Salt spray differentially affects water status, necrosis, and growth in coastal sandplain heathland species

Sandplain heathlands are disturbance-dependent plant communities that occur infrequently in coastal areas of the northeastern United States. We hypothesize that salt spray plays a role in maintaining the composition of the heathland community by excluding salt-intolerant species close to the ocean. We examined the distributions of Solidago nemoralis, Myrica pensylvanica, Pinus rigida, and Quercus spp. in heathlands and conducted greenhouse studies to determine whether different levels of salt spray tolerance explain patterns found in the field. We found that common heathland forb and shrub species grow closer to the ocean than successional woody species. In greenhouse experiments, these species differ in their water status, necrosis, and growth responses to salt spray. The tree species P. rigida and Q. rubra are more susceptible to salt spray than the common heathland species M. pensylvanica. Our results suggest that salt spray may prevent tree species in heathlands from growing close to the ocean and therefore might be an important factor in maintaining the characteristic community composition of these dwarf shrublands in coastal habitats.     

Responses of common and successional heathland species to manipulated salt spray and water availability

Coastal sandplain heathlands are a rare plant community in the northeastern United States. Salt spray and water availability are likely important factors determining heathland distribution. Field surveys and manipulative experiments were performed to examine heathland species' responses to salt spray and water availability. We surveyed field distributions of four typical heathland species: Solidago puberula, Solidago rugosa, Gaylussacia baccata, and Myrica pensylvanica. The distributions of two native tree species, Pinus rigida and Quercus ilicifolia, were also surveyed because they succeed into coastal heathlands with low disturbance frequency. We then manipulated salt spray and water in the field and measured species' water status, necrosis, and growth responses to the treatments. Predawn xylem pressure potential and necrosis were strongly affected by high salt spray and low water availability. Shoot elongation was also limited in S. puberula and S. rugosa grown in high salt, low water treatments. Gaylussacia baccata and Q. ilicifolia were particularly sensitive to high salt spray and low water, suggesting that they might excluded be from areas with those conditions. The interaction between salt spray and water availability could affect the landscape scale and should be incorporated into conservation management plans.     

Effects of Montane Heathland Restoration on Leafhopper Assemblages (Insecta: Auchenorrhyncha)

At the beginning of the 20th century, many montane heathlands were abandoned and became subject to natural succession or afforestation by humans. Thus, the formerly large montane heathlands slowly degraded into small and isolated patches. In this study, we evaluate the influence of restoration measures on leafhopper (Auchenorrhyncha) assemblages of montane heathland ecosystems in Central Europe. Our analyses comprised three different site types that were adjacent to each other: (1) montane heathlands, (2) restoration sites, and (3) control sites. Leafhoppers showed a clear response to montane heathland restoration. Thus, after 4–5 years since implementation of restoration measurements restoration sites were characterized by the highest species richness. However, detailed analyses of leafhopper diversity, species composition, and environmental parameters on the three site types revealed that restoration sites were rather similar to control sites and significantly differing from montane heathlands. We conclude that leafhoppers are excellent bioindicators for restoration measurements because they reflected environmental differences between the three site types. Restoration measurements might only be a useful instrument to promote typical montane heathland leafhopper communities in the long run. Colonization by leafhoppers is, however, dependent on many different factors such as leafhopper mobility, vegetation structure, microclimate, and the establishment of ericaceous dwarf shrubs. Practitioners should establish a management regime (grazing and sod‐cutting) that creates a mosaic of different habitat structures and increases typical heathland vegetation, thus, favoring the colonization of typical heathland leafhoppers.     

Montane heathland rejuvenation by choppering—Effects on vascular plant and arthropod assemblages

