The conservation of ground layer lichen communities in alvar grasslands and the relevance of substitution habitats

Semi-natural calcareous grasslands (alvars) are biodiversity hotspots in Northern Europe, particularly for herb layer plants. In the last century, traditional management has ceased, and the area of grasslands has declined due to extensive encroachment. We were interested in the drivers of ground layer (alias terricolous or epigeic) lichen communities. Our survey consisted of 86 habitat fragments in western Estonia, covering four types of historic alvar grasslands and three types of alvar-like habitats. We found that the ground lichen communities were primarily soil-type-specific, but were also affected by historic disturbances and land use change. In contrast to knowledge about herb layer communities, for which shrub encroachment has been shown to be main driver, the increased density of the herb layer and the reduced diversity of microhabitats were major drivers for the ground layer lichen community. These drivers caused a decrease in species richness, but only within the species of conservation value, and also led to a shift in the composition of lichen growth form from the dominance of squamulose and crustose towards fruticose lichens. We conclude that the traditional practice of restoring alvars by cutting shrubs is insufficient to maintain ground layer lichen biodiversity. Alvar maintenance practices should include grazing, which creates various small-scale ground disturbances and increases microhabitat heterogeneity. Alvar-like habitats originating from large-scale historic disturbances appeared to be suitable for calcicolous epigeic lichens, and can therefore be considered to be temporary substitution habitats, i.e. refugia for the regional species pool.     

Environmental factors and ground disturbance affecting the composition of species and functional traits of ground layer lichens on grey dunes and dune heaths of Estonia

A unique, species‐rich and endangered lichen biota can be found on European coastal and inland sand dunes. However, it is increasingly affected by natural succession as well as by anthropogenic disturbances. We studied lichen diversity on the grey dunes and dune heaths of coastal and inland regions of Estonia. A total of 28 study plots were investigated; in each 0.1 ha study plot general environmental variables and anthropogenic disturbances were described and all epigeic lichen species were identified. We found 66 lichenized fungus (lichen) species, including several rare and ten red‐listed lichens. Multivariate analysis (DCA, CCA) was performed to examine gradients in species composition and to relate variation in species data to environmental factors. In addition, we used redundancy analysis (RDA) to relate variation in species’ trait composition to environmental factors. Species composition on grey dunes differed significantly from that on dune heaths. The characteristic species for grey dunes are, besides several Cladonia species, foliose lichens, e.g. Hypogymnia physodes, Parmelia sulcata and Peltigera spp. Also species’ traits composition was different for either habitat, indicating that sorediate lichens, foliose lichens, lichens with cyanobacterium as the main photobiont, and sparsely branched Cladonia species dominate on grey dunes, while esorediate, green‐algal, crustose and richly branched fruticose lichens are common on dune heaths. Soil pH was the most essential environmental variable for determining both species composition and species’ traits composition. The composition of lichen species was also significantly influenced by forest closeness, soil Mg content and cover of bare sand; the effect of ground disturbances was low compared to the effect of these environmental factors. To protect and conserve the species‐rich lichen biota, it is necessary to protect the dune habitats from building activity, to avoid overtrampling in recreation areas and to regularly remove shrubs and trees.     

Epiphytic lichens on Juniperus communis – an unexplored component of biodiversity in threatened alvar grassland

Semi‐natural grasslands, among them thin‐soil calcareous grasslands (alvars), have great conservation value but have become increasingly rare in Europe. The main threat to alvar grasslands is the encroachment by juniper Juniperus communis and therefore it is usually removed during the restoration practice. Juniper can also be a host plant for many epiphytic lichens, but its role as a phorophyte is poorly known. We studied epiphytic lichen diversity on 126 junipers in 17 sites in western Estonia and found 140 lichenized taxa including several rare and red‐listed species. Using indirect and direct multivariate analyses (DCA, pCCA) and general linear models we revealed that both habitat and phorophyte properties affect lichen assemblies on juniper. Lichen species richness per site showed a unimodal relationship with compound factors of site productivity and juniper characteristics (stem circumference and juniper width). Lichen species richness per phorophyte was increasing with its size and with the proportion of dead branches, and was twice higher in plate alvars than in ryhk alvars. Also, the species composition in plate alvars differed from ryhk alvars by having 42 characteristic lichen species in plate alvars vs three indicators of ryhk alvars. The composition of lichens was significantly influenced by encroachment of alvars, e.g. by high juniper cover and shrub layer height, as well as by the proportion of dead branches and stem circumference of juniper. We conclude that the epiphytic lichen assemblies on junipers are threatened by grassland encroachment similarly to ground layer lichen assemblies. We suggest that some old and scencent junipers should be preserved during the restoration of alvar grasslands.     

