Competition as a demolition derby: why tolerating competitors is more important than suppressing them

Tolerance and suppression are distinct components of competition among plants, and recognizing how they affect competitive outcomes is important for understanding the mechanisms and consequences of competition. We used simulations informed by experimental trials to ask whether tolerance or suppression of competitors was more important for the survival of native plants experiencing competition with an exotic invasive species. When competition was pairwise, tolerance and suppression contributed equally to competitive rank in simulations. However, when multiple native genotypes competed together against an invader, the ability to tolerate competition was up to 50 times more important than the ability to suppress the invader. In two-competitor communities the chief advantage of suppressing competitors was a global decrease in their abundance, but this advantage did not exist in communities of multiple competitors – which is more representative of natural conditions – because decreased competitor abundance benefited all plants regardless of their competitive ability. We suggest that this concept is analogous to a ‘demolition derby,' an automotive contest where participants attempt to have the last functional vehicle on the playing field. Because strong suppressors share the benefits of eliminating competitors with other remaining competitors, we propose that tolerance of competitors is more beneficial than suppression when competition occurs in a multiplayer scenario – in a demolition derby and in nature. This finding has implications for our understanding of how competition influences plant species coexistence, plant community structure and invasion outcomes.     

EVOLUTION OF BROAD AND SPECIFIC COMPETITIVE ABILITY IN NOVEL VERSUS FAMILIAR ENVIRONMENTS IN DROSOPHILA SPECIES

We used nine pairs of competing Drosophila melanogaster and Drosophila simulans populations to test three hypotheses. (1) Weaker competitors undergo greater evolutionary increases in competitive ability, compared with stronger ones. (2) Increased competitive ability against a specific competitor population causes a correlated increase in competitive ability against other competitor populations. (3) In a novel environment, adaptation to the abiotic environment contributes more to competitive ability than adaptation to the competitor population. After 11 generations of competition, initially weaker competitor populations showed relatively greater increases in competitive ability. Broad and specific competitive abilities, the latter being specific to a particular competitor population, were positively correlated in both familiar and novel environments. Adaptation to the abiotic environment seemed to be a more important component of competitive ability in the novel environments. We conclude that in geographically structured species, biotic and abiotic factors affecting the evolution of competitive ability may interact to help create a mosaic of outcomes that can affect the evolutionary dynamics of the interaction over the range of the competing species.     

Competitive exclusion of a worldwide invasive pest by a native. Quantifying competition between two phytophagous insects on two host plant species

1. High competitive ability is believed to be an important characteristic of invasive species. Many animal studies have compared the competitive ability of invasive species with a native species that is being displaced, but few have looked at systems where an invasive species has failed to establish itself. These types of studies are important to determine if competition is relevant not only to invading species but also to the biotic resistance of a community. 2. The thrips species F. occidentalis is a highly invasive pest that has spread from its original range (the western states of the USA) to a worldwide distribution. Despite this, F. occidentalis is largely absent or occurs in low numbers in the eastern states of the USA, where the native F. tritici dominates. It is possible that F. tritici is competitively excluding F. occidentalis from this region. 3. Larval competition between these two thrips species was tested on two known plant hosts, Capsicum annuum (a crop plant), and Raphanus raphanistrum (an invasive weed), using a response surface design with number of larvae surviving as the response variable. The response surface design allowed competition models to be fit to data using maximum likelihood estimation, thus generating quantitative values for interspecific competition. 4. On both plant hosts, the native F. tritici did not experience significant interspecific competition from the invasive F. occidentalis. In contrast, F. occidentalis did experience significant interspecific competition from F. tritici. Competition from F. tritici larvae on F. occidentalis larvae was estimated to be 1.72 times (on C. annuum) and 1.76 times (on R. raphanistrum) the effect of intraspecific competition. The invasive F. occidentalis appears to be competitively excluded by the native F. tritici. 5. This study confirms the importance of competition in the biotic resistance of a community and is one of the few animal studies to not only test for competition in an apparently resistant ecosystem but also to quantify the level of interspecific competition between two animal species.     

