What window traps can tell us: effect of placement, forest openness and beetle reproduction in retention trees

The use of flight interception traps (window traps) has been criticized for catching too many species without affinity to the immediate surroundings. We study aspen retention trees left for conservation reasons in a boreal forest in south-eastern Norway, and investigate how placement of window traps affects the beetle species assemblage, abundance of habitat specialists, saproxylic species and vagrant species. We also test the correlation between beetle trappings and beetle exit holes in wood. The window traps clearly responded to the immediate surroundings of the trap. Traps located on tree trunks had a different species assemblage than traps hanging freely. Traps mounted on trees caught more aspen associated beetles and less vagrant species than their free-hanging counterparts. The differences were larger when trees were dead than alive. There was a significant positive correlation between presence of individuals in the trunk-window traps and presence of exit holes for three aspen associated species. Thus, the trapping results indicated successful reproduction, showing that aspen associated beetles are not only attracted to but also utilise aspen retention trees/high stumps left in clear-cuts. This indicates that this conservation measure in forest management can have positive, alleviating effects concerning the dead wood deficit in managed boreal forest.     

Tree selection by the endangered beetle <Emphasis Type="Italic">Rosalia alpina</Emphasis> in a lapsed pollard beech forest

Abandoned pollard beech forests are particular habitats that may require special conservation efforts to preserve the endangered beetle Rosalia alpina, a model species whose protection may perpetuate the habitat of many other saproxylic species. Forest use can determine the tree selection and population size of R. alpina, variables previously not researched in pollard forests. Selected tree traits and population size, indicated by presence of adult specimens and recent exit holes, were determined in a pollard beech forest before the implementation of habitat manipulations targeted to preserve the species. Multivariate analysis showed tree condition (living, snag, fallen) and clearing size and aspect as influential variables, with the former explaining more variance than the latter. R. alpina positively selected pollard snags and trees located in big and dry clearings, avoiding those in small clearings or shaded areas. Snags showed most adults (90 %) and recent emergence holes (84 %). Distance to the nearest occupied tree, trunk diameter, bark thickness and presence of sap leaks had no effect on the occurrence of R. alpina. Population size was estimated in 0.1 individuals day^?1 ha^?1, and 38 % of available habitat trees showed exit holes, values lower than those observed in other European countries. These results show the need to start re-pollardings in the forest to extend pollard standing life and enhance exposure to sunlight. Pinpointing pollard snags and clearing the surrounding vegetation are recommended as the first measures to be taken in order to favor R. alpina populations on pollard stands with closed canopies.     

Colonisation of ephemeral forest habitats by specialised species: beetles and bugs associated with recently dead aspen wood

The most appropriate strategy for preserving fragmented populations depends on a species’ ability to colonise distant habitat patches. Insects associated with early decay stages of dead wood are expected to have a high capacity to colonise new habitat patches. To study the dispersal ranges of beetles (Coleoptera) and flat bugs (Hemiptera: Aradidae) dependent on recently dead aspen (Populus tremula) wood in Finland, we set out 58 piles of recently cut aspen logs at various distances up to 1.6 km from forests that contained a high density of old aspen trees. We captured insects by trunk window-traps, and counted beetles’ exit holes. Habitat connectivity was measured in terms of the amount of suitable aspen-wood in the surrounding environment, with the closest dead wood items up-weighted by a negative-exponential function. The log-piles attracted many saproxylic insects including four red-listed aspen-specialist species. The exposure of log-piles to the sun, and high levels of habitat connectivity increased the species richness of aspen-specialists, whereas bark peeling by moose decreased richness. The spatial scale at which species richness had its strongest response to habitat was 93 m. Among individual species there was a wide variability in spatial scale of response. This study supports the view that conservation efforts in boreal forests should be concentrated on sites where colonisation by target species is most likely. Restoration of habitat by re-locating logs may be useful at localities with a rich and specialised fauna but which have too low rate of formation of dead wood by natural processes.     

