Competition and coexistence in plant communities

Few ecologists today doubt that competition is an important structuring factor in plant communities, but researchers disagree on the circumstances where it is most intense, and on which traits can be considered to contribute to competitive ability in different species. The distinction between a species' effect on resources and its response to reduced resource levels might help to solve these questions. Whereas classical competition theory predicts competitive exclusion of species with similar requirements, recent ideas stress that species diversity may be explained by a multitude of processes acting at different scales, and that similarities in competitive abilities often may facilitate coexistence.     

Positive interactions, discontinuous transitions and species coexistence in plant communities

The population and community level consequences of positive interactions between plants remain poorly explored. In this study we incorporate positive resource-mediated interactions in classic resource competition theory and investigate the main consequences for plant population dynamics and species coexistence. We focus on plant communities for which water infiltration rates exhibit positive dependency on plant biomass and where plant responses can be improved by shading, particularly under water limiting conditions. We show that the effects of these two resource-mediated positive interactions are similar and additive. We predict that positive interactions shift the transition points between different species compositions along environmental gradients and that realized niche widths will expand or shrink. Furthermore, continuous transitions between different community compositions can become discontinuous and bistability or tristability can occur. Moreover, increased infiltration rates may give rise to a new potential coexistence mechanism that we call controlled facilitation.     

Experimental manipulation of natural plant communities

Attempts to elucidate the factors controlling the structure of plant communities have relied increasingly on field experiments. This is a powerful approach for testing theoretical predictions that offers important advantages over observational and comparative studies. However, field experiments suffer from intrinsic difficulties as well as more-easily remediable limitations. Recent progress has been made by new approaches including the use of multifactor experiments, and the development and dissemination of better statistical tools.     

Interacting coexistence mechanisms in annual plant communities: Frequency-dependent predation and the storage effect

We study frequency-dependent seed predation (FDP) in a model of competing annual plant species in a variable environment. The combination of a variable environment and competition leads to the storage-effect coexistence mechanism (SE), which is a leading hypothesis for coexistence of desert annual plants. However, seed predation in such systems demands attention to coexistence mechanisms associated with predation. FDP is one such mechanism, which promotes coexistence by shifting predation to more abundant plant species, facilitating the recovery of species perturbed to low density. When present together, FDP and SE interact, undermining each other’s effects. Predation weakens competition, and therefore weakens mechanisms associated with competition: here SE. However, the direct effect of FDP in promoting coexistence can compensate or more than compensate for this weakening of SE. On the other hand, the environmental variation necessary for SE weakens FDP. With high survival of dormant seeds, SE can be strong enough to compensate, or overcompensate, for the decline in FDP, provided predation is not too strong. Although FDP and SE may simultaneously contribute to coexistence, their combined effect is less than the sum of their separate effects, and is often less than the effect of the stronger mechanism when present alone.     

Competition and coexistence in plant communities: intraspecific competition is stronger than interspecific competition

Theory predicts that intraspecific competition should be stronger than interspecific competition for any pair of stably coexisting species, yet previous literature reviews found little support for this pattern. We screened over 5400 publications and identified 39 studies that quantified phenomenological intraspecific and interspecific interactions in terrestrial plant communities. Of the 67% of species pairs in which both intra‐ and interspecific effects were negative (competitive), intraspecific competition was, on average, four to five‐fold stronger than interspecific competition. Of the remaining pairs, 93% featured intraspecific competition and interspecific facilitation, a situation that stabilises coexistence. The difference between intra‐ and interspecific effects tended to be larger in observational than experimental data sets, in field than greenhouse studies, and in studies that quantified population growth over the full life cycle rather than single fitness components. Our results imply that processes promoting stable coexistence at local scales are common and consequential across terrestrial plant communities.     

A mathematical approach to spatial distribution and temporal succession in plant communities

The Volterra equations which represent competitions between two species are utilized to examine the phenomenon of boundary formation between two species of plants. The set of stable stationary points for these equations is determined and is illustrated in a product space of parameters and dynamical variables. The stages of boundary appearance and succession are visualized by considering slow changes of the parameters as functions of time and space.     