Land-use changes and atmospheric nitrogen deposition have negatively affected heathland biota. Active habitat management is one possible way of counteracting the biodiversity loss associated with these habitat alterations. However, management practices for lowland heathlands often have been transferred to montane heathlands, irrespective of the differences in environmental conditions or assemblage composition. The objective of this study was to evaluate the effects of so-called choppering for the rejuvenation of montane heathland. Choppering involves chaffing and removing the largest part of the organic layer down to the mineral soil. In this study, we compared montane heathlands that were rejuvenated through the application of choppering (CHOPPER) to old-growth montane heathlands (CONTROL). Thirteen years after the rejuvenation measures had been conducted, the environmental conditions between CHOPPER and CONTROL still differed. CHOPPER was characterised by shorter vegetation (herbs/grasses and dwarf shrubs), more bare soil, less litter and higher temperatures. Although, the vascular plants and all studied arthropod groups were affected by the environmental changes, their responses were somewhat different. CHOPPER had a unique assemblage of each taxonomic group that included at least a few heathland species that mainly occurred in this treatment. However, choppering was most beneficial for vascular plants, grasshoppers and carabid beetles. As shown for lowland heathlands, choppering is also a suitable management measure for montane heathland to rejuvenate vegetation with its characteristic arthropod fauna. Due to the intact seed banks and Ericaceae root systems with their mycorrhizas, in combination with the availability of bare soil, heathland vegetation can rapidly regenerate after choppering. The keystone structures that explain the high relevance of CHOPPER, especially for vascular plants, grasshoppers and carabid beetles, are low-growing vegetation and bare soil, which result in light and warm microclimatic conditions. Based on the results of our study, we recommend choppering as a regular management measure to rejuvenate montane heathland.     

Influence of soil and microclimate on species composition and grass encroachment in heath succession

Aims and Methods Mostly due to land use changes, European heathlands have become increasingly rare. In addition, the increasing amount of atmospheric nitrogen deposition has resulted in an encroachment of grasses and a loss in species diversity. Despite many investigations, information about the precise environmental parameters that determine the development and maintenance of heathland vegetation is still insufficient. In order to determine the environmental factors that control heath succession and grass encroachment, and to develop appropriate management schemes, we studied the influence of several soil and microclimate parameters on species composition and vegetation characteristics in five successional stages in a coastal heathland on the island of Hiddensee, north-east Germany, where the encroachment of Carex arenaria has become a major problem.Important findings We recorded the highest plant species richness in grey dune and birch forest plots, while the encroachment of C. arenaria let to a significant decline in plant species richness. The most important environmental factors influencing species richness and distribution of single species were microclimate, soil moisture, soil pH and the C/N ratio. While many studies reported the importance of differences in nutrient availability, we found no significant correlations between soil nutrient availability and vegetation pattern. Environmental conditions in dense C. arenaria stands, especially soil properties (e.g. soil pH), showed great differences in comparison to the other successional stages. However, no correlations between the encroachment of C. arenaria and single environmental factors were found. Our results show that not only soil nutrients are important abiotic factors in heaths but that also microclimate and soil moisture play an important role and that many factors are involved in heath succession and in the promotion of grass encroachment. Management plans for the conservation and restoration of heathlands should therefore focus on the specific site conditions and should take several abiotic and biotic factors into account.     

Rapid response of Orthoptera to restoration of montane heathland

Understanding how to restore threatened ecosystems is of special relevance for nature conservation. The aim of this study was to use Orthoptera as ecological indicators for the effects of montane heathland restoration in Central Europe. We analysed the three following treatments: (i) montane heathlands (MONHEATH) (N = 7), (ii) restoration sites (RESSITE) (N = 3) and (iii) clear-cuts of spruce forests as unprocessed and ungrazed control sites (CONTROL) (N = 3). Vegetation structure and microclimate differed considerably between MONHEATH on the one hand and RESSITE and CONTROL on the other hand. Orthoptera species richness and density did so too. MONHEATH was characterised by a high-growing dense dwarf-shrub and moss layer having a cool microclimate and high soil moisture. In contrast, RESSITE and CONTROL had sparse vegetation and a warm microclimate; Orthoptera species richness and density was highest on these sites. Our study clearly showed that heathland Orthoptera responded rapidly to restoration measures, while Ericaceae dwarf shrubs slowly established. The vast majority of Orthoptera species found on the restoration sites are early and mid-successional species. The colonization of the sites by late-successional Orthoptera species in the future will depend on the further development of the heathland vegetation; that is, if Ericaceae will expand to the sites. We conclude that the realised restoration measures are suitable to promote heathland Orthoptera of early and mid-successional stages. However, the current management of montane heathlands is insufficient and needs to be intensified in order to provide structurally diverse habitats with their characteristic orthopteran assemblages.     

