Spatio-temporal patterns in post-burn succession on Brittany heathlands

Heathlands in the Paimpont area, Brittany, were ravaged by numerous fires in the summer of 1976. The period in which the fires occurred, and their intensity had important consequences for recolonization processes. In the tall Ulex europaeus heathland described in this paper, fire occurred relatively late (mid-July), and was of high intensity. Colonisation was mainly from propagules, often from species foreign to the original heathland. Plant succession together with vertical and horizontal structure was studied in permanent plots over 13 yr. Three stages were distinguished. Vertical structure varied greatly among the three stages. It was very simple in the colonisation phase, became more complicated during the competition phase, but became simpler again in the stabilisation phase. This was particularly evident for the moss layer, which, 13 yr after the fire, was made up of Polytrichum commune alone. Mosses showed interspecific competition with size as the decisive factor: the large mosses gradually eliminated the smaller ones. Horizontal structure was studied using spatial autocorrelation, which provided some indication of the distinctive -ness and scale of spatial pattern. Some species were weakly aggregated such as Ceratodon purpureus, Polytrichum pili-ferum, and P. juniperinum, whileothers, like the P. formosum-P. commune complex, Agrostis curtisii and Ulex minor were strongly aggregated. In the current community, 13 yr after the fire, a mosaic can be distinguished with three components: (1) a moss layer of Polytrichum commune on which a herbaceous layer, dominated by Agrostis curtisii has developed; (2) a birch wood (both Betulapéndula and B. pubescens), which established on the Polytrichum - Agrostis combination; (3) patches of Ulex minor heathland, with a few Frica and Calluna plants. Extreme simplicity and stability are the two prominent features of the present vegetation state.     

Disturbance Type and Plant Successional Communities in Bahamian Dry Forests

Different disturbances in similar habitats can produce unique successional assemblages of plants. We collected plant species composition and cover data to investigate the effects of three common types of disturbances—fire, anthropogenic clearing (‘cleared’), and clearing followed by goat grazing (‘cleared‐and‐grazed’)—on early‐successional coppice (dry forest) community structure and development on Eleuthera, Bahamas. For each disturbance type, both the ground layer (<0.5 m height) and shrub layer (>0.5 m height) were sampled in eight patches (>1 ha) of varying age (1–28 yr) since large‐scale mature coppice disturbance. Overall, plant communities differed among disturbance types; several common species had significantly higher cover in the shrub layer of fire patches, and cleared‐and‐grazed patches exhibited higher woody ground cover. Total percent cover in the shrub layer increased in a similar linear fashion along the investigated chronosequence of each disturbance type; however, cover of the common tree species, Bursera simaruba, increased at a notably slower rate in cleared‐and‐grazed patches. The pattern of increase and subsequent decrease in cover of Lantana spp. and Zanthoxylum fagara in the shrub layer was characterized by longer persistence and higher covers, respectively, in cleared‐and‐grazed patches, which also exhibited low peak cover and fast decline of nonwoody ground cover. Our results suggest that goats may accelerate some aspects of succession (e.g., quickly removing nonwoody ground cover) and retard other aspects (e.g., inhibiting growth of tree species and maintaining early‐successional shrubs in the shrub layer). These effects may lead to different successional trajectories, and have important conservation implications.     

Plant species richness and composition of heathland relics in north‐western Belgium: evidence for a rescue‐effect?

