Should Environmental Filtering be Abandoned? Cadotte, Marc W.; Tucker, Caroline M.
Trends in ecology & evolution (Amsterdam),
June 2017, 2017-06-00, 20170601, Letnik:
32, Številka:
6
Journal Article
Recenzirano
Environmental filtering, where the environment selects against certain species, is thought to be a major mechanism structuring communities. However, recent criticisms cast doubt on our ability to ...accurately infer filtering because competition can give rise to patterns identical to those caused by environmental filtering. While experiments can distinguish mechanisms, observational patterns are especially problematic. The environment determines community composition not only directly via survival, but also by influencing competition. If species population growth rates covary with environmental gradients, then outcomes of competitive exclusion will also vary with the environment. Here, we argue that observational studies remain valuable, but inferences about the importance of the environment cannot rely on compositional data alone, and that species abundances, population growth, or traits must be correlated with the environment.
The role of the environment for shaping community composition has been criticized recently.
Other mechanisms, such as competition, could produce similar patterns to environmental filtering.
We recognize that competition and the environment do not have separable affects on species and their interactions.
Researchers should use additional information to determine whether fitness–environment covariances influence composition.
The order of species arrival during community assembly can greatly affect species coexistence, but the strength of these effects, known as priority effects, appears highly variable across species and ...ecosystems. Furthermore, the causes of this variation remain unclear despite their fundamental importance in understanding species coexistence. Here, we show that one potential cause is environmental variability. In laboratory experiments using nectar-inhabiting microorganisms as a model system, we manipulated spatial and temporal variability of temperature, and examined consequences for priority effects. If species arrived sequentially, multiple species coexisted under variable temperature, but not under constant temperature. Temperature variability prevented extinction of late-arriving species that would have been excluded owing to priority effects if temperature had been constant. By contrast, if species arrived simultaneously, species coexisted under both variable and constant temperatures. We propose possible mechanisms underlying these results using a mathematical model that incorporates contrasting effects of microbial species on nectar pH and amino acids. Overall, our findings suggest that understanding consequences of priority effects for species coexistence requires explicit consideration of environmental variability.
ABSTRACT
The use of phylogenies in ecology is increasingly common and has broadened our understanding of biological diversity. Ecological sub‐disciplines, particularly conservation, community ecology ...and macroecology, all recognize the value of evolutionary relationships but the resulting development of phylogenetic approaches has led to a proliferation of phylogenetic diversity metrics. The use of many metrics across the sub‐disciplines hampers potential meta‐analyses, syntheses, and generalizations of existing results. Further, there is no guide for selecting the appropriate metric for a given question, and different metrics are frequently used to address similar questions. To improve the choice, application, and interpretation of phylo‐diversity metrics, we organize existing metrics by expanding on a unifying framework for phylogenetic information.
Generally, questions about phylogenetic relationships within or between assemblages tend to ask three types of question: how much; how different; or how regular? We show that these questions reflect three dimensions of a phylogenetic tree: richness, divergence, and regularity. We classify 70 existing phylo‐diversity metrics based on their mathematical form within these three dimensions and identify ‘anchor’ representatives: for α‐diversity metrics these are PD (Faith's phylogenetic diversity), MPD (mean pairwise distance), and VPD (variation of pairwise distances). By analysing mathematical formulae and using simulations, we use this framework to identify metrics that mix dimensions, and we provide a guide to choosing and using the most appropriate metrics. We show that metric choice requires connecting the research question with the correct dimension of the framework and that there are logical approaches to selecting and interpreting metrics. The guide outlined herein will help researchers navigate the current jungle of indices.
Emphasis has been put in recent ecological research on investigating phylogenetic, functional and taxonomic facets of biological diversity. While a flourishing number of indices have been proposed ...for assessing functional diversity, surprisingly few options are available to characterize functional rarity. Functional rarity can play a key role in community and ecosystem dynamics. We introduce here the funrar R package to quantify functional rarity based on species trait differences and species frequencies at local and regional scales. Because of the increasing availability of big datasets in macroecology and biogeography, we optimized funrar to work with large datasets of thousands of species and sites. We illustrate the use of the package to investigate the functional rarity of North and Central American mammals.