Fallen retention aspen trees on clear-cuts can be important habitats for red-listed polypores: a case study in Finland

Green-tree retention is a relatively new forestry application, which aims at decreasing the negative effects of clear-cut logging on forest biodiversity. In this study, the value of retained aspens in maintaining diverse assemblages of wood-decaying fungi (polypores; Basidiomycota) on clear-cuts was investigated, after the retention trees had died, fallen and started to decay. A total of 110 fallen aspen trunks were investigated on clear-cuts and within old-growth forests in eastern Finland, southern boreal zone; and 499 records of polypores belonging to 46 species were made. The intermediately decayed trunks on a clear-cut area hosted more species and more red-listed species than did trunks within forests. Most of the polypore species with more than two records were found in both habitats. These results suggest that many aspen-associated polypores are able to live and reproduce in sun-exposed habitats, if the quality and quantity of dead wood fulfill the species-specific requirements. This unexpected result, however, may be partly due to the exceptionally great abundance of aspen in the study area. Furthermore, in the long term, the local benefits of fallen retention trees can be limited, unless the local continuity of large aspens, both living and dead, is ensured.     

Long-term persistence of aspen – a key host for many threatened species – is endangered in old-growth conservation areas in Finland

Large, dead and dying European aspens (Populus tremula L.) host many threatened species in Fennoscandian boreal forests. Large aspen trees have mostly disappeared and are being harvested from the managed forests that cover 95% of the forest area in Finland. Due to the small area protected (4.1%), the aspen-associated species may encounter major difficulties in the protected areas if aspen trees disappear due to natural forest succession. The availability of aspens was assessed in the old-growth conservation area network in eastern Finland. We mapped all the living and dead aspens in 15 protected old-growth forests. The total number of counted trees was 32 903 individuals. Current amounts of living (2.7 m³/ha) and especially dead aspens (2.8 m³/ha) in the protected areas were higher than in the surrounding managed forests (1.1 and 0.1 m³/ha for living and dead trees, respectively). However, while saplings (dbh<5 cm) occur in most of the areas (12 individuals/ha on average) they survive poorly and young aspen cohorts (5 cm <dbh<15 cm) are lacking or are very rare. The most likely reason for the poor sapling survival is high browsing pressure by the mammalian herbivores, especially the moose. The moose population has increased many times in Finland during the past decades. The poor regeneration of aspens implies that the value of the old-growth conservation areas for aspen-associated species will face a serious bottleneck within a few decades when the currently middle-aged tree cohorts disappear. If the current high browsing pressure and lack of natural disturbances continue the obligatory aspen-associated species may disappear both locally and regionally from the network of the protected areas.     

Habitat availability in forests and clearcuts for saproxylic beetles associated with aspen

To maintain biodiversity in managed, boreal forest in Scandinavia, aspen trees (Populus tremula) are often retained at clearcutting. In this study, the habitat availability for beetles associated with aspen coarse woody debris (CWD) was predicted for forests and clearcuts with a model of CWD dynamics. Habitat requirements of eight beetle species (Agathidium bicolor, Cerylon ferrugineum, Cyphaea curtula, Endomychus coccineus, Homalota plana, Mycetophagus fulvicollis, Ptilinus fuscus and Xylotrechus rusticus) were obtained from their occurrence patterns in relation to characteristics of CWD objects in forest and on clearcuts in a study landscape in central Sweden. Three species were more frequent in forest and three at clearcuts. Five species increased with increasing girth of the CWD. Three were more frequent on standing CWD, and two on lying CWD. From the same study area, we also obtained field data on the recruitment of CWD (i.e., tree mortality) and amounts of different types of CWD. Annual tree mortality of aspen was higher for recent clearcuts (6.3%) compared with older clearcuts (1.1%). For all species, the habitat availability was higher on clearcuts, because enhanced tree mortality increased the amount of recently dead CWD. As a conclusion, green-tree retention of aspen is a conservation effort that is beneficial for species associated with aspen CWD.     

