Coexistence of competitive species with a stage-structured life cycle

Ecological theory provides explanations for exclusion or coexistence of competing species. Most theoretical works on competition dynamics that have shaped current perspectives on coexistence assume a simple life cycle. This simplification, however, may omit important realities. We present a simple two-stage structured competition model to investigate the effects of life-history characteristics on coexistence. The achievement and the stability of coexistence depend not only on competition coefficients but also on a set of life-history parameters that reflect the viability of an individual, namely, adult death rate, maturation rate, and birth rate. High individual viability is necessary for a species to persist, but it does not necessarily facilitate coexistence. Intense competition at the juvenile or adult stage may require higher or lower viability, respectively, for stable coexistence to be possible. The stability mechanism can be explained by the refuge effect of the less competitive stage, and the birth performance, which preserves the less competitive stage as a refuge. Coexistence might readily collapse if the life-history characteristics, which together constitute individual viability, change, even though two species have an inherent competitive relation conducive to stable coexistence.     

Long-Term Competitive Dynamics of Two Cryptic Rotifer Species: Diapause and Fluctuating Conditions

Life-history traits may have an important role in promoting species coexistence. However, the complexity of certain life cycles makes it difficult to draw conclusions about the conditions for coexistence or exclusion based on the study of short-term competitive dynamics. Brachionus plicatilis and B. manjavacasare two cryptic rotifer species co-occurring in many lakes on the Iberian Peninsula. They have a complex life cycle in which cyclical parthenogenesis occurs with diapausing stages being the result of sexual reproduction. B. plicatilis and B. manjavacasare identical in morphology and size, their biotic niches are broadly overlapping, and they have similar competitive abilities. However, the species differ in life-history traits involving sexual reproduction and diapause, and respond differently to salinity and temperature. As in the case of certain other species that are extremely similar in morphology, a fluctuating environment are considered to be important for their coexistence. We studied the long-term competitive dynamics of B. plicatilis and B. manjavacas under different salinity regimes (constant and fluctuating). Moreover, we focused on the dynamics of the diapausing egg bank to explore how the outcome of the entire life cycle of these rotifers can work to mediate stable coexistence. We demonstrated that these species do not coexist under constant-salinity environment, as the outcome of competition is affected by the level of salinity—at low salinity, B. plicatilis excluded B. manjavacas, and the opposite outcome occurred at high salinity. Competitive dynamics under fluctuating salinity showed that the dominance of one species over the other also tended to fluctuate. The duration of co-occurrence of these species was favoured by salinity fluctuation and perhaps by the existence of a diapausing egg bank. Stable coexistence was not found in our system, which suggests that other factors or other salinity fluctuation patterns might act as stabilizing processes in the wild.     

Life-cycle switching and coexistence of species with no niche differentiation

The increasing evidence of coexistence of cryptic species with no recognized niche differentiation has called attention to mechanisms reducing competition that are not based on niche-differentiation. Only sex-based mechanisms have been shown to create the negative feedback needed for stable coexistence of competitors with completely overlapping niches. Here we show that density-dependent sexual and diapause investment can mediate coexistence of facultative sexual species having identical niches. We modelled the dynamics of two competing cyclical parthenogens with species-specific density-dependent sexual and diapause investment and either equal or different competitive abilities. We show that investment in sexual reproduction creates an opportunity for other species to invade and become established. This may happen even if the invading species is an inferior competitor. Our results suggests a previously unnoticed mechanism for species coexistence and can be extended to other facultative sexual species and species investing in diapause where similar density-dependent life-history switches could act to promote coexistence.     

Evolution of life-history traits collapses competitive coexistence

Trade-offs between competitive ability and the other life-history traits are considered to be a major mechanism of competitive coexistence. Many theoretical studies have demonstrated the robustness of such a coexistence mechanism ecologically; however, it is unknown whether the coexistence is robust evolutionarily. Here, we report that evolution of life-history traits not directly related to competition, such as longevity, and predator avoidance, easily collapses competitive coexistence in several competition systems: spatially structured, and predator-mediated two-species competition systems. In addition, we found that a superior competitor can be excluded by an inferior one by common mechanisms among the models. Our results suggest that ecological competitive coexistence due to a life-history trait trade-off balance may not be balanced on an evolutionary timescale, that is, it may be evolutionarily fragile.     

