Plant community patterns in Moroccan temporary ponds along latitudinal and anthropogenic disturbance gradients

Background: Temporary ponds, an abundant habitat in the Maghreb region and notably in Morocco, have a high conservation value. However, they are mainly known from the north of the country.

Aims: The aim of this work was to characterise the vegetation of Moroccan temporary ponds along a combined gradient of latitude and anthropogenic pressure.

Methods: Eighty-five ponds distributed along a north–south gradient of 750 km were sampled. For each pond, all vegetation was surveyed (flooded and dry parts) and the local abiotic characteristics were measured during two successive hydrological cycles. The prevailing anthropogenic pressures were also identified and were attributed an impact score.

Results: Eighty-one characteristic pond species (including 17 rare species) were recorded, with several new distribution data in the southern part of the latitudinal gradient. Plant communities were related to climatic and anthropogenic factors, but mostly to local factors, such as maximum water depth and soil pH. The northern ponds (wettest macroclimate) were rich in characteristic species and rare species, while the southern (driest macroclimate) ponds were more species poor.

Conclusions: In addition to the direct impact of increasing human activity, a further reduction of the floristic richness of temporary ponds is expected due to climatic changes. This is particularly the case for characteristic species which have a high conservation value.     

Spatial indicators of plant community assembly from a 453-year sand-dune chronosequence

Background : We explored evidence for spatial patterning in vegetation across a sand-dune chronosequence spanning 453 years of primary succession to test for indications of a temporal signature of niche versus neutral processes during community assembly.

Methods : The study was conducted on a series of dune-capped beach ridges located in Wilderness State Park in Emmet County of northern Lower Michigan, United States (45° 43′ N, 84° 56′ E). Nearest-neighbour survey data were subjected to species dispersion and association analyses.

Results : Both species richness and species evenness showed the classic ‘hump-shaped’ relationship across successional time, suggesting that competition is an important force in structuring this community. However, the only evidence of intra-specific aggregation occurred in the youngest dune. Older dunes displayed random spatial structure and the oldest dunes tended toward a mix of random patterns and intra-specific segregation, i.e. with nearest neighbours more likely than by chance to be inter-specific. In addition, the frequency of species having lower neighbour-species richness than expected by chance was higher in younger, not older dunes. Species with significantly low neighbour richness (based on absolute nearest neighbours) on later dunes tended to have a high proportion of nearest neighbours that were intra-specific.

Conclusions : The data provided no evidence that resident species in later succession have greater intra-specific aggregation or more niche differentiation based on horizontal niche space. The data are more consistent with theories of community assembly involving competitive equivalence and/or effects of facilitation.     

Species-rich Nardus stricta grasslands host a higher vascular plant diversity on calcareous than on siliceous bedrock

Background: Species-rich Nardus stricta grasslands are a priority habitat for conservation in Europe. They typically occur on siliceous substrates and less frequently are found on calcareous bedrock.

Aims: The present paper aimed to identify the environmental factors (i.e. bedrock type, topographic, and climatic factors) that are related with community diversity and to assess if differences in plant diversity between N. stricta communities on calcareous and siliceous bedrock occur. We hypothesised that Nardus grasslands on calcareous bedrock hosted a higher vascular plant diversity than those on siliceous bedrock.

Methods: Based on 579 vegetation surveys carried out in the south-western Alps, we assessed vascular plant diversity (species richness, Shannon diversity, and Pielou’s equitability index) of species-rich Nardus grasslands and compared it between N. stricta communities on calcareous and siliceous bedrock.

Results: Elevation was identified as the main factor related to species composition, while species diversity was mostly related to mean annual precipitation and bedrock type. Species richness, Shannon diversity, and Pielou’s equitability index were higher within the communities on calcareous rather than on siliceous bedrock and a total of 89 and 34 indicator species were detected, respectively.

Conclusions: Based on our results, we suggest to protect primarily, as a habitat of priority interest, N. stricta grasslands on calcareous substrates for the higher vascular plant diversity hosted.     

Grain size affects the relationship between species richness and above-ground biomass in semi-arid rangelands

ABSTRACT

Background: Discrepancies in the shape of the productivity–diversity relationship may arise from differences in spatial scale. We hypothesised that there is a grain size effect on the productivity–diversity relationship.

Aims: To determine the effect of three sampling grain sizes on the productivity–diversity relationship.