Aim To study the effects of the degree of fragmentation of heathland patches on their species richness and species composition, and to infer the ecological mechanisms behind the observed patterns. Location The heathland patches of the north‐western part of Flanders, Belgium. During the last 200 years, the heathland area in this study area has been reduced from c. 10,000 to c. 40 ha, yielding c. 150 small and highly isolated relic fragments. Methods Different isolation measures were calculated for each of the 153 inventoried heathland patches. The influence of isolation, area and habitat diversity on species richness was investigated using correlation techniques. Community composition of the patches was tested for nestedness, and the mechanisms potentially underlying this pattern were determined. Results and main conclusions Both the analyses at the species richness and community composition level yielded evidence for a positive species–area relation. This relation was not caused by higher habitat heterogeneity in larger patches. Patch isolation, measured in different ways, however seemed much more important in explaining species richness and community composition than fragment area. Our results indicate that area effects are overcompensated by the rescue‐effect: if a patch is close to other patches, species can disperse between them and prevent the species from going extinct. Species having a short living seed bank were also more sensitive to isolation than species with a long living seed bank, indicating that the latter most probably depend on the seed bank to survive periods when environmental conditions are harsh. Analogously to the spatial rescue‐effect, the existence of a persistent seed bank may lead to a sort of temporal rescue‐effect, where the extinction of a plant species is prevented through survival in the seed bank of a patch.     

Coastal heathland succession influences butterfly community composition and threatens endangered butterfly species

Succession has a strong influence on species diversity and composition of semi-natural open terrestrial ecosystems. While several studies examined the effects of succession on butterflies in grassland and forest ecosystems, the response of heathland butterflies to succession had not been investigated so far. To address this issue we sampled butterfly abundance and environmental parameters on the Baltic island of Hiddensee (NE Germany) along a gradient of coastal heathland succession from grey dunes to birch forest. Our results provide evidence that succession of coastal heathland has a strong influence on butterfly diversity, abundance, and species composition. Thereby grass and tree encroachment present the main threats for heathland butterflies. Diversity and abundance of butterflies were highest in shrub-encroached heath directly followed by early stages of coastal heathland succession (dwarf-shrub heath, grey dune). Both observed threatened species (Hipparchia semele, Plebeius argus) were negatively affected by succession: abundance decreased with increasing vegetation density (both species) and grass cover (P. argus); consequently, the two later successional stages (shrub, birch forest) were not occupied. Our findings highlight the importance of the preservation of early stages of coastal heathland succession for endangered butterfly species. For coastal heathland management we therefore suggest to maintain early successional stages by sheep grazing, mowing or, in case of high nutrient contents, intensive techniques such as sod-cutting or choppering. To a lower extend shrub-encroached sites should also be present, which might be beneficial for overall species richness.     

Plant strategies and secondary succession on Brittany heathlands after severe fire

Abstract. Plant succession on heathlands subjected to major fire disturbance and humus burn in 1976 was studied over twelve years following burning. Life history strategies of principal heathland species are described with reference to concepts outlined by Grime (1979) and Whittaker & Goodman (1979). Heathlands and closely related communities are characterised by dominance of speciestolerant of physical stress (‘S’ strategists) whereas species which colonise disturbed sites are closer to rude-rals (‘R’ strategists). After severe burning three main successional patterns were identified. They depend on water and nutrient availability relative to temporal population dynamics. Recovery of heathland is often retarded due to monospecific dominance, e.g. of Polytrichum commune, Molinia caerulea and Betula pubescens. These patterns of secondary succession illustrate the inhibition model advanced by Connell & Slatyer (1977).     

Leaf-nutrient accumulation and turnover at three stages of succession from heathland to forest

Abstract. Accumulation of nutrients in leaves of the dominating species of three ecosystems, characterizing the secondary succession from Genisto-Callunetum heathland through Leucobryo-Pinetum birch-pine woodland to mature Querco-Fagetum oak-beech forest, as well as nutrient turnover within these ecosystems was studied. The objective of the study was to establish potential variations in quantity and quality of nutrient supply to the plants with respect to succession dynamics. The results show very low leaf nutrient concentrations of all species investigated, coinciding with low nutrient availability in the soil. However, the nutrient content of leaves and leaf litter of Quercus petraea and Fagus sylvatica, which dominate the late succession stages, and in Betulapéndula are higher than in the photosynthetic organs (leaves and young shoots) of Calluna vulgaris and Pinus sylvestris. The combination of the higher nutrient content of the leaves and an increasing leaf-litter production during succession results in an increased nutrient turnover via leaf-litter fall. However, due to the high leaf biomass, the storage of nutrients in the leaf biomass is highest within the birch-pine woodland. From this, it may be assumed that the low demand and the low loss of nutrients via leaf-litter fall are favourable for Pinus at the early stages of forest succession on poor sandy soils. In contrast, Quercus and Fagus are provided with better growth conditions at the later stages of succession resulting from the accumulation of plant-available nutrients in the ecosystem by Pinus sylvestris, combined with a higher nutrient turnover as compared with the heathland.     