Interactive effects between plant functional types and soil factors on tundra species diversity and community composition

Plant communities are coupled with abiotic factors, as species diversity and community composition both respond to and influence climate and soil characteristics. Interactions between vegetation and abiotic factors depend on plant functional types (PFT) as different growth forms will have differential responses to and effects on site characteristics. However, despite the importance of different PFT for community assembly and ecosystem functioning, research has mainly focused on vascular plants. Here, we established a set of observational plots in two contrasting habitats in northeastern Siberia in order to assess the relationship between species diversity and community composition with soil variables, as well as the relationship between vegetation cover and species diversity for two PFT (nonvascular and vascular). We found that nonvascular species diversity decreased with soil acidity and moisture and, to a lesser extent, with soil temperature and active layer thickness. In contrast, no such correlation was found for vascular species diversity. Differences in community composition were found mainly along soil acidity and moisture gradients. However, the proportion of variation in composition explained by the measured soil variables was much lower for nonvascular than for vascular species when considering the PFT separately. We also found different relationships between vegetation cover and species diversity according the PFT and habitat. In support of niche differentiation theory, species diversity and community composition were related to edaphic factors. The distinct relationships found for nonvascular and vascular species suggest the importance of considering multiple PFT when assessing species diversity and composition and their interaction with edaphic factors. Synthesis: Identifying vegetation responses to edaphic factors is a first step toward a better understanding of vegetation–soil feedbacks under climate change. Our results suggest that incorporating differential responses of PFT is important for predicting vegetation shifts, primary productivity, and in turn, ecosystem functioning in a changing climate.     

Spider communities as evaluation tools for wet heathland restoration

Over the last two centuries wet heathlands and associated habitats, such as poor fens and bogs, have suffered extensive fragmentation. Recently, large-scale projects aim to restore these rare habitats throughout Europe. To evaluate post-restoration trajectories of wet heathlands, suitable monitoring tools are urgently needed. Here, we investigated whether spider communities are useful tools for evaluating the restoration success of wet heathlands. Ordination revealed that dissimilarity in spider communities between patches of different age classes resulted mainly from vegetation cover and soil moisture. By using a functional trait-approach, we tested if the time since restoration affects trait distribution of spiders in wet heathlands. Typical wet heathland spider species were less common with increasing vegetation encroachment and lower water content. New patches were inhabited by summer active, eurytopic (non-heathland) spiders, while more typical heathland species were found in middle-aged and old patches. Our results suggest that time-related changes in vegetation structure and moistness of restored wet heathlands are clearly reflected by spider communities. Although mobile spiders quickly recolonize the restored heathlands, it takes time for typical heathland spiders to settle. Restoration measures should prevent the negative effects of a vegetation encroachment and a high density of forested edges and should rehabilitate the hydrological cycle in order to preserve rare heathland spiders. We discuss that accounting for responses of spiders provides additional information to guide wet heathlands restoration.     

Species richness, abundance, rarity and environmental gradients in coastal barren vegetation

Coastal barrens in Nova Scotia are heathlands characterised by short, predominantly ericaceous vegetation, sparse tree cover, exposed bedrock, pockets of Sphagnum bog, and stressful climatic conditions. Although coastal barrens are prominent in the physical and cultural landscape, they are largely unprotected. We selected six barrens along the Atlantic coast, and surveyed 20 1-m² plots at each barren for vascular plants, macrolichens, mosses and environmental factors. We recorded 173 species (105 vascular, 41 macrolichen, 27 moss), including six provincially rare vascular species found predominantly in nearshore areas with high levels of substrate salt and nutrients, variable substrate depth, and short vegetation. Although vascular plant and moss richness were similarly correlated with vegetation height, substrate depth, organic matter content, and rock exposure, there were no clear correlations between vascular plant, macrolichen and moss richness across all sites. Vascular plant rarity and species richness were not correlated, but had inverse relationships with key environmental gradients. Tailoring conservation efforts to protect areas of high richness may thus mean that rare species are missed, and vice versa. Ordination and ANOSIM show that barrens vegetation differs widely among sites; therefore, protecting any singular coastal barren will not protect the entire range of vegetation communities and species in this heathland type. Conservation planning should emphasize protecting environmental gradients correlated with richness, rarity and plant community structure, including substrate depth and moisture, and vegetation height. Additionally, protected areas should include a coastal-inland gradient and a diversity of substrate types, including exposed rock and trees.     

