A Spatial Signal of Niche Differentiation in Tropical Forests.

AbstractExplaining diversity in tropical forests remains a challenge in community ecology. Theory tells us that species differences can stabilize communities by reducing competition, while species similarities can promote diversity by reducing fitness differences and thus prolonging the time to competitive exclusion. Combined, these processes may lead to clustering of species such that species are niche differentiated across clusters and share a niche within each cluster. Here, we characterize this partial niche differentiation in a tropical forest in Panama by measuring spatial clustering of woody plants and relating these clusters to local soil conditions. We find that species were spatially clustered and the clusters were associated with specific concentrations of soil nutrients, reflecting the existence of nutrient niches. Species were almost twice as likely to recruit in their own nutrient niche. A decision tree algorithm showed that local soil conditions correctly predicted the niche of the trees with up to 85% accuracy. Iron, zinc, phosphorus, manganese, and soil pH were among the best predictors of species clusters.

Reciprocal inhibition and competitive hierarchy cause negative biodiversity-ecosystem function relationships.

The relationship between biodiversity and ecosystem function (BEF) captivates ecologists, but the factors responsible for the direction of this relationship remain unclear. While higher ecosystem functioning at higher biodiversity levels ('positive BEF') is not universal in nature, negative BEF relationships seem puzzlingly rare. Here, we develop a dynamical consumer-resource model inspired by microbial decomposer communities in pitcher plant leaves to investigate BEF. We manipulate microbial diversity via controlled colonization and measure their function as total ammonia production. We test how niche partitioning among bacteria and other ecological processes influence BEF in the leaves. We find that a negative BEF can emerge from reciprocal interspecific inhibition in ammonia production causing a negative complementarity effect, or from competitive hierarchies causing a negative selection effect. Absent these factors, a positive BEF was the typical outcome. Our findings provide a potential explanation for the rarity of negative BEF in empirical data.

Niche theory for positive plant-soil feedbacks.

Interactions between plants and the soil are an important ecological process in terrestrial ecosystems as they affect plant community structure: when and where we find different plant species. Those interactions are typically thought of as one-directional: local soil conditions filter through dispersing species to produce a community of locally adapted plants. However, plants can modify local physicochemical soil conditions via their roots and associations with soil microbes. These may in turn affect the local fitness of other plants, making plant-soil interactions bidirectional. In order to understand how they differ from other ecological processes that structure plant communities, we need a theory connecting these individual-level plant-soil feedbacks to community-level patterns. Here, we build this theory with a mathematical model of plant community dynamics in which soil conditioning is explicitly modeled over time and depends on the density of the plants. We analyze this model to describe the long-term composition and spatial distribution of the plant community. Our main result is that positive plant-soil feedbacks will create clustering of species with similar soil preferences. The composition of these clusters is further influenced by niche width and conditioning strength. In contrast with competitive dynamics driven by niche overlap, only species belonging to the same cluster can maintain high relative abundance in the community. Spatial heterogeneity in the form of an environmental gradient generates patches, each representing a single cluster. However, such patchiness is disfavored when species differ in dispersal ability. We show that stronger dispersers cannot take over the habitat as long as an exogenous driver favors soil conditions that benefit the other species. If exogenous drivers supersede soil conditioning by plants, we retrieve classic habitat filtering, where species are selected based on their suitability to the local environment. Overall, we provide a novel mathematical model for positive plant-soil feedback that we use to describe the spatial patterns of plant abundance and traits related to soil preference and conditioning ability.

Emergent neutrality in consumer-resource dynamics.

Neutral theory assumes all species and individuals in a community are ecologically equivalent. This controversial hypothesis has been tested across many taxonomic groups and environmental contexts, and successfully predicts species abundance distributions across multiple high-diversity communities. However, it has been critiqued for its failure to predict a broader range of community properties, particularly regarding community dynamics from generational to geological timescales. Moreover, it is unclear whether neutrality can ever be a true description of a community given the ubiquity of interspecific differences, which presumably lead to ecological inequivalences. Here we derive analytical predictions for when and why non-neutral communities of consumers and resources may present neutral-like outcomes, which we verify using numerical simulations. Our results, which span both static and dynamical community properties, demonstrate the limitations of summarizing distributions to detect non-neutrality, and provide a potential explanation for the successes of neutral theory as a description of macroecological pattern.

