Saproxylic Beetles in Standing Dead Wood Density in Relation to Substrate Sun-exposure and Diameter

Species-specific densities of saproxylic beetles in cut 4-m high, medium to large diameter stumps of spruce, birch, aspen and oak were monitored for 7 years after cutting, by trapping emerging insects within seasonally applied net enclosures. A total of 47,038 individuals, representing 316 saproxylic beetle species including 40 red-listed species, were recorded. Densities of 86 species were modelled in relation to stump diameter and measured levels of sun-exposure. Two thirds of the species favoured semi- or fully exposed substrates, while one third performed better in shade. Few species seemed to be specifically adapted to semi-exposed conditions. Diameter was generally of less importance than exposure, and similar numbers of species favoured medium and large diameter substrates. Group mean diameter and exposure calculated from means per individual and species were similar for groups of modelled and non-modelled species, and for groups of red-listed and non-red-listed species, respectively. This indicates that the proportions found in the models are representative for the diverse regional fauna of saproxylic beetles that utilise high stumps of spruce, birch, aspen and oak in early stages of decay. The results suggest that sun-exposed dead wood substrates generated and retained in managed forest landscapes are potentially important for many species. However, stands of unmanaged self-thinning successions of deciduous trees, as well as substantial areas of old-growth spruce-forests are also necessary to address the wide spectrum of saproxylic beetles.     

Saproxylic beetles in artificially created high-stumps of spruce and birch within and outside hotspot areas

Artificially created high-stumps (snags) are created regularly during forest felling operations in Swedish coniferous production forests. The saproxylic beetle fauna in high-stumps of spruce and birch, on 20 clearcuts in Southern Sweden, were sampled by bark sieving. Ten of the clearcuts were located in hotspot areas with documented occurrence of many red-listed saproxylic beetle species. The other ten clearcuts were located in a typical production forest landscape (i.e. the matrix). Our aim was to investigate whether the benefit of creating high-stumps differs if the clearcuts is located in a hotspot area or in the matrix. In total 4,179 saproxylic beetles were found, belonging to 66 species, 9 of which were red-listed. Birch high-stumps hosted more species, on average, than spruce high-stumps. In an ordination analysis, tree species had the strongest explanatory effect among the environmental variables. No difference in beetle fauna could be found between the hotspot and matrix clearcuts, for neither birch nor spruce, according to all parameters: species numbers, species composition and red-listed species. The study does not indicate that conservation efforts in coniferous production forests should be concentrated to hotspot areas.     

Increased openness around retained oaks increases species richness of saproxylic beetles

The decrease of old deciduous trees in northern Europe is a threat to the saproxylic fauna. In northern Europe, oak sustains the highest richness of saproxylic invertebrates, among which beetles is a large group. In order to preserve species associated with old trees, it has become common practice in commercial forestry to retain such trees at final felling. However, to create beneficial conditions for species associated with retained trees, the surrounding plantation has to be managed with regard to their specific demands. In the case of oak-associated species, including many red-listed species, several studies have shown that light is an important factor. The aim of this study was to analyze the effects of increased openness around oaks (Quercus robur) in spruce plantations (Picea abies) on species richness and abundance of oak-associated saproxylic beetles. The study was performed in nine spruce plantations located in southern Sweden, with mature oaks standing in a gradient of canopy openness. Beetles were collected from 54 oaks from May to September during two seasons, using window traps. The analyses revealed that increased openness around oaks increases species richness and abundance of oak-associated beetles. By including insolation angle in the analysis, we found that it is mainly the degree of openness directed south from the oak that has positive effects on beetle richness. These findings imply that it is desirable to maintain an open area around retained oaks, and that this area should be placed on the southern side of the oak to maximize the biodiversity benefit in relation to costs for the forest owner.     

