Long‐term successional forest dynamics: species and community responses to climatic variability

Question: Are trees sensitive to climatic variability, and do tree species differ in their responses to climatic variability? Does sensitivity of forest communities to climatic variability depend on stand composition? Location: Mixed young forest at Walker Branch Watershed near Oak Ridge, East Tennessee, USA. Methods: Using a long‐term dataset (1967–2006), we analyzed temporal forest dynamics at the tree and species level, and community dynamics for forest stands that differed in initial species composition (i.e., chestnut oak, oak–hickory, pine, and yellow poplar stands). Using summer drought and growing season temperature as defined climate drivers, we evaluated relationships between forest dynamics and climate across levels of organization. Results: Over the four‐decade study period, forest communities underwent successional change and substantially increased in biomass. Variation in summer drought and growing season temperature contributed to temporal biomass dynamics for some tree species, but not for others. Stand‐level responses to climatic variability were related to the responses of component species, except in pine stands. Pinus echinata, the dominant species in pine stands, decreased over time due to periodic outbreaks of pine bark beetle (Dendroctonus frontalis). These outbreaks at Walker Branch could not be directly related to climatic conditions. Conclusions: The results indicate that sensitivity of developing forests to climatic variability is stand type‐dependent, and hence is a function of species composition. However, in the long term, direct effects of climatic variability on forest dynamics may be small relative to autogenic successional processes or climate‐related insect outbreaks. Empirical studies testing for interactions between forest succession and climatic variability are needed.     

Diversity and succession of epiphytic macrolichen communities in low‐elevation managed conifer forests in Western Oregon

Abstract. We examined epiphytic macrolichen communities in Pseudotsuga menziesii (Douglas‐fir) forests across the western Oregon landscape for relationships to environmental gradients, stand age and structure, and commercial thinning. We used a retrospective, blocked design through the Coast and the western Cascade ranges of Oregon. Each of our 17 blocks consisted of a young, unthinned stand (age 50–110 yr); an adjacent, thinned stand of equivalent age; and an old‐growth stand (age > 200 yr). We found 110 epiphytic macrolichen taxa in the stands. Forage‐providing alectorioid lichens and the nitrogen‐fixing cyanolichen Lobaria oregana associated strongly with old‐growth stands and remnant old trees in younger stands (unthinned + thinned). Relative to unthinned stands, thinned stands had a slightly higher abundance of alectorioid lichens and a greater presence of Hypogymnia imshaugii. However, thinned stands hosted a lower landscape‐level (γ) diversity, lacking many species that occurred infrequently in the unthinned stands. Patterns in the lichen community composition correlated strongly with climatic gradients; the greatest variation in composition was between the Coast and Cascade ranges. The difference in communities between mountain ranges was greatest among stands 70–110 yr old, suggesting a difference in lichen successional dynamics between the ranges.     

Aspen (<Emphasis Type="Italic">Populus tremuloides</Emphasis>) stands and their contribution to plant diversity in a semiarid coniferous landscape

We conducted a field study to determine the relative contributions of aspen (Populus tremuloides), meadow, and conifer communities to local and landscape-level plant species diversity in the Sierra Nevada and southern Cascade Range, northeastern California, USA. We surveyed plant assemblages at 30 sites that included adjacent aspen, conifer, and meadow communities across a 10,000-km² region. We statistically investigated patterns in local and landscape-scale plant diversity within and among the three vegetation types. Summing across sites, aspen stands supported more plant species overall and more unique plant species than either meadow or conifer communities. Local richness and diversity did not differ between aspen and meadow plots; conifer forest plots were significantly lower in both measures. Heterogeneity in species composition was higher for aspen forest than for meadows or conifer forest, both within sites and between sites. Plant communities in aspen stands shared less than 25% of their species with adjacent vegetation in conifer and meadow plots. Within aspen forest, we found a negative relationship between total canopy cover and plant diversity. Our results strongly support the idea that plant communities of aspen stands are compositionally distinct from adjacent meadows and conifer forest, and that aspen forests are a major contributor to plant species diversity in the study region. Current patterns of aspen stand succession to conifer forest on many sites in the semiarid western US are likely to reduce local and landscape-level plant species diversity, and may also have negative effects on other ecosystem functions and services provided by aspen forest.     

