Old trees contribute bio-available nitrogen through canopy bryophytes

Symbiotic cyanobacteria??bryophyte associations on the forest floor are shown to contribute significantly to stand-level nitrogen budgets through the process of biological nitrogen fixation (BNF), but few studies have considered the role of canopy bryophytes. Given the high biomass of epiphytic bryophytes in many tree species of the North American temperate rain forest, we suggest that canopy bryophytes may contribute substantially to stand-level N dynamics. We confirm the presence of cyanobacteria and measure rates of BNF at three heights (0, 15 and 30 m) in Sitka spruce trees across three watershed estuaries of Clayoquot Sound, British Columbia, Canada. This study is the first to report BNF by cyanobacteria associated with epiphytic and forest floor bryophytes in the coastal temperate rain forest of North America. Cyanobacteria density was significantly greater in epiphytic bryophytes compared to mosses on the forest floor, and rates of BNF were highest at 30 m in the canopy. The majority of total stand-level BNF (0.76 kg N · ha^-1 · yr^-1) occurs in the canopy, rather than on the forest floor (0.26 kg N · ha^-1 · yr^-1). We suggest that BNF by cyanobacterial-bryophyte associations in the canopy of coastal temperate rain forests is a unique source of ecosystem N, which is dependent on large, old trees with high epiphytic bryophyte biomass.     

Response of epiphytic bryophytes to simulated N deposition in a subtropical montane cloud forest in southwestern China

A field manipulation experiment was conducted in a subtropical montane cloud forest in southwestern China to determine the possible responses of epiphytic bryophytes to increasing nitrogen (N) deposition from community to physiology level, and to find sensitive epiphytic bryophytes that may be used as indicators for assessing the degree of N pollution. N addition had significantly negative effects on species richness and cover of the epiphytic bryophyte community. Harmful effects of high N loads were recorded for chlorophyll, growth, and vitality of the species tested. The decline of some epiphytic bryophytes may result from detrimental effects on degradation to photosynthetic pigments. Bazzania himalayana (Mitt.) Schiffn., Bazzania ovistipula (Steph.) Mizut., and Homaliodendron flabellatum (Sm.) Fleisch. are candidates in atmospheric nitrogen monitoring. Epiphytic bryophytes in the montane cloud forest are very sensitive to increasing N deposition and often difficult to recover once they have been destroyed, providing early detection of enhanced N pollution for trees or even the whole forest ecosystem. The inference that increasing N pollution may lead to loss of biodiversity is a concern to the developing economy in western China, and should alert the government to the adverse impacts caused by increased industrial pollution during the process of China’s West Development.     

Species richness of vascular plants, bryophytes and lichens in dry grasslands: The effects of environment, landscape structure and competition

We studied the relative importance of local habitat conditions and landscape structure for species richness of vascular plants, bryophytes and lichens in dry grasslands on the Baltic island of Öland (Sweden). In addition, we tested whether relationships between species richness and vegetation cover indicate that competition within and between the studied taxonomic groups is important. We recorded species numbers of vascular plants, bryophytes and lichens in 4 m² plots (n=452), distributed over dry grassland patches differing in size and degree of isolation. Structural and environmental data were collected for each plot. We tested effects of local environmental conditions, landscape structure and vegetation cover on species richness using generalized linear mixed models. Different environmental variables explained species richness of vascular plants, bryophytes and lichens. Environmental effects, particularly soil pH, were more important than landscape structure. Interaction effects of soil pH with other environmental variables were significant in vascular plants. Plot heterogeneity enhanced species richness. Size and degree of isolation of dry grassland patches significantly affected bryophyte and lichen species richness, but not that of vascular plants. We observed negative relationships between bryophyte and lichen species richness and the cover of vascular plants. To conclude, effects of single environmental variables on species richness depend both on the taxonomic group and on the combination of environmental factors on a whole. Dispersal limitation in bryophytes and lichens confined to dry grasslands may be more common than is often assumed. Our study further suggests that competition between vascular plants and cryptogams is rather asymmetric.     