A COEVOLUTIONARY ISOMORPHISM APPLIED TO LABORATORY STUDIES OF COMPETITION

Some empirical consequences of an isomorphism between the Lotka-Volterra competitive model and a coevolutionary competitive model are developed. In both the Lotka-Volterra and coevolutionary models, four competitive outcomes are possible: 1) species one wins, 2) species two wins, 3) indeterminate outcome, and 4) stable coexistence. These two models are isomorphic in the sense that the inequalities associated with a particular competitive outcome of the Lotka-Volterra model correspond in a one-to-one manner with similar inequalities associated with the same competitive outcome of the coevolutionary model. The inequalities of the Lotka-Volterra model involve the competition coefficients themselves, while the inequalities of the coevolutionary model involve the genetic variances and covariances of the competition coefficients. The isomorphism suggests some alternative interpretations of the results of classical laboratory studies of competition. The Lotka-Volterra (or ecological) hypotheses postulate that the competition coefficients are constant and that genetic considerations play no role in determining the competitive outcome. By contrast, the evolutionary hypotheses derived from the coevolutionary model postulate that the competition coefficients are variables and that the genetic variances and covariances of the competition coefficients determine the competitive outcome. The isomorphism is applied to competitive exclusion and coexistence, and to competitive indeterminacy in Tribolium. In particular, the evolutionary hypotheses isomorphic to the two classical explanations of competitive indeterminacy, the demographic stochasticity and genetic founder effect hypotheses, are constructed. The theory developed here and in a previous paper (Pease, 1984) provides one perspective on the relation among the Lotka-Volterra competition theory, quantitative genetics, competitive exclusion, the reversal of competitive dominance, coexistence, competitive indeterminacy in Tribolium, and experiments investigating the relation between genetic variability and the rate of evolution of fitness.     

Evolution of the maturation rate collapses competitive coexistence

Most theoretical studies on character displacement and the coexistence of competing species have focused attention on the evolution of competitive traits driven by inter-specific competition. We investigated the evolution of the maturation rate which is not directly related to competition and trades off with the birth rate and how it influences competitive outcomes. Evolution may result in the superior competitor becoming extinct if, initially, the inferior competitor has a lower, and the superior one a higher, maturation rate at the coexistence equilibrium. This counterintuitive result is explained by an explosive increase in the adult population of the inferior competitor as a result of the more rapid evolution of its maturation rate, which is caused by differences in the intensity and direction of selection on the maturation rates of the two species and in their adult densities, which are related to differences in their life histories. Thus, a life history trait trade-off with a competitive trait may cause a competitive ecological coexistence to collapse.     

Competitive exclusion through reproductive interference

A simple differential equation model was developed to describe the competitive interaction that may occur between species through reproductive interference. The model has the form comparable to Volterra's competition equations, and the graphical analysis of the outcome of the two-species interaction based on its zero-growth isoclines proved that: (1) The possible outcome in this model, as in usual models of resource competition, is either stable coexistence of both species or gradual exclusion of one species by the other, depending critically upon the values of the activity overlapping coefficient c_ij; (2) but, for the same c_ij-values, competitive exclusion is much more ready to occur here than in resource competition; (3) and moreover, the final result of the competition is always dependent on the initial-condition due to its non-linear isoclines, i.e., even under the parameter condition that generally allows both species to coexist, an extreme bias in intial density to one species can readily cause subsequent complete exclusion of its counterparts. Thus, it may follow that the reproductive interference is likely to be working in nature as an efficient mechanism to bring about habitat partitioning in either time or space between some closely related species in insect communities, even though they inhabit heterogeneous habitats where resource competition rarely occurs so that they could otherwise attain steady coexistence.     

Starve a competitor: evolution of luxury consumption as a competitive strategy

Organisms are often observed to acquire an excess of non-limiting resources, a process known as luxury consumption. Luxury consumption has been largely treated as a bet hedging strategy for temporal variation in resource supply, but may also function as a competitive strategy. We incorporate luxury resource consumption into a derivation of the classic resource ratio model for competition between terrestrial plant, and explore its consequences for population dynamics and competition. We show that luxury consumption reduces the potential for coexistence between two species competing for two resources. Furthermore, we demonstrate that luxury consumption can be selected for because of the competitive advantage that luxury consumers gain. Luxury consumption evolves when competition for resources is local rather than global, there is potential for coexistence between the two species and the competitive environment remains stable over a sufficient period of time to allow selection to act. The evolutionary outcome can be either extinction of one of the competing species or coexistence of the two species with maximum luxury consumption. The potential for selection to favor luxury consumption is well predicted by the competitive outcome between individuals of the two species with and without luxury consumption.     

