Different long-term and short-term responses of land snails to clear-cutting of boreal stream-side forests

Effects of clear-cutting on biodiversity have mainly been studied in the short-term, although knowledge of longer term effects are often more important for managers of forest biodiversity. We assessed relatively long-term effects of clear-cutting on litter dwelling land snails, a group with slow active dispersal and considered to be intolerant to microclimate changes. In a pair wise design we compared snail abundance, species density, and species composition between 13 old seminatural stream-side stands and 13 matched young stands developed 40-60 years after clear-cutting. Using a standardized semi-quantitative method, we identified all snail specimens in a 1.5 l subsample of a pooled litter sample collected from small patches within a 20 × 5 m plot in each stream-side stand. From the young stands a mean of 135 shells and 9.5 species was extracted which was significantly higher than the 58.1 shells and 6.9 species found in old forests. Only two of the 16 species encountered showed a stronger affinity to old than to young forests. In short-term studies of boreal stream-side forests land snail abundance is reduced by clear-cutting. Our results indicate that this decline is transient for most species and within a few decades replaced by an increase. We suggest that local survival in moist stream-side refugia makes the land snails able to benefit from the higher pH and more abundant non-conifer litter in young than in old boreal forests. Our results highlight the importance of longer term studies as a basis for management guidelines for biodiversity conservation.     

Effects of forest management on carabid beetles in Belgium: implications for biodiversity conservation

We sampled the carabid beetles in 22 forests managed by six different silvicultural systems, defined by treatment and tree species composition: even-aged conifer, even-aged beech, even-aged oak, uneven-aged conifer, uneven-aged beech and group mixed (beech + conifer). In each of these forests, we placed pitfall traps in young, medium-aged and mature stands (3 stages). We evaluated the effect of treatment, tree species composition, silvicultural system, stage and habitat type (silvicultural system + stage) on indicators of community conservation value and ecological structure. The species composition and the ecological structure of carabid beetles of the managed stands were then compared to that of nine unmanaged stands (without tree exploitation). In the managed forests, species richness was highest in large young stands (3-10 years old) and in forests managed by even-aged systems (with large clear-cuts), mainly due to eurytopic and opportunist carabid species with high dispersal abilities. Oak and beech, uneven-aged, and mature stands were mainly inhabited by typical forest species, and even-aged conifer stands mainly by ubiquitous species. Several typical forest species recorded in unmanaged stands were lacking from the managed forests. Large scale clear-cutting allows open-habitat species to enter the forest, which increases the species richness at a landscape level but can disfavour typical forest species by competition. Long rotations should be implemented and more areas left unmanaged in Belgium, in order to help typical forest species to re-colonise managed forests.     

Openness in management: Hands-off vs partial cutting in conservation forests, and the response of beetles

The appropriate management of forest reserves is debated; two major alternatives are succession to ‘wild’ state, or management to produce semi-open stands. For temperate conservation stands, there are no strong experiments replicated at landscape level. In each of 22 forests rich in oaks (Quercus spp.) in Sweden, we set up a closed-canopy wild plot (1 ha), and a cutting plot (1 ha) to produce semi-open conditions, studying them before and after cutting. About 25% of the tree basal area was cut (large trees and dead wood retained) and harvested as bio-fuel, a CO₂ - neutral energy source. We examined the response of beetles and trapped 59,000 individuals (1174 species; 100 red-listed species). For both the guild of herbivorous beetles (222 species) and of saproxylic beetles connected to oak wood (267 species), species richness increased by about 35% in the harvested plots, relative to the wild reference plots. Species composition within the groups changed, though not strongly. Thirteen saproxylic species of 50 analysed, and three herbivores of 12 increased in cutting plots. For red-listed saproxylic beetles, species richness did not change significantly. Regression analyses suggest that more open cutting plots disfavour the red-listed beetles of this forest type. Thus, partial cutting increased species diversity of two beetle groups, probably due to changed microclimate and increase in herbaceous plants, but some red-listed saproxylic beetles may be disfavoured. A hands-off alternative may through storms and other disturbances produce open patches, more dead wood, and favour some species. Combinations of these alternatives, carefully planned at the landscape level, need to be considered.     

