Influence of tree hollow characteristics on saproxylic beetle diversity in a managed forest

Tree hollows are key structures in forest ecosystems constituting long-lasting habitats and nutritional resources for many saproxylic arthropod species. Due to diverse microhabitat structures and conditions in tree hollows, they can support a broad range of species. However, in the past intensive management practices in parts of Europe reduced the abundance of tree hollows, resulting in a decrease and endangerment of species specialised in this tree habitat. We investigated 40 beech trees with hollows in 2014 and a subset of 23 of these trees in 2015 in a managed forest landscape in Germany. Using emergence traps we collected 89 beetle species of which 33% were on the Bavarian Red List. We described the tree characteristics, physical hollow characteristics, and their surrounding environment investigating their influence on α-diversity of non-Red List and Red List species. Furthermore, we investigated spatial (between tree hollows) and temporal (same tree hollow but different years) β-diversity, considering the importance of turnover and nestedness components on β-diversity. α-Diversity decreased with increasing decomposition of wood mould and increased with increasing area of hollow entrance in both years. Additional characteristics differed between years and between non-Red List and Red List species. β-Diversity was related to diameter at breast height, number of surrounding tree hollows, area of hollow entrance and a temperature gradient. We found a higher species turnover than nestedness between tree hollows and between years, indicating highly dynamic beetle communities spatially as well as temporally. To support and maintain the diversity of saproxylic beetles inhabiting tree hollows, the heterogeneity of microhabitats is important and should be supported by maintaining the diversity of differently structured and sized tree hollows.     

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.     

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.     

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.     

Influence of tree hollow characteristics on the diversity of saproxylic insect guilds in Iberian Mediterranean woodlands

Saproxylic diversity assessment is a major goal for conservation strategies in woodlands and it should consider woodland composition and configuration at site and tree level as key modelling factors. However, in Mediterranean woodlands little is known about the relation with the environmental factors that structure their assemblages, especially those linked to tree hollow microhabitats. We assessed the diversity of Syrphidae (Diptera) and Coleoptera saproxylic guilds that co-occurred in tree hollows located in three different Iberian Mediterranean woodlands in the Cabañeros National Park (Spain). Furthermore, we evaluated how differences in tree hollow microenvironmental variables (understood as the physical and biotic characteristics of a hollow and tree individual) influenced saproxylic guild diversity both within and among woodland sites. We found that woodland sites that provided greater heterogeneity of trees and hollow microhabitats determined higher saproxylic guild diversity. Nevertheless, certain species or even complete guilds can be favoured in woodlands where some hollow microhabitats predominate as a consequence of historical tree management. In general, hollow volume was the main determining factor for saproxylic guild richness and abundance in woodland sites, and large hollow volume was usually related to higher diversity, which highlighted the importance of multi-habitat hollow trees. Moreover, saproxylic guilds also responded to other different microenvironmental variables, which indicated different ecological preferences among guilds. The conservation of saproxylic insects in Iberian Mediterranean areas must be addressed to protect woodland sites that provide high diversity and large numbers of tree hollow microhabitats, and practices to enhance microhabitat heterogeneity should even be encouraged.     

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

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Influence of sampling effort on saproxylic beetle diversity assessment: implications for insect monitoring studies in European temperate forests

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Influence of snag characteristics on saproxylic beetle assemblages in a south Swedish beech forest

Effects of snag characteristics on saproxylic beetles were studied in an area of managed beech forest in southern Sweden. A snag survey was combined with a beetle survey using 30 small window traps directly attached to beech snags. The total number of species was lower in the snags which were most decayed than in the three younger decay classes, while the number of red-listed species remained the same regardless of the stage of decay. The number of fresh wood species declined and the number of rot hole species increased with increasing snag decay. The diversity of fungicolous and decayed wood species peaked at the intermediate stage of decay in the snags. CCA ordination confirmed that the stage of decay in the snags was most important for species composition, followed by sun exposure. There were no general differences in species density and composition between managed and unmanaged stands. Our study suggests that most species are able to find suitable habitat within a radius of a few kilometres and that the total amount of habitat in an unfragmented forest area is more important for species diversity than the spatial distribution of this habitat. Our data also shows that species diversity increases with habitat diversity. Snags formed from giant beech trees seem to be particularly important for rare species living in rot holes. We conclude that for a high species diversity there is a requirement for snags in different stages of decay, size and degree of sun exposure.     