The impact of land abandonment on the plant diversity of olive groves

We investigated the effect of woody species’ encroachment on plant diversity changes with regard to vascular plants and bryophytes in traditional olive groves of the Maremma Regional Park (Tuscany, Italy) and assessed cross-taxon correlation between these two taxa. We classified the olive groves into four land use types, representing different successional stages. To describe the evenness of species distribution within a community, we plotted rank-abundance diagrams for each taxon and each land use type. The relationship between the number and cover of vascular plants, therophytes, bryophytes, colonists and phanerophytes in each plot was examined using linear regression. The effects of land use type on vascular plant and bryophyte richness and assemblages were assessed by permutational uni- and multivariate analysis of variance. The congruence in species composition between the two taxa was evaluated using Procrustes analysis. The number of vascular plants, bryophytes and therophytes decreased linearly with increased phanerophyte species cover. The number of species belonging to Thero-Brachypodietea progressively decreased throughout succession. Rank-abundance diagrams and multivariate analysis showed differences between the land use types, which were statistically significant for vascular plants between the traditional olive groves and the other land use types, and for bryophytes between the traditional olive groves and woodlands. PROTEST analysis and NMDS graphs showed a correlation between vascular plant and bryophyte communities. The results suggested that conservation measures are needed in the study area in order to ensure both the maintenance of traditional olive groves of conservation interest and high levels of environmental heterogeneity.     

Effects of management and adjacent forest on the heathland bryophyte layer

Heathlands are severely threatened by decreasing habitat quantity and quality. In these habitats, bryophytes form an important component of plant diversity. Therefore, the aims of this study were to investigate the effects of management and adjacent forest land use on the heathland bryophyte layer. Furthermore, effects of these factors on invasive bryophyte species were studied.

Bryophyte communities of 11 dry heathland patches bordering forest were surveyed. In these heathlands, management turned out to be an important factor explaining community composition of bryophytes. Furthermore, significant effects of adjacent land use on bryophyte community composition and species richness were found. However, these effects were rather limited in extent, occurring only close to habitat borders, i.e., up to maximum 5 m into the heathland patch. Also for individual bryophyte species, effects of adjacent forest could only be observed within maximum 5 m from the edge. One remarkable edge effect was the increased dominance of the invasive species Campylopus introflexus near the forest edge, especially at grazed sites. Effects of adjacent land use seemed to predominate management effects. Consequently, management cannot serve as a tool to mitigate edge effects on bryophyte species in heathland patches.     

浙江西天目山主要森林类型的苔藓多样性比较

苔藓是森林的重要组分, 是森林保护区的重要保护对象, 在物种资源和生态系统功能维护中有重要作用。该研究以浙江西天目山国家自然保护区内7种主要森林类型(落叶矮林、落叶阔叶林、常绿-落叶阔叶混交林、常绿阔叶林、针阔混交林、针叶林和竹林)内的苔藓植物为对象, 调查了32个10 m × 10 m的样地, 记录地面生苔藓植物盖度和树附生苔藓植物多度, 采用重要值、相似性系数、多样性指数分析了森林类型间的苔藓植物多样性差异。共采集969份标本, 隶属41科84属142种, 其中苔类植物13科18属33种, 藓类植物28科66属109种, 优势科为灰藓科、青藓科和羽藓科。2种混交林(常绿-落叶阔叶混交林和针阔混交林)的物种丰富度和多样性指数均高于其余5种森林, 其中物种丰富度以针阔混交林最高, 苔藓植物多样性则以常绿-落叶阔叶混交林最高, 竹林两者均为最低。海拔等环境因子较为接近的植被类型的苔藓植物多样性相似性较高, 常绿阔叶林与针叶林相似性最高, 而落叶矮林和竹林相似性最小。     

辽东山地古石河冰缘地貌森林生态系统苔藓物种多样性研究

基于9个20 m×30 m森林群落样地的调查数据,采用物种丰富度、α和β多样性指数,对辽东山地古石河冰缘地貌不同林型石生、树生苔藓植物物种多样性进行定量研究,采用皮尔逊相关分析方法对其影响因素进行分析。结果显示,古石河冰缘地貌苔藓植物共有26科46属59种;不同林型石生、树生苔藓植物物种丰富度和α多样性指数均为:暗针叶林针阔混交林落叶阔叶林;石生苔藓植物β多样性指数最高为落叶阔叶林-针阔混交林间(0.44),最低为落叶阔叶林-暗针叶林间(0.33);树生苔藓植物β多样性指数最高为针阔混交林-暗针叶林间(0.40),最低为落叶阔叶林-暗针叶林间(0.25);分析表明,林冠层郁闭度、海拔高度是影响辽东山地古石河冰缘地貌森林生态系统苔藓物种多样性的重要因子。     

Indirect effects of hiking trails on the community structure and diversity of trunk-epiphytic bryophytes in an old-growth fir forest

