Impact of competition on plant performances along a flooding gradient: a multi‐species experiment

Question: 1. How do the competitive response and the importance of competition vary between species and along a flooding gradient? 2. How does the role of competition in constraining species distribution limits along the gradient vary between lower and upper limits? Location: A 1‐ha meadow within the Alzette floodplain in Luxembourg. Methods: Competitive response and importance of competition were assessed on seven meadow species differing in their tolerance to flooding. Species were cultured in monocultures and in mixtures, in three water treatments reflecting the wet, the middle and the dry end of a natural flooding gradient. We developed two models based on a multiple regression in order to express each component of competition as a function of the neighbour biomass. Results: Five species showed variations in their competitive response across water treatments; however, these species achieved either their highest or their worst competitive response in their optimal water treatment (i.e. the treatment in which the species had the highest biomass in monoculture). Competition was more important for the flood‐tolerant species in the dry treatment than for the flood‐intolerant species in the wet treatment. Conclusions: 1. Variations in species competitive responses along flooding gradients may be the result of either an amplified effect between competition and hydrological stresses, or a hierarchical effect of stress over competition. 2. The role of competition is more important in constraining the upper distribution limits of the flood‐tolerant species than the lower limits of the flood‐intolerant species along flooding gradients.     

Effects of neighbouring vegetation on eucalypt seedlings at two sites subject to different levels of abiotic stress

Abstract In spite of numerous studies on the effect of nutrient levels and/or standing crop on the intensity of resource competition the debate has not been resolved. Field studies that have used natural productivity gradients have generally supported the argument that competitive intensity and resource availability are positively correlated, whereas studies that have used artificial resource gradients have generally refuted the same argument. Here we report the results from study in which both approaches were used within the same system. We studied two species of eucalypt that occupy contrasting parts of the same landscape: Eucalyptus camaldulensis, found mostly along creek lines and in valleys with deep alluvial soils, and Eucalyptus microcarpa, found on hillsides and ridges with shallow soils. We studied the response of seedlings of the two species to the combined effects of competition and manipulated nutrient levels in a glasshouse experiment, and also investigated their responses to removal of neighbouring plants in the field. Eucalyptus microcarpa was less responsive to increased resource availability, which is consistent with one of the principal assumptions of Grime’s C‐S‐R model. In the glasshouse experiment both species of eucalypt responded in a qualitatively similar fashion to the combined effects of resource availability and competition: release from competition resulted in increased growth, but only in pots that received additional resources. In the field we found that neighbouring vegetation could severely limit the establishment of E. camaldulensis but the removal of neighbouring vegetation did not affect the performance of E. microcarpa seedlings. Eucalyptus camaldulensis seedlings suffered high levels of damage from herbivores. Our results thus generally support the predications of the C‐S‐R model, however, they indicate that the effects of competition and herbivory may be heavily confounded.     

Quantifying relationships between traits and explicitly measured gradients of stress and disturbance in early successional plant communities

Questions: How can one explicitly quantify, and separately measure, stress and disturbance gradients? How do these gradients affect functional composition in early successional plant communities and to what extent? Can we accurately predict trait composition from knowledge of these gradients? Location: Southern Quebec, Canada. Methods: Using eight environmental variables measured in 48 early successional plant communities, we estimated stress and disturbance gradients through structural equation modelling. We then measured 10 functional traits on the most abundant species of these 48 communities and calculated their community‐level mean and variance weighted by the relative abundance of each species. Finally, we related these community‐weighted means and variances to the estimated stress and disturbance gradients using general linear models or generalized additive models. Results: We obtained a well‐fitting measurement model of the stress and disturbance gradients existing in our sites. Of the 10 studied traits, only average plant reproductive height was strongly correlated with the stress (r²=0.464) and disturbance (r²=0.543) gradients. Leaf traits were not significantly related to either the stress or disturbance gradients. Conclusions: The well‐fitting measurement model of the stress and disturbance gradients, combined with the generally weak trait–environment linkages, suggests that community assembly in these early successional plant communities is driven primarily by stochastic processes linked to the history of arrival of propagules and not to trait‐based environmental filtering.     