Effects of management and adjacent forest on the heathland bryophyte layer

Heathlands are severely threatened by decreasing habitat quantity and quality. In these habitats, bryophytes form an important component of plant diversity. Therefore, the aims of this study were to investigate the effects of management and adjacent forest land use on the heathland bryophyte layer. Furthermore, effects of these factors on invasive bryophyte species were studied.

Bryophyte communities of 11 dry heathland patches bordering forest were surveyed. In these heathlands, management turned out to be an important factor explaining community composition of bryophytes. Furthermore, significant effects of adjacent land use on bryophyte community composition and species richness were found. However, these effects were rather limited in extent, occurring only close to habitat borders, i.e., up to maximum 5 m into the heathland patch. Also for individual bryophyte species, effects of adjacent forest could only be observed within maximum 5 m from the edge. One remarkable edge effect was the increased dominance of the invasive species Campylopus introflexus near the forest edge, especially at grazed sites. Effects of adjacent land use seemed to predominate management effects. Consequently, management cannot serve as a tool to mitigate edge effects on bryophyte species in heathland patches.     

Waiting for fungi: the ectomycorrhizal invasion of lowland heathlands

1.  In England, the loss of lowland heathland, a habitat of global conservation importance, is primarily due to the invasion of birch and pine. This encroachment has been researched in depth from a plant perspective but little is known about the role of mycorrhizal fungi. In lowland heathlands the resident dwarf shrubs form ericoid mycorrhizas whereas invading trees form ectomycorrhizas. Therefore, tree encroachment into heathlands can be regarded as the replacement of a resident mycorrhizal community by an invading one.
2 . This study examined how fungi form mycorrhizas with Betula and Pinus in lowland heathlands. We addressed the question of whether there are mycorrhizal fungi that mediate invasion using a molecular ecology approach to compare the mycorrhizal inoculum potential of soil at three levels of invasion (uninvaded heathland, invaded heathland and woodland) and the fungi forming mycorrhizas on tree seedlings and trees across diverse sites.
3.  We show that in lowland heathlands: (i) seedlings have severely limited access to ectomycorrhizal fungi relative to woodlands, (ii) there are few keystone spore-dispersed ectomycorrhizal fungi that can mediate tree invasion, (iii) tree seedlings can remain non-mycorrhizal for at least one year when no inoculum is present, even near saplings, and (iv) mycorrhizal seedlings achieve greater biomass than non-mycorrhizal seedlings. Within uninvaded heathland we detected only Rhizopogon luteolus , Suillus variegatus , S. bovinus ( Pinus symbionts) and Laccaria proxima (primarily a Betula symbiont).
4. Synthesis . Overall, ectomycorrhizal inoculum in lowland heathlands is rare; most tree seedlings growing in heathland soil are not mycorrhizal due to limited spore dispersal, poorly developed spore banks and weak common mycorrhizal networks. These seedlings can persist awaiting mycorrhization to boost their growth.     

Post‐fire regeneration in alpine heathland: Does fire severity matter?

Fire severity is thought to be an important determinant of landscape patterns of post‐fire regeneration, yet there have been few studies of the effects of variation in fire severity at landscape scales on floristic diversity and composition, and none within alpine vegetation. Understanding how fire severity affects alpine vegetation is important because fire is relatively infrequent in alpine environments. Globally, alpine ecosystems are at risk from climate change, which, in addition to warming, is likely to increase the severity and frequency of fire in south‐eastern Australia. Here we examine the effects of variation in fire severity on plant diversity and vegetation composition, 5 years after the widespread fires of 2003. We used floristic data from two wide‐spread vegetation types on the Bogong High Plains: open heathland and closed heathland. Three alternative models were tested relating variation in plant community attributes (e.g. diversity, ground cover of dominant species, amount of bare ground) to variation in fire severity. The models were (i) ‘linear’, attributes vary linearly with fire severity; (ii) ‘intermediate disturbance’, attributes are highest at intermediate fire severity and lowest at both low‐ and high‐severity; and (iii) ‘null’, attributes are unaffected by fire severity. In both heathlands, there were few differences in floristic diversity, cover of dominant species and community composition, across the strong fire severity gradient. The null model was most supported in the vast majority of cases, with only limited support for either the linear and intermediate disturbance models. Our data indicate that in both heathlands, vegetation attributes in burnt vegetation were converging towards that of the unburnt state. We conclude that fire severity had little impact on post‐fire regeneration, and that both closed and open alpine heathlands are resilient to variation in fire severity during landscape scale fires.     