Abstract

Forest hiking trails may influence local microclimate and biodiversity, but the effects on community structure and diversity of epiphytic bryophytes on trees are currently unknown. Epiphytic bryophytes on 82 Abies faxoniana Rehder & Wilson tree trunks (41 along the hiking trail edge and 41 controls in the forest interior) were investigated at four heights from the ground (10, 50, 120, and 180 cm). At each site, air temperature and humidity were monitored for 1 year. The light radiation levels and air temperature were higher, and the canopy leaf area index and air humidity lower at the trail edge, indicating deterioration in microclimate, resulting from the trail establishment. The epiphytic bryophyte species richness, community cover, and mean cover of dendroid and pendent growth forms on trunks were significantly lower at the trail edge than the control site, suggesting that trail construction caused these reductions. One marked effect of the presence of the trail was the increase in some sun-loving species and decrease in shade-tolerant species. Moreover, the trail also slightly influenced species richness and epiphytic bryophyte cover at both community and species population levels along the height gradient. Comprehensive analyses showed that microclimate deterioration was mainly driven by the trail establishment, and that the change in micro-climate along the trail, rather than any host traits, played an important role in the declining epiphytic bryophyte community structure and diversity at the trail edge, confirming the initial hypothesis that the presence of a raised boardwalk (hiking trail) indirectly influences epiphytic bryophyte community and diversity by altering the microclimate.     

Impact of Land-Use Intensity and Productivity on Bryophyte Diversity in Agricultural Grasslands

While bryophytes greatly contribute to plant diversity of semi-natural grasslands, little is known about the relationships between land-use intensity, productivity, and bryophyte diversity in these habitats. We recorded vascular plant and bryophyte vegetation in 85 agricultural used grasslands in two regions in northern and central Germany and gathered information on land-use intensity. To assess grassland productivity, we harvested aboveground vascular plant biomass and analyzed nutrient concentrations of N, P, K, Ca and Mg. Further we calculated mean Ellenberg indicator values of vascular plant vegetation. We tested for effects of land-use intensity and productivity on total bryophyte species richness and on the species richness of acrocarpous (small & erect) and pleurocarpous (creeping, including liverworts) growth forms separately. Bryophyte species were found in almost all studied grasslands, but species richness differed considerably between study regions in northern Germany (2.8 species per 16 m²) and central Germany (6.4 species per 16 m²) due environmental differences as well as land-use history. Increased fertilizer application, coinciding with high mowing frequency, reduced bryophyte species richness significantly. Accordingly, productivity estimates such as plant biomass and nitrogen concentration were strongly negatively related to bryophyte species richness, although productivity decreased only pleurocarpous species. Ellenberg indicator values for nutrients proved to be useful indicators of species richness and productivity. In conclusion, bryophyte composition was strongly dependent on productivity, with smaller bryophytes that were likely negatively affected by greater competition for light. Intensive land-use, however, can also indirectly decrease bryophyte species richness by promoting grassland productivity. Thus, increasing productivity is likely to cause a loss of bryophyte species and a decrease in species diversity.     

Change in the bryophyte diversity and species composition of Central European temperate broad-leaved forests since the late nineteenth century

The diversity and species composition of the bryophyte flora colonizing the forest floor, live trees and deadwood in semi-natural broad-leaved forests of northern Germany was compared between a recent survey in 2013 and historical records from around 1900. The survey was based on the comparison of presence/absence data and their interpretation with help of ecological indicator values. Total species richness has declined in bryophytes growing on the forest floor but not in the other guilds, whereas a dramatic species turnover was found for all three guilds. This turnover was apparently primarily driven by the increased atmospheric load of reactive nitrogen from anthropogenic emissions. Sensitive species were replaced by more eutrophication-tolerant bryophytes; liverworts are overrepresented among the sensitive and thus declining species. Promotion of the competitive strength of vascular plants due to increased nitrogen levels is the likely cause of the reduction of bryophyte species richness on the forest floor. Former acidification by high atmospheric sulfur dioxide loads has left an imprint in the bryophyte vegetation by having favored acidophytic species and discriminating against basiphytic species. An increase in the mean indicator value for temperature suggests a beginning effect of climate warming on the bryophyte vegetation. Change in forest structure had an apparently smaller imprint on the bryophyte diversity of the studied semi-natural forests than atmospheric chemistry and climate. In this respect, bryophytes differ from the ecologically similar lichens, where published studies from the same region showed a dramatic decline of species richness and a stronger susceptibility to forest management.     

Bryophytes and their microhabitats in coniferous forest pastures: should they be considered in the pasture management?