Contrasting the effects of environment,dispersal and biotic interactions to explain the distribution of invasive plants in alpine communities

Despite considerable efforts devoted to investigate the community assembly processes driving plant invasions, few general conclusions have been drawn so far. Three main processes, generally acting as successive filters, are thought to be of prime importance. The invader has to disperse (1st filter) into a suitable environment (2nd filter) and succeed in establishing in recipient communities through competitive interactions (3rd filter) using two strategies: competition avoidance by the use of different resources (resource opportunity), or competitive exclusion of native species. Surprisingly, despite the general consensus on the importance of investigating these three processes and their interplay, they are usually studied independently. Here we aim to analyse these three filters together, by including them all: abiotic environment, dispersal and biotic interactions, into models of invasive species distributions. We first propose a suite of indices (based on species functional dissimilarities) supposed to reflect the two competitive strategies (resource opportunity and competition exclusion). Then, we use a set of generalised linear models to explain the distribution of seven herbaceous invaders in natural communities (using a large vegetation database for the French Alps containing 5,000 community-plots). Finally, we measure the relative importance of competitive interaction indices, identify the type of coexistence mechanism involved and how this varies along environmental gradients. Adding competition indices significantly improved model’s performance, but neither resource opportunity nor competitive exclusion were common strategies among the seven species. Overall, we show that combining environmental, dispersal and biotic information to model invasions has excellent potential for improving our understanding of invader success.     

养分资源脉冲供给对几种微藻种间竞争的影响

以淡水生态系统常见的5种微藻为研究对象,通过稳态条件下单一培养的方式获取各微藻在氮素或磷素缺乏条件下对应的生长特征参数和R*值,同时将5种微藻在养分资源脉冲供给的方式下进行混合培养,进而检测养分资源脉冲供给对几种微藻种间竞争的影响作用,并与基于稳态条件下所预测的微藻种间竞争结果进行比较。研究结果显示,在稳态条件下具有最小R*值的纤细角星鼓藻在与其它微藻的种间竞争过程中始终保持优势地位,而其余4种微藻在混合培养状态下的相对比重亦与其稳态条件下的R*值紧密相关。然而,资源脉冲供给条件下的竞争优势种纤细角星鼓藻能与其它种群数量较小的微藻共存。此外,在氮素和磷素的不同供给比例情况下,对应微藻群落的结构也会发生相应的变化。实验养分资源脉冲作用下多种微藻的共存现象与自然水体中的观测现象相一致,显示了资源脉冲可能是维持生物多样性水平的一个重要机制。     

Do biotic interactions modulate ecosystem functioning along stress gradients? Insights from semi-arid plant and biological soil crust communities

Climate change will exacerbate the degree of abiotic stress experienced by semi-arid ecosystems. While abiotic stress profoundly affects biotic interactions, their potential role as modulators of ecosystem responses to climate change is largely unknown. Using plants and biological soil crusts, we tested the relative importance of facilitative–competitive interactions and other community attributes (cover, species richness and species evenness) as drivers of ecosystem functioning along stress gradients in semi-arid Mediterranean ecosystems. Biotic interactions shifted from facilitation to competition along stress gradients driven by water availability and temperature. These changes were, however, dependent on the spatial scale and the community considered. We found little evidence to suggest that biotic interactions are a major direct influence upon indicators of ecosystem functioning (soil respiration, organic carbon, water-holding capacity, compaction and the activity of enzymes related to the carbon, nitrogen and phosphorus cycles) along stress gradients. However, attributes such as cover and species richness showed a direct effect on ecosystem functioning. Our results do not agree with predictions emphasizing that the importance of plant–plant interactions will be increased under climate change in dry environments, and indicate that reductions in the cover of plant and biological soil crust communities will negatively impact ecosystems under future climatic conditions.     

