Learning from a “benign neglect strategy” in a national park: Response of saproxylic beetles to dead wood accumulation

Increasing demands for firewood owing to rising energy costs have accelerated discussions about the amount of dead wood needed for conservation. A sharp increase in dead wood caused by bark beetles in a German national park provides lessons for management of commercial and protected forests. We investigated the effects of dead wood due to bark beetle infestation as well as tree senility on abundance and richness of saproxylic species of beetles. Increasing amounts of spruce dead wood and opening of the canopy by bark beetles had positive effects on the abundance of host-generalist, conifer-specialist, and red-listed saproxylic beetles. Broadleaf specialists were positively associated with the amount of broadleaf dead wood and negatively associated with canopy openness. Gradient analysis of beetle assemblages revealed two major environmental axes: canopy openness and amount of dead wood. We found a threshold for community divergence at a canopy openness of 23% (confidence interval CI: 11-49) and at an amount of dead wood of 64 m³ ha⁻¹ (CI: 35-160). Critically endangered species served as indicators of dense and open forests, but only when the amount of dead wood was high. Our results suggest that, to maintain saproxylic beetle assemblages, the amount of dead wood in commercial montane forests (at present ≈15 m³ ha⁻¹) needs to be tripled, with a focus on broadleaf wood in dense stands and spruce wood in open stands. For large protected areas in Europe, our data suggest that bark beetle infestation and senescence without active forest management improves habitat conditions for saproxylic beetles.     

Openness in management: Hands-off vs partial cutting in conservation forests, and the response of beetles

The appropriate management of forest reserves is debated; two major alternatives are succession to ‘wild’ state, or management to produce semi-open stands. For temperate conservation stands, there are no strong experiments replicated at landscape level. In each of 22 forests rich in oaks (Quercus spp.) in Sweden, we set up a closed-canopy wild plot (1 ha), and a cutting plot (1 ha) to produce semi-open conditions, studying them before and after cutting. About 25% of the tree basal area was cut (large trees and dead wood retained) and harvested as bio-fuel, a CO₂ - neutral energy source. We examined the response of beetles and trapped 59,000 individuals (1174 species; 100 red-listed species). For both the guild of herbivorous beetles (222 species) and of saproxylic beetles connected to oak wood (267 species), species richness increased by about 35% in the harvested plots, relative to the wild reference plots. Species composition within the groups changed, though not strongly. Thirteen saproxylic species of 50 analysed, and three herbivores of 12 increased in cutting plots. For red-listed saproxylic beetles, species richness did not change significantly. Regression analyses suggest that more open cutting plots disfavour the red-listed beetles of this forest type. Thus, partial cutting increased species diversity of two beetle groups, probably due to changed microclimate and increase in herbaceous plants, but some red-listed saproxylic beetles may be disfavoured. A hands-off alternative may through storms and other disturbances produce open patches, more dead wood, and favour some species. Combinations of these alternatives, carefully planned at the landscape level, need to be considered.     

Conservation-oriented forestry and early successional saproxylic beetles: Responses of functional groups to manipulated dead wood substrates

New conservation-oriented forestry aims to maintain intact populations of forest organisms by improving the conservation value of managed forests and providing protected areas. We tested the conservation value of treatments of dead wood for assemblages of early successional saproxylic beetles. In nine areas in northern Sweden, we selected one clear-cut, one mature managed forest and one reserve. In 2001-2002, we placed three blocks of spruce logs, each containing control, burned and shaded logs and a high stump (“snag”) at each site. Saproxylic beetles emerging from the dead wood were collected using emergence traps and beetles flying close to it were collected using flight-intercept traps. After one year of exposure, assemblage composition was examined, with respect to nutritionally-defined functional groups, red-listed species and fire-favoured species. Experimental snags were most complementary to control logs, supporting different assemblages of cambium consumers and fungivores and supporting more red-listed individuals. Burned logs supported depauperate assemblages, particularly with respect to cambium consumers, while shading of logs affected assemblages of fungivores, but only on clear-cuts. Despite containing less dead wood, managed forests provided valuable habitat, supporting similar assemblages of saproxylic beetles to reserves. Most functional groups were less abundant on clear-cuts than in older forests, but fire-favoured species were more common on clear-cuts, suggesting that clear-cuts may support assemblages of species associated with natural disturbances, if suitable substrates are available. Utilization of logs by saproxylic beetles changes over time, so long-term monitoring of our experimental logs will determine their lifetime conservation value.     