Competitive abilities of three narrowly endemic plant species in experimental neighborhoods along a fire gradient

We conducted field experiments manipulating lichens, shrubs, and herbs along a time-since-fire gradient and assessing effects on three endemic herbaceous species of Florida scrub: Eryngium cuneifolium, Hypericum cumulicola, and Polygonella basiramia. Responses included seed germination, survival, biomass, and fecundity. Transplants into recently burned patches generally had higher survival, larger biomass, and greater reproductive output than transplants into long-unburned patches. Open areas and sites near oaks frequently were more favorable than sites near Florida rosemary. Ground lichens did not affect germination but increased mortality rate of seedlings. Neighboring small shrubby and herbaceous species did not affect the performance of these species. Of the three species, naturally occurring E. cuneifolium were farthest from large shrubs, and their microhabitats had the least ground lichens and shrubs. Eryngium cuneifolium and H. cumulicola are capable of forming persistent seed banks and their recruitment after fire depends mostly on these dormant seeds. Polygonella basiramia relies on seed dispersal and immediate seed germination to colonize recently burned patches. Management for these species should involve variable fire regimes to allow all three species to persist along with many other scrub endemics.     

Vegetation,environmental characteristics and ideas on the maintenance of alvars on the Bruce Peninsula,Canada

Abstract. Despite international recognition that alvar habitats are important reservoirs of biodiversity, they remain little studied in North America. In this paper, the results are reported on an investigation of alvars in the central portion of their known distribution on this continent. 210 plots were distributed among seven sites and were quantitatively sampled for vascular plants, lichens, bryophytes and a suite of environmental variables. Detrended and Canonical Correspondence Analyses and other methods were used to investigate differences among alvars, within alvars and between alvars and adjacent forested habitats. The plant communities and environmental conditions were highly similar among sites in the study region, yet very different from surrounding habitats. There were abrupt changes in vegetation and environmental conditions from alvar to forest, without the presence of transition zones in the vegetation or environmental gradients as the forest was approached. The environmental factors associated with the change from alvar to forest and with variation within alvar habitat were examined. Some alvars in the study were found to contain stunted, slow-growing trees reaching ages of 524 yr. These same sites appear to have remained unburned for several centuries, while other sites likely burned 90 yr ago. The plant communities were very similar between the alvars that lacked a major, biomass-removing disturbance in centuries and alvars that had experienced catastrophic fire relatively recently. Maintenance of the plant communities and open nature of alvars appears site-specific rather than habitat specific.     

Restoration of alvar vegetation on Öland,Sweden

Abstract. Alvars in the Baltic region, particularly on the Swedish islands of Öland and Gotland and in western Estonia, are well‐known for their plant species richness and extensive populations of rare species. Grasslands make up most of alvar vegetation. The extent of these grasslands decreases because of bush enchroachment which takes place in most alvars when traditional land use practice changes, notably when grazing is ceased. The main threat for the alvar grassland is formed by the rapid expansion of Juniperus communis and Potentilla fruticosa in dry and wet sites, respectively. Applied research has been carried out during several years in order to develop plans for the restoration of alvar grassland. Thus it was shown that species richness in areas with a mixture of grassland and shrubland decreases in relation to the increased cover by shrubs. In the case of Juniperus there is a distinct drop in species number as soon as the cover exceeds 75‐80%, while the decrease is more gradual with increased cover of Potentilla. The seed banks under dense stands of these shrubs only contain ca. 20% of the species found in dry and wet alvar meadows. This was confirmed by clearing experiments. Long‐term recordings have shown that establishment of juniper seedlings takes place mainly in half‐open areas between already existing junipers. Intermediate‐sized junipers expanded faster than old and big shrubs. Potentilla shrubs recover fast from cutting or burning. After 2‐4 yr they have almost attained their former size. This recovery can be prevented when cattle grazing is introduced. In the framework of a local alvar restoration project on Öland, covering 7000 ha, grazing regimes are re‐established, fences erected and large‐scale clearings carried out. Recently established juniper shrubs are being cleared, intermediate sized junipers (30‐50 yr old) are selectively removed while creating a mosaic landscape with high biological diversity. Older dense juniper stands are left alone or are only partly cleared. Potentilla stands in moist areas are cut to create moist meadows which are breeding grounds for waders, and to establish corridors between remaining open areas. Three items are discussed (1) the importance of the Stora Alvar area; (2) re‐introduction of grazing and (3) re‐introduction of species. The outstanding importance of the area regarding species richness and endemism should be recognized through a ‘golden list’ to be used along with red and blue lists.     

