Bryophyte and Lichen Diversity Under Simulated Environmental Change Compared with Observed Variation in Unmanipulated Alpine Tundra

Effects of simulated environmental change on bryophyte and lichen species richness and diversity in alpine tundra were investigated in a 5-year experiment at Latnjajaure, northern Sweden. The experiment had a factorial design including fertilisation and temperature enhancement in one meadow and one heath plant community. Responses in species richness, biodiversity, and species composition of bryophytes and lichens to experimental treatments were compared to the observed variation in six naturally occurring plant communities. The combination of fertilisation and enhanced temperature resulted in a species impoverishment, for bryophytes in the bryophyte-dominated community, and for lichens in the lichen-dominated communities, but the species composition stayed within the observed natural variation. During the course of the study, no species new to the investigated mid-alpine landscape were recorded, but that scenario is realistic within a decade when comparing with the processes seen in vascular plants.     

Bryophyte colonisation in experimental microcosms: the role of nutrients, defoliation and vascular vegetation

A three-year multi-factorial microcosm experiment simulating fertilisation, defoliation and the composition of vascular vegetation in a dry grassland succession was used to test four hypotheses concerning the establishment and survival of bryophytes in grassland vegetation. H₁: bryophyte cover may be used to predict bryophyte species richness. H₂: bryophyte richness is suppressed at high nutrient levels and promoted by defoliation of vascular plants. H₃: species richness of bryophytes is influenced by the species composition of the vascular vegetation. H₄: bryophyte species richness is negatively correlated with vascular plant biomass.
The relationship between bryophyte richness and bryophyte cover was found to follow the classical species-area richness curve. Bryophyte species richness responded positively to defoliation and negatively to fertilisation. The species composition of vascular vegetation had no significant effect on bryophyte richness. Bryophyte species richness was lower at high vascular plant biomass and vascular plant dry weight above 400 g m⁻² appeared fatal to bryophytes. At high nutrient levels, defoliation increased bryophyte richness, but defoliation did not fully compensate for the negative effect of fertilisation. The study reinforces the concern for short lived shuttle bryophytes in the agricultural landscape.     

Endozoochory by slugs can increase bryophyte establishment and species richness

Herbivory can affect plant community composition and diversity by removing biomass and reducing light competition. Herbivory may particularly benefit low growing species such as bryophytes, which are frequently limited by light competition. Gastropods are important herbivores of seed plants and cryptogams, furthermore, they can disperse propagules such as seeds and spores via endozoochory. However, whether gastropod herbivory can reduce the dominance of vascular plants and thereby promote the germination and establishment of endozoochorously dispersed bryophyte spores has never been tested experimentally. Moreover, it is unclear whether these possible interacting effects can influence bryophyte species richness. Here, we tested for endozoochorous spore dispersal by slugs, in combination with sowing of vascular plants, in a fully factorial common garden experiment. Enclosures contained either slugs previously fed with bryophyte sporophytes, control slugs, or no slugs. After 21 days the bryophyte cover was on average 2.8 times higher (3.9% versus 1.4%) and after eight months the bryophyte species richness 2.6 times higher (5.8 versus 2.2) in enclosures containing slugs previously fed with bryophyte sporophytes than in the other treatments. Furthermore, after eight months high vascular plant cover reduced bryophyte diversity. On average enclosures without seed sowing harboured 1.6 times more bryophyte species than the ones with seed sowing (4.2 versus 2.6), indicating competitive effects of vascular plants on bryophytes. Our findings suggest that slugs are important dispersal vectors for bryophytes and that they can increase bryophyte populations and maintain bryophyte diversity by reducing the dominance of vascular plants.     

Is vascular plant species diversity a predictor of bryophyte species diversity in Mediterranean forests?