Counting niches: Abundance-by-trait patterns reveal niche partitioning in a Neotropical forest.

Tropical forests challenge us to understand biodiversity, as numerous seemingly similar species persist on only a handful of shared resources. Recent ecological theory posits that biodiversity is sustained by a combination of species differences reducing interspecific competition and species similarities increasing time to competitive exclusion. Together, these mechanisms counterintuitively predict that competing species should cluster by traits, in contrast with traditional expectations of trait overdispersion. Here, we show for the first time that trees in a tropical forest exhibit a clustering pattern. In a 50-ha plot on Barro Colorado Island in Panama, species abundances exhibit clusters in two traits connected to light capture strategy, suggesting that competition for light structures community composition. Notably, we find four clusters by maximum height, quantitatively supporting the classical grouping of Neotropical woody plants into shrubs, understory, midstory, and canopy layers.

Generalizing clusters of similar species as a signature of coexistence under competition.

Patterns of trait distribution among competing species can potentially reveal the processes that allow them to coexist. It has been recently proposed that competition may drive the spontaneous emergence of niches comprising clusters of similar species, in contrast with the dominant paradigm of greater-than-chance species differences. However, current clustering theory relies largely on heuristic rather than mechanistic models. Furthermore, studies of models incorporating demographic stochasticity and immigration, two key players in community assembly, did not observe clusters. Here we demonstrate clustering under partitioning of resources, partitioning of environmental gradients, and a competition-colonization tradeoff. We show that clusters are robust to demographic stochasticity, and can persist under immigration. While immigration may sustain clusters that are otherwise transient, too much dilutes the pattern. In order to detect and quantify clusters in nature, we introduce and validate metrics which have no free parameters nor require arbitrary trait binning, and weigh species by their abundances rather than relying on a presence-absence count. By generalizing beyond the circumstances where clusters have been observed, our study contributes to establishing them as an update to classical trait patterning theory.

Emergent niche structuring leads to increased differences from neutrality in species abundance distributions.

Species abundance distributions must reflect the dynamic processes involved in community assembly, but whether and when specific processes lead to distinguishable signals is not well understood. Biodiversity and species abundances may be shaped by a variety of influences, but particular attention has been paid to competition, which can involve neutral dynamics, where competitor abundances are governed only by demographic stochasticity and immigration, and dynamics driven by trait differences that enable stable coexistence through the formation of niches. Key recent studies of the species abundance patterns of communities with niches employ simple models with pre-imposed niche structure. These studies suggest that species abundance distributions are insensitive to the relative contributions of niche and neutral processes, especially when diversity is much higher than the number of niches. Here we analyze results from a stochastic population model with competition driven by trait differences. With this model, niche structure emerges as clumps of species that persist along the trait axis, and leads to more substantial differences from neutral species abundance distributions than have been previously shown. We show that heterogeneity in "between-niche" interaction strength (i.e., in the strength of competition between species in different niches) plays the dominant role in shaping the species abundances along the trait axis, acting as a biotic filter favoring species at the centers of niches. Furthermore, we show that heterogeneity in "within-niche" interactions (i.e., in the competition between species in the same niche) counteracts the influence of heterogeneity in "between-niche" interactions on the SAD to some degree. Our results suggest that competitive interactions that produce niches can also influence the shapes of SADs.

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.

Can editors save peer review from peer reviewers?

