Productivity–diversity patterns in arctic tundra vegetation

Productivity has long been argued to be a major driver of species richness patterns. In the present study we test alternative productivity–diversity hypotheses using vegetation data from the vast Eurasian tundra. The productivity–species pool hypothesis predicts positive relationships at both fine and coarse grain sizes, whereas the productivity–interaction hypothesis predicts unimodal patterns at fine grain size, and monotonic positive patterns at coarse grain size. We furthermore expect to find flatter positive (productivity–species pool hypothesis) or more strongly negative (productivity–interaction hypothesis) relationships for lichens and bryophytes than for vascular plants, because as a group, lichens and bryophytes are better adapted to extreme arctic conditions and more vulnerable to competition for light than the taller‐growing vascular plants. The normalised difference vegetation index (NDVI) was used as a proxy of productivity. The generally unimodal productivity–diversity patterns were most consistent with the productivity–interaction hypothesis. There was a general trend of decreasing species richness from moderately to maximally productive tundra, in agreement with an increasing importance of competitive interactions. High richness of vascular plants and lichens occurred in moderately low productive tundra areas, whereas that of bryophytes occurred in the least productive tundra habitats covered by this study. The fine and coarse grain richness trends were surprisingly uniform and no variation in beta diversity along the productivity gradient was seen for vascular plants or bryophytes. However, lichen beta diversity varied along the productivity gradient, probably reflecting their sensitivity to habitat conditions and biotic interactions. Overall, the results show evidence that productivity–diversity gradients exist in tundra and that these appear to be largely driven by competitive interactions. Our results also imply that climate warming‐driven increases in productivity will strongly affect arctic plant diversity patterns.     

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.     

Bryophytes in grassland vegetation sample plots: What is their correlation with vascular plants?

The correlation pattern between vascular plants and bryophytes was studied, comparing variance proportions accounted for by the main PCA axes of sets containing only vascular plants and sets containing random selection of both bryophytes and vascular plants with the same number of species. In material rich in bryophytes, inclusion of bryophytes almost invariably lowered the overall correlation expressed by the variance proportion of the main PCA axes. This indicates that distribution of bryophytes is largely determined by other factors than that of vascular plants. It is suggested that this may be due to the opportunistic strategy of grassland bryophytes which respond to factors of much shorter duration and greater fluctuations than vascular plants.     

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.     

Comparison of in vitro decomposition of bryophytic and tracheophytic plant material

The fossil record of bryophytes is generally poor and infertile plants attributable to liverworts and mosses could also be thalloid vascular plant gametophytes or herbaceous lycophytes respectively. The paucity of the bryophytic record could be the result of relatively rapid degradation of bryophytic material in comparison to that of vascular plants, the absence of lignified cells in bryophytes sustaining this belief. However, certain organs of bryophytes are as robust as those of vascular plants (e.g. spores) and the hydroxybenzofuran polymers produced by some mosses could be of similar preservational resistance to lignins. A simplistic experiment to test the relative resistance to decay of bryophytic material has been undertaken. Its findings suggest that bryophytic plant material has a similar preservational potential to that of a selected vascular plant, particularly in organic rich sediments. This result may be construed to imply that bryophytes were indeed less abundant in the preservational environments of the Coal Measures than might be expected on the basis of current palaeoenvironmental reconstructions.     

Shifts in bryophyte carbon isotope ratio across an elevation?��?soil age matrix on Mauna Loa, Hawaii: do bryophytes behave like vascular plants?

The carbon isotope ratio (δ(13)C) of vascular plant leaf tissue is determined by isotope discrimination, primarily mediated by stomatal and mesophyll diffusion resistances and by photosynthetic rate. These effects lead to predictable trends in leaf δ(13)C across natural gradients of elevation, irradiance and nutrient supply. Less is known about shifts in δ(13)C for bryophytes at landscape scale, as bryophytes lack stomata in the dominant gametophyte phase, and thus lack active control over CO(2) diffusion. Twelve bryophyte species were sampled across a matrix of elevation and soil ages on Mauna Loa, Hawaii Island. We tested hypotheses based on previous findings for vascular plants, which tend to have less negative δ(13)C at higher elevations or irradiances, and for leaves with higher leaf mass per area (LMA). Across the matrix, bryophytes spanned the range of δ(13)C values typical of C(3) vascular plants. Bryophytes were remarkably similar to vascular plants in exhibiting less negative δ(13)C with increasing elevation, and with lower overstory cover; additionally δ(13)C was related to bryophyte canopy projected mass per area, a trait analogous to LMA in vascular plants, also correlated negatively with overstory cover. The similarity of responses of δ(13)C in bryophytes and vascular plants to environmental factors, despite differing morphologies and diffusion pathways, points to a strong direct role of photosynthetic rate in determining δ(13)C variation at the landscape scale.     