Hollow oaks and beetle conservation: the significance of the surroundings

In this study we investigated hollow oaks (Quercus robur, Q. petrea) situated in open landscapes and in forests in Norway in northern Europe, and compared their importance for rare and threatened beetles (Coleoptera). Old, hollow oak trees, both in parks and in forests, were extremely rich in red-listed beetles, and hosted a high proportion of threatened species. The proportion of oak associated species and the mean number of red-listed beetle species per tree was similar in the two site types, but rarefaction showed that for a certain number of individuals, oaks in forests had more threatened and near-threatened species than oaks in parks. The species composition also differed between site types: Park oaks had a higher proportion of species associated with hollows and animal nests, whereas in forests, there was a higher proportion of species depending on dead oak wood in general. Four factors were significant in explaining the richness of red-listed beetles in our study: Tree circumference, cavity decay stage, proportion of oak in the surroundings, and coarse woody debris (CWD) in the surroundings. Forest oaks were smaller, but they still trapped a species richness comparable to that of the larger park oaks—probably a result of high amounts of CWD in the surroundings. We show that oaks in open landscapes and oaks in forest have only partly overlapping beetle assemblages and, thus, cannot be substituted in conservation. Planning for conservation of red-listed beetles associated with this key habitat demands a large scale perspective, both in space and time, as the surroundings have important effects on associated threatened and near threatened species.     

Can 'continuity indicator species' predict species richness or red-listed species of saproxylic beetles?

This study investigates the relationship between the abundance of wood-rotting fungus suggested as 'continuity indicator species' and environmental variables for the assemblage of saproxylic (wood-living) beetles associated with Fomitopsis pinicola fruiting bodies in a mature spruce forest in southeastern Norway. The presence of species thought to indicate continuity in old growth is one of the criteria used when finding and delineating small protected areas ('woodland key habitats') in Scandinavian forestry. Although it is clear that remnants of old-growth forest are important for many taxa, documentation as to which entities or species the indicator species indeed indicate is scarce. If stands with a continuous and unbroken input of dead wood have a unique assemblage of wood-rotting fungi, it seems relevant to ask if these stands also have a unique assemblage of rare saproxylic beetles. I find that the indicator species exhibit no significant correlations with beetle species richness or with the presence of red-listed saproxylic beetles as a group. The different characteristics of dead wood conditions are the most important environmental variables that explain both the species richness and the presence of red-listed beetles. Single-species analyses reveal contrasting relationships. The red-listed beetle Atomaria alpina shows a significant and positive association to the abundance of indicator species. Contrary, a group of three red-listed species with similar ecology in the family Cisidae exhibits a significant and negative association to indicator species abundance. This indicates that important patterns are concealed when considering general measures such as overall presence of red-listed beetles. Single-species studies are necessary in order to correctly understand how rare beetles respond to forestry activities and to develop a policy that can secure their continuing existence in the boreal forest.     

An indicator system for identification of sites of high conservation value for saproxylic oak (Quercus spp.) beetles in southern Sweden

The saproxylic beetle fauna on old oaks was sampled in four regions of southern Sweden using two methods: window and pitfall trapping. The aim was to test a way of finding indicator species which can be used to identify sites with high species number or that scored high on a conservation priority species index, based on occurrence of red-listed species. From 92 sites surveyed, in total 164 species of saproxylic beetles were identified. Different sets of indicator species were selected based upon 22 sites from a centrally located region. Predictions of species number and the index for 30 other sites from the same province were made. The correlation between observed and predicted species number and the index increased with increasing number of indicators. When comparing different treatment of species indata, the explanatory power of predictions was strongest for presence/absence data. Indicator sets of species effectively caught with pitfall traps gave overall the best predictions of both species number and the index. Predictions of species number and the index worked well within the same regions but gave varied result for the three other regions which shows that transferability of indicators between regions may be doubtful.     

