The contribution of insects to global forest deadwood decomposition.

The amount of carbon stored in deadwood is equivalent to about 8 per cent of the global forest carbon stocks1. The decomposition of deadwood is largely governed by climate2-5 with decomposer groups-such as microorganisms and insects-contributing to variations in the decomposition rates2,6,7. At the global scale, the contribution of insects to the decomposition of deadwood and carbon release remains poorly understood7. Here we present a field experiment of wood decomposition across 55 forest sites and 6 continents. We find that the deadwood decomposition rates increase with temperature, and the strongest temperature effect is found at high precipitation levels. Precipitation affects the decomposition rates negatively at low temperatures and positively at high temperatures. As a net effect-including the direct consumption by insects and indirect effects through interactions with microorganisms-insects accelerate the decomposition in tropical forests (3.9% median mass loss per year). In temperate and boreal forests, we find weak positive and negative effects with a median mass loss of 0.9 per cent and -0.1 per cent per year, respectively. Furthermore, we apply the experimentally derived decomposition function to a global map of deadwood carbon synthesized from empirical and remote-sensing data, obtaining an estimate of 10.9 ± 3.2 petagram of carbon per year released from deadwood globally, with 93 per cent originating from tropical forests. Globally, the net effect of insects may account for 29 per cent of the carbon flux from deadwood, which suggests a functional importance of insects in the decomposition of deadwood and the carbon cycle.

Trait-environment interactions of saproxylic beetles as a guide to biodiversity conservation strategies.

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.

Ambient and substrate energy influence decomposer diversity differentially across trophic levels.

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.

Uneven-aged silviculture can enhance within stand heterogeneity and beetle diversity.

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.

Translocation of deadwood in ecological compensation: A novel way to compensate for habitat loss.

Restoration of degraded habitat is frequently used in ecological compensation. However, ecological restoration suffers from innate problems of long delivery times of features shown to be good proxies for biodiversity, e.g., large dead trees. We tested a possible way to circumvent this problem; the translocation of hard-to-come deadwood substrates from an impact area to a compensation area. Following translocation, deadwood density in the compensation area was locally equivalent to the impact area, around 20 m3 ha-1, a threshold for supporting high biodiversity of rare and red-listed species. However, deadwood composition differed between the impact and compensation area, showing a need to include more deadwood types, e.g., late decomposition deadwood, in the translocation scheme. To guide future compensation efforts, the cost for translocation at different spatial scales was calculated. We conclude that translocation of deadwood could provide a cost-efficient new tool for ecological compensation/restoration but that the method needs refinement.

Evidence-based knowledge versus negotiated indicators for assessment of ecological sustainability: the Swedish Forest Stewardship Council standard as a case study.

Assessing ecological sustainability involves monitoring of indicators and comparison of their states with performance targets that are deemed sustainable. First, a normative model was developed centered on evidence-based knowledge about (a) forest composition, structure, and function at multiple scales, and (b) performance targets derived by quantifying the habitat amount in naturally dynamic forests, and as required for presence of populations of specialized focal species. Second, we compared the Forest Stewardship Council (FSC) certification standards' ecological indicators from 1998 and 2010 in Sweden to the normative model using a Specific, Measurable, Accurate, Realistic, and Timebound (SMART) indicator approach. Indicator variables and targets for riparian and aquatic ecosystems were clearly under-represented compared to terrestrial ones. FSC's ecological indicators expanded over time from composition and structure towards function, and from finer to coarser spatial scales. However, SMART indicators were few. Moreover, they poorly reflected quantitative evidence-based knowledge, a consequence of the fact that forest certification mirrors the outcome of a complex social negotiation process.

Short-term effects of continuous cover forestry on forest biomass production and biodiversity: Applying single-tree selection in forests dominated by Picea abies.

The rotation forestry system provides high biomass production, but could also have a negative impact on species sensitive to disturbance. Continuous cover forestry (CCF) could contribute to solving these conflicting goals, but its feasibility in nutrient limited boreal forests is yet unresolved. In a unique experiment, we simultaneously assessed the short-term effect of single-tree selection on both biomass production and biodiversity (vascular plants, bryophytes, wood-inhabiting fungi), and tested fertilization as a way to mediate growth-biodiversity trade-offs. We found that unharvested stands and stands subjected to single-tree selection had a similar species assemblage of vascular plants, bryophytes, and wood-inhabiting fungi. Fertilization increased growth by 37% and induced shifts in two understory species (favoring the grass Avenella flexuosa and disfavoring the bryophyte Hylocomium splendens). We conclude that single-tree selection may become a useful tool to enhance biodiversity in managed forests.