Methods: We applied generalised linear mixed effect models on community data from 735 vegetation plots in the Taleghan rangelands, Iran, sampled at three grain sizes (0.25, 1 and 2 m²) to ascertain plant productivity-diversity patterns, while accounting for the effects of site, plant community type, disturbance, and life form.

Results: Overall, relationships between biomass and plant species richness were unimodal at grain sizes of 0.25 and 1 m², and asymptotical at 2 m². The spurious occurrence of a single large shrub may overwhelm a small-sized sampling unit, resulting in a high estimate of the sample’s biomass relative to species richness. However, the relationship between biomass and species richness at larger grain sizes is more likely to reach an asymptote.

Conclusions: Shrubs are partly responsible for driving the relationship between plant biomass and species richness. Given that the frequency of shrubs is highly variable between small plots but not so in large plots, their presence may result in unimodal productivity–diversity relationships at small but not at large grain sizes.     

The effects of micro-habitats and grazing intensity on the vegetation of burial mounds in the Kazakh steppes

Background: Burial mounds (kurgans) of Eurasian steppes are man-made habitat islands that have the potential to harbour rich plant diversity due to micro-habitats associated with their topography.

Aims: We assessed whether kurgan micro-habitats harboured different species pools and functional groups from those found on the surrounding steppes. In addition, we asked if these mounds were affected by different grazing intensities from those on the surrounding vegetation.

Methods: We surveyed kurgan micro-habitats (northern and southern slopes, surrounding ditch) and adjacent steppe plains in non-grazed, moderately grazed and heavily grazed sites in northern Kazakhstan. We analysed differences in species composition of four habitats under three grazing regimes using Generalised Linear Mixed Models, PCA ordination and indicator species analysis.

Results: Kurgan micro-habitats had diverse vegetation and supported the co-existence of plant species with different environmental needs. We identified 16 steppe specialists confined to kurgan micro-habitats. Steppe vegetation was well-adapted to extensive grazing, although heavy grazing supported ruderals and a decline in steppe specialists. There was a significant interaction between grazing intensity and habitat type: heavy grazing supported ruderals and suppressed steppe specialists especially on the slopes.

Conclusions: We highlighted that kurgans play an important role as maintaining high plant diversity locally in extensive steppe plains in Central-Asia by increasing environmental heterogeneity and supporting specialist species confined to these micro-habitats.     

The relationship between soil water storage capacity and plant species diversity in high alpine vegetation

Background: In those alpine regions where growing season precipitation is decreasing due to climate change, the capacity of soils to retain water may become an important factor for the persistence of plant species. However, the importance of soil water storage capacity (WSC) for plant species diversity has not been studied so far.

Aims: We aim to evaluate the relevance of WSC for species diversity of alpine plant communities in relation to temperature and length of growing season.

Methods: Species diversity was determined in 150 plots from a broad range of alpine vegetation types in the calcareous western part of the central Swiss Alps. WSC of soil cores sampled in every plot was determined, as well as rooting zone temperature and snowmelt date. Linear mixed models were used to assess the relationship between environmental data and species diversity.

Results: Species diversity was most strongly and positively related to WSC, followed by mean daily minimum temperature (T_min) of the growing season. Species diversity was significantly related to date of snowmelt only in sites with high WSC and/or T_min.

Conclusions: WSC represents an integrative measure for habitat quality and accounts for differences in species diversity within the study region. In order to understand and predict responses of plant species to climate change in high mountain regions, it may be crucial to also take changes in plant water supply into account.     

Notes On The Occurrence Of Clathrus Cancellatus,Tournf., In Argyllshire

Background: Estuaries are characterised by salinity gradients and regular flooding events. These environmental factors form stress gradients, along which species composition changes.

Aims: Analyse and compare patterns of plant species diversity along the estuarine salinity and flooding gradients of the Elbe and Connecticut Rivers.

Methods: Vegetation was sampled at three elevations (low, mid, high) in five sites of each marsh type (fresh, brackish, salt) in both estuaries. Patterns of species density (SD) and evenness (E) along the gradients were analysed and compared between the two estuaries with three-factor ANOVAs.

