The impact of beech thickets on biodiversity

Beech bark disease has dramatically altered hardwood forest structure and composition across northeastern North America. Extensive overstory mortality has resulted in prolific root-sprouting in some stands leading to the development of understory thickets of clonal small-stemmed beech. Beech thickets may impact local forest biodiversity, but this has not been adequately evaluated. We hypothesized significant differences in diversity of groundcover flora, craneflies, amphibians, and small mammals between plots with and without beech thickets. Paired plots were established in uneven-aged northern hardwood forest stands with no recent management history at two sites in the Adirondack Mountains of New York State. Groundcover plants, terrestrial craneflies, amphibians and small mammals were sampled on twenty paired plots. Discriminant analysis showed a significant difference between thicket and non-thicket (control) areas; significant variables in plot type separation were beech sapling abundance, leaf litter depth, and coarse woody debris volume. Groundcover plant cover, richness, and diversity were significantly lower in thicket compared to non-thicket plots, while beech sapling density explained 17–38 % in groundcover plant species diversity. There were no significant differences between the diversity of cranefly, amphibian and small mammal communities of each plot type. Beech thickets are important determinants of local biodiversity.     

Conservation of Heathland Ground Beetles (Coleoptera,Carabidae): The Value of Lowland Coniferous Plantations

The value of conifer-dominated plantation forestry for heathland carabids is examined, in Breckland, eastern England. Historically dominated by heathland habitats supporting scarce species of restricted distribution within the UK, approximately 47% of Breckland was afforested in the early 20th century. The carabid fauna of this forest, previously little known, was sampled by intensive pitfall trapping, and results compared to samples from heathland and arable. The results affirm that Breckland supports many range-restricted carabid beetles, including scarce species restricted to lowland heathland or sandy habitats. Approximately half of the heathland-associated species, and also half of the Nationally Scarce carabid species recorded in Breckland, were found within the forest landscape. Carabid species composition differed between closed-canopy forest, heathland and arable, while open forest habitats (clear-felled stands, young restocked stands, and track margins) contained a diverse fauna with elements from all habitats. The proportion of heathland-associated species was significantly greater in open forest habitats than in closed-canopy forest and was similar to that of heathland assemblages. Wing morphology did not differ between heathland species solely recorded from heathland and those also recorded in the forest. Nationally Scarce species from the forest had a wider UK distribution than those recorded only from heathland. Our study shows that, within conifer forest planted in lowland heathland areas, open habitats provided by trackway networks and clear-felling management can have significant value for the conservation of open-ground carabids, including species of conservation importance.     

Coastal heathland succession influences butterfly community composition and threatens endangered butterfly species

Succession has a strong influence on species diversity and composition of semi-natural open terrestrial ecosystems. While several studies examined the effects of succession on butterflies in grassland and forest ecosystems, the response of heathland butterflies to succession had not been investigated so far. To address this issue we sampled butterfly abundance and environmental parameters on the Baltic island of Hiddensee (NE Germany) along a gradient of coastal heathland succession from grey dunes to birch forest. Our results provide evidence that succession of coastal heathland has a strong influence on butterfly diversity, abundance, and species composition. Thereby grass and tree encroachment present the main threats for heathland butterflies. Diversity and abundance of butterflies were highest in shrub-encroached heath directly followed by early stages of coastal heathland succession (dwarf-shrub heath, grey dune). Both observed threatened species (Hipparchia semele, Plebeius argus) were negatively affected by succession: abundance decreased with increasing vegetation density (both species) and grass cover (P. argus); consequently, the two later successional stages (shrub, birch forest) were not occupied. Our findings highlight the importance of the preservation of early stages of coastal heathland succession for endangered butterfly species. For coastal heathland management we therefore suggest to maintain early successional stages by sheep grazing, mowing or, in case of high nutrient contents, intensive techniques such as sod-cutting or choppering. To a lower extend shrub-encroached sites should also be present, which might be beneficial for overall species richness.     

