Impacts of succession and grass encroachment on heathland Orthoptera

For the conservation of biodiversity, heathlands present important ecosystems throughout Europe. The formerly widespread habitats are nowadays restricted to small and isolated remnants. Without land use heathland vegetation undergoes succession and, in addition, the increasing amount of atmospheric nitrogen deposition has resulted in an encroachment of grasses. In the present study we analysed the effects of succession and grass encroachment on Orthoptera in a coastal heathland on the Baltic island of Hiddensee, Germany. Vegetation, microclimate, soil humidity and Orthoptera were sampled in the five main stages of heathland succession, namely grey dunes, dwarf-shrub heath, grassy heath, heath with shrubs, and birch forest. Vegetation and environmental parameters showed strong differences among the successional stages. Orthoptera species richness was highest in transitional stages. The high proportion of grasses offer favourable habitat conditions for graminivorous, chorto- and thamnobiont species. Orthoptera density was highest in grey dunes. Threatened and specialised species were restricted to the young stages grey dunes and dwarf-shrub heath. Hence, in order to maintain a high diversity of Orthoptera in heathlands, maintaining different successional stages is of critical importance and this should be integrated into heathland management practices.     

Colonization and succession in deep-sea ecosystems

Experimental studies reveal generally slow but variable colonization rates in deep-sea soft bottoms. Species successions following biological disturbance appear to be complex and unpredictable, potentially playing a role in structuring these diverse communities. In contrast, physical processes probably play a prominent role in the rapid species turnovers observed at hydrothermal vents. Better information concerning disturbance regimes, recruitment processes and resource utilization is required to elaborate successional mechanisms in both soft-bottom and vent environments.     

Markov models in the study of post-fire succession in heathland communities

Data on the presence of a number of vegetation states (defined in terms of species dominance in areas of 10×10 cm) and transition probabilities were derived from permanent quadrats in a number of recently burned heath stands. Data were taken from a species-rich community, a species-poor type and a high-level Calluna-Eriophorum bog. Simple Markovian models were constructed using these data, and the model predictions were compared with known or expected trends. Models for species-rich heath yielded poor simulations of expected trends since matrices derived from data for the first years after fire did not contain sufficient information on transitions to states important later in the developmental sequence. Model results for the simpler species-poor and bog communities were more satisfactory and simulated expected trends. In these types all species recovered quickly after fire and less rearrangement of species abundances took place. Maximum likelihood statistics carried out on the transition matrices produced inconclusive results for the species-rich and species-poor types, but indicated that the data from the Calluna-Eriophorum bog approximated a first-order time-homogeneous Markov chain. It was concluded that Markov models lack predictive ability except in relatively simple systems, but that they may be useful in illustrating variations in short-term community dynamics.Nomenclature follows Tutin et al. (1964–80) for vascular plants and Smith (1978) for mosses.I am grateful to Prof. C. H. Gimingham for help and advice during this study, which was carried out during tenure of a Natural Environment Research Council studentship. Thanks are also due to Dr M. B. Usher and an anonymous referee for their comments on the draft.     

Soil seed bank dynamics in post-fire heathland succession in south-eastern Australia

Soil seed banks can exert a strong influence on the path of vegetation succession following fire, with species varying in their capacity to persist in the seed bank over time, leading to changes in seed bank composition and propagules available for post-fire colonisation. This study examined the effect of time since fire on soil seed bank dynamics in a chronosequence of seven sites spanning 26 years in a south-eastern Australian sand heathland. No significant change was evident in the species richness and density of the germinable soil seed bank, but species composition differed significantly among young (0–6 years since fire), intermediate (10–17 years since fire) and old-aged (24–26 years since fire) sites (using presence/absence data). No significant trend was observed in the similarity between the extant vegetation and the soil seed bank with time since fire. A total of 32% of the species recorded in the soil seed bank were not present in the above-ground vegetation at the same site, which suggests that species requiring fire for germination may be present in the seed bank. Most species present in the extant vegetation were not recorded (63%) or were in very low abundances in the soil seed bank (29%). The mode of regeneration appears to be the major determinant of species absence in the soil seed bank, as 66% of species occurring in the extant vegetation but not in the seed bank have the capacity to regenerate by resprouting. This study shows that a major shift in the successional pathway after fire due to altered seed bank composition is unlikely in this vegetation; most species not recorded in the seed bank are either resprouters (obligate or facultative) or serotinous, suggesting that they will readily regenerate following fire. Unless fire frequencies are high and kill fire-sensitive obligate seeders before they reach maturity, the chance that the soil seed bank could substantially alter vegetation composition within the study area after fire is low. However, it is unclear how successional pathways may alter in response to severe fires with the potential to kill both seeders and resprouters.     