Competition intensity at local versus regional spatial scales

Studies of competition intensity over natural (i.e. topographic) gradients often contradict the results from studies where artificial (i.e. fertilizer) gradients have been used. Why should the type of gradient matter? To explore the possibilities, we performed experiments to measure competition intensity experienced by tree seedlings from grass competitors across a natural resource gradient, and simultaneously across artificial soil nutrient (fertiliser) gradients. We measured various functional traits (i.e. specific leaf area, leaf area, leaf nitrogen content, δ¹⁵N, δ¹³C, RGR) to gain mechanistic insight into the nature of competition across these gradients. Competition intensity increased with increasing resource availability, unequivocally at the local scale (i.e. with fertilizer application) but not at the regional scale (i.e. across the natural productivity gradient). Our measurements of plant traits were generally consistent with measurements of competition intensity, and demonstrate that competition occurs even when resource levels are low. Competition mainly acted to reduce the growth of Eucalyptus seedlings. Functional (physiological) traits in the Eucalyptus seedlings were not strongly affected by competitors, with the possible exception of δ¹⁵N, which may effectively integrate information on soil nutrient, moisture and leaf processes.     

Responses of Fraxinus excelsior seedlings to grass-induced above- and below-ground competition

The competitive interactions between woody seedlings and herbaceous vegetation have received increasing interest in recent years. However, little is known about the relative contributions and underlying mechanisms of above- and below-ground competition between species. We used a novel experimental approach to assess the responses of Fraxinus excelsior seedlings to different combinations of root and shoot competition imposed by the grass Dactylis glomerata under greenhouse conditions. Seedling growth was significantly reduced by competition for soil resources, but neither biomass nor height were significantly affected by shoot competition for light. Competitive response indices based on biomass confirmed that below-ground competition was more important than above-ground competition, and indicated that root and shoot competition did not interact to influence plant growth. Fraxinus biomass allocation and seedling traits were almost all significantly affected by root competition; these responses varied depending on the trait examined. In contrast, morphological responses to shoot competition were limited. In the absence of root competition, seedlings showed a significant increase in the biomass allocated to leaves and a greater leaf area ratio in response to shoot competition. Our findings suggest that morphological modifications help to mitigate the negative effects of competition, but the expression of plasticity may be suboptimal due to resource constraints. The present study also highlights the importance of appropriate experimental controls and analysis to avoid confounding effects of experimental design and ontogeny on the interpretation of competitive responses.     

Empirical evidence for competition-driven semelparity in wild medaka

Metabolic theory predicts that maintenance rate increases faster with animal body size than food intake rate, such that the critical resource density R* at which ingested energy exactly covers maintenance requirements increases with body size. Small-sized (low R*) juveniles may thus exclude their larger-sized (high R*) parents in resource competition, resulting in apparent semelparous life histories and non-overlapping generations. However, empirical support for such a competition-driven semelparity (CDS) remains scarce. Here, we report a high consistency of cohort dynamics with CDS in wild medaka (Oryzias latipes). As predicted by the theory, there was a strong juvenile-adult diet overlap, and all individuals died after reproduction as semelparous age-1 adults, synchronous with a rapid somatic growth of age-0 juveniles into the adult stage and with dropping abundances of zooplankton food resources. In addition to the theory, we found evidence for increased reproductive allocation under food stress, translating into immune depression and elevated parasite prevalence. Therefore, CDS in medaka emerges both from intercohort competitive exclusion and from food-dependent energy reallocation from maintenance to reproduction, the later presumably representing an adaptive response to the former. The literature data show that the strengths of both intercohort competition and reproductive allocation increase at higher temperatures in many ectotherms, pointing to climate warming as a potentially powerful magnifier of CDS in the wild.     

Intensity and Importance of Competition for a Grass (Festuca rubra) and a Legume (Trifolium pratense) Vary with Environmental Changes

How plant competition varies across environmental gradients has been a long debate among ecologists. We conducted a growth chamber experiment to determine the intensity and importance of competition for plants grown in changed environmental conditions. Festuca rubra and Trifolium pratense were grown in monoculturs and in two- and/or three-species mixtures under three environmental treatments. The measured competitive variations in terms of growth (height and biomass) were species-dependent. Competition intensity for Festuca increased with decreased productivity, whilst competition importance displayed a humpback response. However, significant response was detected in neither competition intensity nor importance for Trifolium. Intensity and importance of competition followed different response patterns, suggesting that they may not be correlated along an environmental gradient. The biological and physiological variables of plants play an important role to determine the interspecific competition associated with competition intensity and importance. However, the competitive feature can be modified by multiple environmental changes which may increase or hinder how competitive a plant is.     

Philip Grime's fourth corner: are there plant species adapted to high disturbance and low productivity?