This study aimed to (i) investigate the congruence among the species composition and diversity of bryophytes and vascular plants in forests; (ii) test if site prioritization for conservation aims by the maximization of the pooled number of vascular plant species is effective to maximize the pooled number of bryophyte species. The study was performed in six forests in Tuscany, Italy. Four-hundred and twenty vascular plant species (61 of which were woody) and 128 bryophyte species were recorded in 109 plots. Despite the good predictive value of the compositional patterns of both woody plants and total vascular with respect to the compositional pattern of bryophytes, the species richness of the latter was only marginally related to the species richness of the former two. Bryophyte rare species were not spatially related to rare plant species and neither coincided with the sites of highest plant species richness. The species accumulation curves of bryophytes behaved differently with respect to those of woody plants or total vascular plants. Reserve selection analysis based on the maximization of the pooled species richness of either woody plants or total vascular plants were not effective in maximizing the pooled species richness of bryophytes. This study indicates that species diversity of vascular plants is not likely to be a good indicator of the bryophyte species diversity in Mediterranean forests.     

Plant diversity in a calcareous wooded meadow – The significance of management continuity

Questions: What is the contribution of management continuity during the last 30–40 years to variation in species diversity and composition of a calcareous wooded meadow plant community? Is tree cover related to species diversity and composition of the herbaceous layer? What are the effects of local soil gradients on species diversity? Location: Laelatu calcareous wooded meadow, Western Estonian coastal zone. Methods: Plant community composition was assessed in 150 1 m × 1 m plots, located at 30 sites with known management history within Laelatu meadow (7 ha). Light and soil conditions and relative altitude were measured at each plot. DCA was used to analyse variation in species composition and general linear mixed models to analyse the effects of management and environmental parameters on diversity. Results: Management continuity was the primary determinant of plant community composition, followed by light conditions and soil parameters. Species richness, diversity and evenness are positively dependent on management continuity. Spatial autocorrelation is important as well. Diversity started to decline under the tree canopy where 50% or less irradiation reached the level of the herbaceous layer. We did not find significant effects of soil conditions on small‐scale diversity. Conclusions: Management continuity, together with the cover of the tree layer, are the most important determinants of diversity. Despite grassland stands with different management history are located side by side, the regeneration of diversity and composition of plant communities after restoring regular management practices is a slow process.     

A comparison of plant communities on the basis of their clonal growth patterns

This study was performed to analyse how a vegetative propagation pattern of plants affects the coexistence of species and subsequent species richness of the community. We compared community average clonal growth in the herbal communities of forests, wooded meadows, and open meadows in Laelatu, Estonia. The parameters used for the calculation of the community averages and measured for each species were ramet life span, rhizome branching, and clonal mobility. We also examined the intrinsic (i.e. independent of the environment) relationship between community clonal growth and plant species density. We found strong correlations between the environmental factors (productivity, light availability, and mowing regime) and community averages of clonal growth parameters, while species density was (negatively) correlated only with community average of rhizome increment. The community average of ramet life span decreased with the increasing biomass of the herb layer. No evidence was found to support the hypothesis that species-rich communities may consist of species with more contrasting mobility compared with species-poor communities. Independent of the effect of the environmental factors, species density was positively correlated with ramet density. There was intrinsic positive relationship between species density and community average of ramet life span at open meadow sites and intrinsic negative relationship between species density and community average of rhizome increment at wooded meadow sites. We conclude that in forest communities the capability of clonal plants to forage for light is favoured, while in unmown meadows a competitively strong phalanx growth form is advantageous. We established that ramet turnover increases and vegetative mobility decreases with increasing species diversity, although these two relationships depend strongly on the type of the studied community.Co-ordinating editor: J. Tuomi     

Plant diversity in wooded moderate-rich fens across boreal western Canada: an ecoregional perspective