A comparative study on nutrient cycling in wet heathland ecosystems

Summary The concept of the relative nutrient requirement (L _n) that was introduced in the first paper of this series is used to analyse the effects of the dominant plant population on nutrient cycling and nutrient mineralization in wet heathland ecosystems. A distinction is made between the effect that the dominant plant species has on (1) the distribution of nutrients over the plant biomass and the soil compartment of the ecosystem and (2) the recirculation rate of nutrients. The first effect of the dominant plant species can be calculated on the basis of the /k ratio (which is the ratio of the relative mortality to the decomposition constant). The second effect can be analysed using the relative nutrient requirement (L _n). The mass loss and the changes in the amounts of N and P in decomposing above-ground and below-ground litter produced by Erica tetralix and Molinia caerulea were measured over three years. The rates of mass loss from both above-ground and below-ground litter of Molinia were higher than those from Erica litter. After an initial leaching phase, litter showed either a net release or a net immobilization of nitrogen or phosphorus that depended on the initial concentrations of these nutrients. At the same sites, mineralization of nitrogen and phosphorus were measured for two years both in communities dominated by Molinia and in communities dominated by Erica. There were no clear differences in the nitrogen mineralization, but in one of the two years, phosphate mineralization in the Molinia-community was significantly higher. On the basis of the theory that was developed, mineralization rates and ratios between amounts of nutrients in plant biomass and in the soil were calculated on the basis of parameters that were independently measured. There was a reasonable agreement between predicted and measured values in the Erica-communities. In the Molinia-communities there were large differences between calculated and measured values, which was explained by the observation that the soil organic matter in these ecosystems still predominantly consisted of Erica-remains.     

A comparative study on nutrient cycling in wet heathland ecosystems

Summary The term relative nutrient requirement is introduced in order to measure and to compare the nutrient losses from different perennial plant populations and the amount of nutrient that they need to absorb to compensate these losses. The relative nutrient requirement (L) is defined as the amount of the growth-limiting nutrient that must be taken up to maintain or replace each unit of biomass during a given time interval (e.g., mgN g^-1 biomass year^-1). It is derived that in a plant community with two competing perennial plant populations, species¹ will become dominant if the relative competition coefficient k ₁₂ (sensu De Wit 1960) exceeds the ratio between the relative nutrient requirements of the two species (L ₁/L ₂), whereas species 2 will become dominant, if k ₁₂ is below this critical ratio. The above-ground litter production was measured inwet heathland communities dominated by Erica tetralix or by Molinia caeruleain order to estimate N and P losses from theaboveground biomass and to calculate the relative N and P requirements of these species. Molinia lost during one year 63% and 34%, respectively, of the amount of N and P present in the above-ground biomass at the end of the growing season. These losses were in Erica 27% and 31%, respectively. The relative N requirements of the two species show the same difference: 7.5 and 2.6 mg N g^-1 yr^-1, respectively, in Molinia and in Erica. Also the relative P requirement of Molinia is higher as well as that of Erica (0.18 versus 0.08 mg P g^-1 yr^-1). The relative competition coefficient of Molinia with respect to Erica (k _me ) is equal to unity under unfertilized conditions but increases with increasing nutrient supply. Under nutrient-poor conditions k _me is below the critical ratio of the relative nutrient requirements of the two species (L _m /L _e =2.9 or 2.3), so that Erica will be the dominant species. After an increase in nutrient availability k _me increases and exceeds this critical limit which results in Molinia replacing Erica. During the last 20 years this replacement of Erica-dominant communities by monocultures of Molinia has been observed in almost all wet heathlands in The Netherlands along with a strong increase in nitrogen availability.     

