Aspen succession and nitrogen loading: a case for epiphytic lichens as bioindicators in the Rocky Mountains,USA

Question: Can lichen communities be used to assess short‐ and long‐term factors affecting seral quaking aspen (Populus tremuloides) communities at the landscape scale? Location: Bear River Range, within the Rocky Mountains, in northern Utah and southern Idaho, USA. Method: Forty‐seven randomly selected mid‐elevation aspen stands were sampled for lichens and stand conditions. Plots were characterized according to tree species cover, basal area, stand age, bole scarring, tree damage, and presence of lichen species. We also recorded ammonia emissions with passive sensors at 25 urban and agricultural sites throughout an adjacent populated valley upwind of the forest stands. Nonmetric multidimensional scaling (NMS) ordination was used to evaluate an array of 20 variables suspected to influence lichen communities. Results: In NMS, forest succession explained most variance in lichen composition and abundance, although atmospheric nitrogen from local agricultural and urban sources also significantly influenced the lichen communities. Abundance of nitrophilous lichen species decreased with distance from peak ammonia sources and the urban center in all aspen succession classes. One lichen, Phaeophyscia nigricans, was found to be an effective bioindicator of nitrogen loading. Conclusions: Lichen communities in this landscape assessment of aspen forests showed clear responses to long‐term (stand succession) and short‐term (nitrogen deposition) influences. At the same time, several environmental factors (e.g. tree damage and scarring, distance to valley, topography, and stand age) had little influence on these same lichen communities. We recommend further use of epiphytic lichens as bioindicators of dynamic forest conditions.     

Vegetation Composition Determines Microbial Activities in a Boreal Forest Soil

Abstract Forest soil ecology was studied in Fennoscandinavian dry Scots pine forests grazed by reindeer to varying extents (ungrazed, lichen-dominated-sites; grazed sites; and bryophyte-dominated sites). We hypothesized that the productivity parameters of the site (i.e., tree growth and soil nutrient concentrations), the vegetation composition, and the microbial activities are directly correlated. Since the productivity of the lichen-dominated ecosystem is low, microbial activities are assumed to be naturally low. Grazing was expected to decrease both the amount of Scots pine fine roots and the soil microbial activities. Several variables on the characteristics of the soil microbial community, Scots pine fine roots, soil nutrients, and tree growth were studied in relation to vegetation composition by using non-metric multidimensional scaling (NMDS). Basal respiration (Bas), metabolic quotient of the microbial community (qCO2), and pine fine root parameters increased toward the ungrazed, nutrient-poor, lichen-dominated sites, which were grouped at one end of the first axis in the NMDS ordination. Soil nutrient and tree growth parameters and thickness of the humus layer increased toward bryophyte-dominated sites, which were grouped at the other end of the first axis in the ordination. The grazed sites fell between them. These were characterized by lower Bas and qCO2 values and longer lag, compared to ungrazed lichen- or bryophyte-dominated sites, probably due to decreased carbon input and microclimatic change (the soil without lichen carpet is exposed to direct sunlight and wind). Microbial biomass (Cmic), fungal biomass (ergosterol concentration), and the specific growth rate (μCO2) were not related to vegetation ordination. The high fine root production is the most plausible explanation for the high microbial activities at nutrient-poor, lichen-dominated sites, which produce qualitatively poor and slowly decomposing litter, as fine roots secrete considerable amounts of organic substances. At bryophyte-dominated sites, the higher soil nutrient concentrations and the higher production of easily decomposable substrates are likely to maintain the microbial activities.     

Analysis of the vertical distribution of epiphytic lichens on Pinus nigra (Mount Olympos,Greece) along an altitudinal gradient

Epiphytic lichen vegetation was studied in 10 sites along an altitude gradient from 750 to 1510 m on NW-facing slopes of Mount Olympos, Greece to assess the main spatial heterogeneity of microhabitats affecting communities and species composition. Community structure along the gradient was studied by using multivariate techniques. The critical factor for spatial heterogeneity seems to be the height at which the lichen community develops on the tree trunks. Changes in the community structure of the epiphytic lichen vegetation were also detected along the altitudinal gradient and the altitude of 1200 m is considered to be an ecotone. A comparative study of epiphytic lichen communities on Mt. Olympos and in the Thessaloniki area revealed species indicators of air pollution.     

