Bryophyte and lichen diversity: A comparative study

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

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.     

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.     

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.     

Patterns and drivers of phytodiversity in steppe grasslands of Central Podolia (Ukraine)

We asked: (i) Which environmental factors determine the level of α-diversity at several scales and β-diversity in steppic grasslands? (ii) How do the effects of environmental factors on α- and β-diversity vary between the different taxonomic groups (vascular plants, bryophytes, lichens)? We sampled nested-plot series ranging from 0.0001 to 100 m² and additional 10-m² plots, covering different vegetation types and management regimes in steppes and semi-natural dry grasslands of Central Podolia (Ukraine). We recorded all terricolous taxa and used topographic, soil, land-use and climatic variables as predictors. Richness-environment relationships at different scales and across taxonomic groups were assessed with multimodel inference. We also fitted power-law species-area relationships, using the exponent (z value) as a measure of β-diversity. In general, the richness values in the study region were intermediate compared to those known from similar grasslands throughout the Palaearctic, but for 1 cm² we found seven species of vascular plants, a new world record. Heat index was the most important factor for vascular plants and bryophytes (negative relation), while lichen diversity depended mainly on stone and rock cover (positive). The explanatory power of climate-related variables increased with increasing grain size, while anthropogenic burning was the most important factor for richness patterns at the finest grain sizes (positive effect). The z values showed more variation at the finest grain sizes, but no significant differences in their mean between scales. The results highlight the importance of integrating scale into ecological analyses and nature conservation assessments in order to understand and manage biological diversity in steppe ecosystems.     

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.     

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.     

Response of calcareous grassland vegetation to mowing and fluctuating weather conditions

Question: What determines the balance between the cover values of vascular plants, lichens and mosses in dry calcareous grassland communities? Location: Western Estonia. Methods: A five‐year (2001–2005) study was conducted in a dry calcareous grassland. The cover of mosses, lichens and vascular plants and all moss species was recorded in permanent plots. Vascular plants were cut in half of the plots. Data from a nearby weather station were used to calculate mean values of different weather parameters and a summer moisture index for the study years. Results: Significant differences in cover values between years were found. The fluctuations of total moss cover and the cover of the dominating moss species Ctenidium molluscum followed changes in annual precipitation. Both cover values were highest in years with high precipitation. The cover change of vascular plants was best characterized by the moisture index of the growth period (three summer months). Summers with high moisture indexes facilitated vascular plant and lichen growth. Annual precipitation and the cover of mosses had a negative influence on the cover of vascular plants. The cutting of vascular plants did not have a significant effect on moss and lichen cover. Conclusions: 1. On dry calcareous grasslands the growth of mosses is enhanced by high annual precipitation, while the growth of vascular plants and lichens is influenced rather by the high summer moisture index. The cover of vascular plants is inhibited by the large moss cover. 2. Mowing of vascular plants does not influence the cover of mosses and lichens.     

Species richness of vascular plants,bryophytes, and lichens along an altitudinal gradient in western Norway

Species richness patterns of ground-dwelling vascular plants, bryophytes, and lichens were compared along an altitudinal gradient (310–1135 m a.s.l.), in western Norway. Total species richness peaked at intermediate altitudes, vascular plant species richness peaked immediately above the forest limit (at 600–700 m a.s.l.), bryophyte species richness had no statistically significant trend, whereas lichen richness increased from the lowest point and up to the forest limit, with no trend above. It is proposed that the pattern in vascular plant species richness is enhanced by an ecotone effect. Bryophyte species richness responds to local scale factors whereas the lichen species richness may be responding to the shading from the forest trees.     

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.     

Associations Between Weevils (Coleoptera: Curculionidea) and Plants, and Conservation Values in Two Tussock Grasslands, Otago, New Zealand

