Ecological functions and ecosystem services provided by Scarabaeinae dung beetles

Clear understanding of the links between ecological functions and biodiversity is needed to assess and predict the true environmental consequences of human activities. Several key ecosystem functions are provided by coprophagous beetles in the subfamily Scarabaeinae (Coleoptera: Scarabaeidae), which feed on animal excreta as both adults and larvae. Through manipulating feces during the feeding process, dung beetles instigate a series of ecosystem functions ranging from secondary seed dispersal to nutrient cycling and parasite suppression. Many of these ecological functions provide valuable ecosystem services such as biological pest control and soil fertilization. Here we summarize the contributions of dung beetles to nutrient cycling, bioturbation, plant growth enhancement, secondary seed dispersal and parasite control, as well as highlight their more limited role in pollination and trophic regulation. We discuss where these ecosystem functions clearly translate into ecosystem services, outline areas in critical need of additional research and describe a research agenda to fill those gaps. Due to the high sensitivity of dung beetles to habitat modification and changing dung resources, many of these ecological processes have already been disrupted or may be affected in the future. Prediction of the functional consequences of dung beetle decline demands functional studies conducted with naturally assembled beetle communities, which broaden the geographic scope of existing work, assess the spatio-temporal distribution of multiple functions, and link these ecosystem processes more clearly to ecosystem services.     

Global dung beetle response to tropical forest modification and fragmentation: A quantitative literature review and meta-analysis

Although insects are crucial for maintaining ecosystem function, our understanding of their overall response to human activity remains limited. This is no less true of dung-burying beetles (Coleoptera: Scarabaeidae: Scarabaeinae), which provide a suite of critical ecosystem functions and services, yet but face multiple conservation threats, particularly from landscape conversion. Here we use a review and meta-analysis to synthesize the current knowledge concerning response to tropical forest modification and fragmentation of dung beetles (Coleoptera: Scarabaeidae: Scarabaeinae). For every modified habitat type and individual forest fragment across 33 studies, we calculated six dung beetle community parameters, standardized relative to intact tropical forest. We organized modified habitats along an approximate disturbance gradient ranging from selectively logged, late and early secondary forest, through agroforestry, tree plantations, to annual crops, cattle pastures and clear-cuts. Secondary forests, selectively logged forest and agroforests supported rich communities with many intact forest species, while cattle pastures and clear-cuts contained fewer species overall with few forest-dwelling species. Abundance generally declined with increasing modification, but was quite variable. Communities in open habitats were often characterized by hyper-abundance of a small number of small-bodied species, leading to low evenness. Across fragmentation studies, dung beetle species richness, abundance and evenness declined in smaller forest fragments. Richness and abundance sometimes declined in more isolated fragments, although this response appeared to depend on matrix quality. Across both habitat modification and fragmentation studies, geographic location and landscape context appeared to modify dung beetle response by influencing the available pool of colonists. We discuss potential underlying mechanisms and conclude with recommendations for management and conservation and for future research.     

Abiotic and vertebrate seed dispersal in the Brazilian Atlantic forest: implications for forest regeneration

This study documents the proportion of woody plant species having different modes of seed dispersal, and classes of seed and fruit size at 16 Atlantic montane forest plots of southeastern Brazil (23-24° S). These plots represent six chronosequences (from 5-year-old to old-growth forest) of forest regeneration following small-scale shifting agriculture. Our results indicate that there is a gradual but predictable increase in the number of woody plant species relying on vertebrate-mediated seed dispersal in increasingly older plots. Moreover, the percentage of plant species bearing small seeds and fruits (< 0.6 cm in length) was more than halved from earlier to older forest plots, while the percentage of plant species producing middle-sized seeds and fruits at least doubled. Plant species belonging to the Melastomataceae, Myrsinaceae, Rubiaceae and Flacourtiaceae comprised a large proportion of the small-seeded and fruit species (33-53%), and largely occurred in early successional forest plots (5-30 years old). In contrast, species belonging to the Myrtaceae and Lauraceae represented most of the species (51-72%) bearing medium-sized seeds and fruits (0.6-1.5 cm in length), and occurred primarily in the old-growth forests. Shifts in the relative importance of dispersal strategies during the regeneration process of Atlantic montane forest, as well as in diaspore size appear to be related to the balance between early successional and shade-tolerant species associated with particular life forms and plant families. Finally, we discuss the relationships between the species richness of Myrtaceae and Lauraceae trees, seed dispersal by medium to large vertebrates, and possible regeneration scenarios for the Atlantic forest.     