Litterfall of epiphytic macrolichens in Nothofagus forests of northern Patagonia,Argentina: Relation to stand age and precipitation

Abstract: The objective of this study was to analyse how stand age and precipitation influence abundance and diversity of epiphytic macrolichens in southern beech Nothofagus forests, estimated by lichen litter sampling. Five sites of Nothofagus dombeyi (Mirbel) Oersted were selected in Nahuel Huapi National Park, Argentina. At each site, lichen fragments from the forest floor were collected at 12.5 m² plots in pairs of young and mature N. dombeyi forest. Additionally, two sites with multi‐aged subalpine Nothofagus pumilio (Poepp. et Endl.) Krasser forest were investigated in a similar manner. Average litterfall biomass per stand varied from less than 1 kg ha^?1 in a young low‐precipitation stand to a maximum of 20 kg ha^?1 in a mature high‐precipitation stand. In places with higher precipitation, litterfall biomass in N. dombeyi forest was considerably higher in old stands as compared with young ones. In places with less than 2000 mm of precipitation, differences in biomass were less pronounced. Old humid stands contained about twice as many taxa in the litter as old low‐precipitation stands and young stands in general. Mature stands in low‐precipitation sites only contained 17% of the litter biomass as compared with mature stands in high‐precipitation sites. Epiphytic lichen composition changed from predominating fruticose lichens (Usnea spp. and Protousnea spp.) in low‐precipitation stands to Pseudocyphellaria spp., Nephroma spp. and other foliose lichens, in the high‐precipitation stands. There were no clear differences in the proportion of fruticose and foliose lichens between young and old stands. Fruticose lichens dominated litter biomass in both N. pumilio sites.     

Edge effects on epiphytic communities in a Mediterranean Quercus pyrenaica forest

Question: What are the edge effect responses of epiphytic lichen communities in Mediterranean Quercus pyrenaica forest? Location: Central Spain. Methods: We established ten transects perpendicular to a road dissecting a well conserved remnant of Q. pyrenaica forest into two sections. Transects extended from the forest/road edge to 100 m into the forest. Data were collected from seven plots in each transect at different distances from the edge. Variables were grouped into stand scale variables (distance to edge, number of trees per plot, mean diameter per plot, irradiance) and tree scale variables (diameter and height of sampled trees, aspect of the sampled square and relative height of the square). We used General Mixed Linear Models and constrained ordination techniques to test the hypothesis that the spatio‐temporal heterogeneity of light and water controls the occurrence of lichens and bryophytes along the edge‐interior gradient in the Q. pyrenaica forest. Results: Microclimatic parameters vary in a non‐linear way; edge and interior stands showed the most divergent and extreme values. Although the micro‐environment within Mediterranean forests is heterogeneous, interior conditions are apparently suitable for the performance of some specific forest epiphytes. Consequently, species richness does not show significant differences along the gradient. Total epiphytic cover increases towards the forest interior, but distance to the edge together with other predictors at the tree scale (aspect and height of the square) are the most relevant predictors for the composition and structure of these communities. Conclusions: Composition and structure of epiphytic communities in a Mediterranean semi‐deciduous forest are affected by the edge between the forest and the road constructed. Since some extremely rare lichens only occur at interior stands, the conservation of these threatened elements requires urgent conservation measures because well preserved and unmanaged forests in the Mediterranean region are very rare.     

Bole epiphytic lichens as potential indicators of environmental change in subtropical forest ecosystems in southwest China