Epiphyte communities on the trunks of retention trees stabilise in 5 years after timber harvesting, but remain threatened due to tree loss

Live retention trees are expected to support the recovery of epiphytes in regenerating stands by retaining a part of the populations in cutover sites and receiving propagules from adjacent forests. So far, the research has been focused on immediate post-harvesting mortality caused by microclimatic stress while a broader perspective on epiphyte community dynamics is lacking. We studied lichen and bryophyte communities on the trunks of retention trees and adjacent forest trees in Estonia, where significant desiccation (particularly of bryophytes) had been documented within 2-3 years after timber harvesting. The resampling 5-6 years after harvesting indicated that, during the 3 years passed, (1) lichen species richness per surviving tree increased and bryophyte species richness stabilised, (2) there were no clear successional changes in the composition of the communities and (3) retention trees were more frequently colonised than forest trees. Most epiphyte extinctions between the sampling years were related to the death of trees (particularly in the forests because of harvesting) and stochastic disappearances of the smallest populations. Also, retention trees were very rarely colonised by species of conservation concern. We conclude that, in addition to addressing the microclimatic stress in the first post-harvesting years, crucial elements in sustaining epiphytic bryophyte and lichen populations in green-tree retention systems include careful selection of the retention trees and a supportive reserve network. The selection of the trees should assure representativity and long-term survival of local populations, while reserves should host the most demanding species and be stable colonisation sources in general.     

Drastic erosion in functional attributes of tree assemblages in Atlantic forest fragments of northeastern Brazil

The long-term dynamics of plant communities remain poorly understood in isolated tropical forest fragments. Here we test the hypothesis that tropical tree assemblages in both small forest fragments and along forest edges of very large fragments are functionally much more similar to stands of secondary growth (5-65-yr old) than to core primary forest patches. The study was carried out in a severely fragmented landscape of the Brazilian Atlantic forest. Nine functional attributes of tree assemblages were quantified by sampling all trees (DBH ? 10 cm) within 75 plots of 0.1 ha distributed in four forest habitats: small forest fragments (3.4-79.6 ha), forest edges, second-growth patches, and primary forest interior areas within a large forest fragment (3500 ha). These habitats were markedly different in terms of tree species richness, and in the proportion of pioneer, large-seeded, and emergent species. Age of second-growth stands explained between 31.4% and 88.2% of the variation in the functional attributes of tree assemblages in this habitat. As expected, most traits associated with forest edges and small forest fragments fell within the range shown by early (<25-yr old) and intermediate-aged secondary forest stands (25-45-yr old). In contrast to habitat type, tree assemblage attributes were not affected by vegetation type, soil type and the spatial location of plots. An ordination analysis documented a striking floristic drift in edge-affected habitats. Our results suggest that conservation policy guidelines will fail to protect aging, hyper-fragmented landscapes from drastic impoverishment if the remaining forest patches are heavily dominated by edge habitat.     

Bryophyte responses to fragmentation in temperate coastal rainforests: A functional group approach

The species richness and frequency of occurrence of bryophytes within taxonomic and functional groups was examined in relation to the size of 20 old-growth patches (size range: 0.6-63.6 ha) remaining after logging in temperate rainforests of coastal British Columbia. At the centre of each remnant patch, bryophytes were sampled in sixty-three 10 cm × 30 cm microplots on three substrate-types (forest floor, downed logs and tree bases). Generalized linear models demonstrated that the species richness and frequencies of some bryophyte functional groups were related to patch size. In particular, some dispersal-limited groups (perennial stayers) and microclimate-sensitive groups (closed canopy species, epixylic (log-dwelling) species, and liverworts) showed significant declines in either richness or frequency as patch size decreased. In contrast, colonists and open canopy species showed little association with patch size. Many, but not all, of the significant patch size relationships disappeared when the three smallest patches (0.6-1.8 ha) were eliminated from the analysis. These results suggest that patches sized 3.5 ha or larger may provide habitat capable of sustaining a diverse array of bryophyte functional groups in temperate rainforest landscapes.     