Use of remnant boreal forest habitats by saproxylic beetle assemblages in even-aged managed landscapes

Saproxylic Coleoptera are diverse insects that depend on dead wood in some or all of their life stages. In even-aged boreal forest management, remnant habitats left as strips and patches contain most of the dead wood available in managed landscapes and are expected to act as refuges for mature forest species during the regeneration phase. However, use of remnant habitats by the saproxylic fauna has rarely been investigated. Our objective was to characterize the saproxylic beetle assemblages using clearcuts and forest remnants in western Québec, Canada, and to explore the effects of forest remnant stand characteristics on saproxylic beetle assemblages. We sampled both beetle adults and larvae, using Lindgren funnels and snag dissection, in five habitat locations (clearcuts, forest interiors of large patches, edges of large patches, small patches and cut-block separators) from three distinct landscapes. Adult saproxylic beetles (all feeding guilds combined) had significantly higher species richness and catch rates in small patches compared to forest interiors of large patches; the phloeophagous/xylophagous group had significantly higher species richness only. Small patches, cut-block separators and edges of large patches also had the highest snag density and basal area, increasing habitat for many saproxylic beetles. No significant differences in density of saproxylic larvae were found between habitat patches, but snag dissection nevertheless suggests that snags in forest remnants are used by comparable densities of insects. Saproxylic beetles appear to readily use habitat remnants in even-aged managed landscapes suggesting that forest remnants can insure the local persistence of these species, at least in the timeframe investigated in our study.     

At which spatial and temporal scales does landscape context affect local density of Red Data Book and Indicator species?

The landscape context is crucial for forest conservation in regions where the natural forest is fragmented. The focus of practical conservation is currently shifting from local stands to a landscape perspective, but few studies have tested the relative effect of different spatial and temporal scales for occurrence and persistence of species of conservation concern. We studied Red Data Book and Indicator species (the latter proposed to indicate presence of Red Data Book species) of vascular plants, lichens, bryophytes and wood-inhabiting fungi in 22 old temperate broadleaved forests in southern Sweden. We analysed at which scales these species respond to habitat proportion in surrounding landscape. The proportion of suitable habitat was measured at two temporal scales (present-day and historic) and at two spatial scales (about 0-1 km and 1-5 km of study sites). Local density of Red Data Book species increased with increasing proportion of suitable habitat in the current landscape (within 1-5 km of study sites) while Indicator species were unaffected. The response to landscape differed between organism groups. Vascular plants (near significantly) and wood-inhabiting fungi showed a time delay of 120 years in their response, indicating a possible regional extinction debt. An appropriate minimum landscape scale for conservation of Red Data Book species in temperate broadleaved forests in Sweden seems to be about 5 km (radius), but smaller landscapes may be important for vascular plants and wood-inhabiting fungi of conservation concern. In addition, restoration is urgent to counteract the effect of time delays in species responses to recent habitat loss.     

Three-dimensional partitioning of diversity informs state-wide strategies for the conservation of saproxylic beetles

Global conventions on biological diversity force governments to develop region-wide conservation strategies. Such strategies are difficult to design for all taxa because little is known about the important spatial scales. Here we applied additive partitioning of the diversity of saproxylic beetles in Bavarian forests in Southern Germany using a nested hierarchical design of five increasingly broader spatial levels: trap, strata, forest stand, forest site, and ecoregion. We consistently found a significantly higher percentage than expected by chance of between ecoregion diversity and significantly lower α diversity within traps. A significant proportion of β diversity was also found between stands. Analysis of species represented by <0.005% of all specimens in our samples and of species classified as threatened revealed similar results. Critical spatial scales for threatened species encompassed the critical levels of common species. Within habitat substratum guilds, the proportion of β diversity increased from species associated with fresh wood to those associated with rotten wood to those associated with fungi. Our results suggest that the most effective way to ensure saproxylic beetle diversity in a state-wide strategy is to add new conservation sites within different ecoregions and to establish new conservation areas in additional forest stands, rather than to enlarge reserves. Our findings further suggest that monitoring of saproxylic beetle diversity on a broader scale in European temperate forests can be restricted to “monitoring species”, i.e. a subgroup of families easy to identify, and that canopy sampling can be neglected without a substantial loss of information.     

