Short‐term effect of selection cutting in boreal balsam fir forest on cerambycid and scolytid beetles

The ecology and management of boreal forest ecosystems are drawing greater attention worldwide as their importance is being increasingly recognized for carbon sequestration or for harbouring the world's largest remaining intact forests. Selection cuts have been introduced as a more socially acceptable silvicultural method to improve the maintenance of habitat structure and functions as they mimic aspects of boreal forest succession dynamics. Many studies have shown that selection cutting helps maintaining arthropod communities in mature forests, but few have examined the increased risks of damage by bark‐ and wood‐boring insects in boreal forests of eastern North America. We used multidirectional flight‐interception traps to quantify the response of these beetles to 25 and 40% selection cutting in a balsam fir–white birch forest of Québec, Canada. The abundance and species number of both cerambycid and scolytid beetles were 5–6 times larger in selectively cut stands than in controls the year following treatments. Analyses revealed that bark‐ and wood‐boring beetles’ response was mostly associated with increased canopy openness in selectively cut stands (and sun‐exposed locations within them) and residual tree injuries caused by harvesting operations. These conditions attracted beetles such as Trypodendron lineatum (Scolytinae) and Rhagium inquisitor (Cerambycidae), two species known for their ability to attack weakened, dying and dead hosts. Most species were more abundant in selection cuts, except for Evodinus m. monticola (Cerambycidae) whose abundance was strongly reduced after treatment. Some beetles can have detrimental effects on residual trees and thus could reduce timber value, but most species found in treated stands do not represent a high risk for healthy trees. Thus, selection cuts do not seem favourable to the establishment of tree‐killing beetles. However, as they were found more active/abundant after selection cutting, it would be wise to further study their population dynamics over mid‐ and long‐term periods, along with the ecological and economic implications associated with this silvicultural treatment.     

Effects of different tree harvesting intensities on saproxylic beetle diversity in coniferous and broad-leaved mixed forest in the Changbai Mountain,Northeast, China

1. Saproxylic beetles have gained increasing attention due to their role in the decomposition of rotting wood in forests. Studying the response of saproxylic beetles to tree harvesting is important for developing harvesting strategies that consider conservation of saproxylic beetle diversity.
2. We report results from a case study in which we designed four treatment stands to test the effects of forest harvesting intensity on saproxylic beetle diversity, harvest intensities of 0% untreated control (CK), 17.2% light harvest (LT), 34.7% moderate harvest (MT) and 51.9% high harvest (HT). Flight intercept traps were used to collect specimens of saproxylic beetles in each stand at 2, 3 and 4 years post-harvest.
3. The richness and abundance of saproxylic beetles were higher in MT and HT than in CK. Twelve of the 15 indicator species were significantly associated with MT or HT, whereas only three species were significantly associated with CK.
4. We found that moderate and high intensity harvesting affected the composition and increased beetle abundance and richness of saproxylic beetles, and light intensity harvesting had no effect on the beetle community in years 2, 3 and 4 post-harvest. There was no difference in the beetle community composition between moderate and high harvest stands.

     

What factors influence the diversity of saproxylic beetles? A multiscaled study from a spruce forest in southern Norway

The diversity in different groups of obligate saproxylic beetles was related to ecological variables at three levels of spatial scale in mature spruce-dominated forest. The variables were connected to: (i) decaying wood, (ii) wood-inhabiting fungi, (iii) the level of disturbance, (iv) landscape ecology, and (v) vegetational structure. Several strong relationships were found at medium (1 km²) and large scales (4 km²), while only weak relationships were found at a small scale (0.16 ha; 1 ha=10⁴ m²). This may be explained by the local variations in habitat parameters and the high mobilities of many beetle species. Factors connected to decaying wood and wood-inhabiting fungi were clearly the most important factors at all scale levels. In particular, the variables diversity of dead tree parts, number of dead trees of large diameter and number of polypore fungi species increased the species richness of many groups and increased the abundance of many species. Eight species were absent below a certain density of decaying wood per 1 or 4 km². Former extensive cutting was a negative factor at large scale, probably because of decreasing recolonization with increasing distance to the source habitats. Thinning reduced the diversity of species associated with birch. The development of guidelines favouring the diversity of saproxylic beetles are discussed below.     