The matrix affects trackway corridor suitability for an arenicolous specialist beetle

In conserving regional insect diversity dispersal corridors are advocated to counteract fragmentation and for resilience to climate change. However, influences of corridor design and management on their function are poorly understood. Effects of contrasting matrix structure on the suitability and function of trackways as corridors for dispersal of an arenicolous carabid beetle, Harpalus rufipalpis (Sturm), was studied within a plantation landscape using mark-release-recapture. A total of 1,120 marked H. rufipalpis were released into four trackways: two “open” trackways surrounded by pine plantations aged 13–16 years and two “shaded” trackways surrounded by plantations aged 26–37 years. Dispersal was monitored by a grid of pitfall transects placed across trackways at intervals of four meters, extending 44 m north and south of the release point. Numbers of resident and marked recaptured H. rufipalpis, their average daily movement rates and numbers of recaptures in the north and south direction were compared between open and shaded trackways using 238 recaptures. The surrounding matrix affected trackway suitability with greater abundance of resident beetles found in open trackways; however H. rufipalpis was also naturally present in shaded trackways. H. rufipalpis were more active in low quality shaded corridors as inferred from the greater number of recaptures and from greater daily movement rates. Corridor edge permeability differed between trackway types, with more individuals leaving the corridor to enter the matrix in the less suitable shaded trackways. Thus matrix type affected the potential habitat suitability and conduit function of trackway corridors.     

Spontaneous Return of Biodiversity in Restored Subtropical Thicket: Portulacaria afra as an Ecosystem Engineer

An accepted criterion for measuring the success of ecosystem restoration is the return of biodiversity relative to intact reference ecosystems. The emerging global carbon economy has made landscape‐scale restoration of severely degraded Portulacaria afra (spekboom)‐dominated subtropical thicket, by planting multiple rows of spekboom truncheons, a viable land‐use option. Although large amounts of carbon are sequestered when planting a monoculture of spekboom, it is unknown whether this is associated with the return of other thicket biodiversity components. We used available carbon stock data from degraded, restored, and intact stands at one site, and sampled carbon stocks at restored stands at another site in the same plant community. We also sampled plant community composition at both sites. The total carbon stock of the oldest (50 years) post‐restoration stand (250.8 ± 14 t C ha^?1) approximated that of intact stands (245 t C ha^?1) and we observed a general increase in carbon content with restoration age (71.4 ± 24 t C ha^?1 after 35 and 167.9 ± 20 t C ha^?1 after 50 years). A multiple correspondence analysis separated degraded stands from stands under restoration based on ground cover, floristic composition, and total carbon stock. Older post‐restoration and intact stands were clustered according to woody canopy recruit abundance. Our results suggest that spekboom is an ecosystem engineer that promotes spontaneous return of canopy species and other components of thicket biodiversity. The spekboom canopy creates a cooler micro‐climate and a dense litter layer, both likely to favor the recruitment of other canopy species.     

Impacts of succession and grass encroachment on heathland Orthoptera

For the conservation of biodiversity, heathlands present important ecosystems throughout Europe. The formerly widespread habitats are nowadays restricted to small and isolated remnants. Without land use heathland vegetation undergoes succession and, in addition, the increasing amount of atmospheric nitrogen deposition has resulted in an encroachment of grasses. In the present study we analysed the effects of succession and grass encroachment on Orthoptera in a coastal heathland on the Baltic island of Hiddensee, Germany. Vegetation, microclimate, soil humidity and Orthoptera were sampled in the five main stages of heathland succession, namely grey dunes, dwarf-shrub heath, grassy heath, heath with shrubs, and birch forest. Vegetation and environmental parameters showed strong differences among the successional stages. Orthoptera species richness was highest in transitional stages. The high proportion of grasses offer favourable habitat conditions for graminivorous, chorto- and thamnobiont species. Orthoptera density was highest in grey dunes. Threatened and specialised species were restricted to the young stages grey dunes and dwarf-shrub heath. Hence, in order to maintain a high diversity of Orthoptera in heathlands, maintaining different successional stages is of critical importance and this should be integrated into heathland management practices.     