Differential effect of inter- and intraspecific competition on the performance of invasive and native Taraxacum species

Inter- and intraspecific competitive abilities are significant determinants of invasive success and the ecological impact of non-native plants. We tested two major hypotheses on the competitive ability of invasive species using invasive (Taraxacum officinale) and native (T. platycarpum) dandelions: differential interspecific competitive ability between invasive and native species and the kin recognition of invasive species. We collected seeds from two field sites where the two dandelion species occurred nearby. Plants were grown alone, with kin (plants from the same maternal genotype) or strangers (plants from different populations) of the same species, or with different species in a growth chamber, and the performance at the early developmental stage between species and treatments was compared. The invasive dandelions outcompeted the native dandelions when competing against each other, although no difference between species was detected without competition or with intraspecific competition. Populations of native species responded to interspecific competition differently. The effect of kinship on plant performance differed between the tested populations in both species. A population produced more biomass than the other populations when grown with a stranger, and this trend was manifested more in native species. Our results support the hypothesis that invasive plants have better competitive ability than native plants, which potentially contributes to the establishment and the range expansion of T. officinale in the introduced range. Although kin recognition is expected to evolve in invasive species, the competitive ability of populations rather than kinship seems to affect plant growth of invasive T. officinale under intraspecific competition.     

Genetic variation in tolerance of competition and neighbour suppression in Arabidopsis thaliana

Intraspecific competitive interactions can profoundly influence phenotypic evolution. However, prior studies have rarely evaluated the evolutionary potential of the two components of competitive ability, tolerance of competition and suppression of neighbours. Here, we grow a set of 20 Arabidopsis thaliana recombinant inbred lines in three competitive treatments (noncompetitive, intra‐genotypic competition and inter‐genotypic competition) to examine if there is genetic variation for the components of competitive ability and whether neighbour relatedness has an effect on fitness. We find evidence for genetic variation in tolerance of competition and neighbour suppression and that these two competitive strategies are correlated, such that genotypes that tolerate competition will also strongly suppress neighbours. We further observe that the effect of neighbour relatedness on fitness of target individuals depends on neighbour identity, i.e. whether target individuals perform better when competing against self vs. nonself individuals depends on the genotypic identity of the nonself neighbour. The results are particularly relevant to evolutionary responses under multi‐level selection.     

外来物种入侵后的多物种竞争共存的集合种群模型

基于多物种竞争共存模型,提出外来物种与本地物种竞争共存途径的两种假想:外来物种通过插队竞争与本地物种实现共存;外来物种通过等位竞争与本地物种实现共存.并提出根据外来物种在两种竞争共存模式下占据生境斑块比例的稳定值大小来判断外来物种和本地物种的竞争共存途径.根据两种假想,分别建立了外来物种插队竞争共存模型和等位竞争共存模型.通过应用数学软件Mathematica 4.0对两个模型进行了模拟,得出以下结论:在外来物种与本地物种竞争共存状态下,如果外来物种通过插队竞争与本地物种实现共存,当本地物种竞争力差异较大时,外来物种极易对本地稀少物种构成危害.虽然外来物种不会直接造成本地稀少物种的灭绝,但是会使本地稀少物种的生境斑块急剧减少,增加本地稀少物种灭绝的可能性,而当本地物种竞争力差异较小时,外来物种对本地所有物种的影响都较小.如果外来物种通过等位竞争与本地物种实现共存,无论本地物种竞争力差异大小与否,外来物种只是影响到与其生态位相同的本地物种,影响程度取决于外来物种侵入时所占据生境斑块的比例大小.     

Drosophyllum lusitanicum, an endangered West Mediterranean endemic carnivorous plant: threats and its ability to control available resources

Drosophyllum lusitanicum (L.) Link (Droseraceae), Erva pinheira orvalhada, an endangered carnivorous plant, is a local endemic of clearings in pine, cork oak and oak forests or their successional shrublands in the western Iberian Peninsula and Morocco. The conservation status, distribution and population dynamics of this species are only partially known, both for Spanish and Portuguese occurrences. Portuguese distribution data from herbarium and bibliographic sources were collected for this study. Field work on 50 populations was undertaken in order to improve knowledge on their conservation status and possible threats. Natural causes, infrastructure and housing construction are stressed as the most important threats. Germination tests were carried out and the distribution patterns, plant height and the number of flowers were studied in an arbitrarily chosen population. A relatively high light requirement and low competitive ability to gain light are considered factors likely for the microdistribution pattern. Drosophyllum lusitanicum cannot compete for light in habitats with intense competition. Seeds from D. lusitanicum have reduced probability of germination when adult plants are already growing in the area. It is hypothesized that seed germination shows a similar pattern to flowering, viz over a considerable time rather than in a sudden flush. This strategy may be a key factor for the species' survival, representing attempts to take advantage of available resources, preventing intraspecific competition and, finally, preventing sudden total disappearance of a population, resulting from any catastrophic events due to natural causes or to human activity. As a result of this study, some conservation measures are suggested.   © The Linnean Society of London, Botanical Journal of the Linnean Society , 2002, 140 , 383–390.     