Recovery of pristine boreal forest floor community after selective removal of understorey, ground and humus layers

In boreal spruce forests that rarely experience extensive disturbances, fine-scale vegetation gaps are important for succession dynamics and species diversity. We examined the community implications of fine-scale gap disturbances by selective removal of vegetation layers in a pristine boreal spruce forest in Northern Finland. The aim was to investigate how the speed of recovery depends on the type of disturbance and the species growth form. We also wanted to know if there appeared changes in species composition after disturbance. Five different treatments were applied in the study: Control, removal of the ground layer (bryophytes and lichens), removal of the understorey layer (dwarf shrubs, herbs and graminoids), removal of both the ground and understorey layers, and complete removal of the vegetation and humus layers above the mineral soil. The vegetation recovery was monitored in terms of cover and species numbers over a 5-year period. Understorey layer cover, composed mainly of clonal dwarf shrubs, recovered completely in 4 years in treatments where the humus layer remained intact, whereas ground layer cover did not reach the control level in plots from where bryophytes and lichens were removed. Recovery was faster in terms of species number than species cover. Bryophytes, graminoids and dominant dwarf shrubs appeared in all disturbed plots quickly after disturbance. Seedlings of trees appeared exclusively in disturbed plots. Graminoids dominated after the removal of humus layer. The results indicate that the regeneration of forest floor after small gap disturbance occurs mainly by re-establishment of the dominant species. Although destruction of the humus layer leaves a long-lasting scar to the forest floor, exposing of mineral soil may enhance the sexual reproduction of dominant species and the colonization of weaker competitors.     

Changes in the cryptogam flora of the Wealden sandrocks, 1688–1984

The cryptogam flora of 15 sandrock sites in the central Weald, U.K. was assessed and compared with previous records beginning in 1688. A total of 264 species was recorded (29 algae, 90 lichens, 165 bryophytes and eight ferns) of which 18 lichens, 21 bryophytes and one fern were not relocated. Air pollution, changes in the microclimate brought about by invasion of trees and shrubs and trampling appear to be the main cause for the decline in the flora, although over-collection of rarities was also a factor.     

Identifying plant functional types using floristic data bases: Ecological correlates of plant range size

Abstract. In an effort to identify ‘plant functional types’, the islands floras of Great Britain and Kríti (Crete, Greece) were examined separately for ecological correlates of plant range size. Plant functional types (PFTs) were defined here as categories into which plants could be grouped on the basis of attributes that predict greater or lesser sensitivity to ecological variability. Plant range size indicates commonness of a species and was assumed to be a proxy for ‘ecological flexibility’, i.e. species of larger range sizes can better withstand environmental change and differences than species of smaller range sizes. Using evolutionary comparative methods that account for the effect of taxonomic relatedness, both floras were investigated for the effects on range size of woodiness vs. non-woodiness, trees vs. shrubs, trees vs. herbs and shrubs vs. herbs. The British flora was examined additionally for the effects of wind- vs. non-wind-pollination, self vs. animal pollination and animal vs. non-animal fruit dispersal on range size. Two analyses showed significant effects on range size: for British species, trees had larger ranges than shrubs, and wind- pollinated species had larger ranges than non-wind-pollinated species. It is suggested that the lack of a similar pattern for shrubs and trees in Kríti is because the lower water availability of Kríti imbues shrubs with an ecophysiological advantage not relevant in plants of Great Britain. That trees have larger range sizes than shrubs in Great Britain is ascribed to the greater importance of competition for light when other factors are not at issue. The greater range of wind-pollinated than non-windpollinated species in Great Britain is postulated to be because both mutualists must be capable of invading new areas. This may be termed a ‘cost of mutualism’. In terms of PFTs, the results indicate that ‘life-form’ is too broad a classification category by which to differentiate relative sensitivity to environmental variability in Great Britain, in that there were significant differences in range size of trees and shrubs, but not between either of the two categories and herbs, or between woody and non-woody plants. Although pollination type may predict relative sensitivity to variation in Great Britain, dispersal type will not. Finally, differences between Great Britain and Kríti in relative range size patterns suggests that plant functional types may be specific to a region or set of conditions.     

