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.     

Hidden crown jewels: the role of tree crowns for bryophyte and lichen species richness in sycamore maple wooded pastures

Tree crowns typically cover the vast majority of the surface area of trees, but they are rarely considered in diversity surveys of epiphytic bryophytes and lichens, especially in temperate Europe. Usually only stems are sampled. We assessed the number of bryophyte and lichen species on stems and in crowns of 80 solitary sycamore maple trees (Acer pseudoplatanus) at six sites in wooded pastures in the northern Alps. The total number of species detected per tree ranged from 13 to 60 for bryophytes, from 25 to 67 for lichens, and from 42 to 104 for bryophytes and lichens considered together. At the tree level, 29 % of bryophyte and 61 % of lichen species were recorded only in the crown. Considering all sampled trees together, only 4 % of bryophyte, compared to 34 % of lichen species, were never recorded on the stem. Five out of 10 red-listed bryophyte species and 29 out of 39 red-listed lichen species were more frequent in crowns. The species richness detected per tree was unexpectedly high, whereas the proportion of exclusive crown species was similar to studies from forest trees. For bryophytes, in contrast to lichens, sampling several stems can give a good estimation of the species present at a site. However, frequency estimates may be highly biased for lichens and bryophytes if crowns are not considered. Our study demonstrates that tree crowns need to be considered in research on these taxa, especially in biodiversity surveys and in conservation tasks involving lichens and to a lesser degree also bryophytes.     

The effects of stand structure on ground-floor bryophyte assemblages in temperate mixed forests

The effect of tree species composition, stand structure characteristics and substrate availability on ground-floor bryophyte assemblages was studied in mixed deciduous forests of Western Hungary. Species composition, species richness and cover of bryophytes occurring on the soil and logs were analysed as dependent variables. The whole assemblage and functional groups defined on the basis of substrate preference were investigated separately. Substrate availability (open soil, logs) was the most prominent factor in determining species composition, cover and diversity positively, while the litter of deciduous trees had a negative effect on the occurrence of forest floor bryophytes. Besides, bryophyte species richness increased with tree species and stand structural diversity, and for specialist epiphytic and epixylic species log volume was essential. Sapling density and light heterogeneity were influential on bryophyte cover, especially for the dominant terricolous species. Many variables of the forest floor bryophyte community can be estimated efficiently by examining stand structure in the studied region. Selective cutting increasing tree species diversity, stand structural heterogeneity and dead wood volume can maintain higher bryophyte diversity in this region than the shelter-wood system producing even-aged, monodominant, structurally homogenous stands.     

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.     

Tree hollows can affect epiphyte species composition

Tree hollows often harbor animals and microorganisms, thereby storing nutritive resources derived from their biological activities. The outflows from tree hollows can create unique microenvironments, which may affect communities of epiphytic organisms on trunk surfaces below the hollows. In this study, we tested whether the species richness and composition of epiphytic bryophytes (liverworts and mosses) and lichens differ above and below tree hollows of Aria japonica and Cercidiphyllum japonicum in a Japanese temperate forest. The species richness of epiphytic bryophytes and lichens did not differ above and below hollows; however, the species composition of bryophytes differed significantly above and below hollows. Indicator species analyses showed that the moss species Anomodon tristis and the liverwort species Porella vernicosa were significantly more common below than above hollows, while the liverwort species Radula japonica and four lichen species, including Leptogium cyanescens, occurred more frequently above than below hollows. Our results highlight that tree hollows can produce unique microenvironments on trunk surfaces that potentially contribute to the maintenance of epiphytic diversity on a local scale.     

A survey method for assessing the richness of epiphytic lichens using growth forms

The species richness of epiphytic lichens is continuously decreasing by degradation and loss of habitat. Considering that taxonomic identification of all species is time and resource consuming, rapid assessment methods to extrapolate the total number of species are needed for practical conservation. This paper describes an alternative method using the correlation between lichens growth forms and species richness. The study was conducted in 406 forest stands located in Central Spain, covering a wide range of mediterranean-climate ecosystem regions, management intensity levels, canopy cover conditions, and tree sizes. The presence/absence of epiphytic lichens was determined in 6090 trees, which were dominated by oak species (Quercus ilex, Q. faginea, and Q. pyrenaica). In all type of forests, the diversity of growth forms was positively correlated with the total epiphytic lichen richness. In all cases, species richness increased in non-managed forest stands with dense canopies. Thus, we propose the use of lichen growth forms as a helpful surrogate of species richness to detect potentially conservation priority areas in the Mediterranean region.     