In the face of the biodiversity crisis, it is argued that we should prioritize species in order to capture high functional diversity (FD). Because species traits often reflect shared evolutionary ...history, many researchers have assumed that maximizing phylogenetic diversity (PD) should indirectly capture FD, a hypothesis that we name the "phylogenetic gambit". Here, we empirically test this gambit using data on ecologically relevant traits from >15,000 vertebrate species. Specifically, we estimate a measure of surrogacy of PD for FD. We find that maximizing PD results in an average gain of 18% of FD relative to random choice. However, this average gain obscures the fact that in over one-third of the comparisons, maximum PD sets contain less FD than randomly chosen sets of species. These results suggest that, while maximizing PD protection can help to protect FD, it represents a risky conservation strategy.
Niche differences are key to understanding the distribution and structure of biodiversity. To examine niche differences, we must first characterize how species occupy niche space, and two approaches ...are commonly used in the ecological literature. The first uses species traits to estimate multivariate trait space (so-called functional trait diversity, FD); the second quantifies the amount of time or evolutionary history captured by a group of species (phylogenetic diversity, PD). It is often—but controversially—assumed that these putative measures of niche space are at a minimum correlated and perhaps redundant, since more evolutionary time allows for greater accumulation of trait changes. This theoretical expectation remains surprisingly poorly evaluated, particularly in the context of multivariate measures of trait diversity. We evaluated the relationship between phylogenetic diversity and trait diversity using analytical and simulation-based methods across common models of trait evolution. We show that PD correlates with FD increasingly strongly as more traits are included in the FD measure. Our results indicate that phylogenetic diversity can be a useful surrogate for high-dimensional trait diversity, but we also show that the correlation weakens when the underlying process of trait evolution includes variation in rate and optima.
Traits differentially adapt plant species to particular conditions generating compositional shifts along environmental gradients. As a result, community‐scale trait values show concomitant shifts, ...termed trait‒environment relationships. Trait‒environment relationships are often assessed by evaluating community‐weighted mean (CWM) traits observed along environmental gradients. Regression‐based approaches (CWMr) assume that local communities exhibit traits centred at a single optimum value and that traits do not covary meaningfully. Evidence suggests that the shape of trait‒abundance relationships can vary widely along environmental gradients—reflecting complex interactions—and traits are usually interrelated. We used a model that accounts for these factors to explore trait‒environment relationships in herbaceous forest plant communities in Wisconsin (USA).
We built a generalized linear mixed model (GLMM) to analyse how abundances of 185 species distributed among 189 forested sites vary in response to four functional traits (vegetative height—VH, leaf size—LS, leaf mass per area—LMA and leaf carbon content), six environmental variables describing overstorey, soil and climate conditions, and their interactions. The GLMM allowed us to assess the nature and relative strength of the resulting 24 trait‒environment relationships. We also compared results between GLMM and CWMr to explore how conclusions differ between approaches.
The GLMM identified five significant trait‒environment relationships that together explain ~40% of variation in species abundances across sites. Temperature appeared as a key environmental driver, with warmer and more seasonal sites favouring taller plants. Soil texture and temperature seasonality affected LS and LMA; seasonality effects on LS and LMA were nonlinear, declining at more seasonal sites. Although often assumed for CWMr, only some traits under certain conditions had centred optimum trait‒abundance relationships. CWMr more liberally identified (13) trait‒environment relationships as significant but failed to detect the temperature seasonality‒LMA relationship identified by the GLMM.
Synthesis. Although GLMM represents a more methodologically complex approach than CWMr, it identified a reduced set of trait‒environment relationships still capable of accounting for the responses of forest understorey herbs to environmental gradients. It also identified separate effects of mean and seasonal temperature on LMA that appear important in these forests, generating useful insights and supporting broader application of GLMM approach to understand trait‒environment relationships.
Trait‒environment relationships emerge from complex interacting effects of multiple traits and environments on species abundance. A GLMM that modelled these complexities identified a simple set of trait‒environment relationships still capable of accounting for the responses of forest understorey herbs to environmental gradients across Wisconsin.
