The importance of host tree age, size and growth rate as determinants of epiphytic lichen diversity in boreal spruce forests

The amount of large and old trees has decreased in the boreal forests during the last centuries of forestry. Such trees are important habitats for epiphytic lichens and there is a growing concern for lichen species that are associated with large and old trees. However, only little is known about the relative importance of tree size versus age as determinants of lichen diversity. Here we have determined the size, age and growth rate of 157 Norway spruce trees and recorded the occurrence of epiphytic lichen species on their branches and lower stems. The study includes crustose lichens and was done in two old-growth forests in SE Norway. Tree age and tree size were the parameters that explained the largest part of epiphytic lichen diversity. Only the growth rate of the most recent time period, i.e. 1984–2004, showed a statistically significant relationship to diversity. There was no indication of a stabilising species number with increasing tree age. Slow-growing and old trees were, however, mainly of importance to the lichen species growing on stems, and this set of species were in general adversely affected by a large amount of branches. The opposite was the case for the species that were confined to branches as their diversity increased when the amount of branches increased. Our study adds empirical data to support the importance of large and old trees as bearers of biodiversity in boreal forests. Site preservation and patch retention of groups of old and large trees is recommended as measures to maintain epiphytic lichen diversity.     

Assessing the effects of forest management on epiphytic lichens in coppiced forests using different indicators

Abstract

Epiphytic lichens are one of the taxonomic groups most sensitive to forest management. Nevertheless, they have not yet been exhaustively included in the assessment of Sustainable Forest Management. This work aimed at evaluating the effects of forest management on epiphytic lichens in coppiced forests, exploring the spatial patterns of diversity and the composition of communities. Moreover, the goal was to compare the performance of four potential indicators for monitoring the effects of forest management on epiphytic lichens: total lichen diversity, species associated with intensive management, species associated with aged coppiced woodlands and Indicator Species Ratio (ISR). In humid Mediterranean Liguria, 50 sampling units were chosen in Castanea sativa and deciduous Quercus spp. forests subjected to different forest management practices: intensively managed coppice and aged coppice/high forest. The effect of forest management was evident in terms of species composition, since it was possible to find significantly associated species for each of the two management types. At each sampling site, the four indicators were calculated using Indicator Value Analysis and compared through correspondence analysis. The ISR was shown to be a more effective indicator, being independent of floristic composition and the occurrence of rare species.     

云南哀牢山原生林及次生林群落附生地衣物种多样性与分布

附生地衣是森林附生植物的重要类群之一, 在维护森林生态系统的物种多样性以及水分和养分循环等方面发挥着重要作用。作者于2005年12月至2006年5月利用树干取样法调查了云南哀牢山徐家坝地区原生山地常绿阔叶林及其次生群落栎类萌生林、滇山杨(Populus bonatii)林和花椒(Zanthoxylum bungeanum)人工林中525株不同种类和径级树木距地面 0–2.0 m处附生地衣的组成和分布, 并收集了各个群落地面上凋落的地衣, 分析了林冠层附生地衣的物种组成。研究结果表明, 该区森林中附生地衣物种比较丰富。共收集到附生地衣61种, 分属17科29属, 其中原生林、栎类萌生林、滇山杨林和花椒人工林分别有51、53、46和23种。在树干距地面 0–2.0 m位置, 各群落中的附生地衣组成明显不同;但在林冠层中, 各群落内的附生地衣基本相似。原生林中附生地衣种类较多, 但分布不均匀。树干附生地衣的Shannon-Wiener和Simpson多样性指数以栎类萌生林最高, 分别为2.71和0.89;花椒林和滇山杨林次之, 分别为2.43–2.45和0.88–0.89;原生林最低, 为1.25和0.67。树干方位、宿主种类和宿主径级等都对附生地衣的物种组成和多样性有着重要影响, 附生地衣更多地出现于树干南向方位, 云南越桔(Vaccinium duclouxii)的附生地衣最为丰富, 胸径5.0–25.0 cm的树木上附生地衣较多。哀牢山山地森林群落中丰富的附生地衣种类及物种多样性在维系本区山地森林生态系统生物多样性格局方面具有重要的作用。     

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.     