The “dehesa”, a key ecosystem in maintaining the diversity of Mediterranean saproxylic insects (Coleoptera and Diptera: Syrphidae)

The “dehesa” is a traditional Iberian agrosilvopastoral ecosystem characterized by the presence of old scattered trees that are considered as “keystone-structures”, which favor the presence of a wide range of biodiversity. We show the high diversity of saproxylic beetles and syrphids (Diptera) in this ecosystem, including red-listed species. We analyzed whether saproxylic species distribution in the “dehesa” was affected by tree density per hectare, dominant tree species or vegetation coverage. Species diversity did not correlate with tree density; however, it was affected by tree species and shrub coverage but in a different way for each taxon. The highest beetle diversity was linked to Quercus pyrenaica, the most managed tree species, with eight indicator species. In contrast, Q. rotundifolia hosted more species of saproxylic syrphids. Regarding vegetation coverage, shrub coverage was the only variable that affected insect richness, again in a different way for both taxa. In contrast, beetle species composition was only affected by dominant tree species whereas syrphid species composition was not affected by tree species or shrub coverage. We concluded that the high diversity of saproxylic insects in the “dehesa” is related to its long history of agrosilvopastoral management, which has generated landscape heterogeneity and preserved old mature trees. However, the richness and composition of different taxa of insects respond in different ways to tree species and vegetation coverage. Consequently, conservation strategies should try to maintain traditional management, and different saproxylic taxa should be used to monitor the effect of management on saproxylic diversity.     

The species richness of click beetles in ancient pasture woodland benefits from a high level of sun exposure

Forests support high concentrations of species and beetles in particular are often used to evaluate forest biodiversity. Ancient pasture woodlands are facing a major decline in Europe mainly due to the abandonment of traditional management and subsequent succession. We studied click beetles (Coleoptera: Elateridae) in one of the largest central-European remnants of pasture woodland in Lány Game Park (Czech Republic) using flight interception traps placed at standing veteran trees. The gradient of sun-exposure, circumference of stem, height and vitality of tree and tree species were studied in relation to the species richness of click beetles and their ecological groups. Total species richness reached nearly one half of the recently documented fauna in the study area and species accumulations showed us that the majority of species were represented. Most species preferred solitary trees in sun-exposed habitats and avoided shaded trees in closed canopies. The same results were obtained for ecological groups, such as saproxylic and non-saproxylic species, functional groups and guilds. Our results showed that the species richness of one of the most ecologically diverse beetle families, click beetles, benefits from a high level of sun exposure. Thus, the long spatial and temporal continuity of sun-exposed veteran trees could be a good predictor for sustainable forest management.     

Conflict between insect conservation and public safety: the case study of a saproxylic beetle (<Emphasis Type="Italic">Osmoderma eremita</Emphasis>) in urban parks

Urban parks can harbour small populations of saproxylic insects of high conservation concern, such as Osmoderma eremita and other rare beetles. These areas often host old trees which have become very uncommon in rural areas where they are threatened by commercial forestry management procedures based on frequent tree cutting. Nevertheless, old trees of urban parks may represent a hazard for public safety and are sometimes cut by management authorities. The aim of this work was to assess the loss of reproductive sites for saproxylic beetles of the Scarabaeidae, Lucanidae and Cerambycidae, when felling plans are adopted according to a Visual Tree Assessment Procedure (VTA), in a Mediterranean urban park. On July–August 2004, 1,247 holm oaks were surveyed within the border of an urban park of Rome (Villa Borghese). The occurrence of saproxylic beetles (i.e. the presence of frass, living insects or their remains) was verified in 66 old holm oaks, 41% of which were doomed to cutting by VTA. Eleven of these trees (41% of the trees doomed to be cut) held fragments of adults and sometimes living larvae of Osmoderma eremita, and four of them (36%) were included in the felling plan. The presence of Osmoderma eremita in tree holes was more frequent in deep cavities. The presence of frass in the cavities was positively associated with tree height and a high degree of damage at the root collar and negatively with the presence of hole-nesting birds.     