Determining the spatial scale for conservation purposes - an example with grizzly bears

This study suggests procedures for determining the spatial scale for conservation guidelines for animals, giving an illustration with an analysis of grizzly bear habitat selection. Bear densities were sampled by identifying hairs at bait stations in British Columbia. Habitat variables were measured using remote sensing. Spatial scale was changed by varying the window size over which the variables were averaged. First, the spatial pattern of bears was studied, measuring the patchiness in bear densities at a variety of spatial scales, by calculating the correlation in bear densities between adjacent windows. This was repeated for the habitat variables. Finally, the overall interaction between bears and habitats was analysed, measuring the strength of habitat selection at different spatial scales. There are three domains of scale: at 2-4 km, bears and habitats are patchy, at 5-10 km, bears select for habitats, and at 40+ km, habitats are patchy and bears select for habitats. At scales of 40+ km, bears selected for: (i) higher slopes, or (ii) higher slopes, and some combination of more avalanche chutes, fewer roads and trees, higher elevations, and less logged land. Within 15 km areas, bears selected for 6 km areas that are either at higher elevations, or at higher elevations and had fewer trees. The relationship of conservation guidelines at different spatial scales should be determined by measuring and comparing hierarchical to non-hierarchical selection. The scales that bears select for habitats roughly correspond to the scales used in present grizzly bear conservation plans in British Columbia.     

Conservation ecology of boreal polypores: A review

Here we quantitatively summarize the conservation ecology of one group of dead-wood-dependent organisms, the polyporous fungi, in boreal Europe. At the substrate scale, the decay stage is the strongest determinant of species richness, with large (>20 cm diameter) downed logs hosting more species than other dead-wood types. At the stand scale, the amount of dead wood is the strongest determinant of polypore species richness; the minimum average amount of dead wood for the occurrence of rare polypores appears to be 20–40 m³/ha. Species-area analysis shows that in mature boreal forests species accumulation levels off at around 20–30 ha. This leads us to suggest a heuristic 20/20/20 rule of thumb: a 20 ha stand, with an average of 20 m³/ha of dead wood of which many are logs >20 cm, is likely to be the minimum for the ecologically justified conservation of polypore diversity at the stand scale in boreal Europe. Equally crucial for polypore diversity, however, is the current and historic extent of suitable habitats at the landscape scale. The time lag between the isolation of a habitat patch and the new equilibrium in the number or occurrence of species seems to be around 100–150 years, indicating that an extinction debt is likely to exist in recently isolated fragments. Only a few studies have addressed the ecological efficiency of the new, biodiversity-oriented forest management tools (retention trees, woodland key habitats). Despite this it seems that the traditional large conservation areas are the most effective means of polypore conservation.     

Effects of forest management on carabid beetles in Belgium: implications for biodiversity conservation

We sampled the carabid beetles in 22 forests managed by six different silvicultural systems, defined by treatment and tree species composition: even-aged conifer, even-aged beech, even-aged oak, uneven-aged conifer, uneven-aged beech and group mixed (beech + conifer). In each of these forests, we placed pitfall traps in young, medium-aged and mature stands (3 stages). We evaluated the effect of treatment, tree species composition, silvicultural system, stage and habitat type (silvicultural system + stage) on indicators of community conservation value and ecological structure. The species composition and the ecological structure of carabid beetles of the managed stands were then compared to that of nine unmanaged stands (without tree exploitation). In the managed forests, species richness was highest in large young stands (3-10 years old) and in forests managed by even-aged systems (with large clear-cuts), mainly due to eurytopic and opportunist carabid species with high dispersal abilities. Oak and beech, uneven-aged, and mature stands were mainly inhabited by typical forest species, and even-aged conifer stands mainly by ubiquitous species. Several typical forest species recorded in unmanaged stands were lacking from the managed forests. Large scale clear-cutting allows open-habitat species to enter the forest, which increases the species richness at a landscape level but can disfavour typical forest species by competition. Long rotations should be implemented and more areas left unmanaged in Belgium, in order to help typical forest species to re-colonise managed forests.     