Lichen epiphytes are applied as excellent environmental indicators worldwide. However, very little is known about epiphytic lichen communities and their response to forest dynamics in subtropical China. This paper proposes the applications of the cover, diversity, and functional traits of epiphytic lichens to assess environmental changes associated with succession in subtropical forests of southwest China. Bole lichens were sampled from 120 plots of eight representative forest types in the Ailao Mountains. Total cover, species richness, diversity and community structure of bole lichens differed significantly among forest types, and the highest cover and diversity occurred in the Populus bonatii secondary forest (PBSF). Sixty-one indicator species were associated with particular forest types and more than 50% occurred in the PBSF. Both cover and diversity of most lichen functional groups varied regularly during forest succession. Lichen pioneer species were not displaced by competitively superior species as succession proceeds and cyanolichens were more prevalent in secondary forests. The results also highlight the importance of habitat variables such as canopy openness, host diversity, forest age, tree size, the size of the largest tree, tree density, and basal area on the lichen community. Consequently, our findings support the notion that epiphytic lichens, in terms of cover, diversity, species composition and functional traits can be used as effective indicators for large-scale and long-term forest monitoring. More importantly, the narrowly lobed foliose group was the best candidate indicator of environmental conditions in this region. The combined application of lichen indicator species and functional groups seemed to be a more reliable and more powerful method for monitoring forest dynamics in subtropical montane ecosystems.     

Contrasting functional traits maintain lichen epiphyte diversity in response to climate and autogenic succession

Aim Lichen epiphytes are important for biodiversity conservation and are also widely applied as environmental indicators. However, biogeographical and ecological knowledge underpinning lichen epiphyte conservation, and the use of lichens as indicators, is based primarily on a limited range of ‘macrolichen’ species. Wider trends in epiphyte biodiversity remain largely unexplored. This paper examines the community structure of lichen epiphytes on aspen (Populus tremula L.) in Scotland, including species across all functional groups and comprising, therefore, taxonomically difficult ‘microlichens’. Location Northern Britain (Scotland). Methods Epiphytes were sampled from 12 sites throughout Scotland and examined at two scales: between and within aspen stands. Species were classified into contrasting functional groups and ordination by detrended correspondence analysis was used to summarize community structure. Results Within aspen stands (between trees) epiphyte communities showed successional patterns related to tree age. These successional patterns changed predictably for stands aligned along a climatic gradient (between stands). Main conclusions A dual climatic–successional trend in epiphyte community structure is presented. Large‐scale trends in epiphyte diversity are explained as the local response of species with contrasting functional traits to climate and autogenic succession. Turnover of functional groups between stands is positively related to β‐diversity, and ecological limits to the frequency of contrasting functional groups are presented. Accordingly, the study and application of lichen species with similar functional traits may inadequately represent patterns of biodiversity. This prompts criticism of the currently accepted conservation strategy, i.e. (1) an emphasis in the conservation literature on ‘macrolichen’ species with similar ecologies and (2) the application of lichen indices over climatically variable geographical areas.     

Tree and stand level variables influencing diversity of lichens on temperate broad-leaved trees in boreo-nemoral floodplain forests

Tree and stand level variables affecting the species richness, cover and composition of epiphytic lichens on temperate broad-leaved trees (Fraxinus excelsior, Quercus robur, Tilia cordata, Ulmus glabra, and U. laevis) were analysed in floodplain forest stands in Estonia. The effect of tree species, substrate characteristics, and stand and regional variables were tested by partial canonical correspondence analysis (pCCA) and by general linear mixed models (GLMM). The most pronounced factors affecting the species richness, cover and composition of epiphytic lichens are acidity of tree bark, bryophyte cover and circumference of tree stems. Stand level characteristics have less effects on the species richness of epiphytic lichens, however, lichen cover and composition was influenced by stand age and light availability. The boreo-nemoral floodplain forests represent valuable habitats for epiphytic lichens. As substrate-related factors influence the species diversity of lichens on temperate broad-leaved trees differently, it is important to consider the effect of each tree species in biodiversity and conservation studies of lichens. Nomenclature Randlane et al. (2007) for lichens; Leht (2007) for vascular plants.     