Habitat requirements of the endangered longhorn beetle <Emphasis Type="Italic">Aegosoma scabricorne</Emphasis> (Coleoptera: Cerambycidae): a possible umbrella species for saproxylic beetles in European lowland forests

Detailed knowledge of habitat requirements is an essential prerequisite for efficient conservation of any endangered species. Despite the grain support beetle Aegosoma scabricorne (Cerambycidae) being one of the largest European beetles, and an endangered, disappearing, species in Central Europe, its bionomics remain relatively poorly known. A. scabricorne is known as a polyphagous species on broadleaved trees; thus, to investigate its habitat preferences 174 broadleaved trees (87 occupied and 87 unoccupied by the species) were surveyed in the area of southern Moravia (Czech Republic) in 2015. The species was found to be strongly associated with declining or freshly dead trees that are, preferably, further damaged (breakage of stem or primary branch, hollows, etc.) and it particularly thrives on large trees (diameter >50 cm). Surprisingly, stem exposure to the sun was shown to be an unimportant characteristic for this species. However, in this study the number of exit holes was significantly smaller on the shaded north-facing quarter of the stem. Our results suggest concrete conservation measures to support the species. We also suggest that this species may be used as an umbrella species for saproxylic beetles of European lowland forests.     

Spatial pattern in the occurrence of the lichen Lobaria pulmonaria in managed and virgin boreal forests

Two populations of the lichen Lobaria pulmonaria. growing on aspens and goat willows in 12 and 20 km² study areas of boreal forest in Finland, were surveyed thoroughly to investigate the factors influencing the spatial distribution of the lichen. In one study area, where forestry has been intensive and old-growth forest is highly fragmented, L, pulmonaria was sparse and grew mostly on willows. In contrast, a large and continuous virgin forest area supported a higher incidence of L. pulmonaria. with the lichen being common on both aspens and willows. In both study areas, the distributions of aspen and willow were clumped over the scales of 100-1000 m. The spatial pattern of L pulmonaria was more clumped in the managed forest than in the virgin forest. The reduced incidence of the lichen on aspens in the managed area was attributed to a disruption of habitat continuity and small average tree size. There was no comparable reduction in the incidence on willows, probably because the willow had a very aggregated distribution in the managed area, which probably facililated local colonization of the lichen. Presence of the lichen was significantly related to size-corrected local density of aspen and willow trees as well as to spatial connectivity to neighboring lichen-occupied trees.     

Occurrence of epiphytic macrolichens in relation to tree species and age in managed boreal forest

For the maintenance of biodiversity some epiphytic macrolichens have been used to identify forests with high conservation value. To understand what these macrolichens indicate, we studied the relationships of tree species and age with the abundance of a set of 33 macrolichens (20 of them in use as indicator species) at 90 localities in a 150 km² area in south-central Sweden.
We sampled the four most common tree species Scots pine Pinus sylvestris. Norway spruce Picea abies , birch Betula spp, and aspen Populus tremula for macrolichens at 0-2 m from the base. Of the 33 target species we found 17, six of them were indicators in use for this region. In 30, out of 40 tested, cases there was a positive relation between lichen abundance and age of the tree.
On all tree species the number of the studied lichen species increased with age. The colonisation rate of the species was fastest on birch. All species occurring on birch were also found on at least one of the coniferous trees, while aspen had six species not found on other trees. Pine and spruce with a slow growth rate had more signal lichen species than fast growing trees.
We could not detect a competitive situation between common and rare species as the most abundant or largest lichens did not exclude other species. The length of the longest fruticose lichen was positively correlated with the number of species on the tree. Shade had a negative effect on the number of species.
We conclude that the indicator macrolichens encountered in this study showed a preference for old trees or aspens. This result, and the low number of indicator species, points at the importance of promoting these trees for the preservation of biodiversity in the boreal forest.     

Importance of habitat patch size for occupancy and density of aspen-associated saproxylic beetles

In Fennoscandian boreal forests, aspen (Populus tremula) is one of the most important tree species for biodiversity. In this study we explore how occupancy and density of beetles associated with dead aspen are related to habitat patch size and connectedness in a 45,000 ha boreal managed forest landscape in central Sweden. Patch size was estimated as amount of breeding substrate and connectedness as crown cover of living aspen in the surrounding landscape. The beetles were sampled by sieving of bark or by inspection of species-characteristic galleries in 56 patches with dead aspen. Six of nine aspen-associated species (Xylotrechus rusticus, Ptilinus fuscus, Mycetophagus fulvicollis, Cyphaea curtula, Homalota plana and Endomychus coccineus) showed a positive significant relationship between habitat patch size and occupancy. For all these species, except C. curtula, there was also a significant positive relationship between patch size and density. Connectedness was not retained as a significant variable in the analyses. Species not defined as aspen-associated constituted a significantly larger proportion of the total density of individuals of saproxylic beetles in smaller habitat patches than in larger patches. Richness of aspen-associated species was positively related to habitat patch size. Efforts in the managed forest should be directed towards preserving and creating larger patches of living and dead aspen trees and increasing the amount of aspen at the landscape level.     