Importance of hedgerows as habitat corridors for forest plants in agricultural landscapes

Hedgerows have been proposed as habitat and conservation corridor for forest plant species, but their importance for the survival of these species is still not clear. The objective of our study was to examine the frequency of occurrence of forest species and total forest species richness in different parts of the hedgerows, and to relate these patterns of occurrence to the species’ habitat requirements and life history traits. We surveyed in total 130 forest-hedgerow transects in North-western Germany, including three age classes.About 77% of all forest plant species (including some endangered taxa) occurring in the neighbouring forests were also found in the adjacent hedgerows. In all age classes, there was a negative relationship between distance from the forest-hedgerow ecotone and the number of species. Ancient hedgerows were not significantly more species-rich than more recent ones. Within 100 m distance from the forest edge, forest species richness increased with an increasing number of species in the nearby forest and with an increasing cover of the tree canopy as well as a decreasing cover of the shrub layer. Species with high frequency had comparatively high Ellenberg indicator values for light, and were mostly associated with anemochorous and epizoochorous seed dispersal.The frequency of occurrence in hedgerows of species with common attributes was partly in agreement, partly in disagreement with the results obtained in previous studies. We conclude that patterns are difficult to generalize, probably due to a strong regional variation in the pool of forest species and in the specific environments of both forests and hedgerows.     

Conservation of beetles in boreal pine forests: the effects of forest age and naturalness on species assemblages

The effort of boreal forest conservation has emphasised the preservation of old-growth forests while the role of young successional stages in maintaining biodiversity has remained largely unstudied. We compared the richness of beetle species and composition of species assemblages between managed and seminatural forests in five stages of forest succession. The sites were in boreal sub-xeric pine-dominated forests in eastern Finland. Seminatural study sites, especially the recently burned sites, were important habitats for threatened and near-threatened species. We propose that young stages of natural succession should be included in the network of protected forest areas. On the other hand, the composition of saproxylic species assemblages in seminatural forests differed from the assemblages in managed forests, indicating also the need to improve the forest management guidelines so that they better address the requirements of species protection. Regeneration methods applied should resemble or mimic the natural disturbances more closely.     

Effect of forest fragmentation on fruit and seed predation of the tropical dry forest tree Ceiba aesculifolia

Forests fragmentation reduces the density of natural plant populations forming patches of the remaining individuals. One of the biotic interactions that can be affected by forest fragmentation and is poorly studied is seed predation. We determined the effects of forest fragmentation on seed and fruit predation in Ceiba aesculifolia by comparing trees in continuous forest with trees in fragmented forest. We compared the following variables: (a) frequency of fruit predation by Collie’s squirrel (Sciurus colliaei) in each habitat; (b) frequency of the cotton-staining bug seed predator (Dysdercus, Orden Hemiptera) in each habitat; (c) the effect of seed predation on germination frequency and time; and (d) the effect of different life stages of Dysdercus on seed viability. In continuous habitat, 100% of the trees presented fruits with squirrel predation while only 34% of trees in fragmented habitats presented fruit predation. In continuous forest 27% of the trees contained fruits with the seed predator Dysdercus, while only 2% of the trees in fragmented forest presented Dysdercus. The initial weight of damaged seeds was greater than seeds that were not damaged indicating that seed predators select heavier seeds to feed upon. Frequency of seed germination was affected by different life stages; pre-adults decreased germination significantly more than nymphs and adults. Seed predation significantly increased the time it took for germination to occur. Our study shows that forest fragmentation significantly affects predation patterns of squirrels and cotton-staining bugs. Reduction of natural seed predators in forest fragments may have long-term consequences on forest structure and diversity.     