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.     

Does prescribed burning result in biotic homogenization of coastal heathlands?

Biotic homogenization due to replacement of native biodiversity by widespread generalist species has been demonstrated in a number of ecosystems and taxonomic groups worldwide, causing growing conservation concern. Human disturbance is a key driver of biotic homogenization, suggesting potential conservation challenges in seminatural ecosystems, where anthropogenic disturbances such as grazing and burning are necessary for maintaining ecological dynamics and functioning. We test whether prescribed burning results in biotic homogenization in the coastal heathlands of north‐western Europe, a seminatural landscape where extensive grazing and burning has constituted the traditional land‐use practice over the past 6000 years. We compare the beta‐diversity before and after fire at three ecological scales: within local vegetation patches, between wet and dry heathland patches within landscapes, and along a 470 km bioclimatic gradient. Within local patches, we found no evidence of homogenization after fire; species richness increased, and the species that entered the burnt Calluna stands were not widespread specialists but native grasses and herbs characteristic of the heathland system. At the landscapes scale, we saw a weak homogenization as wet and dry heathland patches become more compositionally similar after fire. This was because of a decrease in habitat‐specific species unique to either wet or dry habitats and postfire colonization by a set of heathland specialists that established in both habitat types. Along the bioclimatic gradient, species that increased after fire generally had more specific environmental requirements and narrower geographical distributions than the prefire flora, resulting in a biotic ‘heterogenisation’ after fire. Our study demonstrates that human disturbance does not necessarily cause biotic homogenization, but that continuation of traditional land‐use practices can instead be crucial for the maintenance of the diversity and ecological function of a seminatural ecosystem. The species that established after prescribed burning were heathland specialists with relatively narrow geographical ranges.     

Age structure and dynamics of Patagonian beech forests in Torres del Paine National Park,Chile

This study documents the stem size and age-structure in forests dominated by different species of Nothofagus in Torres del Paine National Park (51° S), in the Chilean Patagonian region. We also explored the relationship between the various types of Nothofagus forest and postglacial succession. Pioneer stands on moraine fields 1–2 km of the glacier front are dominated by Nothofagus betuloides and Nothofagus antarctica. Moraines appear to be first colonized by the evergreen N. betuloides, followed within 5–7 years by deciduous N. antarctica. Nothofagus antarctica may replace the former species and develop monospecific stands on glacial valleys. Most trees in the N. antarctica stand studied were older than 40 years and floods may cause a significant mortality of young trees. Recruitment from seed seems to be infrequent. Old-growth stands dominated by deciduous Nothofagus pumilio occupy more stable substrates, and probably represent the last stage of postglacial succession. This long-lived tree species had recorded ages over 200 years. The canopy of N. pumilio forests appears to be a mosaic of even-aged, old-growth patches. We propose that regeneration episodes follow the blowdown of a large portion of the canopy, with long intervals with little or no regeneration. Windstorms may be an important force influencing the regeneration of N. pumilio. Exogenous disturbances, such as floods and windstorms, are an integral part of the forest cycle in the Patagonian region.     

