using flight intercept traps placed in harvest corridors and retention strips in 9 replicated uneven-aged spruce stands in central Sweden. We compared abundance, species richness and composition between harvest corridors and retention strips using generalized linear models, rarefaction, permutational multivariate analysis of variance and indicator species analysis. Canopy openness doubled, mean temperature and variability in daily temperature increased and humidity decreased on harvest trails. Beetle richness and abundance were greater in harvests trails than in retention strips and the beetle species composition differed significantly between habitats. Twenty-five species were associated with harvest trails, including three old-growth specialists such as Agathidium discoideum (Erichson), currently red-listed. We observed only one species, Xylechinus pilosus (Ratzeburg) that strongly favored retention strips. Harvest trails foster both open habitat species and old-growth species while retention strips harbored forest interior specialists. The combination of closed canopy, stratified forest in the retention strips and gap-like conditions on the harvest trails thus increases overall species richness and maintains more diverse assemblages at the stand level than would otherwise be seen in less heterogeneous stand types. This suggests that uneven-aged silviculture may provide added conservation benefits for both open habitat and old-growth specialists than silvicultural approaches that reduce stand heterogeneity. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

The species‐energy hypothesis predicts increasing biodiversity with increasing energy in ecosystems. Proxies for energy availability are often grouped into ambient energy (i.e., solar radiation) and substrate energy (i.e., non‐structural carbohydrates or nutritional content). The relative importance of substrate energy is thought to decrease with increasing trophic level from primary consumers to predators, with reciprocal effects of ambient energy. Yet, empirical tests are lacking. We compiled data on 332,557 deadwood‐inhabiting beetles of 901 species reared from wood of 49 tree species across Europe. Using host‐phylogeny‐controlled models, we show that the relative importance of substrate energy versus ambient energy decreases with increasing trophic levels: the diversity of zoophagous and mycetophagous beetles was determined by ambient energy, while non‐structural carbohydrate content in woody tissues determined that of xylophagous beetles. Our study thus overall supports the species‐energy hypothesis and specifies that the relative importance of ambient temperature increases with increasing trophic level with opposite effects for substrate energy. [ABSTRACT FROM AUTHOR]

Topographic heterogeneity causes gradients in altitude, potential solar radiation and soil humidity on a range of scales from micro- to macro-habitat. These gradients are important determinants for the distribution of many organisms but have been largely neglected in studies of species associated with dead wood, a group of great conservation concern. In this study, we evaluated the effects of topography-related gradients (altitude, potential solar radiation and soil humidity) and habitat characteristics (bark cover, ground contact and dead wood volume) on saproxylic (wood-inhabiting) beetle assemblages. We sampled boreal saproxylic beetles hatching from 750 experimentally exposed spruce and birch logs in a regional scale field experiment including 10 landscapes and spanning gradients of altitude (range 85–510 m asl), potential solar radiation (based on slope and slope aspect and highest in equator-facing slopes, PADIR, range 0.24–0.71) and soil humidity (humidity index, range 2.3–3.1). The logs were placed in north Swedish forests and clear-cuts and beetles were sampled with emergence traps the 4th summer after the logs were introduced. Saproxylic assemblage composition varied considerably in response to altitude, potential solar radiation and soil humidity. The response was evident in both forests and clear-cuts and for both birch and spruce logs. Species density and abundance responses differed among trophic groups. For birch logs, fungivore species density increased significantly with increased potential solar radiation. For spruce logs, altitude affected total species density and density of cambium consumers positively and abundance of fungivores negatively, suggesting a delay in succession due to slower decomposition at higher altitudes. In addition to the topography-related gradients, ground contact and bark cover of logs as well as the availability of dead wood in the vicinity influenced the beetle assemblages. Our results clearly show that topography-related gradients affect assemblage composition of saproxylic beetles. These factors should be considered in the management of saproxylic biodiversity. To ensure the safeguarding of intact saproxylic assemblages it is important that the full range of dead wood habitats is conserved. Topographic heterogeneity may provide opportunities for landscape scale survival in response to rapid climate change but there is also a risk that some niches will disappear. Some of the negative effects of climate change could be mitigated by allocating set asides and restoration efforts in areas where gradient in altitude, solar radiation and soil humidity are particularly well developed over short distances. [ABSTRACT FROM AUTHOR]