Explaining the saproxylic beetle diversity of a protected Mediterranean area

Saproxylic beetle diversity is high at the Cabañeros National Park (central Spain), where woodland habitats exhibit remarkable heterogeneity. Our aim was to explain the diversity of saproxylic beetles, focusing on species turnover among mature woodland types. We surveyed five woodland types that represented the heterogeneity of the park’s woodland habitats. Beetles were collected using window traps over a period of 20 months. The Jaccard Similarity Index was used as indirect value of beta diversity among woodlands and to test the relation between species turnover and geographical distance. We also identified the contribution of species turnover to landscape diversity by using a partitioning model. Moreover, the presence of mixed woodlands (more than one tree species) allowed us to attempt to valorise the effect of tree species (coupled with their historical management) on species turnover among woodlands. Finally, we looked for different saproxylic beetle preferences for habitat and tree species using an indicator value method. We found that saproxylic beetle species composition varied significantly among the studied woodlands. The variation in species turnover was independent from the distance among woodlands, which suggested that beetle dispersal abilities could not explain this high turnover. Tree species within woodlands were a key factor that increased diversity turnover in woodlands and, consequently, the diversity of the park. Moreover, we found saproxylic beetle species that had different habitat and tree species preferences. We conclude that woodland heterogeneity (highly affected by woodland composition) seems to be the driving force for saproxylic beetle diversity in this protected area.     

Restoration of beech forest for saproxylic beetles—effects of habitat fragmentation and substrate density on species diversity and distribution

The influence of spatial location and density of beech snags on species diversity and distribution patterns of saproxylic beetles was studied in a 2,400 ha forest landscape in southern Sweden. Complete snag surveys were combined with a beetle survey using small window traps directly attached to the beech snags. The density of beech snags ≥30 cm dbh varied between one and seven snags per ha within the study area, corresponding to 1.1–5.1 m³/ha. A total of 2,610 specimens of 180 saproxylic beetles species were trapped, of which 19 species were red-listed. Within the study area, the number of red-listed and formerly red-listed species was highest around traps in old-growth stands, intermediate in managed stands contiguous with old-growth and lowest in managed stands isolated from old-growth by a two km-wide zone without beech forest. Logistic regressions revealed negative relationships between distance to old-growth forest and occurrence of eleven species, among them six red-listed or formerly red-listed species. The number of non red-listed species was not correlated with isolation from old-growth forest. The number of red-listed species also increased with snag density within 200–300 m around the traps. Our results suggest that red-listed species generally have a lower dispersal capacity than other saproxylic beetles. We conclude that retention of dead wood close to existing populations is more beneficial for red-listed species than an even distribution of snags across the forest landscape.     

Saproxylic beetle assemblages related to silvicultural management intensity and stand structures in a beech forest in Southern Germany

Compared to agricultural land and spruce plantations, central European beech-oak forests are often relatively close to natural conditions. However, forest management may alter these conditions. In Steigerwald, southern Germany, a large beech-dominated forest area, three management intensities were applied during the past 30–70 years. Here, we examined the influence of management intensity on saproxylic beetles in >100-year old mature stands at 69 sampling plots in 2004. We sampled beetles using flight-window traps and time standard direct searches. The community structure based on presence/absence data changed remarkably along the gradient from unmanaged to low-intensity to high-intensity management, but these differences were not evident using abundance data from flight interception traps. Saproxylic species richness decreased in intensively managed forests. Elateridae and threatened species richness peaked in unmanaged forests and in forests under low-intensity management. Saproxylic species richness was dependent on certain micro-habitat factors. These factors were (1) the amount of dead wood for Elateridae, overall and threatened saproxylic beetle richness; (2) the amount of flowering plants for Cerambycidae; (3) the richness of wood-inhabiting fungi for Staphylinidae, Melandryidae and overall saproxylic beetle richness; and (4) the frequency of Fomes fomentarius for threatened species. Species richness was better explained by plot factors, such as dead wood or fungi, than by management intensity. These results suggest that the natural variation of dead wood niches (decay stages, snag sizes, tree cavities and wood-inhabiting fungi species) must be maintained to efficiently conserve the whole saproxylic beetle fauna of beech forests. Also, intensive management may alter the specialised saproxylic beetle community even if the initial tree-species composition is maintained, which was the case in our study. For monitoring the ecological sustainability of forest management we must focus on threatened species. If structures alone are sampled then the amount of dead wood is the best indicator for a rich saproxylic beetle fauna.     