Assemblages of saproxylic beetles on large downed trunks of oak

Old living oaks (Quercus robur) are known as a very species‐rich habitat for saproxylic beetles, but it is less clear to what extent such veteran trees differ from an even rarer feature: downed trunks of large oaks. In this study, we set out to sample this habitat, using window traps, with two aims: (1) to describe the variation of assemblages among downed trunks of different type and (2) to compare beetles on downed oaks with data from veteran standing trees. The results showed that trunk volume and sun exposure better explained assemblages as well as species numbers on downed trunks than did decay stage. Furthermore, species classified as facultative saproxylic species showed weak or no differentiation among downed trunks. Species with different feeding habits showed no apparent differentiation among downed trunks. Furthermore, species composition on dead, downed oak trunks differed sharply from that of living, veteran oaks. Wood or bark feeders were more common on veterans than downed trunks, but there was no difference for those species feeding on fungi or those feeding on insects and their remains. In conclusion, for a successful conservation of the saproxylic beetle fauna it is important to keep downed oak trunks, and particularly large ones, in forest and pastures as they constitute a saproxylic habitat that differs from that of living trees.     

Increasing insect abundance by killing deciduous trees: a method of improving the food situation for endangered woodpeckers

A total of 135 trees were girdled or notched in a forest area close to the river Dalälven in central Sweden. The purpose was to see whether good microhabitats for wood-living insects could be created and if so, could be used as food for the endangered white-backed woodpecker Dendrocopos leucotos. The chosen tree species were three of the most frequent used for foraging by the white-backed woodpecker, i.e. aspen Populus tremula , birch Betula alba and common alder Alnus glutinosa.
The process of dying and decay in the manipulated trees was followed for eight to nine years, and the following variables were studied: tree condition, tree structure, wood-condition, consistency of bark and cambium, frequency of polypori, foraging marks made by woodpeckers and abundance of insect species of importance as food for woodpeckers. Notched aspen died faster and were more quickly colonized by wood-living insects than were girdled aspen. Since the notched aspen was also reduced more quickly to low stumps or lost their bark, their value for wood-living insects was short-lived. Notched birch died almost as quickly as girdled birch but were populated by slightly fewer insect species. Notching, in general, seemed to create good foraging trees in the short-term whereas girdling produced trees of a longer-term value. In order to improve the local food situation for the white-backed woodpecker, a combination of the two methods is suggested. Nonetheless, both must be seen as emergency measures and cannot compensate for a large-scale reduction in the abundance of old deciduous trees in the forest.     

Hybrid aspen is not preferred by the large poplar borer (Saperda carcharias)

An industrially valuable tree, aspen, suffers from several fungal diseases and insect pests such as the large poplar borer (Saperda carcharias). The role of the beetle as a pest of aspen and hybrid aspen was investigated in five aspen and three hybrid aspen stands in Finland. Approximately 50% of the trees in the study had signs of the large poplar borer. Of these trees, 5% had insect exit holes. Approximately 70% of the trees with larval galleries had only one or two larval galleries. The trees with larval galleries were on average 2 m shorter than those without larval galleries. No significant difference could be detected in the diameter growth between these two groups. The proportion of decay was greater in hybrid aspen (27%) than in aspen (14%). These results show that the large poplar borer is an important pest of both aspen and hybrid aspen in Finland. No significant difference was observed between the aspen and hybrid aspen stands in the number of trees with larval galleries. Hybrid aspen therefore does not appear to be more sensitive to damage caused by the large poplar borer. Thus, a change from hybrid aspen to regular aspen in aspen cultivation is unnecessary, presuming that healthy saplings and an optimal rotation time are used.     