1. Suppression of wildfires in boreal landscapes has become widespread and has seriously affected many fire favoured species. However, little is known about the response of organism assemblages to large wildfires in regions with a long history of effective fire suppression, such as Scandinavia. 2. We studied the short-term effects of a >1600 ha wildfire on beetle assemblages in northern Sweden. The first summer after fire, beetles were sampled in 12 sites using 36 large window traps, half in old pine forest stands in the burned area and half in similar, but unburned control stands. The entire beetle assemblage and eight subgroups were analysed: saproxylics, non-saproxylics, moderately fire favoured, strongly fire favoured, fungivores, predators, cambium consumers and red-listed species. 3. Species composition differed markedly between burned and unburned forests in all nine groups. Furthermore, beetle abundance was higher in the burned area for the entire assemblage and for saproxylics, both groups of fire favoured species, predators and cambium consumers. Species number was higher only for non-saproxylics, strongly fire favoured species and cambium consumers. 4. Our results show that wildfire has rapid and strong effects on a wide range of beetles. However, we only trapped two individuals of fire-dependent beetles, which may suggest a lack of such species in the region, possibly due to >100 years of fire suppression. At the regional scale, the studied wildfire may potentially increase the abundance of these beetles after a longer period of reproduction in the burned area. [ABSTRACT FROM AUTHOR]

In managed landscapes, conservation planning requires effective methods to identify high-biodiversity areas. The objective of this study was to evaluate the potential of airborne laser scanning (ALS) and forest estimates derived from satellite images extracted at two spatial scales for predicting the stand-scale abundance and species richness of birds and beetles in a managed boreal forest landscape. Multiple regression models based on forest data from a 50-m radius (i.e., corresponding to a homogenous forest stand) had better explanatory power than those based on a 200-m radius (i.e., including also parts of adjacent stands). Bird abundance and species richness were best explained by the ALS variables "maximum vegetation height" and "vegetation cover between 0.5 and 3 m" (both positive). Flying beetle abundance and species richness, as well as epigaeic (i.e., ground-living) beetle richness were best explained by a model including the ALS variable "maximum vegetation height" (positive) and the satellite-derived variable "proportion of pine" (negative). Epigaeic beetle abundance was best explained by "maximum vegetation height" at 50 m (positive) and "stem volume" at 200 m (positive). Our results show that forest estimates derived from satellite images and ALS data provide complementary information for explaining forest biodiversity patterns. We conclude that these types of remote sensing data may provide an efficient tool for conservation planning in managed boreal landscapes. [ABSTRACT FROM AUTHOR]

In the original published article, the sentence "Nevertheless, semi-natural forest remnants continue to be harvested and fragmented (Svensson et al. 2018; Jonsson et al. 2019), and over 2000 forest-associated species (of 15 000 assessed) are listed as threatened on Sweden's red-list, largely represented by macro-fungi, beetles, lichens and butterflies (Sandström 2015)."under the section Introduction was incorrect. The correct version of the sentence is "Nevertheless, semi-natural forest remnants continue to be harvested and fragmented (Svensson et al. 2018; Jonsson et al. 2019), and approximately 2000 forest-associated species (of 15 000 assessed) are on Sweden's red-list, largely represented by macro-fungi, beetles, lichens and butterflies (Sandström 2015)." [ABSTRACT FROM AUTHOR]

Species diversity commonly increases with succession and this relationship is an important justification for conserving large areas of old-growth habitats. However, species with different ecological roles respond differently to succession. We examined the relationship between a range of diversity measures and time since disturbance for boreal forest beetles collected over a 285 year forest chronosequence. We compared responses of “functional” groups related to threat status, dependence on dead wood habitats, diet and the type of trap in which they were collected (indicative of the breadth of ecologies of species). We examined fits of commonly used rank-abundance models for each age class and traditional and derived diversity indices. Rank abundance distributions were closest to the Zipf-Mandelbrot distribution, suggesting little role for competition in structuring most assemblages. Diversity measures for most functional groups increased with succession, but differences in slopes were common. Evenness declined with succession; more so for red-listed species than common species. Saproxylic species increased in diversity with succession while non-saproxylic species did not. Slopes for fungivores were steeper than other diet groups, while detritivores were not strongly affected by succession. Species trapped using emergence traps (log specialists) responded more weakly to succession than those trapped using flight intercept traps (representing a broader set of ecologies). Species associated with microhabitats that accumulate with succession (fungi and dead wood) thus showed the strongest diversity responses to succession. These clear differences between functional group responses to forest succession should be considered in planning landscapes for optimum conservation value, particularly functional resilience. [ABSTRACT FROM AUTHOR]