Ecoregions are increasingly being used as a framework for conservation planning. The Mid-Boreal Uplands Ecoregion stretches across Canada from Manitoba to British Columbia. From the perspective of conservation and to understand the dynamics of plant diversity and community composition in a common wetland type, we examined the plant communities and environmental variables in 80 wooded moderate-rich fens within this ecoregion. Regional diversity totalled 273 species, with 86 bryophytes and 187 vascular plants. Total diversity was greatest in Manitoba and decreased in a longitudinal trend west through Saskatchewan and Alberta. This may be related, in part, to orographic precipitation at Manitoba sites and a gradient of growing degree days. Richness of locally rare vascular plants exhibited a clear west to east gradient. Ten species of provincially rare vascular plants were observed across the ecoregion, but without pattern. Ordinations and other analyses revealed distinct plant communities for all three locations, with vascular plant assemblages more discrete than bryophyte assemblages. Bryophyte diversity increased with latitude and longitude, whereas vascular plant diversity decreased. Additionally, elevation, precipitation, surface water alkalinity, water temperature, percent overstory density, and peat organic C played a role in determining species richness and community composition. Overall, species composition and diversity in a single wetland type exhibited continuous change across multiple political jurisdictions at the ecoregion scale. Conservation plans for wetlands at the ecodistrict scale may be preferable.     

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。     

Bryophyte and vascular plant species richness in boreo-nemoral moist forests and mires

We compare species richness of bryophytes and vascular plants in Estonian moist forests and mires. The material was collected from two wetland nature reserves. Bryophyte and vascular plant species were recorded in 338 homogeneous stands of approximately 1 ha in nine forest and two mire types. Regional species pools for bryophytes and vascular plants were significantly correlated. The correlations between the species richnesses of bryophytes and vascular plants per stand were positive in all community types. The relative richnesses (local richness divided by the regional species pool size) were similar for bryophyte species and for vascular plant species. This shows that on larger scales, conservation of the communities rich in species of one taxonomic plant group, maintains also the species richness of the other. The minimum number of stands needed for the maintenance of the regional species pool of typical species for the every community type was calculated using the species richness accumulation curves. Less stands are needed to maintain the bryophyte species pools (300–5300 for bryophytes and 400–35 000 for vascular plants).     

High species richness in an Estonian wooded meadow

Abstract. A wooded meadow at Laelatu in western Estonia was found to be very rich in vascular plants. The maximum number of species found was 25 in a 10 cm x 10 cm plot, 42 in a 20 cm x 20 cm plot and 63 in a 1 m² plot. Species richness is related to the management history of the site. The highest richness was found in sites with the most regular long-term mowing. Species density was lower in the case of fertilization or temporary cessation of mowing. The richest community had an LAI of 2.8 and an above-ground dry biomass of 175 g/m². With increasing height of the herb layer the number of species decreases. Small disturbances cause a remarkable decrease in species density.     

How do bryophytes govern generative recruitment of vascular plants?

Interactions between vascular plants and bryophytes determine plant community composition in many ecosystems. Yet, little is known about the importance of interspecific differences between bryophytes with respect to their effects on vascular plants. We compared the extent to which species-specific bryophyte effects on vascular plant generative recruitment depend on the following underlying mechanisms: allelopathy, mechanical obstruction, soil moisture and temperature control. We sowed 10 vascular plant species into monospecific mats of six chemically and structurally diverse bryophytes, and examined 1-yr seedling recruitment. Allelopathic effects were also assessed in a laboratory phyto-assay. Although all bryophytes suppressed vascular plant regeneration, there were significant differences between the bryophyte species. The lack of interactions indicated the absence of species-specific adaptations of vascular plants for recruitment in bryophyte mats. Differences between bryophyte species were best explained by alterations in temperature regime under bryophyte mats, mostly by reduced temperature amplitudes during germination. The temperature regime under bryophyte mats was well predicted by species-specific bryophyte cushion thickness. The fitness of established seedlings was not affected by the presence of bryophytes. Our results suggest that climatically or anthropogenically driven changes in the species' composition of bryophyte communities have knock-on effects on vascular plant populations via generative reproduction.     