The effect of stand structure and tree species composition on epiphytic bryophytes in mixed deciduous-coniferous forests of Western Hungary

The effect of tree species, stand structure, landscape and historical variables was studied on the species composition, species richness and cover of epiphytic bryophyte assemblages in mixed deciduous-coniferous forests of Western Hungary. Stand and tree level assemblages were analyzed by ordinations and generalized linear modeling in 35 70-110 year old stands of different management regimes.Bryophytes showed a considerable preference to different host trees, so that stand level diversity of bryophyte assemblages was determined mainly by tree species diversity, and their composition by tree species composition. Cover and diversity of epiphytic bryophytes were the highest on oaks (Quercus petraea and Quercus robur), and the lowest on Scotch pine (Pinus sylvestris). The presence of sapling (shrub) layer increased, whereas a large number of medium sized trees decreased bryophyte species richness in this study. Tree size was much less influential which is explained by the lack of large, veteran trees. Forest management maintaining tree species diversity, structural heterogeneity and temporal continuity of the stands could considerably contribute to the conservation of this organism group. Selective cutting is more appropriate for these conservational purposes than shelterwood management system.     

Land use and habitat gradients determine bird community diversity and abundance in suburban, rural and reserve landscapes of Minnesota, USA

Bird species’ community responses to land use in the suburbanizing Twin Cities, Minnesota, USA, were contrasted among reserves, rural lands, and suburbs. For each land use type, bird composition, diversity, and abundance were recorded for 2 years in ≈99 plots in three sampling units (each ≈4500 ha). A habitat gradient defined by canopy structure (grasslands to savannas to forests) was influenced by land use, so ≈300 plots were used to characterize simultaneous variation in bird communities along land use and habitat gradients. At broad scales (aggregate of 33 plots covering ≈4500 ha) suburbs supported the lowest bird richness and diversity and rural landscapes the most, with reserves slightly below rural. Although reserves were like rural lands in diversity of bird communities, they supported more species of conservation concern, particularly of grasslands and savannas. Differences among land use types varied with habitat structure. Suburbs, rural lands, and reserves had similar forest bird communities, but differed in grassland and savanna bird communities. The extensive rural forests are important for the region’s forest birds. Suburban grasslands and savannas had low shrub abundance, low native bird richness and high non-native bird richness and abundance. However, total bird richness and diversity were as high in suburban as in rural and reserve plots because high native richness in suburban forests and high non-native species richness in suburban grasslands and savannas compensated for lower native richness in suburban grasslands and savannas. Bird conservation here and in the Midwest USA should protect rural forests, expand grasslands and savannas in reserves, and improve habitat quality overall.     

Polypore diversity in the herb-rich woodland key habitats of Koli National Park in eastern Finland

Species diversity of polypores (Basidiomycetes) was compared between woodland key habitats (WKHs) and old-growth forest controls in boreal forests in eastern Finland. WKHs, which were set aside for their rich vascular plant flora, turned out not to be hot spots for the species richness of polypores, nor did their species composition represent the overall species richness of the area. Differences in the total volume of CWD, tree species composition and several CWD qualities were reflected as differences in the species assemblages between the groups. The results indicated that only a fraction of the overall polypore diversity was represented in the small-size WKHs (<0.5 ha), and the protection of red-listed and indicator polypores in WKHs was random. However, rare old-growth forest indicators were found even in small-size WKHs, if the CWD quality was appropriate for them. Nevertheless, our study does not answer the question whether the occurrences of rare and red-listed species will survive in small-size WKHs surrounded by altered environments such as clear-cuts, because our study sites were located in undisturbed areas. The results stress the importance to protect typical old-growth forest patches with high CWD volume and quality in boreal forests.     

Bird community structure across riparian buffer strips of varying width in a coastal temperate forest

Buffer strips are strips of forest retained along streamsides after harvesting to mitigate negative impacts of forestry on aquatic and riparian fauna and water quality. The capacity of riparian buffer strips of old-growth forest to maintain species richness and abundance of natural bird communities was explored in coastal montane forest on Vancouver Island, Canada. Breeding bird communities in buffer strips of varying widths along rivers were compared with controls of equivalent area in uncut old-growth riparian forest to identify shifts in species richness, diversity, abundance and composition. We observed that effects on riparian bird communities were greatest in very narrow buffers with high amounts of edge habitat. Several forest-interior species were found almost exclusively in wider buffers and abundances dropped dramatically between wide (125 m) and medium (41 m) width buffers with replacement by open-edge species in narrow buffers. Species composition of communities in wide buffers were very similar to controls while narrow buffers shared less than half of their species with controls. Species richness and diversity increased in buffers over the three years while remaining constant in controls. Increases in species richness and abundance within buffers were positively correlated with similar increases in the adjacent clearcuts, suggesting that regeneration in clearcuts may facilitate recolonization of forest in remnant buffers. For the forest-interior species found primarily in wide buffers, buffers >100 m may need to be retained.     