How to tell a shrub from a tree: A life‐history perspective from a South African savanna

The ecological differences between ‘shrubs’ and ‘trees’ are surprisingly poorly understood and clear ecological definitions of these two constructs do not exist. It is not clear whether a shrub is simply a small tree or whether shrubs represent a distinct life‐history strategy. This question is of special interest in African savannas, where shrubs and trees often co‐dominate, but are often treated uniformly as ‘woody plants’ even though the tree to shrub ratio is an important determinant of ecosystem functioning. In this study we use data from a long‐term fire experiment, together with a trait‐based approach to test (i) if woody species usually classified as shrubs or trees in African savanna differ in key traits related to disturbance and resource use; and (ii) if these differences justify the interpretation of the two growth forms as distinct life‐history strategies. We measured for 22 of the most common woody plant species of a South African savanna 27 plant traits related to plant architecture, life‐history, leaf characteristics, photosynthesis and resprouting capacity. Furthermore we evaluated their performance during a long‐term fire experiment. We found that woody plants authors call (i) shrubs; (ii) shrubs sometimes small trees; and (3) trees responded differently to long‐term fire treatments. We additionally found significant differences in architecture, diameter‐height‐allometry, foliage density, resprouting vigour after fire, minimum fruiting height and foliar δ¹³C between these three woody plant types. We interpret these findings as evidence for at least two different life‐history‐strategies: an avoidance/adaptation strategy for shrubs (early reproduction + adaptation to minor disturbance) and an escape strategy for trees (promoted investment in height growth + delayed reproduction).     

The differences in carbon dynamics between boreal dwarf shrubs and Scots pine seedlings in a microcosm study

Aims The ground level of boreal pine forests consists of a dense layer of ericaceous shrubs, herbs, grasses, mosses and lichens. The primary productivity of this forest floor vegetation is notable but the role the most common ericoid dwarf shrub plant species,Calluna vulgaris,Vaccinium myrtillusandVaccinium vitis-idaea, play in carbon (C) cycling in these ecosystems is poorly understood. Here, we studied their C dynamics in detail using plants of similar size (age 14–19 months) in a microcosm study.     

Vertical gradients in leaf trait diversity in a New Zealand forest

Leaves come in a remarkable diversity of sizes and shapes. However, spatial patterns in leaf trait diversity are rarely investigated and poorly resolved. We used a hierarchical approach to evaluate vertical variability in leaf morphology (i.e., leaf trait diversity) in 16 common tree and shrub species inhabiting a New Zealand forest. Height-related heterogeneity in leaf area, specific leaf area, circularity and length to width ratio was analyzed at three scales: (1) among leaves within plants, (2) among plants within species and (3) among species within functional groups (i.e., trees vs. shrubs). Results were scale dependent. Among-leaf morphological diversity was unrelated to plant height. Among-individual morphological diversity increased with the average height of each species, indicating that taller plant species express a greater range of leaf traits than shorter species. Among-species morphological diversity was higher in shrubs than in trees. We hypothesize that scale-dependent patterns in leaf trait diversity result from scale-dependent adaptations to forest environmental conditions. As trees grow from the forest floor into the canopy, they are exposed to a range of environmental conditions, which may select for a range of leaf traits through ontogeny. Conversely, shrubs never reach the forest canopy and may instead be differentially adapted to suites of environmental conditions associated with different stages of forest recovery from tree-fall disturbances. Overall results indicate that vertical patterns in leaf trait diversity exist. However, their strength and directionality are strongly scale-dependent, suggesting that different processes govern leaf shape diversity at different levels of ecological organization.     

The response of lichens inhabiting exposed wood of spruce logs to post-hurricane disturbances in Western Carpathian forests

We investigated which of the following environmental factors: the number of years since the windthrow of the tree (the age of dead wood), the phytocenosis (the type of forest community), altitude, exposure, wood hardness and the spatial scale of forest disturbances (small gaps with a few fallen spruces vs large-area windthrows) contributed to the diversity and abundance of lichens inhabiting the exposed wood of windthrown spruce trees in Polish Western Carpathian forests. Both Shannon H index and sum of coverage coefficients rose with increasing age of the wood, levelling off after 11–14 y (diversity) and 14–17 y (abundance). This factor appeared to be the most important for this group of lichens, but the significant positive impact of large-area windthrows on the lichen abundance was also demonstrated by using a GLM model. The age of the wood we precisely determined on the basis of data on Norway spruce mortality collected annually in permanent plots of the Gorce National Park since 2000. Using the Shore durometer we linked the course of the wood-inhabiting lichen succession with wood decay more precisely than before. The largest number of species was associated with medium hard wood, i.e., 51 < x ≤ 80 on the Shore scale. Based on the NMDS analysis, we distinguished four age groups of logs, differing in lichen abundance and defined by the dominance of distinctive species. A large number of usually corticolous lichen species used the wood of windthrown spruce logs as an optional habitat to survive large-scale, post-hurricane forest disturbances.     