Dispersal limitation of saproxylic insects in a managed forest? A population genetics approach

Dispersal is a key trait of species enabling gene flow among populations. For species persistence dispersal may therefore be crucial especially in a patchy or changing environment. Tree hollows are a patchy habitat as their number is locally limited and in addition to that, habitat quality of tree hollows differs. Both factors are important for colonization by saproxylic insects as species may be specialized on a particular age or quality of a tree hollow and may be dispersal-limited. Intensive forest management may further decrease the number of tree hollows and increase distances between them, if trees with hollows are removed. However, not much is known about the dispersal abilities of most saproxylic species, even though such knowledge could improve conservation efforts. To investigate the genetic population structure of saproxylic species, we collected saproxylic organisms with emergence traps from 40 hollows in beech trees in a managed forest in Germany (approximately 14 × 14 km). We focused on three threatened species, one coleopteran and two dipteran (Anaspis ruficollis, Scraptiidae; Criorhina floccosa, Syrphidae; Xylomya maculata, Xylomyidae) emerging from our tree hollows. Microsatellite analysis was used to assess gene flow among tree hollows and population genetic structure. In contrast to other studies reporting limited dispersal in saproxylic insects, we found no indication for restricted gene flow in all three species investigated. However, we studied relatively abundant species and our study site may not have been large enough to detect genetic substructure. This study indicates that the amount and quality of a suitable habitat may at least in some cases be more limiting than the physical ability to disperse.     

亚热带森林树种多样性对凋落叶分解胞外酶活性的影响

森林生物多样性与生态系统功能关系是当前群落生态学的热点研究领域。然而, 以往研究更多聚焦在森林植物多样性丧失对群落生产力的影响, 而对森林凋落物分解的相关研究稍显不足。森林凋落叶分解的快慢直接受控于凋落物分解者分泌的胞外酶的活性, 后者更是指示森林生态系统养分循环的重要指标之一。本研究依托我国江西亚热带森林生物多样性与生态系统功能控制实验, 通过对不同植物多样性梯度样方内目标树种凋落叶胞外酶活性、理化性质以及腐生真菌的分析, 探索树种多样性丧失对胞外酶活性的影响及其调控机制, 以探讨森林树种多样性对地表、地下生态过程和功能的影响。结果表明, 样方水平树种多样性丧失显著影响胞外酶的活性, 除单种样方外, 随着样方水平树种丰富度的增加, 胞外酶活性呈现出增长趋势; 与碳周转相关的α-葡萄糖苷酶(AG)、β-葡萄糖苷酶(BG)、纤维二糖水解酶(CB)在树种多样性最大时活性达到最高; 而木糖苷酶(XS)以及与氮、磷和顽拗有机养分分解相关的N-乙酰-β-氨基葡萄糖苷酶(NAG)、酸性磷酸酶(AP)和多酚氧化酶(PPO)在树种多样性较低时活性较高。针对目标树种周围的邻居树种多样性进一步分析发现, 各胞外酶活性随着邻居树种多样性的变化呈“单峰”响应趋势, 酶活性大多在邻居树种丰富度为6时呈现峰值。研究发现真菌分解者在胞外酶活性对植物多样性的响应上可能存在重要的调控作用, 可以推测树种多样性通过改变腐生真菌分解者的群落结构和多度, 从而影响胞外酶活性。     

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.     

Spatial heterogeneity of tree diversity response to climate warming in montane forests

Many studies reported biotic change along a continental warming gradient. However, the temporal and spatial change of tree diversity and their sensitivity to climate warming might differ from region to region. Understanding of the variation among studies with regard to the magnitude of such biotic changes is minimal, especially in montane ecosystems. Our aim is to better understand changes in spatial heterogeneity and temporal dynamics of mountain tree communities under climate warming over the past four decades. In 2017, we resurveyed and recorded all tree species from 107 long‐term monitoring plots that were first studied between 1974 and 1976. These plots were located in montane forests in the Giant Panda National Park (GPNP), China. Our results showed that spatial differences were found in tree species diversity changes response to mean annual temperature change over the past four decades. Tree species richness increased significantly under climate warming in Minshan (MS) and Xiaoxiangling (XXL) with higher warming rate than Qionglai (QLS) and Liangshan (LS). The trees species diversity in MS and XXL were more sensitive to climatic warming. MS and XXL should receive priority protection in the next conservation plan of the GPNP. The GPNP should avoid taking a “one‐size‐fits‐all” approach for diversity conservation due to spatial heterogeneity in plant community dynamics.     

