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.     

Boxes mimicking tree hollows can help conservation of saproxylic beetles

Old hollow trees have declined in Europe and many saproxylic (i.e. wood-dwelling) invertebrates living on them are threatened. The aim of this study was to investigate to what extent artificial habitats can be exploited by saproxylic beetles. To mimic the conditions in tree hollows, we constructed wooden boxes filled with different combinations of substrates like oak saw dust, oak leaves, a dead hen (Gallus domesticus), chicken dung, lucerne flour or potatoes and placed them on tree trunks. To investigate the importance of distance from dispersal sources, we placed boxes at different distances (0–1,800 m) from three species-rich sites with high densities of hollow oaks. Over 3 years, 3,423 specimens of 105 saproxylic beetle species were caught in 47 boxes. Among beetles found in hollow oaks that were either tree-hollow species, bird nest species, or wood rot species, 70% were also found in the boxes. A dead hen added to the artificial wood mould gave a higher number of beetle specimens. The number of species associated with tree hollows in oak decreased with distance from sites with hollow oaks. In conclusion, the prospects for using artificial environments for boosting substrate availability, or to fill spatial and temporal gaps therein, for saproxylic beetles are good.     

Hollow oaks and beetle functional diversity: Significance of surroundings extends beyond taxonomy

Veteran hollow oaks (Quercus spp.) are keystone structures hosting high insect diversity but are declining in numbers due to intensification of land use and the abandonment of traditional management. The loss of this vital habitat is resulting in a reduction of biodiversity, and this likely has consequences for ecosystem functioning, especially if functional diversity is reduced. A considerable amount of research has been done on predictors of beetle taxonomic diversity in veteran oaks, but predictors of functional diversity have remained largely unexplored. The aim of this study was to establish whether the features and surroundings of veteran oaks are related to functional diversity within three functional groups of beetles (decomposers, predators, and flower visitors) and determine whether species richness and functional diversity within the groups are dependent on the same predictors. Sampling was carried out intermittently between 2004 and 2011 on 61 veteran oaks in Southern Norway. Of the 876 beetle species that were collected, 359 were determined to be decomposers, 284 were predators, and 85 were flower visitors. Species richness and functional diversity in all groups were consistently higher in traps mounted on veteran oaks in forests than in open landscapes. However, additional predictors differed between groups, and for species richness and functional diversity. Decomposer species richness responded to tree vitality, while functional diversity responded to habitat connectivity, predator species richness responded to regrowth of shrubs while functional diversity responded to tree circumference, and flower visitor richness and functional diversity did not respond to any additional predictors. Previous studies have found that the features and surroundings of veteran oaks are important for conservation of taxonomic diversity, and the results from this study indicate that they are also important for functional diversity within multiple functional groups.     

Boxing for biodiversity: evaluation of an artificially created decaying wood habitat

Many saproxylic species are threatened in Europe because of habitat decline. Hollow trees represent an important habitat for saproxylic species. Artificial habitats may need to be created to maintain or increase the amount of habitat due to natural habitat decline. This study investigated the extent to which saproxylic beetles use artificial habitats in wooden boxes. The boxes were placed at various distances (0–1800 m) from known biodiversity hotspots with hollow oaks and studied over 10 years. Boxes were mainly filled with oak saw dust, oak leaves, hay and lucerne flour. In total, 2170 specimens of 91 saproxylic beetle species were sampled in 43 boxes. The abundance of species associated with tree hollows, wood rot and animal nests increased from the fourth to the final year, but species richness declined for all groups. This study shows that wooden boxes can function as saproxylic species habitats. The artificial habitats developed into a more hollow-like environment during the decade long experiment with fewer but more abundant tree hollow specialists.     

Development of secondary woodland in oak wood pastures reduces the richness of rare epiphytic lichens

Wooded pastures with ancient trees were formerly abundant throughout Europe, but during the last century, grazing has largely been abandoned often resulting in dense forests. Ancient trees constitute habitat for many declining and threatened species, but the effects of secondary woodland on the biodiversity associated with these trees are largely unknown. We tested for difference in species richness, occurrence, and abundance of a set of nationally and regionally red-listed epiphytic lichens between ancient oaks located in secondary woodland and ancient oaks located in open conditions. We refined the test of the effect of secondary woodland by also including other explanatory variables. Species occurrence and abundance were modelled jointly using overdispersed zero-inflated Poisson models. The richness of the red-listed lichens on ancient oaks in secondary woodland was half of that compared with oaks growing in open conditions. The species-level analyses revealed that this was mainly the result of lower occupancy of two of the study species. The tree-level abundance of one species was also lower in secondary woodland. Potential explanations for this pattern are that the study lichens are adapted to desiccating conditions enhancing their population persistence by low competition or that open, windy conditions enhance their colonisation rate. This means that the development of secondary woodland is a threat to red-listed epiphytic lichens. We therefore suggest that woody vegetation is cleared and grazing resumed in abandoned oak pastures. Importantly, this will also benefit the vitality of the oaks.     