Changes in food resources and conservation of scarab beetles: from sheep to dog dung in a green urban area of Rome (Coleoptera, Scarabaeoidea)

The aim of the research was to show how a change in land use influences the structure of a dung beetle assemblage and affect its conservation. In the Pineto Urban Regional Park (Rome), dog dung is the sole food resource currently available for scarab dung beetles, after the recent removal of wild and domestic herbivores. A one-year sampling was conducted to study the scarab assemblage in dog scats (1999) and to compare it with the previous assemblage associated with sheep droppings (1986). Richness, evenness and similarity parameters were compared between the two allochronic assemblages. From sheep to dog dung, an impoverishment of the total richness was observed (from 19 to 9 species) together with an increase of individuals (by 7 times). Dog dung harboured 20% of the current scarab dung beetle fauna of Rome, probably as a consequence of the dog mixed diet, rich in cellulose. Both the communities showed a high percentage of tunnellers, probably because of the food shortage and, for dog scats, of the high dehydration rate. A comparison with other Roman scarab communities enhanced that: (1) the change in food resource determined a higher difference in species composition respect to other parameters (size and habitat diversity); (2) dog dung provided a temporary refuge for species that otherwise may encounter local extinction in urban environments.     

Parasitoids (Hymenoptera, Ichneumonoidea) of Saproxylic beetles are affected by forest successional stage and dead wood characteristics in boreal spruce forest

The habitat requirements and effects of forest management on insects belonging to higher trophic levels are relatively unknown in forest ecosystems. We tested the effect of forest successional stage and dead wood characteristics on the saproxylic parasitoid (Hymenoptera, Ichneumonoidea) assemblage in boreal spruce-dominated forests in northern Sweden. Within each of nine areas, we selected three sites with different management histories: (1) a clear-cut (2) a mature managed forest and (3) an old-growth forest. Parasitoids were collected in 2003 using eclector traps mounted on fresh logs, which were either untreated (control), burned, inoculated with fungi, or naturally shaded, and on artificially-created snags.Both forest type and dead wood characteristics had a significant effect on parasitoid assemblages. Grouped idiobionts and some species, such as Bracon obscurator and Ontsira antica, preferred clear-cuts, while others, such as Cosmophorus regius (Hym., Braconidae) and other koinobionts, were associated with older successional forest stages. No single dead wood substrate was sufficient to support the entire community of parasitoids in any forest type, even when the regular host was present. In particular, snags hosted a different assemblage of species from other types of dead wood, with parasitoids of Tetropium spp. such as Rhimphoctona spp. (Hym., Ichneumonidae) and Helconidea dentator (Hym., Braconidae) being abundant. These results indicate that a diversity of dead wood habitats is necessary to support complete assemblages of beetle-associated parasitoids from early successional stages of dead wood and that parasitoids may be more sensitive to habitat change than their hosts.     

Rove beetles and ground beetles (Coleoptera: Staphylinidae, Carabidae) as indicators of harvest and regeneration practices in western Canadian foothills forests

Staphylinid beetle assemblages from coniferous foothills forest in west-central Alberta, Canada were studied via pitfall trapping to examine the effects of stand age and possible edge effects. Sites included a chronosequence of stands from 1 to 27 years post-harvest, and four types of mature forest that had not been disturbed by fire for at least 80 years. In all, 19 sites were sampled between 1989 and 1991. A total of 98 species were identified, nine of which are reported for the first time in Alberta. Staphylinids were more abundant in mature forest stands but assemblages were more diverse in regenerating stands. Thirty-four rove beetle species showed significant indicator value for particular stands or groups of stands, including mature forest, young forest, and open ground specialists. After harvesting, the catch rate of many forest species decreased dramatically, and open ground species were more commonly collected. Populations of some forest species remained active on logged sites for one or 2 years before disappearing. As stands regenerated, they were colonized by species characteristic of young stands, but true forest species were found only in older unharvested stands. The beetle assemblages from regenerating stands became more similar to those from mature stands as they aged, but still differed considerably from them 27 years after harvesting. Transects across forest-clearcut edges revealed a significant beetle response to habitat edges. Staphylinids assemblages were compared to the ground beetle (Carabidae) assemblage sampled via the same pitfall trapping regime. Mature forest specialists are threatened by fragmentation and loss of habitat. In order to conserve these beetle assemblages, forest managers should retain adequate patches of older successional stages on working landscapes.     