Soil pH and phosphorus drive species composition and richness in semi-natural heathlands and grasslands unaffected by twentieth-century agricultural intensification

Background: Increased soil phosphorus (P) caused by agricultural intensification has been associated with decreased plant species richness (SR) in central Europe. How plant communities and soil P gradients are related in unimproved open habitats remains unclear.

Aims: The aim of this article was to characterise the relationship between soil chemical parameters and plant species composition and richness in unimproved open habitats.

Methods: The influence of soil chemical parameters (pH, P, K, Mg) on species composition was assessed, using data from 40 heathland and 54 grassland plots, by non-metric multidimensional scaling and permutational multivariate analysis of variance. The relationship between soil chemical parameters and SR was tested by linear mixed effects models.

Results: A direct relationship between heathland community composition and pH was observed, explaining 10% of variation in species composition, while P, Mg and pH together explained 17% of variation in grassland composition. In heathlands, SR increased with increasing pH, whereas in grasslands, SR decreased with increasing soil P.

Conclusions: Soil chemical parameters were substantially related to plant community composition and richness. In an area spared from a century of agricultural intensification, reduced pH appeared to constrain SR in heathlands, while even slight P increases (<10 mg kg^?1) depressed plant SR in semi-natural grasslands.     

Plant communities,species richness and their environmental correlates in the sandy heaths of Little Desert National Park,Victoria

Plant community composition and its likely environmental controls were investigated for 200 sample plots (each 100 m²) from Mediterranean-type vegetation in the Little Desert National Park, Victoria. TWINSPAN classification revealed four readily identifiable vegetation types; mallee-broombush, heathland, stringybark open woodland, and an assemblage intermediate between mallee-broombush and heathland referred to here as broom-heath. Mallee-broombush was found on Parilla Sands characterized by high Ca levels relative to heathland and stringybark open woodlands on unconsolidated Lowan Sands. The first axis of a 2 dimensional non-metric MDS ordination also divided heathlands (high axis scores) from mallee-broombush (low scores), while the second separated these vegetation types from stringybark woodlands and broom-heath. Vector-fitting revealed significant correlations between the locations of samples in ordination space and exchangeable soil Ca, soil colour, aspect and Shannon–Weiner diversity. Highest species richness/diversity was associated with the ecotonal area between Parilla and Lowan Sands (i.e. broom-heath) where a number of species characteristic of different assemblages had overlapping ranges. The fire-sensitive conifer, Callitris rhomboidea, was preferentially located in stringybark woodland and broom-heath vegetation types. Its presence was positively associated with high species richness and aspects having a southerly component. Four Callitris stands sampled for population structure were all > 40 years old and showed evidence of interfire recruitment from seeds released by old, serotinous cones. Overall, results suggest that variations in plant community composition and structure in the eastern block of the Little Desert are primarily due to variations in soil properties associated with the distribution of the two dominant substrate types, Parilla Sand and Lowan Sand. However, the interplay of topography and fire behaviour has probably been more important than substrate type in determining the distribution and population structure of longer-lived, fire-sensitive species such as Callitris rhomboidea.     

Extreme winter warming events more negatively impact small rather than large soil fauna: shift in community composition explained by traits not taxa

