Experimental manipulation of succession in heathland ecosystems

An experiment was carried out in two heathland ecosystems, one dominated by Calluna vulgaris and the other by Molinia caerulea, to analyse the effects of soil organic matter accumulation and nutrient mineralization on plant species dynamics during succession. The experiment included one treatment that received nutrient solution and two treatments where the rate of soil organic matter accumulation was reduced by removing litter or accelerated by adding litter. In a fourth treatment the C. vulgaris litter produced in the C. vulgaris-dominated plots was replaced by litter of M. caerulea and vice versa. Treatments were applied over 8 years. Addition of nutrient solution caused C. vulgaris to decline, and grass species to increase sharply, compared to the control plots. Addition of litter enhanced both N mineralization and the biomass of M. caerulea and Deschampsia flexuosa but reduced the biomass of C. vulgaris. The effects of replacing C. vulgaris litter by M. caerulea litter, or vice versa, on N mineralization and species dynamics could not be attributed to differences between the decomposability of the different litter materials that were transferred. The results confirm the hypothesis that increased litter inputs accelerate the rate of species replacement during succession.     

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.     

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.     

Oscillating vegetation dynamics in a wet heathland

Abstract. Question: The significance of disturbances caused by periodical inundation was investigated with respect to its effects on vegetation dynamics, species richness and fluctuations, and to the relevance of certain plant properties. Location and Method: At a sod‐cut stand within nutrient‐poor inland sand dunes, permanent plots along a transect were surveyed over a period of up to nine years after sod cutting. Results: In contrast to never inundated plots, periodically inundated plots were characterized by low vegetation cover and by high numbers of species belonging to many different communities, each of them with a low cover. Periodical inundations favoured the presence of pioneers, species tolerant of disturbances, species adapted to wet conditions and stoloni‐ferous species. Furthermore, annual fluctuations of species within each plot were higher and most species occurred only sporadically. Discussion: A comprehensive model is presented describing the relevant processes identified in the littoral zone. Changing water tables result in the creation of gaps. The re‐colonization of these gaps follows mainly from vegetative regeneration and less to the dispersion of diaspores. Highest species numbers in the zone of moderate disturbances result from a high rate of re‐colonization in spite of local extinctions following each disturbance event. It is suggested that colonization abilities are among the most important features for species occurrence at a site rich in disturbances (more important than competitive abilities and more important than a slow rate of displacement). For nature conservation such sites are very important, because they allow (rare) pioneer species to survive for longer periods of time.     

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.     

Interspecific competition in metapopulations

The assumptions and predictions of metapopulation models for competing species are discussed in relation to empirical studies of colonization and extinction in metapopulations. In three species of Daphnia in rockpools, interspecific competition increased local extinction rates, while no effects on colonization rates were detected. Distributional patterns were consistent with several predictions of the competition model; for example, the number of species on an island increased with the number of pools and the proportion of pools occupied by each species decreased with increasing species number. It is concluded that interspecific competition is important for the distributional dynamics of Daphnia species in rockpools, but the question whether the coexistence of these species depends on metapopulation dynamics is still unresolved. Other studies on the effects of interspecific competition on colonization and extinction rates are discussed.     

Secondary succession after fire in a Mediterranean dwarf-shrub community

Abstract. Changes in vegetation composition after fire in a Sarcopoterium spinosum dwarf-shrub community on phosphorus-deficient terra rossa soil overlying limestone were studied over a 10-yr period. Herbicide application and phosphate fertilizer were combined to provide a range of initial conditions for recovery. The site was undisturbed during the growing season in winter but was grazed by cattle in summer. Encroachment rate of S. spinosum was mainly a function of the density of viable buds on plants that survived the fire and/or the shrub control treatments. This, together with the very low rate of shrub establishment from seed set a short-to-medium term limit to the attainable cover of S. spinosum. The leguminous shrub Calycotome villosa appeared only in the third year after the fire, apparently from seed. After shrub control, the encroachment rate of C. villosa was low, even after 9 yr. On the other hand, where no herbicide was applied, and with the cessation of goat grazing, C. villosa became increasingly prominent, to the extent that by the 8th year after the fire, it inhibited the continued expansion of S. spinosum. Phosphorus nutrition tended to reduce the cover of S. spinosum because of its stimulating effect on the predominantly leguminous herbaceous vegetation. C. villosa also appeared to benefit from greater phosphorus availability, despite greater competition from the annual vegetation. These findings show that, depending on the context, secondary succession after fire can take different courses. Whereas the combined effect of herbicide control and P-enrichment cannot prevent shrub encroachment, it can slow down the rate of succession for more than a decade, despite the presence of ample shrub seed sources in the area.     