Riverscape-level patterns of riparian plant diversity along a successional gradient, Willamette river, Oregon

Riparian forest communities dominated by Populus balsamifera ssp. trichocarpa L. (Torr. and Gray ex Hook.) Brayshaw are important contributors to biodiversity in terrestrial and aquatic ecosystems of the Western United States. Species composition along a successional gradient from stand initiation to late-succession of P. balsamifera-dominated riparian forests was investigated along 145 km of the Willamette River, Oregon. There were 151 total species encountered across 28 stands and a mean species richness of 33.3 species per stand. Young stands were dominated by P. balsamifera and Salix tree spp. and opportunistic herbaceous species. Understory trees, shrubs, and herbaceous species as well as late-successional tree species established 12–15 years after stand initiation. Fraxinus latifolia Benth. was the dominant late-successional tree species. Vertical structural diversity, P. balsamifera mean diameter at breast height, large tree biomass, and stand age were strongly correlated with understory species presence and abundance based on non-metric multidimensional scaling (NMS) ordination. There were no young stands on mid and high terraces and this was reflected in geomorphic position being strongly correlated with the stand age gradient. Abundance of Phalaris arundinacea L. an invasive grass species, was also significantly correlated with plant species composition and abundance. This study indicates that Willamette River riparian forests are diverse and therefore important to the biodiversity of the Willamette River valley and that their presence as a mosaic of communities of different successional stages may be threatened by human interventions, including influences exerted by introduced plant species.     

Implications of floristic and environmental variation for carbon cycle dynamics in boreal forest ecosystems of central Canada

Abstract. Species composition, detritus, and soil data from 97 boreal forest stands along a transect in central Canada were analysed using Correspondence Analysis to determine the dominant environmental/site variables that differentiate these forest stands. Picea mariana stands were densely clustered together on the understorey DCA plot, suggesting a consistent understorey species composition (feather mosses and Ericaceae), whereas Populus tremuloides stands had the most diverse understorey species composition (ca. 30 species, mostly shrubs and herbs). Pinus banksiana stands had several characteristic species of reindeer lichens (Cladina spp.), but saplings and Pinus seedlings were rare. Although climatic variables showed large variation along the transect, the CCA results indicated that site conditions are more important in determining species composition and differentiating the stand types. Forest floor characteristics (litter and humus layer, woody debris, and drainage) appear to be among the most important site variables. Stands of Picea had significantly higher average carbon (C) densities in the combined litter and humus layer (43530 kg‐C.ha^‐1) than either Populus (25 500 kg‐C.ha^‐1) or Pinus (19 400 kg‐C.ha^‐1). The thick surface organic layer in lowland Picea stands plays an important role in regulating soil temperature and moisture, and organic‐matter decomposition, which in turn affect the ecosystem C‐dynamics. During forest succession after a stand‐replacing disturbance (e.g. fires), tree biomass and surface organic layer thickness increase in all stand types as forests recover; however, woody biomass detritus first decreases and then increases after ca. 80 yr. Soil C densities show slight decrease with ages in Populus stands, but increase in other stand types. These results indicate the complex C‐transfer processes among different components (tree biomass, detritus, forest floor, and soil) of boreal ecosystems at various stages of succession.     

The effect of traditional management burning on lichen diversity

Question: How does regular management burning of a northern, Calluna vulgaris‐dominated heathland affect the lichen diversity at the patch and landscape scale? Location: Mar Lodge Estate, Scottish Highlands, United Kingdom. Methods: 26 fire sites of different ages and 11 long‐term unburnt stands were surveyed to create a chronosequence of changing lichen diversity following burning. Data were analysed graphically, with a GLM and using a CCA. Results: Though the immediate effect of fire was to significantly reduce lichen diversity, it generally recovered within 20 years. There was a significant difference in the population dynamics between wet and dry moorland areas with terricolous lichens in the former site being replaced by pleurocarpous mosses. Older stands, unburnt for 25 years or more, generally had lower diversity than stands 10 to 15 years old. Changes in lichen diversity and community composition can be attributed to the development of Calluna stand structure following burning. Conclusions: Fire can be seen to play an important role in maintaining the diversity of lichens in heathland areas by providing a variety of stand‐structures and ages across the landscape that favours the development of greater beta‐diversity.     