Invasion and persistence of bird-dispersed,subtropical thicket and forest species in fire-prone coastal fynbos

Abstract. Seedling abundance at four microsites (open fynbos, beneath emergent fynbos shrubs, beneath thicket, and beneath forest) was determined at three coastal dune landscapes, located along a gradient of increasing summer rainfall and where fire-dependent fynbos was the predominant vegetation. At all sites thicket seedlings were most common beneath emergent fynbos shrubs and under thicket clumps; seedlings of forest species were most abundant at forest microsites although some individuals were recorded beneath thicket. Very few thicket seedlings were observed in open fynbos. Birds play a keystone role in facilitating establishment of the fleshy fruit-bearing thicket flora. Seedling abundance at microsites of different thicket and forest species was generally unrelated to fruit abundance. Germination success of most species was highest under shaded conditions; soil organic content had no effect on germination. Removal of pulp and birdingestion enhanced the germination, relative to untreated controls, of two out of three species tested. A simple Markov model predicted a gradual increase in cover of the thicket and forest component and a gradual decline in fynbos under a ‘normal’ (20-yr interval) fire regime simulated over 10 cycles. Although inter-fire seedling establishment under emergent fynbos shrubs is important in the initial colonisation of fynbos by obligate resprouting thicket shrubs, these species persist and expand by vegetative recruitment after and between fires, respectively. In the prolonged absence of fire, the endemic-rich and fire-dependent fynbos flora would be replaced by species-poor forest and thicket.     

Do riparian forest strips in modified forest landscapes aid in conserving bat diversity?

Agricultural practices lead to losses of natural resources and biodiversity. Maintaining forests alongside streams (riparian forest strips) has been used as a mechanism to minimize the impact of clearing for agriculture on biodiversity. To test the contribution of riparian forest strips to conserve biodiversity in production landscapes, we selected bats as a biodiversity model system and examined two dimensions of diversity: taxonomic and functional. We compared bat diversity and composition in forest, with and without stream habitat, and in narrow forest riparian strips surrounded by areas cleared for agriculture. We tested the hypothesis that riparian forest strips provide potential conservation value by providing habitat and serving as movement corridors for forest bat species. Riparian forest strips maintained 75% of the bat species registered in forested habitats. We found assemblage in sites with riparian forest strips were dominated by a few species with high abundance and included several species with low abundance. Bat species assemblage was more similar between sites with streams than between those sites to forests without stream habitat. These results highlight the importance of stream habitat in predicting presence of bat species. We registered similar number of guilds between forest sites and riparian forest strips sites. Relative to matrix habitats, stream and edge habitats in riparian forest strips sites were functionally more diverse, supporting our hypothesis about the potential conservation value of riparian forest strips. Results from this study suggest that maintaining riparian forest strips within cleared areas for agricultural areas helps conserve the taxonomic and functional diversity of bats. Also, it provides basic data to evaluate the efficacy of maintaining these landscape features for mitigating impacts of agricultural development on biodiversity. However, we caution that riparian forest strips alone are not sufficient for biodiversity maintenance; their value depends on maintenance of larger forest areas in their vicinity.     

Response of ground-dwelling beetle (Coleoptera) assemblages to giant knotweed (<Emphasis Type="Italic">Reynoutria</Emphasis> spp.) invasion