Biodiversity conservation in old-growth boreal forest: black spruce and balsam fir snags harbour distinct assemblages of saproxylic beetles

Biodiversity conservation of forest ecosystems strongly relies on effective dead wood management. However, the responses of saproxylic communities to variations in dead wood characteristics remains poorly documented, a lack of knowledge that may impede the development of efficient management strategies. We established the relationship between saproxylic beetles—at the species and community levels—and attributes of black spruce and balsam fir in old-growth boreal forests. The relationship was first evaluated for individual snag bole segments, and then for forest stands. A total of 168 bole sections were collected in summer 2006 along a compositional gradient ranging from black spruce-dominated stands to balsam fir-dominated ones, in a boreal forest dominated by >90-year-old stands. A total of 16,804 beetles belonging to 47 species emerged from bole segments, with 21% of the species being found exclusively in black spruce snags and 36% exclusively in balsam fir snags. Black spruce and balsam fir snags thus contributed differently to forest biodiversity by being inhabited by different saproxylic communities. Wood density was an important attribute in the host-use patterns for several species of saproxylic beetles, but no relationship was found between snag availability within stands and abundance of beetles strongly linked to either black spruce or balsam fir. Our study outlines the relative contribution of tree compositional diversity to saproxylic species, while highlighting the contribution of black spruce and balsam fir to animal diversity in old-growth boreal forests.     

A study of fungus flies (Dipteral Mycetophilidae) in beech woodland

Abstract. 1. A study was made of larval and adult Mycetophilidae in coppiced beech woodland in southern England. Adult mycetophilids were trapped throughout 1968 using sticky traps at ground level. Larvae inhabiting terrestrial fungi were collected from all sporophores from a defined area of the woodland floor between August and October 1968.
2. Larval mycetophilids were extracted from only 12.6% of all sporophores collected and from fifteen of thirty-eight species of fungi present. Some species of fungi were never inhabited by larval mycetophilids while others appeared to be highly attractive to them. Since it was not possible to identify the larvae beyond the level of tribe it was impossible to ascertain host preferences for different species. Peak numbers of larvae occurred at the beginning of September and the beginning of October.
3. Adult mycetophilids had two peaks of activity, the first in March and April when the dominant species were Boletina gripha and Phronia basalis and the second in autumn when species of the genus Mycetophila were dominant. Phronia basalis is a species whose larvae inhabit dead wood and the abundance of this species (40% of all adults) probably reflects the amount of rotting wood in this type of habitat. The autumn peak of adult activity was due to species known to inhabit agaric sporophores and came about 1 month after the peak of larval numbers in fungi.
4. The adult fauna was rich with 107 species representing about a quarter of the known British fauna. However, only twelve species were trapped in sufficient numbers to allow deductions concerning seasonal activity.
5. The results were discussed with respect to the problems of assessing populations of insects inhabiting fungal sporophores and to the previous work on this neglected group.     

Contrasting Responses of Saproxylic Insects to Focal Habitat Resources: The Example of Longhorn Beetles and Hoverflies in Belgian Deciduous Forests

