Biomass and water storage dynamics of epiphytes in old-growth and secondary montane cloud forest stands in Costa Rica

The way competition structures plant communities has been investigated intensely over many decades. Dominance structures due to competitive hierarchies, with consequences for species richness, have not received as much experimental attention, since their manipulation is a large logistic undertaking. Here the data from a model system based on bryophytes are presented to investigate competition structure in a three-species system. Grown in monocultures, pairwise and three-species mixtures under no and high nitrogen supply, the three moss species responded strongly to treatment conditions. One of them suffered from nitrogen fertilisation and hence performed better in mixtures, where the dominant species provided physical shelter from apparently toxic nitrogen spray. Accordingly, no linear competitive hierarchy emerged and qualitative transitivity remains restricted to the unfertilised treatments. Faciliation also affected other properties of the competition structure. The reciprocity of competition effects could not be observed. Moreover, the performances in three-species mixtures were not well predictable from the knowledge of monocultures and pairwise mixtures because competitive effects were not additive. This had implications for community stability at equilibrium: all two-species systems were stable, both fertilised and unfertilised, while the three-species system was only stable when fertilised. This stability under fertilisation has probably to do with the facilitative effect of the two dominant species on the third. In this experiment, little support for commonly held ideas was found about the way competition in plant communities is structured. On the other hand, this study shows that moss communities are ideal model systems to test predictions of theoretical models concerning properties and consequences of competition in plant communities.     

A spatially explicit trait-based approach uncovers changes in assembly processes under warming

The re-assembly of plant communities during climate warming depends on several concurrent processes. Here, we present a novel framework that integrates spatially explicit sampling, plant trait information and a warming experiment to quantify shifts in these assembly processes. By accounting for spatial distance between individuals, our framework allows separation of potential signals of environmental filtering from those of different types of competition. When applied to an elevational transplant experiment in the French Alps, we found common signals of environmental filtering and competition in all communities. Signals of environmental filtering were generally stronger in alpine than in subalpine control communities, and warming reduced this filter. Competition signals depended on treatments and traits: Symmetrical competition was dominant in control and warmed alpine communities, while hierarchical competition was present in subalpine communities. Our study highlights how distance-dependent frameworks can contribute to a better understanding of transient re-assembly dynamics during environmental change.     

Dispersal spectra,diaspore size and the importance of endozoochory in the equatorial Andean montane forests

The dispersal syndrome hypothesis states that plant diaspores show morphological features that are the results of adaptation for dispersal by a particular vector. This can enable to identify the relative importance of dispersal agents within plant communities. Nevertheless, there is still little information about seed dispersal spectra and diaspore traits related to different dispersal agents in the equatorial montane forests, despite their high biodiversity and important ecosystem services as watersheds for human communities. Due to an increase in environmental stress at high elevations a reduction in the prevalence of endozoochory, and a reduction in the size of endozoochorous diaspores in plant assemblages could be expected. We reviewed published data from 64 Andean cloud forest plots to assess the dispersal spectra, the incidence of different traits related to seed dispersal, and the distribution of dispersal syndromes within cloud forests of northern South America. We then evaluated two questions related to seed dispersal in these forests: (1) Does the number and percentage of endozoochorous species in woody plant assemblages decrease at higher elevation? and (2) Does the mean diaspore size of endozoochorously dispersed tree assemblages decrease with elevation?     

Experimental evidence for root competition effects on community evenness in one of two phytometer species

Aims Plant–plant interactions, being positive or negative, are recognized to be key factors in structuring plant communities. However, it is thought that root competition may be less important than shoot competition due to greater size symmetry belowground. Because direct experimental tests on the importance of root competition are scarce, we aim at elucidating whether root competition may have direct or indirect effects on community structure. Indirect effects may occur by altering the overall size asymmetry of competition through root–shoot competitive interactions.     

Does plant diversity increase top–down control of herbivorous insects in tropical forest?