Fire-induced shifts in overstory tree species composition and associated understory plant composition in Glacier National Park,Montana

In Rocky Mountain forests, fire can act as a mechanism of change in plant community composition if postfire conditions favor establishment of species other than those that dominated prefire tree communities. We sampled pre and postfire overstory and postfire understory species following recent (1988–2006) stand-replacing fires in Glacier National Park (GNP), Montana. We identified changes in relative density of tree species and groups of species (xerophytes vs. mesophytes and reseeders vs. resprouters) in early succession. Postfire tree seedling densities were adequate to maintain prefire forest structure, but relative densities among species were variously changed. Changes were directly related to individual species’ response to severe fires. Most notably, relative density of the mesophytic resprouter quaking aspen (Populus tremuloides) and the xerophytic reseeder lodgepole pine (Pinus contorta) increased substantially following fire, with a concomitant decline in proportional abundance of other tree species that, in some cases, dominated stands before fire. Trends identified in our study suggest that forest community shifts toward those dominated by lodgepole pine and quaking aspen are occurring in GNP. Cover of understory species was not affected by tree species composition or density. These forest communities will likely change throughout succession with the addition of shade-intolerant species in early seral stages and shade-tolerant species later in succession. However, with increased fire frequency, the lodgepole pine-dominated postfire communities observed in our study may become more common throughout time.     

Epiphytic lichen synusiae and functional trait groups in boreo‐nemoral deciduous forests are influenced by host tree and environmental factors

Deciduous forests with temperate broad‐leaved tree species are particularily important in terms of biodiversity and its protection, but are threatened habitats in northern Europe. Using multivariate analyses we studied the effect of forest site type, environmental variables and host tree properties on epiphytic lichen synusiae as well as on the composition of species‐specific functional traits. Epiphytic lichens were examined on Acer platanoides, Fraxinus excelsior, Quercus robur, Tilia cordata, Ulmus glabra and U. laevis in two types of forests: Humulus‐type floodplain forests and Lunaria‐type boreo‐nemoral forests on the talus slopes of limestone escarpment (klint forests). Klint forests located near the seashore were under greater maritime influence compared to floodplain forests located in inland Estonia which experience stronger air temperature contrasts. In addition to stand level and climatic variables, tree level factors (bark pH, trunk circumference and cover of bryophytes) considerably affected the species composition of the lichen synusiae. Overall, 137 lichen species were recorded, including 14 red‐listed species characteristic of deciduous trees. We defined 13 lichen societies and showed their preference to forests of a specific site type and/or host tree properties. In forests of both types, most of the epiphytic lichens were crustose, and had apothecia as the fruit bodies and chlorococcoid algae as the photobiont. However, the proportion of lichens with a foliose or fruticose growth form, as well as the proportion of lichens with vegatative diaspores, were higher in floodplain forests. In klint forests with a stronger influence from the wind, crustose species completely dominated, while species with vegetative diaspores were rare and most species dispersed sexually. Lichens with Trentepohlia as the photobiont were characteristic of these forests, and lichens with lirellate ascomata were prevailing, indicating the great uniqueness of the kint forests for epiphytic lichens in the boreo‐nemoral region.     

An Analysis of Vegetation Restoration on Opencast Oil Shale Mines in Estonia

We compared four types of 30‐year‐old forest stands growing on spoil of opencast oil shale mines in Estonia. The stand types were: (1) natural stands formed by spontaneous succession, and plantations of (2) Pinus sylvestris (Scots pine), (3) Betula pendula (silver birch), and (4) Alnus glutinosa (European black alder). In all stands we measured properties of the tree layer (species richness, stand density, and volume of growing stock), understory (density and species richness of shrubs and tree saplings), and ground vegetation (aboveground biomass, species richness, and species diversity). The tree layer was most diverse though sparse in the natural stands. Understory species richness per 100‐m² plot was highest in the natural stand, but total stand richness was equal in the natural and alder stands, which were higher than the birch and pine stands. The understory sapling density was lower than 50 saplings/100 m² in the plantations, while it varied between 50 and 180 saplings/100 m² in the natural stands. Growing stock volume was the least in natural stands and greatest in birch stands. The aboveground biomass of ground vegetation was highest in alder stands and lowest in the pine stands. We can conclude that spontaneous succession promotes establishment of diverse vegetation. In plantations the establishment of diverse ground vegetation depends on planted tree species.     