Epiphytic lichens and bryophytes of forest ecosystems in Tuscany (Central Italy)

The distribution of epiphytic bryophytes and lichens on three species of trees (Quercus ilex, Quercus cerris and Fagus sylvatica, representing the forest ecosystems of central Italy at low, medium and high elevations, respectively) was examined with respect to host specificity and habitat differences. Although most species were host-specific, the results suggested that habitat characteristics are more important than phorophyte properties in determining the distribution of lichens and bryophytes. Factors affecting the diversity of the epiphytic flora were different for lichens and bryophytes. In the case of lichens, climatic parameters and forest monospecificity were important factors, whereas tree cover (i.e. shade) and to a certain extent the number of available trees seemed more important for bryophytes.     

Development of secondary woodland in oak wood pastures reduces the richness of rare epiphytic lichens

Wooded pastures with ancient trees were formerly abundant throughout Europe, but during the last century, grazing has largely been abandoned often resulting in dense forests. Ancient trees constitute habitat for many declining and threatened species, but the effects of secondary woodland on the biodiversity associated with these trees are largely unknown. We tested for difference in species richness, occurrence, and abundance of a set of nationally and regionally red-listed epiphytic lichens between ancient oaks located in secondary woodland and ancient oaks located in open conditions. We refined the test of the effect of secondary woodland by also including other explanatory variables. Species occurrence and abundance were modelled jointly using overdispersed zero-inflated Poisson models. The richness of the red-listed lichens on ancient oaks in secondary woodland was half of that compared with oaks growing in open conditions. The species-level analyses revealed that this was mainly the result of lower occupancy of two of the study species. The tree-level abundance of one species was also lower in secondary woodland. Potential explanations for this pattern are that the study lichens are adapted to desiccating conditions enhancing their population persistence by low competition or that open, windy conditions enhance their colonisation rate. This means that the development of secondary woodland is a threat to red-listed epiphytic lichens. We therefore suggest that woody vegetation is cleared and grazing resumed in abandoned oak pastures. Importantly, this will also benefit the vitality of the oaks.     

影响广东黑石顶树附生苔藓分布的环境因子

通过对广东省黑石顶自然保护区内5个2500 m2样地内树附生苔藓的调查及有关环境因子的测定,研究了树附生苔藓的分布格局及其与环境因子的关系.树附生苔藓在不同高度的分布存在一定梯度,20 cm高处树附生苔藓的种类数与盖度均大于60 cm及更高处,且其群落优势种的数量组成与后者存在较大差异.不同树种附生苔藓盖度和种数差异较大.基于附生苔藓植物的盖度进行DCA排序及聚类分析将树种分成四组,马尾松(Pinus massoniana)因具有两种特有的网藓(巴西网藓Syrrhopodon prolifer和鞘刺网藓S.armatus)单独一组,福建青冈(Quercus chungii)亦与其余各种的差异均较大,形成一组,其余阔叶树种根据其所处的森林类型分成两组,针阔叶混交林内的阔叶树种和次生阔叶林内的阔叶树种各形成一组.对环境因子及树皮含水量和pH的分析显示,垂直梯度上空气湿度的差异可能是造成附生苔藓在不同高度分布差异的主要影响因子之一,不同树种附生苔藓的差异在一定程度上受树皮pH的影响,而与树皮含水量无关.同一树种上树附生苔藓的分布又在一定程度上受森林类型的影响.     

宝天曼地区栓皮栎林恢复过程中高等植物物种多样性变化

对宝天曼地区不同恢复阶段栓皮栎林的高等植物物种多样性特征进行了初步分析。结果表明,各层物种丰富度和多样性指数在不同恢复阶段栓皮栎林表现出草本层＞灌木层＞乔木层的趋势。乔木层物种丰富度和多样性指数基本上随恢复时间的增加而增加;灌木层在未破坏的栓皮栎林内较低,而在其他恢复阶段的栓皮栎林内基本相同;草本层在恢复５年、１５年和２５年的栓皮栎林中较高,而在恢复４５年和未破坏的栓皮栎林内较低。乔木层物种均匀度指数在未破坏的栓皮栎林中较高,在恢复２５年的栓皮栎林中较低;灌木层在恢复４５年的栓皮栎林中较高,在恢复１５年的栓皮栎林较低;草本层在恢复５年的栓皮栎林中较高,在恢复１５年的栓皮栎林中较低。不同恢复阶段栓皮栎林各层次间的物种多样性差异大多不显著,只有乔木层和草本层及灌木层和草本层之间的物种丰富度指数有显著差异。不同恢复阶段栓皮栎林在乔木层物种多样性特征上的差异最大,在灌木层物种多样性特征上有一定差异,在草本层物种多样性特征上没有明显的差异。     