Higher trophic level interactions are key mediators of ecosystem functioning in tropical forests. A rich body of theory has been developed to predict the effects of plant diversity on communities at higher trophic levels and the mechanisms underlying such effects. The 'enemies hypothesis’ states that predators exert more effective top–down control of herbivorous insects with increasing plant diversity. Support for this hypothesis has been found in temperate forests and agroecosystems, but remains understudied in tropical forests. We compared incidence of attacks of different natural enemies using artificial caterpillars in a tropical forest landscape and investigated the role of plant community structure (i.e. species richness, composition and density), and the role of forest fragmentation (i.e. patch size, edge distance and canopy openness) on predation intensity. Plant community effects were tested with respect to three vegetation strata: trees, saplings and herbs. Observed predation was substantially due to ants. Predation rates increased with plant species richness for trees and herbs. Density of saplings, herb cover and herb species composition were important factors for predation. No significant patterns were found for fragmentation parameters, suggesting that forest fragmentation has not altered predation intensity. We conclude that in tropical forests, top–down control of herbivorous insects in the understory vegetation is affected by a combination of plant diversity, plant species composition and structural features of the plant community.     

Functional and phylogenetic approaches reveal the evolution of diversity in a hyper diverse biota

Ant communities in tropical forests may be governed by varying assembly mechanisms, depending on the particular habitat investigated. We compared phylogenetic diversity and structure across two forest biomes (dry and humid) and two vertical layers (arboreal and terricolous) in ant communities in Madagascar, and assessed the influence of invasive species on this community structure. We estimated phylogenetic signal and correlated evolution for habitat and several functional traits and tested for conservatism in relevant functional and habitat traits. Ancestral states were reconstructed to illuminate the evolution of habitat traits. All analyses utilized phylogenies estimated from newly generated data from three nuclear markers for 290 Malagasy ant taxa. Dry forests, although lower in species richness, were found to support equally high lineage diversity as humid forests. In contrast, phylogenetic diversity was much lower in arboreal than in terricolous communities. We observed significant phylogenetic clustering in the combined humid forest and in the arboreal–humid, arboreal–dry and terricolous–humid communities, whereas the combined dry forest community was overdispersed. Among ant communities in Madagascar, overdispersion and competition therefore may be more prevalent in dry forest, and habitat filtering may be more dominant in humid forest. Excluding invasive ant species had little overall effect on community structure. All investigated traits showed low to intermediate conservatism; strong support for correlated evolution was found for increased eye size and an arboreal lifestyle. Habitat transitions from humid to dry and from terricolous to arboreal occurred more frequently, and ancestors of most lineages were predicted to be terricolous or humid‐forest adapted. We conclude that most Malagasy ant clades first colonized humid forests and subsequently transitioned into dry forests, indicating that previous hypotheses on the evolution of Madagascar's hyperdiverse biota may not apply to ants and other arthropods.     

Anthropogenic disturbance changes the structure of arboreal tropical ant communities

We investigated the influence of anthropogenic disturbance on the structure of arboreal Formicidae communities in SE Asian lowland forests. Included were a primary forest and three disturbed forests which had been cut for crop planting and abandoned 5. 15, and 40 yr after agricultural use for natural regeneration. Ant communities of at least 10 individuals of one tree species were sampled from each forest type by fogging. Diversity and community structure differed clearly among forest types. During the course of forest regeneration ant communities became more and more similar to those of the primary forest. A surrogate analysis shows that ant communities of the primary forest cannot be distinguished from randomly composed communities. This is in contrast to the theoretical expectations according to which ant communities should be structured by interspecific competition that lead to a large degree of predictability. However, a deterministic pattern of ant communities is found in the disturbed forest. This indicates that human disturbance not only changes the faunistic composition of ant communities but could also change the dynamics of the whole system. The transition from stochastic to deterministic communities might be of general importance for understanding the mechanisms structuring communities in disturbed habitats.     

Challenges to the generality of WBE theory

The West, Brown and Enquist (WBE) theory has attracted great interest because it makes general predictions about scaling of ecological processes with body size. Recent research by Muller-Landau and co-workers challenges the generality of this theory by showing that demographic processes in natural forests do not scale in the way that the theory predicts. For WBE theory to be relevant to plant community dynamics, more complex models are required to deal with the influences of competition for light, nutrient supply and disturbance experienced by such communities.     