Evaluation of Swedish woodland key habitats using red-listed bryophytes and lichens

All Swedish forest land is at present being surveyed with the aim of mapping woodland key habitats which have an estimated number of ca. 70 000. An untested requirement of these habitats is that they should contain red-listed species. In order to investigate if they fulfill their aim, an inventory of nationally red-listed bryophytes and lichens was performed in ca. 120 randomly selected woodland key habitats distributed throughout the country. The species were recorded in line transects, covering the whole surface of the habitats. The mean number of red-listed species per habitat was 0.7 for bryophytes and 1.4 for lichens with 11 species at the richest site and 71% of the sites having at least one species. Nemoral forests and coniferous forests were the most species-rich habitat types. The habitats of northern Sweden were more species rich than the southern ones. There were weak correlations between presence of bryophytes and lichens. The study shows that a majority of the woodland key habitats contain red-listed bryophyte and lichen species. Further studies are needed in order to record more organism groups and to compare the woodland key habitats with the surrounding matrix forests.     

Functional response of lignicolous fungal guilds to bark beetle deforestation

Conifer-dominated forests in the northern hemisphere are prone to large-scale natural disturbances, yet our understanding of their effects beyond changes in species diversity is limited. Bark beetle disturbances provide dead wood for lignicolous fungal guilds and increase insolation but also desiccation. We investigated whether species richness of these guilds increases and functional diversity decreases after bark beetle disturbance, which would promote through habitat filtering the coexistence of species adapted to harsh conditions, i.e. light stress for lichens and substrate desiccation for wood-inhabiting fungi.We sampled epixylic and epiphytic lichens (primary producers) and wood-inhabiting fungi (mainly wood decomposers, some form ectomycorrhizas) in the Bohemian Forest (Long Term Ecological Research – LTER – Site Bavarian Forest National Park), an area in Central Europe most heavily affected by the bark beetle Ips typographus, on undisturbed plots and disturbed plots with spruce (Picea abies) dieback 8 years ago. We analysed species diversity, functional diversity (optimized by phylogeny), and functional compositions.Species richness of lichens but not that of wood-inhabiting fungi was higher on disturbed plots than on undisturbed plots. Community compositions of both guilds differed considerably on disturbed and undisturbed plots. On both types of plots, lichen communities were clustered according to functional diversity, which indicated habitat filtering, and fungal communities were overdispersed, which indicated competition. Disturbance increased the strength of these two patterns only slightly and was significant only for fungi. Single-trait analysis revealed changes in the functional composition; on disturbed plots, lichenous species with larger and more complex growth forms and fungi with large, perennial fruit bodies were favoured. Although the forest canopy changed tremendously because of the bark beetle disturbance, the most important driver of lichen and fungal diversity and mean trait assemblages seemed to be the enrichment of dead wood. The changes in insolation and moisture did not act as habitat filters for either guild. This indicated that the assembly patterns of lichen and fungal communities in coniferous forests are not affected by stand-replacing disturbances in contrast to the predictions for other disturbance regimes.     

Response of ant functional composition to fire

Little is known about the impact of disturbances on functional diversity and the long‐term provisioning of ecosystem services, especially in animals. In this work we analyze the effect of wildfire on the functional composition of Mediterranean ant communities. In particular, we asked whether a) fire changes functional composition (mean and dissimilarity of trait values) at the community level; and b) such fire‐induced functional modification is driven by changes in the relative abundance‐dominance of species or by a replacement of species with different traits. We sampled ant communities in burned and unburned plots along 22 sites in a western Mediterranean region, and we computed two complementary functional trait composition indices (‘trait average’ and ‘trait dissimilarity’) for 12 functional traits (related to resource exploitation, social structure and reproduction) and with two different datasets varying in the way species abundance is considered (i.e. abundance and occurrence data). Our results suggest a set of functional responses that seem to be related to direct mortality by fire as well as to indirect fire‐induced modifications in environmental conditions relevant for ants. Trait average of colony size, worker size, worker polymorphism and the ratio between queen and worker size, as well as the trait dissimilarity of the proportion of behaviorally dominant species and of liquid food consumption, and overall functional diversity, were higher in burned than in unburned areas. Interestingly, different patterns arise when comparing results from abundance and occurrence data. While the response to fire in trait averages is quite similar, in the case of trait dissimilarity, the higher values in response to fire are much more marked when considering occurrence rather than abundance data. Our results suggest that changes in trait average are driven at the same time by replacement of species with different traits and by changes in the relative abundance‐dominance of species, while fire promotes a higher diversity of functions that is primarily driven by rare species that are functionally unique. Overall, we observed major fire‐induced changes in functional composition in Mediterranean ant communities that might have relevant consequences for ecosystem processes and services.     