Giant knotweeds (Reynoutria spp.) are highly productive and aggressive invaders in riparian wetlands of Europe and North America. We sampled ground-dwelling beetles by pitfall traps from six sites comparing monotypic Reynoutria stands with the invaded native Urtica-dominated stands. Three sites were located in a semi-natural softwood forest and three sites were on a ruderal embankment. The analyses are based on a total of 13,244 individuals from 218 species. Location and site significantly influenced beetle assemblages. Moreover, there were pronounced differences between vegetation stands. The monotypic Reynoutria stands exhibited lower beetle abundance, species richness and rarefaction diversity irrespective of location. However, the negative effect on species richness, abundance and assemblage similarities were stronger on the transformed ruderal embankment than in the semi-natural softwood forest. Reynoutria invasion seems to influence microclimatic conditions. We found a higher abundance of silvicolous and a lower abundance of xerophilous ground beetles in the Reyountria stands than in the Urtica-dominated stands. Feeding guilds reacted differently to Reynoutria invasion that reduced the abundance of predators and herbivores but enhanced that of detritivores. Detritivores assumingly profit from the perennial presence of the large quantities of Reynoutria litter. We conclude that highly productive invaders pauperise the arthropod fauna and alter link strengths in trophic cascades shifting primary producer-based food webs to detritus-based food webs.     

Response of carabid beetles (Coleoptera: Carabidae) and spiders (Araneae) to coastal heathland succession

We analyzed the impacts of succession and grass encroachment on carabid beetle and spider assemblages in a coastal heathland. Further, indicator species for different successional stages (grey dunes, dwarf-shrub heath, grassy heath, heath with shrubs, birch forest) were identified, and their relations to habitat parameters were analyzed. The study was conducted on the Baltic island of Hiddensee, Germany. Ground-dwelling arthropods were sampled using pitfall traps along a successional gradient containing five stages. Ordination by nonmetric multidimensional scaling (NMDS) and analysis of similarities (ANOSIM) revealed a clear separation of species composition among the successional stages. Both in carabid beetles and spiders, most indicator species were obtained in the youngest stage (grey dunes) and fewest in the intermediate stages (grassy heath, heath with shrubs). Also the proportion of endangered species was highest in grey dunes. Based on our results, conservation management of coastal heathlands should preserve a mosaic of different successional stages with a clear preference on younger stages (grey dunes and dwarf-shrub heath).     

The age of Calluna stands moderates post‐fire regeneration rate and trends in northern Calluna heathlands

Questions: Does stand age influence the direction and rate of post‐fire successional dynamics in coastal Calluna heaths and can old degraded heath vegetation be restored through reintroduction of fire? Location: Coastal heaths in the Tarva archipelago, central Norway. Methods: We investigated revegetation dynamics after experimental fires set in young (8 years since last fire) and old (>50 years since last fire) grazed heath stands. A repeated measures design was used, with floristic data recorded in permanent plots in the post‐fire successions (n=12) over a 7‐year period. The data were analysed using multivariate ordination techniques (PCA, RDA and PRC) and mixed effects models. Results: The age of Calluna stands strongly influenced post‐fire succession, different trends due to age explained 10.4% of variation in floristic data. Young heath showed faster succession towards pre‐fire community composition than old heath, and this could partially be explained by succession‐related factors: young heath had lower cover of mosses and lichens in the pre‐burned vegetation, and lower cover of litter early in succession. Young heath had a less pronounced overall community response to fire than old heath. Vegetative regeneration of C. vulgaris was absent in both old and young heath, but Calluna still re‐established as the dominant species within 5–7 years in both young and old stands. Regeneration dynamics were also affected by habitat conditions, different trends due to habitat explained 6% of variation. Conclusions: Our study demonstrates that old stands do develop characteristic heathland vegetation and structure after fire, and while potential invasives into the system such as trees and rhizomatous species are present, they do not impair Calluna regeneration or vegetation development towards the target heathland community composition and structure. Further, as our young stands are only in their second fire rotation after restoration, we suggest that characteristic dynamics of managed heathlands can re‐establish relatively rapidly, even in severely degenerated sites (>50 years since last fire). Site‐specific factors also need to be considered. We conclude that there is restoration potential in old heaths, despite slow dynamics in the first rotation.     