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.     

The effects of windthrow on plant species richness in a Central European beech forest

The effects of soil disturbance caused by the uprooting of a single or a few canopy trees on species richness and composition of vascular plant species and bryophytes were examined in a temperate beech forest (Fagus sylvatica) in northern Germany. We recorded the vegetation in 57 pairs of disturbed and adjacent undisturbed plots and established a chronosequence of mound ages to study the effect of time since microsite formation on plant species richness and composition. We found significant differences in plant species richness and composition between disturbed and adjacent undisturbed plots. Species richness of both vascular plants and bryophytes was higher in the disturbed than in the undisturbed plots, but these differences were more pronounced for bryophytes. We suggest that three main factors are responsible for this differential response. The availability of microsites on the forest floor that are suitable for the recruitment of bryophytes is lower than for vascular plants. Establishment of bryophytes in disturbed microsites is favoured by a greater abundance of propagules in the close vicinity and in the soil of the disturbed microsites, as well as by a greater variety of regeneration strategies in bryophytes than in vascular plants. Time since mound formation was a major factor determining plant species richness and composition. A significant decrease in the mean number of species was found from young mounds to intermediate and old mounds. However, differences were observed between vascular plants and bryophytes in the course of changes through time in species richness and composition. A large number of exclusive and infrequent vascular plant species was observed on young mounds, among them several disturbance specialists. We suggest that the establishment of many vascular plant species was infrequent and short-lived due to unfavourable light conditions and a low abundance of propagules. By contrast, the development of a litter layer was the main reason for the decreased mean number of bryophytes on old mounds. Our study supports the view that groups of species differing in important life history traits exhibit different responses to soil disturbance.     

Comparative cryptogam ecology: a review of bryophyte and lichen traits that drive biogeochemistry

BACKGROUND: Recent decades have seen a major surge in the study of interspecific variation in functional traits in comparative plant ecology, as a tool to understanding and predicting ecosystem functions and their responses to environmental change. However, this research has been biased almost exclusively towards vascular plants. Very little is known about the role and applicability of functional traits of non-vascular cryptogams, particularly bryophytes and lichens, with respect to biogeochemical cycling. Yet these organisms are paramount determinants of biogeochemistry in several biomes, particularly cold biomes and tropical rainforests, where they: (1) contribute substantially to above-ground biomass (lichens, bryophytes); (2) host nitrogen-fixing bacteria, providing major soil N input (lichens, bryophytes); (3) control soil chemistry and nutrition through the accumulation of recalcitrant polyphenols (bryophytes) and through their control over soil and vegetation hydrology and temperatures; (4) both promote erosion (rock weathering by lichens) and prevent it (biological crusts in deserts); (5) provide a staple food to mammals such as reindeer (lichens) and arthropodes, with important feedbacks to soils and biota; and (6) both facilitate and compete with vascular plants. APPROACH: Here we review current knowledge about interspecific variation in cryptogam traits with respect to biogeochemical cycling and discuss to what extent traits and measuring protocols needed for bryophytes and lichens correspond with those applied to vascular plants. We also propose and discuss several new or recently introduced traits that may help us understand and predict the control of cryptogams over several aspects of the biogeochemistry of ecosystems. CONCLUSIONS: Whilst many methodological challenges lie ahead, comparative cryptogam ecology has the potential to meet some of the important challenges of understanding and predicting the biogeochemical and climate consequences of large-scale environmental changes driving shifts in the cryptogam components of vegetation composition.     

Impact of climate change on alpine vegetation of mountain summits in Norway

Climate change is affecting the composition and functioning of ecosystems across the globe. Mountain ecosystems are particularly sensitive to climate warming since their biota is generally limited by low temperatures. Cryptogams such as lichens and bryophytes are important for the biodiversity and functioning of these ecosystems, but have not often been incorporated in vegetation resurvey studies. Hence, we lack a good understanding of how vascular plants, lichens and bryophytes respond interactively to climate warming in alpine communities. Here we quantified long-term changes in species richness, cover, composition and thermophilization (i.e. the increasing dominance of warm-adapted species) of vascular plants, lichens and bryophytes on four summits at Dovrefjell, Norway. These summits are situated along an elevational gradient from the low alpine to high alpine zone and were surveyed for all species in 2001, 2008 and 2015. During the 15-year period, a decline in lichen richness and increase in bryophyte richness was detected, whereas no change in vascular plant richness was found. Dwarf-shrub abundance progressively increased at the expense of lichens, and thermophilization was most pronounced for vascular plants, but occurred only on the lowest summits and northern aspects. Lichens showed less thermophilization and, for the bryophytes, no significant thermophilization was found. Although recent climate change may have primarily caused the observed changes in vegetation, combined effects with non-climatic factors (e.g. grazing and trampling) are likely important as well. At a larger scale, alpine vegetation shifts could have a profound impact on biosphere functioning with feedbacks to the global climate.     

