Assessing the roles of nitrogen, biomass, and niche dimensionality as drivers of species loss in grassland communities.

Eutrophication is a major driver of species loss in plant communities worldwide. However, the underlying mechanisms of this phenomenon are controversial. Previous studies have raised three main explanations: 1) High levels of soil resources increase standing biomass, thereby intensifying competitive interactions (the "biomass-driven competition hypothesis"). 2) High levels of soil resources reduce the potential for resource-based niche partitioning (the "niche dimension hypothesis"). 3) Increasing soil nitrogen causes stress by changing the abiotic or biotic conditions (the "nitrogen detriment hypothesis"). Despite several syntheses of resource addition experiments, so far, no study has tested all of the hypotheses together. This is a major shortcoming, since the mechanisms underlying the three hypotheses are not independent. Here, we conduct a simultaneous test of the three hypotheses by integrating data from 630 resource addition experiments located in 99 sites worldwide. Our results provide strong support for the nitrogen detriment hypothesis, weaker support for the biomass-driven competition hypothesis, and negligible support for the niche dimension hypothesis. The results further show that the indirect effect of nitrogen through its effect on biomass is minor compared to its direct effect and is much larger than that of all other resources (phosphorus, potassium, and water). Thus, we conclude that nitrogen-specific mechanisms are more important than biomass or niche dimensionality as drivers of species loss under high levels of soil resources. This conclusion is highly relevant for future attempts to reduce biodiversity loss caused by global eutrophication.

An experimental test of the area-heterogeneity tradeoff.

A fundamental property of ecosystems is a tradeoff between the number and size of habitats: as the number of habitats within a fixed area increases, the average area per habitat must decrease. This tradeoff is termed the "area-heterogeneity tradeoff." Theoretical models suggest that the reduction in habitat sizes under high levels of heterogeneity may cause a decline in species richness because it reduces the amount of effective area available for individual species under high levels of heterogeneity, thereby increasing the likelihood of stochastic extinctions. Here, we test this prediction using an experiment that allows us to separate the effect of the area-heterogeneity tradeoff from the total effect of habitat heterogeneity. Surprisingly, despite considerable extinctions, reduction in the amount of effective area available per species facilitated rather than reduced richness in the study communities. Our data suggest that the mechanism behind this positive effect was a decrease in the probability of deterministic competitive exclusion. We conclude that the area-heterogeneity tradeoff may have both negative and positive implications for biodiversity and that its net effect depends on the relative importance of stochastic vs. deterministic drivers of extinction in the relevant system. Our finding that the area-heterogeneity tradeoff may contribute to biodiversity adds a dimension to existing ecological theory and is highly relevant for understanding and predicting biodiversity responses to natural and anthropogenic variations in the environment.

Heterogeneity-diversity relationships in sessile organisms: a unified framework.

The hypothesis that environmental heterogeneity promotes species richness by increasing opportunities for niche partitioning is a fundamental paradigm in ecology. However, recent studies suggest that heterogeneity-diversity relationships (HDR) are more complex than expected from this niche-based perspective, and often show a decrease in richness at high levels of heterogeneity. These findings have motivated ecologists to propose new mechanisms that may explain such deviations. Here we provide an overview of currently recognised mechanisms affecting the shape of HDRs and present a conceptual model that integrates all previously proposed mechanisms within a unified framework. We also translate the proposed framework into an explicit community dynamic model and use the model as a tool for generating testable predictions concerning how landscape properties interact with species traits in determining the shape of HDRs. Our main finding is that, despite the enormous complexity of such interactions, the predicted HDRs are rather simple, ranging from positive to unimodal patterns in a highly consistent and predictable manner.

Mechanisms of seed mass variation along resource gradients.

The enormous variation in seed mass along gradients of soil resources has fascinated plant ecologists for decades. However, so far, this research has focused on the description of such variation, rather than its underlying mechanisms. Here we experimentally test a recent model relating such variation to two fundamental properties of plant growth: allometry of biomass growth and size-asymmetry of light competition. According to the model, mean seed mass should increase, and the variance of seed mass should show a unimodal response, to increasing soil resource availability (productivity). We test these predictions and their underlying assumptions using a combination of field observations, mesocosm experiments and greenhouse experiments focusing on Mediterranean annual plants. Our results support the predictions and assumptions of the model, and allow us to reject alternative models of seed mass variation. We conclude that growth-allometry and size-asymmetric light competition are key drivers of seed-mass variation along soil resource gradients.

Dispersal increases ecological selection by increasing effective community size.