Ecosystem level processes and species interactions have become important concepts in conservation and land management. Despite being New Zealand’s greatest contributors to global diversity, native invertebrates have been largely overlooked in the assessment of land values, and their diversity has often been assumed to reflect native plant diversity without justification. Invertebrates can in fact affect plant species composition, and in ecosystems such as New Zealand’s remaining indigenous and semi-modified tussock grasslands can do so in excess of more conspicuous vertebrate grazers. An understanding of the interactions between invertebrates and their plant hosts may be informative in assessing land conservation values, increase the efficiency of rapid inventory analyses and would be applicable across the production-conservation spectrum. This study considers the Curculionoidea, vascular plant, bryophyte and lichen communities of two semi-modified tussock grasslands in the Otago region of southern New Zealand. Quantitative plant and invertebrate sampling were carried out in January 2001. Data were analysed using ANalysis Of SIMilarity (ANOSIM) and Multi-Dimensional Scaling (MDS) ordination techniques. Vascular plant, bryophyte and lichen species richness was highest in the same site and plots as native weevil species richness, however the proportion of native vegetation in these locations was lower. Associations identified between Curculionoidea and vascular plants were dismissed due to the confounding effect of species frequency across samples. This appeared to have little influence on associations with bryophytes and/or lichens and these were given more weighting. The ecological implications of associations are considered and variability in weevil composition is discussed in reference to the tussock grassland environment. The importance of plant–invertebrate relationships to conservation and the uses and limitations of the PRIMER MDS ordination technique for determining associations are discussed.     

Factors influencing epiphytic bryophyte and lichen species richness at different spatial scales in managed temperate forests

The effect of management related factors on species richness of epiphytic bryophytes and lichens was studied in managed deciduous-coniferous mixed forests in Western-Hungary. At the stand level, the potential explanatory variables were tree species composition, stand structure, microclimate and light conditions, landscape and historical variables; while at tree level host tree species, tree size and light were studied. Species richness of the two epiphyte groups was positively correlated. Both for lichen and bryophyte plot level richness, the composition and diversity of tree species and the abundance of shrub layer were the most influential positive factors. Besides, for bryophytes the presence of large trees, while for lichens amount and heterogeneity of light were important. Tree level richness was mainly determined by host tree species for both groups. For bryophytes oaks, while for lichens oaks and hornbeam turned out the most favourable hosts. Tree size generally increased tree level species richness, except on pine for bryophytes and on hornbeam for lichens. The key variables for epiphytic diversity of the region were directly influenced by recent forest management; historical and landscape variables were not influential. Forest management oriented to the conservation of epiphytes should focus on: (i) the maintenance of tree species diversity in mixed stands; (ii) increment the proportion of deciduous trees (mainly oaks); (iii) conserving large trees within the stands; (iv) providing the presence of shrub and regeneration layer; (v) creating heterogeneous light conditions. For these purposes tree selection and selective cutting management seem more appropriate than shelterwood system.     

The natural exclusion of red deer from large boulder grazing refugia and the consequences for saxicolous bryophyte and lichen ecology

Large boulder grazing refugia permitted comparison of saxicolous bryophyte and lichen assemblages with those boulder tops accessible to red deer (Cervus elaphus) on a sporting estate in northwest Scotland. Plant succession was predicted to occur unchecked by grazing on the tops of these large boulders with cascading effects on bryophytes and lichens—assuming boulders had been in place over the same time period. Fifty pairs of boulders (one ≥2 m and the other accessible to red deer) were selected at random from various locations below north-facing crags. Percentage cover of each bryophyte and lichen species was estimated from three randomly placed quadrats on each boulder top. Due consideration was given to the influence of island biogeography theory in subsequent model simplification. Mean shrub cover and height, leaf-litter, bryophyte cover and bryophyte species richness were significantly higher within quadrats on large boulder tops that naturally excluded red deer. Lichen cover and lichen species richness were significantly higher on boulder tops accessible to red deer. Lichen cover was in a significant negative relationship with bryophyte cover, shrub cover and litter cover. Bryophyte cover showed a significant positive relationship with shrub height but there was an optimum shrub cover. Natural exclusion of red deer from the tops of large boulders has facilitated plant succession. The results suggest that grazing arrests the lithosere on boulder tops accessible to red deer at an early plagioclimax favouring saxicolous lichens. The results are relevant to situations where red deer might be excluded from boulder fields that hold lichen assemblages of conservation value.     

Tree and stand level variables influencing diversity of lichens on temperate broad-leaved trees in boreo-nemoral floodplain forests

Tree and stand level variables affecting the species richness, cover and composition of epiphytic lichens on temperate broad-leaved trees (Fraxinus excelsior, Quercus robur, Tilia cordata, Ulmus glabra, and U. laevis) were analysed in floodplain forest stands in Estonia. The effect of tree species, substrate characteristics, and stand and regional variables were tested by partial canonical correspondence analysis (pCCA) and by general linear mixed models (GLMM). The most pronounced factors affecting the species richness, cover and composition of epiphytic lichens are acidity of tree bark, bryophyte cover and circumference of tree stems. Stand level characteristics have less effects on the species richness of epiphytic lichens, however, lichen cover and composition was influenced by stand age and light availability. The boreo-nemoral floodplain forests represent valuable habitats for epiphytic lichens. As substrate-related factors influence the species diversity of lichens on temperate broad-leaved trees differently, it is important to consider the effect of each tree species in biodiversity and conservation studies of lichens. Nomenclature Randlane et al. (2007) for lichens; Leht (2007) for vascular plants.     