Secondary metabolites and nutrients of woody plants in relation to browsing intensity in African savannas.

Carbon-based secondary metabolites (CBSMs) are assumed to function as defences that contribute to herbivore-avoidance strategies of woody plants. Severe browsing has been reported to reduce concentrations of CBSMs and increase N concentrations in individual plants, causing heavily browsed plants to be characterised by N-rich/C-poor tissues. We hypothesised that concentrations of condensed tannins (CT) and total polyphenols (TP) should decrease, or N increase, in relation to increasing intensity of browsing, rendering severely browsed plants potentially more palatable (increased N:CT) and less N-limited (increased N:P) than lightly browsed ones. We sampled naturally browsed trees (taller than 2 m) of four abundant species in southern Kruger National Park, South Africa. Species-specific relationships between N:CT, CT, TP and P concentrations and increasing browsing intensity were detected, but N and N:P were consistently invariable. We developed a conceptual post-hoc model to explain diverse species-specific CBSM responses on the basis of relative allocation of C to total C-based defence traits (e.g. spines/thorns, tough/evergreen leaves, phenolic compounds). The model suggests that species with low allocation of C to C-based defence traits become C-limited (potentially more palatable) at higher browsing intensity than species with high allocation of C to C-based defences. The model also suggests that when N availability is high, plants become C-limited at higher browsing intensity than when N availability is low.

Saproxylic and non-saproxylic beetle assemblages in boreal spruce forests of different age and forestry intensity.

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.

Biodiversity differences between managed and unmanaged forests: meta-analysis of species richness in Europe.

Past and present pressures on forest resources have led to a drastic decrease in the surface area of unmanaged forests in Europe. Changes in forest structure, composition, and dynamics inevitably lead to changes in the biodiversity of forest-dwelling species. The possible biodiversity gains and losses due to forest management (i.e., anthropogenic pressures related to direct forest resource use), however, have never been assessed at a pan-European scale. We used meta-analysis to review 49 published papers containing 120 individual comparisons of species richness between unmanaged and managed forests throughout Europe. We explored the response of different taxonomic groups and the variability of their response with respect to time since abandonment and intensity of forest management. Species richness was slightly higher in unmanaged than in managed forests. Species dependent on forest cover continuity, deadwood, and large trees (bryophytes, lichens, fungi, saproxylic beetles) and carabids were negatively affected by forest management. In contrast, vascular plant species were favored. The response for birds was heterogeneous and probably depended more on factors such as landscape patterns. The global difference in species richness between unmanaged and managed forests increased with time since abandonment and indicated a gradual recovery of biodiversity. Clearcut forests in which the composition of tree species changed had the strongest effect on species richness, but the effects of different types of management on taxa could not be assessed in a robust way because of low numbers of replications in the management-intensity classes. Our results show that some taxa are more affected by forestry than others, but there is a need for research into poorly studied species groups in Europe and in particular locations. Our meta-analysis supports the need for a coordinated European research network to study and monitor the biodiversity of different taxa in managed and unmanaged forests.

Keeping pace with forestry: Multi-scale conservation in a changing production forest matrix.

The multi-scale approach to conserving forest biodiversity has been used in Sweden since the 1980s, a period defined by increased reserve area and conservation actions within production forests. However, two thousand forest-associated species remain on Sweden's red-list, and Sweden's 2020 goals for sustainable forests are not being met. We argue that ongoing changes in the production forest matrix require more consideration, and that multi-scale conservation must be adapted to, and integrated with, production forest development. To make this case, we summarize trends in habitat provision by Sweden's protected and production forests, and the variety of ways silviculture can affect biodiversity. We discuss how different forestry trajectories affect the type and extent of conservation approaches needed to secure biodiversity, and suggest leverage points for aiding the adoption of diversified silviculture. Sweden's long-term experience with multi-scale conservation and intensive forestry provides insights for other countries trying to conserve species within production landscapes.

Correction to: Keeping pace with forestry: Multi-scale conservation in a changing production forest matrix.

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)."

Forest management strategy affects saproxylic beetle assemblages: A comparison of even and uneven-aged silviculture using direct and indirect sampling.

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.