Impact of selective logging on genetic composition and demographic structure of four tropical tree species

Over-exploitation and fragmentation are serious problems for tropical forests. Most sustainable forest management practices avoid clear-cuts and apply selective logging systems focused on a few commercial species. We applied a simulation model to estimate the impact of such selective logging scenarios on the genetic diversity and demography of four tropical tree species from French Guiana. The simulations used data on genetic and demographic composition, growth, phenology and pollen and seed dispersal obtained for Dicorynia guianensis, Sextonia rubra, Symphonia globulifera and Vouacapoua americana at the experimental site in Paracou. Whereas Symphonia globulifera serves as a model for a species with low logging pressure, the other three species represent the most exploited tree species in French Guiana. In simulations with moderate logging, typical for French Guiana, with large cutting diameter (>60 cm diameter) and long cutting cycles (65 years), the two species V. americana and Sextonia rubra were not able to recover their initial stock at the end of the rotation period, with a large decrease in the number of individuals and in basal area. Under a more intensive logging system (cutting diameter >45 cm diameter, cutting cycles of 30 years) that is common practice in the Brazilian Amazon, only Symphonia globulifera showed no negative impact. Generally, the differences between the genetic parameters in the control scenarios without logging and the logging scenarios were surprisingly small. The main reasons for this were the overlapping of generations and the effective dispersal ability of gene vectors in all species, which guarantee relative homogeneity of the genetic structure in different age classes. Nevertheless, decreasing the population size by logging reduced the number of genotypes and caused higher genetic distances between the original population and the population at the end of the logging cycles. Sensitivity analysis showed that genetic changes in the logging scenarios were principally determined by the growth, densities and cutting diameter of each species, and only to a very small extent by the reproductive system including factors such as pollen and seed dispersal and flowering phenology.     

Three decades of deforestation in southwest Sumatra: Have protected areas halted forest loss and logging, and promoted re-growth?

Much of the forest cover in southern Sumatra, Indonesia has been cleared since the early 1970s, but accurate estimates of the scales and rates of loss are lacking. This study combined high-quality remote sensing applications and extensive field surveys, both to provide an accurate picture of deforestation patterns across an area of 1.17 million ha in southwest Sumatra and to assess whether southwest Sumatra’s Bukit Barisan Selatan National Park (BBSNP) has halted forest loss and logging, and promoted re-growth, since its creation in 1984. Of the single large (692,850 ha) contiguous area of forest standing across our study area in 1972, nearly half (344,409 ha) has been cleared from 1972 to 2002, at an average rate per original forest cover of 1.69% y⁻¹. In Gunung Raya Wildlife Sanctuary (GRWS) and Hydrological Reserves (HR), forests have shrunk by 28,696 ha and 113,105 ha, at an average rate of 2.74% y⁻¹ and 2.13% y⁻¹, respectively. In contrast, forests in BBSNP have reduced four times more slowly than those in GRWS and HR, and have shrunk by 57,344 ha, at an average rate of 0.64% y⁻¹. Nevertheless, the forests within BBSNP were cleared almost as rapidly during the post-establishment, as during the pre-establishment, period (0.65% y⁻¹ and 0.63% y⁻¹, respectively) despite the introduction of protection measures during the post-establishment period, following the government’s pledge to expand and protect Indonesia’s network of Protected Areas (PAs) at the 1982 Bali World Parks Congress. While these protection measures failed to slow down rates of forest loss caused by agricultural encroachments they reduced large-scale mechanised logging by a factor of 4.2 and stabilized some 8610 ha of agricultural encroachments, enabling forest re-growth.     