Linking woodland key habitat inventory and forest inventory data to prioritize districts needing conservation efforts

Woodland key habitat (WKH) inventories have been conducted in northern European countries, with the aim to create networks of minimally disturbed forest stands for protection. The goal of national forest inventory is to provide information relevant to forest management, such as on forest types, trees species composition, age structure and wood volume. The aim of this study was to link these two inventory databases to identify districts of Latvia most deficient in connectivity and habitat quality, in order to prioritize districts needing conservation effort. As an example, the area of deciduous forest with nemoral tree species (oak, ash, lime, maple and elm) and aspen was chosen. These forests provide habitat for a specific community of epiphytes. Using information in the WKH database, habitat quality in different districts of Latvia was estimated by the frequencies of occurrence of structural elements and selected indicator epiphyte species in nemoral tree species and aspen WKHs. Using digital data in the national forest inventory database, fragmentation metrics were determined for forests that, according to age and tree species composition, could potentially be nemoral tree and aspen WKHs. On a regional level, the lowest habitat quality in WKH occurred in districts that had the least fragmentation of potential WKH forest. In the less fragmented areas, the habitat quality of the existing WKH will likely increase in the future, and could be promoted by management to create structural elements typical of natural forests. The districts with the most fragmented nemoral and aspen forests, contained WKHs with the best habitat quality. A focus on protection should be given to these stands as they are the most likely to support source populations, and there is a need to improve spatial continuity of suitable tree substrate in these areas.     

The impact of available water content in previous agricultural soils on tree growth and nutritional status in young hybrid aspen plantations in Estonia

The impact of soil available water content (AWC) on the growth and foliar nutrient concentrations of trees in 7-yr-old hybrid aspen (Populus tremula L. x P. tremuloides Michx.) plantations established on abandoned agricultural lands was studied. AWC in the topmost 75 cm soil layer was significantly related to height growth and foliar N concentration of hybrid aspens. The correlations were stronger on well-drained automorphic soils and in the case of a dry growing season with insufficient precipitation. AWC over 150–160 mm can be considered sufficient, and below 120 mm insufficient for fast growth of hybrid aspen. The differences in AWC were less pronounced in the top 25 cm soil layer but were more noticeable in the 25–50 and 50–75 cm soil layers. AWC estimated as a function of soil specific surface area and bulk density was shown here to be a significant indicator for site selection for establishing hybrid aspen plantations. Foliar N concentration over 2.7% can be considered optimal and below 2.4%, insufficient for hybrid aspen on the studied soils. Foliar concentrations of P (on average 0.24%) and K (on average 0.79%) varied little and, thus, did not correlate with tree growth. The most suitable previous agricultural soils for afforestation with hybrid aspen would be moderately drained Albeluvisols, Luvisols and Planosols.     

Tree and stand-scale factors affecting richness and composition of epiphytic bryophytes and lichens in deciduous woodland key habitats

Conservation and sustainable forestry are essential in a multi-functional landscape. In this respect, ecological studies on epiphytes are needed to determine abiotic and biotic factors associated with high diversity. The aim of the present study was to evaluate relative sensitivity of conservation targets (epiphytic bryophytes and lichens) in relation to contrasting environmental variables (tree species, tree diameter at breast height, bark crevice depth, pH, tree inclination, pH, forest stand age, area and type) in boreo-nemoral forests. The study was conducted in Latvian 34 woodland key habitat (WKH) boreo-nemoral forest stands. Generalized linear mixed models and canonical correspondence analysis showed that tree species and tree bark pH were the most important variables explaining epiphytic bryophyte and lichen composition and richness (total, Red-listed, WKH indicator species). Forest stand level factors, such as stand size and habitat type, had only minor influence on epiphytic species composition and richness. The results of the present study indicate a need to maintain the diversity of tree species and large trees, particularly Acer platanoides, Carpinus betulus, Fraxinus excelsior, Populus tremula, Tilia cordata, Ulmus glabra and Ulmus laevis in conservation of epiphytic bryophyte and lichen communities in the future.     

Habitat of Osmoderma eremita Scop. (Coleoptera: Scarabaeidae), a beetle living in hollow trees

The habitat of Osmoderma eremita, a vulnerable species in Europe restricted to tree cavities, was examined in southeastern Sweden. The occurrence of O. eremita larvae and fragments, larval frass and imagines were investigated in 135, 72 and 21 living oak trees with wood mould cavities, respectively. Living individuals and fragments were only found in hollows with frass. The correlation between different characteristics of the oaks and the occurrence of the beetle were examined by building multi-variate models with logistic regression. The frequency of O. eremita is higher in hollows with openings directed towards the sun (S or W) and in cavities with large amounts of wood mould. In one area the frequency was higher in trees which stand in an open or half open surrounding. The tendency to prefer sun exposed sites implies that the forestation of oak meadows, caused by cessation of traditional management, might be detrimental to the species.     