Using exclusion rate to unify niche and neutral perspectives on coexistence

The competitive exclusion principle is one of the most influential concepts in ecology. The classical formulation suggests a correlation between competitor species similarity and competition severity, leading to rapid competitive exclusion where species are very similar; yet neutral models show that identical species can persist in competition for long periods. Here, we resolve the conflict by examining two components of similarity – niche overlap and competitive similarity – and modeling the effects of each on exclusion rate (defined as the inverse of time to exclusion). Studying exclusion rate, rather than the traditional focus on binary outcomes (coexistence versus exclusion), allows us to examine classical niche and neutral perspectives using the same currency. High niche overlap speeds exclusion, but high similarity in competitive ability slows it. These predictions are confirmed by a well‐known model of two species competing for two resources. Under ecologically plausible scenarios of correlation between these two factors, the strongest exclusion rates may be among moderately similar species, while very similar and highly dissimilar competitors have very low exclusion rates. Adding even small amounts of demographic stochasticity to the model blurs the line between deterministic and probabilistic coexistence still further. Thus, focusing on exclusion rate, instead of on the binary outcome of coexistence versus exclusion, allows a variety of outcomes to result from competitive interactions. This approach may help explain species coexistence in diverse competitive communities and raises novel issues for future work.     

Interactions between frequency and size of disturbance affect competitive outcomes

Disturbance has many effects on ecological communities, and it is often suggested that disturbance can affect species diversity by altering competitive outcomes. However, disturbance regimes have many distinct aspects that may act, and interact, to influence species diversity. While there are many theoretical models of disturbance-prone communities, few have specifically documented how interactions between different aspects of a disturbance regime change competitive outcomes. Here, we present a model of two plant species subject to disturbance which we then use to examine species coexistence over varying levels of two aspects of disturbance: frequency, and spatial extent (i.e., area disturbed). We show that the competitive outcome is affected differently by changes in each aspect and that the effect of disturbance frequency on species coexistence depends strongly on the spatial extent of the disturbance, and vice versa. We classify the nature of these interactions between disturbance frequency and extent on the basis of the shape of the resulting coexistence regions in a frequency?Cextent parameter plane. Our results illustrate that different types of interaction can result from differences in life-history traits that control species-specific sensitivity to frequency and extent of disturbance. Thus, our analysis shows that the various aspects of disturbance must be carefully considered in concert with the life-history traits of the community members in order to assess the consequences of disturbance.     

Density dependence across multiple life stages in a temperate old-growth forest of northeast China

Recent studies on species coexistence suggest that density dependence is an important mechanism regulating plant populations. However, there have been few studies of density dependence conducted for more than one life-history stage or that control for habitat heterogeneity, which may influence spatial patterns of survival and mask density dependence. We explored the prevalence of density dependence across multiple life stages, and the effects of controlling for habitat heterogeneity, in a temperate forest in northeast China. We used generalized linear mixed-effects models to test for density-dependent mortality of seedlings and spatial point pattern analysis to detect density dependence for sapling-to-juvenile transitions. Conspecific neighbors had a negative effect on survival of plants in both life stages. At the seedling stage, we found a negative effect of conspecific seedling neighbors on survival when analyzing all species combined. However, in species-level analyses, only 2 of 11 focal species were negatively impacted by conspecific neighbors, indicating wide variation among species in the strength of density dependence. Controlling for habitat heterogeneity did not alter our findings of density dependence at the seedling stage. For the sapling-to-juvenile transition stage, 11 of 15 focal species showed patterns of local scale (<10 m) conspecific thinning, consistent with negative density dependence. The results varied depending on whether we controlled for habitat heterogeneity, indicating that a failure to account for habitat heterogeneity can obscure patterns of density dependence. We conclude that density dependence may promote tree species coexistence by acting across multiple life-history stages in this temperate forest.     

Spatial patterns and associations of four species in an old-growth temperate forest

The spatial pattern of a tree species is an important characteristic of plant communities, providing critical information to explain species coexistence. The spatial distribution and association of four different successional species were analyzed among different life-history stages in an old-temperate forest. Significant aggregation patterns were found, and the degree of aggregation decreased with the scales and life-history stages. Significant interspecific spatial associations were detected. In comparing the relationships among the different life-history stages, positive associations were found at small scales in all of the juvenile species pairs. In the adult stage, negative associations were detected in coniferous vs. deciduous species pairs, while the deciduous species pairs, which have identical resource requirements, showed a positive association in this study. The coniferous species pairs showed a positive association at small scales. We infer that seed dispersal, competitive ability, or the requirement for specific topographic and light environments may contribute to the coexistence of these species.     

