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Cavities made by birds are an important microhabitat for many taxa in forests. Long-term dynamics of cavity patterns and the effect of forest management on cavities are, however, largely unknown. We studied cavity production, measured as nest cavity production rates (CPR = no. of new cavities/km2/year), of woodpeckers and tits in forests with different management intensity in southern Finland, based on a data from 37 years. Forests were divided into managed, seminatural and natural stands. The data covered 56 forest stands with the total area of 1690 ha. Stands were inventoried annually for new cavities. The total numbers of woodpecker and tit cavities were 2238 and 329, respectively. There were large differences in CPRs between forest stands with different management intensity. For woodpeckers, the CPR was highest in natural forests (5.7) and lowest in managed forests (1.5). For the tit species, the respective numbers were 0.9 and 0.3. The CPRs of different cavity-making bird species and cavity tree characteristics (e.g. tree condition and species) were consistent, suggesting that different cavity-makers benefit from similar forest and tree characteristics. The results also suggest that forests managed with currently prevailing methods limit the production of cavities. To promote cavities, the results from this and other studies suggest that managed forests should include more features of natural forests, such as more diverse tree species and within-stand structural variability distribution (tree-level heterogeneity), larger amount of decayed wood, more retention trees and snags and longer rotation periods. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

This document is a correction notice for an article titled "Translocation of deadwood in ecological compensation: A novel way to compensate for habitat loss." The correction addresses an error in one of the equations presented in the article, specifically related to the calculation of the geometrical form of a truncated cone. The correct version of the equation is provided in the correction notice. The original article has been corrected. The correction notice is authored by Olov Tranberg, Anne-Maarit Hekkala, Ola Lindroos, Therese Löfroth, Mari Jönsson, Jörgen Sjögren, and Joakim Hjältén. [Extracted from the article]

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. [ABSTRACT FROM AUTHOR]

Intensive management of boreal forests has replaced natural disturbances, leading to losses of residual deadwood and saproxylic biodiversity. Declining population sizes have been well documented for insect groups such as bark and wood-boring beetles, whilst other groups such as flat bugs (Heteroptera: Aradidae) are less well-studied and little is known about their response to forest restoration. We conducted two restoration experiments addressing this lack of knowledge. We predicted the following responses to restoration: (1) artificial gap-creation and prescribed burning will attract more flat bugs than control stands, (2) increasing pre-burning fuel loads will have a positive influence on flat bug response to fire restoration. To test these predictions we conducted two field experiments; in Sweden we compared prescribed burning of standing forest with artificial gap-creation and in Finland we compared differing levels of fuel-load prior to burning. We found partial support for our first prediction, total numbers of flat bugs collected was higher in burned forest stands compared to controls in both countries. By contrast, gap-cutting only resulted in a marginal increase in the numbers and species of flat bugs collected. We found no support for our second hypothesis. The red-listed Aradus angularis and A. laeviusculus were primarily found in burned stands. Aradus betulae, A. betulinus and A. lugubris were more frequent in burned stands than gap-cuttings and untreated controls. Our study therefore suggests that ecological restoration, especially burning, is an efficient way of attracting flat bugs to suitable habitat, we therefore recommend ecological restoration to be incorporated in boreal forest management. [ABSTRACT FROM AUTHOR]

Carbon-based secondary metabolites (CBSMs) such as tannins are assumed to function as plant defences against herbivores. CBSMs are thought to be inversely related to growth rate and nutrient concentrations because a physiological trade-off exists between cellular growth and differentiation, but CBSM concentrations can be altered by herbivory-induced changes in the trade-off. We predicted that a significant interaction exists between herbivory and growth phase, such that the effects of large herbivores (or their exclusion) on nutrient or CBSM concentrations are greatest during phases of rapid shoot or leaf growth. Leaf samples were collected during phases of different growth rate from six woody species 4 years after establishment of a large-scale long-term herbivore exclusion experiment in Kruger National Park, South Africa. Samples were analysed for N, P, condensed tannins and total phenolics. Interactions between growth phase and herbivores were rare. However, the assumption that elevated nutrients and reduced CBSMs occurs during fast phases of growth was supported by four species (consistent with the growth-differentiation balance hypothesis), but not the other two. Large herbivores generally did not affect nutrients, but CBSMs in four species were reduced by large herbivores other than elephants, while CBSMs in two species were reduced by elephants. Carbon limitation ultimately prevailed among woody plants taller than 2 m under long-term browsing. Large herbivores and plant growth phase are independent and important determinants of nutrients or CBSMs in African savannas, but the effects depend on the interacting assemblages of species, which poses challenges to the application of current general hypotheses of plant defence. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