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.     

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).     

Plant colonisation in small forest–floor patches: importance of plant group and disturbance traits

It has been proposed that the interaction between life–history attributes of different organisms and distrbance characteristics play an important role in determining the successional pattern following a disturbance event. We compared the responses of vascular plants and bryophytes (mosses and hepatics) to variation in disturbance size and severity in an old-growth boreal forest during a four–year period. The experiment included two patch sizes (0.25 and 2.5 m²) and two levels of severity: humus patches (removal of vegetation) and mineral soil patches (removal of both vegetation and humus layer). Treatments were chosen to reflect some aspects of disturbance by uprooting. Species richness was significantly affected by both disturbance size and severity but the response differed among plant groups. In vascular plants, species numbers were highest in humus patches while mosses were more numerous in mineral soil patches, the most severe disturbance. In contrast, severity had no effect on hepatics. Plant recovery was more rapid in bryophytes than in vascular plants. Species richness of bryophytes had exceeded that of adjacent, undisturbed vegetation after 2-3 yr. We attribute the contrasting response of the plant groups to differences in regeneration strategies. As a group, bryophytes had a greater variety of regeneration methods than vascular plants, with several types of asexual propagules and abundant production of spores in some species. In contrast, clonal ingrowth dominated in vascular plants while seedlings were rare. Thus, our analysis supports the view that plant response to patchy disturbance is strongly dependent on the interplay among disturbance traits and specific attribtites of different plant groups.     

Community structure and diversity of bryophytes and vascular plants in abandoned fen meadows

We examined effects of abandonment on species diversity and species composition by comparing 21 calcareous fen meadows in the pre-alpine zone of central and northeastern Switzerland. Meadows were divided into three classes of successional stages (mown: annually mown in late summer, young fallow: 4–15 years, and old fallow: >15 years of abandonment). In each fen, we measured litter mass in four 20 cm×20 cm plots, as well as (aboveground) biomass and species density (number of species per unit area) of bryophytes and vascular plants. Bryophyte biomass was reduced in abandoned fens, whereas litter mass and aboveground biomass of vascular plants increased. Species density of both taxonomic groups was lower in abandoned than in mown fens. Young and old successional stages were not different except for bryophytes, for which old successional stages had higher species density than young stages. We used litter mass and aboveground biomass of vascular plants as covariables in analyses of variance to reveal their effects on species density of both taxonomic groups. For bryophytes, litter mass was more important than vascular plant biomass in explaining variance of species density. This indicates severe effects of burying by litter on bryophyte species density. For species density of vascular plants, both vascular plant biomass and litter mass were of similar importance in explaining the decreased species density. Canonical correspondence analyses showed that abandonment also had an effect on species composition of both bryophytes and vascular plants. However, young and old successional stages were not different indicating fast initial changes after abandonment, but slow secondary succession afterwards. Furthermore, indicator species analysis showed that there was no establishment of new species after abandonment that might dramatically alter fen communities. Re-introduction of mowing as a nature conservation strategy may thus be very promising – even for old fallows.     

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。     

Fungal biomass associated with the phyllosphere of bryophytes and vascular plants

Little is known about the amount of fungal biomass in the phyllosphere of bryophytes compared to higher plants. In this study, fungal biomass associated with the phyllosphere of three bryophytes (Hylocomium splendens, Pleurozium schreberi, Polytrichum commune) and three vascular plants (Avenella flexuosa, Gymnocarpium dryopteris, Vaccinium myrtillus) was investigated using ergosterol content as a proxy for fungal biomass. Phyllosphere fungi accounted for 0.2-4.0 % of the dry mass of moss gametophytes, representing the first estimation of fungal biomass associated with bryophytes. Significantly more fungal biomass was associated with the phyllosphere of bryophytes than co-occurring vascular plants. The ergosterol present in moss gametophytic tissues differed significantly between species, while the ergosterol present in vascular plant leaf tissues did not. The photosynthetic tissues of mosses had less associated fungal biomass than their senescent tissues, and the magnitude of this difference varied in a species-specific manner. The fungal biomass associated with the vascular plants studied varied significantly between localities, while that of mosses did not. The observed differences in phyllosphere community biomass suggest their size could be affected by host anatomical and physiological attributes, including micro-niche availability and chemical host defenses, in addition to abiotic factors like moisture and nutrient availability.     

