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.     

Mechanisms of coexistence in competitive metacommunities

Although there is a large body of theory on spatial competitive coexistence, very little of it involves comparative analyses of alternative mechanisms. We thus have limited knowledge of the conditions under which multiple spatial mechanisms can operate or of emergent properties arising from interactions between mechanisms. Here we present a mathematical framework that allows for comparative analysis of spatial coexistence mechanisms. The basis for comparison is mechanisms operating in spatially homogeneous competitive environments (e.g., life-history trade-offs) versus mechanisms operating in spatially heterogeneous competitive environments (e.g., source-sink dynamics). Our comparative approach leads to several new insights about spatial coexistence. First, we show that spatial variation in the expression of a life-history trade-off leads to a unique regional pattern that cannot be predicted by considering trade-offs or source-sink dynamics alone. This result represents an instance where spatial heterogeneity constrains rather than promotes coexistence, and it illustrates the kind of counterintuitive emergent properties that arise due to interactions between different classes of mechanisms. Second, we clarify the role of dispersal mortality in spatial coexistence. Previous studies have shown that coexistence can be constrained or facilitated by dispersal mortality. Our broader analysis distinguishes situations where dispersal mortality is not necessary for coexistence from those where such mortality is essential for coexistence because it preserves spatial variation in the strength of competition. These results form the basis for two important future directions: evolution of life-history traits in spatially heterogeneous environments and elucidation of the cause and effect relationship(s) between biodiversity and ecosystem functioning.     

The age-structured lottery model

The lottery model of competition between species in a variable environmental has been influential in understanding how coexistence may result from interactions between fluctuating environmental and competitive factors. Of most importance, it has led to the concept of the storage effect as a mechanism of species coexistence. Interactions between environment and competition in the lottery model stem from the life-history assumption that environmental variation and competition affect recruitment to the adult population, but not adult survival. The strong role of life-history attributes in this coexistence mechanism implies that its robustness should be checked for a variety of life-history scenarios. Here, age structure is added to the adult population, and the results are compared with the original lottery model. This investigation uses recently developed shape characteristics for mortality and fecundity schedules to quantify the effects of age structure on the long-term low-density growth rate of a species in competition with its competitor when applying the standard invasibility coexistence criterion. Coexistence conditions are found to be affected to a small degree by the presence of age structure in the adult population: Type III mortality broadens coexistence conditions, and type I mortality makes them narrower. The rates of recovery from low density for coexisting species, and the rates of competitive exclusion in other cases, are modified to a greater degree by age structure. The absolute rates of recovery or decline of a species from low density are increased by type I mortality or early peak reproduction, but reduced by type III mortality or late peak reproduction. Analytical approximations show how the most important effects can be considered as simple modifications of the long-term low-density growth rates for the original lottery model.     

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.     

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.     

Competition and coexistence with multiple life-history stages

Do complex life histories affect the conditions under which competitors can coexist? We investigated this using a two-species, two-stage Ricker model. With complex life cycles, the competition coefficients associated with each life-history stage suggest one of three competitive outcomes-coexistence, alternate stable states, or competitive exclusion-that depend on the relative magnitudes of intraspecific and interspecific competition. When the two stages suggest the same outcome, only that outcome can occur. When the stages suggest different outcomes, either one may prevail. It is also possible to have emergent outcomes, in which the outcome is not suggested by either stage. This can occur when the two stages suggest competitive exclusion by opposite species or when one stage suggests alternate stable states and the other suggests coexistence. Therefore, determining the mechanisms of coexistence in species with complex life histories may require consideration of competitive interactions within all life-history stages.     

Host availability and the evolution of parasite life-history strategies

Parasites exploit an inherently patchy resource, their hosts, which are discrete entities that may only be available for infection within a relatively short time window. However, there has been little consideration of how heterogeneities in host availability may affect the phenotypic or genotypic composition of parasite populations or how parasites may evolve to cope with them. Here we conduct a selection experiment involving an entomopathogenic nematode (Steinernema feltiae) and show for the first time that the infection rate of a parasite can evolve rapidly to maximize the chances of infecting within an environment characterized by the rate of host availability. Furthermore, we show that the parasite's infection rate trades off with other fitness traits, such as fecundity and survival. Crucially, the outcome of competition between strains with different infection strategies depends on the rate of host availability; frequently available hosts favor "fast" infecting nematodes, whereas infrequently available hosts favor "slow" infecting nematodes. A simple evolutionarily stable strategy (ESS) analysis based on classic epidemiological models fails to capture this behavior, predicting instead that the fastest infecting phenotype should always dominate. However, a novel model incorporating more realistic, discrete bouts of host availability shows that strain coexistence is highly likely. Our results demonstrate that heterogeneities in host availability play a key role in the evolution of parasite life-history traits and in the maintenance of phenotypic variability. Parasite life-history strategies are likely to evolve rapidly in response to changes in host availability induced by disease management programs or by natural dynamics in host abundance. Incorporating parasite evolution in response to host availability would therefore enhance the predictive ability of current epidemiological models of infectious disease.     