Bioassays with a non-target slug ( Deroceras spp.) and chemical analyses were conducted using leaf tissue from already existing genetically modified insect-resistant aspen trees to examine whether genetic modifications to produce Bacillus thuringiensis (Bt) toxins could affect plant phytochemistry, which in turn might influence plant-herbivore interactions. Three major patterns emerged. First, two independent modifications for Bt resistance affected the phytochemical profiles of leaves such that both were different from the isogenic wild-type (Wt) control leaves, but also different from each other. Among the contributors to these differences are substances with a presumed involvement in resistance, such as salicortin and soluble condensed tannins. Second, bioassays with one Bt line suggest that the modification somehow affected innate resistance ('Innate' is used here in opposition to the 'acquired' Bt resistance) in ways such that slugs preferred Bt over Wt leaves. Third, the preference test suggests that the innate resistance in Bt relative to Wt plants may not be uniformly expressed throughout the whole plant and that leaf ontogeny interacts with the modification to affect resistance. This was manifested through an ontogenetic determined increase in leaf consumption that was more than four times higher in Bt compared to Wt leaves. Our result are of principal importance, as these indicate that genetic modifications can affect innate resistance and thus non-target herbivores in ways that may have commercial and/or environmental consequences. The finding of a modification-ontogeny interaction effect on innate resistance may be especially important in assessments of GM plants with a long lifespan such as trees. [ABSTRACT FROM AUTHOR]

In addition to potential benefits, biotechnology in silviculture may also be associated with environmental considerations, including effects on organisms associated with the living tree and on ecosystems and processes dependent on tree residue. We examined whether genetic modification of lignin characteristics (CAD and COMT) in Populus sp. affected leaf litter quality, the decomposition of leaf litter, and the assemblages of aquatic insects colonizing the litter in three natural streams. The decomposition of leaf litter from one of the genetically modified (GM) lines (CAD) was affected in ways that were comparable over streams and harvest dates. After 84 days in streams, CAD-litter had lost approximately 6.1% less mass than the non-GM litter. Genetic modification also affected the concentration of phenolics and carbon in the litter but this only partially explained the decomposition differences, suggesting that other factors were also involved. Insect community analyses comparing GM and non-GM litter showed no significant differences, and the two GM litters showed differences only in the 84-day litterbags. The total abundance and species richness of insects were also similar on GM and non-GM litter. The results presented here suggest that genetic modifications in trees can influence litter quality and thus have a potential to generate effects that can cross ecosystem boundaries and influence ecosystem processes not directly associated with the tree. Overall, the realized ecological effects of the GM tree varieties used here were nevertheless shown to be relatively small. [ABSTRACT FROM AUTHOR]

Non-random distribution patterns of specialized phytophagous insects on their hosts may depend on intraspecific differences in plant tissue quality, including nutrients and secondary compounds. Secondary compounds are involved in plant resistance, but are also important for the recognition and acceptability of plants as resources by specialized insects. If individuals within a plant species vary in their content of such secondary substances, there may also be qualitative differences between them. In such cases, natural selection will favor insects with the ability to distinguish and prefer the more suitable plants. In Sweden, the leaf beetle Gonioctena linnaeana Schrank (Coleoptera, Chrysomelidae) is highly specialized on one host, the native willow Salix triandra L (Salicaceae). Field observations reveal that some host plants in a population harbor many feeding larvae, causing severe defoliation, whereas neighboring plants may have few or no feeding larvae. Our hypothesis is that the distribution pattern of G. linnaeana larvae in this population results from qualitative differences between individual host plants in combination with the ability of G. linnaeana females to distinguish between plants that are suitable and not suitable for offspring performance. We examine whether larval survival differs depending on diet and whether the content of secondary chemical compounds explains female preference. Based on the higher survival rate of larvae reared on leaves from preferred hosts, we conclude that G. linnaeana females have evolved a behavior that maximizes offspring performance and thus positively affects female fitness. A chemical survey of the plants indicates that luteolin-7-glucoside and an unidentified flavonoid are important for separating the preferred from the non-preferred plants. [ABSTRACT FROM AUTHOR]

Boreal flat bugs include a high proportion of species that are considered negatively affected by forestry. Knowledge on the biology and habitat demands of individual species is generally limited. We examined the influence on flat bugs of stand-age and clear-cutting, comparing five classes of spruce stands. The five classes were: clear-cut, unthinned, and thinned (all three products of current clear-cutting forestry), mature managed and old-growth stands (these two had never been clear-cut). We also compared unburned and recently burned mature pine forest. Fire, but not stand age, had a pronounced effect on species richness and total abundance. Aradus depressus showed a significant association with older forest stands. Aradus betulae occurred only in clear-cuts and burned forest indicating that this species is favored by disturbance in general. Aradus lugubris, Aradus crenaticollis and Aradus brevicollis were found only in the burned forest. Aradus brevicollis has not previously been shown to be associated with fire. [ABSTRACT FROM AUTHOR]