Life-history trade-offs and ecological dynamics in the evolution of longevity

Longevity is a life-history trait that is shaped by natural selection. An unexplored consequence is how selection on this trait affects diversity and diversification in species assemblages. Motivated by the diverse rockfish (Sebastes) assemblage in the North Pacific, the effects of trade-offs in longevity against competitive ability are explored. A competition model is developed and used to explore the potential for species diversification and coexistence. Invasion analyses highlight that life-history trait trade-offs in longevity can mitigate the effects of competitive ability and favour the coexistence of a finite number of species. Our results have implications for niche differentiation, limiting similarity and assembly dynamics in multispecies interactions.     

Temporal resource partitioning mitigates interspecific competition and promotes coexistence among insect parasites

A key to understanding life's great diversity is discerning how competing organisms divide limiting resources to coexist in diverse communities. While temporal resource partitioning has long been hypothesized to reduce the negative effects of interspecific competition, empirical evidence suggests that time may not often be an axis along which animal species routinely subdivide resources. Here, we present evidence to the contrary in the world's most biodiverse group of animals: insect parasites (parasitoids). Specifically, we conducted a meta-analysis of 64 studies from 41 publications to determine if temporal resource partitioning via variation in the timing of a key life-history trait, egg deposition (oviposition), mitigates interspecific competition between species pairs sharing the same insect host. When competing species were manipulated to oviposit at (or near) the same time in or on a single host in the laboratory, competition was common, and one species was typically inherently superior (i.e. survived to adulthood a greater proportion of the time). In most cases, however, the inferior competitor could gain a survivorship advantage by ovipositing earlier (or in a smaller number of cases later) into shared hosts. Moreover, this positive (or in a few cases negative) priority advantage gained by the inferior competitor increased as the interval between oviposition times became greater. The results from manipulative experiments were also correlated with patterns of life-history timing and demography in nature: the more inherently competitively inferior a species was in the laboratory, the greater the interval between oviposition times of taxa in co-occurring populations. Additionally, the larger the interval between oviposition times of competing taxa, the more abundant the inferior species was in populations where competitors were known to coexist. Overall, our findings suggest that temporal resource partitioning via variation in oviposition timing may help to facilitate species coexistence and structures diverse insect communities by altering demographic measures of species success. We argue that the lack of evidence for a more prominent role of temporal resource partitioning in promoting species coexistence may reflect taxonomic differences, with a bias towards larger-sized animals. For smaller species like parasitic insects that are specialized to attack one or a group of closely related hosts, have short adult lifespans and discrete generation times, compete directly for limited resources in small, closed arenas and have life histories constrained by host phenology, temporal resource subdivision via variation in life history may play a critical role in allowing species to coexist by alleviating the negative effects of interspecific competition.     

Coexistence of competing juvenile–adult structured populations

The Leslie-Gower model is a discrete time analog of the competition Lotka–Volterra model and is known to possess the same dynamic scenarios of that famous model. The Leslie–Gower model played a historically significant role in the history of competition theory in its application to classic laboratory experiments of two competing species of flour beetles (carried out by Park in the 1940s–1960s). While these experiments generally supported what became the Competitive Exclusion Principle, Park observed an anomalous coexistence case. Recent literature has discussed Park’s ‘coexistence case’ by means of non-Lotka–Volterra, non-equilibrium dynamics that occur in a high dimensional model with life cycle stages. We study this dynamic possibility in the lowest possible dimension, that is to say, by means of a model involving only two species each with two life cycle stages. We do this by extending the Leslie–Gower model so as to describe the competitive interaction of two species with juvenile and adult classes. We give a complete account of the global dynamics of the resulting model and show that it allows for non-equilibrium competitive coexistence as competition coefficients are increased. We also show that this phenomenon occurs in a general class of models for competing populations structured by juvenile and adult life cycle stages.     