Extreme weather events can have negative impacts on species survival and community structure when surpassing lethal thresholds. Extreme winter warming events in the Arctic rapidly melt snow and expose ecosystems to unseasonably warm air (2–10 °C for 2–14 days), but returning to cold winter climate exposes the ecosystem to lower temperatures by the loss of insulating snow. Soil animals, which play an integral part in soil processes, may be very susceptible to such events depending on the intensity of soil warming and low temperatures following these events. We simulated week‐long extreme winter warming events – using infrared heating lamps, alone or with soil warming cables – for two consecutive years in a sub‐Arctic dwarf shrub heathland. Minimum temperatures were lower and freeze‐thaw cycles were 2–11 times more frequent in treatment plots compared with control plots. Following the second event, Acari populations decreased by 39%; primarily driven by declines of Prostigmata (69%) and the Mesostigmatic nymphs (74%). A community‐weighted vertical stratification shift occurred from smaller soil dwelling (eu‐edaphic) Collembola species dominance to larger litter dwelling (hemi‐edaphic) species dominance in the canopy‐with‐soil warming plots compared with controls. The most susceptible groups to these winter warming events were the smallest individuals (Prostigmata and eu‐edaphic Collembola). This was not apparent from abundance data at the Collembola taxon level, indicating that life forms and species traits play a major role in community assembly following extreme events. The observed shift in soil community can cascade down to the micro‐flora affecting plant productivity and mineralization rates. Short‐term extreme weather events have the potential to shift community composition through trait composition with potentially large consequences for ecosystem development.     

Fire and regeneration: the role of seed banks in the dynamics of northern heathlands

Questions: How do species composition and abundance of soil seed bank and standing vegetation vary over the course of a post‐fire succession in northern heathlands? What is the role of seed banks – do they act as a refuge for early successional species or can they simply be seen as a spillover from the extant local vegetation? Location: Coastal Calluna heathlands, Western Norway. Methods: We analysed vegetation and seed bank along a 24‐year post‐fire chronosequence. Patterns in community composition, similarity and abundances were tested using multivariate analyses, Sørensen's index of similarity, vegetation cover (%) and seedling counts. Results: The total diversity of vegetation and seed bank were 60 and 54 vascular plant taxa, respectively, with 39 shared species, resulting in 68% similarity overall. Over 24 years, the heathland community progressed from open newly burned ground via species rich graminoid‐ and herb‐dominated vegetation to mature Calluna heath. Post‐fire succession was not reflected in the seed bank. The 10 most abundant species constituted 98% of the germinated seeds. The most abundant were Calluna vulgaris (49%; 12 018 seeds m^?2) and Erica tetralix (34%; 8 414 seeds m^?2). Calluna showed significantly higher germination the first 2 years following fire. Conclusions: Vegetation species richness, ranging from 23 to 46 species yr^?1, showed a unimodal pattern over the post‐fire succession. In contrast, the seed bank species richness, ranging from 21 to 31 species yr^?1, showed no trend. This suggests that the seed bank act as a refuge; providing a constant source of recruits for species that colonise newly burned areas. The traditional management regime has not depleted or destroyed the seed banks and continued management is needed to ensure sustainability of northern heathlands.     

Use of vegetation classification and plant indicators to assess grazing abandonment in Estonian coastal wetlands

Question: What are the effects of grazing abandonment on the vegetation composition of Estonian coastal wetlands? Location: Vormsi Island and Silma Nature Reserve in western Estonia, Europe. Methods: Local knowledge and field reconnaissance were used to identify current and historical management levels of wetland sites within the west Estonian study area. Nine study sites, with varying management histories, were selected comprising an area of 287 ha. A total of 198 quadrats were taken from 43 distinct vegetation patches in five of the sites. TWINSPAN analysis was used to identify community type, and a phytosociological key was constructed for character taxa. This vegetation classification was then applied within a GIS‐based context to classify all the study sites, using a ground survey technique and 1:2000 scale air photos. Results: We identified 11 different brackish coastal wetland community types. Indicator species were defined with community characteristics for the seven main vegetation types readily recognisable in the field. Coastal wet grasslands were most extensive in grazed sites, or sites that had been more intensively grazed, while abandoned sites were largely composed of Phragmites australis stands, tall grassland, and scrub. Site variations based on vegetation composition were significantly correlated with past grazing intensity. Plant community types showed significant edaphic differences, with particularly low soil moisture and high conductivity and pH for open pioneer patches compared to other vegetation types. Conclusion: Abandonment of traditionally grazed coastal grasslands threatens their characteristic biodiversity. This study found that grazing abandonment reduced the extent of coastal wetland grasslands of particular conservation value. Nevertheless, plant species of conservation interest were found across the sequence of community types described. The study shows that grazing is an important factor influencing coastal wetland plant communities but suggests that vegetation distribution is affected by environmental variables, such as topography.     

