The importance of substrate type, shading and scorching for the attractiveness of dead wood to saproxylic beetles

Modern forestry management has reduced the amount of dead wood in forest ecosystems and this has become a serious threat to flora and fauna. Efforts are therefore being made to reverse this trend but one problem is that we still lack detailed knowledge regarding the substrate requirements of many saproxylic species. In a field experiment, conducted in three forest types (forest reserve, mature managed forest and clear-cut), we evaluated the value, from a conservation perspective, of different substrate types (logs, snags and tops) of Norway Spruce, Picea abies, and if the quality of spruce logs as saproxylic habitats can be improved by simple log treatments (scorching and shading). We collected 9982 individuals representing 262 saproxylic beetle species in window traps. Both substrate type and, to a lesser extent, log treatment had a significant effect on the abundance and species richness of saproxylic beetles attracted to the different dead wood substrates. However, more importantly, the composition of the beetle assemblages differed significantly between both substrates and log treatments. Snags, logs and tops all attracted significantly different beetle assemblages and scorched logs differed from untreated control logs. Sixteen red listed species were trapped, with the highest number (11 species) being found on scorched logs. We found strong evidence that some species preferred a specific substrate type, mainly logs, in some cases treated logs (scorched or shaded), but not snags, the substrate commonly provided for conservation purposes on e.g. clear-cuts. This stresses the importance of conducting forestry in such a way that a multitude of both forest habitats and dead wood substrates are available continuously in the forest landscape to maintain biodiversity.     

The small mammal community of Mukogodo Forest, Kenya

Species richness and diversity of rodents and insectivores were investigated at relict forest patches of Mukogodo, Laikipia, Kenya using Sherman's live traps and pitfall traps. Two hundred and nineteen individuals were captured in 3021 trap‐nights. There were eleven species in two taxonomic groups, Rodentia and Insectivora. Two other rodent species were sighted but not captured. Thirteen bats belonging to four species (Epomophorous wahlbergi, Pipistrellus kuhlii, Scotophilus dingani and Nycteris thebaica) were opportunistically trapped using mist nets. Two of the four species accumulation curves for forest patches did not reach an asymptote. Species richness and diversity were highest at Kurikuri compared with other patches because of habitat variability. The results support the prediction that forest disturbance and degradation lead to an increase in generalist species as compared with specialists and highlight the importance of relict afromontane forests in the conservation of small mammals in Kenya.     

Habitat determinants of the taxonomic and functional diversity of parasitoid wasps

Parasitoid wasps (Hymenoptera) are a highly diverse component of terrestrial habitats, yet very little is known about how their community structure and functional diversity is influenced by habitat variables. Parasitoids were sampled using Malaise traps in early and late summer across ten forest sites in the Waitakere Ranges, New Zealand. At each trap, a range of local habitat variables were measured in a 20 × 20 m plot and landscape cover data were obtained for a 2-km radius. Species from two families, Ichneumonidae and Braconidae, were identified using morphological and molecular methods. Functional groups (idiobiosis, koinobiosis), key life history traits (endoparasitism, ectoparasitism, host taxa), and morphological traits (ovipositor length, body size) were measured to calculate functional diversity indices. The abundance, species richness, composition, and functional groups of parasitoids were chiefly influenced by vegetation type, plant diversity, coarse woody debris, and seasonal sampling. However, different taxa showed different patterns to habitat variables. Functional diversity measures were strongly linked to vegetation type. Kauri conifer forest was found to support a specialised parasitoid community, with lower abundance and species richness than broadleaved forest, but with higher functional evenness and high functional dispersion; indicating a community of co-existing species with a high utilisation of resources. The lack of congruence in the habitat relationships of different parasitoid groups highlights that preservation of a wide range of different forest habitats is required to adequately conserve parasitoid diversity.     