Forest pastures, like many other semi-natural (traditional) rural biotopes, have undergone a drastic decline both in area and quality during the last century in many areas. We explored the bryophyte flora of Finnish coniferous forest pastures on acidic soil and aimed to recognize the most important microsites (rocks, coarse woody debris, tree bases, mineral soil patches and closed vegetation) for bryophyte diversity. The effects of microhabitat heterogeneity (microsite entropy) on bryophytes was also examined. We found altogether 83 bryophyte species. The only red-listed species, Tayloria tenuis, was frequently found on dung patches and a few rare ruderal species grew exclusively on bare mineral soil. Rocks comprised the most species rich microsite and many common forest floor species showed preference for this microsite. Microhabitat heterogeneity explained bryophyte species richness on both alpha (plot average) and gamma (pasture total) scales. The results suggest that certain individual bryophyte species and their microsites should be taken into account in the management of this biotope, rather than guiding the management solely on the basis of the overall bryophyte diversity.     

赤水河上游主要森林植被中苔藓物种多样性研究

对赤水河上游典型森林植被中苔藓植物多样性调查分析。(1)苔藓植物有15科19属28种(含变种、亚种);生活型有3种：交织型(64.29%)、矮丛集型(28.57%)和平铺型(7.14%)。(2)丰富度以竹叶林中最大(2.131),残存阔叶林中次之(0.687),针叶林中最小(-1.444)。(3)α多样性呈现一定的规律,竹叶林（3.170）＞残存阔叶林（3.000）＞人工植被地（2.807）＞灌木林（2.322）＞针叶林=谷顶区(1.585),α多样性的排序规律与丰富度基本保持一致,均出现在竹叶林中最大。(4)β多样性出现灌木林—残存阔叶林生境中最高（1.000）,竹叶林—残存阔叶林中次之（0.938）,灌木林—针叶林（0.857）中最小的变化差异趋势。分析得出了植被型、湿度和荫蔽度是制约着该地区苔藓植物多样性的主要生态因素。通过苔藓植物多样性的探究,为保护该流域生态环境和生物多样性提供参考价值。     

Bryophytes as indicators of ancient woodlands in Schleswig-Holstein (Northern Germany)

Ancient woodlands, with their long ecological continuity, frequently harbor a high number of typical, rare and threatened species, and are therefore of particular importance for nature conservation. To pinpoint these habitats, a common application is the use of plants as “ancient woodland indicators”. The occurrence of these particular species allows for evaluating the continuity of woodland cover in time. While lists of ancient woodland vascular plants have been derived for many regions, the identification and use of bryophytes as ancient woodland indicators has been widely neglected. This is a bit surprising because certain woodland bryophytes are very sensitive to varying environmental conditions or changes in land management. It therefore appeared promising to compile an ecologically grounded list of ancient woodland indicator bryophytes for practical use.In this study, we present a set of ancient woodland indicator bryophytes based on the analysis of datasets from the North German federal state of Schleswig-Holstein. To compile this list, we systematically evaluated the bryophyte distribution data from floristic surveys in relation to ancient woodland cover data from state-wide inventories. In this way, we were able to determine ancient woodland bryophytes using consistent and repeatable statistical methods.The presented list of 31 ancient woodland indicator bryophytes is ecologically sound and corresponds well with data from the sparse literature. We could distinguish two groups of ancient woodland indicator bryophytes. The first group is linked to base-rich, semi-natural deciduous woodlands with high soil and air humidity. The second group comprises acidophilic bryophytes that occur not only in acidic beech and oak woods, but also in acidic mixed or coniferous forests on ancient woodland sites. Apart from the ancient woodland indicator bryophytes, we could identify one group of recent woodland bryophytes and four groups of bryophytes that are more or less indifferent with respect to woodland continuity.Finally, we provide recommendations for the application of ancient woodland indicator bryophytes in nature conservation practice. Management suggestions for the conservation of the typical bryophyte diversity of ancient semi-natural woodlands are also given.     

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.     

广东黑石顶森林苔藓群落特征初探

通过对5个面积为2500m2的样地中树附生和地面苔藓植物的定量调查,初步研究了广东黑石顶森林中苔藓植物多样性特征。本次调查共发现苔藓植物50种,包括30种苔类和20种藓类。样地中树附生苔藓植物40种,地面苔藓植物24种,前者远高于后者。各样地中地面苔藓植物的种类为8-13种,盖度0.59%-1.12%,坡度及小生境的多样性对地面苔藓植物物种多样性及盖度均有很大影响;树附生苔藓植物12-20种,盖度0.63%-1.63%。结果显示30a的次生林中苔藓植物的物种丰富度及盖度均可恢复到与成熟阔叶林接近的水平。相似性分析表明,成熟阔叶林内苔藓植物种类组成与针阔叶混交林及30a的次生阔叶林差异不明显,但前者的苔藓植物群落结构与后者的差异较大。     