Polypore diversity in managed and old-growth boreal Picea abies forests in southern Finland

Polypore communities were compared between mature managed, overmature managed, and old-growth spruce-dominated forests in southern Finland. A total of 85 polypore species, with 6000 records, were found in 16 sample plots, each 4 ha in size. Old-growth stands had on average 80% more species than mature stands, and 38% more species than overmature managed stands. Variation in polypore species richness was best explained by diversity of dead wood and the number and volume of dead trees. The best predictor for the number of threatened polypore species was the number of cut stumps. Threatened species were practically confined to old-growth forests and to stands in which the amount of dead wood exceeded 20 m³/ha. This figure appears to represent a stand-level threshold value for the amount of dead wood, below which the persistence of threatened species becomes unlikely. Our results suggest that a significant increase in the amount of dead wood (e.g. by leaving large retention trees and even by killing trees) is needed in managed forests before they become suitable habitats for threatened polypores.     

Does forest continuity matter in conservation? - A study of epiphytic lichens and bryophytes in beech forests of southern Sweden

The effects of forest continuity at local scale for red-listed and indicator species of epiphytic lichens and bryophytes were investigated in 150 Fagus sylvatica stands in southern Sweden. Stands having forest continuity (n = 106) had continuous forest cover more than 350 years, whereas stands lacking continuity (n = 44) had forest cover less than 160 years. Forest continuity was identified by comparing a sequence of historical maps with a modern survey of beech forests. In the field woody beech substrates were searched for the epiphytes of interest. A number of environmental and spatial variables were inventoried and compiled for each stand. In all 64 species (51 lichens, 13 bryophytes) were found in the stands having continuity, and 21 (14 lichens, 7 bryophytes) in the stands lacking continuity. Controlling for the different number of surveyed stands, stands having continuity had significantly more species of lichens, but not of bryophytes. In the stands lacking continuity we did not find lichens associated with the very late succession stage. The quantity of substrates, stand age and forest continuity were the three most important factors explaining species richness as well as composition of studied epiphytes. The effect of continuity was probably due to a combination of a higher substrate quality, mainly old beeches, and a longer time available for colonization. Also, we found strong positive correlations between number of indicator and red-listed epiphyte species. In short-term conservation old stands having continuity, containing suitable substrates and indicator species are target areas.     

Does closure of traditionally managed open woodlands threaten epigeic invertebrates? Effects of coppicing and high deer densities

The demise of traditional woodland management techniques, such as coppicing or woodland pasture, is causing a gradual closure of formerly sparse lowland woods across Central Europe. It is established that these processes threaten such organisms as butterflies and higher plants. Effects on other groups, such as epigeic invertebrates, are little known, hindering rational conservation decisions. We investigated the effects of stand openness on three epigeic groups, carabids (Carabidae), arachnids (Araneae + Opiliones) and myriapods-isopods (Chilopoda + Diplopoda + Oniscidea), in a lowland deciduous wood in the Czech Republic. Situating some of the traps in an intensive deer park allowed a simultaneous assessment of effects of high vs. low ungulate densities. Carabids reached the highest species richness in either sparse stands with low game or dense stands, high game. More arachnids occurred under low game and in sparse stands than under high game density and in dense stands. The highest richness of myriapods-isopods was in sparse stands with low game. Ordinations revealed that species of conservation concern (‘relic species’ according to C. European authors) tended to be associated with sparse stands and low game. A considerable proportion of epigeic woodland invertebrates, including many species of conservation concern, depends on preserving highly heterogeneous sparse canopy conditions. Restoring such conditions in selected areas will benefit these sensitive open woodland specialists while causing minimum harm to specialists of dense stands, that likely prosper in commercially managed high forests.     