Comparison of old-field and forest revegetation dynamics in Provence

Abstract. This paper describes vegetation changes on cultivation terraces after abandonment, considered as secondary succession, and vegetation responses after clear-cutting, considered as regeneration. Species were grouped in life form and dynamic categories in order to infer dynamic patterns. By applying the Shannon diversity index to these characteristics, an index of functional diversity is obtained. Regenerative succession is considered here as a particular case of secondary succession, characterized by a fast vegetative regeneration of the dominant woody species, which controls the succession of the understorey species. Post-agricultural succession and forest regeneration succession show a similar relation between the diversity index and a forest index, indicating the relative amount of forest species. It appeared that the old-field dynamics resemble the deciduous oak-forest regeneration after the woody species have established. The species diversity at the end of the post-agricultural succession and in the forest is controlled by the nature of the dominant tree species.     

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.     

Significance of woodland key habitats for polypore diversity and red-listed species in boreal forests

Preservation of woodland key habitats has become an integral part of biodiversity-oriented forest management in northern Europe. In Finland, brook-side spruce forests constitute the most important key-habitat type in terms of total area and timber volume. Our aim was to compare polypore diversity and the occurrence of red-listed species between brook-side key habitats and comparable sites of managed forest as controls. Furthermore, we assessed the importance of stand structural features to polypore communities, and contrasted regions with different land-use history. Altogether 69 key habitats and 70 controls were inventoried in four regions across southern Finland. A total of 28,023 surveyed dead-wood units had 3,307 occurrences of 114 species of which 25 species were red-listed. Key habitats hosted on average 28% (about two species) more polypore species (mainly deciduous-specialists) than controls because of a larger average volume of dead wood and a higher proportion of dead deciduous wood. However, the average number of red-listed species was low and did not differ between the two forest categories. We detected a landscape effect connected with the intensity and length of forest management history, with the highest numbers of all and red-listed species per site in eastern Finland. Volume and diversity of dead wood were the most important variables explaining variation in species richness. Presently, key habitats in managed forests appear to contribute only little to the conservation of red-listed polypore species.     

Succession in myxomycete communities on dead <Emphasis Type="Italic">Pinus densiflora</Emphasis> wood in a secondary forest in southwestern Japan

Changes in myxomycete communities and species were investigated over an 8-year period in relation to the decay state of dead Pinus densiflora Siebold & Zucc. wood on which myxomycete fruiting bodies occurred. The study was carried out during three different seasons in a pine forest in southwestern Japan. A total of 44 species and seven varieties of myxomycetes were recorded. The species richness and diversity of the annual myxomycete communities did not clearly change in relation to the series of years, but the percent similarity of the myxomycete community from the beginning of the survey through the following years tended to decrease every season. The ordination of the annual communities, analyzed using non-metric multidimensional scaling (NMDS), indicated that seasonal factors on the first axis and the decay state of the wood on the second axis were significantly related. Species colonization patterns were arranged using succession indices and the distribution of certain species at particular times of the year: Arcyria ferruginea, A. obvelata, Lamproderma arcyrionema, and Physarum viride early in the year and Stemonitopsis hyperopta, Cribraria intricata, Lindbladia cribrarioides, Lamproderma columbinum, Tubifera ferruginosa, and Trichia verrucosa later on. Changes in the relative abundance of colony sizes of several species showed annual trends. Species using slightly decayed wood at the beginning were replaced by those using more brittle wood as the years progressed. Myxomycete succession on dead wood changed through time as the wood decayed, based on species preferences for particular decay stages.     

Cryptogam communities on decaying deciduous wood – does tree species diversity matter?

Bryophyte and fungal communities were investigated on fallen trees representing seven deciduous tree species in a mixed near natural nemoral forest. Bryophytes were represented by 41 taxa, including several very frequent species. Of the 296 fungal species, most were recorded with very low frequency and the share of high frequent species was much lower than among the bryophytes. Species turnover was bigger in the fungal communities, compared to the bryophyte communities, and related to a higher extent to measured differences in environmental conditions. Tree species diversity was found to be an important factor for fungal species composition, while only small differences in bryophyte species composition were found between the different tree species. On the other hand bryophyte species richness showed distinct relations to tree species and microclimatic variables, a tendency which was not evident for fungal diversity. It is concluded that the two organism groups to some extent differ in their conservation demands. Thus, conservation of wood-inhabiting bryophytes requires prioritising of large, coherent forest stands in which a stable humid microclimate and a reasonable supply of dead wood is secured. Successful conservation of fungi requires that substantial amounts of dead wood are left for natural decay in a variety of natural forest environments representing different tree species, so that heterogeneity in dead wood types is secured.     