Selection and drift are universally accepted as the cornerstones of evolutionary changes. Recent theories extend this view to ecological changes, arguing that any change in species composition is driven by deterministic fitness differences among species (enhancing selection) and/or stochasticity in birth and death rates of individuals within species (enhancing drift). These forces have contrasting effects on the predictability of ecological dynamics, and thus understanding the factors affecting their relative importance is crucial for understanding ecological dynamics. Here we test the hypothesis that dispersal increases the relative importance of ecological selection by increasing the effective size of the community (i.e., the size relevant for competitive interactions and drift). According to our hypothesis, dispersal increases the effective size of the community by mixing individuals from different localities. This effect diminishes the relative importance of demographic stochasticity, thereby reducing drift and increasing the relative importance of selective forces as drivers of species composition. We tested our hypothesis, which we term the "effective community size" hypothesis, using two independent experiments focusing on annual plants: a field experiment in which we manipulated the magnitude of dispersal and a mesocosm experiment in which we directly manipulated the effective size of the communities. Both experiments, as well as related model simulations, were consistent with the hypothesis that increasing dispersal increases the role of selective forces as drivers of species composition. This finding has important implications for our understanding of the fundamental forces affecting community dynamics, as well as the management of species diversity, particularly in patchy and fragmented environments.

Seed mass diversity along resource gradients: the role of allometric growth rate and size-asymmetric competition.

The large variation in seed mass among species inspired a vast array of theoretical and empirical research attempting to explain this variation. So far, seed mass variation was investigated by two classes of studies. One class focuses on species varying in seed mass within communities, while the second focuses on variation between communities, most often with respect to resource gradients. Here, we develop a model capable of simultaneously explaining variation in seed mass within and between communities. The model describes resource competition (for both soil and light resources) in annual communities and incorporates two fundamental aspects: light asymmetry (higher light acquisition per unit biomass for larger individuals) and growth allometry (negative dependency of relative growth rate on plant biomass). Results show that both factors are critical in determining patterns of seed mass variation. In general, growth allometry increases the reproductive success of small-seeded species while light asymmetry increases the reproductive success of large-seeded species. Increasing availability of soil resources increases light competition, thereby increasing the reproductive success of large-seeded species and ultimately the community (weighted) mean seed mass. An unexpected prediction of the model is that maximum variation in community seed mass (a measure of functional diversity) occurs under intermediate levels of soil resources. Extensions of the model incorporating size-dependent seed survival and disturbance also show patterns consistent with empirical observations. These overall results suggest that the mechanisms captured by the model are important in determining patterns of species and functional diversity.

Light asymmetry explains the effect of nutrient enrichment on grassland diversity.

One of the most ubiquitous patterns in plant ecology is species loss following nutrient enrichment. A common explanation for this universal pattern is an increase in the size asymmetry of light partitioning (the degree to which large plants receive more light per unit biomass than smaller plants), which accelerates the rates of competitive exclusions. This 'light asymmetry hypothesis' has been confirmed by mathematical models, but has never been tested in natural communities due to the lack of appropriate methodology for measuring the size asymmetry of light partitioning in natural communities. Here, we use a novel approach for quantifying the asymmetry of light competition which is based on measurements of the vertical distribution of light below the canopy. Using our approach, we demonstrate that an increase in light asymmetry is the main mechanism behind the negative effect of nutrient enrichment on species richness. Our results provide a possible explanation for one of the main sources of contemporary species loss in terrestrial plant communities.

Quantifying Competitive Exclusion and Competitive Release in Ecological Communities: A Conceptual Framework and a Case Study.

A fundamental notion in community ecology is that local species diversity reflects some balance between the contrasting forces of competitive exclusion and competitive release. Quantifying this balance is not trivial, and requires data on the magnitude of both processes in the same system, as well as appropriate methodology to integrate and interpret such data. Here we present a novel framework for empirical studies of the balance between competitive exclusion and competitive release and demonstrate its applicability using data from a Mediterranean annual grassland where grazing is a major mechanism of competitive release. Empirical data on the balance between competitive exclusion and competitive release are crucial for understanding observed patterns of variation in local species diversity and the proposed approach provides a simple framework for the collection, interpretation, and synthesis of such data.

A neutral theory with environmental stochasticity explains static and dynamic properties of ecological communities.

Understanding the forces shaping ecological communities is crucial to basic science and conservation. Neutral theory has made considerable progress in explaining static properties of communities, like species abundance distributions (SADs), with a simple and generic model, but was criticised for making unrealistic predictions of fundamental dynamic patterns and for being sensitive to interspecific differences in fitness. Here, we show that a generalised neutral theory incorporating environmental stochasticity may resolve these limitations. We apply the theory to real data (the tropical forest of Barro Colorado Island) and demonstrate that it much better explains the properties of short-term population fluctuations and the decay of compositional similarity with time, while retaining the ability to explain SADs. Furthermore, the predictions are considerably more robust to interspecific fitness differences. Our results suggest that this integration of niches and stochasticity may serve as a minimalistic framework explaining fundamental static and dynamic characteristics of ecological communities.

