Latitudinal variation in the competition‐colonisation trade‐off reveals rate‐mediated mechanisms of coexistence

Theories of species coexistence often describe a trade‐off between colonising and competitive abilities. In sessile marine invertebrates, this trade‐off can manifest as trends in species distributions relative to the size of isolated patches of substrate. Based on their abilities to find available substrate and competitively exclude neighbours, good colonisers tend to dominate smaller patches, whereas better competitors tend to monopolise larger patches. In theory, species with equivalent colonising and competitive abilities should display similar distributions across patch sizes. We used patch size to observe this manifestation of the competition‐colonisation trade‐off over 20° of latitude. The trade‐off was more readily observed at lower latitudes and was proportional to the ‘ecological age’ of communities (i.e. the degree of resource acquisition and likelihood of species interactions). Results suggest that ecological age may mediate the prominence of stochastic or deterministic coexistence mechanisms and will depend on the rate of ecological processes.     

The competition–dispersal trade‐off exists in forbs but not in graminoids: A case study from multispecies alpine grassland communities

Much theoretical evidence has demonstrated that a trade‐off between competitive and dispersal ability plays an important role in facilitating species coexistence. However, experimental evidence from natural communities is still rare. Here, we tested the competition–dispersal trade‐off hypothesis in an alpine grassland in the Tianshan Mountains, Xinjiang, China, by quantifying competitive and dispersal ability using a combination of 4 plant traits (seed mass, ramet mass, height, and dispersal mode). Our results show that the competition–dispersal trade‐off exists in the alpine grassland community and that this pattern was primarily demonstrated by forbs. The results suggest that most forb species are constrained to be either good competitors or good dispersers but not both, while there was no significant trade‐off between competitive and dispersal ability for most graminoids. This might occur because graminoids undergo clonal reproduction, which allows them to find more benign microenvironments, forage for nutrients across a large area and store resources in clonal structures, and they are thus not strictly limited by the particular resources at our study site. To the best of our knowledge, this is the first time the CD trade‐off has been tested for plants across the whole life cycle in a natural multispecies plant community, and more comprehensive studies are still needed to explore the underlying mechanisms and the linkage between the CD trade‐off and community composition.     

EVOLUTION OF THE STORAGE EFFECT

The storage effect, a mechanism that promotes species coexistence in temporally variable environments, poses a dilemma to evolutionary ecologists. Ecological studies have demonstrated its importance in natural communities, but evolutionary models have predicted that selection either impedes coexistence or diminishes the storage effect if there is coexistence. Here, we develop a lottery model of competition in which two species experience a trade‐off in competitive ability between two types of years. We use an adaptive evolution framework to determine conditions favoring the evolution of the storage effect. Storage evolves via divergence of relative performance in the two environments under a wide range of biologically realistic conditions. It evolves between two initially identical species (or lineages) when the trade‐off in performance is strong enough. It evolves for species having different initial trade‐offs for both weak and strong trade‐offs. Our simple 2‐species‐2‐environment scenario can be extended to multiple species and environmental conditions. Results indicate that the storage effect should evolve in a broad range of situations that involve a trade‐off in competitive ability among years, and are consistent with empirical observations. The findings show that storage can evolve in a manner and under conditions similar to other types of resource partitioning.     

Local range boundaries vs. large‐scale trade‐offs: climatic and competitive constraints on tree growth

Species often respond to human‐caused climate change by shifting where they occur on the landscape. To anticipate these shifts, we need to understand the forces that determine where species currently occur. We tested whether a long‐hypothesised trade‐off between climate and competitive constraints explains where tree species grow on mountain slopes. Using tree rings, we reconstructed growth sensitivity to climate and competition in range centre and range margin tree populations in three climatically distinct regions. We found that climate often constrains growth at environmentally harsh elevational range boundaries, and that climatic and competitive constraints trade‐off at large spatial scales. However, there was less evidence that competition consistently constrained growth at benign elevational range boundaries; thus, local‐scale climate‐competition trade‐offs were infrequent. Our work underscores the difficulty of predicting local‐scale range dynamics, but suggests that the constraints on tree performance at a large‐scale (e.g. latitudinal) may be predicted from ecological theory.     