Resource availability at three presumed stages of a heathland succession on the Lüneburger Heide,Germany

Abstract. The availability of maj or plant resources was investigated in three vegetation types that were assumed to represent different stages of a secondary succession on heathland on the Lüneburger Heide, northwestern Germany. Canopy transmission and absorption of photosynthetically active radiation (PAR), soil-water availability, and nutrient (Ca, K, Mg, N, P) availability were monitored with high spatial and temporal resolution in (1) a Calluna vulgaris heathland, (2) a pioneer birch-pine forest and (3) a late-successional oak-beech forest, situated close to each other on comparable geological substrate (diluvial). Mean fractional transmission of PAR during summer decreased from 0.48 in the heathland to 0.04 in the oak-beech forest while the fractional canopy absorption increased from 0.49 to 0.92. Soil-water availability as indicated by the soil-water potential, was significantly influenced by differential canopy interception loss and characteristic rooting patterns in the three vegetation types. Annual mean nutrient concentrations in the equilibrium soil solution were similar or, for some elements, increased from the heathland to the birch-pine and the oak-beech forest despite a growing demand. A marked increase was found for the total nutrient pools in the soil-organic layer between early and late successional vegetation types. On the poor glacial parent material, nutrient pools seem to be strongly dependent on stand productivity and litter production which increased with succession. Thus, for nutrients, facilitation seems to be important in this type of succession.     

Massive floral display affects insect visits but not pollinator‐mediated pollen transfer in Rhododendron ferrugineum

Fragmentation of natural vegetation creates one of the largest threats to plant–pollinator interactions. Although fragmentation impacts on plant populations have been explored in many, mainly herbaceous, species, the response of wild mass‐flowering species is poorly known. Here, we studied 28 heathland patches dominated by the mass‐flowering shrub Rhododendron ferrugineum, each presenting different R. ferrugineum floral display sizes (total inflorescence number per patch) and patch isolation (median distance to the three nearest patches). We assessed the impacts of these two factors on (i) heathland patch visitor assemblage (considering R. ferrugineum versus surrounding community) and (ii) R. ferrugineum flower visitation rate and pollen transfer limitation (comparing seed set from emasculated to pollen‐supplemented flowers). We found that diversity and abundance of bees visiting R. ferrugineum in heathland patches significantly decreased with decreasing R. ferrugineum floral display, while overall visitor density per patch and flower visitation rate increased. Moreover, a decrease in massive floral display and increase in patch isolation resulted in reduced visitor density in the surrounding community. Even in patches with few individuals, we found disproportionate visitor abundance in R. ferrugineum compared to the surrounding community. Finally, pollen transfer limitation in R. ferrugineum was neither affected by visitation rate nor by patch attributes. By disproportionally attracting pollinators from co‐flowering species, and probably promoting geitonogamous pollen transfer, the mass‐flowering trait appears adequate to compensate, in terms of conspecific pollen transfer, for the decrease in visitor diversity and abundance and in mate availability, which usually result from population fragmentation.     

Limited by the host: Host age hampers establishment of holoparasite Cuscuta epithymum

A good understanding of the relationship between plant establishment and the ecosystem of which they are part of is needed to conserve rare plant species. Introduction experiments offer a direct test of recruitment limitation, but generally only the seed germination and seedling phases are monitored. Thus the relative importance of different establishment stages in the process of recruitment is not considered. This is particularly true for parasitic plants where empirical data are generally missing. During two consecutive growing seasons we examined the effect of heathland management applications, degree of heathland succession (pioneer, building and mature phase) and seed-density on the recruitment and establishment of the endangered holoparasite Cuscuta epithymum. In general, recruitment after two growing seasons was low with 4.79% of the sown seeds that successfully emerged to the seedling stage and a final establishment of 89 flowering adults (i.e. <1.5% of the sown seeds). Although a higher seed-density resulted in a higher number of seedlings, seed-density did not significantly affected relative germination percentages. The management type and subsequent heath succession had no significant effect on seedling emergence; whereas, seedling attachment to the host, establishment and growth to full-grown size were hampered in older heath vegetation (i.e. high, dense, and mature canopy). Establishment was most successful in turf-cut pioneer heathland, characterised by a relatively open and low vegetation of young Calluna vulgaris. The age of C. vulgaris, C. epithymum's main host, proved to be the most limiting factor. These results emphasise the importance of site quality (i.e. successional phase of its host) on recruitment success of C. epithymum, which is directly affected by the management applied to the vegetation. Lack of any heathland management will thus seriously restrict establishment of the endangered parasite.     