Vegetation-environment relationships in a heavy metal-dry grassland complex

Heavy-metal content is assumed to be the most important edaphic factor that determines vegetation composition on contaminated soil. We compared the effects of heavy metals on species composition and species richness in the heavy metal-dry grassland complex of the Bottendorf Hills (Central Germany) with those of other environmental factors. Based on 206 relevés, we distinguished nine communities of the classesKoelerio-Corynephoretea andFestuco-Brometea. Four communities in which the metallophytesArmeria maritima subsp.halleri andMinuartia verna subsp.hercynica occurred with high frequency were classified as heavy metal subassociations of four different dry grassland associations because of the dominance of dry grassland species. We measured the soil content of copper, zinc and lead, and the carbonate content, C/N ratio, pH and conductivity of the soil, soil depth and incident radiation per site. The first axis resulting from a DCA was positively correlated with the cover and height of the herb layer, the soil depth and soil carbonate content, and negatively with the soil content of copper, the proportion of rocks, the soil C/N ratio and incident radiation per site. The number of vascular plants, bryophyte and lichen species per plot increased with pH up to 7.5 and then decreased slightly. Species richness increased with carbonate content and conductivity of the soil and decreased with the soil C/N ratio. Heavy metal content of the soil and species richness were not correlated. The occurrence of the metallophytes was strongly related to the copper content of the soil. In conclusion our study has shown that heavy metal content is not necessarily the main factor determining the total composition and richness of grasslands on soil containing heavy metals. Heavy metal grasslands are not necessarily floristically distinct from “normal” dry grasslands.     

Plants, insects and birds in semi-natural pastures in relation to local habitat and landscape factors

The preservation of remaining semi-natural grasslands in Europe has a high conservation priority. Previously, the effects of artificial fertilisation and grazing intensity on grassland animal and plant taxa have been extensively investigated. In contrast, little is known of the effects of tree and shrub cover within semi-natural grasslands and composition of habitats in the surrounding landscape on grassland taxa. We evaluated the effect that each of these factors has on species richness and community structure of vascular plants, butterflies, bumble bees, ground beetles, dung beetles and birds surveyed simultaneously in 31 semi-natural pastures in a farmland landscape in south-central Sweden. Partial correlation analyses showed that increasing proportion of the pasture area covered by shrubs and trees had a positive effect on species richness on most taxa. Furthermore, species richness of nectar seeking butterflies and bumble bees were negatively associated with grazing intensity as reflected by grass height. At the landscape level, species richness of all taxa decreased (butterflies and birds significantly so) with increasing proportion of urban elements in a 1-km² landscape area centred on each pasture, while the number of plant and bird species were lower in landscapes with large proportion of arable fields. Our results differed markedly depending on whether the focus was on species richness or community structure. Canonical correspondence analyses (CCA) showed that the abundance of most taxa was ordered along a gradient describing tree cover within pastures and proportion of arable fields in the landscape. However, subsets of grassland birds and vascular plants, respectively, showed markedly different distribution patterns along axis one of the CCA. In contrast to current conservation policy of semi-natural pastures in Sweden, our results strongly advise against using a single-taxon approach (i.e., grassland vascular plants) to design management and conservation actions in semi-natural pastures. Careful consideration of conservation values linked to the tree and shrub layers in grasslands should always precede decisions to remove trees and shrubs on the grounds of promoting richness of vascular plants confined to semi-natural grasslands. Finally, the importance of landscape composition for mobile organisms such as birds entails that management activities should focus on the wider countryside and not exclusively on single pastures.     