Grime's CSR species life‐strategy theory (competition–stress–ruderality) provides a conceptual framework to classify species into competitive (growing under high productivity, low disturbance), stress‐tolerant (low productivity, low disturbance) and ruderal (high productivity, high disturbance). Importantly, this classification is based on the assumption that the niche space of disturbance and productivity is filled unevenly: while in productive habitats species can adapt to different disturbance regimes, species of low‐productivity and disturbed habitats do not exist, resulting in a triangular distribution of species optima along axes of disturbance and productivity. This assumption has often been criticised, but it has not yet been put under a rigorous test. Here we use existing data on niche positions of central European plant species to test this hypothesis, namely its prediction that species adapted to jointly stressed (low‐productive) and disturbed habitats do not exist. We use Ellenberg indicator values and newly developed indicator values for disturbance as proxies of species positions in the space of productivity and disturbance. We found that positions of species optima along the gradients of productivity and disturbance severity are not independent of each other, with very few species adapted to low‐productive and severely disturbed habitats. In contrast, there is no relationship between productivity and disturbance frequency; a number of species occur in low‐productive and frequently disturbed habitats. The relationship between productivity and disturbance severity can be either due to tradeoffs between life history traits responsible for response to disturbance and productivity (as originally assumed by Grime) or due to historical rarity of severely disturbed habitats in unproductive conditions and consequent absence of evolution of species adapted to them. Our data are based on one specific flora, shaped by glaciations and early introduction of agriculture, but the question of what causes this pattern can be resolved by future analyses of floras with different evolutionary and ecological histories.     

HOW CRITICAL IS THE CRITICAL N:P RATIO?1

The value of cellular N:P that corresponds to co‐limitation by N and P, the critical (R_crit) or optimum ratio, has been used to infer the competitive advantage of phytoplankton growing in P‐impoverished systems. Using a revised quota model, with a normalized quota function and capable of simulating surge transport, the interactions between the minimum P‐quota (PCo), the shape of the P‐quota–cell growth relationship (affected by constant KQP), and transport kinetics in affecting the utility of R_crit are considered. For a low PCo to endow an organism with a high R_crit over a wide range of growth rates, the P‐quota curve must be more hyperbolic (KQP low) rather than linear (KQP high). PCo and KQP also affect the half saturation constant for growth, KgP ; this and the capacity to transport nutrients at rates above those required to sustain steady‐state growth endows a competitive advantage. However, the kinetics of transport into the organism have a greater potential for affecting KgP than changing the kinetics of internal P usage. Thus, the value of R_crit is not a critical factor affecting competition except in extreme oligotrophic conditions. For competition between species, nutrient transport, accumulation, and resource utilization are all important. However, the efficiency of internal resource utilization is of lesser importance, and certainly not of greater importance, than resource acquisition. Multinutrient models intended to describe competition need to recognize these interactions; the traditional quota model is poorly equipped to do so.     

Relative importance of competition and plant–soil feedback,their synergy,context dependency and implications for coexistence

Plants interact simultaneously with each other and with soil biota, yet the relative importance of competition vs. plant–soil feedback (PSF) on plant performance is poorly understood. Using a meta‐analysis of 38 published studies and 150 plant species, we show that effects of interspecific competition (either growing plants with a competitor or singly, or comparing inter‐ vs. intraspecific competition) and PSF (comparing home vs. away soil, live vs. sterile soil, or control vs. fungicide‐treated soil) depended on treatments but were predominantly negative, broadly comparable in magnitude, and additive or synergistic. Stronger competitors experienced more negative PSF than weaker competitors when controlling for density (inter‐ to intraspecific competition), suggesting that PSF could prevent competitive dominance and promote coexistence. When competition was measured against plants growing singly, the strength of competition overwhelmed PSF, indicating that the relative importance of PSF may depend not only on neighbour identity but also density. We evaluate how competition and PSFs might interact across resource gradients; PSF will likely strengthen competitive interactions in high resource environments and enhance facilitative interactions in low‐resource environments. Finally, we provide a framework for filling key knowledge gaps and advancing our understanding of how these biotic interactions influence community structure.     