Genetic erosion in a stenotopic heathland ground beetle (Coleoptera: Carabidae): a matter of habitat size?

Two centuries ago large areas of north-west Europe were covered by coherent heathlands which hosted numerous specialized species. Changes in land use made heathlands fragmented and rare, consequently, they are in the focus of nature conservation efforts today. But how large should remaining heathland patches be in order to secure the survival of populations of specialized species? We investigated the genetic diversity at five allozyme loci of Poecilus lepidus, a flightless and stenotopic heathland ground beetle. 29 populations from differently sized heathland patches in north-west Germany were analyzed. Results show a weak but significant genetic differentiation and no evidence for isolation by distance or other patterns of spatial autocorrelation. Linear regression analysis revealed significant relationships between patch size, allelic richness, number of alleles and expected heterozygosity. These findings are explained by severe habitat fragmentation together with strong fluctuations in population size which have been reported for this species in the past. To conserve the vast majority of the species’ genetic diversity for a period of 100 years we suggest to maintain heathland patches of at least 50 ha in size.     

Two centuries of vegetation succession in an inland sand dune area,central Netherlands

Questions: (a) What are the rates and directions of vegetation succession in an inland sand‐dune system? (b) What are the differences in successional trajectories in different relief types? and (c) Is it possible to preserve the last areas of still active dunes and under what circumstances? Location: The study sites were located in the northern part of the Veluwe Region, central Netherlands; longitude 5°44′ E, latitude 52°20′ N, altitude 9 to 24 m a.s.l. Methods: Vegetation and relief mapping was conducted in three permanent plots, 200 m × 200 m in size, in 1988 and 2003. Phytosociological relevés (2400) were recorded in each 10 m × 10 m subplots. Age of woody species was determined by wood coring. Geographic Information System, ordination analyses, and TWINSPAN were used for data exploration and elaboration. Results: A total of 70 vascular plants and 19 bryophytes were recorded over successional stages spanning approximately190 years. The following dominant species formed the sequence of successional stages, but not all participated in all relief types: Ammophila arenaria, Festuca arenaria, Corynephorus canescens, Festuca ovina and Agrostis capillaris, and pine forest dominated in its herb layer at first by Deschampsia flexuosa and later by either Empetrum nigrum, Vaccinium myrtillus or Vaccinium vitis‐idaea. Conclusions: The successional trajectory is basically unidirectional for more than 100 years; no clear multiple successional pathways were observed, as is frequent in coastal dunes. Successional divergence was observed after approximately 130 years in the composition of the herb layer in the closed pine forest. The obvious vegetation heterogeneity in the still active sand‐blown area is related to differences in timing of vegetation establishment on particular relief types, thus the succession exhibits a terrain‐dependent asynchronous character. We conclude that the last patches of still‐active sand dunes can be preserved only by repeated strong artificial disturbances.     

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.     

中国苦苣苔科植物的多样性与地理分布（附表）

物种多样性编目是开展生物多样性保护的重要基础,该研究结合最新分子系统学研究成果以及近年来发表的新资料,对中国苦苣苔科植物多样性和地理分布数据进行了统计和分析.结果表明:中国苦苣苔科植物共有44属671种(含种下单位,下同),其中特有属11个;特有种573种,占总种数的85.39%.种数最多的10个属依次为广义报春苣苔属(180种)、广义马铃苣苔属(122种)、石蝴蝶属(39种)、半蒴苣苔属(39种)、芒毛苣苔属(38种)、长蒴苣苔属(35种)、石山苣苔属(31种)、吊石苣苔属(31种)、蛛毛苣苔属(28种)、汉克苣苔属(22种).在地理分布上,种数排名前10的省份(区)有广西(260种,33属)、云南(236种,30属)、贵州(96种,28属)、广东(93种,17属)、四川(85种,21属)、湖南(58种,13属)、西藏(39种,9属)、湖北(29种,15属)、福建(26种,13属)、江西(25种,9属).含中国特有苦苣苔科植物的属中排前10位的分别为广义报春苣苔属(178种)、广义马铃苣苔属(119种)、石蝴蝶属(37种)、半蒴苣苔属(35种)、石山苣苔属(30种)、长蒴苣苔属(29种)、吊石苣苔属(23种)、蛛毛苣苔属(19种)、芒毛苣苔属(19种)、汉克苣苔属(11种).这表明中国南部和西南部是苦苣苔科植物的一个分布中心,特别是石灰岩地区有着高度的物种多样性和特有性,广义报春苣苔属、广义马铃苣苔属、石蝴蝶属、半蒴苣苔属、石山苣苔属、吊石苣苔属等为我国典型的优势属.此外,根据目前的研究现状,还对我国苦苣苔科植物资源的调查、分类学和系统发育研究、保护和可持续利用等进行了讨论.     