Woodpeckers as reliable indicators of bird richness, forest health and harvest

Woodpeckers (family Picidae) show promise as indicators of avian diversity in forests because their populations can be reliably monitored, and their foraging and nesting activities can positively influence the abundance and richness of other forest birds. A correlation between woodpecker richness and richness of forest birds is known to exist at the landscape scale, but uncertainty remains whether this correlation occurs at the smaller stand-level spatial scales where forest management activities take place. We used data collected under a diverse range of forest types, harvest treatments, and forest health conditions during a long-term study of bird communities in interior British Columbia, Canada, to examine two basic questions: (1) at the level of individual forest stands, is woodpecker richness correlated with bird richness (measured as richness of all other bird species)? and (2) do woodpecker richness and bird richness have similar habitat correlates? Bird richness was positively correlated with woodpecker richness (β = 0.59, SE = 0.22, 95% CI = [0.14 1.03]). Richness of both woodpeckers and all other birds were positively correlated with tree species richness and negatively correlated with density of pines, and the effect for forest harvest type was similar for both measures of avian richness (uncut < partial harvest < clearcut with reserves). The effect of density of lodgepole pines killed by mountain pine beetles differed between the two richness measures, being positive for woodpecker richness and negative for forest bird richness. We conclude that the richness of woodpeckers is indeed correlated with the richness of other birds at the stand-level, and can serve as a reliable indicator of overall bird richness in most forest stands and conditions, except during insect outbreaks when differential responses by woodpeckers and the rest of the avian community may decouple the relationship between bird richness and woodpecker richness.     

Isolation and edge effects among woodland key habitats in Sweden: Is forest policy promoting fragmentation?

Fragmentation of natural forests is a major threat to forest biodiversity. In areas with a long history of forestry, the remaining patches of old forests constitute only a minor part of the landscape. In such situations small stands may be valuable and important for conservation. However, as they may suffer from strong edge effects and isolation, their value may be lower than anticipated. In Sweden a national inventory of woodland key habitats (WKHs) has identified about 1% of the forest landscape as sites where red-listed species occur or may occur. Most are small (national median 1.4 ha) and isolated stands within an intensively managed landscape. The present analyses calculate WKH core area based on a range of depths of edge influence, and isolation based on both distance to nearest WKH and a weighted isolation measure that includes all neighboring WKHs and protected forest. These analyses are done on the WKHs in Norrbotten County in northern Sweden and include almost 5000 stands. The actual core area in the WKHs is about 30% given a 50 m edge influence. The degree of isolation is species dependent but the results indicate that only species with high dispersal abilities may effectively utilize the network of WKHs. For species with effective dispersal distances of less than 2 km the network is probably insufficient. The results emphasize the need to create buffer zones, to increase reserve areas and to manage the matrix so that species dispersal is promoted. This likely includes a necessity to aggregate biodiversity efforts on the landscape scale.     

Effect of hydrologic regime and forest age on Collembola in riparian forests

The principal objective of the study was to assess factors of primary importance for collembolan community variability measured in riparian forest stands of different age after traditional clear cutting within natural and altered hydrologic regimes. The study was conducted in the Ukrainian part of the Latorica river floodplain where the largest stands of the highly protected medio-European Querco—Ulmetum minoris fluvial forests can be found. Six oak forest stands were investigated on the river floodplain: three in a natural section (inside levee) of the floodplain, which have their original features preserved with periodical inundation, and three in a section separated by a flood control embankment (outside levee) and subjected to serious damage by drainage. In each section the three stands sampled were represented by different ages regenerated with clear-cutting (>3 years, >8 years and >112 years).Differences in hydrologic regimes were central to identification of a predictable and explainable percentage of variation in species composition of Collembola communities in fluvial forests. The hydrologic regime factor was correlated with the basic structural attributes of collembolan communities, such as abundance, species richness and some ecological traits. Season accounted for a higher degree of variation in collembolan communities than did the age of forest stands. We conclude that differences in hydrologic regime are of much higher importance in structuring collembolan communities in riparian forests than are the ages of stands.     

Enhancing vegetation diversity in glades, rides and roads in plantation forests

Permanent open spaces in plantation forests provide an opportunity for enhancing biodiversity in plantations and in the wider landscape. We surveyed vegetation in 60 glades, rides and roads in 12 Irish plantation forests, and collected data on solar radiation, soil and management. Variation in species richness and diversity of vascular plants and bryophytes was determined principally by soil factors and open space management. Light levels were positively associated with vascular plant species richness and negatively associated with richness of bryophytes. In rides, the most important plantation feature in determining solar radiation levels and plant diversity was ride width. Increasing edge-to-area ratio corresponded with a decrease in β-diversity within glades. Roadside plots most strongly influenced by road gravel and disturbance supported the highest vascular plant, open-habitat and ruderal species richness and vegetation evenness. For open spaces to contribute significantly to the biodiversity of plantation forests, rides and roads should be a minimum of 15 m in width and glades should be a minimum of 625-900 m² in area.     