Tree Species Composition Predicts Epiphytic Lichen Communities in an African Montane Rain Forest

The ecology of many tropical rain forest organisms, not the least in Africa, remains poorly understood. Here, we present a detailed ecological study of epiphytic lichens in the equatorial montane rain forest of Bwindi National Park (331 km²), Uganda. We evaluated all major lichen growth forms, including selected groups of crustose lichens. In 14 transects at elevations of 1290 m to 2500 m, we sampled 276 trees belonging to 60 species. We recorded all lichen species on each tree trunk between ground level and 2 m above the ground, yielding 191 lichen species in 67 genera, with a mean of 4.7 species per tree. We used non‐metric multi‐dimensional scaling to separate epiphytic lichen assemblages according to tree species composition and elevation. Structural equation modeling indicated that elevation influenced tree species composition and that tree species composition largely determined lichen species composition. Thus, elevation acted indirectly on the lichen assemblages. Further studies examining factors such as bark properties and lichen colonization ecology may clarify what determines the association between tree species and lichen assemblages. The link between lichen assemblages and large‐scale elevation patterns, as well as disturbance and regrowth histories, warrants further study. An analysis of lichen species composition on individual tree species that occur over large elevation ranges would distinguish the effect of tree species on lichen assemblages from the effect of elevation and thus climate. Our study highlights the limited extent of our knowledge of tropical epiphytic lichens.     

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.     

Vegetative Community Compositional Gradients of Tropical Hardwood Hammocks along the Florida Keys

Tropical hardwood hammocks, among the rarest and most threatened vegetative communities in the United States, occur throughout the 225-km Florida Keys archipelago as it extends toward the Caribbean from the southeast tip of peninsular Florida. Compounding their critical conservation status, tropical hardwood hammocks and the dynamics that support their peculiar species diversity in the region are poorly understood. The goal of this study was to explore the dynamics of the species compositional gradient of the hammocks along the Florida Keys, and to identify significant ecological units within the gradient. The primary data for this research were assembled from the Institute for Regional Conservation's floristic database of South Florida. We were able to extract presence/absence data for 295 species from comprehensive surveys of 23 study sites. Nonmetric multidimensional scaling was used to deconstruct the compositional trends into a reduced ordination space. Cluster analysis was subsequently used to identify discrete ecological units. Additionally, we used vector fitting to interpret the significant correlated ancillary variables. Our main results were three well-fitted nonmetric multidimensional scaling axes with three nonoverlapping ecological units. Of the ancillary variables, latitude, longitude, percent composition from biogeographical regions, richness, and area were correlated to the nonmetric multidimensional scaling results. These results increase our understanding of the community structure of the hammocks along the Florida Keys, and can contribute to increasing our ability to adequately protect and restore tropical hardwood hammocks and other similar tropical dry forest communities throughout the Caribbean.     

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.     

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.     

Isolation affects tree‐scale epiphytic lichen community structure on New Zealand mountain beech trees

Question: Is epiphytic lichen community structure significantly affected by isolation from source community? Location: Foothills of the Southern Alps, South Island, New Zealand. Methods: Epiphytic lichen richness and environmental variables were measured on 382, young Nothofagus solandri var. cliffortioides (Hook. f.) Poole (mountain beech) trees that had recently colonized grassland adjacent to a forest remnant. Richness and the presence of individual lichen taxa were modelled as a function of isolation from the forest fragment, tree size and other habitat conditions. Results: Richness of epiphytic lichen communities was negatively related to tree isolation, although this effect was much smaller than the effects of tree size and other local (tree‐scale) habitat conditions. Different lichen taxa responded in different ways to isolation, area effects and local habitat conditions. Conclusions: This study shows that many epiphytic lichens on mountain beech are limited in their ability to colonize new substrate, even over distances of less than 1 km, which may be due to limitation in dispersal and/or establishment. Lichens are greatly influenced by local habitat conditions, such as tree size, and in this particular environment their negative interaction with sooty moulds is an important driver of community structure.     