Epiphytic and epixylic lichen species diversity in Estonian natural forests

The epiphytic and epixylic lichen flora of natural forests was recorded in different parts of Estonia. Altogether 232 taxa of lichens, lichenicolous fungi, or non-lichenized fungi were recorded, 10 of them listed in the Estonian Red Data Book. We found regional differences in lichen species composition and diversity caused by differences in the forest types. The tree-species-rich boreo-nemoral forests had the most diverse lichen flora, while the boreal forest dominated by coniferous trees or birch had the lowest diversity. The stand age proved to be significant in regard to the number of lichen species in a forest. The most remarkable effect on the diversity of forest lichen species was caused by the presence of Populus tremula. Aspen had the highest number of lichen species on the basal trunk and twigs, and also the highest number of host-specific lichen species.     

Non-dominant trees significantly enhance species richness of epiphytic lichens in subtropical forests of southwest China

Host species has an important influence on the distribution of epiphytic lichens in forests. However, the importance of non-dominant trees in shaping lichen communities has been poorly studied owing to the relative rarity of individuals. The importance of dominant and non-dominant trees for distribution of epiphytic lichens was determined in eight subtropical forests in southwestern China. Dominant trees supported more abundant total and exclusive lichen species only in secondary forests. The occurrence of non-dominant trees promoted lichen diversity within forest types and influenced lichen communities on both tree groups. The effects of total tree species on lichen distribution largely resulted from the presence of non-dominant trees. Dominant and non-dominant trees supported distinct lichen assemblages within forest types, and ordination analyses showed a clear separation. Our study, therefore, reinforces the importance of non-dominant trees for conserving epiphytic lichens, and highlights that lichen assemblages are shaped by both dominant and non-dominant trees.     

Forest edge-interior differentiation in the epiphytic lichen diversity of the forest steppe in the Khangai Mountains,Mongolia

Aims The effects of traditional land use by mobile livestock keepers on biodiversity in forest steppe ecotones are insufficiently studied. Epiphytes are an important part of forest plant diversity. Here we analyze differences in the diversity and composition of the epiphytic lichen vegetation between the edge and the interior of Siberian larch forests in the Khangai Mountains, western Mongolia, which are highly subdivided into patches. We asked whether the epiphytic lichen vegetation at the forest edge differs significantly from that in the interior, whether the edge is inhabited by more nitrophilous species than the interior and whether the density of nomad camps around the forest affects epiphytic lichen diversity.Methods Cover percentages of epiphytic lichen species were recorded from 20 trees per plot on 6 plots in the interior and 6 plots at the edge of Larix sibirica forests. The position of nomad summer camps was surveyed using Global Positioning System. Data were analyzed with pairwise significance tests, analysis of similarities, nonmetric multidimensional scaling and canonical correspondence analysis.Important findings The composition of the epiphytic lichen vegetation clearly differed between the two habitats, with more species being more frequent at the edge than in the interior. However, there was no difference in species richness (α-diversity). The epiphyte vegetation at the edge was more uniform and characterized by lower variation of tree-level α-diversity and lower β-diversity than in the interior. At the edge, only nitrophytic lichens were dominant, whereas in the interior, nitrophytes and acidophytes were among the dominant species. This pattern is probably attributable to the spatial heterogeneity of the intensity of forest grazing and was shown to be influenced by the density of nomad summer camps in the vicinity of the forests. Tree-level α-diversity increased with stem diameter, but high-diameter trees were rare. The results suggest that the present level of forest patchiness and the effect of forest grazing increases the diversity of epiphytic lichens on the landscape level, while logging of high-diameter trees reduces lichen diversity.     

The significance of precipitation and substrate chemistry for epiphytic lichen diversity in spruce-fir forests of the Salish Mountains, northwestern Montana

The relevance of chemical site factors for the abundance of epiphytic lichens was studied in Picea engelmannii-Abies lasiocarpa forests of the Salish Mountains, northwestern Montana, USA. The Salish Mountains are an area with relatively low atmospheric pollutant load and low precipitation. Canonical correspondence analysis (CCA) suggests that cover of several lichen species was limited by high Mn concentrations of bark or by high ratios of Mn to Ca, Mg and Fe. Mn in the bark is known primarily to derive from the soil. An effect of Mn concentration or Mn/Ca and Mn/Mg ratios was not found on A. lasiocarpa. This suggests that A. lasiocarpa deposits Mn in the bark in a physiologically inactive form as already known from A. balsamea. Precipitation chemistry was apparently less relevant for epiphytic lichen distribution in the Salish Mountains, as no correlations between element concentrations in stemflow and cover values were found and as amounts of stemflow were small. However, precipitation in the study year was less than average. The lacking significance of precipitation chemistry is probably the cause why epiphytic lichen vegetation differed less between living and dead trees in the Salish Mountains than in highly polluted coniferous forests studied by our group in Germany; in Germany, the difference between living and dead trees was attributed to reduced interception of pollutants from the atmosphere by trees with reduced crown surface. The result of the present study that small-scale variation of epiphytic lichen abundance is only partly explainable by chemical parameters gives rise to the assumption that microclimate (e.g., moisture), which has not been systematically explored, could be an important site factor for epiphytic lichens in the Salish Mountains. Furthermore, tree age was identified by CCA as a relevant site factor for lichens on P. engelmannii.     