Decaying wood in tree trunk hollows as a natural substrate for Cryptococcus neoformans and other yeast-like fungi of clinical interest

The occurrence of Cryptococcus neoformans var. neoformans and other yeast-like fungi of clinical interest in decaying wood inside tree trunk hollows, bark and other plant materials is reported. The var. neoformans was isolated from 3 of 45 (6.6%) wood and one of 390 Eucalyptus bark samples. Two of the positive wood samples came from a tree trunk hollow of Butea monosperma (Family: Papilionaceae) growing in Roshan Ara Garden, Old Delhi whereas the third was from a trunk hollow of Tamarindus indica (Family: Papilionaceae) growing outside of Talkatora Garden, New Delhi. The solitary positive Eucalyptus bark sample originated from Amritsar. The isolations of var. neoformans from decaying wood inside trunk hollows of B. monosperma and T. indica constitute the first record of the natural occurrence of this pathogen in association with these trees. The observation reinforces the recent evidence for decaying wood inside trunk hollows of some trees to be a new natural habitat of the variety neoformans. Besides, in consonance with their essentially saprobic character, a number of other yeast-like fungi were sporadically isolated. This includes, Cryptoccus laurentii Cryptococcus albidus Candida lusitaniae C. guilliermondii C. krusei C. tropicalis C. zeylanoides Trichosporon cutaneum Rhodotorula mucilaginosa R. glutinis Geotrichum capitatum G. klebahnii and Sporobolomyces salmonicolor. Cryptococcus neoformans var. gattii was not found in any of the 702 samples of plant materials, including the bark and detritus of Eucalyptus camaldulensis and E. tereticornis trees. A more extensive environmental survey, covering divergent climatic regions, is warranted to identify the natural reservoirs of var. gattii in India. This revised version was published online in August 2006 with corrections to the Cover Date.     

Influence of forest development phase on centipede diversity in managed beech forests in Slovenia

Biodiversity in managed forests depends on management practices that determine the forest structure. The influence of stand structure on centipede diversity was investigated by comparing centipede communities from different forest development phases. Using the quadrat counts method, soil samples were taken thrice during 2000 (in spring, summer and autumn) from a deforested area and three evenly aged beech stands (juvenile, pole and timber phases) in managed beech forests in Slovenia. The characteristics of centipede communities and the similarities among them were estimated. The collection comprised 1150 centipedes representing 32 species. At a single site, 10–24 species were found. The average density was between 22 and 808 individuals per m². Differences among communities from different sites were found. The highest species number and abundance were found in the juvenile phase and the lowest in the deforested area. The occurrence of species differed among phases, but none of the species found with a high incidence in one development phase was restricted to that phase. The composition of the centipede community was more sensitive to the successional status of the forest than to the season. A comparison with differently structured forest was made, which suggests that homogenisation leads to reduced centipede diversity. To preserve biodiversity and healthy forest functioning, the maintenance of heterogeneity within a forest stand should be an important focus of management operations.     

元谋干热河谷不同人工林中鞘翅目甲虫多样性比较

在元谋干热河谷采用网扫法调查了云南松林、桉树林、新银合欢林、桉树＋新银合欢林、印楝林及多树种混交的人工林鞘翅目昆虫多样性。结果表明,鞘翅目甲虫标本925号,计71种,分属18个科,其中叶甲科种类最丰富,象甲科数量最丰富。人工林鞘翅目昆虫群落物种丰富度在7～23,Shannon—Wiener多样性指数在1．249～2．562,昆虫多样性总体较低。各样地鞘翅目群落之间为不相似水平。云南松林鞘翅目昆虫群落物种丰富度、Shannon—Wiener指数、Simpson指数及Pielou指数分别为20、2．562、0．104和0．855,其多样性最高,多树种混交林较高,印楝林较低,而桉树林、新银合欢林、桉树＋新银合欢林多样性极低。多树种混交的恢复对昆虫多样性提高有显著促进作用。     