COMPETITION AS A CONTROL MECHANISM IN SUBALPINE MEADOWS

Interspecific competition in a stable subalpine meadow was investigated experimentally. Vegetation responds to gradients of moisture and disturbance which control species composition. Moisture and snow melt patterns produce a productivity gradient that affects the importance of competition in this vegetation mosaic. Reciprocal transplant and selective removal experiments demonstrated that the intensity of competition is directly related to productivity and that seedlings are more susceptible to competitive effects than are established plants. Competition may reduce diversity in stable, productive habitats, while disturbance and stress limit diversity directly. Habitats of moderate productivity and minor disturbance have maximum diversity. The experiments and observations combine to suggest that habitat heterogeneity and local disturbance interact to control biotic interference and that the relative importance of each varies with productivity.     

基于树冠重叠面积的天然混交林林木竞争指数

构建基于树冠重叠面积和林木混交关系的竞争指数,探究天然混交林林分及主要树种竞争的表达方法,并以甘肃省小陇山林区锐齿槲栎天然混交林为研究对象,采用胸径、树高、冠幅、距离等信息的2期调查数据,对新构建的基于树冠重叠面积的天然混交林林木竞争(CIM)指数进行验证,并选取Spearman系数,对CIM竞争指数和传统竞争指数进行相关性分析。结果表明,采用树冠投影重叠和考虑树高关系确定竞争木的方法能有效避免竞争木多选或漏选; CIM1、CIM2、CIM3能够表达出混交林竞争压力变化,表现出竞争压力越大林木蓄积生长量越小的趋势,将树种混交因子引入竞争指数能够提高蓄积生长量与竞争指数的相关性,更加客观的描述林分中竞争状况。竞争指数与蓄积生长量相关性由大到小排序为CIM2>CIM3>CIM1>Bella竞争指数(CIO)>简单竞争指数(CI)>CIO2>CIO1,表明竞争指数 CIM2对混交林竞争表达更具参考意义。林分中主要树种锐齿槲栎由于相邻木中同树种相邻木增加,导致竞争压力增大。天然混交林中树种组成复杂多样,林木分布不均匀,在竞争指数中引入树种混交因子能够较为真实的反映林木竞争状况。     

大别山五针松种内和种间竞争强度

通过对安徽省岳西县大别山五针松群落内的53株对象木及2079株竞争木的调查,运用Hegyi单木竞争模型分析了大别山五针松的种内和种间竞争强度。结果表明,大别山五针松的种内和种间竞争强度分别为17.11%和82.89%,说明竞争主要来自种间。大别山五针松的伴生种较多,种内与主要伴生种间的竞争关系为短柄枹大别山五针松种内黄山松满山红茅栗金缕梅紫茎四照花灯台树鹅耳枥。竞争强度随对象木胸径的增大而减小,当对象木的胸径小于25cm时,所受到的竞争强度较大;当胸径在25cm以上时,竞争强度变化很小,二者符合幂函数关系(CI=AD-B),所得的预测模型能很好的预测大别山五针松种内和种间竞争强度。     

Habitat disturbance modifies dominance,coexistence, and competitive interactions in tropical ant communities

1. Interspecific competition is a major structuring principle in ecological communities. Despite their prevalence, the outcome of competitive interactions is hard to predict, highly context-dependent, and multiple factors can modulate such interactions. 2. We tested predictions concerning how competitive interactions are modified by anthropogenic habitat disturbance in ground-foraging ant assemblages inhabiting fragmented Inter-Andean tropical dry forests in southwestern Colombia, and investigated ant assemblages recruiting to baits in 10 forest fragments exposed to varying level of human disturbance. 3. Specifically, we evaluated how different components of competitive interactions (patterns of species co-occurrence, resource partitioning, numerical dominance, and interspecific trade-offs between discovery and dominance competition) varied with level of habitat disturbance in a human-dominated ecosystem. 4. Multiple lines of evidence suggest that the role of competitive interactions in structuring ground-foraging ant communities at baits varied with respect to habitat disturbance. As disturbance increased, community structure was more likely to exhibit random co-occurrence patterns, higher levels of monopolization of food resources by dominant ants, and disproportionate dominance of a single species, the little fire ant (Wasmannia auropunctata). At a regional scale, we found evidence for a trade-off between dominance and discovery abilities of the 15 most common species at baits. 5. Together, these results suggest that human disturbance modifies the outcome of competitive interactions in ground-foraging ant assemblages and may promote dominant species that reduce diversity and coexistence in tropical ecosystems.     

