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Abstract The competitive effect of one individual on another can have impacts beyond just reductions in performance. Because species plastically respond to their environment, competition can also induce changes in species traits, and in turn, these modified traits can then affect interactions with yet other individuals. In this context, plasticity is often argued to favor species coexistence by increasing the niche differentiation between species, though experimental evidence for this hypothesis that explicitly projects competitive outcomes is largely lacking. Here, we transiently subjected four annual plant species to early‐season intraspecific or interspecific competition to explicitly induce plastic responses and then examined the response of these individuals to competitors faced later in life. Competing with nearby individuals early in the growing season tended to amplify the sensitivity of individuals to competition, and particularly so for interspecific competition, but the strength of this effect depended on the identity of the focal species. This increase in interspecific relative to intraspecific competition caused plasticity to decrease the predicted likelihood of pairwise coexistence. By combining recent theory with a new experimental approach, we provide a pathway toward integrating phenotypic plasticity into our quantitative understanding of coexistence. 

Abstract Ecological communities frequently exhibit remarkable taxonomic and trait diversity, and this diversity is consistently shown to regulate ecosystem function and resilience. However, ecologists lack a synthetic theory for how this diversity is maintained when species compete for limited resources, hampering our ability to project the future of biodiversity under climate change. Water‐limited plant communities are an ideal system in which to study these questions given (1) the diversity of hydraulic traits they exhibit, (2) the importance of this diversity for ecosystem productivity and drought resilience, and (3) forecast changes to precipitation and evapotranspiration under climate change. We developed an analytically tractable model of water and light competition in age‐structured perennial plant communities and demonstrated that high diversity is maintained through phenological division of the time between storms. We modeled a system where water arrives in the form of intermittent storms, between which plants consume the limited pool of soil water until it becomes dry enough that they must physiologically shut down to avoid embolism. Competition occurs because individuals, by consuming the shared water pool, cause their competitors to shut down earlier, harming their long‐term growth and reproduction. When total precipitation is low, plants in the model compete only for water. However, increases in precipitation can cause the canopy to close and individuals to begin competing for light. Variation among species in the minimum soil water content at which they can sustain growth without embolizing leads to emergent phenological variation, as species will shut down at varying points between storm events. When this variation is paired with a trade‐off such that species that shut down early are compensated by faster biomass accumulation, higher fecundity, or lower mortality, there is no limit to the number that can coexist. These results are robust to variation in both total precipitation and the time between storms. The model therefore offers a plausible explanation for how hydraulic trait diversity is maintained in a wide array of natural systems. More broadly, this work illustrates how the phenological division of an apparently singular resource can emerge because of common trade‐offs and ultimately foster high taxonomic and trait diversity. 

Abstract Most ecological models are based on the assumption that species interact in pairs. Diverse communities, however, can have higher‐order interactions, in which two or more species jointly impact the growth of a third species. A pitfall of the common pairwise approach is that it misses the higher‐order interactions potentially responsible for maintaining natural diversity. Here, we explore the stability properties of systems where higher‐order interactions guarantee that a specified set of abundances is a feasible equilibrium of the dynamics. Even these higher‐order interactions which lead to equilibria do not necessarily produce stable coexistence. Instead, these systems are more likely to be stable when the pairwise interactions are weak or facilitative. Correlations between the pairwise and higher‐order interactions, however, do permit robust coexistence even in diverse systems. Our work not only reveals the challenges in generating stable coexistence through higher‐order interactions but also uncovers interaction patterns that can enable diversity. 

Abstract Understanding how diversity is maintained in plant communities requires that we first understand the mechanisms of competition for limiting resources. In ecology, there is an underappreciated but fundamental distinction between systems in which the depletion of limiting resources reduces the growth rates of competitors and systems in which resource depletion reduces the time available for competitors to grow, a mechanism we call ‘competition for time’. Importantly, modern community ecology and our framing of the coexistence problem are built on the implicit assumption that competition reduces the growth rate. However, recent theoretical work suggests competition for time may be the predominant competitive mechanism in a broad array of natural communities, a significant advance given that when species compete for time, diversity‐maintaining trade‐offs emerge organically. In this study, we first introduce competition for time conceptually using a simple model of interacting species. Then, we perform an experiment in a Mediterranean annual grassland to determine whether competition for time is an important competitive mechanism in a field system. Indeed, we find that species respond to increased competition through reductions in their lifespan rather than their rate of growth. In total, our study suggests competition for time may be overlooked as a mechanism of biodiversity maintenance. 

Abstract When species simultaneously compete with two or more species of competitor, higher‐order interactions (HOIs) can lead to emergent properties not present when species interact in isolated pairs. To extend ecological theory to multi‐competitor communities, ecologists must confront the challenges of measuring and interpreting HOIs in models of competition fit to data from nature. Such efforts are hindered by the fact that different studies use different definitions, and these definitions have unclear relationships to one another. Here, we propose a distinction between ‘soft’ HOIs, which identify possible interaction modification by competitors, and ‘hard’ HOIs, which identify interactions uniquely emerging in systems with three or more competitors. We show how these two classes of HOI differ in their motivation and interpretation, as well as the tests one uses to identify them in models fit to data. We then show how to operationalise this structure of definitions by analysing the results of a simulated competition experiment underlain by a consumer resource model. In the course of doing so, we clarify the challenges of interpreting HOIs in nature, and suggest a more precise framing of this research endeavour to catalyse further investigations. 

Observations of pathogen community structure provide evidence for both the coexistence and replacement of related strains. Despite many studies of specific host-pathogen systems, a unifying framework for predicting the outcomes of interactions among pathogens has remained elusive. We address this gap by developing a pathogen invasion theory (PIT) based on modern ecological coexistence theory and testing the resulting framework against empirical systems. Across major human pathogens, PIT predicts near-universal mutual susceptibility of one strain to invasion by another strain. However, predicting co-circulation from mutual invasion also depends on the degree to which susceptible abundance is reduced below the invasion threshold by overcompensatory epidemic dynamics, and the time it takes for susceptibles to replenish. The transmission advantage of an invading strain and the strength and duration of immunity are key determinants of susceptible dynamics. PIT unifies existing ideas about pathogen co-circulation, offering a quantitative framework for predicting the emergence of novel pathogen strains. 

A central assumption in most ecological models is that the interactions in a community operate only between pairs of species. However, two species may interactively affect the growth of a focal species. Although interactions among three or more species, called higher-order interactions, have the potential to modify our theoretical understanding of coexistence, ecologists lack clear expectations for how these interactions shape community structure. Here we analytically predict and numerically confirm how the variability and strength of higher-order interactions affect species coexistence. We found that as higher-order interaction strengths became more variable across species, fewer species could coexist, echoing the behavior of pairwise models. If interspecific higher-order interactions became too harmful relative to self-regulation, coexistence in diverse communities was destabilized, but coexistence was also lost when these interactions were too weak and mutualistic higher-order effects became prevalent. This behavior depended on the functional form of the interactions as the destabilizing effects of the mutualistic higher-order interactions were ameliorated when their strength saturated with species’ densities. Last, we showed that more species-rich communities structured by higher-order interactions lose species more readily than their species-poor counterparts, generalizing classic results for community stability. Our work provides needed theoretical expectations for how higher-order interactions impact species coexistence in diverse communities.