Fire is important in maintaining a diversity of communities and landscapes in boreal ecosystems, but, in recent times, forest management has suppressed fires. Managers now commonly burn a proportion of clear-cuts in Sweden to conserve components of the fire-favoured fauna, but effects of these treatments on the ant fauna are unknown. We compared ant communities between burned and control clear-cuts to determine their value for this important faunal group. Ants were collected in 1995 and 1996 from 5 burned and 5 control clear-cuts in mid-boreal Sweden. We compared abundances of workers and reproductive females of common species between the treatments. Burned and control clear-cuts supported similar abundances of common species, differing only in the abundance of Leptothorax acervorum queens collected in 1995. This suggests that L. acervorum was attracted to the burning, but we did not detect an increase in the abundance of its workers the following year. The abundance of dominant red wood ants ( F. rufa group) was not correlated with that of any other species, suggesting that their patchy occurrence did not conceal differences between treatments. Our findings suggest that nests of many ant species survive low severity fires. Habitat structure is likely to be important in determining ant community structure and low severity burning in disturbed open habitats, such as the clear-cuts examined in this study, may not significantly alter habitat structure relative to unburned habitats in the short term. [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]

The distribution and ecology of insects in arctic regions are poorly known. The aim of this study was to determine the distribution of galling sawflies in the Canadian arctic and their oviposition preference. The Swedish Tundra Northwest 1999 expedition visited 17 sites in the Canadian arctic. We determined the occurrence of galling sawflies at all the sites and studied the oviposition preference of two leaf-galling sawflies, Eupontania arctica and Pontania nivalis, on Salix reticulata and S. glauca, respectively. Galling sawflies were abundant at only one site, the mainland site at Ivvavik National Park. Only a few galls in total were found at the remaining sites, suggesting that galling sawflies are rare in the higher arctic, and potential explanations for this pattern are discussed. Shoots with leaf galls were longer than shoots without galls on both S. reticulata and S. glauca. These differences could not be explained by a higher number of leaves on longer shoots. This suggests that long shoots are preferred by sawflies because of faster development and better survival of larvae on long shoots. [ABSTRACT FROM AUTHOR]

The aim of this study was to determine the inheritance pattern of phenolic secondary compounds in pure and hybrid willows and its consequences for plant resistance to leaf-feeding insects. F1, F2, and back cross hybrids along with pure species were produced by hand pollination of pure, naturally growing Salix caprea (L., Salicaceae) and S. repens (L.) plants. Leaf concentrations of condensed tannins and seven different phenolic glucosides were determined by rising butanol-HCl and HPLC analyses. Insect herbivore leaf damage was measured on the same leaves as used for chemical analyses. We found hybrids to be approximately intermediate between the parental species: S. caprea with high levels of condensed tannins and no phenolic glucosides, and S. repens with low levels of condensed tannins and high levels of phenolic glucosides. We also found a negative correlation between concentrations of condensed tannins and phenolic glucosides, suggesting a trade-off in production of these two substances. F2 hybrids and the hybrid back-crossed to S. caprea were significantly more damaged by insect herbivores than the parental species and the F1 hybrid, indicating reduced resistance and possibly a selective disadvantage for these hybrid categories. [ABSTRACT FROM AUTHOR]

The aim of this study was to assess the responses of herbivores and pathogens to hybrid plants under controlled conditions. F1 hybrids and parental species, produced by hand-pollinating willows in the field, were potted and kept in an experimental field under controlled conditions. In 1997, plant growth and survival were measured along with densities of insects and the degree of pathogen infection on the willows. The survival rate was higher for S. repens than for the hybrids and lowest for S. caprea. Densities of the sawflies Pontania pedunculi and P. brigmanii and the leaf-galling midge Iteomyia capreae were higher on hybrids and on S. caprea than on S. repens. The densities of Crepidodera fulvicornis (Chrysomelidae), chrysomelid larvae and the bud-galling midge Dasineura rosaria did not differ between any of the plant categories. Hybrids were more severely infected by rust ( Melampsora sp.) than S. caprea and the totally resistant S. repens. Densities of herbivores on hybrid willows were consistent with the dominance hypothesis (i.e. herbivore densities were similar to densities on one of the parental species) or supported the no-difference hypothesis. Furthermore, herbivore densities on hybrid plants were most similar to densities on the more susceptible parent. The breakdown in rust resistance in hybrid plants suggests that resistance traits are severely disrupted by the genetic re-arrangement in hybrids and that this increased susceptibility could select against hybridisation. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

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). [ABSTRACT FROM AUTHOR]