Plant–soil feedbacks and the coexistence of competing plants

Plant–soil feedbacks can have important implications for the interactions among plants. Understanding these effects is a major challenge since it is inherently difficult to measure and manipulate highly diverse soil communities. Mathematical models may advance this understanding by making the interplay of the various processes affecting plant–soil interaction explicit and by quantifying the relative importance of the factors involved. The aim of this paper is to provide a complete analysis of a pioneering plant–soil feedback model developed by Bever and colleagues (J Ecol 85: 561–573, 1997; Ecol Lett 2: 52–62, 1999; New Phytol 157: 465–473, 2003) to fully understand the range of possible impacts of plant–soil feedbacks on plant communities within this framework. We analyze this model by means of a new graphical method that provides a complete classification of the potential effects of soil communities on plant competition. Due to the graphical character of the method, the results are relatively easy to obtain and understand. We show that plant diversity depends crucially on two key parameters that may be viewed as measures of the intensity of plant competition and the direction and strength of plant–soil feedback, respectively. Our analysis provides a formal underpinning of earlier claims that plant–soil feedbacks, especially when they are negative, may enhance the diversity of plant communities. In particular, negative plant–soil feedbacks can enhance the range of plant coexistence by inducing competitive oscillations. However, these oscillations can also destabilize plant coexistence, leading to low population densities and extinctions. In addition, positive feedbacks can allow locally stable forms of plant coexistence by inducing alternative stable states. Our findings highlight that the inclusion of plant–soil interactions may fundamentally alter the predictions on the structure and functioning of above-ground ecosystems. The scenarios presented in this study can be used to formulate hypotheses about the ways soil community effects may influence plant competition that can be tested with empirical studies. This will advance our understanding of the role of plant–soil feedback in ecological communities.     

Analyzing the Spatial Structure of Broughtonia cubensis (Orchidaceae) Populations in the Dry Forests of Guanahacabibes,Cuba

A current key issue in ecology is the role of spatial effects on population and community dynamics. In this paper, we test several hypotheses related to spatial structures and coexistence of epiphytic tropical orchid species with special emphasis on the endemic species Broughtonia cubensis. More specifically, we explored the spatial structure of orchid–host plant communities at three different levels of organization (occupied vs. nonoccupied host trees, trees with B. cubensis vs. other orchids, and reproductive vs. nonreproductive B. cubensis plants). We mapped all potential host trees and orchids at three 20 × 20 m plots and applied techniques of spatial point pattern analysis such as mark connection and mark correlation functions to evaluate departures from randomized communities. We found spatial aggregation of trees with epiphytic orchids and segregation between trees with and without epiphytic orchids, and that there was an intraspecific spatial aggregation of B. cubensis in relation to the other seven epiphytic orchid species. Furthermore, we found spatial aggregation of reproductive B. cubensis individuals and segregation between reproductive and nonreproductive individuals on their phorophytes. Thus, orchid–host plant communities show hierarchical spatial structuring with aggregation and segregation at different levels of organization. Our results point to an enhancement of local species in the coexistence of tropical epiphytic orchid communities, by reducing competition through niche differentiation.     

Competitive Asymmetry Reduces Spatial Effects on Size-Structure Dynamics in Plant Populations