Temperature-mediated coexistence in temperate forest ant communities

Patterns of ant species diversity are well documented and yet the mechanisms promoting species coexistence among communities are often elusive. Two emerging hypotheses that account for coexistence in ant communities are the discovery-dominance tradeoff and the dominance-thermal tolerance tradeoff. Here we used behavioural assays and community-level sampling from ant assemblages in the southern Appalachians, USA to test for the discovery-dominance and dominance-thermal tolerance tradeoffs. Species that were behaviorally dominant during interspecific interactions tended to forage in a narrow window of generally warmer temperatures, whereas subordinate species tended to forage in a wide range of temperatures, including colder temperatures. Species that foraged at lower temperature tended to be behaviourally subordinate, suggesting that a dominance-thermal tolerance tradeoff promotes coexistence in this system. Species richness was positively related to site average annual temperature and within-site variation in ground temperature, suggesting that temperature also shapes the structure of ant communities and regulates diversity. There was no relationship between the ability of a species to discover food resources and its behavioural dominance, contrary to the predictions of the discovery-dominance tradeoff hypothesis. In sum, our results show that temperature plays numerous roles in promoting regional coexistence in this system.     

Dispersal and life span spectra in plant communities: a key to safe site dynamics, species coexistence and conservation

Dispersal and life span of individual plant species within five plant communities were assessed to obtain a characterization of these communities in this respect. Such a characterization is important in the context of restoration and maintenance. The most frequent species of five communities were ranked in eight classes according to their level of seed dispersal capability, their seed bank formation (dispersal in time and space) and their individual life span. In the communities, all eight classes were found, but communities differed in the distribution of the species over the classes.
A theoretical framework was constructed to use the level of specialization of plant species in terms of dispersal in space and time, and life span, to define the characteristics of safe site dynamics within communities. Following simple rules, the relative reliability of the occurrence of safe sites in space and time was defined. After that, the relative reliability of the habitat was linked to the best fitting combination of trait specialization level. Having defined this link, communities could be characterized in a comparative way by their level and pattern of reliability of the opportunities for recruitment in space and time.
The meaning of the coexistence of a range of trait combinations in one community was discussed. It was postulated that habitat reliability can explain this by assuming that the habitat of the community is part of a larger system, or consists of several “subsystems”. These insights need to be considered in nature conservation. Succession and also specializations beyond the scope of dispersal and life span may influence the occurrence of species in a seemingly unfit habitat (for instance the occurrence of semi parasitic annuals in a community of perennials, because they use the perennial root system of other species). Finally, the meaning of safe site reliability in space and time in the context of restoration of communities was discussed. The reliability in space and time may be different today from that of the past, which restricts the feasibility of restoration of communities.     

Experimental analysis of patch dynamics in tallgrass prairie plant communities

Abstract. Previous research has indicated that patch structure at small spatial scales (<100m²) in tallgrass prairies was defined by a diverse array of infrequent species because dominant species occurred in all samples at this scale. Also, patch structure was not significantly different from that derived from random species associations. Based on these results, we hypothesized that remo val of a dominant species would have no effect on patch structure in these prairies. We tested this hypothesis by removing a dominant grass, Schizachyrium scoparium (Poaceae), from half of each of four 10 m × 10 m study blocks, and comparing differences in patch structure between control and removal halves before and after removal. The minimum resolution in our study was 1 m². Patches of similar species composition were defined by cluster analysis of presence/absence data and cover data. Patch sizes ranged from 1 to 34 m². Following the removal of S. scoparium there was an overall increase in the number of species in the removal half of each block compared to pre-treatment levels. However, the number of patch types and number of spatially mapped groups, based on presence/absence or cover data, did not change between control and removal plots after the removal of S. scoparium. This supports the hypothesis that removal of a large, dominant species would have no effect on patch structure at this scale of resolution in these prairies. Thus, patch structure, as defined here, is an emergent property in these grasslands that is not predictable from changes in species composition. This property of stochastic patch structure results from interactions of processes operating at scales both larger and smaller than our scale of resolution. Stochastic models may provide a reasonable approach to modelling small-scale patch dynamics in tallgrass prairie communities.     