Assessing the extinction vulnerability of wood-inhabiting fungal species in fragmented northern Swedish boreal forests

Fragmentation of old-growth forests and greatly reduced amounts of coarse dead wood in managed forests threat the persistence of many saproxylic species in boreal Fennoscandia. Individual old-growth forest remnants may lose species over time as they pay off their extinction debt. We tested this by comparing the observed site occupancy of individual wood-inhabiting fungal species in isolated old-growth stands (i.e. woodland key habitats; WKHs) with statistical predictions of their occupancy assuming potential extinction debt had already been paid off. The occupancy of species was analysed in two sets of WKHs differing in time since isolation (i.e. recent and old isolates).Few species occurred more frequently than expected in WKHs. However, patterns across species and across all WKHs masked important differences among species in their risk of facing future extinction. The site occupancy decreased significantly between recent and old isolates for a group of annual, red-listed specialist fungal species, suggesting that an extinction debt in WKHs may exist among specific species confined to coarse dead wood and old-growth forest habitat. Generalist species that also occur in the surrounding matrix showed no negative trends, or actually increased in site occupancy, making future extinctions less likely. Thus, continuing loss of threatened species are likely if not preservation of WKHs are combined with other conservation efforts in managed forest landscapes. Natural forest landscapes may serve as important references when aiming to identify species in risk of future extinction but more detailed knowledge about the biology of the most vulnerable species is also required.     

Conserving ground-dwelling beetles in an endangered woodland community: Multi-scale habitat effects on assemblage diversity

Patterns of biodiversity are influenced by habitat features at multiple spatial scales, yet few studies have used a multi-scale approach to examine ground-dwelling beetle diversity patterns. We trapped and quantified ground-dwelling beetle assemblages at two spatial scales: (1) microhabitat elements, represented by open ground, ground under trees and ground next to logs and (2) macrohabitat, represented by three vegetation types in a box-gum grassy woodland in south-eastern Australia. Species richness and evenness was highest at samples from under trees and lowest at samples in the open. At the macrohabitat scale, species richness and evenness did not differ among vegetation types. Assemblage composition was significantly different between trees, logs and open elements. Assemblage composition was different only between vegetation types with contrasting high and low shrub cover. Estimation of true species richness indicated assemblages at logs may have a higher number of species compared to trees and open elements, and implied greater spatial heterogeneity in assemblages at logs. Significant spatial autocorrelation in beetle assemblages was detected for logs at up to 400 m, but not for ground under trees or in the open. In agreement with previous studies, a mix of vegetation types at the macrohabitat scale is important for beetle conservation. Assemblage composition, however, appears to be more closely linked with habitat elements at the microhabitat scale, where logs support a high diversity of beetle species. This strongly supports the idea that restoring logs to box-gum grassy woodlands would be useful for increasing beetle species richness and assemblage heterogeneity.     

Sampling saproxylic beetles: lessons from a 10-year monitoring study

We sampled saproxylic beetles using trunk window traps in two birch-dominated forests in Finland during 1990-1999. The sampling scheme, 10 traps attached to living fruiting bodies of Fomes fomentarius growing on dead birches, remained unchanged in both forests throughout the study period. Beetles belonging to 32 selected families were identified every year, whereas all species were identified during the last 4 years. Total number of identified individuals was 40,294 and number of species 583, of which 258 were saproxylic (dependent on dead wood). Species richness of rare and threatened saproxylic beetles in the samples varied a lot between the years and did not fluctuate synchronously between the forests. Variation between years was smaller when all saproxylic species were pooled together and some abundant species fluctuated synchronously in the two forests. Similarity indices and DCA-ordination did not generally suggest decreasing similarity between samples with increasing temporal isolation. Incidence-based similarities of common saproxylic species within and between forests and years were high (means 0.7-0.8), whereas those of rare saproxylics were roughly 50% smaller and much more variable. More than 75% of the common saproxylic species found during the entire 10-year period were detected already after 3 years of sampling but accumulation of rare and threatened species was much slower. Our results suggest that: (1) occurrence of rare and threatened species in samples is much less predictable than that of common species and, e.g. reserve selection based on rarities should be made cautiously; (2) estimation of total number of threatened species in a forest is very difficult, because such species accumulate slowly in the samples; (3) samples from different years can be comparable in certain cases.     