The growth of each individual in plant populations was simulatedby a spatial competition model for five density levels and fourdifferent spatial distribution patterns of individuals, varyingfrom highly clumped to regular. The simulation results wereanalysed using the diffusion model for evaluating the effectsof density and distribution pattern on the size-structure dynamicsin relation to the degree of competitive asymmetry. At low densities,changes in statistics of plant weight over time such as mean,coefficient of variation, skewness, and Box-Cox-transformedkurtosis differed greatly among spatial patterns, irrespectiveof the degree of competitive asymmetry. In completely symmetriccompetition, the spatial effect on size-structure dynamics remainedrelatively large irrespective of densities, although mean plantweight became similar among the spatial patterns with increasingdensity. However, the spatial effect diminished with increaseddensity in strongly asymmetric competition, when similar sizedistributions were realized irrespective of the spatial patterns.Therefore, it was concluded that: (1) irrespective of the degreeof competitive asymmetry, spatial pattern is important for size-structuredynamics at low densities; (2) spatial pattern is nearly immaterialunder strongly asymmetric competition at high densities; and(3) under crowded conditions, neighbourhood effects are muchmore apparent at the population level in less asymmetric competition.These processes and outcomes are linked to the forms of thefunctions of mean growth rate of individuals [G(t,x) function]and variance in growth rate [D(t,x) function]. These functionsare variable depending on the spatial pattern under symmetriccompetition, but are rather stable under strongly asymmetriccompetition at high densities irrespective of the spatial patterns.Therefore, size structure under strongly asymmetric competitioncan be regarded as a stable system, whereas that under symmetriccompetition is regarded as a variable system in relation tothe spatial pattern and process. From this, it was inferredthat: (1) the goodness-of-fit of spatial competition modelsfor crowded plant populations is higher in less asymmetric competition;and (2) higher species diversity in plant communities is associatedwith the lower degree of competitive asymmetry.Copyright 1994,1999 Academic Press Asymmetric competition, diffusion model, neighbourhood effect, size-structure stability, spatial competition model, spatial distribution pattern, species diversity, symmetric competition     

Structured environments foster competitor coexistence by manipulating interspecies interfaces

Natural environments, like soils or the mammalian gut, frequently contain microbial consortia competing within a niche, wherein many species contain genetically encoded mechanisms of interspecies competition. Recent computational work suggests that physical structures in the environment can stabilize local competition between species that would otherwise be subject to competitive exclusion under isotropic conditions. Here we employ Lotka-Volterra models to show that interfacial competition localizes to physical structures, stabilizing competitive ecological networks of many species, even with significant differences in the strength of competitive interactions between species. Within a limited range of parameter space, we show that for stable communities the length-scale of physical structure inversely correlates with the width of the distribution of competitive fitness, such that physical environments with finer structure can sustain a broader spectrum of interspecific competition. These results highlight the potentially stabilizing effects of physical structure on microbial communities and lay groundwork for engineering structures that stabilize and/or select for diverse communities of ecological, medical, or industrial utility.     

热带森林植物功能群及其动态研究进展

热带森林极高的物种多样性和结构复杂性给生态学研究带来了很多挑战。植物功能群是对特定环境响应相似或对主要的生态过程具有相似作用的物种组合。应用植物功能群的方法是有效减少热带森林群落复杂性,并揭示其格局和过程的良好途径。在介绍植物功能群概念和划分途径的基础上,分析了热带森林植物功能群的时空动态规律。一般来讲,划分植物功能群通常有3种途径,并可通过5个步骤来完成。热带森林植物功能群的空间分布常受景观格局的制约,而环境异质性往往是影响不同植物功能群组配比例变化的直接原因。不同类型的植物功能群随演替过程发生显著的替代,而干扰体系和全球气候变化对功能群的动态过程具有重要的驱动作用。以功能群为基础的动态模型在模拟热带林群落动态和预测植被潜在分布等方面具有广阔的发展前景。探索有效的植物功能分类方法、构建完善的植物功能性状数据库、开发更为精确的功能群动态模型以及加强基于景观水平的植物功能群动态机制的认识等是未来热带森林植物功能群及其动态研究的重要方向。     

Reproductive allocation patterns in different density populations of spring wheat