Understorey vegetation change in a Picea mariana chronosequence

Eighteen black spruce (Picea mariana) stands, representing postfire ages of 26 to 120 yr, were surveyed for understorey vegetation and site/microsite characteristics at two spatial scales. This enabled comparison of within- versus among-stand compositional variation.Detrended correspondence analysis (DCA) ordination among the 18 stands revealed a complex age/moisture gradient. DCA ordination among 1 800 quadrats within the stands indicated a similar gradient with much compositional overlap. Quadrats were grouped, using two-way indicator species analysis (TWINSPAN), into 9 classes each representing a phase in understorey vegetation composition. These phases shifted in abundance from young to old stands with a high degree of concordance among replicates in the same age class. Understorey succession is strongly linked to the stages in tree growth, mortality and thinning coupled with the accumulation of site moisture.Abbreviations DCA Detrended Corrospondence Analysis     

Report of the Annual Conference of the Cryptogamic Section, 1948

Background: Trait-based assembly rules are a powerful tool in community ecology, used to explore the pattern and process of community structure (richness and composition).

Aims: A preliminary test for the utility of trait-based assembly rules in explaining cryptogamic epiphyte communities (lichens and bryophytes).

Methods: We sampled epiphytes from three different tree species (aspen, birch and pine), and from trees of contrasting age. The community composition of epiphyte species (taxon analysis) and functional groups (trait analysis) was summarised using multivariate ordination (nonmetric multidimensional scaling, NMDS).

Results: Ordination documented a widely observed pattern in which different tree species have taxonomically different epiphyte communities. However, NMDS sample scores were correlated to tree age in the trait-based analysis, but not in the taxon analysis.

Conclusions: Our results point to the existence of a common pattern in community traits during succession (on trees of different age) when measured for epiphyte communities with contrasting taxonomic composition. This pattern is evidenced by consistent trends in lichen growth form and reproductive strategy (sexual vs. asexual).     

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.     

亚热带山地森林附生地衣的移植生长及其对环境变化的响应

本研究选取云南哀牢山亚热带森林系统6种常见的附生大型地衣进行为期2年的移植实验,分析其在原生林林外、林缘以及林内3种生境下的生物量增长速率和健康率等的差异,并探讨其生长对环境因子变化的响应。结果显示,网肺衣Lobaria retigera和平滑牛皮叶Sticta nylanderiana在林缘处生物量增长最快;而槽枝Sulcaria sulcata、多花松萝Usnea florida、皮革肾岛衣Nephromopsis pallescens、针芽肺衣Lobaria isidiophora 4种地衣则都是在林外光照更强、湿度更低的条件下生长得最好,林内最差。除网肺衣外,所有地衣的健康率均在林外较好,林缘次之,林内最差。非参数相关分析表明,6种地衣的生长速率和光照以及温度呈正相关,与大气湿度呈负相关;温度和光照是影响地衣生长最主要的生境因子。     

Environmental drivers of tree community turnover in western Amazonian forests

A more comprehensive understanding of the factors governing tropical tree community turnover at different spatial scales is needed to support land‐management and biodiversity conservation. We used new forest inventory data from 263 permanent plots in the Carnegie Biodiversity‐Biomass Forest Plot Network spanning the eastern Andes to the western Amazonian lowlands of Peru to examine environmental factors driving genus‐level canopy tree compositional variation at regional and landscape scales. Across the full plot network, constrained ordination analysis indicated that all environmental variables together explained 23.8% of the variation in community composition, while soil, topographic, and climatic variables each explained 15.2, 10.9, and 17.0%, respectively. A satellite‐derived metric of cloudiness was the single strongest predictor of community turnover, and constrained ordination revealed a primary gradient of environmentally‐driven community turnover spanning from cloudy, high elevation sites to warm, wet, lowland sites. For three focal landscapes within the region, local environmental variation explained 13.4–30.8% of compositional variation. Community turnover at the landscape scale was strongly driven by topo‐edaphic factors in the two lowland landscapes examined and strongly driven by potential insolation and topography in the montane landscape. At the regional scale, we found that the portion of compositional variation that was uniquely explained by spatial variation was relatively small (2.7%), and was effectively zero within the three focal landscapes. Overall, our results show strong canopy tree compositional turnover in response to environmental gradients at both regional and landscape scales, though the most important environmental drivers differed between scales and among landscapes. Our results also highlight the usefulness of key satellite‐derived environmental covariates that should be considered when conducting biodiversity analyses in tropical forests.     