Habitat factors associated with bank voles (Clethrionomys glareolus) and concomitant hantavirus in northern Sweden.

Puumala virus (PUUV), genus hantavirus, causes nephropathia epidemica, a mild form of hemorrhagic fever with renal syndrome in humans. In this study, bank voles, the natural reservoir of PUUV, were captured at locations of previous human PUUV exposure and paired controls within a region of high incidence in northern Sweden. The aim of the study was to evaluate the influence of environmental factors on the abundance of bank voles and the occurrence of PUUV. The total number of voles and the number of PUUV-infected voles did not differ between locations of previous human PUUV exposure and paired controls. The number of bank voles expressing antibodies to PUUV infection increased linearly with total bank vole abundance implying density independent transmission. Using principal component and partial correlation analysis, we found that particular environmental characteristics associated with old-growth moist forests (i.e., those dominated by Alectoria spp., Picea abies, fallen wood, and Vaccinium myrtillus) were also associated with increased abundance of bank vole and hence the number of PUUV-infected bank voles, whereas there were no correlations with factors associated with dry environments (i.e., Pinus sylvestris and V. vitis-idea). This suggests that circulation and persistence of PUUV within bank vole populations was influenced by habitat factors. Future modeling of risk of exposure to hantavirus and transmission of PUUV within vole populations should include the influence of these factors.

GM trees with increased resistance to herbivores: trait efficiency and their potential to promote tree growth.

Climate change, as well as a more intensive forestry, is expected to increase the risk of damage by pests and pathogens on trees, which can already be a severe problem in tree plantations. Recent development of biotechnology theoretically allows for resistance enhancement that could help reduce these risks but we still lack a comprehensive understanding of benefits and tradeoffs with pest resistant GM (genetically modified) trees. We synthesized the current knowledge on the effectiveness of GM forest trees with increased resistance to herbivores. There is ample evidence that induction of exogenous Bacillus thuringiensis genes reduce performance of target pests whereas upregulation of endogenous resistance traits e.g., phenolics, generates variable results. Our review identified very few studies estimating the realized benefits in tree growth of GM trees in the field. This is concerning as the realized benefit with insect resistant GM plants seems to be context-dependent and likely manifested only if herbivore pressure is sufficiently high. Future studies of secondary pest species and resistance evolution in pest to GM trees should be prioritized. But most importantly we need more long-term field tests to evaluate the benefits and risks with pest resistant GM trees.

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.

Plant sex and hare feeding preferences.

To evaluate the general extent to which sex-related differences in palatability occur in boreal dioecious woody plants, males and females of five dioecious woody plant species were presented to free-ranging mountain hares (Lepus timidus) during winter. Hares strongly preferred branches from male plants when feeding on Populus tremula and Salix caprea and weakly preferred male S. pentandra. However, they did not show any sex-related preference when feeding on the other two species studied (Myrica gale and Juniperus communis). Nitrogen concentration and, to some degree, digestibility showed strong relationships with hare food preferences. By contrast, the concentration of phenolics was only weakly related to feeding preference. Phenolics could, nevertheless, still be important if only one or a few specific compounds deter hare feeding. These results indicate that sex-related differences in plant palatability in the boreal forest might be more widespread than previously believed, particularly for species of the family Salicaceae. Thus, herbivores might be responsible for the female-biased sex ratios found in willow populations in northern Scandinavia (e.g. Elmqvist et al. 1988).

Willow hybrids and herbivory: a test of hypotheses of phytophage response to hybrid plants using the generalist leaf-feeder Lochmaea caprea (Chrysomelidae).

To test hypotheses of herbivore response to hybrid plants, I determined the preference of the generalist beetle Lochmeae capreae for individuals of Salix caprea, Salix phylicifolia and F1 hybrids between these species. F1 hybrids and pure individuals were created by hand-pollination of willow individuals in the field. The preference tests were performed in 1994 and 1995 and in both years there was a tendency for higher leaf area consumption from hybrids than from the parental species. However, statistically significant differences in consumption were only found in 1995 between hybrids and S. phylicifolia. The result from this study therefore lends most support to the dominance hypothesis: hybrid susceptibility was similar to that of one of the parental species. This is one of the first strict preference tests showing that a generalist herbivore can detect differences in palatability between hybrids and parental species. Furthermore, the experimental design used in the study minimized environmental variation. The results should therefore primarily reflect genetically based quality differences between hybrids and parental species.