生态群落物种共存的进化机制

本文概述了目前对生态群落的物种共存研究中存在的若干问题及动、植物群落物种共存机制的研究进展。植物群落的物种共存主要介绍与环境、种子再迁移、生态位分化、竞争平衡理论、种库假设、再生生态位等有关的几种假设、生态学上相似种的共存及“原”群落概念等。动物群落的物种共存机制主要从以下几方面叙述：（1）异质环境中的资源分割,主要指动物斑状滋养的不同利用;（2）避免竞争排斥的行为机制,如边缘效应、聚群效应、扩散行为、相互作用和干扰;（3）特化者和泛化者的共存,包括：竞争是物种向多功能进化的作用力、最佳觅食理论与生态学特化及特化概念的发展。最后指出进一步研究的方向。     

Competition and coexistence in submerged aquatic plant communities: the effects of species interactions versus abiotic factors

SUMMARY. 1. Aquatic plant biomass and species composition were studied at three sites in Long Lake, Alberta, Canada, to examine the role of biotic and abiotic factors in determining species diversity.
2. Results of controlled in situ experiments to test for interspecific competition showed that biomass of plants in four different mixed communities did not increase in response to selected species removals ( P >0.1, n =16).
3. In contrast, biomass of the dominant taxa (Myriophyllum exalbescens, Ceratophyllum demersum and Chara sp.) were correlated ( P <0.05) with abiotic factors (i.e. distance from shore, water depth, sediment exchangeable phosphorus concentration and/or sediment organic content).
4. These results suggest that interspecific competition between naturally coexisting species of submerged aquatic plants is slight and that spatial heterogeneity or differential utilization of abiotic resources promotes species diversity in submerged plant communities.     

Experimental evidence for long-term coexistence of copiotrophic and oligotrophic bacteria in pelagic surface seawater

Most marine copiotrophic bacteria can produce extracellular enzymes to degrade biopolymers into bio-available smaller solutes, while oligotrophic bacteria usually cannot. Bacterial extracellular enzymes and enzymatic products can be a common resource that could be utilized by both copiotrophs and oligotrophs; when present, oligotrophs may outcompete the enzyme-producing copiotrophs. However, copiotrophs and oligotrophs consistently coexist in the ocean. How they maintain coexistence has still not been experimentally studied. In this study, the interaction and coexistence of a copiotroph and an oligotroph, isolated from the same surface seawater sample and utilizing the same proteinaceous substrate, were experimentally investigated. The copiotroph could secrete extracellular proteases to degrade and then utilize the proteinaceous substrate. The oligotroph was unable to utilize the proteinaceous substrate by itself, but could grow by using the hydrolysate amino acids. The copiotroph outcompeted the oligotroph by adsorbing the amino acids quickly and having a higher growth rate in the rich medium. The oligotroph survived by adapting to low concentration of nutrients. The copiotroph and oligotroph were able to maintain long-term (up to 142 days) coexistence in the laboratory. This study indicates that differences in the utilization of different concentrations of nutrients can drive the coexistence of marine copiotrophs and oligotrophs.     

An approach to modelling in immunology

Like most other fields in biology, immunology has been revolutionised by the techniques of molecular biology and the resulting explosion in available experimental data. It is argued that efforts to integrate the data to gain insight into how various subsystems in the immune system interact and function require mathematical modelling and computer simulation in close collaboration with experimentalists. This paper illustrates some of the techniques available for modelling immune systems, and highlights the issues that should be borne in mind by anyone starting down the modelling path.     