Waterlogging and fire impacts on nitrogen availability and utilization in a subtropical wet heathland (wallum)

Protein, amino acids and ammonium were the main forms of soluble soil nitrogen in the soil solution of a subtropical heathland (wallum). After fire, soil ammonium and nitrate increased 90- and 60-fold, respectively. Despite this increase in nitrate availability after fire, wallum species exhibited uniformly low nitrate reductase activities and low leaf and xylem nitrate. During waterlogging soil amino acids increased, particularly γ-aminobutyric acid (GABA) which accounted for over 50% of amino nitrogen. Non-mycorrhizal wallum species were significantly (P < 0.05) ¹⁵N-enriched (0.3–4.3‰) compared to species with mycorrhizal associations (ericoid-type, ecto-, va-mycorrhizal) which were strongly depleted in ¹⁵N (-6.3 to -1.8‰). Lignotubers and roots had δ¹⁵N signatures similar to that of the leaves of respective species. The exceptions were fine roots of ecto-, ecto/va-, and ericoid type mycorrhizal species which were enriched in ¹⁵N (0.1–2.4‰). The 5¹⁵N signatures of δ¹⁵N_{total soil N} and δ¹⁵N_{soil NH4+} were in the range 3.7–4.5‰, whereas δ¹⁵N_{soil NO3?} was significantly (P < 0.05) more enriched in ¹⁵N (9.2–9.8‰). It is proposed that there is discrimination against ¹⁵N during transfer of nitrogen from fungal to plant partner. Roots of selected species incorporated nitrogen sources in the order of preference: ammonium > glycine > nitrate. The exception were proteoid roots of Hakea (Proteaceae) which incorporated equal amounts of glycine and ammonium.     

Interspecific competition between mice: A reciprocal field manipulation experiment

Previous studies have demonstrated a species replacement sequence in disturbed forest and heathland, with the native New Holland mouse (Pseudomys novaehollandiae) replacing the introduced house mouse (Mus musculus) in the earliest part of the succession. The replacement appeared to be competitive displacement and this has been confirmed by controlled experimental removal of P. novaehollandiae from field sites, which produced significant increases in the abundance and habitat use of Mus musculus. This paper reports the results of the controlled reciprocal manipulation experiment in the field. A pulsed addition of M. musculus to three treatment plots produced significant reductions in the abundance and habitat use of P. novaehollandiae, when compared with control plots with no addition of M. musculus. This effect was observed only at high M. musculus densities, and the return to the control conditions within 3 months of the pulsed addition is consistent with asymmetric interspecific competition from competitively superior P. novaehollandiae. The importance of considering the regional abundance of M. musculus in relation to its competitive ability is stressed, as at high densities M. musculus is able to create its own ecological space contrary to its more usual competitively submissive status.     

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.     

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.     

Vegetation succession after fire in sclerophyll woodland communities in south-eastern Australia

The dynamics of vegetation regeneration after burning were examined in three dry sclerophyll communities near Canberra, in south-eastern Australia. Changes in seedling and regrowth populations were followed in permanent quadrats during the first two years after burning, compared with both the preburn vegetation and population changes over the same period in adjacent, unburnt plots. All species represented either by living plants in the tree and shrub strata and/or by seed in the soil and litter prior to burning regenerated during the first year after the fire treatments. No new species invaded the areas after burning. Species varied in their regenerative strategy and recovered after the fires either by germination of seed residual in the soil and ash or released from trees after burning, by regrowth from surviving vegetative organs, or by a combination of germination and regrowth. Both seedling input and the vegetative recovery of populations were higher during the first than second year after burning. The vegetative multiplication and seed gertnination of many species were stimulated by fire. It was concluded that the regeneration of the communities studied, as well as the post-fire recovery of numerous different communities reported in the literature, closely resembled an initial floristic composition model. It is considered that the process of vegetation redevelopment after a disturbance (i.e. secondary succession) will be influenced greatly by the species composition at the time of disturbance, and by the type of disturbing agent. A single model would not be expected to adequately describe secondary succession following disturbance by agents imposing different stress conditions on a community.     

Fire,rodent herbivory,and plant competition: implications for invasion and altered fire regimes in the Mojave Desert

Oecologia - Biological invasions are responsive to changing wildfire regimes related to human activities that are altering biological communities. Our objective was to investigate how fire, rodent...     

Shoot regeneration after fire or freezing temperatures and its relation to plant life-form for some heathland species

Shoot regeneration after prescribed burning or following the freezing temperatures of winter was monitored for nineteen heathland species present in an Arctostaphyleto-Callunetum community in northeast Scotland. Species whose renewal buds were near the surface of the ground started to grow earlier in the spring than species with renewal buds above the surface, but grouping species according to the position of their renewal bud (i.e. their life-form) did not account for all of the interspecific variation apparent. In the case of shoot regeneration after fire, species whose renewal buds were destroyed by fire because they were above-ground started to regenerate about the same time as species with belowground buds, protected from fire, but reached their maximum frequency of occurrence later. Grouping species by life-form was of limited value as a means of interpreting this interspecific variation in the timing of shoot regeneration after fire. It would be unwise to use plant life-form as the sole basis for interpreting or predicting a species' response to temperature stress when extreme temperatures occur regularly, as they do in heathland. The possible use of other plant traits to interpret and predict interspecific variation in the regeneration rate of heathland plants is discussed.Nomenclature follows Tutin et al. (1964–1980) for vascular plants. Acknowledgements. The Nature Conservancy Council and Mr J. J. Humphries kindly allowed Dinnet Moor to be used for the work presented here. One of us (RJR) received financial support for field work from the Natural Sciences and Engineering Research Council of Canada.     