The effects of increased intraspecific competition on size hierarchies （size inequality） and reproductive allocation were investigated in populations of the annual plant, spring wheat （Triticurn aestivurn）. A series of densities （100, 300, 1 000, 3 000 and 10 000 plants/m^2） along a gradient of competition intensity were designed in this experiment. The results showed that average shoot biomass decreased with increased density. Reproductive allocation was negatively correlated to Gini coefficient （R^2 = 0.927）, which suggested that reproductive allocation is inclined to decrease as size inequality increases. These results suggest that both vegetative and reproductive structures were significantly affected by intensive competition. However, results also indicated that there were different relationships between plant size and reproductive allocation pattern in different densities. In the lowest density population, lacking competition （100 plants/m^2）, individual reproductive allocation was size independent but, in high density populations （300, 1 000, 3 000 and 10 000 plants/m^2）, where competition occurred, individual reproductive allocation was size dependent： the small proportion of larger individuals were winners in competition and got higher reproductive allocation （lower marginal reproductive allocation; MRA）, and the larger proportion of smaller individuals were suppressed and got lower reproductive allocation （higher MRA）. In conclusion, our results support the prediction that elevated intraspecific competition would result in higher levels of size inequality and decreased reproductive allocation （with a negative relationship between them）. However, deeper analysis indicated that these frequency- and size-dependent reproductive strategies were not evolutionarily stable strategies.     

Effect of Canopy Structure on Degree of Asymmetry of Competition in Two Forest Stands in Northern Japan

The canopy structure in terms of the vertical distribution of leaf mass and the degree of asymmetry of competition between individual trees was studied in two types of forest stand in Hokkaido, northern Japan: a naturally regenerated stand ofBetulaspp. and an artificial plantation ofPicea abies.The canopy structure in theBetulastand was more hierarchical; larger individuals were not heavily shaded even in the lowest part of their crowns and smaller individuals were heavily shaded by their larger neighbours. The canopy structure in thePiceastand was less hierarchical; even larger individuals were shaded in the lowest part of their crowns and smaller individuals were not heavily shaded by their neighbours. Application of the general formula of size-dependent mean growth rate revealed that competition in theBetulastand was more one-sided than that in thePiceastand. This result was consistent with the trends in the change over time in size equality in both stands.Even if competition is mediated by light, which often makes competition one-sided, the degree of one-sidedness in competition can be variable depending on canopy structure.     

The nutrient-load hypothesis: patterns of resource limitation and community structure driven by competition for nutrients and light

Resource competition theory predicts that the outcome of competition for two nutrients depends on the ratio at which these nutrients are supplied. Yet there is considerable debate whether nutrient ratios or absolute nutrient loads determine the species composition of phytoplankton and plant communities. Here we extend the classical resource competition model for two nutrients by including light as additional resource. Our results suggest the nutrient-load hypothesis, which predicts that nutrient ratios determine the species composition in oligotrophic environments, whereas nutrient loads are decisive in eutrophic environments. The underlying mechanism is that nutrient enrichment shifts the species interactions from competition for nutrients to competition for light, which favors the dominance of superior light competitors overshadowing all other species. Intermediate nutrient loads can generate high biodiversity through a fine-grained patchwork of two-species and three-species coexistence equilibria. Depending on the species traits, however, competition for nutrients and light may also produce multiple alternative stable states, suppressing the predictability of the species composition. The nutrient-load hypothesis offers a solution for several discrepancies between classical resource competition theory and field observations, explains why eutrophication often leads to diversity loss, and provides a simple conceptual framework for patterns of biodiversity and community structure observed in nature.     

Effects of variation in individual growth on plant species coexistence

Abstract. Both size structure and variability (spatial heterogeneity, disturbance, stochasticity, variation in species attributes, etc.) are regarded as regulatory mechanisms of species coexistence. However, none of the models so far proposed consider both size structure and variability simultaneously. A size-structured variation model for plant-community dynamics is proposed, which is based on the diffusion model for growth dynamics of plant populations. This model has four functions: (1) mean growth rate of individuals of size x at time t, G(t, x) (species-specific mean traits, e.g. competitive ability); (2) variance in growth rate of individuals of size x at time t, D(t, x) (stochastic factors due to genetic variation, environmental heterogeneity, spatial variation of individuals, etc.); (3) mortality rate of individuals of size x at time t, M(t, x); and (4) recruitment rate at time t, R(t), as a boundary condition. The interference function for individuals of size x at time t, C(t, x), is introduced, which expresses the degree of interactions between individuals and hence averaged effects of local neighbourhood competition; the G(t, x), D(t, x), M(t, x) and R(t) functions are given in terms of C(t, x). These four functions describe the growth dynamics of individuals of each species in the plant community. Effects of the G(t, x), D(t, x), M(t, x) and R(t) functions on species coexistence in plant communities were evaluated by simulation and the relative importance of the D(t, x) function as well as size structure was shown for species coexistence especially in plant communities where competition among species is non-transitive or niche limitation does not work.     