Competitive strategies between Anastrepha fraterculus and Anastrepha obliqua (Diptera: Tephritidae) regulating the use of host fruits

Interspecific competition between individuals of different species can result in reductions in their fecundity, growth or survival, reflecting differential exploitation of resources that become intensified due to spatial co-occurrence, ecological similarity and increased population densities. As two species cannot occupy the same niche, coexistence is only possible if the available resources are used in non-overlapping manners such as niche partitioning or the use of refuges. Among agricultural insect pests, such as fruit flies of the family Tephritidae, competitive interactions can result in competitive displacement, host changes, or the expansion or restriction of the numbers of hosts utilized that can have negative consequences for human agricultural activities. We evaluated the competitive interactions between two fruit fly species of the genus Anastrepha, Anastrepha obliqua (Macquart, 1835) and Anastrepha fraterculus (Wiedmann, 1830), on their respective preferred hosts (mangoes and guava). Experiments of larval competition and competition for ovipositioning sites by adult females were performed to compare the parameters of larval development time, numbers of pupae and emerged adults and numbers of ovipositions in the presence or absence of interspecific competition. We observed that the interactions between those species were asymmetrical and hierarchical, and our results suggest a competitive displacement of A. fraterculus by A. obliqua when those two species are present on the same fruit, whether mangoes or guavas.     

Biodiversity and belowground interactions mediate community invasion resistance against a tall herb invader

Aims Species-rich plant communities are hypothesized to be more resistant against plant invasions because they use resources in a more efficient way. However, the relative contributions of aboveground competition and belowground interactions for invasion resistance are still poorly understood.Methods We compared the performance of Knautia arvensis transplants growing in plots differing in plant diversity both under full competition and with shoots of neighbors tied back to determine the relative strength of aboveground competition in suppressing this test invader without the confounding effect of shading. In addition, we assessed the effects of belowground competition and soil-borne pathogens on transplant performance.Important findings Both aboveground competition and plant species richness strongly and independently affected invader performance. Aboveground biomass, height, leaf mass per area and flowering of transplanted individuals of K. arvensis decreased with increasing species richness of the host community. Species-rich and species-poor communities both imposed equally strong aboveground competition on K. arvensis. However, belowground interactions (especially belowground root competition) had strong negative effects on transplant performance. In addition, the presence of grasses in a plant community further reduced the performance of K. arvensis. Our results suggest that belowground competition can render species-rich host communities more suppressive to newly arriving species, thus enhancing community invasion resistance.     

Resource use and co-existence: Experimental tests from a sequence of different-aged pastures

Summary Competitive relationships were estimated for plants from young and old pasture populations. Experimental treatments were conducted under both common garden field station conditions and actual grazed pasture conditions. The results suggest four conclusions. (1) Grazing and other pasture conditions did not prevent plant competition. (2) Interspecific competition appears to promote the use of different resources (niche divergence). (3) Intraspecific competition may lead to a broadening of resource use by a species and, thus, indirectly increase both niche overlap and interspecific competition. No evidence was found to support an alternative theory of co-existence in which interspecific competition is argued to promote a balancing of competitive abilities without changes to relative niche overlap. (4) Changes in resource use appears to occur within a few decades after pasture formation.     

不同迁移能力和竞争能力的集合种群竞争模式及其模拟

种群竞争是影响生态系统演化的莺要生态过程之一。本文是在徐彩琳和Tilman研究工作的基础上,将竞争系数引入集合种群动力模式,建立了集合种群之间竞争的数学模型,并对5-集合种群的竞争动态进行了计算机模拟研究。结果表明：种群竞争排除与共存受迁移扩散能力和竞争能力影响很大,排除原理在理论上是存在的,在广域集合种群和实际中物种是竞争共存的．共存的条件是其竞争能力与扩散能力呈非线性负相关关系,竞争的结果使物种的强弱序列发生变化。     

Direct Interference on Competition in a Chemostat

Thispaperstudiestheglobaldynamicsofcompetitioninchemostatinwhichtwopopulationsofmicrooganismscompeteexploitativelyforasingle,essential,nonreproducing,growth-limitingsubstrateandthereisadirectinterferencebetweencompetitors.Inordertounderstandthedifferencesintheeffectsofintraspecificandinterspecificinterference,thebothcasesareconsideredrespectively.Keywords:##4Populationdynamicsecology;;chemostat;;competition;;interference;;interspecific;;intraspecific;;principalofcompetitiveexclusion     

Temperature-dependent competition hierarchy: a mechanism stabilizing the phenological strategy in the scorpionfly Panorpa communis L.