Trait‐based tests of coexistence mechanisms

Recent functional trait studies have shown that trait differences may favour certain species (environmental filtering) while simultaneously preventing competitive exclusion (niche partitioning). However, phenomenological trait‐dispersion analyses do not identify the mechanisms that generate niche partitioning, preventing trait‐based prediction of future changes in biodiversity. We argue that such predictions require linking functional traits with recognised coexistence mechanisms involving spatial or temporal environmental heterogeneity, resource partitioning and natural enemies. We first demonstrate the limitations of phenomenological approaches using simulations, and then (1) propose trait‐based tests of coexistence, (2) generate hypotheses about which plant functional traits are likely to interact with particular mechanisms and (3) review the literature for evidence for these hypotheses. Theory and data suggest that all four classes of coexistence mechanisms could act on functional trait variation, but some mechanisms will be stronger and more widespread than others. The highest priority for future research is studies of interactions between environmental heterogeneity and trait variation that measure environmental variables at within‐community scales and quantify species' responses to the environment in the absence of competition. Evidence that similar trait‐based coexistence mechanisms operate in many ecosystems would simplify biodiversity forecasting and represent a rare victory for generality over contingency in community ecology.     

Heteromyopia and the spatial coexistence of similar competitors

Most spatial models of competing species assume symmetries in the spatial scales of dispersal and interactions. This makes analysis tractable, and has led to the conclusion that segregation of species in space does not promote coexistence. However, these symmetries leave parts of the parameter space uninvestigated. Using a moment‐approximation method, we present a spatial version of the Lotka–Volterra competition equations to investigate effects of removing symmetries in the distances over which individuals disperse and interact. Some spatial segregation of the species always comes about due to competition, and such segregation does not necessarily lead to coexistence. But, if interspecific competition occurs over shorter distances than intraspecific competition (heteromyopia), spatial segregation becomes strong enough to promote coexistence. Such coexistence is most likely when the species have similar dynamics, in contrast to the competition–colonization trade‐off that requires large competitive differences between species.     

Trait adaptation promotes species coexistence in diverse predator and prey communities

Species can adjust their traits in response to selection which may strongly influence species coexistence. Nevertheless, current theory mainly assumes distinct and time‐invariant trait values. We examined the combined effects of the range and the speed of trait adaptation on species coexistence using an innovative multispecies predator–prey model. It allows for temporal trait changes of all predator and prey species and thus simultaneous coadaptation within and among trophic levels. We show that very small or slow trait adaptation did not facilitate coexistence because the stabilizing niche differences were not sufficient to offset the fitness differences. In contrast, sufficiently large and fast trait adaptation jointly promoted stable or neutrally stable species coexistence. Continuous trait adjustments in response to selection enabled a temporally variable convergence and divergence of species traits; that is, species became temporally more similar (neutral theory) or dissimilar (niche theory) depending on the selection pressure, resulting over time in a balance between niche differences stabilizing coexistence and fitness differences promoting competitive exclusion. Furthermore, coadaptation allowed prey and predator species to cluster into different functional groups. This equalized the fitness of similar species while maintaining sufficient niche differences among functionally different species delaying or preventing competitive exclusion. In contrast to previous studies, the emergent feedback between biomass and trait dynamics enabled supersaturated coexistence for a broad range of potential trait adaptation and parameters. We conclude that accounting for trait adaptation may explain stable and supersaturated species coexistence for a broad range of environmental conditions in natural systems when the absence of such adaptive changes would preclude it. Small trait changes, coincident with those that may occur within many natural populations, greatly enlarged the number of coexisting species.     

Survival rates indicate that correlations between community‐weighted mean traits and environments can be unreliable estimates of the adaptive value of traits

Correlations between community‐weighted mean (CWM) traits and environmental gradients are often assumed to quantify the adaptive value of traits. We tested this assumption by comparing these correlations with models of survival probability using 46 perennial species from long‐term permanent plots in pine forests of Arizona. Survival was modelled as a function of trait × environment interactions, plant size, climatic variation and neighbourhood competition. The effect of traits on survival depended on the environmental conditions, but the two statistical approaches were inconsistent. For example, CWM‐specific leaf area (SLA) and soil fertility were uncorrelated. However, survival was highest for species with low SLA in infertile soil, a result which agreed with expectations derived from the physiological trade‐off underpinning leaf economic theory. CWM trait–environment relationships were unreliable estimates of how traits affected survival, and should only be used in predictive models when there is empirical support for an evolutionary trade‐off that affects vital rates.     