Using Sex Pheromone and a Multi-Scale Approach to Predict the Distribution of a Rare Saproxylic Beetle

The European red click beetle, Elater ferrugineus L., is associated with wood mould in old hollow deciduous trees. As a result of severe habitat fragmentation caused by human disturbance, it is threatened throughout its distribution range. A new pheromone-based survey method, which is very efficient in detecting the species, was used in the present study to relate the occurrence of E. ferrugineus to the density of deciduous trees. The latter data were from a recently completed regional survey in SE Sweden recording >120,000 deciduous trees. The occurrence of E. ferrugineus increased with increasing amount of large hollow and large non-hollow trees in the surrounding landscape. Quercus robur (oak) was found to be the most important substrate for E. ferrugineus, whereas two groups of tree species (Carpinus betulus, Fagus sylvatica, Ulmus glabra, vs. Acer platanoides, Aesculus hippocastanum, Fraxinus excelsior, Tilia cordata) were less important but may be a complement to oak in sustaining populations of the beetle. The occurrence of E. ferrugineus was explained by the density of oaks at two different spatial scales, within the circle radii 327 m and 4658 m. In conclusion, priority should be given to oaks in conservation management of E. ferrugineus, and then to the deciduous trees in the genera listed above. Conservation planning at large spatial and temporal scales appears to be essential for long-term persistence of E. ferrugineus. We also show that occurrence models based on strategic sampling might result in pessimistic predictions. This study demonstrates how pheromone-based monitoring make insects excellent tools for sustained feedback to models for landscape conservation management.     

SPATIAL PATTERNING AND COMPETITION IN AN ASPEN-WHITE PINE SUCCESSIONAL SYSTEM

The spatial patterning of and competition between established large-toothed aspen (Populus grandidentata) and invading white pine (Pinus strobus) were examined in a 65-yr-old forest system at the AuSable Trails Institute of Environmental Studies, Kalkaska County, Michigan. Density, diameter, and distance measures were recorded for tree species on two 40 × 40 m grids containing 64 quadrats each 5 × 5 m. Several indices of dispersion were used to assess pattern and regression analysis of nearest neighbor distance to tree diameter provided an index of competition. The results indicate strong aggregation for both aspen and pine on a large scale, but regularity of pattern for pines in relation to their nearest neighbor. This regularity appears to be due to competition with established aspens and suggests that the spatial patterning of the aspens acts to control the spacing and subsequent development of the invading white pines.     

Habitat selection by the highly endangered long-horned beetle <Emphasis Type="Italic">Rosalia alpina</Emphasis> in Southern Europe: a multiple spatial scale assessment

Despite the popularity of the saproxylic cerambycid Rosalia alpina as a flagship species, its ecology is still poorly know, especially in the southern part of its range. Detailed information on its habitat preferences is needed to plan appropriate management. We set our multiple spatial scale assessment of habitat preferences in a beech forest of central Italy whose landscape, featuring both unmanaged forest and two types of grazed open forest, allowed us to look at the influence of different land uses. Preferred trees occurred in open sites, and those close to tall undergrowth were avoided. A range of moribund or dead trees were used: those preferred had a lower percentage canopy closure, significantly thicker bark, and were more sun-exposed, than the average. Logistic regression showed that the most important variables for selection were distance from nearest occupied tree, bark thickness, undergrowth height and irradiation. Occurrence likelihood augmented as the distance from nearest other occupied tree increased. Despite being mostly unmanaged, forest was avoided, whereas open forest (with trees pruned by ‘shredding’) was used more than expected. Although intensive forestry limits the availability of dead wood, closed forest may be unsuitable when shadowing useful substrate. The disappearance of traditional forms of forest management as shredding and moderate cattle grazing may lead to woody vegetation expansion and habitat closure eventually threatening the persistence of R. alpina. The return to traditional habitat management would be beneficial to R. alpina, an issue that conservation plans should carefully take it into account.