Modelling the multidimensional niche by linking functional traits to competitive performance

Linking competitive outcomes to environmental conditions is necessary for understanding species'' distributions and responses to environmental change. Despite this importance, generalizable approaches for predicting competitive outcomes across abiotic gradients are lacking, driven largely by the highly complex and context-dependent nature of biotic interactions. Here, we present and empirically test a novel niche model that uses functional traits to model the niche space of organisms and predict competitive outcomes of co-occurring populations across multiple resource gradients. The model makes no assumptions about the underlying mode of competition and instead applies to those settings where relative competitive ability across environments correlates with a quantifiable performance metric. To test the model, a series of controlled microcosm experiments were conducted using genetically related strains of a widespread microbe. The model identified trait microevolution and performance differences among strains, with the predicted competitive ability of each organism mapped across a two-dimensional carbon and nitrogen resource space. Areas of coexistence and competitive dominance between strains were identified, and the predicted competitive outcomes were validated in approximately 95% of the pairings. By linking trait variation to competitive ability, our work demonstrates a generalizable approach for predicting and modelling competitive outcomes across changing environmental contexts.     

Functional and performance comparisons of invasive Hieracium lepidulum and co‐occurring species in New Zealand

Abstract One of the key environmental factors affecting plant species abundance, including that of invasive exotics, is nutrient resource availability. Plant functional response to nutrient availability, and what this tells us about plant interactions with associated species, may therefore give us clues about underlying processes related to plant abundance and invasion. Patterns of abundance of Hieracium lepidulum, a European herbaceous invader of subalpine New Zealand, appear to be related to soil fertility/nutrient availability, however, abundance may be influenced by other factors including disturbance. In this study we compare H. lepidulum and field co‐occurring species for growth performance across artificial nutrient concentration gradients, for relative competitiveness and for response to disturbance, to construct a functional profile of the species. Hieracium lepidulum was found to be significantly different in its functional response to nutrient concentration gradients. Hieracium lepidulum had high relative growth rate, high yield and root plasticity in response to nutrient concentration dilution, relatively low absolute yield, low competitive yield and a positive response to clipping disturbance relative to other species. Based on overall functional response to nutrient concentration gradients, compared with other species found at the same field sites, we hypothesize that H. lepidulum invasion is not related to competitive domination. Relatively low tolerance of nutrient dilution leads us to predict that H. lepidulum is likely to be restricted from invading low fertility sites, including sites within alpine vegetation or where intact high biomass plant communities are found. Positive response to clipping disturbance and relatively high nutrient requirement, despite poor competitive performance, leads us to predict that H. lepidulum may respond to selective grazing disturbance of associated vegetation. These results are discussed in relation to published observations of H. lepidulum in New Zealand and possible tests for the hypotheses raised here.     

Interspecific competition among macroparasites in a density-dependent host population

 We analyze the dynamics of a community of macroparasite species that share the same host. Our work extends an earlier framework for a host species that would grow exponentially in the absence of parasitism, to one where an uninfected host population is regulated by factors other than parasites. The model consists of one differential equation for each parasite species and a single density-dependent nonlinear equation for the host. We assume that each parasite species has a negative binomial distribution within the host and there is zero covariance between the species (exploitation competition). New threshold conditions on model parameters for the coexistence and competitive exclusion of parasite species are derived via invadibility and stability analysis of corresponding equilibria. The main finding is that the community of parasite species coexisting at the stable equilibrium is obtained by ranking the species according t! o th e minimum host density H ^* above which a parasite species can grow when rare: the lower H ^* , the higher the competitive ability. We also show that ranking according to the basic reproduction number Q ₀ does not in general coincide with ranking according to H ^* . The second result is that the type of interaction between host and parasites is crucial in determining the competitive success of a parasite species, because frequency-dependent transmission of free-living stages enhances the invading ability of a parasite species while density-dependent transmission makes a parasite very sensitive to other competing species. Finally, we show that density dependence in the host population entails a simplification of the portrait of possible outcomes with respect to previous studies, because all the cases resulting in the exponential growth of host and parasite populations are eliminated.. Received: 24 June 1996 / Revised version: 28 April 1998