Nitrogen mineralization in heathland ecosystems dominated by different plant species

Net N mineralization rates were measured in heathlands still dominated by ericaceous dwarf shrubs (Calluna vulgaris or Erica tetralix) and in heathlands that have become dominated by grasses (Molinia caerulea or Deschampsia flexuosa). Net N mineralization was measuredin situ by sequential soil incubations during the year. In the wet area (gravimetric soil moisture content 74–130%), the net N mineralization rates were 4.4 g N m^–2 yr^–1 in the Erica soil and 7.8 g N m^–2 yr^–1 in the Molinia soil. The net nitrification rate was negligibly slow in either soil. In the dry area (gravimetric soil moisture content 7–38%), net N mineralization rates were 6.2 g N M-2 yr^–1 in the Calluna soil, 10.9 g N m^–2 yr^–1 in the Molinia soil and 12.6 g N m^–2 yr^–1 in the Deschampsia soil. The Calluna soil was consistently drier throughout the year, which may partly explain its slower mineralization rate. Net nitrification was 0.3 g N m^–2 yr^–1 in the Calluna soil, 3.6 g N m^–2 yr^–1 in the Molinia soil and 5.4 g N m^–2 yr^–1 in the Deschampsia soil. The net nitrification rate increased proportionally with the net N mineralization rate suggesting ammonium availability may control nitrification rates in these soils. In the dry area, the faster net N mineralization rates in sites dominated by grasses than in the site dominated by Calluna may be explained by the greater amounts of organic N in the soil of sites dominated by grasses. In both areas, however, the net amount of N mineralized per gram total soil N was greater in sites dominated by Molinia or Deschampsia than in sites dominated by Calluna or Erica. This suggests that in heathlands invaded by grasses the quality of the soil organic matter may be increased resulting in more rapid rates of soil N cycling.     

Phylogenetic and functional dissimilarity does not increase during temporal heathland succession

Succession has been a focal point of ecological research for over a century, but thus far has been poorly explored through the lens of modern phylogenetic and trait-based approaches to community assembly. The vast majority of studies conducted to date have comprised static analyses where communities are observed at a single snapshot in time. Long-term datasets present a vantage point to compare established and emerging theoretical predictions on the phylogenetic and functional trajectory of communities through succession. We investigated within, and between, community measures of phylogenetic and functional diversity in a fire-prone heathland along a 21 year time series. Contrary to widely held expectations that increased competition through succession should inhibit the coexistence of species with high niche overlap, plots became more phylogenetically and functionally clustered with time since fire. There were significant directional shifts in individual traits through time indicating deterministic successional processes associated with changing abiotic and/or biotic conditions. However, relative to the observed temporal rate of taxonomic turnover, both phylogenetic and functional turnover were comparatively low, suggesting a degree of functional redundancy among close relatives. These results contribute to an emerging body of evidence indicating that limits to the similarity of coexisting species are rarely observed at fine spatial scales.     

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.     