The maximum power principle predicts the outcomes of two-species competition experiments

The maximum power principle (MPP) states that biological systems organize to increase power whenever the system constraints allow. The MPP has the potential to explain a variety of ecological patterns because biological power (metabolism) is a component of all ecological interactions. I empirically tested the MPP by reanalyzing three two-species competition experiments by Gause, Vandermeer, and Fox and Morin. These experiments investigated competitive outcomes in microcosms of heterotrophic protists. I introduce metabolic state-space graphs to portray the metabolic trajectories of the communities and show that the steady-state outcomes of these experiments are consistent with the MPP. Winning species were successfully predicted a priori from their status as the species with the highest power when alone. In addition, periods of coexistence, although not predictable a priori, were consistent with the MPP because coexistence states had community-level power that was higher than either species could achieve alone. Thus, the outcomes of all ten trials were the maximum power states, given the options. The results suggest that the maximum power principle may represent a useful energetic organizing principle for communities.     

Size-dependent mortality induces life-history changes mediated through population dynamical feedbacks

The majority of taxa grow significantly during life history, which often leads to individuals of the same species having different ecological roles, depending on their size or life stage. One aspect of life history that changes during ontogeny is mortality. When individual growth and development are resource dependent, changes in mortality can affect the outcome of size-dependent intraspecific resource competition, in turn affecting both life history and population dynamics. We study the outcome of varying size-dependent mortality on two life-history types, one that feeds on the same resource throughout life history and another that can alternatively cannibalize smaller conspecifics. Compensatory responses in the life history dampen the effect of certain types of size-dependent mortality, while other types of mortality lead to dramatic changes in life history and population dynamics, including population (de-)stabilization, and the growth of cannibalistic giants. These responses differ strongly among the two life-history types. Our analysis provides a mechanistic understanding of the population-level effects that come about through the interaction between individual growth and size-dependent mortality, mediated by resource dependence in individual vital rates.     

基于竞争和扩散能力的非捕食-被捕食集合种群系统的竞争机制

对于非捕食 被捕食(食饵)生态系统,强弱物种之间存在一定的竞争影响.在不考虑栖息地毁坏的情况下,引进双向竞争机制,将Tilman的单向竞争模式推广为n集合种群双向竞争模型,并对6-集合种群的竞争动态进行了计算机模拟研究.结果表明,在平衡态,种群竞争共存的条件是其竞争能力与扩散能力呈现指数型负相关关系,竞争的结果使物种的强弱序列发生变化;物种竞争排除与共存受迁移扩散能力和竞争能力影响很大,在局域斑块上竞争排斥的集合种群在广域尺度上可以竞争共存,即逃亡共存.     

Theoretical dynamics of competitors under predation

Summary Continuous population models of two prey species and a predator were explored by isocline analysis. When predator satiation and substitution between prey (with or without switching) are introduced in the models, many qualitatively different kinds of dynamic behaviour become possible. These depend in a complex but predictable way on competitive relations between prey and on predator feeding behaviour and efficiency. Under constant predation many cases of threshold responses between two or more alternate stable states are possibly; the numerical response of the predator population reduces some of the possibilities.Apparently contradictory community phenomena previously proposed, e.g. prey coexistence versus exclusion by addition of predator, exclusion versus stabilization by addition of alternate prey, are all possible as special cases. A prey which is relatively tolerant to predation can act as a keystone species, on which the existence of other prey species in the community depends, in either a positive or a negative sense. In certain conditions predator-induced obligatory mutualism between two prey species is theoretically possible.To Michael Evenari, pioneer, teacher and friend     

Coexistence of competing juvenile-adult structured populations

The Leslie-Gower model is a discrete time analog of the competition Lotka-Volterra model and is known to possess the same dynamic scenarios of that famous model. The Leslie-Gower model played a historically significant role in the history of competition theory in its application to classic laboratory experiments of two competing species of flour beetles (carried out by Park in the 1940s-1960s). While these experiments generally supported what became the Competitive Exclusion Principle, Park observed an anomalous coexistence case. Recent literature has discussed Park's 'coexistence case' by means of non-Lotka-Volterra, non-equilibrium dynamics that occur in a high dimensional model with life cycle stages. We study this dynamic possibility in the lowest possible dimension, that is to say, by means of a model involving only two species each with two life cycle stages. We do this by extending the Leslie-Gower model so as to describe the competitive interaction of two species with juvenile and adult classes. We give a complete account of the global dynamics of the resulting model and show that it allows for non-equilibrium competitive coexistence as competition coefficients are increased. We also show that this phenomenon occurs in a general class of models for competing populations structured by juvenile and adult life cycle stages.