Old park trees as habitat for saproxylic beetle species

Very old trees harbour a diverse fauna of saproxylic insects, many of which are classified as threatened due to the scarcity of this kind of habitat. Parks, which often contain many old trees, are therefore considered to be important sites for this fauna. However parks are intensively managed and dead wood is often removed. Therefore this study compares if the saproxylic beetle fauna in parks is as diverse as it is in more natural stands. Eight ‘Park’ sites at manor houses around lake Mälaren, Sweden were compared with trees in wooded meadows: eight grazed sites, here termed ‘Open’, and 11 sites regrown with younger trees, termed ‘Re-grown’. The comparison was made on lime trees (Tilia spp.): one of the most frequent tree species in old parks which host a diverse beetle fauna. Beetles were sampled with window traps, which in total caught 14,460 saproxylic beetles belonging to 323 species, of which 50 were red-listed. When comparing all saproxylic species, ‘Park’ sites had significantly fewer species than ‘Open’ sites. However, for beetles in hollow trees and for red-listed species there was no significant difference, the number in ‘Park’ being intermediate between ‘Open’ and ‘Re-grown’. Species composition differed between sites, but only marginally so. Therefore, the conclusion is that old park trees on average are as valuable for faunal diversity as trees in more natural sites. Large conservation benefits can be obtained by combining cultural and conservational values in the management of park habitats.     

Saproxylic beetle diversity in a managed boreal forest: importance of stand characteristics and forestry conservation measures

Saproxylic beetles constitute a significant proportion of boreal forest biodiversity. However, the long history of timber production in Fennoscandia has significantly reduced the availability of dead wood and is considered a threat to the conservation of saproxylic beetle assemblages. Therefore, since the mid‐1990s dead wood retention in harvested stands has formed an integral part of silvicultural practices. However, the contribution of this biodiversity‐orientated management approach to conserving saproxylic beetle assemblages in boreal forest landscapes that include production forestry remains largely untested. We examined differences in resident saproxylic beetle assemblages among stands under different management in a boreal forest landscape in Central Sweden, and in particular stands managed according to new conservation‐orientated practices. We also investigated the relationship between beetle diversity and forest stand characteristics. Bark of coarse woody debris (CWD) was sieved for beetles in old managed stands, unmanaged nature reserves, and set‐aside areas, and clear‐cut stands harvested according to certification guidelines [new forestry (NF) clear‐cuts]. All stand types contributed significantly to the total diversity of beetles found. While stand size, position, and distance to nearest reserve were unimportant, both the quality and the quantity of CWD in stands contributed significantly to explaining beetle abundance and species richness. This extends the previous findings for red‐listed invertebrates, and shows that heterogeneous substrate quality and a range of management practices are necessary to maintain saproxylic beetle diversity in boreal forest landscapes that include production forestry. The unique abiotic conditions in combination with the abundant and varied CWD associated with NF clear‐cuts form an important component of forest stand heterogeneity for saproxylic beetles. It is thus essential that sufficient, diverse, CWD is retained in managed boreal landscapes to ensure the conservation of boreal saproxylic beetle assemblages.     

Litter-dwelling beetles in primeval forests of Central Europe: Does deadwood matter?

We investigated the distribution pattern of litter dwelling beetles in four primeval forests of the Western Carpathians. The forests are situated in two mountain ranges and are either southerly exposed oak forests or northerly exposed beech forests. Beetles were extracted from leaf litter of plots close to coarse woody debris (c-CWD) and distant from coarse woody debris (d-CWD). We collected 2946 individuals of 172 species. Plots close to CWD usually were hot spots of species richness and beetle density, which could be increased two-fold and five-fold, respectively, when compared to d-CWD sites. The influence of CWD within each forest type was stronger than that of any other environmental factor. The c-CWD plots were characterised by both a higher presence of common species and a higher number of rare species. Especially, zoophagous and mycetophagous beetles were enhanced. The assemblage wide carrying capacity at c-CWD sites varied in respect to the gradients of temperature and moisture within a forest. A longer gradient provided a higher carrying capacity. We assume that the influence of downed deadwood on litter dwelling beetles will also be effective in commercial forests and stop the downward spiral with continuing loss of species.     