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.     

Forest microsite effects on community composition of ectomycorrhizal fungi on seedlings of Picea abies and Betula pendula

Niche differentiation in soil horizons, host species and natural nutrient gradients contribute to the high diversity of ectomycorrhizal fungi in boreal forests. This study aims at documenting the diversity and community composition of ectomycorrhizal fungi of Norway spruce ( Picea abies ) and silver birch ( Betula pendula ) seedlings in five most abundant microsites in three Estonian old-growth forests. Undisturbed forest floor, windthrow mounds and pits harboured more species than brown- and white-rotted wood. Several species of ectomycorrhizal fungi were differentially represented on either hosts, microsites and sites. Generally, the most frequent species in dead wood were also common in forest floor soil. Ordination analyses suggested that decay type determined the composition of EcM fungal community in dead wood. Root connections with in-growing mature tree roots from below affected the occurrence of certain fungal species on seedling roots systems in dead wood. This study demonstrates that ectomycorrhizal fungi differentially establish in certain forest microsites that is attributable to their dispersal and competitive abilities. Elevated microsites, especially decayed wood, act as seed beds for both ectomycorrhizal forest trees and fungi, thus affecting the succession of boreal forest ecosystems.     

Positive edge effects on forest-interior cryptogams in clear-cuts

Biological edge effects are often assessed in high quality focal habitats that are negatively influenced by human-modified low quality matrix habitats. A deeper understanding of the possibilities for positive edge effects in matrix habitats bordering focal habitats (e.g. spillover effects) is, however, essential for enhancing landscape-level resilience to human alterations. We surveyed epixylic (dead wood inhabiting) forest-interior cryptogams (lichens, bryophytes, and fungi) associated with mature old-growth forests in 30 young managed Swedish boreal forest stands bordering a mature forest of high conservation value. In each young stand we registered species occurrences on coarse dead wood in transects 0-50 m from the border between stand types. We quantified the effect of distance from the mature forest on the occurrence of forest-interior species in the young stands, while accounting for local environment and propagule sources. For comparison we also surveyed epixylic open-habitat (associated with open forests) and generalist cryptogams. Species composition of epixylic cryptogams in young stands differed with distance from the mature forest: the frequency of occurrence of forest-interior species decreased with increasing distance whereas it increased for open-habitat species. Generalists were unaffected by distance. Epixylic, boreal forest-interior cryptogams do occur in matrix habitats such as clear-cuts. In addition, they are associated with the matrix edge because of a favourable microclimate closer to the mature forest on southern matrix edges. Retention and creation of dead wood in clear-cuts along the edges to focal habitats is a feasible way to enhance the long-term persistence of epixylic habitat specialists in fragmented landscapes. The proposed management measures should be performed in the whole stand as it matures, since microclimatic edge effects diminish as the matrix habitat matures. We argue that management that aims to increase habitat quality in matrix habitats bordering focal habitats should increase the probability of long-term persistence of habitat specialists.     

Functional structure of ant and termite assemblages in old growth forest,logged forest and oil palm plantation in Malaysian Borneo

Forested tropical landscapes around the world are being extensively logged and converted to agriculture, with serious consequences for biodiversity and potentially ecosystem functioning. Here we investigate associations between habitat disturbance and functional diversity of ants and termites—two numerically dominant and functionally important taxa in tropical rain forests that perform key roles in predation, decomposition, nutrient cycling and seed dispersal. We compared ant and termite occurrence and composition within standardised volumes of soil and dead wood in old growth forest, logged forest and oil palm plantation in Sabah, Malaysian Borneo. Termites occurred substantially less frequently in converted habitats than in old growth forest, whereas ant occurrences were highest in logged forest and lowest in old growth forest. All termite feeding groups had low occurrence in disturbed habitats, with soil feeders occurring even less frequently than wood feeders. Ant functional groups showed more variable associations, with some opportunist and behaviourally dominant groups being more abundant in degraded habitats. The importance of ants and termites in tropical ecosystems and such differing patterns of assemblage variation suggest that ecosystem functioning may be significantly altered in converted habitats.     