Competition and regeneration in quaking aspen–white fir (Populus tremuloides–Abies concolor) forests in the Northern Sierra Nevada,USA

Question: How does competition between quaking aspen (Populus tremuloides) and white fir (Abies concolor) affect growth and spatial pattern of each species? Location: The northern Sierra Nevada, California, USA. Methods: In paired plots in mixed aspen‐ (n=3) or white fir‐dominated (n=2) stands, we mapped trees and saplings and recorded DBH, height, species, and condition and took increment cores. We tallied seedlings by species. Tree ring widths were used as a measure of basal area change over the last decade, and canopy openness was identified using hemispherical photographs. Linear mixed models were used to relate neighborhood indices of competition, stand, and tree‐level variables to diameter increment. Spatial patterns of stems were identified using the Neighborhood Density Function. Results: White fir radial growth was higher in aspen‐ than white fir‐dominated plots. Individual‐level variables were more important for white fir than for aspen growth, while variables representing competitive neighborhood were important only for aspen. The forest canopy was more open in aspen‐ than white fir‐dominated stands, but ample aspen seedlings were observed in all stands. Canopy stems of aspen and white fir were randomly distributed, but saplings and small trees were clumped. Aspen saplings were repelled by canopy aspen stems. Conclusions: Variation in canopy openness explained more stand–stand variation in white fir than aspen growth, but high light levels were correlated with recruitment of aspen seedlings to the sapling class. Radial growth of aspen was predicted by indices of neighborhood competition but not radial growth of white fir, indicating that spacing and stem arrangement was more important for aspen than white fir growth. Fire suppression has removed a major disturbance mechanism that promoted aspen persistence and reduced competition from encroaching conifers, and current forests favor species that regenerate best by advance regeneration (white fir).     

Influence of tree age, tree size and crown structure on lichen communities in mature Alpine spruce forests

Testing the relations between tree parameters and the richness and composition of lichen communities in near-natural stands could be a first step to gather information for forest managers interested in conservation and in biodiversity assessment and monitoring. This work aims at evaluating the influence of tree age and age-related parameters on tree-level richness and community composition of lichens on spruce in an Alpine forest. The lichen survey was carried out in four sites used for long-term monitoring. In each site, tree age, diameter at breast height, tree height, the first branch height, and crown projection area were measured for each tree. Trees were stratified into three age classes: (1) <100 years old, immature trees usually not suitable for felling, (2) 100–200 years old, mature trees suitable for felling, and (3) >200 years old, over-mature trees normally rare or absent in managed stands. In each site, seven trees in each age class were selected randomly. Tree age and related parameters proved to influence both tree-level species richness and composition of lichen communities. Species richness increased with tree age and related parameters indicative of tree size. This relation could be interpreted as the result of different joint effects of age per se and tree size with its area-effect. Species turnover is also suspected to improve species richness on over-mature trees. Similarly to species richness, tree-level species composition can be partially explained by tree-related parameters. Species composition changed from young to old trees, several lichens being associated with over-mature trees. This pool of species, including nationally rare lichens, represents a community which is probably poorly developed in managed forests. In accordance to the general aims of near-to-nature forestry, the presence of over-mature trees should be enhanced in the future forest landscape of the Alps especially in protected areas and Natura 2,000 sites, where conservation purposes are explicitly included in the management guidelines.     

Stand Composition,Tree Proximity and Size Have Minimal Effects on Leaf Function of Coexisting Aspen and Subalpine Fir

Forest structural heterogeneity due to species composition, spatial relationships and tree size are widely studied patterns in forest systems, but their impacts on tree function are not as well documented. The objective of this study was to examine how stand composition, tree proximity relationships and tree size influence the leaf functional traits of aspen, an early successional species, and subalpine fir, a climax species. We measured foliar nutrients, nonstructural carbohydrates (aspen only), defense chemistry and xylem water potential of aspen and subalpine fir trees in three size classes growing in close proximity or independently from other trees under three stand conditions: aspen dominant, aspen-conifer mixed, and conifer dominant stands. Close proximity of subalpine fir to aspen reduced aspen’s storage of starch in foliar tissue by 17% suggesting that competition between these species may have small effects on carbon metabolism in aspen leaves. Simple sugar (glucose + sucrose) concentrations in aspen leaves were slightly higher in larger aspen trees than smaller trees. However, no differences were found in stem water potential, foliar concentrations of nitrogen, phosphorus, or secondary defense chemicals of aspen or subalpine fir across the gradients of stand composition, tree proximity or tree size. These results suggest that mechanisms of coexistence allow both aspen and subalpine fir to maintain leaf function across a wide range of stand structural characteristics. For aspen, resource sharing through its clonal root system and high resource storage capacity may partially contribute to its functional stability in mixed aspen-conifer stands.     