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.     

Wildfire severity reduces richness and alters composition of soil fungal communities in boreal forests of western Canada

Wildfire is the dominant disturbance in boreal forests and fire activity is increasing in these regions. Soil fungal communities are important for plant growth and nutrient cycling postfire but there is little understanding of how fires impact fungal communities across landscapes, fire severity gradients, and stand types in boreal forests. Understanding relationships between fungal community composition, particularly mycorrhizas, and understory plant composition is therefore important in predicting how future fire regimes may affect vegetation. We used an extreme wildfire event in boreal forests of Canada's Northwest Territories to test drivers of fungal communities and assess relationships with plant communities. We sampled soils from 39 plots 1 year after fire and 8 unburned plots. High‐throughput sequencing (MiSeq, ITS) revealed 2,034 fungal operational taxonomic units. We found soil pH and fire severity (proportion soil organic layer combusted), and interactions between these drivers were important for fungal community structure (composition, richness, diversity, functional groups). Where fire severity was low, samples with low pH had higher total fungal, mycorrhizal, and saprotroph richness compared to where severity was high. Increased fire severity caused declines in richness of total fungi, mycorrhizas, and saprotrophs, and declines in diversity of total fungi and mycorrhizas. The importance of stand age (a surrogate for fire return interval) for fungal composition suggests we could detect long‐term successional patterns even after fire. Mycorrhizal and plant community composition, richness, and diversity were weakly but significantly correlated. These weak relationships and the distribution of fungi across plots suggest that the underlying driver of fungal community structure is pH, which is modified by fire severity. This study shows the importance of edaphic factors in determining fungal community structure at large scales, but suggests these patterns are mediated by interactions between fire and forest stand composition.     

亚热带常绿阔叶林附生地衣凋落物的物种多样性和生物量的边缘效应

地衣是亚热带山地森林系统附生生物类群的重要组成部分之一,对环境变化极其敏感。为了更好地了解附生地衣对森林边缘效应的响应,我们在云南哀牢山地区原生山地常绿阔叶林中,分别在距林缘5m、25m、50m和 100m处设立样地,收集附生大型地衣的凋落物1年;分析附生地衣凋落物的物种多样性和生物量、功能群特征和组成结构对林缘深度变化的响应特征。研究结果显示,边缘效应能够显著提高林缘附生地衣群落的物种多样性和生物量;其发生的距离最深不超过25m。林缘-林内梯度上,不同地衣功能群的响应模式具有各自的特征。排序分析表明仅在5m样地与其他样地之间存在显著差异,指示种分析则发现仅5m样地具有指示种。哀牢山原生林中边缘效应促进林缘附生地衣生长和分布的现象,可能与当地高湿环境削减了地衣的高光损伤以及以叶状和灌状类群为主的地衣个体受到风力破坏的程度相对较低有关。     

Tree and stand-scale factors affecting richness and composition of epiphytic bryophytes and lichens in deciduous woodland key habitats

Conservation and sustainable forestry are essential in a multi-functional landscape. In this respect, ecological studies on epiphytes are needed to determine abiotic and biotic factors associated with high diversity. The aim of the present study was to evaluate relative sensitivity of conservation targets (epiphytic bryophytes and lichens) in relation to contrasting environmental variables (tree species, tree diameter at breast height, bark crevice depth, pH, tree inclination, pH, forest stand age, area and type) in boreo-nemoral forests. The study was conducted in Latvian 34 woodland key habitat (WKH) boreo-nemoral forest stands. Generalized linear mixed models and canonical correspondence analysis showed that tree species and tree bark pH were the most important variables explaining epiphytic bryophyte and lichen composition and richness (total, Red-listed, WKH indicator species). Forest stand level factors, such as stand size and habitat type, had only minor influence on epiphytic species composition and richness. The results of the present study indicate a need to maintain the diversity of tree species and large trees, particularly Acer platanoides, Carpinus betulus, Fraxinus excelsior, Populus tremula, Tilia cordata, Ulmus glabra and Ulmus laevis in conservation of epiphytic bryophyte and lichen communities in the future.     