Functional trait trade‐off and species abundance: insights from a multi‐decadal study

Phylogenetically informed trait comparisons across entire communities show promise in advancing community ecology. We use this approach to better understand the composition of a community of winter annual plants with multiple decades of monitoring and detailed morphological, phenological and physiological measurements. Previous research on this system revealed a physiological trade‐off among dominant species that accurately predicts population and community dynamics. Here we expanded our investigation to 51 species, representing 96% of individual plants recorded over 30 years, and analysed trait relationships in the context of species abundance and phylogenetic relationships. We found that the functional‐trait trade‐off scales to the entire community, albeit with diminished strength. It is strongest for dominant species and weakens as progressively rarer species are included. The trade‐off has been consistently expressed over three decades of environmental change despite some turnover in the identity of dominant species.     

Environmental context alters ecological trade‐offs controlling ant coexistence in a spatially heterogeneous region

1. Ecological trade‐offs in ant (Hymenoptera: Formicidae) assemblages and their implications for coexistence boast a rich history in entomology. Yet investigations of trade‐offs have largely been limited to homogeneous environments. We examined how environmental context modifies trade‐off expression in an ant assemblage spanning a heterogeneous region in central Florida, U.S.A. 2. We examined how trade‐off expression is altered among two contrasting habitat types: open shrub and forest. We tested for the presence of the dominance‐discovery trade‐off and two dominance‐thermal tolerance trade‐offs by estimating behavioral dominance, discovery ability, and thermal tolerance (foraging thermal limit, lethal temperature, and maximal abundance temperature) for a wide range of interacting ant species. 3. We found significantly linear dominance hierarchies in both shrub and forest habitats, showing dominant species out‐compete subordinates for food resources. In thermally stressful shrub habitats, subordinates exhibit higher thermal tolerances, take greater thermal risks, and reach maximum forager abundances at higher temperatures than do dominant species. This suggests temperature mediated trade‐offs control coexistence in shrub habitat. In thermally moderate forest habitat, we found limited evidence for trade‐offs between competitive dominance and resource discovery or between dominance and thermal traits, implying other processes control coexistence. These results demonstrate that trade‐offs controlling ant coexistence may be contingent on environmental context.     

Competitive response of savanna tree seedlings to C4 grasses is negatively related to photosynthesis rate

Savanna tree species vary in the magnitude of their response to grass competition, but the functional traits that explain this variation remain largely unknown. To address this gap, we grew seedlings of 10 savanna tree species with and without grasses in a controlled greenhouse experiment. We found strong interspecific differences in tree competitive response, which was positively related to photosynthesis rates, suggesting a trade‐off between the ability to grow well under conditions of low and high grass biomass across tree species. We also found no competitive effect of tree seedlings on grass, suggesting strong tree‐grass competitive asymmetry. Our results identify a potentially important trade‐off that enhances our ability to predict how savanna tree communities might respond to variation in grass competition.     

Characterizing the contribution of plasticity and genetic differentiation to community‐level trait responses to environmental change

The match between functional trait variation in communities and environmental gradients is maintained by three processes: phenotypic plasticity and genetic differentiation (intraspecific processes), and species turnover (interspecific). Recently, evidence has emerged suggesting that intraspecific variation might have a potentially large role in driving functional community composition and response to environmental change. However, empirical evidence quantifying the respective importance of phenotypic plasticity and genetic differentiation relative to species turnover is still lacking. We performed a reciprocal transplant experiment using a common herbaceous plant species (Oxalis montana) among low‐, mid‐, and high‐elevation sites to first quantify the contributions of plasticity and genetic differentiation in driving intraspecific variation in three traits: height, specific leaf area, and leaf area. We next compared the contributions of these intraspecific drivers of community trait–environment matching to that of species turnover, which had been previously assessed along the same elevational gradient. Plasticity was the dominant driver of intraspecific trait variation across elevation in all traits, with only a small contribution of genetic differentiation among populations. Local adaptation was not detected to a major extent along the gradient. Fitness components were greatest in O. montana plants with trait values closest to the local community‐weighted means, thus supporting the common assumption that community‐weighted mean trait values represent selective optima. Our results suggest that community‐level trait responses to ongoing climate change should be mostly mediated by species turnover, even at the small spatial scale of our study, with an especially small contribution of evolutionary adaptation within species.     