树种及多样性组配对南亚热带人工林早期树木生长的影响

为科学筛选提升南亚热带人工林生产力的树种配置模式,选择南亚热带8个乡土树种,采用随机区组的试验设计,建立了树种多样性梯度(1、2、4、6个树种)人工新造林试验平台,研究树种多样性及不同功能特性树种混交对人工林早期树木生长的影响。结果表明: 在树木生长第5年,树木生长并没有随树种多样性增加而增加;速生树种马尾松和米老排纯林生长量是珍贵树种红椎和格木纯林的2.5～4.5倍;2个树种混交和4个树种混交情况下,针阔树种混交、速生树种与固氮树种混交显著提高树木生长量51.5%～132.8%,而当6个树种混交时,不同树种组配对生长量没有显著影响。不同树种配置模式下土壤氮、磷养分是影响树木早期生长的主要因素。针阔树种混交、速生和固氮树种混交能显著提高南亚热带人工林树木的早期生长。     

Short-term development of ambrosia and bark beetle assemblages following a windstorm in French broadleaved temperate forests

Abstract: In most temperate deciduous forests, windstorm is the main source of dead wood. However, the effects of this natural disturbance on ambrosia and bark beetle communities are poorly known. In managed oak‐hornbeam forests storm‐damaged in France in 1999, we sampled ambrosia (and second bark beetles) by ethanol‐baited window‐flight traps in 2001. By comparing uncleared gaps, undisturbed closed‐canopy controls and seedling‐sapling stands, we investigated the short‐term effects of gap formation, gap size and surrounding landscape to provide a snapshot of scolytid response. Contrary to expectations, neither the abundance nor the richness of ambrosia beetle species was significantly higher in gaps than in undisturbed stands. Few responses in abundance at the species level and only a slight difference in assemblage composition were detected between gaps and closed‐canopy controls. Gaps were more dissimilar from seedling‐sapling stands, than from closed‐canopy controls. More scolytid individuals and species were caught in gaps than in seedling‐sapling stands. Mean local and cumulative richness peaked in mid‐size gaps. Only mid‐size gaps differed from closed‐canopy controls in terms of species composition. We identified generalist gap species (Xyleborus saxesenii, X. cryptographus), but also species significantly more abundant in mid‐size gaps (Platypus cylindrus, Xyloterus signatus). The faunistic peculiarity of mid‐size gaps seemed to be partly related to a bias in oak density among gap size classes. Few landscape effects were observed. Only the scolytids on the whole and X. dispar were slightly favoured by an increasing density in fellings at the 78 ha scale. We did not find any correlation between scolytid abundance and the surrounding closed‐forest percentage area. We confirmed that temperate, deciduous, managed stands did not come under threat by ambrosia and bark beetle pests after the 1999 windstorm. Nonetheless, our data stressed the current expansion in Western Europe of two invasive species, X. peregrinus and especially X. germanus, now the predominant scolytid in the three oak forests studied.     

Influence of tree age, tree size and crown structure on lichen communities in mature Alpine spruce forests

Testing the relations between tree parameters and the richness and composition of lichen communities in near-natural stands could be a first step to gather information for forest managers interested in conservation and in biodiversity assessment and monitoring. This work aims at evaluating the influence of tree age and age-related parameters on tree-level richness and community composition of lichens on spruce in an Alpine forest. The lichen survey was carried out in four sites used for long-term monitoring. In each site, tree age, diameter at breast height, tree height, the first branch height, and crown projection area were measured for each tree. Trees were stratified into three age classes: (1) <100 years old, immature trees usually not suitable for felling, (2) 100–200 years old, mature trees suitable for felling, and (3) >200 years old, over-mature trees normally rare or absent in managed stands. In each site, seven trees in each age class were selected randomly. Tree age and related parameters proved to influence both tree-level species richness and composition of lichen communities. Species richness increased with tree age and related parameters indicative of tree size. This relation could be interpreted as the result of different joint effects of age per se and tree size with its area-effect. Species turnover is also suspected to improve species richness on over-mature trees. Similarly to species richness, tree-level species composition can be partially explained by tree-related parameters. Species composition changed from young to old trees, several lichens being associated with over-mature trees. This pool of species, including nationally rare lichens, represents a community which is probably poorly developed in managed forests. In accordance to the general aims of near-to-nature forestry, the presence of over-mature trees should be enhanced in the future forest landscape of the Alps especially in protected areas and Natura 2,000 sites, where conservation purposes are explicitly included in the management guidelines.