Niche versus neutrality: a dynamical analysis.

Understanding the forces shaping ecological communities is of crucial importance for basic science and conservation. After 50 years in which ecological theory has focused on either stable communities driven by niche-based forces or nonstable "neutral" communities driven by demographic stochasticity, contemporary theories suggest that ecological communities are driven by the simultaneous effects of both types of mechanisms. Here we examine this paradigm using the longest available records for the dynamics of tropical trees and breeding birds. Applying a macroecological approach and fluctuation analysis techniques borrowed from statistical physics, we show that both stabilizing mechanisms and demographic stochasticity fail to play a dominant role in shaping assemblages over time. Rather, community dynamics in these two very different systems is predominantly driven by environmental stochasticity. Clearly, the current melding of niche and neutral theories cannot account for such dynamics. Our results highlight the need for a new theory of community dynamics integrating environmental stochasticity with weak stabilizing forces and suggest that such theory may better describe the dynamics of ecological communities than current neutral theories, deterministic niche-based theories, or recent hybrids.

Competitive exclusion, beta diversity, and deterministic vs. stochastic drivers of community assembly.

Species diversity has two components - number of species and spatial turnover in species composition (beta-diversity). Using a field experiment focusing on a system of Mediterranean grasslands, we show that interspecific competition may influence the two components in the same direction or in opposite directions, depending on whether competitive exclusions are deterministic or stochastic. Deterministic exclusions reduce both patch-scale richness and beta-diversity, thereby homogenising the community. Stochastic extinctions reduce richness at the patch scale, but increase the differences in species composition among patches. These results indicate that studies of competitive effects on beta diversity may help to distinguish between deterministic and stochastic components of competitive exclusion. Such distinction is crucial for understanding the causal relationship between competition and species diversity, one of the oldest and most fundamental questions in ecology.

Temporal fluctuation scaling in populations and communities.

Taylor's law, one of the most widely accepted generalizations in ecology, states that the variance of a population abundance time series scales as a power law of its mean. Here we reexamine this law and the empirical evidence presented in support of it. Specifically, we show that the exponent generally depends on the length of the time series, and its value reflects the combined effect of many underlying mechanisms. Moreover, sampling errors alone, when presented on a double logarithmic scale, are sufficient to produce an apparent power law. This raises questions regarding the usefulness of Taylor's law for understanding ecological processes. As an alternative approach, we focus on short-term fluctuations and derive a generic null model for the variance-to-mean ratio in population time series from a demographic model that incorporates the combined effects of demographic and environmental stochasticity. After comparing the predictions of the proposed null model with the fluctuations observed in empirical data sets, we suggest an alternative expression for fluctuation scaling in population time series. Analyzing population fluctuations as we have proposed here may provide new applied (e.g., estimation of species persistence times) and theoretical (e.g., the neutral theory of biodiversity) insights that can be derived from more generally available short-term monitoring data.

Area-heterogeneity tradeoff and the diversity of ecological communities.

For more than 50 y ecologists have believed that spatial heterogeneity in habitat conditions promotes species richness by increasing opportunities for niche partitioning. However, a recent stochastic model combining the main elements of niche theory and island biogeography theory suggests that environmental heterogeneity has a general unimodal rather than a positive effect on species richness. This result was explained by an inherent tradeoff between environmental heterogeneity and the amount of suitable area available for individual species: for a given area, as heterogeneity increases, the amount of effective area available for individual species decreases, thereby reducing population sizes and increasing the likelihood of stochastic extinctions. Here we provide a comprehensive evaluation of this hypothesis. First we analyze an extensive database of breeding bird distribution in Catalonia and show that patterns of species richness, species abundance, and extinction rates are consistent with the predictions of the area-heterogeneity tradeoff and its proposed mechanisms. We then perform a metaanalysis of heterogeneity-diversity relationships in 54 published datasets and show that empirical data better fit the unimodal pattern predicted by the area-heterogeneity tradeoff than the positive pattern predicted by classic niche theory. Simulations in which species may have variable niche widths along a continuous environmental gradient are consistent with all empirical findings. The area-heterogeneity tradeoff brings a unique perspective to current theories of species diversity and has important implications for biodiversity conservation.