浙江天目山种植林管理历史及粗死木残体特征影响柳杉腐木甲虫多样性（英文）

不合理的森林管理是导致腐木甲虫多样性丧失的重要原因。在中国亚热带地区, 多样性较高的天然林已被大面积的人工种植林取代, 然而, 这些人工林对腐木甲虫多样性的影响还研究甚少。本研究对浙江天目山自然保护区人工幼龄林（30~40年）、 人工老熟林（80~100年）和半天然混合林（>200年）中柳杉枯立木上的腐木甲虫群落及多样性进行比较。结果表明： 半天然混合林腐木甲虫个体数量（97.4±66.7）显著高于幼龄林（39.9±16.3）和老熟林（21.9±5.9）, 但半天然林混合林(27.9±11.2)与幼龄林(24.1±3.7)腐木甲虫物种数差异并不显著(P>0.05), 而幼龄林的腐木甲虫物种数和个体数量则显著高于老熟林(P<0.05)。腐木甲虫物种数和个体数量与样地粗死木残体体积相关性显著(P<0.05)。典范对应分析和多响应置换过程分析表明腐木甲虫群落特征在不同林型间差异显著(P<0.001)。柳杉枯立木直径、 粗死木残体的直径和数量以及林冠盖度均对腐木甲虫物种组成具有显著影响(P<0.05)。腐木甲虫营养级组成分析也表明, 半天然混合林菌食性甲虫数量显著高于种植林(P<0.001)。结果提示, 提高种植林粗死木残体的数量和质量可以增加腐木甲虫的物种丰富度, 但种植林腐木甲虫多样性可能在随后的演替阶段有所下降, 而且种植林与天然林在腐木甲虫群落组成上差异十分明显。     

Micro-habitat drivers of saproxylic beetle assemblages in old woodlands of Mediterranean cork oak (Quercus suber)

1. Cork oak landscapes are fascinating ecosystems, historically managed for cork extraction. The persistence in this habitat of many hollow veteran trees provides suitable micro-habitats for saproxylic beetles.
2. We investigated the saproxylic beetle community of two isolated cork oak woodlands of central Italy with different degree of recovery after human transformation: (1) an open woodland and (2) a dense mixed woodland, both dominated by cork oak trees.
3. We found endemic, rare and threatened saproxylic beetles in both the areas, confirming the important conservation value of cork oak landscapes. In the open woodland we observed a higher number of species in all trophic categories, except for mycophagous specialists. Several microhabitat variables reflected the different stage of recovery of the two woodlands.
4. Our findings suggest the crucial role of diversified environments in protected areas: even a small difference in the degree of recovery (i.e., tree closeness) can affect the number of beetle species. Specifically, we found (1) more xerophilous species in the open woodland and (2) more mesophilous species in the dense mixed woodland.

     

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.     

Beetle species richness along the forest productivity gradient in northern Finland

The occurrence and habitat associations of the majority of invertebrate groups in boreal forests are poorly known, even though these groups represent perhaps over 99% of the animal species diversity in the forests. We studied the beetle (Coleoptera) fauna of four forest site types in northern Finland: in spruce mires, herb rich, mesic and sub-xeric forests. We sampled beetles in 32 study sites with five window and five pitfall traps in each. We describe the species abundance and diversity patterns within and among forest types and relate these patterns to structural components of the forests. The volume of decaying wood varied from 14 to 93 m³ ha⁻¹ among sampling sites. The total beetle catch consisted of 100 333 individuals and 435 species. The beetle species richness did not vary according to site fertility but the number of specimens increased with increasing fertility in heath forest sites. The richness of beetle species correlated only weakly with any of the stand structure characteristics at the stand level. Nevertheless, the detrended correspondence analysis (DCA) indicated that different beetle assemblages are characteristic of different forest types. The high level of beta-diversity in beetles among forest types indicates that focusing exclusively on, for example, key-biotopes (presumed biodiversity hotspots) when selecting areas to be set aside would result in a situation where a large proportion of species, even of the rare and threatened ones, is not included in this network of protected areas. This suggests that the complementary set of different forest types may be the best general strategy to maintain the overall beetle species diversity in boreal forests.     

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.     