TRADE‐OFFS,SPATIAL HETEROGENEITY,AND THE MAINTENANCE OF MICROBIAL DIVERSITY

Specialization and concomitant trade‐offs are assumed to underlie the non‐neutral coexistence of lineages. Trade‐offs across heterogeneous environments can promote diversity by preventing competitive exclusion. However, the importance of trade‐offs in maintaining diversity in natural microbial assemblages is unclear, as trade‐offs are frequently not detected in artificial evolution experiments. Stressful conditions associated with patches of heavy‐metal enriched serpentine soils provide excellent opportunities for examining how heterogeneity may foster genetic diversity. Using a spatially replicated design, we demonstrate that rhizobium bacteria symbiotic with legumes inhabiting contrasting serpentine and nonserpentine soils exhibit a trade‐off between a genotype's nickel tolerance and its ability to replicate rapidly. Furthermore, we detected adaptive divergence in rhizobial assemblages across soil type heterogeneity at multiple sites, suggesting that this trade‐off may promote the coexistence of phenotypically distinct bacterial lineages. Trade‐offs and adaptive divergence may be important factors maintaining the tremendous diversity within natural assemblages of bacteria.     

Environmental heterogeneity does not affect levels of phenotypic plasticity in natural populations of three Drosophila species

Adaptation of natural populations to variable environmental conditions may occur by changes in trait means and/or in the levels of plasticity. Theory predicts that environmental heterogeneity favors plasticity of adaptive traits. Here we investigated the performance in several traits of three sympatric Drosophila species freshly collected in two environments that differ in the heterogeneity of environmental conditions. Differences in trait means within species were found in several traits, indicating that populations differed in their evolutionary response to the environmental conditions of their origin. Different species showed distinct adaptation with a very different role of plasticity across species for coping with environmental changes. However, geographically distinct populations of the same species generally displayed the same levels of plasticity as induced by fluctuating thermal regimes. This indicates a weak and trait‐specific effect of environmental heterogeneity on plasticity. Furthermore, similar levels of plasticity were found in a laboratory‐adapted population of Drosophila melanogaster with a common geographic origin but adapted to the laboratory conditions for more than 100 generations. Thus, this study does not confirm theoretical predictions on the degree of adaptive plasticity among populations in relation to environmental heterogeneity but shows a very distinct role of species‐specific plasticity.     

Phenotypic constraints and community structure: Linking trade‐offs within and among species

Trade‐offs are central to many topics in biology, from the evolution of life histories to ecological mechanisms of species coexistence. Trade‐offs observed among species may reflect pervasive constraints on phenotypes that are achievable given biophysical and resource limitations. If so, then among‐species trade‐offs should be consistent with trade‐offs within species. Alternatively, trait variation among co‐occurring species may reflect historical contingencies during community assembly rather than within‐species constraints. Here, we test whether a key trade‐off between relative growth rate (RGR) and water‐use efficiency (WUE) among Sonoran Desert winter annual plants is apparent within four species representing different strategies in the system. We grew progeny of maternal families from multiple populations in a greenhouse common garden. One species, Pectocarya recurvata, displayed the expected RGR–WUE trade‐off among families within populations. For other species, although RGR and WUE often varied clinally among populations, among‐family variation within populations was lacking, implicating a role for past selection on these traits. Our results suggest that a combination of limited genetic variation in single traits and negative trait correlations could pose constraints on the evolution of a high‐RGR and high‐WUE phenotype within species, providing a microevolutionary explanation for phenotypes that influence community‐level patterns of abundance and coexistence.     

Species coexistence in resource-limited patterned ecosystems is facilitated by the interplay of spatial self-organisation and intraspecific competition

The exploration of mechanisms that enable species coexistence under competition for a sole limiting resource is widespread across ecology. Two examples of such facilitative processes are intraspecific competition and spatial self-organisation. These processes determine the outcome of competitive dynamics in many resource-limited patterned ecosystems, classical examples of which include dryland vegetation patterns, intertidal mussel beds and subalpine ribbon forests. Previous theoretical investigations have explained coexistence within patterned ecosystems by making strong assumptions on the differences between species (e.g. contrasting dispersal behaviours or different functional responses to resource availability). In this paper, I show that the interplay between the detrimental effects of intraspecific competition and the facilitative nature of self-organisation forms a coexistence mechanism that does not rely on species-specific assumptions and captures coexistence across a wide range of the environmental stress gradient. I use a theoretical model that captures the interactions of two generic consumer species with an explicitly modelled resource to show that coexistence relies on a balance between species' colonisation abilities and their local competitiveness, provided intraspecific competition is sufficiently strong. Crucially, the requirements on species' self-limitation for coexistence to occur differ on opposite ends of the resource input spectrum. For low resource levels, coexistence is facilitated by strong intraspecific dynamics of the species superior in its colonisation abilities, but for larger volumes of resource input, strong intraspecific competition of the locally superior species enables coexistence. Results presented in this paper also highlight the importance of hysteresis in understanding tipping points, in particular extinction events. Finally, the theoretical framework provides insights into spatial species distributions within single patches, supporting verbal hypotheses on coexistence of herbaceous and woody species in dryland vegetation patterns and suggesting potential empirical tests in the context of other patterned ecosystems.     