Measurement and prediction of nitrogen loss by simulated erosion events on cultivated forest soils of contrasting structure

The objectives of this study were to determine nitrogen (N) loss associated with erosion of forest soils and to explore the role of soil structure and other factors governing N enrichment of sediment to aid prediction of N loss. We measured erosion, size distribution of aggregates in the sediment and N distribution in various aggregate fractions using simulated rainfall on samples of three cultivated forest soils of contrasting structure (repacked in trays) exposed to four erosion conditions. Both sediment loss and N loss increased with slope and kinetic energy of rainfall suggesting greater dependency of N loss on sediment loss than on N concentration in the sediment. Irrespective of erosion treatments and soil type, the bulk sediment and its size fractions were mostly richer in N than those of the uneroded soils. The enrichment ratio (E_R) and concentration ratio (C_R) of N for sediment cast some doubt on the application of raindrop stripping as a mechanism of N enrichment of sediment for soils of widely differing characteristics. Previously published models of N enrichment of sediment did not predict E_R for our soils satisfactorily. However, an empirical model using published data on erosion and N loss agreed well with the results. This predictive method only requires information on sediment loss that can be easily obtained from an erosion model.     

Influence of tree characteristics and forest management on tree microhabitats

Higher densities of tree microhabitats in unmanaged forests may explain biodiversity differences with managed forests. To better understand the determinants of this potential biodiversity indicator, we studied the influence of tree characteristics on a set of tree microhabitats (e.g. cavities, cracks, bark features) on 75 plots in managed and unmanaged French forests. We hypothesized that the number of different microhabitat types per tree and the occurrence of a given microhabitat type on a tree would be higher in unmanaged than in managed forests, and that this difference could be linked to individual tree characteristics: diameter, vitality and species. We show that unmanaged forests contained more trees likely to host microhabitats (i.e. large trees, snags) at the stand level. However, at the tree level, forest management did not influence microhabitats; only tree characteristics did: large trees and snags contained more microhabitats. The number and occurrence of microhabitats also varied with tree species: oaks and beech generally hosted more microhabitats, but occurrence of certain types of microhabitats was higher on fir and spruce. We conclude that, even though microhabitats are not equally distributed between managed and unmanaged forests, two trees with similar characteristics in similar site conditions have the same number and probability of occurrence of microhabitats, whatever the management type. In order to preserve biodiversity, foresters could reproduce unmanaged forest features in managed forests through the conservation of specific tree types (e.g. veteran trees, snags). Tree microhabitats could also be more often targeted in sustainable forest management monitoring.     

Impacts of selective logging on canopy and ground assemblages of tropical forest butterflies: Implications for sampling

Commercial selective logging is a major cause of habitat disturbance in Southeast Asian rainforests, yet despite much research there is little consensus on impacts of disturbance on biodiversity. There is also little consensus on the most appropriate methods for sampling tropical species, making it difficult to draw general conclusions from published studies. For example, many studies have used butterflies to assess impacts of selective logging but sampling has usually been conducted at ground level and the canopy fauna has often been ignored. In this study, we investigate the importance of sampling in the canopy by using fruit-baited traps to investigate impacts of selective logging on Nymphalid butterflies in primary forest and forest selectively logged 15 years previously in Sabah (Malaysian Borneo). Analysing combined data from canopy (≈25 m above ground level) and ground-level traps showed significantly lower diversity in selectively logged forest. However, this difference was not observed when data from only ground traps were analysed. Primary forest supported a butterfly assemblage comprising species with more restricted geographical ranges, and thus higher conservation value, compared with selectively logged forest. This result was observed regardless of whether or not canopy data were included in this analysis. We conclude that sampling in the canopy is critical when producing species inventories, but of little importance when determining the impacts of selective logging on restricted-range species.     

The responses of understorey birds to forest fragmentation, logging and wildfires: An Amazonian synthesis

We combine mist-net data from 24 disturbance treatments taken from seven studies on the responses of understorey Amazonian birds to selective logging, single and recurrent wildfires, and habitat fragmentation. The different disturbance treatments had distinct effects on avian guild structure, and fire disturbance and the isolation of forest patches resulted in bird communities that were most divergent from those in continuous, undisturbed forest in terms of their species composition. Although low-intensity logging treatments had the least noticeable effects, the composition of understorey birds was still markedly different from the composition in undisturbed forest. This analysis demonstrates the importance of preventing habitat fragmentation and the spread of fires in humid tropical forests, and highlights the need for more research to determine the long-term suitability of large areas of degraded forest for forest birds.     