Revisitation of sites surveyed 19 years ago reveals impoverishment of longhorned beetles in natural and planted forests

As planted forests expand in area, they are beginning to dominate landscapes as a matrix and cause the fragmentation of remaining natural forests. To understand and predict the responses of biological assemblages to maturing planted landscapes, examining the effects of forest type (natural vs planted) and forest age on such assemblages is particularly important. Therefore, to document the effects of forest type and age on longhorned beetle assemblages, in 2008 we collected beetles in broad‐leaved natural and cedar planted forests where beetles had also been collected in 1989. Beetle species composition differed greatly between the two forest types in 1989, whereas this difference was less pronounced in 2008. Species richness and total abundance were higher in natural forests than in planted forests in 1989. In 2008, species richness had decreased in both forest types, but the difference between the two forest types had been maintained. Total abundance was also markedly lower in 2008, and the difference between forest types was much smaller. Although larval host plants were not associated with the responses of species to year (forest age or maturation), beetle species whose larvae fed on either broad‐leaved or coniferous trees (or both) exhibited slight preferences for natural forests. These results suggest that longhorned beetle assemblages become impoverished in planted landscapes as the planted matrix matures. Changes in species composition with forest maturation may be difficult to predict based on larval host plants. However, consideration of larval host plants may enable the prediction of changes in species composition caused by the replacement of natural forests by planted forests.     

Ten years after: Release cutting around old oaks still affects oak vitality and saproxylic beetles in a Norway spruce stand

1. It is often suggested to release oaks (Quercus robur) from competition, to ensure their survival and boost their conservational value. However, few studies have explored how long-lasting this effect is and how it affects saproxylic beetles.
2. Ten years after cutting, we investigated effects of different release cutting levels (high, medium, and no release) around 140-year-old retained oaks in a commercial forest with Norway spruce (Picea abies).
3. We evaluated oak vitality using crown and dead wood measurements. Saproxylic beetles were caught in window traps, identified to species level, and grouped according to their association with oak and/or Norway spruce.
4. Released oaks had more light, higher temperatures, greater crown growth, and less dead wood in the crown compared to the no-release control.
5. After 10 years we still found a higher abundance of oak-associated beetles and higher overall species richness of saproxylic beetles in the released oaks. Beetle species composition significantly differed between released oaks and control.
6. We suggest avoiding planting trees beneath retained oaks when regenerating conifer forests and to proceed with conservation management during subsequent thinning by removing regeneration under the oak crowns. We emphasize the benefits of monitoring the retained oaks to maintain tree vitality, habitat quality, and insolation.

     

Non-native plantation forests as alternative habitat for native forest beetles in a heavily modified landscape

The once extensive native forests of New Zealand’s central North Island are heavily fragmented, and the scattered remnants are now surrounded by a matrix of exotic pastoral grasslands and Pinus radiata plantation forests. The importance of these exotic habitats for native biodiversity is poorly understood. This study examines the utilisation of exotic plantation forests by native beetles in a heavily modified landscape. The diversity of selected beetle taxa was compared at multiple distances across edge gradients between each of the six possible combinations of adjacent pastoral, plantation, clearfell and native forest land-use types. Estimated species richness (Michaelis–Menten) was greater in production habitats than native forest; however this was largely due to the absence of exotic species in native forest. Beetle relative abundance was highest in clearfell-harvested areas, mainly due to colonisation by open-habitat, disturbance-adapted species. More importantly, though, of all the non-native habitats sampled, beetle species composition in mature P. radiata was most similar to native forest. Understanding the influence of key environmental factors and stand level management is important for enhancing biodiversity values within the landscape. Native habitat proximity was the most significant environmental correlate of beetle community composition, highlighting the importance of retaining native remnants within plantation landscapes. The proportion of exotic beetles was consistently low in mature plantation stands, however it increased in pasture sites at increasing distances from native forest. These results suggest that exotic plantation forests may provide important alternative habitat for native forest beetles in landscapes with a low proportion of native forest cover.     

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.     

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.     

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.     

Specialists in ancient trees are more affected by climate than generalists

Ancient trees are considered one of the most important habitats for biodiversity in Europe and North America. They support exceptional numbers of specialized species, including a range of rare and endangered wood‐living insects. In this study, we use a dataset of 105 sites spanning a climatic gradient along the oak range of Norway and Sweden to investigate the importance of temperature and precipitation on beetle species richness in ancient, hollow oak trees. We expected that increased summer temperature would positively influence all wood‐living beetle species whereas precipitation would be less important with a negligible or negative impact. Surprisingly, only oak‐specialist beetles with a northern distribution increased in species richness with temperature. Few specialist beetles and no generalist beetles responded to the rise of 4°C in summer as covered by our climatic gradient. The negative effect of precipitation affected more specialist species than did temperature, whereas the generalists remained unaffected. In summary, we suggest that increased summer temperature is likely to benefit a few specialist beetles within this dead wood community, but a larger number of specialists are likely to decline due to increased precipitation. In addition, generalist species will remain unaffected. To minimize adverse impacts of climate change on this important community, long‐term management plans for ancient trees are important.