Application of central-place foraging theory shows the importance of Mediterranean dehesas for the conservation of the cinereous vulture, Aegypius monachus

The dehesa (oak woodland) is an extensive agro-pastoral ecosystem characteristic of the Western Mediterranean countries which is suffering a great transformation process since 1950. Although its distribution largely overlaps with several endangered species, there is scarce information on how they use this human-transformed habitat. We studied the foraging habitat selection of one of them, the cinereous vulture Aegypius monachus. We radio-tracked 14 cinereous vultures in one of the largest European colonies from 1998 to 2000. Used and available habitats were compared at two scales using compositional analysis. Moreover, we developed a distance-based GLMM for assessing habitat selection in this central-place forager species, by taking into account the spatial distribution of habitat patches in relation to the location of the colony. Home ranges overlapped over a total surface of 592,527 ha around the colony, and both individual home ranges and travel foraging distances (mean 27.86 km, maximum 86 km) were larger during the breeding season. All cinereous vultures avoided agricultural lands within their home ranges throughout the year. Habitat use in relation to the distance to the colony pointed out that dehesas were positively selected in spite of being on average far away from the colony than other habitats, a result that was consistent among individuals and seasons. The cinereous vulture thus depends for its conservation not only on the protection of breeding areas, as has been so far considered, but also on the maintenance of well-conserved dehesas close to the colonies. Preserving the cinereous vultures could contribute to the economic sustainability of dehesas by attracting PAC funds for their traditional low-intensity exploitation. Although other species may also benefit from this study since cinereous vulture could be a “flagship” for the large-scale conservation of Mediterranean oak woodlands and associated biodiversity, more fine local management guidelines should be performed on the basis of studies on more sensitive species.     

Vascular plants as a surrogate species group in complementary site selection for bryophytes, macrolichens, spiders, carabids, staphylinids, snails, and wood living polypore fungi in a northern forest

Vascular plants were investigated as a potential surrogate group in complementary small scale site selection, such as woodland key habitats in Scandinavia. We compared the response of vascular plants to environmental gradients to that of seven other plant, fungal and animal groups within a forest reserve in western Norway using data from 59 plots of 0.25 ha. We also examined whether the spatial changes in species (beta-2 index) of vascular plants matched that of the other groups. All seven groups responded to the same gradients in nutrient richness and humidity as the vascular plants. Furthermore, changes in species composition of vascular plants were reflected in comparable degrees of change among the “target“ groups. The lower the degree of change in species composition between plots in the “target“ groups relative to that of vascular plants, the higher the percentage “target“ species encompassed in a complementary selection of sites based on vascular plants. We conclude that in practical site selection of small scale sites of conservation value, such as woodland key habitats, vascular plants may be used in combination with an inventory of important habitats for rare and/or redlisted forest species, such as dead wood, old trees, deciduous trees, and cliffs.     