贵州桫椤群落的初步研究

桫椤(Alsophila spinulosa)是国家一级保护植物,在贵州分布较多,局部地段形成以桫椤为主的植物群落。本文就贵州赤水县金沙沟桫椤群落的区系组成、群落外貌、结构等特征进行了初步研究。通过分析表明赤水桫椤群落是保存完好、结构典型、类型多样的珍贵群落,应注意很好保护。     

Self-recovery of plant communities: examples, mechanisms, and approaches for description

Autogenic (self-recovering) plant communities that need no transitional stages of succession for their recovery after damage are widespread in different areas of the Globe. They are typical for tundras, but also common in boreal forests, alpine belts, deserts and Mediterranean biomes. Such communities usually dominate the landscape. The authors examine in details the communities of cottongrass (Eriophorum vaginatum) wet tussock tundras of North-Eastern Asia and Alaska, the dryad (Dryas punctata) lichen dry patchy tundras of the same region and Scots pine (Pinus sylvestris) lichen-feathermoss forests on fluvial-glacial deposits of Fennoscandia and Russian Plain. Although authogenic plant communities are serial, they exist side by side with proper edaphic variants of climaxes but don't appear to be replaced by the latter, being even more stable. Moreover, they even seem to "compete" for the space with the climax communities. Stabilization of autogenic communities is primarily induced by the impact of abiotic disturbance factors, like periodic fires, permafrost action or scree sliding, and afterwards maintained by several mechanisms of community structure organization. Such mechanisms of self-recovery act at the levels of both plant population and community as a whole. Specific adaptions of dominant species to typical kinds of disturbance (at a scale of either the community, or its local parts) illustrate the first level, whereas close co-adaptations between dominants and accompanying species refer to the second one. Such co-adaptations are present as complementarity of plant life strategies and growth forms within a community and/or as a shifting-mosaic steady state which leads to quick recovery of local disturbances. Among different possible approaches to simulation of community autogenesis those of combinatorial simulation seem to be especially perspective.     

Plant communities in broad-leaved deciduous forests in Hordaland county, Western Norway

The vegetation of broad-leaved deciduous forests in Hordaland, Western Norway is described with regards to species composition and structure. The investigation is based on phytosociological analyses of native forest stands dominated by Corylus avellana, Fraxinus excelsior, Ulmus glabra , or Tilia cordata. Two-way-indicator analysis (TWINSPAN) and correspondence analysis (CA) are used to distinguish different vegetation types and assess possible gradients in the vegetation data. The vegetation types are discussed in relation to equivalent forest communities and to syntaxonomical units. Two forest types are described at the first hierarchical TWINSPAN level: 1) A hygrophilous and slightly thermophilous Fraxinus excelsior-Cirriphyllum piliferum forest characterised by an open tree canopy, a dense field layer of tall herbs and ferns, and a high cover of bryophytes. 2) A thermophilous and less hygrophilous Corylus avellana-Brachypodium sylvaticum forest characterised by a dense tree layer, a more open field layer with larger elements of small herbs, and a somewhat lower bryophyte cover. The CA analysis clearly separates the samples from the first TWINSPAN division along the first ordination axis. Five forest types have been described at the second hierarchical level, mainly associated with differences in mesotrophic and eutrophic species, but there is clearly a gradient structure in the species composition between these plant communities. In relation to syntaxonomy, the first TWINSPAN division supports the separation of the west Norwegian broad-leaved deciduous forests into a hygrophilous plant community, the Eurhynchio-Fraxinetum (Blom 1982) Øvstedal 1985 and a drier and more thermophilous community, the Primulo-Ulmetum (Blom 1982) Øvstedal 1985.