Results: The regional species pool was 33% higher for the Connecticut than for the Elbe. SD of fresh marshes (19 ± 2.2) was more than twice in the Connecticut than in the Elbe. We found an overall increase in SD from low to high elevation and from salt to freshwater marshes in both estuaries. However, SD and E were strongly depressed at intermediate elevations in the Elbe fresh and brackish marshes.

Conclusions: Although diversity patterns in the two estuaries show overall similarities, patterns of SD and E differ, when particular elevational zones and marsh types are compared. We hypothesise this to be due to evolutionary and historical influences on the regional species pools, shaping the impact of local biotic and abiotic processes.     

Plant richness and composition in hardwood forest understories vary along an acidic deposition and soil-chemical gradient in the northeastern United States

Aims

A century of atmospheric deposition of sulfur and nitrogen has acidified soils and undermined the health and recruitment of foundational tree species in the northeastern US. However, effects of acidic deposition on the forest understory plant communities of this region are poorly documented. We investigated how forest understory plant species composition and richness varied across gradients of acidic deposition and soil acidity in the Adirondack Mountains of New York State.

Methods

We surveyed understory vegetation and soils in hardwood forests on 20 small watersheds and built models of community composition and richness as functions of soil chemistry, nitrogen and sulfur deposition, and other environmental variables.

Results

Community composition varied significantly with gradients of acidic deposition, soil acidity, and base cation availability (63% variance explained). Several species increased with soil acidity while others decreased. Understory plant richness decreased significantly with increasing soil acidity (r?=?0.60). The best multivariate regression model to predict richness (p?<?0.001, adjusted-R²?=?0.60) reflected positive effects of pH and carbon-to-nitrogen ratio (C:N).

Conclusions

The relationship we found between understory plant communities and a soil-chemical gradient, suggests that soil acidification can reduce diversity and alter the composition of these communities in northern hardwood forests exposed to acidic deposition.

     

Spatial distribution of functional traits of bryophytes along environmental gradients in an Atlantic Forest remnant in north-eastern Brazil

ABSTRACT

Background: Functional trait-based approaches have been used to identify patterns of plant species diversity and composition related to environmental variability/changes. Bryophytes are rapidly affected by perturbations and thus their traits and distribution are expected to show well-defined relationships with environmental variability.

Aim: To quantify the impacts of fragmentation on the diversity and structure of epiphytic bryophytes to vertical and horizontal environmental gradients in an Atlantic Forest remnant.

Methods: Six functional traits related to water relations and light tolerance were recorded and one of them, the leaf lobule area in liverworts, was selected for morphometric measurements along the gradients analysed. Functional diversity and composition metrics of bryophytes along gradients were compared.

Results: Functional diversity changed little along the vertical and horizontal gradients. Conversely, the functional composition of traits changed markedly. Traits related to water storage, such as the presence and area of lobules, and to protection against excess light incidence, such as dark pigments, were more numerous in the canopy and at the fragment edge.

Conclusions: Functional composition is more correlated with the vertical and horizontal gradients than functional diversity. The lobule of liverworts stood out as the most relevant trait to explain the adaptive strategies of bryophytes.     

Vegetation change at high elevation: scale dependence and interactive effects on Niwot Ridge

Background: High-elevation mountain systems may be particularly responsive to climate change.

Aims: Here we investigate how changes along elevation gradients in mountain systems can aid in predicting vegetation distributional changes in time, focusing on how changing climatic controls affect meso-scale transitions at the lower and upper boundaries of alpine vegetation (with forest and subnival zones, respectively) as well as micro-scale transitions among plant communities within the alpine belt. We focus on climate-related drivers, particularly in relation to climate change, but also consider how species interactions, dispersal and responses to disturbance may influence plant responses to these abiotic drivers.

Results: Empirical observations and experimental studies indicate that changing climatic controls influence both meso-scale transitions at the upper and lower boundaries of alpine vegetation and micro-scale transitions among plant communities within tundra. Micro-scale heterogeneity appears to buffer response in many cases, while interactions between climate and other changes may often accelerate change.

Conclusions: Interactions with microtopography and larger edaphic gradients have the capacity to both facilitate rapid changes and reinforce stability, and that these interactions will affect the responsiveness of vegetation to climate change at different spatial scales.     

Vegetation pattern of mountains in West Greenland – a baseline for long-term surveillance of global warming impacts

Background: Steep environmental gradients, coupled with predicted high temperature rises in the Arctic make arctic mountain vegetation highly suitable for surveillance of changes related to global warming. However, guidelines and baselines for such a purpose are widely lacking since arctic mountain vegetation has been little explored.