Diversity of dead wood inhabiting fungi and bryophytes in semi-natural beech forests in Europe

Saproxylic organisms are among the most threatened species in Europe and constitute a major conservation problem because they depend on the most important forestry product - dead wood. Diversity of fungal and bryophyte communities occurring on dead beech trees was analyzed in five European countries (Slovenia, Hungary, The Netherlands, Belgium and Denmark) considering tree level species richness (TLSR), country level species richness (CLSR), frequency distributions of species, occurrence of threatened species and relations between TLSR and decay stage, tree size and countries. Altogether 1009 trees were inventoried in 19 beech dominated forest reserves.The number of fungi in the full dataset was approximately three times larger (456 versus 161 species) and the proportion of low frequent species was higher than among bryophytes. The species richness of bryophytes and fungi was significantly different among countries considering both TLSR and CLSR. In addition the diversity patterns deviated considerably between the two groups of organisms. Slovenian sites appeared to be biodiversity hotspots for bryophytes characterized by high TLSR and CLSR and a high fraction of threatened species. Hungarian sites had somewhat lower bryophyte diversity, while the Atlantic region had deteriorated assemblages. Fungal species richness was very high in Denmark, but the Hungarian and Slovenian sites were richer in threatened and low frequency species. Tree size was better able to explain variation in TLSR in both organism groups than decay stage. TLSR was found to vary significantly between countries but the difference was most considerable in the case of bryophytes.The diversity patterns of both organism groups along the investigated geographical gradient appear to be influenced by both climatic and management related factors (forest history, dead wood availability and continuity, habitat fragmentation). There is no doubt that an increase in the abundance of dead wood in European beech forests will benefit diversity of saproxylic fungi and bryophytes, especially if a continuous presence of large diameter logs are secured within individual stands.     

The effects of forest patch size and matrix type on changes in carabid beetle assemblages in an urbanized landscape

In this study we compared ground beetles (Carabidae) from a range of different forest fragments along an urbanization gradient in Brussels, Belgium. We address the following questions: (i) How does the degree of urbanization in the surrounding habitat affect forest beetles, and does it interact with the effects of patch size and distance to forest edge? (ii) Do these factors have a different effect at the level of individual species, habitat affinity groups or total community? During 2002 we sampled 13 forest plots in 10 forest patches, ranging in size from 5.27 to 4383 ha. The beetles were captured using transects of pitfall traps from the edge to a distance of 100 m into each woodland and identified to species level. Effects of urbanization, forest size and forest edge were evaluated on total species number, abundance and habitat affinity groups and ten abundant, widespread model carabid species. Overall, the effects of urbanization, forest size and edge effects slightly influenced total species richness and abundance but appeared to have a major effect on ground beetle assemblages through species specific responses. More urbanized sites had significantly fewer forest specialists and more generalist species. Large forest fragments were favoured by forest specialist species while generalist species and species frequently associated with forest (forest generalists) dominated the smaller forests. Forest edges mainly harboured generalist species while forest specialist species were more frequent into the forests if the forest patches were large enough, otherwise they disappeared due to the destruction or impoverishment of their habitat. Our results show the importance of differentiating between habitat affinity, especially habitat generalists versus specialists, the latter having a higher value in nature conservation, and merely the quantity of species represented in human-dominated areas.     

Effects of landscape-scale broadleaved woodland configuration and extent on roost location for six bat species across the UK

Although forest fragmentation can greatly affect biodiversity, responses to landscape-scale measures of woodland configuration in Europe have been examined for only a limited range of taxa. Almost all European bat species utilise woodland, however little is known about how they are affected by the spatial arrangement of woodland patches. Here we quantify landscape structure surrounding 1129 roosts of six bat species and a corresponding number of control locations across the UK, to examine associations between roost location and landscape composition, woodland proximity and the size of the nearest broadleaved woodland patch. Analyses are performed at two spatial scales: within 1 km of the roost and within a radius equivalent to the colony home-range (3–7 km). For four species, models at the 1 km scale were better able to predict roost occurrence than those at the home-range scale, although this difference was only significant for Pipistrellus pipistrellus. For all species roost location was positively associated with either the extent or proximity of broadleaved woodland, with the greatest effect of increasing woodland extent seen between 0% and 20% woodland cover. P. pipistrellus, Pipistrellus pygmaeus, Rhinolophus hipposideros, Eptesicus serotinus and Myotis nattereri all selected roosts closer to broadleaved woodland than expected by chance, with 90% of roosts located within 440 m of broadleaved woodland. Roost location was not affected by the size of the nearest broadleaved patch (patches ranged from 0.06–2798 ha ± 126 SD). These findings suggest that the bat species assessed here will benefit from the creation of an extensive network of woodland patches, including small patches, in landscapes with little existing woodland cover.     