新疆石人沟山区岩面生地衣群落分布格局的环境解释

地衣作为真菌和绿藻/蓝绿藻的成功共生体,广泛分布在陆地生态中的各种栖息地。岩面生地衣作为陆地生态系统的主要组成部分,在干旱和半干旱地区陆地食物链中具有重要地位,同时对岩石的生物腐蚀和土壤的形成有重要作用。岩面生地衣的多样性和分布格局强烈地受到海拔、湿度、温度、降水量、太阳辐射强度和基物的特征(岩石类型、岩石大小、岩石的化学成分和营养成分)等多种因素的影响。为了研究乌鲁木齐县石人沟山区岩面生地衣群落与基物间的关系,该研究在乌鲁木齐县石人沟山区设立16个样地,计测样地中岩面生地衣的盖度,包括坡度、坡向、光照强度等7个环境因子,采用典范对应分析法(CCA)对各群落的物种分布格局与环境因子的关系进行了探讨。结果表明:石人沟山区的岩面生地衣共有27种,隶属于7目9科15属。其中,黄枝衣目、茶渍目和鸡皮衣目的种类较多,占该地区岩面生地衣总数的74.07%。CCA排序结果显示坡度、坡向、光照强度、湿度、岩石pH值是5个影响岩面生地衣种类分布格局的主要环境因子,并显示了岩面生地衣与样地间的对应性。     

Winners and losers in a long-term study of vegetation change at Moor House NNR: Effects of sheep-grazing and its removal on British upland vegetation

We analysed data collected between 1954 and 2000 from nine long-term experiments designed to assess the effects of sheep-grazing versus no-sheep-grazing at Moor House NNR, an Environmental Change Network site. The experiments were set up between 1954 and 1972 across a range of vegetation types typical of much of upland Britain. Data from this type of experiment are often difficult to analyse and we describe the procedures undertaken to prepare the data for analysis. We fitted the resultant data to the British National Vegetation Classification and used ordination techniques to assess the relative positions of the experiments to each other. Finally, we used Generalized Linear Mixed-effects Modelling within a Bayesian framework to model change of species taxonomic/physiognomic groups through time in both sheep-grazed and ungrazed treatments across all nine experiments; variables included species diversity, Shannon–Weiner index and derived data on occurrence and abundance of species groups based on taxonomy and physiognomy. Hurdle analysis was used to model the species groups; this analysis separated the change through time in both probability of occurrence (binomial distribution) and abundance (Poisson distribution).In the sheep-grazed plots (the “business-as-usual” treatment hence here designated the “control”) there was a reduction in species diversity and a decrease in abundance of vascular plants, grasses, lichens, liverworts and mosses; whereas herbs, sedges and shrubs increased. When probability of occurrence was considered, there was a reduction in number of presences of both lichens and liverworts. Thus, the status quo management of continuous sheep-grazing, even though reduced since 1972, has resulted in a marked change in species composition of these plant communities, with some winners and some losers, but overall they support the concept of biotic homogenization. It is likely that some of these changes were driven by external factors such as elevated atmospheric nutrient deposition. Removal of sheep grazing had some positive benefits; with the herbs, mosses, sedges and shrubs increasing, but faster reductions in grasses and liverworts. Sedges + rushes were stable. It suggested that future monitoring schemes might use either the probability of occurrence of liverworts and lichens, or the abundance of lichens, liverworts, grasses and mosses as sensitive indicators of change in upland Britain.Moreover, during the period that Moor House has been protected as a nature reserve some key plant species groups have declined in spite of reductions in grazing pressure. To reverse this trend requires some form of interventionist management. In order to increase the diversity of vascular plants some form of disturbance will probably be needed, but for bryophytes and lichens this remains a research question. If the results from these small-scale experiments are replicated at the landscape scale a reduction of sheep grazing pressure, for example in rewilding schemes, will have little effect on species composition over a 28–44 year period.