Comparative seed morphology and character evolution in the genus Lysimachia (Myrsinaceae) and related taxa

Summary We investigated seed morphology in 34 species of the genus Lysimachia and in 14 species and two subspecies of six additional genera (Anagallis, Ardisiandra, Asterolinon, Glaux, Pelletiera, Trientalis), which have been shown to be closely related to, or are placed within Lysimachia in previous molecular studies. We studied seed shape, seed coat structure, and seed coat surface patterns. Three major types of seed shape were identified: (1) sectoroid, (2) polyhedral, and (3) coarsely rugose with concave hilar area. In addition, seeds may be keeled or winged. The outer layer of the seed coat is either sponge-like and adhering only loosely to the inner seed coat or it is thin and tightly adhering to the underlying tissue. Seed surface patterns can be divided into six main types: (1) reticulate, (2) tuberculate, (3) vesiculose, (4) colliculate, (5) undulate, or (6) poroid-alveolate. Seed surface patterns are mostly congruent with molecular phylogenetic relationships. A reticulate surface pattern is diagnostic of, e.g. Lysimachia subgenera Palladia and Hawaiian Lysimachiopsis. Mapping seed characters onto a recent phylogenetic tree, reveals that they provide potentially synapomorphic character states for various subclades of Lysimachia. Salient examples include a rugose seed shape, which turns out to be synapomorphic for the clade comprising the genus Pelletiera plus Asterolinon linum-stellatum and a sponge-like outer seed coat layer, which characterizes a clade with Lysimachia vulgaris, L. thyrsiflora, and L. terrestris, with an analogue that apparently evolved in parallel in Trientalis europaea. We also discuss possible habitat factors that may have favored the independent evolution of particular seed types such as winged seeds in various lineages.     

The age of Calluna stands moderates post‐fire regeneration rate and trends in northern Calluna heathlands

Questions: Does stand age influence the direction and rate of post‐fire successional dynamics in coastal Calluna heaths and can old degraded heath vegetation be restored through reintroduction of fire? Location: Coastal heaths in the Tarva archipelago, central Norway. Methods: We investigated revegetation dynamics after experimental fires set in young (8 years since last fire) and old (>50 years since last fire) grazed heath stands. A repeated measures design was used, with floristic data recorded in permanent plots in the post‐fire successions (n=12) over a 7‐year period. The data were analysed using multivariate ordination techniques (PCA, RDA and PRC) and mixed effects models. Results: The age of Calluna stands strongly influenced post‐fire succession, different trends due to age explained 10.4% of variation in floristic data. Young heath showed faster succession towards pre‐fire community composition than old heath, and this could partially be explained by succession‐related factors: young heath had lower cover of mosses and lichens in the pre‐burned vegetation, and lower cover of litter early in succession. Young heath had a less pronounced overall community response to fire than old heath. Vegetative regeneration of C. vulgaris was absent in both old and young heath, but Calluna still re‐established as the dominant species within 5–7 years in both young and old stands. Regeneration dynamics were also affected by habitat conditions, different trends due to habitat explained 6% of variation. Conclusions: Our study demonstrates that old stands do develop characteristic heathland vegetation and structure after fire, and while potential invasives into the system such as trees and rhizomatous species are present, they do not impair Calluna regeneration or vegetation development towards the target heathland community composition and structure. Further, as our young stands are only in their second fire rotation after restoration, we suggest that characteristic dynamics of managed heathlands can re‐establish relatively rapidly, even in severely degenerated sites (>50 years since last fire). Site‐specific factors also need to be considered. We conclude that there is restoration potential in old heaths, despite slow dynamics in the first rotation.     