Translucent windows: how uncertainty in competitive interactions impacts detection of community pattern

Traits can provide a window into the mechanisms that maintain coexistence among competing species. Recent theory suggests that competitive interactions will lead to groups, or clusters, of species with similar traits. However, theoretical predictions typically assume complete knowledge of the map between competition and measured traits. These assumptions limit the plausible application of these patterns for inferring competitive interactions in nature. Here, we relax these restrictions and find that the clustering pattern is robust to contributions of unknown or unobserved niche axes. However, it may not be visible unless measured traits are close proxies for niche strategies. We conclude that patterns along single niche axes may reveal properties of interspecific competition in nature, but detecting these patterns requires natural history expertise firmly tying traits to niches.     

Competition between invasive and indigenous species: an insular case study of subterranean termites

An important requirement for the management of invasive species is to identify the biological and ecological factors that influence the ability of such species to become established and spread within a new environment. Although competition is one of the key interactions determining the coexistence of species and exclusion, few studies directly examine the mechanism of competitive interactions within invasive communities. This study focused on putative competition in a social insect invader, R. flavipes, an American termite introduced into France, and an indigenous European termite, R. grassei. We first characterized and mapped a zone of sympatry between these two species. We then evaluated the degree of direct and indirect competition by comparing several life-history traits: behavioral aggression, chemical recognition and dispersion modes. Interspecific competition revealed that R. flavipes was dominant over R. grassei. Intraspecific competition was not found in R. flavipes while it appeared in varying degrees in R. grassei. These findings seemed to be correlated with the remarkable chemical homogeneity found in R. flavipes in comparison with R. grassei. Genetic analyses revealed that R. flavipes foraged over a greater distance than R. grassei colonies and might suggest a difference in the capacity to produce secondary reproductives. These findings suggest that R. flavipes has a significant advantage owing to competitive asymmetry that may enable the species to become dominant. The interspecific superiority, lack of intraspecific aggression and large extensive colonies, seem to be some of the reasons for its invasive success.     

Coexistence in metacommunities: a tree-species model

Simple patch-occupancy models of competitive metacommunities have shown that coexistence is possible as long as there is a competition-colonization tradeoff such as that of superior competitors and dispersers. In this paper, we present a model of competition between three species in a dynamic landscape, where patches are being created and destroyed at a different rate. In our model, species interact according to a linear non-transitive hierarchy, such that species Y(3) outcompetes and can invade patches occupied by species Y(2) and this species in turn can outcompete and invade patches occupied by the inferior competitor Y(1). In this hierarchy, inferior competitors cannot invade patches of species with higher competitive ability. Analytical results show that there are regions in the parameter space where coexistence can occur, as well as regions where each of the species exists in isolation depending on species' life-history traits associated with their colonization abilities and extinction proneness as well as with the dynamics of habitat patches. In our model, the condition for coexistence depends explicitly on patch dynamics, which in turn modulate the limiting similarity for species coexistence. Coexistence in metacommunities inhabiting dynamic landscapes although possible is harder to attain than in static ones.     

Eco-Evolutionary dynamics enable coexistence via neighbor-dependent selection

Recent studies suggest that selection can allow coexistence in situations where ecological dynamics lead to competitive exclusion, provided that there is a trade-off between traits optimal for interacting with conspecifics and traits optimal for interacting with heterospecifics. Despite compelling empirical evidence, there is no general framework for elucidating how and when selection will allow coexistence in natural communities. Here we develop such a framework for a mechanism that we term "neighbor-dependent selection." We show that this mechanism can both augment coexistence when ecological conditions allow for niche partitioning and enable coexistence when ecological conditions lead to competitive exclusion. The novel insight is that when ecological conditions lead to exclusion, neighbor-dependent selection can allow coexistence via cycles driven by an intransitive loop; selection causes one species to be a superior interspecific competitor when it is rare and an inferior interspecific competitor when it is abundant. Our framework predicts the conditions under which selection can enable coexistence, as opposed to merely augmenting it, and elucidates the effects of heritability on the eco-evolutionary feedbacks that drive coexistence. Given increasing evidence that evolution operates on ecological timescales, our approach provides one means for evaluating the role of selection and trait evolution in species coexistence.     

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.     

Evolution restricts the coexistence of specialists and generalists: the role of trade-off structure

Environmental variability and adaptive foraging behavior have been shown to favor coexistence of specialists and generalists on an ecological timescale. This leaves unaddressed the question of whether such coexistence can also be expected on an evolutionary timescale. In this article, we study the attainability, through gradual evolution, of specialist-generalist coexistence, as well as the evolutionary stability of such communities when allowing for immigration. Our analysis shows that the potential for specialist-generalist coexistence is much more restricted than originally thought and strongly depends on the trade-off structure assumed. We establish that ecological coexistence is less likely for species facing a trade-off between per capita reproduction in different habitats than when the trade-off acts on carrying capacities alone. We also demonstrate that coexistence is evolutionarily stable whenever it is ecologically stable but that in most cases, such coexistence cannot be reached through gradual evolution. We conclude that an evolutionarily stable community of specialists and generalists may be created only through immigration from elsewhere or through mutations of large effect. Our results highlight that trade-offs in fitness-determining traits can have counterintuitive effects on the evolution of specialization.     