北京百花山自然保护区不同植被地面生苔藓植物物种多样性

采用相似性系数和物种多样性指数对北京百花山自然保护区8种植被内地面生苔藓植物的物种多样性进行了研究,得知8种植被中地面生苔藓植物有13科31属65种（含种以下的单位）,优势科6科,优势种12种。不同植被中苔藓植物的物种组成、优势科和优势种不同,丛藓科和真藓科植物在草甸和灌丛内占绝对优势,而森林植被内的绢藓科为绝对优势科;油松林和落叶阔叶混交林的物种相似性最高为51.63,草甸和荆条灌丛与大部分植被的物种相似为0。地面生苔藓植物的分布与其生长的基质相关性最大,另外水分条件、小生境、草本层盖度和凋落物盖度等也影响地面生苔藓植物的分布。白桦林和落叶阔叶混交林应成为生物多样性保护的重点植被类型。     

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.     

Nutrient status of black spruce (Picea mariana [Mill.] BSP) forest soils dominated by Kalmia angustifolia L.

A study was conducted to determine soil chemistry in an uncut black spruce (Picea mariana) forest with and without the ericaceous understory shrub Kalmia angustifolia, as well as on a cut black spruce forest currently dominated by Kalmia. The organic (humus) and mineral (Ae, upper and lower B horizons) soils associated with Kalmia from uncut and cut forests, and non-Kalmia soils from uncut forest, were analyzed for selected soil properties. In general, mineral soils (B horizon) associated with Kalmia in uncut forest have lower values for organic matter (3.25%), organic nitrogen (0.66 mg·g⁻¹), Fe³⁺ (95.4 μg·g⁻¹) and Mn²⁺ (9 μg·g⁻¹), and higher values for pH (4.12) and Ca²⁺ (27 μg·g⁻¹) compared to non-Kalmia (organic matter, 3.43%; organic-N, 1.15 mg·g⁻¹; Fe³⁺, 431 μg·g⁻¹; Mn²⁺, 23.2 μg·g⁻¹; pH, 3.14; Ca²⁺, 15.6 μg·g⁻¹) and cut black spruce-Kalmia (organic matter, 3.74%; organic-N, 0.94 mg·g⁻¹; Fe³⁺, 379 μg·g⁻¹; Mn²⁺, 27 μg·g⁻¹; pH, 2.87; Ca²⁺, 25.2 μg·g⁻¹) forest. The high C:N ratio in Kalmia mineral soil from upper B (29.73) and lower B (identified as B+) (33.08) in uncut black spruce forest was recorded compared to non-Kalmia soils in B (18.17) and B+ (17.05) horizons in uncut black spruce forest. Phenolics leached out from Kalmia litter were lower in Kalmia associated soils than the non-Kalmia soils from the uncut forest, and Kalmia associated soils from the cut forest area. Results indicate that soils associated with Kalmia were nutrient poor particularly for nitrogen, phosphorus, iron and manganese, and provide some basis for the hypothesis that Kalmia has dominated microsites that were nutrient poor prior to Kalmia colonization.     

Bryophyte biomass and species richness on the Park Grass Experiment,Rothamsted, UK

The relationships between bryophyte biomass and species richness and soil pH, nutrient applications and vascular plant biomass and species richness were analyzed for the Park Grass Experiment (Rothamsted, UK). The study examined the abundance of bryophytes in relation to long-term fertilizer and lime application and to fertilizer treatments recently being ceased on some plots. The probability of bryophytes being present on a plot increased with increasing soil pH, and on plots at soil pH 3.3–4.5, the lowest values in this experiment, there were virtually no mosses present. Total bryophyte biomass decreased with increasing vascular plant biomass and vascular plant richness. Both bryophyte biomass and species richness showed a curvilinear response to soil pH. Bryophyte biomass was markedly increased on plots where nitrogen (N) fertilization had recently been ceased. The abundance of the common bryophyte species showed individualistic responses to treatments. N had a negative effect on the abundance of Brachythecium rutabulum. Increasing soil pH, and the application of phosphorus (P) and potassium (K) fertilizer together, had a positive effect on Eurhynchium praelongum. This species was also negatively affected by N, but tolerated larger amounts of it (100–150 kg ha^–1 N) than B. rutabulum. An ephemeral moss, Bryum subapiculatum, had a unimodal response to soil pH but showed no response to N, P, K or other explanatory variables.