An ericoid shrub plays a dual role in recruiting both pines and their fungal symbionts along primary succession gradients

Relative importance of positive and negative interactions between plant species may change along disturbance and resource gradients. Positive interactions are suggested to prevail in low resource, low productivity (high stress) conditions and negative interactions in high resource availability. A dwarf shrub, mountain crowberry Empetrum nigrum ssp. hermaphroditum, is known to have allelopathic impacts on both Scots pine Pinus sylvestris and its ectomycorrhizal symbionts. We aimed to study if the outcome of Empetrum impacts on Scots pine changes along primary succession gradients on the dune shores of Bothnian Bay, in Finland, where abiotic stress gradually changes to biotic stress along the succession. We found that Empetrum may act as a facilitator despite its allelopathic effects, since the proportion of Scots pine seedlings established in Empetrum patches was higher than in patches without the shrub in early and mid succession stages, whereas patches without Empetrum were preferred in late succession. The amount of mycelial fungal biomass (ergosterol) in the soil in the vicinity of the seedling roots was higher in Empetrum patches than in patches without Empetrum and it increased along the succession gradient. Proportion of pine root tips colonised by suilloid morphotypes with abundant external mycelia and the diversity of ectomycorrhizal morphotypes were higher in mid successional stage in Empetrum patches compared to patches without Empetrum. Our results suggest that in the harsh physical conditions of the dune shore Empetrum facilitates pine seedling establishment in the early and mid stages of succession by providing mechanical and physical shelter whereas in late succession negative interactions (competition and allelopathy) between the shrub and the pine are dominating. To our knowledge we present the first finding that an ericoid mycorrhizal shrub could enhance both the performance of the ectomycorrhizal host tree and the tree's fungal symbionts.     

Effects of prescribed burning on plant available nutrients in dry heathland ecosystems

Heathland management is an important tool with which to modify ecosystem impacts caused by atmospheric nutrient deposition. Since changes in nutrient availability as a result of management measures affect the outcomes of heathland succession and species competition, studies on this issue are important from both a nature conservation and management point of view. This study reports the effects of prescribed burning on nutrient availability in dry heathland soils and the nutrient content of the two competing heathland species Calluna vulgaris and Deschampsia flexuosa, with particular reference to N and P. We hypothesise that winter prescribed burning leads to additional N availability, which enhances the importance of P in the context of nutrient limitation in heathland ecosystems. In the nature reserve “Lueneburg Heath” (NW Germany) we examined the availability of nutrients in the humus horizons and in the leachate as well as the relevant C:element ratios in Calluna and Deschampsia before and after a burning experiment. Our results show that prescribed burning resulted in drastically increased NH₄⁺ availability in the O-horizon. We observed only short-term effects (for NO₃⁻, PO₄³⁻, Mg) and insignificant effects on the availability of other nutrients (K, Ca). As a consequence of an increased nutrient availability in the humus horizons and a limited nutrient uptake by plants after burning, leaching increased significantly for N, Ca, K, and Mg after burning treatment. No significant changes were found in the foliar C:N ratios for either species after prescribed burning, although Deschampsia showed an increased deficiency for all the other nutrients, particularly for P, as expressed by increased foliar C:P and N:P ratios. By contrast, the nutrient content of Calluna did not change significantly, suggesting that prescribed burning favours the competitive capacity of Calluna as against Deschampsia. We assume that water shortage as a result of changes in the microclimate was mainly responsible for the deterioration of the nutrient content of Deschampsia. This gives Calluna a competitive advantage, enabling it to out-compete Deschampsia on burned heathlands, with respect to the key factor P-limitation.