Influence of boreal forest succession and dead wood qualities on saproxylic beetles

1 Saproxylic insects, a functional group dominated by beetles, are dependent on dead or moribund trees as habitat elements. 2 Although there are few studies of saproxylic insects from the North American boreal zone, European studies demonstrate that forest harvest can lead to a biologically significant decrease in saproxylic beetle diversity. 3 We studied saproxylic beetles in the North American boreal mixedwood forest using flight intercept traps established on naturally dead and girdled trembling aspen and spruce trees along a successional gradient of undisturbed stands from deciduous to coniferous overstory trees. 4 Composition and diversity of beetle assemblages differed among forest successional types. 5 Snag age class was an important determinant of composition for saproxylic beetle assemblages. 6 Multivariate regression analysis of these data indicated that saproxylic beetles are responding to changes in coarse woody debris, and not to the relative densities of canopy tree species, although these variables are strongly correlated. 7 Coarse woody debris management should be a primary concern in forest management plans seeking to conserve saproxylic organisms and the critical ecosystem functions (i.e. nutrient cycling) in which they participate.     

Mimicking natural disturbances of boreal forests: the effects of controlled burning and creating dead wood on beetle diversity

The young successional stages of boreal forests are an important habitat for many saproxylic species. These habitats are formed by disturbances such as forest fires and they are characterized by large volumes of dead wood and sun-exposed conditions. Today, young successional stages of natural origin are very rare in Fennoscandia and there is need for restoration. We constructed a large-scale field experiment in which we studied the effects of two restoration practices on beetle diversity: controlled burning and partial harvesting with creating different volumes of dead wood. We sampled beetles with flight-intercept traps recording a total of 56,031 individuals and 755 species. The species richness and abundance of both saproxylic and non-saproxylic beetles were increased by burning and harvesting but the volume of dead wood created on harvested sites had no short-term effect on species richness or abundance. Rare species, especially saproxylic ones, preferred burned sites and a similar trend was observed among red-listed and pyrophilous species. Burning and harvesting also resulted in different species assemblages and there were some additional differences according to the volume of dead wood. We conclude that fire can be successfully used in restoration of managed boreal forests to increase species diversity and to facilitate the recovery of declined species. However, long-term monitoring is needed to clarify the effects of the restoration practices, in particular those of creating dead wood without using fire.     

A comparison of three methods to survey saproxylic beetles in hollow oaks

One of the most endangered assemblages of species in Europe is insectsassociated with old trees. For that reason there is a need of developing methodsto survey this fauna. This study aims at comparing three methods – windowtrapping, pitfall trapping and wood mould sampling – to assess speciesrichness and composition of the saproxylic beetle fauna in living, hollow oaks.We have used these methods at the same site, and to a large extent in the sametrees. Useful information was obtained from all methods, but they partiallytarget different assemblages of species. Window trapping collected the highestnumber of species. Pitfall trapping collected beetles associated with treehollows which rarely are collected by window traps and therefore it isprofitable to combine these two methods. As wood mould sampling is the cheapestmethod to use, indicator species should preferably be chosen among specieswhich are efficiently collected with this method.     

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.     

Micro and macro-habitat associations in saproxylic beetles: implications for biodiversity management