Effects of forest continuity on flying saproxylic beetle assemblages in small woodlots embedded in agricultural landscapes

Ancient forests are of considerable interest for strategies for biodiversity conservation. However, in European forest landscapes fragmented and harvested for a long time forest continuity might be no longer a key driver for flying organisms such as saproxylic beetles. In a study based on paired samples (n = 60 stands, p = 180 traps) of ancient and recent forests, we investigated the effects of forest continuity on saproxylic beetle assemblages in two French regions. Mean species richness was significantly related with deadwood volume in ancient forests, but not in recent forests. This loss of relationship between assemblages and their environment suggests that dispersal limitation is at work, at least for some species. Forest continuity had a significant effect on mean species richness and on the mean number of common species, but not on rare species. Forest continuity had a significant effect on assemblage composition in one out of the four cases tested. In both regions, we identified species associated with either recent or ancient forests. Finally, mean body size of species was significantly smaller in recent forests compared with ancient ones, as was their tree diameter preference, despite a higher volume of large deadwood in recent forests. These results lend support to using forest continuity as a criterion to identify sites of conservation importance, even in highly fragmented landscapes.     

Altitudinal effects on habitat selection of dung beetles (Scarabaeoidea: Aphodiidae) in the northern Iberian peninsula

The effects or vegetation cover on the abundance and species richness of dung beetles (Aphodiidae) were examined in three altitudinal zones: lowland (520–600 m), mid-altitude (950–963 m) and highland (1230–1275 m). Pitfall traps baited with fresh cattle dung were placed in three vegetation types at each altitudinal zone: forest, heathland and pasture. Species richness was lower in forests than in the other vegetation types. The total abundance of beetles also varied with vegetation type, but the effect was dependent on altitude. Number of individuals was higher in pasture at highlands and in heathland and forest at mid-altitude and lowlands. An altitudinal dependence of distribution of individuals between vegetation types was also found for the more abundant species in the study area. These results could be explained if the local distribution of individuals between vegetation types was determined by microcli-matic factors, mainly temperature. Differences in altitude between sites are associated with temperature changes, determining the habitat selection of species at each altitude.     

Water hyacinth,Eichhornia crassipes (Pontederiaceae), reduces benthic macroinvertebrate diversity in a protected subtropical lake in South Africa

The socio-economic impacts of the free-floating aquatic plant water hyacinth, Eichhornia crassipes (Pontederiaceae), on aquatic systems are well documented, yet the impacts on aquatic biodiversity, particularly invertebrate biodiversity, are less well understood. This study aimed to determine whether the presence of water hyacinth altered the diversity and assemblage structure of benthic macroinvertebrates in a conservation area. The benthic macroinvertebrate assemblage was sampled over 1 year at five sites under water hyacinth mats and at five sites without water hyacinth at Lake Nsezi—Nseleni River in the vicinity of Richards Bay, KwaZulu-Natal, South Africa. Artificial substrates were placed beneath water hyacinth mats or in the open water to allow for colonization by freshwater macroinvertebrates, and left for a period of 6 weeks, repeated on seven occasions. Twenty nine families comprising 18,797 individuals were collected, 817 (13 families) individuals were from under water hyacinth mat sites compared to 17,980 (27 families) individuals from open water sites. Ninety-eight percent of individuals collected were, however, the invasive snail, Tarebia granifera. Open water samples were separated from samples beneath the water hyacinth mat by non-metric Multidimensional Scaling, indicating reduced biodiversity associated with the presence of water hyacinth. Exclusion of the dominant Thiaridae from the analysis did not alter the groupings. Family richness(s) and abundance (N) were significantly higher in open water communities(S: H₃ = 21.09; P = 0.0001; N: H₃ = 22.58; P = 0.00001), while evenness (J’) was higher under water hyacinth mats (H₃ = 20.13; P = 0.0002). The presence of water hyacinth had a significantly negative impact on aquatic macroinvertebrate biodiversity in a conservation area, and therefore the control of this invasive aquatic plant must play a major role in catchment management.     

Communities of ground-living spiders in deciduous forests: Does tree species diversity matter?