Effects of restoration measures on plant communities of wet heathland ecosystems

Question: Which are the success and failure of restoration measures, particularly sod‐cutting and hydrological measures, in small wetlands on mineral soils in The Netherlands. Location: Twente, in the eastern part of The Netherlands. Methods: Success or failure of restoration measures has been assessed by comparing experimental plots with that in reference plots for (1) species richness, (2) re‐establishment of endangered species and (3) species composition, including life forms and Red List species. In total 119 samples were taken in 42 permanent plots in fen meadows, small‐sedge marshes, wet heathlands, and soft‐water pools. Topsoil samples were analysed for pH, Ca, Mg, Na, K and Cation Exchange Capacity. Gradient analysis was carried out by means of Canonical Correspondence Analysis. Results: Sod‐cutting, in combination with re‐wetting measures (infilling of ditches and drains), led to restoration of communities of soft‐water pools and small sedge marshes within five years. This rapid recovery is probably related to the presence of persistent seed banks of the component species of these communities. Complete restoration of fen meadows took longer (10–15 yr). Apparently, many species of fen meadows have short‐lived seed banks. Sod‐cutting of a degraded wet heathland and a soft‐water pool was only successful temporarily, probably as a consequence of low water tables. Conclusions: To prevent depletion of (persistent) soil seed banks, sod‐cutting in nutrient‐poor wetlands is not recommended in areas where the groundwater regime and the base status of the soil can not be restored to levels required by plant communities of wet heathlands.     

Fate of airborne nitrogen in heathland ecosystems: a 15N tracer study

In the present study, we analyze the fate of airborne nitrogen in heathland ecosystems (NW Germany) by means of a ¹⁵N tracer experiment. Our objective was to quantify N sequestration and N allocation patterns in an ecosystem that is naturally limited by N, but that has been exposed to airborne N inputs exceeding critical loads for more than 3 decades. We hypothesized that the system has a tendency towards N saturation, which should be indicated by low N sequestration and high N leaching. We analyzed ¹⁵N partitioning (aboveground biomass and soil horizons) and investigated ¹⁵N leaching over 2 years following a ¹⁵N tracer pulse addition. ¹⁵N tracer recovery was 90% and 76% in the first and second year, respectively. Contrary to our expectations, more than 99% of the tracer recovered was sequestered in the biomass and soil, while leaching losses were <0.05% after 2 years. Mosses were the most important short‐term sink for ¹⁵N (64% recovery in the first year), followed by the organic layer. In the second year, the moss layer developed from a sink to a source (23% losses), and soil compartments were the most important sink (gains of 11.2% in the second year). Low ¹⁵N recovery in the current year's shoots of Calluna vulgaris (<2%) indicated minor availability of ¹⁵N tracer sequestered in the organic layer. N partitioning patterns showed that the investigated heaths still have conservative N cycling, even after several decades of high N loads. This finding is mainly attributable to the high immobilization capacities for N of podzols in soil compartments. In the long term, the podzol A‐ and B‐horizons in particular may immobilize considerable amounts of incoming N. Since N compounds of these horizons are not readily bio‐available, podzols have a high potential to withdraw airborne N from the system's N cycle.     

Interspecific competition: A mechanism for rodent succession after fire in wet heathland

Abstract A competitive interaction between two species of sympatric native rodents was demonstrated experimentally in Myall Lakes National Park (NSW, Australia). The previously documented replacement of Pseudomys gracilicaudatus by Rattus lutreolus in areas of wet heath in this region implied that competition may be the mechanism facilitating this succession. We present experimental evidence to support this suggestion. Removal of the larger R. lutreolus caused a significant increase in abundance of P. gracilicaudatus on five experimental sites in comparison to five unmanipulated control sites. The sites used in this experiment were chosen from three ages of wet heath regenerating after fire. These three ages represent key stages in the replacement of P. gracilicaudatus by R. lutreolus and the occurrence of competition in each of these ages indicates that competition plays an important role in this succession. In the initial stages of the experiment when R. lutreolus were removed they were replaced by new R. lutreolus individuals. This has been interpreted as evidence of strong intraspecific competition. After removal of R. lutreolus, P. gracilicaudatus expanded its habitat range into microhabitats that had formerly been occupied by R. lutreolus and reduced its range in microhabitats previously occupied by P. gracilicaudatus. This leads us to believe that P. gracilicaudatus was occupying inferior microhabitat before the removal of R. lutreolus, which had excluded it from more preferable microhabitat.     