The β‐null deviation measure, developed as a null model for β‐diversity, is increasingly used in empirical studies to detect the underlying structuring mechanisms in communities (e.g. niche versus ...neutral and stochastic versus deterministic). Despite growing use, the ecological interpretation of the presence/absence and abundance‐based versions of the β‐null diversity measure have not been tested against communities with known assembly mechanisms, and thus have not been validated as an appropriate tool for inferring assembly mechanisms. Using a mechanistic model with known assembly mechanisms, we simulated replicate metacommunities and examined β‐null deviation values 1) across a gradient of niche (species‐sorting) to neutrally structured metacommunities, 2) through time, and 3) we compared the effect of changes in assembly mechanism on the performance of the β‐null deviation measures. The impact of stochasticity on assembly outcomes was also considered. β‐null deviation measures proved to be interpretable as a measure of niche or neutral assembly. However, the presence/absence version of the β‐null deviation measure could not differentiate between niche and neutral metacommunities if demographic stochasticity were present. The abundance‐based β‐null deviation measure was successful in distinguishing between niche and neutral metacommunities and was robust to the presence of stochasticity, changes through time, and changing assembly mechanisms. However, we suggest that it is not robust to changing abundance evenness distributions or sampling of communities, and so its interpretation still requires some care. We encourage the testing of the assumptions behind null models for ecology and care in their application.
Aim: Biogeographical theory and conservation valuation schemes necessarily involve assessing how biodiversity is distributed through space and 'biodiversity' encapsulates many different aspects of ...biological organization and information. While biogeography may try to explain biodiversity patterns, successful conservation strategies should attempt to maximize different aspects of diversity. Ultimately, diversity patterns are the product of evolutionary history, and research and conservation efforts seek to understand the unequal distribution of evolutionary history. For conservation efforts, results have been inconsistent as to whether species richness (SR) provides sufficient surrogacy for evolutionary history. Here, we provide a conceptual framework allowing for the direct comparison of taxonomic richness and phylogenetic diversity (PD), both in terms of their mechanistic relationship and the relationship between their spatial distributions. Location: Global. Methods: We present a framework that relates regional SR, PD, biogeographically weighted evolutionary distinctiveness and biogeographically weighted SR. Further, we use simulations to illustrate how the size of the species pool, topological patterns within the phylogeny and autocorrelation in spatial distributions affect the correlation among metrics. Results: In regions that include both recently diversified groups and ancient species poor lineages, large species pools and low spatial autocorrelation, the correlation between biodiversity measures is lower than regions with low richness, balanced phylogenetic trees and high spatial autocorrelation. Main conclusions: We can now understand and predict when regional richness and PD should be strongly correlated. This congruency is the product of evolutionary and ecological processes that determine species pool membership and community assembly. Further, in regions where SR is not expected to be congruent with phylogenetic distinctiveness, re-examining how existing reserve networks protect the multiple aspects of biodiversity is critically important.
ABSTRACT
It is often claimed that conserving evolutionary history is more efficient than species‐based approaches for capturing the attributes of biodiversity that benefit people. This claim ...underpins academic analyses and recommendations about the distribution and prioritization of species and areas for conservation, but evolutionary history is rarely considered in practical conservation activities. One impediment to implementation is that arguments related to the human‐centric benefits of evolutionary history are often vague and the underlying mechanisms poorly explored. Herein we identify the arguments linking the prioritization of evolutionary history with benefits to people, and for each we explicate the purported mechanism, and evaluate its theoretical and empirical support. We find that, even after 25 years of academic research, the strength of evidence linking evolutionary history to human benefits is still fragile.
Most – but not all – arguments rely on the assumption that evolutionary history is a useful surrogate for phenotypic diversity. This surrogacy relationship in turn underlies additional arguments, particularly that, by capturing more phenotypic diversity, evolutionary history will preserve greater ecosystem functioning, capture more of the natural variety that humans prefer, and allow the maintenance of future benefits to humans. A surrogate relationship between evolutionary history and phenotypic diversity appears reasonable given theoretical and empirical results, but the strength of this relationship varies greatly. To the extent that evolutionary history captures unmeasured phenotypic diversity, maximizing the representation of evolutionary history should capture variation in species characteristics that are otherwise unknown, supporting some of the existing arguments. However, there is great variation in the strength and availability of evidence for benefits associated with protecting phenotypic diversity. There are many studies finding positive biodiversity–ecosystem functioning relationships, but little work exists on the maintenance of future benefits or the degree to which humans prefer sets of species with high phenotypic diversity or evolutionary history. Although several arguments link the protection of evolutionary history directly with the reduction of extinction rates, and with the production of relatively greater future biodiversity via increased adaptation or diversification, there are few direct tests. Several of these putative benefits have mismatches between the relevant spatial scales for conservation actions and the spatial scales at which benefits to humans are realized. It will be important for future work to fill in some of these gaps through direct tests of the arguments we define here.
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