Hedgerows in an agri-natural landscape: Potential habitat value for native bees

Recent declines in North American honeybee populations have highlighted the importance of native bee conservation, and the need for research on the ecological requirements of native bees in farmland. In this study, we investigated the value of hedgerows as foraging habitat for native bees in mosaics of small-scale agriculture and natural vegetation in two riparian landscapes in southeast Arizona, USA. In the summers of 2002 and 2003, we surveyed bees and flowers in four habitats: hedgerows, agricultural fields, woodlots, and native woodland. We asked: (1) How do hedgerows compare to other available habitats in bee abundance and species richness? (2) How does bee species composition in hedgerows compare to species composition in agricultural fields and woodland? (3) How do flower resources in hedgerows compare to those in fields and woodland?We found that hedgerows were attractive foraging habitat for native bees, especially in early summer, when hedgerows tended to have higher species richness than other agricultural or natural habitats. Cumulative species richness was highest in agricultural fields, although cumulative species richness did not significantly differ among fields, hedgerows, and woodland. While bee faunas overlapped among habitats, bee assemblages in hedgerows were more similar to those in woodland than to those in fields. The hedgerow herbaceous flora was roughly intermediate to that of fields and woodland; hedgerows also supported high densities of woodland-characteristic shrubs. These flowering shrubs were important in attracting bees that were otherwise uncommon in the landscape, including some species that are potentially valuable pollinators of agricultural crops.     

Parasitoids (Hymenoptera, Ichneumonoidea) of Saproxylic beetles are affected by forest successional stage and dead wood characteristics in boreal spruce forest

The habitat requirements and effects of forest management on insects belonging to higher trophic levels are relatively unknown in forest ecosystems. We tested the effect of forest successional stage and dead wood characteristics on the saproxylic parasitoid (Hymenoptera, Ichneumonoidea) assemblage in boreal spruce-dominated forests in northern Sweden. Within each of nine areas, we selected three sites with different management histories: (1) a clear-cut (2) a mature managed forest and (3) an old-growth forest. Parasitoids were collected in 2003 using eclector traps mounted on fresh logs, which were either untreated (control), burned, inoculated with fungi, or naturally shaded, and on artificially-created snags.Both forest type and dead wood characteristics had a significant effect on parasitoid assemblages. Grouped idiobionts and some species, such as Bracon obscurator and Ontsira antica, preferred clear-cuts, while others, such as Cosmophorus regius (Hym., Braconidae) and other koinobionts, were associated with older successional forest stages. No single dead wood substrate was sufficient to support the entire community of parasitoids in any forest type, even when the regular host was present. In particular, snags hosted a different assemblage of species from other types of dead wood, with parasitoids of Tetropium spp. such as Rhimphoctona spp. (Hym., Ichneumonidae) and Helconidea dentator (Hym., Braconidae) being abundant. These results indicate that a diversity of dead wood habitats is necessary to support complete assemblages of beetle-associated parasitoids from early successional stages of dead wood and that parasitoids may be more sensitive to habitat change than their hosts.     

The effects of windthrow on forest insect communities: a literature review

This paper reviews the effects that windstorm-induced drastic changes (micro-climate, soil, vegetation, and ground structural heterogeneity) have on forest insect communities. In the current context of shady and CWD-deprived managed forests, windthrow gaps act as regional biodiversity hotspots by maintaining habitat continuity in a mosaic landscape, and by facilitating the breeding and population growth of clearing specialists and saproxylic species. Windthrow gaps are dead-wood islands where forest protection and habitat conservation goals may stand against each other. Besides the quantitative effect of dead wood on bark beetle outbreaks and saproxylic diversity, the latter is favoured by key dead-wood micro-habitats such as large logs, snags and sun-exposed coarse woody debris. The role of natural enemies and sanitation operations in regulating pest outbreaks is discussed. Heterogeneous openings provide many micro-habitats favouring flower-visiting insects, phytophages on saplings, on fallen tree crowns, and on diverse understory flora, as well as ground insects on specific micro-sites.