Nutrient leaching in dry heathland ecosystems: effects of atmospheric deposition and management

Atmospheric nutrient deposition has contributed to widespread changes in sensitive seminatural ecosystems throughout Europe. For an understanding of underlying processes it is important to quantify input–output flows in relation to ongoing atmospheric inputs and current management strategies. In this study we quantified losses of N, P, Ca, Mg, and K via leaching in heathland ecosystems (Lüneburger Heide, NW Germany) as a function of current deposition rates and different management measures (mowing, prescribed burning, choppering, sod-cutting) which aim to prevent shrub and tree encroachment. Leaching was only moderately related to atmospheric input rates, indicating that leaching was mostly affected by internal turnover processes. Leaching significantly increased for most of the nutrients after the application of management measures, particularly in the choppered and sod-cut plots. However, atmospheric nutrient inputs exceeded leaching outputs for most of the nutrients, even in the plots subjected to management. Despite high deposition rates (20–25 kg N ha⁻¹ year⁻¹), retention of atmospheric N input ranged between 74% and 92% in the control plots. In the treated plots, N retention decreased to 59–80%. However, in the study area mean N leaching in the controls has almost doubled since 1980 and currently amounts to 3.7 kg ha⁻¹ year⁻¹, indicating an early stage of N saturation. Our study provides evidence that leaching did not compensate for atmospheric nutrient deposition, particularly as regards N. Management, thus, will be an indispensable tool for the maintenance of the low-nutrient status as a prerequisite for the long-term preservation of heathland ecosystems.     

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.     

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.     

Ecological succession and the rehabilitation of disturbed terrestrial ecosystems

Plant and Soil - Soil and vegetation development on surface-mined coal sites in a mixed grass prairie region were studied as (a) naturally revegetated chronosequences of 1, 7, 17, 30 and 45-year...     

Methanogen communities along a primary succession transect of mire ecosystems

Peat accumulating mires are important sources of the greenhouse gas methane. Methane emissions and methanogenic Archaea communities have been shown to differ between fens and bogs, implying that mire succession includes an ecological succession in methanogen communities. We investigated methane production and the methanogen communities along a chronosequence of mires (ca. 100-2,500 years), which consisted of five sites (1-5) located on the land-uplift coast of the Gulf of Bothnia. Methane production was measured in a laboratory incubation experiment. Methanogen communities were determined by amplification of a methyl coenzyme M-reductase (mcr) gene marker and analyzed by terminal-restriction fragment length polymorphism. The terminal-restriction fragment length polymorphism fingerprinting resulted in 15 terminal restriction fragments. The ordination configuration of the terminal restriction fragments data, using nonmetric multidimensional scaling, showed a clear gradient in the methanogen community structure along the mire chronosequence. In addition, fingerprint patterns of samples from the water table level and 40 cm below differed from one another in the bog site (site 5). Methane production was negligible in the three youngest fen sites (sites 1-3) and showed the highest rates in the oligotrophic fen site (site 4). Successful PCR amplification using mcr gene primers revealed the presence of a methanogen community in all five sites along the study transect.     

Element interactions in forest ecosystems: succession,allometry and input-output budgets

Element interactions within forests differ from those in other major ecosystems for three major reasons: — a greater allocation of carbon to structural material; — a greater element storage within biomass; and — the diversity of carbon- and nutrient-containing metabolites produced. The most important of these differences is structural material, which can lead to C: element ratios in biomass (as a whole) 100 × greater than those in unicellular organisms. Stand allometry causes the amount of carbon stored and C:element ratios in biomass to change in predictable ways in the course of secondary succession. Such changes affect microbial dynamics and C: element interactions within soils. Bicarbonate, organic acids, nitrate, phosphate, and sulfate are major anions within forest soils: they control leaching of both anions and cations. Biotic interactions of C, N, P, and S during both uptake and mineralization control the potential for production of these anions within forests, and geochemical interactions regulate their mobility and loss.