Although both saproxylic longhorn beetles and hoverflies benefit from the presence of woody substrates for reproduction, they differ in their requirements for floral resources and for microbiotopes of overmature and senescent trees. This led us to expect contrasting responses between the two species groups in relation to these essential resources. We examined this prediction in 22 mature oak- and beech-dominated stands of southern Belgium by relating their species assemblages to local vegetation structure and composition, altitude and landscape composition. Stands were organised in pairs as a function of their overall dead wood supply. Free-hanging window traps, stump emergence traps and Malaise traps produced 30 longhorn beetle species (1637 individuals) and 106 hoverfly species (3020 individuals). Paired-comparisons controlling for annual variation in captures showed that, unlike saproxylic hoverflies, stands with dead wood hosted more species and individuals of longhorn beetles. Accordingly, the two species groups were found to be independent on ordination axes, responding to different sets of environmental conditions. While stands dominated by oaks with a high snag volume were highly favoured by longhorn beetles, saproxylic and threatened syrphids were limited to open-stands with large trees and a well-developed, species rich herb layer providing the floral resources required for their reproduction. Our results suggest that, when defining criteria to identify or restore important habitats for saproxylic insect conservation, variables related to different aspects of dead wood supply should not be the only criteria taken into account.     

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.     

Forest restoration by burning and gap cutting of voluntary set-asides yield distinct immediate effects on saproxylic beetles

Today, the importance of restoring natural forest disturbance regimes and habitat structures for biodiversity is widely recognized. We evaluated the immediate effects of two restoration methods on wood-inhabiting (saproxylic) beetles in boreal forest voluntary set-asides. We used a before-after control-impact experimental set-up in 15 set-asides; each assigned to one of three treatments: (1) restoration burning, (2) gap cutting and (3) no-treatment reference stands. Before treatment, abundance, species richness and assemblage composition of trapped beetles did not differ significantly among treatments. Burning resulted in a significant change in assemblage composition and increased species richness and abundance compared to reference stands. As predicted, saproxylic species known to be fire favoured increased dramatically after burning. The immediate response shows that, initially, fire favoured species are attracted from the surrounding landscape and not produced on site. Gap cutting increased the abundance of cambium consumers but had no significant effect on total species richness or assemblage composition of saproxylic beetles. The stronger effect of burning compared to gap cutting on saproxylic assemblages is probably due to the very specific conditions created by fires that attracts many disturbance-dependent species, but that at the same time disfavour some disturbance-sensitive species. By contrast, gap cutting maintained assemblage composition, increased abundances and is likely to increase species richness in the years to follow, due to elevated level of dead wood. The restoration methods applied in this study may prove particularly useful, partly because of positive effect on saproxylic beetles, but also due to the cost-efficiency of the measures; the voluntary set-asides were already established and the restoration costs fully covered by revenue from the extracted timber.     

Saproxylic insects and fungi in deciduous forests along a rural–urban gradient

Urbanization is increasing worldwide and is regarded a major threat to biodiversity in forests. As consequences of intensive human use, the vegetation structure of naturally growing urban forests and their amount of deadwood can be reduced. Deadwood is an essential resource for various saproxylic insects and fungi. We assessed the effects of urbanization and forest characteristics on saproxylic insects and fungi. We exposed standardized bundles consisting of each three freshly cut beech and oak branches in 25 forests along a rural–urban gradient in Basel (Switzerland). After an exposure of 8 months, we extracted the saproxylic insects for 10 months using an emergence trap for each bundle. We used drilling chips from each branch to determine fungal operational taxonomic units (OTUs). In all, 193,534 insect individuals emerged from the experimental bundles. Our study showed that the abundance of total saproxylic insects, bark beetles, longhorn beetles, total flies, moths, and ichneumonid wasps decreased with increasing degree of urbanization, but not their species richness. However, the taxonomic composition of all insect groups combined was altered by wood moisture of branches and that of saproxylic beetles was influenced by the degree of urbanization. Unexpectedly, forest size and local forest characteristics had a minor effect on saproxylic insects. ITS (internal transcribed spacer of rDNA) analysis with fungal specific primers revealed a total of 97 fungal OTUs on the bundles. The number of total fungal OTUs decreased with increasing degree of urbanization and was affected by the volume of naturally occurring fine woody debris. The composition of fungal OTUs was altered by the degree of urbanization and pH of the branch wood. As a consequence of the altered compositions of saproxylics, the association between total saproxylic insects and fungi changed along the rural–urban gradient. Our study shows that urbanization can negatively impact saproxylic insects and fungi.     