Topographic and forest-stand variables determining epiphytic lichen diversity in the primeval beech forest in the Ukrainian Carpathians

The Uholka-Shyrokyi Luh area of the Carpathian Biosphere Reserve is considered the largest and the most valuable primeval beech forest in Europe for biodiversity conservation. To study the impact of different topographic and forest-stand variables on epiphytic lichen diversity a total of 294 systematically distributed sampling plots were surveyed and 198 epiphytic lichen species recorded in this forest landscape, which has an uneven-aged structure. The obtained data were analysed using a non-metric multidimensional ordination and a generalized linear model. The epiphytic lichen species density at the plot level was mainly influenced by altitude and forest-stand variables. These variables are related to both the light availability i.e. canopy closure, and the habitat diversity, i.e. the developmental stage of the forest stands and the mean stem diameter. We found that lichen species density on plots with a relatively open canopy was significantly higher than on plots with a fairly loose or closed canopy structure. The late developmental stage of forest stands, which is characterized by a large number of old trees with rough and creviced bark, had a strong positive effect on lichen species density. In the Uholka-Shyrokyi Luh primeval forest the mean stem diameter of beech trees significantly correlated with lichen species density per plot. Similar trends in the species diversity of nationally red-listed lichens were revealed. Epiphytic lichens with a high conservation value nationally and internationally were found to be rather abundant in the Uholka-Shyrokyi Luh area, which shows its international importance for the conservation of forest-bound lichens.     

Epiphytes of Sitka spruce (<Emphasis Type="Italic">Picea sitchensis</Emphasis>) plantations in Ireland and the effects of open spaces

The epiphytes of the trunks and branches of mature Sitka spruce (Picea sitchensis) trees were studied in twelve plantations containing open spaces (glades, rides and roads) in the east and southwest of Ireland. A pair of trees was studied at each site: one tree at the south-facing edge of an open space and one in the forest interior. Spruce trees were found to support a moderately diverse range of bryophytes and lichens, including two relatively rare bryophyte species. Clear patterns in vertical distribution were identified, with bryophyte richness and cover decreasing and lichen richness and cover increasing from the tree base to the upper trunk. The open spaces themselves did not appear to affect overall epiphyte diversity, with no significant differences in any of the diversity measures between edge and interior trees. The main effect of open spaces was on the epiphyte cover of the edge trees. This was related to increased light levels combined with the presence of live branches from close to ground level on the south sides of the edge trees, which produced optimum conditions for bryophytes at the tree base and lichens in the upper plots. However, this dense side-canopy negatively affected epiphyte diversity on the north sides of the edge trees. Further research is required to assess the effects of open spaces within forestry plantations on epiphyte diversity.     

Epiphytes of Sitka spruce (<Emphasis Type="Italic">Picea sitchensis</Emphasis>) plantations in Ireland and the effects of open spaces

The epiphytes of the trunks and branches of mature Sitka spruce (Picea sitchensis) trees were studied in twelve plantations containing open spaces (glades, rides and roads) in the east and southwest of Ireland. A pair of trees was studied at each site: one tree at the south-facing edge of an open space and one in the forest interior. Spruce trees were found to support a moderately diverse range of bryophytes and lichens, including two relatively rare bryophyte species. Clear patterns in vertical distribution were identified, with bryophyte richness and cover decreasing and lichen richness and cover increasing from the tree base to the upper trunk. The open spaces themselves did not appear to affect overall epiphyte diversity, with no significant differences in any of the diversity measures between edge and interior trees. The main effect of open spaces was on the epiphyte cover of the edge trees. This was related to increased light levels combined with the presence of live branches from close to ground level on the south sides of the edge trees, which produced optimum conditions for bryophytes at the tree base and lichens in the upper plots. However, this dense side-canopy negatively affected epiphyte diversity on the north sides of the edge trees. Further research is required to assess the effects of open spaces within forestry plantations on epiphyte diversity.     