Structural equation modelling detects unexpected differences between bryophyte and vascular plant richness along multiple environmental gradients

Aim The aim of this study was to develop and evaluate a structural equation model explaining the species richness of bryophytes and vascular plants along multiple environmental gradients. Our primary interests in developing this model were: (1) to evaluate the effect of tree canopy along an altitudinal gradient on bryophyte and vascular plant richness; (2) to determine to what extent lithology was able to explain richness in the two groups of plants; (3) to assess whether anthropogenic disturbances decrease richness; and (4) to explore, comparing competing models, the causal links connecting spring area, discharge and spring complexity and to assess how these variables are related to richness. Location Eighty‐six springs were sampled in the south‐eastern Alps of Italy, from the lowlands to high‐mountain regions, and on different lithologies. Methods A structural equation model (SEM) was used to test certain hypotheses about the direct and indirect effects of altitude, canopy, lithology, disturbance, spring complexity, discharge and spring area on bryophyte and vascular plant richness. Competing models were evaluated and bootstrap simulation was used to determine the stability of parameter estimates. Results SEM analysis made it possible to disentangle the different effects of canopy and lithology on bryophyte and vascular plant richness: for the former it was demonstrated that the increased richness with altitude was related primarily to lithology, whereas the latter increased because of the reduction of canopy cover. In addition, the model predicted that spring discharge determined the size of the spring area and even the complexity of the spring; these latter two variables influenced to different degrees both bryophyte and vascular plant richness. Anthropogenic disturbances affected the richness of bryophytes more than that of the vascular plants. Main conclusions Our study demonstrates that several similarities in the patterns of bryophyte and vascular plant richness are in fact induced by different environmental variables.     

Influence of experimental soil disturbances on the diversity of plants in agricultural grasslands

Aims and Methods Disturbance is supposed to play an important role for biodiversity and ecosystem stability as described by the intermediate disturbance hypothesis (IDH), which predicts highest species richness at intermediate levels of disturbances. In this study, we tested the effects of artificial soil disturbances on diversity of annual and perennial vascular plants and bryophytes in a field experiment in 86 agricultural grasslands differing in land use in two regions of Germany. On each grassland, we implemented four treatments: three treatments differing in application time of soil disturbances and one control. One year after experimental disturbance, we recorded vegetation and measured biomass productivity and bare ground. We analysed the disturbance response taking effects of region and land-use-accompanied disturbance regimes into account.Important findings Region and land-use type strongly determined plant species richness. Experimental disturbances had small positive effects on the species richness of annuals, but none on perennials or bryophytes. Bare ground was positively related to species richness of bryophytes. However, exceeding the creation of 12% bare ground further disturbance had a detrimental effect on bryophyte species richness, which corresponds to the IDH. As biomass productivity was unaffected by disturbance our results indicate that the disturbance effect on species richness of annuals was not due to decreased overall productivity, but rather due to short-term lowered inter- and intraspecific competition at the newly created microsites. Generally, our results highlight the importance of soil disturbances for species richness of annual plants and bryophytes in agricultural grasslands. However, most grasslands were disturbed naturally or by land-use practices and our additional experimental soil disturbances only had a small short-term effect. Overall, total plant diversity in grasslands seemed to be more limited by the availability of propagules rather than by suitable microsites for germination. Thus, nature conservation efforts to increase grassland diversity should focus on overcoming propagule limitation, for instance by additional sowing of seeds, while the creation of additional open patches by disturbance might only be appropriate where natural disturbances are scarce.     

Lichen species richness is highest in non-intensively used grasslands promoting suitable microhabitats and low vascular plant competition

Lichens are very sensitive to habitat changes and their species richness is likely to decline under intensive land use. Currently, a comprehensive study analyzing lichen species richness in relation to land-use types, extending over different regions and including information on habitat variables, is missing for temperate grasslands. In three German regions we studied lichen species richness in 490 plots of 16 m² representing different land-use types, livestock types, and habitat variables. Due to the absence of low-intensity pastures and substrates such as woody plants, deadwood and stones, there were no lichens in the 78 plots in Schorfheide-Chorin. In the two other regions, the richness of lichen species was 45 % higher in pastures than in meadows, and 77 % higher than in mown pastures, respectively. Among the pastures, the richness of all lichen species was on average 10 times higher in sheep-grazed pastures than in the ones grazed by cattle or horses. On average, the richness of all lichen species increased by 3.3 species per additional microhabitat. Furthermore, the richness of corticolous lichens increased by 1.2 species with 10 % higher cover of woody plants, lignicolous lichen species richness increased by 4.8 species with 1 % higher cover of deadwood, and saxicolous lichen species richness increased by 1.0 species with 1 % higher cover of stones. Our findings highlight the importance of low-intensity land use for lichen conservation. In particular, the degradation of grasslands rich in microhabitats and the destruction of lichen substrates by intensification, and conversion of unfertilized pastures formerly grazed at low intensity to meadows should be avoided to maintain lichen diversity.     

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.