长白山针阔混交林不同演替阶段的昆虫多样性

昆虫多样性变化对生态系统健康有重要的指示作用, 为研究昆虫群落变化与生境演替之间的关系, 本研究采用网捕、灯诱和诱捕法系统调查了长白山针阔混交林不同演替阶段（次生白桦林、次生针阔混交林、原始阔叶红松林）昆虫群落的组成和多样性, 分析了昆虫在森林演替过程中的规律及与植被群落之间的关系。系统调查共采集昆虫标本8 183头, 隶属于14个目699种, 其中鳞翅目和鞘翅目是主要优势类群。次生针阔混交林昆虫的个体数量最多, 原始阔叶红松林中物种数最多。不同演替阶段昆虫群落的物种数和个体数差异不显著, 但次生针阔混交林、原始阔叶红松林的Fisher’s α指数显著高于次生白桦林。目水平上的昆虫多样性未表现出显著性差异. 昆虫多样性在森林演替过程中和草本植物多样性的变化趋势相同;由于食性和生境选择的不同, 森林演替过程中鳞翅目昆虫多样性逐渐升高, 而鞘翅目多样性逐渐降低。     

Importance of high quality early-successional habitats in managed forest landscapes to rare beetle species

Species adapted to early-successional forest habitats are in managed landscapes largely confined to clearcuts. To improve habitat quality on clearcuts, green tree and dead wood retention is widely applied in forestry; however, its effects on rare early-successional species have rarely been shown. We repeatedly surveyed two red-listed beetle species (Upis ceramboides and Platysoma minus) on clearcuts in a managed boreal forest landscape. We found that U. ceramboides decreased its occupancy over time while P. minus increased, indicating that red-listed species vary in their ability to successfully utilise managed habitats. We found no effect of connectivity on probability of occurrence, colonisation or extinction per clearcut. Trees retained alive improved habitat quality of clearcuts, since both species were more frequent in dead wood of such trees, in comparison to logging residues. We suggest that retention can be improved by protecting and creating dead wood as intact trees during harvesting. Rare specialist species require habitat of high quality, and consequently it is impossible to meet the requirements of these species on every clearcut. To preserve all early-successional species at a regional scale, we recommend focusing retention of green trees and dead wood to one or a few trees species on each clearcut and in each landscape.     

Trophic level modulates carabid beetle responses to habitat and landscape structure: a pan‐European study

1. Anthropogenic pressures have produced heterogeneous landscapes expected to influence diversity differently across trophic levels and spatial scales. 2. We tested how activity density and species richness of carabid trophic groups responded to local habitat and landscape structure (forest percentage cover and habitat richness) in 48 landscape parcels (1 km²) across eight European countries. 3. Local habitat affected activity density, but not species richness, of both trophic groups. Activity densities were greater in rotational cropping compared with other habitats; phytophage densities were also greater in grassland than forest habitats. 4. Controlling for country and habitat effects, we found general trophic group responses to landscape structure. Activity densities of phytophages were positively correlated, and zoophages uncorrelated, with increasing habitat richness. This differential functional group response to landscape structure was consistent across Europe, indicated by a lack of a country × habitat richness interaction. Species richness was unaffected by landscape structure. 5. Phytophage sensitivity to landscape structure may arise from relative dependency on seed from ruderal plants. This trophic adaptation, rare in Carabidae, leads to lower phytophage numbers, increasing vulnerability to demographic and stochastic processes that the greater abundance, species richness, and broader diet of the zoophage group may insure against.     

