The tremendous diversity of species in ecological communities has motivated a century of research into the mechanisms that maintain biodiversity. However, much of this work examines the coexistence ...of just pairs of competitors. This approach ignores those mechanisms of coexistence that emerge only in diverse competitive networks. Despite the potential for these mechanisms to create conditions under which the loss of one competitor triggers the loss of others, we lack the knowledge needed to judge their importance for coexistence in nature. Progress requires borrowing insight from the study of multitrophic interaction networks, and coupling empirical data to models of competition.
Niche differences and average fitness differences jointly determine coexistence. However, little empirical information about the magnitude of these two mechanisms is available. Using multispecies ...population models fit to long-term demographic data for common, co-occurring species in five grassland and shrubland plant communities in western North America, we estimated the strength of stabilizing niche differences and average fitness differences. In all five communities, both pairwise and full community comparisons showed evidence for strong stabilizing mechanisms and relatively small average fitness differences. For a total of 17 species pairs, a measure of niche differences based on simulations of invasion growth rates ranged from 0.59 to 0.93 with a mean of 0.81, where 0 indicates complete niche overlap and 1 indicates zero niche overlap. A corresponding measure of average fitness differences ranged from 1.02 to 2.54 with a mean of 1.53, where 1 indicates identical fitness and a value of 2 indicates a fourfold difference in sensitivity to competition. Comparisons of full communities displayed similar patterns: niche differences ranged from 0.58 to 0.69 with a mean of 0.64, and the average fitness differences ranged from 1.42 to 1.63 with a mean of 1.47. In almost every case, the stabilizing mechanisms were much stronger than minimally necessary to prevent competitive exclusion. Considering that all but one of the species we studied are perennial grasses, which are often grouped in the same functional type, the magnitude of these niche differences is surprising. In all five communities, differences between intra- and interspecific effects at the recruitment stage contributed far more to stabilization than interactions affecting growth and survival. Our results indicate that for these abundant, co-occurring species (1) dynamics are far from neutral, with strong niche differences and weak fitness differences combining to stabilize coexistence, and (2) processes operating at early life stages account for a large proportion of the stabilizing effect. Given the limitations of our inductive approach, both these findings represent hypotheses in need of experimental testing.
Summary
One of the most pervasive concepts in the study of community assembly is the metaphor of the environmental filter, which refers to abiotic factors that prevent the establishment or ...persistence of species in a particular location. The metaphor has its origins in the study of community change during succession and in plant community dynamics, although it has gained considerable attention recently as part of a surge of interest in functional trait and phylogenetic‐based approaches to the study of communities.
While the filtering metaphor has clear utility in some circumstances, it has been challenging to reconcile the environmental filtering concept with recent developments in ecological theory related to species coexistence. These advances suggest that the evidence used in many studies to assess environmental filtering is insufficient to distinguish filtering from the outcome of biotic interactions.
We re‐examine the environmental filtering metaphor from the perspective of coexistence theory. In an effort to move the discussion forward, we present a simple framework for considering the role of the environment in shaping community membership, review the literature to document the evidence typically used in environmental filtering studies and highlight research challenges to address in coming years.
The current usage of the environmental filtering term in empirical studies likely overstates the role abiotic tolerances play in shaping community structure. We recommend that the term ‘environmental filtering’ only be used to refer to cases where the abiotic environment prevents establishment or persistence in the absence of biotic interactions, although only 15% of the studies in our review presented such evidence. Finally, we urge community ecologists to consider additional mechanisms aside from environmental filtering by which the abiotic environment can shape community pattern.
Lay Summary
Selecting among competing statistical models is a core challenge in science. However, the many possible approaches and techniques for model selection, and the conflicting recommendations for their ...use, can be confusing. We contend that much confusion surrounding statistical model selection results from failing to first clearly specify the purpose of the analysis. We argue that there are three distinct goals for statistical modeling in ecology: data exploration, inference, and prediction. Once the modeling goal is clearly articulated, an appropriate model selection procedure is easier to identify. We review model selection approaches and highlight their strengths and weaknesses relative to each of the three modeling goals. We then present examples of modeling for exploration, inference, and prediction using a time series of butterfly population counts. These show how a model selection approach flows naturally from the modeling goal, leading to different models selected for different purposes, even with exactly the same data set. This review illustrates best practices for ecologists and should serve as a reminder that statistical recipes cannot substitute for critical thinking or for the use of independent data to test hypotheses and validate predictions.
Understanding long‐term coexistence of numerous competing species is a longstanding challenge in ecology. Progress requires determining which processes and species differences are most important for ...coexistence when multiple processes operate and species differ in many ways. Modern coexistence theory (MCT), formalised by Chesson, holds out the promise of doing that, but empirical applications remain scarce. We argue that MCT's mathematical complexity and subtlety have obscured the simplicity and power of its underlying ideas and hindered applications. We present a general computational approach that extends our previous solution for the storage effect to all of standard MCT's spatial and temporal coexistence mechanisms, and also process‐defined mechanisms amenable to direct study such as resource partitioning, indirect competition, and life history trade‐offs. The main components are a method to partition population growth rates into contributions from different mechanisms and their interactions, and numerical calculations in which some mechanisms are removed and others retained. We illustrate how our approach handles features that have not been analysed in the standard framework through several case studies: competing diatom species under fluctuating temperature, plant–soil feedbacks in grasslands, facilitation in a beach grass community, and niche differences with independent effects on recruitment, survival and growth in sagebrush steppe.