Aims: We explore options for long-term surveillance on the basis of a detailed analysis of extant plant community patterns and their underlying environmental conditions in the mountainous inland of West Greenland.

Methods: Distribution, abundance and site conditions of vegetation types were analysed, using 664 vegetation samples and detailed vegetation maps in four altitudinal belts.

Results: Most plant communities had a restricted elevation distribution and were confined to special habitats predominantly defined by mesotopography and soil moisture.

Conclusions: Based on the strong linkage to habitat conditions, horizontal and vertical changes of species distribution and vegetation pattern are excellent indicators for inferring underlying environmental changes on three different scales. The recommendations given concerning climate sensitive species and plant communities, ecotones for setting up observation sites as well as stratification of analysis by habitats can be the basis for establishing long-term surveillance programmes on arctic mountain vegetation.     

Litter and its interaction with standing vegetation affect seedling recruitment in Tibetan alpine grasslands

Background: Seedling recruitment is widely recognised to be important in the maintenance of diversity and coexistence of species. It is not clear how local abiotic factors mediated by litter and biotic interactions influence plant species recruitment in alpine grasslands.

Aims: To determine how litter and standing vegetation affected seedling recruitment in plant communities in Tibetan alpine grasslands.

Methods: Seedling recruitment was quantified in response to experimental treatments: (1) removal of standing vegetation and litter; (2) removal of standing vegetation with litter intact; (3) intact standing vegetation and removal of litter and (4) control: intact standing vegetation and litter.

Results: Litter removal increased seedling numbers, while standing vegetation removal had no effect. An interaction between litter and standing vegetation removal marginally increased seedling number. Species richness of seedlings was not related to either litter removal or standing vegetation removal. Species composition of seedling differed significantly between removal of both litter and standing vegetation and control.

Conclusions: Litter and its interaction with standing vegetation has an important role in affecting plant species recruitment in this alpine plant community. In contrast, biotic interactions, such as competition or facilitation from standing vegetation, appear to have only minor effects on recruitment.     

Seed-shedding in heather (Calluna vulgaris (L.) Hull

Abstract

Background: Accumulating evidence, mainly from small-scale experiments, suggests that species diversity helps stabilise ecological communities; however, this relationship needs testing at larger scales in a wider range of natural communities.

Aims: In experimentally burnt ericaceous shrub stands, we aimed to determine whether more diverse stands had more stable vegetation cover.

Methods: Using two prescribed fire management experiments – one on heathland and one in forest – at scales of 100–700 m², we compared pre-disturbance vegetation characteristics (measures of diversity, traits and composition) with resistance (degree of perturbation on disturbance), resilience (here defined as post-disturbance recovery rate) and stability (the inverse of temporal variability).

Results: Responses to disturbance were usually best explained by a measure of dominance: Simpson's index. High dominance was associated with high resilience, but low resistance and low stability. Within the forest, the shrub community had lower dominance and higher stability than it did on heathland. Dominance and diversity were strongly linked as opposite poles of a principal axis of vegetation variation; therefore, more diverse vegetation was more stable.

Conclusions: Ericaceous shrub cover is known to support important ecosystem services, such as soil protection, carbon sequestration, flood prevention and livestock grazing. Our results suggest that high shrub diversity would support stable provision of these services. Land managers who prioritise this aim in British heathlands and pinewoods should consider a wider range of management approaches than those of typical current practice.     

Trait dynamics of Mediterranean xerophytic shrub communities growing on stabilised inland dunes respond to nutrient and aridity gradients

Background: Understanding the processes that determine community assembly and their dynamics is a central issue in ecology. The analysis of functional diversity can improve our understanding of these dynamics by identifying community assembly processes.

Aims: We studied the effect of environment–community covariations on both functional diversity and functional structure of xerophytic shrub communities for inferring the community assembly processes shaping this vegetation type.

Methods: Functional diversity was quantified using (1) community-weighted mean of the studied traits, (2) functional groups, defined using Ward’s hierarchical agglomerative clustering method and (3) Rao’s quadratic entropy. Relationships between functional diversity and environmental gradients were identified by Spearman correlations and modelled using generalised additive models.