Stand characteristics and biodiversity indicators along the productivity gradient in boreal forests: Defining a critical set of indicators for the monitoring of habitat nature quality

Abstract

We quantified the effects of anthropogenic disturbances on the structure and biodiversity of boreal forests on acidic soils and created a statistically supported rational set of indicators to monitor the stand “naturalness”. For that, we surveyed various traits of tree layer, understory, herb layer, forest floor and several widely accepted biodiversity epiphytic indicators in 252 old‐aged boreal stands in Estonia, mostly dominated by Scots pine or Norway spruce. Multifactorial general linear model analyses showed that many forest characteristics and potential indicators were confounded by the gradient of soil productivity (reflected by the forest site type), local biogeographic gradients and also by stand age. Considering confounding effects, boreal forests in a near‐natural state have more large‐diameter trees (diameter at breast height >40 cm) and larger variety of diameter classes, higher proportion of spruce or deciduous trees, a larger amount of coarse woody debris in various stages, a more closed tree canopy and denser understory than managed mature forests. By increasing light availability above the field layer, forest management indirectly increases the coverage of herbs and lichens on the forest floor but reduces the alpha‐ and beta‐diversity of herbs and the proportion of graminoids. Human disturbances reduce the relative incidence of many commonly accepted biodiversity indicators such as indicator lichens, woodpeckers, wood‐dwelling insects or fungi on trees. The test for the predictive power of characteristics reacting on disturbance revealed that only a fraction of them appeared to be included in a diagnostic easy‐to‐apply set of indicators to assess the nature quality of boreal forest: the amount of dead wood, the proportion of deciduous trees, the presence of specially shaped trees and woodpeckers and, as an indicator of disturbances, the forest herb Melampyrum pratensis. Many of these indicators have already been implemented in practice.     

The ecology and distribution of lichens in tropical deciduous and evergreen forests of northern Thailand

During three years of research on epiphytic lichen communities as indicators of environmental change in northern Thailand plots were set up in a range of forest types between 400 and 1600 m in 1991/2 and revisited in 1993. Other areas were visited in 1993 and collections made in a wider range of geographical, altitudinal and vegetation conditions in Thailand. From this data factors influencing the distribution of lichens in a monsoon climate are outlined and characteristic components of the lichen flora given at family, generic and where possible specific level. Dominant taxa of the montane forests include hygrophilous macrolichens of the 'Lobarion' that are also a characteristic component of old growth fagaceous forests in Europe, whereas the evergreen forests are dominated by moisture-dependent crustose taxa with a trentepohlioid photobiont, and the deciduous dipterocarp forests by often brightly coloured xerophytic lichens with a trebouxioid photobiont. Taxa are proposed as indicators of forest type including those that are indicators of old-growth forests and of disturbance. Quantitative recording of selected taxa at genus and species level is suggested to estimate rates of change in monsoon forests in southeast Asia.     

Post-fire recolonization of dominant epiphytic lichen species on Quercus hypoleucoides (Fagaceae)

Following a forest fire (27 500 ha) in 1994, post-fire recolonization of Quercus hypoeleucoides by epiphytic lichens was documented as changes in lichen cover, number of small thalli, specific factors that affected reestablishment of lichens, and modes of dispersal. Three sites in the Chiricahua Mountains (Arizona, USA) were chosen according to the severity of fire damage—unburned, moderately burned, and severely burned. From 1994 through 1997, the amount of dead lichen cover significantly increased at the moderately burned site. For the same time period, the amount of live lichen cover significantly increased at the severely burned site. Numbers of new thalli increased significantly at the severely burned site each year but only in the last year (1996–1997) for the moderately burned site. Bark texture and proximity to trees with lichens were among the most important physical factors for recolonization. The most important means of dispersal for Flavopunctelia praesignis was fragmentation. For Punctelia hypoleucites, the primary means of dispersal was spores. Increases in live lichen cover and numbers of new thalli occur faster in severely burned areas probably due to the loss of lichens on tree trunks, which provides space and a lack of competition.     

Phosphorus Fertilization Increases the Abundance and Nitrogenase Activity of the Cyanolichen Pseudocyphellaria crocata in Hawaiian Montane Forests

Nitrogen-fixing epiphytes (especially lichens with a cyanobacterial symbiont—cyanolichens) have the potential to contribute significant amounts of nitrogen (N) to montane tropical forests, which are typically low in N—but the factors controlling the abundance and distribution of epiphytic cyanolichens are poorly understood. In long-term fertilization experiments in montane forests on a 4.l million-yr-old Oxisol on the island of Kauà'i and on a 152-yr-old lava flow on the island of Hawai'i, the epiphytic cyanolichen Pseudocyphellaria crocata increased significantly in abundance in canopies of host trees fertilized with phosphorus (P). There was no response to fertilization with N or other essential elements. Nitrogen-fixation rates were also elevated in lichens in P-fertilized plots at both sites. Phosphorus supply to host trees may be an important factor controlling N inputs to montane tropical forests by N-fixing epiphytes.     