Plant trait responses to the environment and effects on ecosystem properties

A combined analysis of plant trait responses to the environment, and their effects on ecosystem properties has recently been proposed. In this study, we related the trait composition of plant communities to soil nutrients and disturbance as environmental drivers and to productivity, decomposition and soil carbon as ecosystem properties. We surveyed two sites, one comprising intensively grazed and fertilized grasslands, the other consisting of semi-natural grassland and open heathland. Species abundance and trait values of 49 species were recorded in 69 plots, as well as parameters describing soil resources, land-use disturbances, and ecosystem properties. Our main goal was to test whether the average or the diversity of the trait values of the vegetation had stronger effects on ecosystem properties (mass ratio vs. diversity hypothesis). Structural equation modeling was used to perform a simultaneous analysis of trait responses and effects. Specific leaf area and leaf nutrient contents were always negatively correlated with stem dry matter content and canopy height, indicating greater investments in supportive and nutrient-conserving tissue as plants increased in size. In the agricultural site, disturbance was the single most important factor decreasing plant height, while leaf traits such as specific leaf area and leaf nutrient contents increased with soil resources in heathlands. Productivity was directly or indirectly driven by leaf traits, and investments in structural tissue increased standing biomass and soil carbon. Different environmental drivers in the two sites produced opposing leaf trait effects on litter decomposition. Ecosystem properties were explained by the community mean trait value as predicted by the mass ratio hypothesis. Evidence for effects of functional diversity on productivity and other ecosystem properties was not detected, suggesting that diversity–productivity relationships depend on the length of the investigated environmental gradients. We conclude that changes in community composition and dominance hierarchies deserve the most attention when ecosystem properties must be maintained.     

Catchment Liming to Restore Degraded, Acidified Heathlands and Moorland Pools

Current restoration measures of degraded, acidified heathland ecosystems have not always been successful in the Netherlands. Positive effects of a restored hydrology are often counteracted by acidification of the soil and the local groundwater system. Liming of the heathlands in the catchment of moorland pools might contribute to the restoration of both habitats. Experimental catchment liming was carried out in two degraded Dutch heathlands, with doses varying between 2 and 6 tons/ha. Catchment liming resulted in increased pH and base cation concentrations in the highest elevated limed parts, as well as in the lower situated, nonlimed heath areas and moorland pools. Generally, catchment liming created suitable conditions for the return of heathland target species, and the positive effects lasted for at least 6 years. The response of the heathland vegetation to the liming has, however, been slow because only a small number of endangered plant species increased in abundance. In contrast, four Red List soft‐water macrophytes strongly increased in abundance in the moorland pool. Our results show that, even with the slow return of Red List plant species, catchment liming can be a successful management tool for the restoration of the acidified heathland landscape.     

Edaphic influences of ophiolitic substrates on vegetation in the Western Italian Alps

Background and aims

Soils derived from serpentinite (serpentine soils) often have low macronutrient concentrations, exceedingly low Ca:Mg molar ratios and high heavy metal concentrations, typically resulting in sparse vegetative cover. This combined suite of edaphic stresses is referred to as the “serpentine syndrome.” Although several plant community-level studies have been conducted to identify the most important edaphic factor limiting plant growth on serpentine, the primary factor identified has often varied by plant community and local climate. Few studies to date have been conducted in serpentine plant communities of alpine or boreal climates. The goal of our study was to determine the primary limiting edaphic factors on plant community species composition and productivity (cover) in the alpine and boreal climate of the Western Alps, Italy.

Methods

Soil properties and vegetation composition were analyzed for several sites underlain by serpentinite, gabbro, and calc-schist substrates and correlated using direct and indirect statistical methods.

Results

Boreal forest soils were well-developed and tended to have low pH throughout the soil profile resulting in high Ni availability. Alpine soils, in comparison, were less developed. The distinct serpentine plant communities of the Western Alps are most strongly correlated with high levels of bioavailable Ni associated with low soil pH. Other factors such as macronutrient deficiency, low Ca:Mg molar ratio and drought appear to be less important.

Conclusions

The strong ecological influence of Ni is caused by environmental conditions which increase metal mobilization.