Results: Variations in community composition and functional diversity correlated with soil nutrient availability and aridity. Increasing nutrient availability resulted in both greater average plant height and higher abundance of plants with green photosynthetic organ colour, whereas the abundance of nanophanerophytes increases with aridity.

Conclusions: The species composition and trait structure of the studied Mediterranean xerophytic shrub communities varies along nutrient and aridity gradients. This supports the importance of environmental filters for the local assembly and dynamics of these inland dune communities.     

Changes in plant taxonomic and functional diversity patterns following treeline advances in the South Urals

Background: Treeline ecotones represent environmental boundaries that fluctuate in space and time and thus induce changes in plant taxonomic and functional diversity.

Aims: To study changes through time in taxonomic and functional plant diversity patterns along the treeline ecotone.

Methods: In 2002, vegetation was sampled along a gradient from upper montane forest to the treeline–alpine transition in the South Ural Mountains, Russia. In 2014, vegetation was resampled and plant functional traits were collected. We studied spatial and temporal changes in plant species composition, functional composition and functional diversity.

Results: Species composition and diversity changed along the elevational gradient. The functional composition in height, leaf area, specific leaf area and leaf nitrogen content decreased with elevation, whereas functional composition of leaf carbon content increased. We found a temporal shift towards shorter plants with smaller leaves in treeline sites. Functional richness varied in several traits along the elevational gradient, while functional dispersion showed a trend towards increased functional dispersion in height, specific leaf area and leaf nitrogen in the treeline–tundra transition.

Conclusions: Tree encroachment across the treeline ecotone has resulted in a shift in plant species relative abundances and functional diversity, possibly affecting plant community assembly patterns.     

The consequences of multiple resource shifts on the productivity and composition of alpine tundra communities: inferences from a long-term snow and nutrient manipulation experiment

Background: Gradients in the amounts and duration of snowpack and resulting soil moisture gradients have been associated with different plant communities across alpine landscapes.

Aims: The extent to which snow additions could alter plant community structure, both alone and in combination with nitrogen (N) and phosphorus (P) additions, provided an empirical assessment of the strength of these variables on structuring the plant communities of the alpine tundra at Niwot Ridge, Colorado Front Range.

Methods: A long-term snow fence was used to study vegetation changes in responses to snowpack, both alone and in conjunction with nutrient amendments, in plots established in dry and moist meadow communities in the alpine belt. Species richness, diversity, evenness and dissimilarity were evaluated after 20 years of treatments.

Results: Snow additions, alone, reduced species richness and altered species composition in dry meadow plots, but not in moist meadow; more plant species were found in the snow-impacted areas than in nearby controls. Changes in plant community structure to N and N + P additions were influenced by snow additions. Above-ground plant productivity in plots not naturally affected by snow accumulation was not increased, and the positive responses of plant species to nutrient additions were reduced by snow addition. Plant species showed individualistic responses to changes in snow and nutrients, and indirect evidence suggested that competitive interactions mediated responses. A Permanova analysis demonstrated that community dissimilarity was affected by snow, N, and P additions, but with these responses differing by community type for snow and N. Snow influenced community patterns generated by N, and finally, the communities impacted by N + P were significantly different than those affected by the individual nutrients.

Conclusions: These results show that changes in snow cover over a 20-year interval produce measureable changes in community composition that concurrently influence and are influenced by soil nutrient availability. Dry meadow communities exhibit more sensitivity to increases in snow cover whereas moist meadow communities appear more sensitive to N enrichment. This study shows that the dynamics of multiple limiting resources influence both the productivity and composition of alpine plant communities, with, species, life form, and functional traits mediating these responses.     

Report of the Alpine Section

Background: Under climate-change scenarios, rock outcrops, by providing microclimatically diverse habitats, different from those of surrounding zonal vegetation matrix, may serve as climatic refugia and thereby facilitate the persistence of specialist species.

Aim: We tested whether rock outcrops of southern temperate latitudes could act as local refugia for a cold-adapted flora in the face of global warming.

Methods: We related species composition of 50 outcrops and that of their surrounding vegetation to climatic data at local and regional scales to establish whether species distributions reflected differences that could indicate the existence of climatic refugia.

Results: While at a regional scale, species composition of the outcrops across the study area was related to mean annual precipitation and mean annual maximum temperature, locally, southern faces of outcrops had different species and lower maximum temperatures than insolated north faces and surrounding vegetation plots.