The relationships between species diversity and ecosystem functions are in the focus of recent ecological research. However, until now the influence of species diversity on ecosystem processes such as decomposition or mineral cycling is not well understood. In deciduous forests, spiders are an integral part of the forest floor food web. In the present study, patterns of spider diversity and community structure are related to diversity of deciduous forest stands in the Hainich National Park (Thuringia). In 2005, pitfall trapping and quantitative forest floor sampling were conducted in nine plots of forest stands with one (Diversity Level 1), three (DL 2) and five (DL 3) major deciduous tree species. Species richness, measured with both methods, as well as spider abundance in forest floor samples were highest in stands with medium diversity (DL 2) and lowest in pure beech stands (DL 1). The Shannon-Wiener index and spider numbers in pitfall traps decreased from DL 1 to DL 3, while the Shannon-Wiener index in forest floor samples increased in the opposite direction. Spider community composition differed more strongly between single plots than between diversity levels. Altogether, no general relationship between increasing tree species diversity and patterns of diversity and abundance in spider communities was found. It appears that there is a strong influence of single tree species dominating a forest stand and modifying structural habitat characteristics such as litter depth and herb cover which are important for ground-living spiders.     

Does selective logging change ground‐dwelling beetle assemblages in a subtropical broad‐leafed forest of China?

Selective logging with natural regeneration is advocated as a near‐to‐nature strategy and has been implemented in many forested systems during the last decades. However, the efficiency of such practices for the maintenance of forest species are poorly understood. We compared the species richness, abundance and composition of ground‐dwelling beetles between selectively logged and unlogged forests to evaluate the possible effects of selective logging in a subtropical broad‐leafed forest in southeastern China. Using pitfall traps, beetles were sampled in two naturally regenerating stands after clearcuts (ca. 50 years old, stem‐exclusion stage: selectively logged 20 years ago) and two mature stands (> 80 years old, understory re‐initiation stage: selectively logged 50 years ago) during 2009 and 2010. Overall, selective logging had no significant effects on total beetle richness and abundance, but saproxylic species group and some abundant forest species significantly decreased in abundance in selectively logged plots compared with unlogged plots in mature stands. Beetle assemblages showed significant differences between selectively logged and unlogged plots in mature stands. Some environmental characteristics associated with selective logging (e.g., logging strategy, stand age, and cover of shrub and moss layers) were the most important variables explaining beetle assemblage structure. Our results conclude that selective logging has no significant impacts on overall richness and abundance of ground‐dwelling beetles. However, the negative effects of selective logging on saproxylic species group and some unlogged forest specialists highlight the need for large intact forested areas for sustaining the existence of forest specialist beetles.     

Trait-specific response of ground beetles (Coleoptera: Carabidae) to forest fragmentation in the temperate region in Korea

The objective of this study was to examine the trait-specific response of ground beetles in terms of abundance, species richness, and composition for habitat fragmentation in temperate forests. In addition, we examined how different ecological groups and species of ground beetles responded to environmental variables. During middle May to early November in 2013, ground beetles were sampled using pitfall traps in 27 sites (18 patches and 9 continuous forests) in central Korea. A total of 51 species were identified from 17,845 ground beetles in the 27 study sites. Continuous forests had generally higher abundance and species richness of ground beetles than forest patches. Canonical correspondence analysis for ground beetles was significant, and the proportion of variance explained by environmental variables, such as patch size, elevation, organic matter (C), and soil moisture and pH, was 43.61%. In addition, species composition of ground beetles in continuous forests was grouped distinctively away from those in medium and small-sized patches. In conclusion, both small (1.1–9.6 ha) and medium patches (12.8–51.2 ha) were failed to preserve similar ground beetle biodiversity compared to continuous forests. However, our study revealed that medium-sized forest patches clearly had higher conservation value for forest specialist ground beetles than small-sized forest patches irrespective of forest type.     