Functional trade-offs increase species diversity in experimental plant communities.

Functional trade-offs have long been recognised as important mechanisms of species coexistence, but direct experimental evidence for such mechanisms is extremely rare. Here, we test the effect of one classical trade-off - a negative correlation between seed size and seed number - by establishing microcosm plant communities with positive, negative and no correlation between seed size and seed number and analysing the effect of the seed size/number correlation on species richness. Consistent with theory, a negative correlation between seed size and seed number led to a higher number of species in the communities and a corresponding wider range of seed size (a measure of functional richness) by promoting coexistence of large- and small-seeded species. Our study provides the first direct evidence that a seed size/number trade-off may contribute to species coexistence, and at a wider context, demonstrates the potential role of functional trade-offs in maintaining species diversity.

A general framework for neutral models of community dynamics.

Neutral models of community dynamics are a powerful tool for ecological research, but their applications are currently limited to unrealistically simple types of dynamics and ignore much of the complexity that characterize natural ecosystems. Here, we present a new analytical framework for neutral models that unifies existing models of neutral communities and extends the applicability of existing models to a much wider spectrum of ecological phenomena. The new framework extends the concept of neutrality to fitness equivalence and in spite of its simplicity explains a wide spectrum of empirical patterns of species diversity including positive, negative and unimodal productivity-diversity relationships; gradual and highly delayed declines in species diversity with habitat loss; and positive and negative responses of species diversity to habitat heterogeneity. Surprisingly, the abundance distribution in all of these cases is given by the dispersal limited multinomial (DLM), the abundance distribution in Hubbell's zero-sum model, showing DLM's robustness and demonstrating that it cannot be used to infer the underlying community dynamics. These results support the hypothesis that ecological communities are regulated by a limited set of fundamental mechanisms much simpler than could be expected from their immense complexity.

Demographic analysis of Hubbell's neutral theory of biodiversity.

Hubbell's neutral model is increasingly applied in both theoretical and empirical studies but so far little attention has been paid to the ecological mechanisms that determine species diversity in neutral communities. In this contribution we use a stochastic individual-based Markovian model to provide an explicit derivation of Hubbell's local community model from the fundamental processes of reproduction, mortality, and immigration, and show that such derivation provides important insights on the mechanisms regulating species diversity that cannot be obtained from the original model and its previous extensions. One important insight is that the basic parameters of Hubbell's model, community size (J) and the probability that a dying individual will be replaced by an immigrant (m), cannot be considered independent and that their interdependency leads to a counterintuitive trade-off between community size and species diversity. We further demonstrate that Hubbell's treatment of community size as a free parameter hides fundamental mechanisms that influence species diversity through their effect on the size of the community. For example, while in Hubbell's model immigration can only increase species diversity by promoting colonization rates, the demographic derivation shows that immigration can also promote species diversity by reducing extinction rates. Our demographic derivation also unifies previous contrasting predictions about the effect of reproduction on species diversity by showing that both positive and negative effects are possible, and that the balance between the two effects depends on the size of the community. The demographic derivation also reconciles an apparent contradiction between Hubbell's theory and patch occupancy theory, and integrates three previously proposed mechanisms of species diversity, the More Individuals Hypothesis, the rescue effect, and the dilution effect, within a single, unified framework.

A movement ecology paradigm for unifying organismal movement research.

Movement of individual organisms is fundamental to life, quilting our planet in a rich tapestry of phenomena with diverse implications for ecosystems and humans. Movement research is both plentiful and insightful, and recent methodological advances facilitate obtaining a detailed view of individual movement. Yet, we lack a general unifying paradigm, derived from first principles, which can place movement studies within a common context and advance the development of a mature scientific discipline. This introductory article to the Movement Ecology Special Feature proposes a paradigm that integrates conceptual, theoretical, methodological, and empirical frameworks for studying movement of all organisms, from microbes to trees to elephants. We introduce a conceptual framework depicting the interplay among four basic mechanistic components of organismal movement: the internal state (why move?), motion (how to move?), and navigation (when and where to move?) capacities of the individual and the external factors affecting movement. We demonstrate how the proposed framework aids the study of various taxa and movement types; promotes the formulation of hypotheses about movement; and complements existing biomechanical, cognitive, random, and optimality paradigms of movement. The proposed framework integrates eclectic research on movement into a structured paradigm and aims at providing a basis for hypothesis generation and a vehicle facilitating the understanding of the causes, mechanisms, and spatiotemporal patterns of movement and their role in various ecological and evolutionary processes. "Now we must consider in general the common reason for moving with any movement whatever." (Aristotle, De Motu Animalium, 4th century B.C.).