Conclusions: South faces of outcrops by providing cool microhabitats, and currently harbouring species not found in the surrounding zonal vegetation matrix, could serve as local refugia for heat-intolerant plants and other microthermic organism. The degree of ‘stability’ of these refugia will depend on the degree of future change in climate. Therefore, it is important to plan protected area networks that maximise local environmental heterogeneity, including the protection of rock outcrops both as refugia for cold-adapted species and as potential stepping stones that would allow dispersal of these species between supportive environments through unfavourable ones.     

Uncoupled changes in tree cover and field layer vegetation at two Pyrenean treeline ecotones over 11 years

Background: The alpine treeline ecotone is regarded as a sensor of the effects of global change on alpine plant communities. However, little is known about how treeline dynamics influence the diversity and composition of alpine plant communities. Such information is necessary to forecast how ascending montane forests may affect the composition of alpine flora.

Aims: We analysed the temporal variations in tree cover, plant diversity and composition, and the effect of tree cover dynamics on field layer vegetation over a period of 11 years, at two alpine treeline ecotones in the central Pyrenees, Spain.

Methods: Tree and field layer vegetation was sampled in permanent transects in 1998 and 2009, using the point-intercept method. Temporal changes in tree cover, plant species richness and abundance were characterised along the ecotone by using a randomisation approach, rarefaction curves, and a non-parametric multivariate test, respectively.

Results: Tree cover increased significantly at one of the sites, whereas plant species richness only increased at the other site where tree cover had not changed. Vegetation composition changed significantly at both sites, but it was not spatially coupled with changes in tree cover along the ecotone.

Conclusions: A change of tree cover does not necessarily trigger changes in the ground flora at the treeline over relatively short periods (decade scale). The results challenge our ability to infer short-term biodiversity impacts from upslope advance of forests. Integrated tree and field layer monitoring approaches are necessary to produce a better understanding of the impact of ongoing global change on treeline ecotones.     

Giant stem rosettes have strong facilitation effects on alpine plant communities in the tropical Andes

ABSTRACT

Background: Giant rosettes constitute one of the most distinctive growth-forms in tropical alpine ecosystems. However, their interactions with other plant species remain unexplored in high Andean páramos.

Aims: Quantify the effect of a dominant rosette (Coespeletia timotensis) on soil and microclimatic conditions and relate the impacts to plant community structure.

Methods: We analysed topsoil temperatures, soil organic matter (SOM) and plant species cover in areas adjacent to C. timotensis and paired areas outside, at three sites (4250–4360 m a.s.l.) in La Culata National Park, Venezuela. Species richness, total vegetation cover and percent cover of each species near and away from the rosettes were compared.

Results: Topsoil temperature amplitudes were lower and SOM greater near C. timotensis stems, compared to areas away from them. C. timotensis had a consistent positive effect on species richness, vegetation cover and the cover of many abundant species (including cushions and herbs).

Conclusions: The facilitation effects of C. timotensis on abiotic conditions and community structure indicate they play a key role as nurse plants, pointing to giant rosettes as foundation species for the maintenance of plant diversity in the alpine tropics.     

Limited effect of hare grazing and short-term climatic variations on the most common alpine vegetation community in the Snowy Mountains,Australia

Background: In the alpine zone of the Snowy Mountains, grazing by mammals is limited. However, introduced European hare numbers have increased since the 1970s.

Aims: To estimate the density of hares and hence grazing pressure among years. To assess the response of biomass, vegetation height and composition to a cessation of hare grazing.

Methods: We used indices of hare abundance based on spotlighting and counts of hare pellets on a transect. The effect of hare grazing on tall alpine herbfield was assessed by using 15 paired exclosure and control quadrats for six years.

Results: The indices of hare abundance suggested densities similar to those in upland areas of Britain. Grazing did not affect the composition, cover of herbs or graminoids or, for 2010, vegetation height or biomass. Variation in vegetation and hare numbers among years was not correlated with climatic variables. Observations of selective grazing suggested that impacts on vegetation may be localised and restricted to certain species. Prior analyses of hare pellets indicated that hares might spread seed of native and exotic species.

Conclusions: Hares are having no general effect on tall alpine herbfield but may affect certain plant species via selective grazing or by spread of viable seed.