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.     

Role of the Siberian flying squirrel as an umbrella species for biodiversity in northern boreal forests

One of the potentially useful indirect shortcut methods in biodiversity conservation is the umbrella species concept. An umbrella species can be seen relatively demanding for the size of the area and probably also for certain habitat types: conservation management for the umbrella species would thus encompass other species preferring similar habitats but with smaller area requirements. As such, it has a comprehensive spatial aspect for landscape planning. We tested the role of the Siberian flying squirrel (Pteromys volans) as an umbrella species for wood dependent species among red-listed and old-growth forest associated polypores, epiphytic lichens and beetles. Flying squirrels inhabit home ranges of several to tens of hectares, and prefer mature spruce-dominated (Picea abies) mixed forests, which often have high amounts of dead wood. We carried out species surveys and trappings during 1 year from 20 mature spruce-dominated forest stands (altogether 162 ha), of which 12 were occupied by the flying squirrel. The amount of dead wood was higher in occupied stands than in unoccupied stands. We also found a tendency for a higher number of species and number of records in occupied stands, a relationship mostly due to the polypore species. The presence of the flying squirrel may reflect the habitat availability for species depending on dead and living wood, and assist in site selection of conservation areas. We suggest that the flying squirrel has potential as an umbrella species to partly enhance maintenance of biodiversity in northern boreal forests in Finland.     

Plants, insects and birds in semi-natural pastures in relation to local habitat and landscape factors

The preservation of remaining semi-natural grasslands in Europe has a high conservation priority. Previously, the effects of artificial fertilisation and grazing intensity on grassland animal and plant taxa have been extensively investigated. In contrast, little is known of the effects of tree and shrub cover within semi-natural grasslands and composition of habitats in the surrounding landscape on grassland taxa. We evaluated the effect that each of these factors has on species richness and community structure of vascular plants, butterflies, bumble bees, ground beetles, dung beetles and birds surveyed simultaneously in 31 semi-natural pastures in a farmland landscape in south-central Sweden. Partial correlation analyses showed that increasing proportion of the pasture area covered by shrubs and trees had a positive effect on species richness on most taxa. Furthermore, species richness of nectar seeking butterflies and bumble bees were negatively associated with grazing intensity as reflected by grass height. At the landscape level, species richness of all taxa decreased (butterflies and birds significantly so) with increasing proportion of urban elements in a 1-km² landscape area centred on each pasture, while the number of plant and bird species were lower in landscapes with large proportion of arable fields. Our results differed markedly depending on whether the focus was on species richness or community structure. Canonical correspondence analyses (CCA) showed that the abundance of most taxa was ordered along a gradient describing tree cover within pastures and proportion of arable fields in the landscape. However, subsets of grassland birds and vascular plants, respectively, showed markedly different distribution patterns along axis one of the CCA. In contrast to current conservation policy of semi-natural pastures in Sweden, our results strongly advise against using a single-taxon approach (i.e., grassland vascular plants) to design management and conservation actions in semi-natural pastures. Careful consideration of conservation values linked to the tree and shrub layers in grasslands should always precede decisions to remove trees and shrubs on the grounds of promoting richness of vascular plants confined to semi-natural grasslands. Finally, the importance of landscape composition for mobile organisms such as birds entails that management activities should focus on the wider countryside and not exclusively on single pastures.     

Hot spots, indicator taxa, complementarity and optimal networks of taiga

If hot spots for different taxa coincide, priority-setting surveys in a region could be carried out more cheaply by focusing on indicator taxa. Several previous studies show that hot spots of different taxa rarely coincide. However, in tropical areas indicator taxa may be used in selecting complementary networks to represent biodiversity as a whole. We studied beetles (Coleoptera), Heteroptera, polypores or bracket fungi (Polyporaceae) and vascular plants of old growth boreal taiga forests. Optimal networks for Heteroptera maximized the high overall species richness of beetles and vascular plants, but these networks were least favourable options for polypores. Polypores are an important group indicating the conservation value of old growth taiga forests. Random selection provided a better option. Thus, certain groups may function as good indicators for maximizing the overall species richness of some taxonomic groups, but all taxa should be examined separately.     