Peer review is the gold standard for scientific communication, but its ability to guarantee the quality of published research remains difficult to verify. Recent modeling studies suggest that peer review is sensitive to reviewer misbehavior, and it has been claimed that referees who sabotage work they perceive as competition may severely undermine the quality of publications. Here we examine which aspects of suboptimal reviewing practices most strongly impact quality, and test different mitigating strategies that editors may employ to counter them. We find that the biggest hazard to the quality of published literature is not selfish rejection of high-quality manuscripts but indifferent acceptance of low-quality ones. Bypassing or blacklisting bad reviewers and consulting additional reviewers to settle disagreements can reduce but not eliminate the impact. The other editorial strategies we tested do not significantly improve quality, but pairing manuscripts to reviewers unlikely to selfishly reject them and allowing revision of rejected manuscripts minimize rejection of above-average manuscripts. In its current form, peer review offers few incentives for impartial reviewing efforts. Editors can help, but structural changes are more likely to have a stronger impact.

Biodiversity maintenance may be lower under partial niche differentiation than under neutrality.

Niche differentiation is normally regarded as a key promoter of species coexistence in competitive systems. One might therefore expect that relative to neutral assemblages, niche-differentiated communities should support more species with longer persistence and lower probability of extinction. Here we compare stochastic niche and neutral dynamics in simulated assemblages, and find that when local dynamics combine with immigration from a regional pool, the effect of niches can be more complex. Trait variation that lessens competition between species will not necessarily give all immigrating species their own niche to occupy. Such partial niche differentiation protects certain species from local extinction, but precipitates exclusion of others. Differences in regional abundances and intrinsic growth rates have similar impacts on persistence times as niche differentiation, and therefore blur the distinction between niche and neutral dynamical patterns-although niche dynamics will influence which species persist longer. Ultimately, unless the number of niches available to species is sufficiently high, niches may actually heighten extinction rates and lower species richness and local persistence times. Our results help make sense of recent observations of community dynamics, and point to the dynamical observations needed to discern the influence of niche differentiation.

Can Clustering in Genotype Space Reveal "Niches"?

Community ecology lacks the success enjoyed by population genetics to quantify the relative roles played by deterministic and stochastic processes. It has been proposed that clustered patterns of abundance in genotype space provide evidence of selection in microbial communities, since no such clustering would arise in the absence of selection. We critique this test for its unrealistic null hypothesis. We show mathematically and with simulations that point mutations alone lead to clustering in genotype space by causing correlations between abundances of similar genotypes. We also show potential deviations from the mutation-only pattern caused by immigration from a source pool. Clustered patterns in genotype space may still be revealing of selection if analyzed quantitatively but only if neutral and selective regimes can be distinguished once mutation and immigration are included in the null model.

The effect of intraspecific variation and heritability on community pattern and robustness.

Intraspecific trait variation is widespread in nature, yet its effects on community dynamics are not well understood. Here we explore the consequences of intraspecific trait variation for coexistence in two- and multispecies competitive communities. For two species, the likelihood of coexistence is in general reduced by intraspecific variation, except when the species have almost equal trait means but different trait variances, such that one is a generalist and the other a specialist consumer. In multispecies communities, the only strong effect of non-heritable intraspecific variation is to reduce expected species richness. However, when intraspecific variation is heritable, allowing for the possibility of trait evolution, communities are much more resilient against environmental disturbance and exhibit far more predictable trait patterns. Our results are robust to varying model parameters and relaxing model assumptions.

Revising the tolerance-fecundity trade-off; or, on the consequences of discontinuous resource use for limiting similarity, species diversity, and trait dispersion.

The recently proposed tolerance-fecundity trade-off model represents a step forward in the study of seed size diversity in plant communities. However, it uses an oversimplified picture of seed tolerance, with an infinitely sharp threshold: the probability that a seed tolerate a given stress level is either 1 or 0. This invites a revision of the model, presented here. We demonstrate that this simplification has large impacts on model behavior, including altering predictions regarding limiting similarity, raising expected diversity levels, and lessening expected spacing between species along the trait axis. Such dramatic impacts ultimately stem from the fact that a discontinuity in the probability of tolerating a site drastically reduces competition between similar species. This is one example of a class of models with a nondifferentiable peak in the competition kernel, which we recently showed is produced by resource use unrealistically modeled as discontinuous and affects fundamental predictions regarding limiting similarity. This article illustrates those general results and offers a revised model of the tolerance-fecundity trade-off.