Restoration of habitats is critically important in preventing full realization of the extinction debt owed as a result of anthropogenic habitat destruction. Although much emphasis has been placed on macrohabitats, suitable microhabitats are also vital for the survival of most species. The aim of this large-scale field experiment was to evaluate the relative importance of manipulated microhabitats, i.e., dead wood substrates of spruce (snags, and logs that were burned, inoculated with wood fungi or shaded) and macrohabitats, i.e., stand types (clear-cuts, mature managed forests, and forest reserves) for species richness, abundance and assemblage composition of all saproxylic and red-listed saproxylic beetles. Beetles were collected in emergence traps in 30 forest stands in 2001, 2003, 2004 and 2006. More individuals emerged from snags and untreated logs than from burned and shaded logs, but species richness did not differ among substrates. Assemblage composition differed among substrates for both all saproxylics and red-listed saproxylic species, mainly attributed to different assemblage composition on snags. This suggests that the practise of leaving snags for conservation purposes should be complemented with log supplementation. Clear-cuts supported fewer species and different assemblages from mature managed forests and reserves. Neither abundance, nor species richness or assemblage composition differed between reserves and mature managed forests. This suggests that managed stands subjected to selective cutting, not clear-felling, maintain sufficient old growth characteristics and continuity to maintain more or less intact assemblages of saproxylic beetles. Thus, alternative management methods, e.g., continuity forestry should be considered for some of these stands to maintain continuity and conservation values. Furthermore, the significantly higher estimated abundance per ha of red-listed beetles in reserves underlines the importance of reserves for maintaining viable populations of rare red-listed species and as source areas for saproxylic species in boreal forest landscapes.     

The saproxylic beetle assemblage associated with different host trees in Southwest China

Dead wood is a habitat for many insects and other small animals, some of which may be rare or endangered and in need of effective protection. In this paper, saproxylic beetle assemblages associated with different host trees in the subtropical forests in southwestern China were investigated. A total of 277 species (1 439 specimens) in 36 beetle families were collected from 117 dead wood samples, of which 101 samples were identified and respectively belonged to 12 tree genera. The number of saproxylic beetle species varied greatly among logs of different tree genera, with the highest diversity on logs of Juglans. Generally, broad‐leaved trees had a higher richness and abundance of saproxylic species than coniferous trees. Cluster analysis revealed that assemblages from broad‐leaved tree genera were generally similar (except for Betula) and assemblages from coniferous trees formed another distinct cluster. The subsequent indicator analysis proposed that there are different characteristic species for different cluster groups of host tree genera. In our study, log diameter has no positive influence on beetle species density. Conversely, comparisons of individual‐based rarefaction curves suggested that beetle species richness was highest in the small diameter class both in coniferous and broad‐leaved tree genera. With increased wood decay, proportion of habitat specialists (saproxylic beetles living on one tree genus) decreased, whereas proportion of habitat generalists (living on more than three tree genera) increased. The beetle species density was found to be higher in early stages, and decreased in later stages as well. A negative influence of altitude on saproxylic beetle species richness and abundance was detected. It was indicated that different tree genera and altitudes possibly display cross effects in modulating the altitudinal distribution and host preference of the beetles.     

Robinia pseudoacacia as a surrogate for native tree species for saproxylic beetles inhabiting the riparian mixed forests of northern Italy

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Fast but ephemeral effects of ecological restoration on forest beetle community

Many protected areas have a long history of human intervention before being protected. In protected forests, the past land use has reduced the amount of natural structures, which are crucial substrates for thousands of species. We evaluate the short-term ecological effect of forest restoration (dead wood creation) on conifer-associated saproxylic (dead-wood dependent) beetles. More specifically, we analyze the effect of dead wood creation on the number of beetle species and individuals 1 and 5 years after restoration in spruce and pine forests, using a large-scale monitoring network over Finland. The number of saproxylic beetle species and individuals was larger at restored than at control plots both 1 and 5 years after restoration in both spruce and pine forests. Community composition in restored plots was different from control plots 1 year after restoration, but had returned towards the control plot composition 5 years after restoration, while control plots remained largely unchanged. Both in spruce and pine forests, there were more red-listed and rare saproxylic beetles in restored than in control plots 1 and 5 years after restoration. Our results indicate that restoration has an overall positive influence on saproxylic beetle diversity immediately after dead wood creation, but this effect is rather short-lived. Long term monitoring of restored dead wood is crucial in investigating successional pathways as well as biotic communities in advanced decay stages, and in fully evaluating the ecological effect of dead wood creation as a forest restoration measure.