Bryophyte species richness on boulders: relationship to area, habitat diversity and canopy tree species

Bryophytes growing on siliceous boulders (diameter 50-200 cm) were studied in two forests in east-central Sweden to investigate the influence of different canopy tree species on the bryophyte species richness. Granite boulders lying below crowns of Ulmus glabra (elm), Fraxinus excelsior (ash), Acer platanoides (maple), Quercus robur (oak), Betula pendula (birch), and Picea abies (Norway spruce) were studied in two mixed stands. Both boulder area and within-boulder habitat diversity were positively related to species richness. Boulders below the base-rich deciduous trees F. excelsior, U. glabra and A. platanoides held roughly twice as many species as those below P. abies, with Q. robur and B. pendula as intermediates. The rank order among trees was as expected from bark pH and litter decomposability in the literature. We also used sample plots to investigate species richness at a smaller scale. Within plots, intermediate levels of litter, inclination and exposed rock promoted species richness, and also the covering tree species had an effect. Individual plots were on average not more species-rich on large boulders than on small ones. This leads us to conclude that population extinctions on the plot-level, and re-colonisations from other parts of the boulder, are less important for species richness than within-boulder habitat diversity. Since species richness was lower under P. abies it is important to favour a diversity of tree species and to include base-rich deciduous trees in otherwise homogeneous Picea forests.     

The conservation value of urban riparian areas for landbirds during spring migration: Land cover, scale, and vegetation effects

Urbanization changes bird community structure during the breeding season but little is known about its effects on migrating birds. We examined patterns of habitat use by birds at the local and landscape level during 2002 spring migration at 71 riparian plots along an urban gradient in Cincinnati, Ohio, USA. Using linear regression, we examined variation in relative density, species richness, and evenness of four migratory guilds associated with natural land covers and building area at four scales (50, 100, 250, 500 m radial buffers). We also examined the influence of local vegetation using multiple regression models. As building area increased, riparian forests tended to be narrower and have fewer native trees and shrubs. In general, native birds were positively associated with tree cover (within 250-500 m of stream) and native vegetation, and negatively with building area (within 250 m); exotic species responded inversely to these measures. Short-distance migrants and permanent residents displayed the weakest responses to landscape and vegetation measures. Neotropical migrants responded strongest to landscape and vegetation measures and were positively correlated with areas of wide riparian forests and less development (>250 m). Resident Neotropical migrants increased with wider riparian forests (>500 m) without buildings, while en-route migrants utilized areas having a wide buffer of tree cover (250-500 m) regardless of buildings; both were positively associated with native vegetation composition and mature trees. Consequently, developed areas incorporating high native tree cover are important for conserving Neotropical migrants during stopover.     

Conservation of epiphytic lichens: Significance of remnant aspen (Populus tremula) trees in clear-cuts

In the boreal forest landscape, aspen has been effectively selected against in favour of conifers. The decrease in aspen is of particular concern since it has more host-specific species associated with it than any other boreal tree species. Recently forest management systems have begun to include green-tree retention in order to maintain structural diversity. We initiated this study in order to investigate whether retained aspen trees provide suitable habitats for epiphytic lichens and, if so, whether this varies according to species characteristics, such as morphology and photobiont. We chose to investigate the abundance and vitality of five species: three foliose cyanobacterial lichens (Collema curtisporum, Collema furfuraceum and Leptogium saturninum) and two crustose green-algal lichens (Biatora epixanthoides and Mycobilimbia carneoalbida). These lichens were examined on aspen within forest stands and on remnant aspen in clear-cuts at eight localities, in northern Sweden. Our results show that the response to exposure following clear-felling differs between species according to their morphology. The two crustose lichens were more abundant in the forest stands compared to the clear-cuts. In the clear-cuts, they mainly occurred on the northern sides of the trunks. Even 24 years after cutting, the three cyanolichens were equally or even more abundant on trees in the clear-cuts than in the forest stands. However, they were mainly found on the northern sides of the remnant aspen. We suggest that remnant aspen trees may function as suitable substrate and as stepping stones for colonisation of new stands, at least for the cyanolichens studied here.