Competitive intransitivity promotes species coexistence

Using a spatially explicit cellular automaton model with local competition, we investigate the potential for varied levels of competitive intransitivity (i.e., nonhierarchical competition) to promote species coexistence. As predicted, on average, increased levels of intransitivity result in more sustained coexistence within simulated communities, although the outcome of competition also becomes increasingly unpredictable. Interestingly, even a moderate degree of intransitivity within a community can promote coexistence, in terms of both the length of time until the first competitive exclusion and the number of species remaining in the community after 500 simulated generations. These results suggest that modest levels of intransitivity in nature, such as those that are thought to be characteristic of plant communities, can contribute to coexistence and, therefore, community-scale biodiversity. We explore a potential connection between competitive intransitivity and neutral theory, whereby competitive intransitivity may represent an important mechanism for "ecological equivalence."     

Preemption of space can lead to intransitive coexistence of competitors

Intransitive competition has the potential to be a powerful contributor to species coexistence, but there are few proposed biological mechanisms that could create intransitivities in natural communities. Using a three‐species model of competition for space, we demonstrate a mechanism for coexistence that combines a colonization–competition tradeoff between two species with the ability of a third species to preempt space from the other competitors. The combination of differential abilities to colonize, preempt, and overtake space creates a community where no single species can exclude both of its competitors. The dynamics of this kind of community are analogous to rock‐paper‐scissors competition, and the three‐species community can persist even though not all pairs of species can coexist in isolation. In distinction to prior results, this is a mechanism of intransitivity that does not require nonhierarchical local interference competition. We present parameter estimates from a subtidal marine community illustrating how documented competitive traits can lead to preemption‐based intransitivities in natural communities, and we describe methods for an empirical test of the occurrence of this mechanism.     

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.     

Evolutionary stability of coexistence due to the storage effect in a two-season model

The question of species coexistence has been central to ecology since its founding. Ever-present environmental variation may be one answer to that question. Previous models have demonstrated that species can exploit this variation to coexist with competitors by having different environmental responses (the storage effect). When traits governing species’ environmental response can evolve, however, coexistence is not assured. In this study, we use a continuous time, two-season model to determine the evolutionary outcome of competing species evolving in their seasonal performance trait. We extend the competitive exclusion principle to show that the storage effect can allow no more than N species to coexist on N discrete seasons with no relative nonlinearity. We find a broad region of parameter space where coexistence is evolutionarily stable. The size of this region depends on the period of fluctuations relative to the individual lifespan. Relatively long period fluctuations yield a large coexistence region, but as the period decreases, the region narrows and disappears asymptotically. Finally, we cast our adaptive dynamics technique in terms of Chesson’s concept of equalizing and stabilizing mechanisms to demonstrate that the breakdown in coexistence at short periods is due to loss of the stabilizing covariance between the environment and competition.     

Coexistence of predator and prey in intraguild predation systems with ontogenetic niche shifts

In basic intraguild predation (IGP) systems, predators and prey also compete for a shared resource. Theory predicts that persistence of these systems is possible when intraguild prey is superior in competition and productivity is not too high. IGP often results from ontogenetic niche shifts, in which the diet of intraguild predators changes as a result of growth in body size (life-history omnivory). As a juvenile, a life-history omnivore competes with the species that becomes its prey later in life. Competition can hence limit growth of young predators, while adult predators can suppress consumers and therewith neutralize negative effects of competition. We formulate and analyze a stage-structured model that captures both basic IGP and life-history omnivory. The model predicts increasing coexistence of predators and consumers when resource use of stage-structured predators becomes more stage specific. This coexistence depends on adult predators requiring consumer biomass for reproduction and is less likely when consumers outcompete juvenile predators, in contrast to basic IGP. Therefore, coexistence occurs when predation structures the community and competition is negligible. Consequently, equilibrium patterns over productivity resemble those of three-species food chains. Life-history omnivory thus provides a mechanism that allows intraguild predators and prey to coexist over a wide range of resource productivity.     

Inbreeding depression in the competitive fertilization success of male crickets

Mating between close relatives generally results in offspring of decreased fitness. Inbreeding depression is generally greater for life-history traits than for morphological traits, and recent studies of traits subject to sexual selection suggest that these may suffer the greatest inbreeding depression. Sexual selection continues after mating in the form of sperm competition and cryptic female choice, imposing strong selection on male competitive fertilization success. Here, I examine the effects of a single generation of full-sib mating on competitive fertilization success in a cricket, Teleogryllus oceanicus. The estimated coefficient of inbreeding depression in competitive fertilization success was 0.37, higher than that for other life-history and morphological traits. Such intense inbreeding depression coupled with little or no additive genetic variance for this trait is consistent with strong directional selection on male competitive fertilization success generating high levels of dominance variance, and provides an adaptive explanation for the evolution of inbreeding avoidance found in this species.     

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.