Nitrogen mineralization in relation to site history and soil properties for a range of Australian forest soils

Rates of N mineralization were measured in 27 forest soils encompassing a wide range of forest types and management treatments in south-east Australia. Undisturbed soil columns were incubated at 20°C for 68 days at near field-capacity water content, and N mineralization was measured in 5-cm depth increments to 30 cm. The soils represented three primary profile forms: gradational, uniform and duplex. They were sampled beneath mature native Eucalyptus sp. forest and from plantations of Pinus radiata of varying age (<1 to 37 years). Several sites had been fertilized, irrigated, or intercropped with lupins. The soils ranged greatly in total soil N concentrations, C:N ratios, total P, and sand, silt, and clay contents. Net N mineralization for individual soil profiles (0–30 cm depth) varied from 2.0 to 66.6 kg ha^-1 over 68 days, with soils from individual depths mineralizing from <0 (immobilization) to 19.3 kg ha^-1 per 5 cm soil depth. Only 0.1–3.1% of the total N present at 0–30 cm in depth was mineralized during the incubation, and both the amount and the percentage of total N mineralized decreased with increasing soil depth. N fertilization, addition of slash residues, or intercropping with lupins in the years prior to sampling increased N mineralization. Several years of irrigation of a sandy soil reduced levels of total N and C, and lowered rates of N mineralization. Considuring all soil depths, the simple linear correlations between soil parameters (C, N, P, C:N, C:P, N:P, coarse sand, fine sand, silt, clay) and N mineralization rates were generally low (r<0.53), but these improved for total N (r=0.82) and organic C (r=0.79) when the soils were grouped into primary profile forms. Prediction of field N-mineralization rates was complicated by the poor correlations between soil properties and N mineralization, and temporal changes in the pools of labile organic-N substrates in the field.     

Changes of lignin phenols and neutral sugars in different soil types of a high-elevation forest ecosystem 25 years after forest dieback

Long-term effects of forest disturbance 25 yr ago on lignin and non-cellulosic polysaccharide pools in an unmanaged high-elevation Norway spruce (Picea abies L. [Karst.]) forest were investigated by comparing three dieback sites with three adjacent control sites with non-infested spruce on identical soils. Samples were taken from the forest floor and the mineral soil; one Ah horizon sample per site was physically fractionated into density and particle size fractions. Additionally, changes in the above- and belowground input of lignin and non-cellulosic polysaccharides after forest dieback were quantified. Lignin and its degree of structural alteration in plant and soil samples were assessed by CuO oxidation and subsequent analysis of the lignin phenols. Non-cellulosic polysaccharides were determined after hydrolysis with trifluoroacetic acid (TFA), derivatisation of their neutral sugar monomers by reduction to alditols, and subsequent acetylation. The total plant-derived input of lignin and non-cellulosic polysaccharides to the soil was similar for the dieback and the control sites. The chemical composition of the input has changed considerably after forest dieback, as shown by significantly higher syringyl/vanillyl (S/V) ratios and significantly lower (galactose+mannose)/(arabinose+xylose) (GM/AX) ratios. This indicates a changed plant input and a higher contribution of microbial sugars. Contents of lignin phenols in the forest floor and coarse particle size fractions of the A horizons were significantly smaller at the dieback sites (p<0.01). Moreover, larger acid-to-aldehyde ratios of vanillyl units (Ac/Al)_v indicated an increased degree of lignin phenol alteration. Also contents of neutral sugars were significantly (p<0.01) smaller in the forest floor, but not in the A horizons of the dieback sites. The GM/AX mass ratios as well as the (rhamnose+fucose)/(arabinose+xylose) (RF/AX) ratios in the forest floor and coarse particle size fractions of the mineral topsoil were significantly (p<0.01) larger after forest dieback, indicating a larger relative contribution of microbial sugars. In general, the lignin phenol and neutral sugar pools of all three soil types exhibited similar response patterns to the changed site conditions. Our results demonstrate that the lignin and neutral sugar pools of humic topsoil horizons are highly sensitive to forest disturbances. However, the two compounds show different patterns in the mineral soil, with the major neutral sugar pool being stabilized against changes whereas the lignin phenol pool decreases significantly.     

