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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., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2024 The Authors. Published by Elsevier Ltd.. All rights reserved.)

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.

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., (Copyright © 2017 Elsevier Ltd. All rights reserved.)

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.

Genetic modifications of trees may provide many benefits, e.g. increase production, and mitigate climate change and herbivore impacts on forests. However, genetic modifications sometimes result in unintended effects on innate traits involved in plant-herbivore interactions. The importance of intentional changes in plant defence relative to unintentional changes and the natural variation among clones used in forestry has not been evaluated. By a combination of biochemical measurements and bioassays we investigated if insect feeding on GM aspens is more affected by intentional (induction Bt toxins) than of unintentional, non-target changes or clonal differences in innate plant defence. We used two hybrid wildtype clones (Populus tremula x P. tremuloides and Populus tremula x P. alba) of aspen that have been genetically modified for 1) insect resistance (two Bt lines) or 2) reduced lignin properties (two lines COMT and CAD), respectively. Our measurements of biochemical properties suggest that unintended changes by GM modifications (occurring due to events in the transformation process) in innate plant defence (phenolic compounds) were generally smaller but fundamentally different than differences seen among different wildtype clones (e.g. quantitative and qualitative, respectively). However, neither clonal differences between the two wildtype clones nor unintended changes in phytochemistry influenced consumption by the leaf beetle (Phratora vitellinae). By contrast, Bt induction had a strong direct intended effect as well as a post experiment effect on leaf beetle consumption. The latter suggested lasting reduction of beetle fitness following Bt exposure that is likely due to intestinal damage suffered by the initial Bt exposure. We conclude that Bt induction clearly have intended effects on a target species. Furthermore, the effect of unintended changes in innate plant defence traits, when they occur, are context dependent and have in comparison to Bt induction probably less pronounced effect on targeted herbivores.

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.

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.

One main aim with genetic modification (GM) of trees is to produce plants that are resistant to various types of pests. The effectiveness of GM-introduced toxins against specific pest species on trees has been shown in the laboratory. However, few attempts have been made to determine if the production of these toxins and reduced herbivory will translate into increased tree productivity. We established an experiment with two lines of potted aspens (Populus tremula×Populus tremuloides) which express Bt (Bacillus thuringiensis) toxins and the isogenic wildtype (Wt) in the lab. The goal was to explore how experimentally controlled levels of a targeted leaf beetle Phratora vitellinae (Coleoptera; Chrysomelidae) influenced leaf damage severity, leaf beetle performance and the growth of aspen. Four patterns emerged. Firstly, we found clear evidence that Bt toxins reduce leaf damage. The damage on the Bt lines was significantly lower than for the Wt line in high and low herbivory treatment, respectively. Secondly, Bt toxins had a significant negative effect on leaf beetle survival. Thirdly, the significant decrease in height of the Wt line with increasing herbivory and the relative increase in height of one of the Bt lines compared with the Wt line in the presence of herbivores suggest that this also might translate into increased biomass production of Bt trees. This realized benefit was context-dependent and is likely to be manifested only if herbivore pressure is sufficiently high. However, these herbivore induced patterns did not translate into significant affect on biomass, instead one Bt line overall produced less biomass than the Wt. Fourthly, compiled results suggest that the growth reduction in one Bt line as indicated here is likely due to events in the transformation process and that a hypothesized cost of producing Bt toxins is of subordinate significance.

Current clear-cutting forestry practices affect many boreal organisms negatively, and those dependent on dead wood (saproxylics) are considered as particularly vulnerable. The succession of species assemblages in managed forest habitats regenerating after clear-cutting is, however, poorly known. We compared beetle assemblages in three successional stages of managed boreal spruce forests established after clear-cutting and two types of older spruce forests that had not been clear-cut. We also assessed whether saproxylic and non-saproxylic beetle assemblages show similar biodiversity patterns among these forest types. Beetles were collected in window traps in nine study areas, each encompassing a protected old-growth forest (mean forest age approximately 160 years, mean dead wood volume 34 m3/ha), an unprotected mature forest (approximately 120 years old, 15 m3/ha), a middle-aged commercially thinned forest (53 years old, 3 m3/ha), a young unthinned forest (30 years old, 4 m3/ha), and a clearcut (5-7 years after harvest, 11 m3/ha). Saproxylic beetles, in particular red-listed species, were more abundant and more species rich in older forest types, whereas no significant differences among forest types in these variables were detected for non-saproxylics. The saproxylic assemblages were clearly differentiated; with increasing forest age, assemblage compositions gradually became more similar to those of protected old-growth forests, but the assemblage composition in thinned forests could not be statistically distinguished from those of the two oldest forest types. Many saproxylic beetles adapted to late-successional stages were present in thinned middle-aged forests but absent from younger unthinned forests. In contrast, non-saproxylics were generally more evenly distributed among the five forest types, and the assemblages were mainly differentiated between clearcuts and forested habitats. The saproxylic beetle assemblages of unprotected mature forests were very similar to those of protected old-growth forests. This indicates a relatively high conservation value of mature boreal forests currently subjected to clear-cutting and raises the question of whether future mature forests will have the same qualities. Our results suggest a high beetle conservation potential of developing managed forests, provided that sufficient amounts and qualities of dead wood are made available (e.g., during thinning operations). Confirming studies of beetle reproduction in dead wood introduced during thinning are, however, lacking.