Herbaria century record of increasing eutrophication in Spanish terrestrial ecosystems

Additional biological evidence is presented for the alteration of biogeochemical cycles by human activities. The leaf δ¹⁵N and the concentrations of nutrients in herbarium specimens of 24 species of vascular plants and 3 species of bryophytes collected in northern and eastern regions of Spain have substantially changed throughout the XX century. In the second half of the century, when anthropogenic nitrogen fixation and mobilization started to increase rapidly, leaf δ¹⁵N values started to decrease strongly, indicating that additional anthropogenic nitrogen is being retained in Spanish terrestrial ecosystems. The concentration of nutrients in vascular plants did not present any clear pattern, but there were increasing concentrations of N and other nutrients (P, K, and S) in the last decades in bryophytes, which are usually better biomonitors of airborne chemicals than vascular plants. Important consequences for ecosystem structure and functioning such as enhancement of the carbon sink or changes in community biodiversity and species distribution may be expected from this increase in eutrophication.     

Possible interactions between environmental factors in determining species optima

Questions: 1. Indicator values, such as those of Ellenberg, for different environmental factors are seen as independent. We tested for the presence of interactions between environmental factors (soil moisture and reaction) to see if this assumption is simplistic. 2. How close are Ellenberg indicator values (IV s) related to the observed optima of species response curves in an area peripheral to those where they have been previously employed and 3. Can the inclusion of bryophytes add to the utility of IVs? Location: South Uist, Outer Hebrides, Scotland, UK. Methods: Two grids (ca. 2000 m × 2000 m) were sampled at 50‐m intervals across the transition from machair to upland communities covering an orthogonal gradient of both soil pH (reaction) and soil moisture content. Percentage cover data for vascular plants, bryophytes and lichens were recorded, along with pH and moisture content of the underlying sand/soil/ peat. Reaction optima, derived from species response curves calculated using HOF models, were compared between wet and dry sites, and moisture optima between acidic and basic samples. Optima for the whole data set were compared to Ellenberg IVs to assess their performance in this area, with and without the inclusion of bryophytes. Results: A number of species showed substantially different pH optima at high and low soil moisture contents (18% of those tested) and different soil moisture optima at high and low pH (49%). For a number of species the IVs were poor predictors of their actual distribution across the sampled area. Bryophytes were poor at explaining local variation in the environmental factors and also their inclusion with vascular plants negatively affected the strength of relationships. Conclusions: A substantial number of species showed an interaction between soil moisture and reaction in determining their optima on the two respective gradients. It should be borne in mind that IVs such as Ellenberg's may not be independent of one another.     

苔藓植物相互作用的研究进展

综述了苔藓植物种内和种间正相互作用、种内和种间竞争、生态位与共存等的研究现状及其进展.苔藓植物常为严酷环境中的优势植物,并在保持水分和获取光及养分资源两方面存在权衡.因严酷环境中竞争的重要性相对较低,苔藓植物的正相互作用较为多见.然而,在苔藓植物种内、种间及其与维管植物之间,均不乏竞争的实例.在严酷性较低的环境如过湿的泥炭地中,竞争的重要性增加.多种苔藓植物间可存在竞争等级,该等级往往随环境的变化而变化.随机过程、物种的殖民本质以及更新策略的差异可导致苔藓植物生态位的重叠与共存,但竞争导致生态位分化也是苔藓植物共存的机理之一.苔藓植物不应简单地归类于耐压-杂草型生活史策略者,竞争仍是构建某些苔藓植物群落及其与维管植物共存群落的重要因子之一.     

Epigeic bryophytes do not improve bioindication by Ellenberg values in mountain forests

Based on a stratified random sample of 93 vegetation plots and coincident measurements of ecological conditions in mountain forests of the Bavarian Alps, the degree to which species composition and Ellenberg indicator values derived thereof were related to measured environmental variables was assessed for vascular understorey plants and epigeic bryophytes. According to Mantel tests vascular composition contained ca. 30% more ecological information than bryophyte composition. When expressed as average Ellenberg or Düll values, vascular plant-based values reflected 60% more of measured variables than bryophyte-based values. The differences remained after rarefaction of the vascular matrix to the gamma diversity of bryophytes, showing that indication is not a function of indicator richness. Analysing vascular plants and bryophytes combined yielded very similar, or even slightly less stringent relationships with the environment than using vascular plants only.Bivariate relationships of indicator values with corresponding ecological measurements confirmed the specific potential of the values to estimate ecological factors from both plant groups, but vascular plants performed better for all factors. Bryophyte indication was particularly poor for light, temperature and base saturation. Bryophyte-based indicator values did not significantly predict the residuals of measured ecological variables against vascular plant-based Ellenberg values.For the study region, it is concluded that indicator values of vascular forest understorey should be used without consideration of Düll's indicator values for epigeic bryopyhtes. There appears to be potential to improve bioindication by recalibrating indicator values of epigeic bryophytes based on ecological measurements and vascular plant indicator values.