Effects of post-fire conditions on soil respiration in boreal forests with special reference to Northeast China forests

Forest fires frequently occur in boreal forests,and their effects on forest ecosystems are often significant in terms of carbon flux related to climate changes.Soil respiration is the second largest carbon flux in boreal forests and the change in soil respiration is not negligible.Environmental factors controlling the soil respiration,for example,soil temperature,are altered by such fires.The abnormal increase in soil temperature has an important negative effect on soil microbes by reducing their activities or even by killing them directly with strong heat.On the other hand,although vegetation is directly disturbed by fires,the indirect changes in soil respiration are followed by changes in root activities and soil microbes.However,there is very limited information on soil respiration in the forests of Northeast China.This review,by combining what is known about fire influence on soil respiration in boreal forests from previous studies of post-fire effects on soil conditions,soil microbes,and forest regeneration,presents possible scenarios of the impact of anticipated post-fire changes in forest soil respiration in Northeast China.     

Edge and land-use effects on epiphytic lichen diversity in the forest-steppe ecotone of the Mongolian Altai

Epiphytic lichen diversity was studied in forests of Siberian larch (Larix sibirica) in the forest-steppe ecotone of the Mongolian Altai. These forests are utilized for livestock grazing, fuelwood collection and occasional logging by pastoral nomads. The density of nomad households in the proximity of the forests influences epiphytic lichen diversity more strongly than the position of sample trees in the forest interior or at the forest line to the steppe. This suggests that land use exerts a stronger effect on lichen diversity than the distinct gradient in microclimate between the forest interior and the forest edge. The co-occurrence of nitrophytes with anitrophytic acidophytes on a small spatial scale as well as higher N and Ca concentrations and pH values in the bark of larch trees at the forest edge than in the forest interior indicate that moderate livestock grazing increases the epiphytic lichen diversity due to an increase of the diversity of chemically different microhabitats. Preference of many lichen species (and of rare species in particular) for overmature and decaying trees suggests that logging and fuelwood collection has adverse effects on epiphytic lichen diversity. This adverse effect is likely to be more crucial for lichen diversity than the putatively positive effect of livestock grazing, since more lichens with a preference for old and decaying trees than for nitrogen-enriched bark were found. The present study is the first one investigating the impact of pastoral nomadism in Central Asia on epiphyte diversity.     

Significant decrease in epiphytic lichen diversity in a remote area in the European Alps,Austria

We present long-term (1993–2010) monitoring results of lichens from a remote site in Austria. Whereas gaseous pollution is negligible in this area, levels of long-range air pollution of S and N via rain and fog are high. Lichen cover on tree trunks have decreased significantly. Accordingly, we found that the population of all species had declined. The analyses demonstrated significant changes in lichen community composition. The lichen flora suffered a significant decrease in diversity. Our hypothesis is that the breakdown of epiphytic lichens is mainly due to eutrophication through long-term N deposition, and more particularly to ammonia in precipitation and in fog. Since many areas in the European Alps receive high wet deposition loads, a decrease in the abundance and diversity of epiphytic lichens is most probably a widespread phenomenon.     

Effects of post-fire conditions on soil respiration in boreal forests with special reference to Northeast China forests

Forest fires frequently occur in boreal forests, and their effects on forest ecosystems are often significant in terms of carbon flux related to climate changes. Soil respiration is the second largest carbon flux in boreal forests and the change in soil respiration is not negligible. Environmental factors controlling the soil respiration, for example, soil temperature, are altered by such fires. The abnormal increase in soil temperature has an important negative effect on soil microbes by reducing their activities or even by killing them directly with strong heat. On the other hand, although vegetation is directly disturbed by fires, the indirect changes in soil respiration are followed by changes in root activities and soil microbes. However, there is very limited information on soil respiration in the forests of Northeast China. This review, by combining what is known about fire influence on soil respiration in boreal forests from previous studies of post-fire effects on soil conditions, soil microbes, and forest regeneration, presents possible scenarios of the impact of anticipated post-fire changes in forest soil respiration in Northeast China.