The role of ectomycorrhizal communities in forest ecosystem processes: New perspectives and emerging concepts

The fungal symbionts forming ectomycorrhizas, as well as their associated bacteria, benefit forest trees in a number of ways although the most important is enhancing soil nutrient mobilization and uptake. This is reciprocated by the allocation of carbohydrates by the tree to the fungus through the root interface, making the relationship a mutualistic association. Many field observations suggest that ectomycorrhizal fungi contribute to a number of key ecosystem functions such as carbon cycling, nutrient mobilization from soil organic matter, nutrient mobilization from soil minerals, and linking trees through common mycorrhizal networks. Until now, it has been very difficult to study trees and their fungal associates in forest ecosystems and most of the work on ECM functioning has been done in laboratory or nursery conditions. In this review with discuss the possibility of working at another scale, in forest settings. Numerous new techniques are emerging that makes possible the in situ study of the functional diversity of ectomycorrhizal communities. This approach should help to integrate developing research on the functional ecology of ectomycorrhizas and their associated bacteria with the potential implications of such research for managing the effects of climate change on forests.     

中亚热带四种森林土壤团聚体及其有机碳分布特征

选择中亚热带地区4种典型森林类型:杉木林、湿地松林、毛竹林和次生林4种森林土壤为研究对象,研究了森林类型对土壤不同发生层水稳性团聚体及其有机碳分布特征的影响。结果表明:不同森林类型对>5 mm和2~5 mm土壤团聚体含量影响显著(p<0.05),表现为次生林>杉木林>毛竹林>湿地松林,而在其他粒径无显著差异。0~30 cm土层内团聚体R0.25和MWD次生林显著高于其他人工林,杉木林次之,湿地松林和毛竹林最低,其他土层无显著差异。各森林类型同土层不同粒径团聚体中有机碳含量随粒径大小变化,团聚体粒径越小,有机碳含量越高。0~10 cm土层同粒径土壤团聚体有机碳含量从大到小依次是次生林、杉木林、湿地松林和毛竹林,而在其他土层各森林类型之间差异不显著。     

History and forest biodiversity of woodland key habitats in south boreal Sweden

Forest exploitation has destructed forest habitats in Fennoscandia during last centuries. Vital components of existing forest biodiversity have been identified as small (a few ha) woodland key habitats (WKHs). Many WKHs in boreal forest landscapes are assumed to represent remnant near-natural forests from before the phase of industrial forest exploitation. Thus, WKHs are supposed to (i) exhibit a low degree of exploitation, (ii) have old-growth characteristics and (iii) host Red-list species. Yet, WKHs’ history and biodiversity have not been systematically investigated. Thus, their conservation values remain unclear. This study investigates history and biodiversity of 15 WKHs in central Sweden with retrospective methods and field studies. We analyze (1) forest structure before, during and after the forest exploitation period, (2) existing biodiversity, i.e. species and structural elements, and (3) to what extent human activities (forestry) have influenced current biodiversity. Our results indicate that forest structure within the areas that currently are classified as WKHs has changed dramatically since the mid-19th century, when forests were unaffected by logging. The results suggest that fire-suppression during last century, gradually increased logging until mid-20th century and abandonment by modern forestry during the late 20th century, are main causes behind this transformation. Less than 10% of studied WKHs had biological features that resembled those of pristine boreal forests. Thus, most studied WKHs cannot be classified as near-natural or remnants of pristine forest. We suggest that the program for WKHs give more focus to pre-industrial forest structure, local historical aspects being seriously regarded and ecological restoration efforts being introduced.