Effects of forest management on epiphytic lichen diversity in Mediterranean forests

Question: What are the responses of epiphytic lichens to the intensity of management along a large environmental gradient in Mediterranean Quercus forests? Location: Central Spain. Methods: This study was carried out on 4590 trees located in 306 forest stands dominated by Quercus faginea or Quercus ilex ssp. ballota. The effect of forest management and other predictor variables on several species diversity indicators were studied. Variables modelled were total species richness, cyanolichen richness and community composition. A large number of predictor variables were included: forest fragmentation (patch size, stand variability), climate and topographic (altitude, slope, sun radiation, annual rainfall and mean annual temperature) and intensity of management. General linear models and constrained ordination techniques were used to model community traits and species composition, respectively. Results: Total richness and especially cyanolichens richness were significantly and negatively affected by the intensity of management. Lichen composition was influenced by management intensity, climatic and topographic variables and stand variability. Conclusions: In Mediterranean forests, human activities related to forestry, agricultural and livestock use cause impoverishment of lichen communities, including the local disappearance of the most demanding species. The conservation of unmanaged forests with a dense canopy is crucial for lichen diversity.     

Factors influencing epiphytic bryophyte and lichen species richness at different spatial scales in managed temperate forests

The effect of management related factors on species richness of epiphytic bryophytes and lichens was studied in managed deciduous-coniferous mixed forests in Western-Hungary. At the stand level, the potential explanatory variables were tree species composition, stand structure, microclimate and light conditions, landscape and historical variables; while at tree level host tree species, tree size and light were studied. Species richness of the two epiphyte groups was positively correlated. Both for lichen and bryophyte plot level richness, the composition and diversity of tree species and the abundance of shrub layer were the most influential positive factors. Besides, for bryophytes the presence of large trees, while for lichens amount and heterogeneity of light were important. Tree level richness was mainly determined by host tree species for both groups. For bryophytes oaks, while for lichens oaks and hornbeam turned out the most favourable hosts. Tree size generally increased tree level species richness, except on pine for bryophytes and on hornbeam for lichens. The key variables for epiphytic diversity of the region were directly influenced by recent forest management; historical and landscape variables were not influential. Forest management oriented to the conservation of epiphytes should focus on: (i) the maintenance of tree species diversity in mixed stands; (ii) increment the proportion of deciduous trees (mainly oaks); (iii) conserving large trees within the stands; (iv) providing the presence of shrub and regeneration layer; (v) creating heterogeneous light conditions. For these purposes tree selection and selective cutting management seem more appropriate than shelterwood system.     

The Finnish forest site type approach: ordination and classification studies of mesic forest sites in southern Finland

A.K. Cajander's forest site type classification system is based on definition of plant communities typical to certain climatical and edaphical site conditions, but the structure and composition of the tree stands in Finland are considered sensitive to random variation and are therefore not used as primary classification criteria. The system has often received criticism, usually that the effects of the tree stand and successional stage of the stand have been underestimated. Most of the present-day forest stands in Finland represent young successional stages and are subjected to intensive management. This should result in an additional difficulty in the application of the forest site types in the field.The present study is based on three independent data sets representing forests on mineral soil in southern part of Finland. TWINSPAN classification, DCA ordination and canonical correspondence analysis (CCA) techniques were applied in successive stages of the data analysis. It was found that the definition of the intermediately fertile, mesic site types was clearly confused by the effects of the tree species and age of the stand. The analyses also revealed that the succession pathways on mesic forest sites are largely determined by the tree species composition. In stands dominated by Pinus sylvestris, the succession follows the competitive hierarchy model, whereas in stands dominated by Picea abies, severe shading of the tree canopy governs the development of understorey vegetation.Abbreviations CCA Canonical correspondence Analysis - DCA Detrended correspondence Analysis - TWINSPAN Two-way indicator species analysis