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.

     

Litter‐dwelling beetles (Insecta: Coleoptera) can survive in clear‐cutting during subsequent soil ploughing

1. A 2‐year study of litter dwelling beetles was conducted in different mature pine stands and clear‐cuts in Lithuania using the litter sifting method. We hypothesized that clear‐cutting and subsequent ploughing would increase species diversity and the abundance of beetles, and also would encourage the immediate appearance of early‐successional beetle species replacing late successional species in the clear‐cuts.
2. We did not confirm a hypothesis regarding increase in the number of species and abundance of beetles in clear‐cuts.
3. Our hypothesis regarding the immediate appearance of early successional species and disappearance of late‐successional species in clear‐cuts was confirmed.
4. We also revealed that subsequent soil ploughing in clear‐cuts did not accelerate this process, which was linked to the possibility of late successional species surviving in the undisturbed spaces between the strips of ploughed soil.
5. The present study shows that late‐successional forest litter‐dwelling coleopteran species of old pine forests have a better chance of survival in a ploughed clear‐cut, at least in northern Europe, than has been reported in other studies.

     

Patterns of local wood use and cutting of Philippine Mangrove Forests

Small-scale wood harvesting from mangrove forests is a commonplace yet barely studied phenomenon. This paper integrates bio-ecological and ethnographic methods to examine local wood use and cutting of mangrove forests in two areas of the Philippines. Findings reveal considerable site variation in cutting intensity, with heavier cutting typically closer to settlements and in forest stands that are not effectively regulated by government or private interests. Overall, cutting is responsible for almost 90% of stem mortality in both natural and plantation forests. Field measurements confirm ethnographic evidence indicating that harvesting for construction wood, but not fuelwood, is both species- and size-selective. Mangrove management and conservation efforts can be made more effective by better understanding how local people are harvesting wood resources from these forests.     

High plant species richness indicates management-related disturbances rather than the conservation status of forests

There is a wealth of smaller-scale studies on the effects of forest management on plant diversity. However, studies comparing plant species diversity in forests with different management types and intensity, extending over different regions and forest stages, and including detailed information on site conditions are missing. We studied vascular plants on 1500 20 m × 20 m forest plots in three regions of Germany (Schwäbische Alb, Hainich-Dün, Schorfheide-Chorin). In all regions, our study plots comprised different management types (unmanaged, selection cutting, deciduous and coniferous age-class forests, which resulted from clear cutting or shelterwood logging), various stand ages, site conditions, and levels of management-related disturbances. We analyzed how overall richness and richness of different plant functional groups (trees, shrubs, herbs, herbaceous species typically growing in forests and herbaceous light-demanding species) responded to the different management types. On average, plant species richness was 13% higher in age-class than in unmanaged forests, and did not differ between deciduous age-class and selection forests. In age-class forests of the Schwäbische Alb and Hainich-Dün, coniferous stands had higher species richness than deciduous stands. Among age-class forests, older stands with large quantities of standing biomass were slightly poorer in shrub and light-demanding herb species than younger stands. Among deciduous forests, the richness of herbaceous forest species was generally lower in unmanaged than in managed forests, and it was even 20% lower in unmanaged than in selection forests in Hainich-Dün. Overall, these findings show that disturbances by management generally increase plant species richness. This suggests that total plant species richness is not suited as an indicator for the conservation status of forests, but rather indicates disturbances.     