An examination of scale of assessment,logging and ENSO-induced fires on butterfly diversity in Borneo

The impact of disturbance on species diversity may be related to the spatial scales over which it occurs. Here I assess the impact of logging and ENSO (El Niño Southern Oscillation) -induced burning and forest isolation on the species richness (477 species out of more than 28,000 individuals) and community composition of butterflies and butterfly guilds using small (0.9 ha) plots nested within large (450 ha) landscapes. The landscapes were located in three habitat classes: (1) continuous, unburned forest; (2) unburned isolates surrounded by burned forest; and (3) burned forest. Plots with different logging histories were sampled within the two unburned habitat classes, allowing for independent assessment of the two disturbance factors (logging and burning). Disturbance within habitat classes (logging) had a very different impact on butterfly diversity than disturbance among habitat classes (due to ENSO-induced burning and isolation). Logging increased species richness, increased evenness, and lowered dominance. Among guilds based on larval food plants, the species richness of tree and herb specialists was higher in logged areas but their abundance was lower. Both generalist species richness and abundance was higher in logged areas. Among habitat classes, species richness was lower in burned forest and isolates than continuous forest but there was no overall difference in evenness or dominance. Among guilds, generalist species richness was significantly lower in burned forest and isolates than continuous forest. Generalist abundance was also very low in the isolates. There was no difference among disturbance classes in herb specialist species richness but abundance was significantly higher in the isolates and burned forest than in continuous forest. Tree specialist species richness was lower in burned forest than continuous forest but did not differ between continuous forest and isolates.The scale of assessment proved important in estimating the impact of disturbance on species richness. Within disturbance classes, the difference in species richness between primary and logged forest was more pronounced at the smaller spatial scale. Among disturbance classes, the difference in species richness between continuous forest and isolates or burned forest was more pronounced at the larger spatial scale. The lower levels of species richness in ENSO-affected areas and at the larger (landscape) spatial scale indicate that future severe ENSO events may prove one of the most serious threats to extant biodiversity.     

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.     

The importance of small-scale heterogeneity in boreal forests: variation in diversity in forest-floor invertebrates across the succession gradient

Heterogeneity in species assemblages of forest-floor arthropods — carabid beetles, ants and spiders — within and between different forest age classes was studied in the southern Finnish taiga. The importance of processes operating on the local scale (within the movement radius of the species) vs on the regional scale (among forest stands) in determining the observed variation was assessed. Four data sets with different spatial resolutions in mesic forests in the same general study area were used. The material consists of 18 283 carabids of 51 species, 48 769 spiders of 212 species, and 126 718 worker ants of 23 species. Analyses of abundance variation and species complementarity among successional stages revealed that in all the three taxa species occurring in the mature forest were prevalent in the younger successional stages as well, constituting more than half of the catch in any age class. A great majority of carabid and spider species were widely distributed across the forest age classes, whereas ants include a higher proportion of species with a narrower amplitude across the succession gradient. Comparisons of similarity between samples at increasing distance from one another on the local scale within forest stands (a few tens of meters to a few hundreds of meters) showed a quite consistent pattern in carabids and spiders: there was more variation between sampling sites in young successional forests than in the mature forest. Furthermore, only in the mature forest a slight, albeit statistically not significant, negative relationship between similarity of samples and distance between sampling sites was detected. In carabids and spiders, comparisons between samples located at a distance of 10–15 m from each other showed considerable heterogeneity, the mean percentage similarity being c. 0.6 (in ants c. 0.8). On the regional scale, systematic variation between young and mature forest stands is a major element increasing the total diversity (species turnover c. 50% in carabids and spiders; compositional similarity c. 0.3–0.4 in carabids, 0.2–0.3 in spiders), but variation within forest stands on a spatial scale of 10–15 meters is another important component in the total heterogeneity. The results suggest that regional abundance variation is a primary factor influencing the composition of local assemblages; a set of hypotheses elaborating this conclusion is formulated. The result implies that maintenance of habitat heterogeneity on a small scale (10–15 m) is needed to preserve biodiversity in managed forests.