Re‐forestation restores native dominance in an island beetle fauna

Native re‐forestation is a widely used restoration tool, typically undertaken with the expectation that planting native trees will initiate succession processes (including the re‐establishment of native fauna) that will eventually return the ecosystem to a native‐dominated state. Invertebrate groups can be used to assess restoration progress, as their life history traits enable them to respond more rapidly to environmental change than many other organisms. In this study, we assessed beetle responses to re‐forestation. Using two trapping methods (flight intercept traps and pitfall traps), we compared beetle assemblages in exotic pasture (pre‐restoration state), <10‐year‐old planted native forest (restoration intervention) and approximately 40‐year‐old unmanaged regenerating native forest (reference state). Analysis of the flight intercept‐trapped beetles suggests that re‐forestation has initiated a transition from an exotic‐dominated pasture fauna toward a native‐dominated fauna: in planted forests, 75% of all flight‐intercept‐trapped beetles were native (compared with 22% in pasture and 87% in unmanaged forest). Flight intercept‐trapped beetles also had higher native diversity and abundance in both forest types than in pasture. Pitfall‐trapped beetle species were predominantly native in both forest types, but there were few statistically significant differences between the forests and pasture in the pit‐fall trap data set. Both trapping methods detected significant compositional differences between the beetle assemblages in planted forest and unmanaged forest. Replanting native forest has increased native beetle diversity, abundance, and dominance (compared with the pre‐restoration state), but convergence with the unmanaged reference forest has not yet been achieved.     

Patterns in woody species diversity, richness and partitioning of diversity in forest communities of tropical deciduous forest biome

The regressive succession model hypothesizes tropical savanna-woodlands to be a degraded stage of primary deciduous forests. Species diversity, richness and evenness of woody species in savanna-woodlands, secondary deciduous forests and mature deciduous forests of central India were compared to test if the regressive succession explained pattern in species richness, diversity, functional diversity and basal area. At the plot scale (0.1 ha) secondary deciduous forests and savanna-woodlands had similar species diversity, a pattern not consistent with the regressive model of deciduous forest succession, and mature deciduous forests had greater species diversity and richness (p<0.05). When examined at a larger scale or community scale by pooling all plots within a community type, the trend in diversity persisted even with greater effort allocated to sampling of secondary deciduous forests. Species richness at the community scale was greatest in secondary deciduous forest as expected from species area relationship. The communities shared 28 woody species but the species composition was significantly different between the communities. I suggest that conservation of tropical deciduous forests based on regressive succession model is problematic.     

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

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

Butterfly community recovery in degraded rainforest habitats in the Upper Guinean Forest Zone (Kakum forest, Ghana)

Successful regeneration of secondary tropical forest might be crucial in the conservation of rainforests, since large areas of primary forest have been destroyed or degraded. Animal communities might play an important role in restoration of biological diversity in these secondary habitats, since some groups have high mobility and capacity for dispersal. Fruit-feeding butterflies were trapped to measure differences between butterfly communities in primary rainforest and disturbed forest habitats of different stage of regeneration including clear-cut, abandoned farmland, newly planted forest and middle-aged secondary growth. 3465 specimens representing 114 species were identified from 56 traps operated for 36 days. Extremely high values of rarefied species richness were estimated in the clear-cut habitat, due to the high number of singletons and doubletons. This was caused by a gap-effect that allowed penetration of canopy and open area species after disturbance. The differences between butterfly communities were best demonstrated through ecological composition, richness and abundance of indicator groups and habitat similarity based on Jaccard’s similarity index. The results show a clear ability of butterfly communities in degraded forest habitats to regenerate in 50–60 years after clearance.     

The effects of stand structure on ground-floor bryophyte assemblages in temperate mixed forests

The effect of tree species composition, stand structure characteristics and substrate availability on ground-floor bryophyte assemblages was studied in mixed deciduous forests of Western Hungary. Species composition, species richness and cover of bryophytes occurring on the soil and logs were analysed as dependent variables. The whole assemblage and functional groups defined on the basis of substrate preference were investigated separately. Substrate availability (open soil, logs) was the most prominent factor in determining species composition, cover and diversity positively, while the litter of deciduous trees had a negative effect on the occurrence of forest floor bryophytes. Besides, bryophyte species richness increased with tree species and stand structural diversity, and for specialist epiphytic and epixylic species log volume was essential. Sapling density and light heterogeneity were influential on bryophyte cover, especially for the dominant terricolous species. Many variables of the forest floor bryophyte community can be estimated efficiently by examining stand structure in the studied region. Selective cutting increasing tree species diversity, stand structural heterogeneity and dead wood volume can maintain higher bryophyte diversity in this region than the shelter-wood system producing even-aged, monodominant, structurally homogenous stands.