Decline in biodiversity have increased the interest in alternative forest management approaches. Uneven-aged silviculture has been proposed as a mean to maintain continuity of forest canopy cover, mimic small-scale disturbances and provide a stratified forest structure similar to that of old-growth forests and therefore better maintain species associated with unmanaged forest. We used a large-scale chronosequence study spanning 50 years to study beetle diversity in uneven-aged silviculture compared with both short-term impacts and the longer-term legacy of even-aged silviculture. We compared: (1) even-aged recently clear-felled stand, (2) even-aged recently thinned stand, (3) uneven-aged stands subjected to selective felling with (4) uneven-aged reference stands to evaluate whether abundance, species richness and composition of beetles (Coleoptera) were affected differently by even-aged than by uneven-aged management. We collected 15,147 beetles from 461 species using flight interception traps in 30 stands. Beetle composition was maintained in uneven-aged managed stands; composition did not differ from unmanaged reference stands, the exception being cambium consumers. Both even-aged silviculture treatments (clear-felling and thinning) had different beetle composition compared to the reference stands, indicating that assemblages had yet to recover even 50 years into the rotation. However, beetle composition did not differ between uneven-aged managed and thinned stands. The result supports our prediction that uneven-aged silviculture better maintains beetles assemblages associated with semi-natural mature forest than even-aged silviculture. The greater temporal continuity in selectively felled stands could benefit species dependent of mature or old growth forest since some of the needed habitat qualities are continuously available. Uneven-aged silviculture could therefore serve as an important tool for landscape planning to benefit biodiversity and thus help fulfil environmental commitments. However, uneven-aged silviculture may still alter the forest and should therefore be viewed as an alternative to even-aged silviculture, rather than to set-asides. [ABSTRACT FROM AUTHOR]

Abstract: Studies on the effects of stump harvesting on forest biodiversity are scarce and studies on long-term effects are until now non-existent. We evaluated such long-term effects by sampling beetles at 14 clear-cuts with and 14 clear-cuts without stump harvesting; harvesting had been done 21–28years before this study. By using window traps, we collected 6429 individuals belonging to 432 species in 55 taxonomic families. To control for potentially confounding effects of among-site variations in landscape setting we also assessed forest age and volume of deciduous trees within the forest surrounding each site. The long-term effects from harvesting on beetle abundance, species richness and species composition was generally small in comparison to the influence of the characteristics of the surrounding forests. The species richness of the beetle family Latridiidae and the functional group fungivores appeared, however, to be negatively affected by the previous stump harvesting, while several other groups showed strong associations to the characteristics of the surrounding forest. We found little support for considerable long-term effects of stump harvesting on beetles flying in the stands. Long-term effects of stump harvesting at the landscape scale accumulated from harvest of many localities may still be severe, and should be the subject of future studies. [Copyright &y& Elsevier]

Abstract: The aim of this study was to evaluate how future harvesting of spruce low stumps as a renewable energy source, might influence the saproxylic (wood living) beetle fauna. Species richness, abundance and assemblages of saproxylic beetles were compared on low stumps, high stumps and logs of spruce on 10 clear-cuts in northern Sweden. The sampling was conducted with emergence traps and a total of 929 individuals and 120 species were collected. No significant differences in beetle abundance or species richness were detected between substrate types. However, there were clear differences in assemblage composition between all substrate types. Our results suggest that low stumps created at final felling support as many species and individuals per volume dead wood as other dead wood substrates and that they are therefore overlooked as substrates for wood living organisms. Consequently, harvesting of low stumps might have negative effects on the diversity of saproxylic beetles, as up to 80% of the dead wood remaining on clear-cuts is in the form of low stumps. Future studies should evaluate if compensatory measures can be performed to minimize negative impacts. [ABSTRACT FROM AUTHOR]