Environmental and historical effects on lichen diversity in managed and unmanaged wooded meadows

Question: Which environmental and historical variables affect epiphytic lichen diversity in managed and unmanaged wooded meadows? Location: The island of Gotland located in the Baltic Sea east of the Swedish mainland. Methods: We examined total epiphytic lichen diversity (crustose, foliose and fruticose species) on 1148 trees in eight grazed, eight traditionally managed (mowing, hay gathering and pollarding) and seven unmanaged wooded meadows. In addition to management, data on site location, habitat structure, history and adjacent habitat were analysed. Results: Lichen species richness increased with wooded meadow area and was greater within managed sites than in unmanaged meadows. Historic crown cover (ca. 1930) also influenced present‐day lichen richness. Geographic location, distance to sea, wooded meadow area and average tree circumference were important determinants of lichen species composition. Tree circumference was the strongest overall predictor of the number of species on individual trees. However, tree circumference interacted with management regime, whereby cessation of management appeared to reduce species richness most on large trees. Traditionally managed sites, on average, supported the greatest richness of red‐listed species. Conclusions: Management regime, wooded meadow area and canopy cover were the main drivers of lichen species richness, while geographic location, wooded meadow area, distance to sea and average tree circumference were important determinants of species composition.     

Protective management of trees against debarking by deer negatively impacts bryophyte diversity

When wildlife populations become too large, they impact other flora and fauna within the ecosystems that they inhabit. For example, the recent rise in population numbers of sika deer in Japan has led to the stripping of bark from tree overstories in forested areas. This has led to protective management actions, such as wrapping the trunks of trees in wire mesh. The present study investigates the impact of this management action on epiphytic diversity at Mt. Ohdaigahara, which is one of the hotspots for bryophyte diversity in Japan. The correlation between the diversity of epiphytic bryophytes and environmental variables was examined, including the presence/absence of wire mesh protection. A generalized linear model showed that species richness and bryophyte cover was significantly correlated with both tree diameter (at 1.5 m height) and tree density (P < 0.01), but negatively correlated with wire mesh protection. Inductively coupled plasma-mass spectrometry analysis showed a significant 3- to 6-fold higher concentration of zinc in bryophytes occupying tree bark under wire mesh protection than for those without wire mesh. Hence, the high sensitivity of bryophytes to zinc accumulation, as a result of toxicity caused by galvanized iron mesh, has led to the loss of species richness and bryophyte cover on tree trunks. Furthermore, other heavy metals found in wire mesh may also contribute to the negative effect on bryophytes. Therefore, to establish best practices for biodiversity conservation that include bryophytes, materials that are free of heavy metals should be preferentially used for tree protection.     

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.     

Up in the Tree – The Overlooked Richness of Bryophytes and Lichens in Tree Crowns

Assessing diversity is among the major tasks in ecology and conservation science. In ecological and conservation studies, epiphytic cryptogams are usually sampled up to accessible heights in forests. Thus, their diversity, especially of canopy specialists, likely is underestimated. If the proportion of those species differs among forest types, plot-based diversity assessments are biased and may result in misleading conservation recommendations. We sampled bryophytes and lichens in 30 forest plots of 20 m × 20 m in three German regions, considering all substrates, and including epiphytic litter fall. First, the sampling of epiphytic species was restricted to the lower 2 m of trees and shrubs. Then, on one representative tree per plot, we additionally recorded epiphytic species in the crown, using tree climbing techniques. Per tree, on average 54% of lichen and 20% of bryophyte species were overlooked if the crown was not been included. After sampling all substrates per plot, including the bark of all shrubs and trees, still 38% of the lichen and 4% of the bryophyte species were overlooked if the tree crown of the sampled tree was not included. The number of overlooked lichen species varied strongly among regions. Furthermore, the number of overlooked bryophyte and lichen species per plot was higher in European beech than in coniferous stands and increased with increasing diameter at breast height of the sampled tree. Thus, our results indicate a bias of comparative studies which might have led to misleading conservation recommendations of plot-based diversity assessments.     

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.     

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.     

Bryophyte and lichen diversity: A comparative study

Abstract We describe the regional species richness, variation in species richness and species turnover of bryophytes and lichens from 36 sites in lowland forests of southeastern Australia. The analyses subdivided the two major taxa into their constituent sub-groups: mosses, liverworts, and crustose, fruticose and foliose lichens. They also explored correlations between selected environmental variables and patterns of diversity. On a regional scale, there were 77 species of bryophytes and 69 species of lichens, giving a total of approximately one-third of the total number of vascular plant species in the region. Mean species richness was higher for lichens than bryophytes. Also, the two taxa were negatively correlated because lichens favoured dry sites and bryophytes favoured moist ones. Species turnover was greater for bryophytes than lichens, largely due to the distribution of liverwort species. Foliose lichens showed higher levels of turnover than crustose lichens. Multiple regression and canonical correspondence analysis showed that both taxa and all sub-groups responded to the same three variables: vascular plant cover, time since last fire and topographic position. Other variables, including time since logging and intensity of logging, explained little variation in bryophyte or lichen diversity. The data suggest that the strategies for the conservation of bryophyte and lichen biodiversity will be different, to reflect the different patterns of species richness and species turnover.