Trait‐based tests of coexistence mechanisms Adler, Peter B; Fajardo, Alex; Kleinhesselink, Andrew R ...
Ecology letters,
October 2013, Volume:
16, Issue:
10
Journal Article
Peer reviewed
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.
Theory predicts that intraspecific competition should be stronger than interspecific competition for any pair of stably coexisting species, yet previous literature reviews found little support for ...this pattern. We screened over 5400 publications and identified 39 studies that quantified phenomenological intraspecific and interspecific interactions in terrestrial plant communities. Of the 67% of species pairs in which both intra‐ and interspecific effects were negative (competitive), intraspecific competition was, on average, four to five‐fold stronger than interspecific competition. Of the remaining pairs, 93% featured intraspecific competition and interspecific facilitation, a situation that stabilises coexistence. The difference between intra‐ and interspecific effects tended to be larger in observational than experimental data sets, in field than greenhouse studies, and in studies that quantified population growth over the full life cycle rather than single fitness components. Our results imply that processes promoting stable coexistence at local scales are common and consequential across terrestrial plant communities.
The Net Effect of Functional Traits on Fitness Laughlin, Daniel C.; Gremer, Jennifer R.; Adler, Peter B. ...
Trends in ecology & evolution (Amsterdam),
November 2020, 2020-11-00, 20201101, Volume:
35, Issue:
11
Journal Article
Peer reviewed
Open access
Generalizing the effect of traits on performance across species may be achievable if traits explain variation in population fitness. However, testing relationships between traits and vital rates to ...infer effects on fitness can be misleading. Demographic trade-offs can generate variation in vital rates that yield equal population growth rates, thereby obscuring the net effect of traits on fitness. To address this problem, we describe a diversity of approaches to quantify intrinsic growth rates of plant populations, including experiments beyond range boundaries, density-dependent population models built from long-term demographic data, theoretical models, and methods that leverage widely available monitoring data. Linking plant traits directly to intrinsic growth rates is a fundamental step toward rigorous predictions of population dynamics and community assembly.
Community ecology seeks to generalize the effects of traits on performance across species, which may be possible if traits explain variation in population fitness.Linking traits to a specific demographic rate, such as survival or reproduction, is an important step but can be misleading because trade-offs among demographic rates obscure the net effect on fitness.Integrating demographic rates (survival, growth, and reproduction) and computing low density population growth rates incorporates these trade-offs into measures of fitness.Several approaches can be used to quantify the net effect of traits on fitness, and these approaches span a trade-off of empirical rigor and logistical ease.The adaptive value of traits is evident when intrinsic growth rates are explained by trait–environment interactions across multiple coexisting species spanning an environmental gradient.
Ecologists seek general explanations for the dramatic variation in species abundances in space and time. An increasingly popular solution is to predict species distributions, dynamics, and responses ...to environmental change based on easily measured anatomical and morphological traits. Trait-based approaches assume that simple functional traits influence fitness and life history evolution, but rigorous tests of this assumption are lacking, because they require quantitative information about the full lifecycles of many species representing different life histories. Here, we link a global traits database with empirical matrix population models for 222 species and report strong relationships between functional traits and plant life histories. Species with large seeds, long-lived leaves, or dense wood have slow life histories, with mean fitness (i.e., population growth rates) more strongly influenced by survival than by growth or fecundity, compared with fast life history species with small seeds, short-lived leaves, or soft wood. In contrast to measures of demographic contributions to fitness based on whole lifecycles, analyses focused on raw demographic rates may underestimate the strength of association between traits and mean fitness. Our results help establish the physiological basis for plant life history evolution and show the potential for trait-based approaches in population dynamics.
In many terrestrial ecosystems, variation in aboveground net primary production (ANPP) is positively correlated with variation in interannual precipitation. Global climate change will alter both the ...mean and the variance of annual precipitation, but the relative impact of these changes in precipitation on mean ANPP remains uncertain. At any given site, the slope of the precipitation‐ANPP relationship determines the sensitivity of mean ANPP to changes in mean precipitation, whereas the curvature of the precipitation‐ANPP relationship determines the sensitivity of ANPP to changes in precipitation variability. We used 58 existing long‐term data sets to characterize precipitation‐ANPP relationships in terrestrial ecosystems and to quantify the sensitivity of mean ANPP to the mean and variance of annual precipitation. We found that most study sites have a nonlinear, saturating relationship between precipitation and ANPP, but these nonlinearities were not strong. As a result of these weak nonlinearities, ANPP was nearly 40 times more sensitive to precipitation mean than variance. A 1% increase in mean precipitation caused a −0.2% to 1.8% change in mean ANPP, with a 0.64% increase on average. Sensitivities to precipitation mean peaked at sites with a mean annual precipitation near 500 mm. Changes in species composition and increased intra‐annual precipitation variability could lead to larger ANPP responses to altered precipitation regimes than predicted by our analysis.
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