Different temporal trends in vascular plant and bryophyte communities along elevational gradients over four decades

Despite many studies showing biodiversity responses to warming, the generality of such responses across taxonomic groups remains unclear. Very few studies have tested for evidence of bryophyte community responses to warming, even though bryophytes are major contributors to diversity and functioning in many ecosystems. Here, we report an empirical study comparing long‐term change in bryophyte and vascular plant communities in two sites with contrasting long‐term warming trends, using “legacy” botanical records as a baseline for comparison with contemporary resurveys. We hypothesized that ecological changes would be greater in sites with a stronger warming trend and that vascular plant communities, with narrower climatic niches, would be more sensitive than bryophyte communities to climate warming. For each taxonomic group in each site, we quantified the magnitude of changes in species'' distributions along the elevation gradient, species richness, and community composition. We found contrasted temporal changes in bryophyte vs. vascular plant communities, which only partially supported the warming hypothesis. In the area with a stronger warming trend, we found a significant increase in local diversity and dissimilarity (β‐diversity) for vascular plants, but not for bryophytes. Presence–absence data did not provide sufficient power to detect elevational shifts in species distributions. The patterns observed for bryophytes are in accordance with recent literature showing that local diversity can remain unchanged despite strong changes in composition. Regardless of whether one taxon is systematically more or less sensitive to environmental change than another, our results suggest that vascular plants cannot be used as a surrogate for bryophytes in terms of predicting the nature and magnitude of responses to warming. Thus, to assess overall biodiversity responses to global change, abundance data from different taxonomic groups and different community properties need to be synthesized.     

祁连山不同景观类型中苔藓植物物种多样性研究

通过对祁连山不同景观类型中的苔藓植物进行样方调查与物种组成统计,采用物种相似性系数、多样性指数对苔藓植物物种多样性进行了分析。结果表明,30个10 m×10 m的样地内共计苔藓植物19科34属48种,其中优势科(≥3种)有6科,含19属31种,分别占本地区苔藓植物总属数和总种数的55.88%和64.58%。在5种景观类型中,针叶林带和高山灌丛带分布的苔藓植物物种相似性最高(相似性系数为0.5357);河岸带和高山草甸带分布的苔藓植物物种相似性最低(为0),表现出祁连山地区苔藓植物分布的丰富性和复杂性;高山灌丛带分布的苔藓植物物种多样性指数最高,但针叶林带苔藓植物最为丰富,是祁连山苔藓物种多样性保护的关键区域。     

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.     

Vascular Plants Facilitated Bryophytes in a Grassland Experiment

In grassland communities vascular plants and bryophytes form two distinct layers. In order to understand the factors responsible for plant community structure, more information about interactions between these plant groups is needed. Often negative correlations between vascular plant and bryophyte covers have been reported, suggesting competition. Here we tested experimentally whether different grassland vascular plant species (Trifolium pratense, Festuca pratensis, Prunella vulgaris) had different influences on the cover of two bryophyte species (Rhytidiadelphus squarrosus, Brachythecium rutabulum). In a two-year garden pot experiment one bryophyte species and one vascular plant species were planted per pot. Bryophytes were planted at a constant density, vascular plants in four densities. The cover of both bryophyte species increased with increasing vascular plant cover, showing the facilitative effect of vascular plants through creating better microclimate, e.g., optimising temperature. Bryophyte responses to vascular plant species were species-specific. Festuca had significantly positive effects on both bryophyte species in the second year, and Trifolium on Brachythecium in both years, whereas Prunella had no significant effect on bryophytes. The facilitative effect of vascular plants was stronger at the second experimental year. In summary, the biotic effects between bryophytes and grassland vascular plants are species-specific and positive interactions are prevailing at low vascular plant densities.     

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.     

Predicting bryophyte diversity in grassland and eucalypt-dominated remnants in subhumid Tasmania

Aim To determine the environmental factors associated with bryophyte diversity in remnants in a fragmented, agricultural landscape. Location Eighty‐two remnants of tussock grassland, eucalypt woodland and eucalypt forest in the subhumid Midlands region of Tasmania, Australia. Methods Remnants were surveyed for bryophytes and predictor variables, such as vascular plant cover, climate, and topography. Management histories for each remnant were compiled using both site observation and landowner surveys. Bryophyte cover, richness, and composition were related to the independent variables using simple correlation and general linear models. Results We found weak relationships between the dependent variables and the fragmentation variables (remnant area, remnant perimeter to area ratio, distance to nearest remnant, distance to nearest larger remnant, and remnant age). Instead, climatic variables were important in predicting bryophytes, in particular those affecting humidity (minimum temperature of the coldest month, precipitation). Despite extensive sheep grazing in this landscape, grazing was not correlated with bryophyte diversity. Bryophyte diversity was not explained by vascular plant richness and was only weakly explained by composition, but was predicted by the cover of vascular plants. There was greater bryophyte cover and richness and different composition where the cover of native vascular plants was lower. Main conclusions The implications of our results are that all remnants, regardless of area, age and isolation, appear to be valuable for bryophyte conservation in this highly altered landscape. Our results also suggest that the cover of the vascular plant community, rather than its diversity, holds promise as a guide to bryophyte diversity. Bryophyte composition was similar between sites and a focus on the most species‐rich sites may be the best conservation strategy in this ecosystem.     