“Primeval forest relict beetles” of Central Europe: a set of 168 umbrella species for the protection of primeval forest remnants

Identification of forest stands with priority for the conservation of biodiversity is of particular importance in landscapes with a long cultural and agricultural history, such as Central Europe. A group of species with a high indicator value for the naturalness of forest ecosystems are saproxylic insects. Some of these species, especially within the order Coleoptera, have been described as primeval forests relicts. Here, we compiled a list of 168 “primeval forest relict species” of saproxylic beetles based on expert knowledge. These species can serve as focal and umbrella species for forest conservation in Central Europe. They were selected because of their dependence on the continuous presence of primeval forest habitat features, such as over-mature trees, high amounts of dead wood, and dead wood diversity, as well as their absence in managed Central European forests. These primeval forest relict species showed a moderately strong clumping pattern within the phylogeny of beetles, as indicated by phylogenetic signal testing using the D-statistic. When we controlled for phylogenetic relatedness, an ordinal linear model revealed that large body size and preference for dead wood and trees of large diameter are the main characteristics of these species. This list of species can be used to identify forest stands of conservation value throughout Central Europe, to prioritize conservation and to raise public awareness for conservation issues related to primeval forests.     

Mycetophilidae (Diptera), an insect group vulnerable to forestry practices? A comparison of clearcut,managed and semi-natural spruce forests in southern Norway

The mycetophilid fauna and environmental variables were studied at 15 sites within a spruce forest in southern Norway. There were five replications of each of the following categories: semi-natural forests, clearcuts, and managed forests (clearcut 70–120 years ago). Clearcutting seems to induce a long lasting effect on the Mycetophilidae fauna. The semi-natural forests were more speciesrich and contained more potentially rare species than the two other categories. Even though managed forests and clearcuts differed in faunal composition, their species richness was not significantly different. Continuity is probably a main factor for maintaining the diversity of Mycetophilidae species. Lumping the 15 sites together, the number of species was strongly correlated with the (temporal) continuity of tree cover and substrates, which may reflect an increased diversity of fungal habitats both in dead wood and on the ground. The degree of continuity was recognized by means of indicator species of fungi, lichen and vascular plants. The amount of dead wood was correlated with the species diversity, but was probably dependent on the continuity factor, since clearcuts with much dead wood had relatively fewer species.Important elements in a strategy for conservation of the diversity of mycetophilid species seem to be: (i) to identify and protect the remnant patches of forests with long continuity, (ii) as far as possible, to practice timber harvesting in earlier clearcut forests instead of semi-natural forests.     

Long‐term development of species richness in a central European beech (Fagus sylvatica) forest affected by windthrow—Support for the intermediate disturbance hypothesis?

On the basis of long‐term surveys of permanent plots and traps, we examined the communities of saproxylic beetles, fungi, herbs, and trees on an untreated 22 ha large beech forest windthrow and asked whether the results lend support to the intermediate disturbance hypothesis (IDH). We studied species richness and the similarity of community composition. Additionally, we grouped species by their frequency trend over time to successional model types to examine whether, corresponding to the IDH, the diversity of these groups explained peak richness at intermediate intervals after the disturbance. In line with the IDH, species richness showed a hump‐backed temporal course for alpha and gamma diversity. We found evidence for a linear succession directly after the disturbance. This, however, did not continue, and in all species groups, a partial recovery of the initial community was observed. In the case of fungi, herbs, and trees, but not for saproxylic beetles, alpha diversity was driven by the diversity of the successional model types. Our results underline that the mechanisms driving species richness after disturbances are more complex than the IDH suggests and that these mechanisms vary with species group. We assumed that, besides competition, legacy effects, facilitation, habitat heterogeneity, and random saturation of the species pool are important. In case of trees and herbs, we found indications for strong legacy and competition effects. For fungi and beetles, substrate heterogeneity and microclimate were assumed to be important. We concluded that disturbances contribute to increasing species richness not only by reducing the effectiveness of competitors but also by increasing the amount and diversity of resources, as well as their rate of change over time.