Competitive asymmetry, foraging area size and coexistence of annuals

Individuals allocate resources to the expansion of their foraging area and those resources are no longer available for the traits that determine how well those individuals are able to protect their foraging area against competitors. The resulting trade‐off between foraging area size and the traits associated with the ability to compete for the resources within the foraging area applies to ecological scenarios as different as territorial defence by individuals and colonies, and light competition in plants. Whether the trade‐off affects species performance in competition for resources at the area of overlap between foraging areas depends on the symmetry of resource division. In symmetric competition resources are divided equally between the competitors, while in asymmetric competition the individual with the smallest foraging area, and consequently the greatest competitive ability, gains all the resources. Competition may also be a combination of the symmetric and asymmetric processes. I studied the effects of competitive asymmetry on population dynamics and coexistence of two annual species with different sized foraging areas using an individual‐based spatially explicit simulation model. Symmetric competition favoured the species with the larger foraging area and did not allow coexistence. Competitive asymmetry favoured the species with smaller foraging area and allowed coexistence, which was due to the consequences of losing an asymmetric competition being more severe than losing a symmetric competition. The mechanism of coexistence is the larger foraging area's superiority in low population densities (little competition) and the smaller foraging area's ability to win a large foraging area when competition was intense. Competitive asymmetry and small size of both foraging areas led to population dynamics dominated by long‐term fluctuations of small intensity. Symmetric competition and large size of the foraging areas led to large short‐term fluctuations, which often resulted in the extinction of one or both of the species due to demographic stochasticity.     

Simple rules for the coexistence and competitive dominance of plants mediated by mycorrhizal fungi

Mycorrhizal fungi have been demonstrated to be important in the makeup of plant communities. Likely, their most important role is in altering mineral nutrition of plants, which, in turn, is thought to be among the most important determinants of plant competitive ability. Using mathematical models we examine what role these fungi can play in determining the competitive outcome between two plants in competition for one mineral resource. Depending on different relationships between the benefit accrued to the plant and the fungi, the presence of mycorrhizal fungi can (i) have no impact on which plant wins in competition, (ii) change the order of competitive dominance or (iii) enable coexistence when compared with the system in the absence of mycorrhizal fungi. Furthermore, environmental conditions, such as light and nutrient levels, can determine if coexistence is possible. We describe the necessary biological trade‐offs for coexistence and experimental tests for these trade‐offs.     

Coexistence introducing regulation of environmental conditions

Interactions between environmental conditions and environment-affecting species have not been investigated extensively. In this study, the population dynamics of species yielding regulative feedback between temperature (a representative of environmental condition) and species with a temperature-altering trait was examined. We considered a simple closed model that described the population of two species (at least one of them had a temperature-altering trait) competing for one resource. The long-term outcomes of the competition and changes of temperature were explored against increasing background temperature. As a result of simulations, the regulation of temperature was accompanied by the coexistence of two species, which was contrary to the 'Gause's exclusion principle'. The steady-state analysis showed that (i) the temperature-altering trait allowed species to coexist and (ii) the coexistence of species with the trait could introduce the regulation of temperature. A 'trade-off' in their ability to utilize a resource plays a key role in this coexistence and homeostasis of temperature. This may imply that actual environmental conditions can be automatically stabilized by resource competition among species in natural ecosystems.     

Life‐history constraints in grassland plant species: a growth‐defence trade‐off is the norm

Plant growth can be limited by resource acquisition and defence against consumers, leading to contrasting trade‐off possibilities. The competition‐defence hypothesis posits a trade‐off between competitive ability and defence against enemies (e.g. herbivores and pathogens). The growth‐defence hypothesis suggests that strong competitors for nutrients are also defended against enemies, at a cost to growth rate. We tested these hypotheses using observations of 706 plant populations of over 500 species before and following identical fertilisation and fencing treatments at 39 grassland sites worldwide. Strong positive covariance in species responses to both treatments provided support for a growth‐defence trade‐off: populations that increased with the removal of nutrient limitation (poor competitors) also increased following removal of consumers. This result held globally across 4 years within plant life‐history groups and within the majority of individual sites. Thus, a growth‐defence trade‐off appears to be the norm, and mechanisms maintaining grassland biodiversity may operate within this constraint.