Previous studies of the phenologies and the different microclimatic patterns of the distribution of the scorpionflies Panorpa communis L. and Panorpa vulgaris Imhoff and Labram 1836 showed different phenological strategies. In P. communis , a species foraging at shadowy and cool places only, a majority of 90% of the individuals are univoltine; however, approximately 10% of the offspring of the first annual generation are bivoltine. This proportion remained unchanged in the Freiburg population over 8 years. Differently, all individuals of P. vulgaris foraging equally frequent at sunny and warm as at shadowy and cool places are bivoltine. The proximate cause of bivoltinism in both species is a heritable variation of different ‘day length thresholds’ triggering diapause‐free development if natural day length exceeds these thresholds. As selection favours maximal temporal exploitation of food availability it remains obscure why in P. communis the number of diapause‐free developing individuals does not increase continuously from year to year although this phenotype reproduces twice a year. Therefore, in the present paper, we focus on the following main questions. Does the competitive inferiority of P. communis in the presence of P. vulgaris at the temperature regime of the late summer function as a mechanism maintaining the majority of individuals of P. communis univoltine, by dramatically reducing the fitness of the bivoltine ones? As a long‐term evolutionary change in the frequency of bivoltine individuals in P. communis solely depends on the lifetime reproductive success of the females, we here consider the influence of interspecific competition and temperature conditions on the reproductive success of the females of P. communis only. Five lines of evidence suggest that the mechanism of maintaining univoltinism in P. communis is primarily because of differences in the ability of each species to exploit dead arthropod resources: (1) these species show complete diet overlap; (2) dead arthropods are limiting resources for both species of scorpionflies as indicated by positive demographic effects with increased food availability; (3) in competition with P. vulgaris at high temperatures, P. communis is competitively inferior in the ability to detect and exploit dead arthropods; (4) this reduced resource acquisition of female P. communis translates into significant reductions in the survivorship, body condition, fecundity and lifetime reproductive success; (5) exploitation competition does account for these negative demographic effects on second‐generation females of P. communis more than interference competition.     

Towards a solution of the plankton paradox: the importance of physiology and life history

Phytoplankton communities reveal an astonishing biodiversity, whereas classical competition theory seems to suggest that only a few competing species can survive. Recently we suggested a new solution to this plankton paradox. In theory, at least, competition between multiple species can generate complex dynamics that can support a large number of species. How likely is it then, in reality, that competitive chaos indeed promotes biodiversity? To obtain some insight, we simulated multispecies competition according to five different physiological scenarios. For random species parameters, biodiversity was generally low. Assuming plausible physiological trade‐offs, the simulations revealed switches back and forth between equilibrium and nonequilibrium dynamics, and a higher biodiversity. An extremely high biodiversity, with sometimes more than 100 species on three resources, was observed in simulations that assumed a cyclic relation between competitive abilities and resource contents. We conclude that physiological and life‐history patterns have a major impact on the likelihood of nonequilibrium dynamics and on the biodiversity of plankton communities.     

Size-uneven competition and resource availability: A factorial experiment on seedling establishment of three Mediterranean species

Abstract

A factorial experiment was conducted in order to test whether the ranking of competitive ability at seedling stage is affected by different resource availability and competition asymmetry levels. Seedlings of three Mediterranean species (Rhamnus alaternus, Ampelodesmos mauritanicus and Cistus incanus) were grown at two competition intensities with different water and nutrient levels. Plants responses were expressed as mortality, biomass production and plant architecture. Data were also analysed by an index of competitive response intensity. A clear ranking of competitive response was observed, with Rhamnus > Ampelodesmos > Cistus. Competition was affected both by water and nutrient availability, but none of these factors modified competitive ranking. Seedling competition behaviour appeared to be related to species successional roles.     

FROM METABOLISM TO POLYMORPHISM IN BACTERIAL POPULATIONS: A THEORETICAL STUDY

Abstract Stable polymorphisms are commonly observed in experimental bacterial populations grown in homogeneous media. Evidence is accumulating that metabolic interactions might be the main mechanism underlying the emergence and maintenance of such polymorphisms. To date, however, attempts to model the evolution of bacterial polymorphism have not considered metabolism as a possible component of polymorphism maintenance. Here, we propose a simulation approach to model the evolution of selected polymorphisms in a bacterial population. Using recent knowledge of the relationship between bacterial fitness and metabolism, we build a simple metabolic model and test the effect of resource competition on polymorphism. Without making an a priori hypothesis on fitness functions, we show that stable polymorphic situations could be observed under high nutrient competition, and we propose a functional, metabolism‐based explanation to the debated issue of polymorphism maintenance.