Interspecific competition in small rodents: from populations to individuals

The role of interspecific competition in shaping animal and plant communities has formed one of the major issues in ecology for decades. Small mammals, mainly rodents, have been among the model systems used for research on interspecific competition. Most studies within small mammal systems in the past have examined effects of competition on population attributes such as on population size, habitat use, or population dynamics. Population-level responses are the cumulative effects of individual responses, however, the influence of competition on individual life-history traits has rarely been studied. Research on life-histories may bridge gaps between population biology and effects of competition on individual behaviour. In this paper, we review recent research approaches to interspecific competition in rodents based on census data and species assemblages, that use regression analysis, time series analysis, removal and exclusion experiments, and showcase our own experimental research on the effects of interspecific competition on individual life-history traits in boreal voles.     

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).     

Interspecific competition in perennial sedentary organisms: An individual-based model

We investigated a mathematical model of the dynamics of the ecological system consisting of two competing perennial species, each of which leads a sedentary life. It is an individual-based model, in which the growth of each individual is described. The rate of this growth is weakened by competition from neighboring individuals. The strength of the competitors' influence depends on their size and distance to them. The conditions, in which the competitive exclusion of one of the competitors and the coexistence of both competitors take place are provided. The influence of the parameters responsible for the strength of competition, the degree of competitive asymmetry, and consideration of the importance of specific elements of the spatial structure of this ecological system on the results of the competition were analyzed. Both species co-exist when they are equal competitors. Permanent coexistence is possible only when interspecific competition is weaker than intraspecific. When interspecific competition is stronger, the coexistence of equal interspecific competitors is random. Both species have equal probability of extinction. If species are not equal competitors, the stronger one wins. This result can be modified by different strengths of intraspecific competition. The weaker interspecific competitor can permanently coexist with stronger one, when its individuals suffer stronger intraspecific competition.     

The implications of succession after fire for the conservation management of moorland invertebrate assemblages

Fire management in protected areas requires an understanding of the consequences of fire regimes. Invertebrates are a key component of biological communities, but studies of fire impacts on diverse invertebrate assemblages over long timeframes are rare. The responses of ground- and foliage-active invertebrate assemblages to fire in buttongrass moorlands were investigated using a space-for-time design. Assemblages in recently burnt moorlands were distinct from those in older moorlands. Contrary to expectations, ground-active invertebrate abundance, but not taxon richness, was greatest in young regrowth (2–3 years since last fire), owing to large populations of Formicidae, Orthoptera, Collembola and Diptera. Foliage-active invertebrate assemblages followed the expected trend with least numbers of invertebrates and taxa in young regrowth. Very few taxa (n?=?9) were absent from young successional stages and none were absent from later successional stages. Invertebrate assemblages in moorlands on low productivity soils took approximately twice as long to return to their pre-fire state than assemblages on moderate productivity soils. The shifts in invertebrate composition were associated with shifts in vegetation composition. Vegetation density was found to be a potentially important predictor of invertebrate compositional variation. Fire in buttongrass moorland appears to have a limited impact on ground-active and foliage-active invertebrate assemblages, suggesting that these components of the invertebrate fauna are resilient to fire (i.e. able to return to the pre-fire state). Given that fire impedes successional processes that convert moorlands into rainforest, and eliminate many of the moorland invertebrate species, conservation management of moorlands should involve the acceptance or imposition of fire.     

Interspecific competition in bats: state of knowledge and research challenges

1. Interspecific competition (IC) is often seen as a main driver of evolutionary patterns and community structure. Bats might compete for key resources, and cases of exaggerated divergence of resource-related characters or trait overdispersion in bat assemblages are often explained in terms of current or past interspecific competition. However, other pressures leading to patterns that mimic the outcome of competition cannot always be ruled out.
2. We present the state of knowledge on IC among bats, providing a critical evaluation of the information available and identifying open questions and challenges.
3. We reviewed 100 documents addressing potential or actual IC in bats and categorised them in terms of the resource for which bats compete (food, foraging habitat, roosts, water, and acoustic space). We also examined the ecomorphological and behavioural traits considered therein to highlight responses to IC or niche partitioning.
4. We found that: although resources should be limiting in order for competition to occur, this is seldom tested; sympatry is sometimes taken as synonymous of syntopy (yet sympatric species that are not syntopic will never experience competition); comparisons between sympatry and allopatry are rare; and testing of objective criteria exploring the existence of niche partitioning or character displacement is not commonly adopted.
5. While morphological examination of food remains in droppings has often led to coarse-grained analysis that proved insufficient to establish the occurrence of food niche overlap or partitioning, new frontiers are being opened by state-of-the-art molecular dietary analysis.
6. A better understanding of IC in bats is paramount, since distributional changes leading to novel bat assemblages driven by climate change are already taking place, and the dramatic decline in insect availability, as well as the global loss or alteration of foraging habitat, may generate new competitive interactions or exacerbate existing interactions in the Anthropocene, and into the future.