辽东栎林不同层植物沿海拔梯度分布的DCCA分析

 为研究北京东灵山辽东栎（Quercus liaotungensis）不同层植物的分布与环境因子之间的关系,对东灵山辽东栎林进行了植物群落学调查。沿 海拔梯度在乔木层、灌木层和草本层各调查119个样方,同时调查了每个样方的环境因子如海拔、坡度和坡位。物种数据采用二元属性数据,对 非数值环境因子进行赋值。对东灵山辽东栎林和各环境因子进行去趋势典范对应分析（Detrended canonical correspondence analysis, DCCA）,结果表明：DCCA 第一轴基本上反映了植物群落所处环境的海拔的梯度变化,第二轴基本上反映了坡位的梯度变化。同时发现辽东栎林 大多数植物分布在低海拔区域。乔木层的物种分布比较均匀。草本层的物种分布重叠率是最大的,说明草本植物的生境相似性很高。低海拔区 域的生物多样性要高于高海拔区域的生物多样性。结合环境因子间的相关关系和DCCA排序结果,发现海拔是影响东灵山辽东栎林物种分布的主 要环境因子。     

Species richness and vertical stratification of epiphytic lichens in subtropical primary and secondary forests in southwest China

An entire-tree investigation was conducted in two primary and three secondary forest types in the subtropical Ailao Mountains of southwest China to determine whether species richness and vertical stratification of epiphytic lichens responded to forest type and host attributes. Lichen species number, composition and cover differed significantly among forest types and tree species, while tree diameter and tree height had a modest influence. Epiphytic lichen species and functional groups showed clear vertical stratification. Epiphytic lichens were richer in canopies than on trunks and exhibited a great preference for the intermediate zones of trees, while five lichen groups showed distinct vertical diversification. The stratification patterns are clearly related to forest type and may reflect the microclimatic requirements of individual species, e.g. light availability and humidity.     

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.     

Crustose lichens as indicators of forest continuity in boreal coniferous forests

The crustose lichen flora of fifteen areas in boreal coniferous forests from southern Sweden to Lapland was surveyed. Many of these areas are included in a national programme for monitoring long-term environmental change, the PMK programme. In all 190 species were recorded, 129 of them occurring on bark of conifers and lignum. Some species were found exclusively in forests with a long continuity. By using ordination and hierarchical classifications further species with similar habitat requirements were identified. An Indicator Species Index of Forest Continuity (ISIFC) was designed. Only two of the twenty index species occur on deciduous trees. The ISIFC was highly correlated with forest continuity and also with occurrence of species listed as threatened. Correlations with macroclimatic variables are considered sampling artifacts, and microclimatic factors are suggested to be of primary importance for species occurring in forests with a long continuity.     

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.     

Multiple-scale environmental modulation of lichen reproduction

It is necessary to understand how environmental changes affect plant fitness to predict survival of a species, but this knowledge is scarce for lichens and complicated by their formation of sexual and asexual reproductive structures. Are the presence and number of reproductive structures in Lobaria pulmonaria, a threatened lichen, dependent on thallus size, and is their formation sequential? Does any size-dependence and sequential formation vary along a climate gradient? Generalized linear mixed models were used to explore the effect of environmental predictors on the size and presence/abundance of each reproductive structure and to determine the probability of a given-sized thallus to develop any reproductive structure. The largest individuals are more likely to develop reproductive structures, and the lichen uses a mixed strategy of early asexual reproduction and late sexual. Macro and microclimatic variables also influenced reproductive capacity. Relationships among climate conditions and lichen size and reproductive capacity can compromise the future viability of the species in the most southern populations of Europe.