武汉市城区苔藓植物多样性和分布及与环境因子的关系

对武汉市7个中心城区和部分远郊城区共26个样点73个样地的苔藓群落进行调查,采集苔藓植物样本共431份。经鉴定调查区内共有苔藓植物20科35属91种,其中狭叶小羽藓（Haplocladium angustifolium（Hampe et C.Muell.）Broth.）和钝叶绢藓（Entodon obtusatus Broth.）等为优势种,分布于其中的24个样点。多样性指数分析结果显示,位于青龙山国家森林公园的α多样性Patrick和Shannon-Wiener指数最高,说明该样点苔藓植物群落的复杂程度高,群落所含信息量大。Patrick指数与环境因子的Pearson相关性分析结果显示,土壤含水率和与主要干道距离是显著影响地面生苔藓植物多样性的环境因子,而显著影响树附生苔藓植物多样性的环境因子仅与距主要干道距离相关。用典范对应分析法（CCA）研究26个样点中苔藓植物的盖度与主要环境因子的关系,结果发现人为干扰程度、草本盖度、与城市主要干道的距离等对地面生和树附生苔藓植物的分布都有显著影响。其中地面生苔藓植物的分布受人为干扰和草本盖度影响最为显著;而树附生苔藓植物的分布受附生树干胸径及与主要干道距离影响最为显著。研究结果表明武汉市区苔藓植物的多样性和分布受人为干扰较大,树附生苔藓对汽车尾气等因素更为敏感。     

Length of the Spacer Rather than its Plasticity Relates to Species Distribution in Various Natural Habitats

Much evidence from laboratory experiments and theoretical studies show that plasticity of clonal growth traits like lateral spread provides advantage in heterogeneous conditions. However, few tests of whether species with plastic clonal growth have an advantage over non-plastic species in natural conditions exist. I analyzed whether spacer length (i.e., length of a stolon or a rhizome branch between two ramets) and the variation of it (as a surrogate to its plasticity) relate to species presence and abundance in open meadow, wooded meadow and forest habitats with varying environmental conditions at Laelatu wooded meadow, Estonia. In fertilized, compared to unfertilized conditions, a weak and non-conclusive advantage of both long spacers and high variability of spacer length was detected. Abandonment in open meadows lead to a prevalence of species with shorter spacers, while on abandoned wooded meadows the species with longer spacers dominated. There was no difference in variability of spacer length between managed and abandoned meadows. In more heterogeneous forest habitats high variability of spacer length was more common, but the effect of lengthy spacers was more pronounced. The results suggest that while the variability of spacer length indeed corresponds with larger abundance in some conditions, the actual length of spacers has a more pronounced relationship with abundance of plants in natural vegetation.     

苔藓植物对植物天然更新的影响

种子是天然更新的基础,幼苗和幼树的生长是天然更新过程中最敏感的阶段。苔藓植物作为许多生态系统的主要地被物,在维管植物天然更新过程中发挥着不容忽视的作用。在许多植被类型中,都有发育良好的苔藓群落,大量的研究证明它们的存在影响到维管植物的萌发、建成和生长,并且可能进一步影响到物种的共生,有一些种类甚至可能指示树种的更新。本文就苔藓植物对物种更新早期阶段的作用和影响进行了综述。包括苔藓植物的生理特性造成的微气候变化;苔藓群落对种子传播、萌发及幼苗建成的影响。并讨论了苔藓植物与更新物种的种间关系及其对幼苗生长的化感作用。