Experimental evidence for the immediate impact of fertilization and irrigation upon the plant and invertebrate communities of mountain grasslands

The response of montane and subalpine hay meadow plant and arthropod communities to the application of liquid manure and aerial irrigation – two novel, rapidly spreading management practices – remains poorly understood, which hampers the formulation of best practice management recommendations for both hay production and biodiversity preservation. In these nutrient-poor mountain grasslands, a moderate management regime could enhance overall conditions for biodiversity. This study experimentally assessed, at the site scale, among low-input montane and subalpine meadows, the short-term effects (1 year) of a moderate intensification (slurry fertilization: 26.7–53.3 kg N·ha⁻¹·year⁻¹; irrigation with sprinklers: 20 mm·week⁻¹; singly or combined together) on plant species richness, vegetation structure, hay production, and arthropod abundance and biomass in the inner European Alps (Valais, SW Switzerland). Results show that (1) montane and subalpine hay meadow ecological communities respond very rapidly to an intensification of management practices; (2) on a short-term basis, a moderate intensification of very low-input hay meadows has positive effects on plant species richness, vegetation structure, hay production, and arthropod abundance and biomass; (3) vegetation structure is likely to be the key factor limiting arthropod abundance and biomass. Our ongoing experiments will in the longer term identify which level of management intensity achieves an optimal balance between biodiversity and hay production.     

The pairwise approach to analysing species co‐occurrence

The analysis of species co‐occurrence patterns continues to be a main pursuit of ecologists, primarily because the coexistence of species is fundamentally important in evaluating various theories, principles and concepts. Examples include community assembly, equilibrium versus non‐equilibrium organization of communities, resource partitioning and ecological character displacement, the local–regional species diversity relationship, and the metacommunity concept. Traditionally, co‐occurrence has been measured and tested at the level of an entire species presence–absence matrix wherein various algorithms are used to randomize matrices and produce statistical null distributions of metrics that quantify structure in the matrix. This approach implicitly recognizes a presence–absence matrix as having some real ecological identity (e.g. a set of species exhibiting nestedness among a set of islands) in addition to being a unit of statistical analysis. An emerging alternative is to test for non‐random co‐occurrence between paired species. The pairwise approach does not analyse matrix‐level structure and thus views a species pair as the fundamental unit of co‐occurrence. Inferring process from pattern is very difficult in analyses of co‐occurrence; however, the pairwise approach may make this task easier by simplifying the analysis and resulting inferences to associations between paired species.     

An evolutionary modelling approach to understanding the factors behind plant invasiveness and community susceptibility to invasion

Ecologists have had limited success in understanding which introduced species may become invasive. An evolutionary model is used to investigate which traits are associated with invasiveness. Translocation experiments were simulated in which species were moved into similar but evolutionarily younger communities. The main findings were that species that had previously been the most abundant in their original communities have significantly higher rates of establishment than did species that had previously occurred at low abundance in their original community. However, if establishment did occur, previously abundant and previously low-abundant species were equally likely to become dominant and were equally likely to exclude other species from their new community. There was a suggestion that the species that were most likely to establish and exclude others were 'genetically' different. When species that had evolved in different simulations (but with identical environmental conditions) were transplanted into communities that had also evolved in different simulations of the same conditions, the outcomes were difficult to predict. Observed rates of establishment and subsequent competitive dominance were observed to be species- and community combination-specific. This evolutionary study represents a novel in silico attempt to tackle invasiveness in an experimental framework and may provide a new methodology for tackling these issues.     

Improved modelling of the impacts of sea level rise on coastal wetland plant communities

This study presents an enhanced methodology for modelling the impacts of sea level rise on coastal wetlands. The tool integrates dGPS-calibrated LiDAR data, isostatic uplift and sediment accretion rates to predict the location and extent of plant communities at three non-contiguous micro-topographical coastal wetlands in Estonia by 2100 in response to global sea level rise. Scenarios were run including sediment accretion, elevated sediment accretion and then discounting sediment accretion and dGPS calibration for comparison. Results showed an increase in surface elevation (related to sediment accretion and isostatic uplift) resulting in a decrease in local sea level in the majority of sites and scenarios in the north of the country, although a rise in local sea level is predicted in sites with limited allochthonous sediment supply, predominantly impacting higher elevation plant communities. Wetlands situated on the west coast are likely to maintain equilibrium with sea level as a result of lower sedimentation and isostatic uplift than more northerly sites. This study shows that dGPS-calibrated LiDAR data and sediment accretion are essential to maintain model validity in Baltic coastal wetlands due to their low relief and could considerably improve current sea level rise impact models for other regions.