Ecosystems worldwide are rapidly losing taxonomic, phylogenetic, genetic, and functional diversity as a result of human appropriation of natural resources, modification of habitats and climate, and the spread of pathogenic, exotic, and domestic plants and animals. Twenty years of intense theoretical and empirical research have shown that such biotic impoverishment can markedly alter the biogeochemical and dynamic properties of ecosystems, but frontiers remain in linking this research to the complexity of wild nature, and in applying it to pressing environmental issues such as food, water, energy, and biosecurity. The question before us is whether these advances can take us beyond merely invoking the precautionary principle of conserving biodiversity to a predictive science that informs practical and specific solutions to mitigate and adapt to its loss.

https://science.sciencemag.org/content/sci/336/6087/1401.full.pdf

The Functions of Biological Diversity in an Age of Extinction

The decomposition of dead organic matter is a major determinant of carbon and nutrient cycling in ecosystems, and of carbon fluxes between the biosphere and the atmosphere. Decomposition is driven by a vast diversity of organisms that are structured in complex food webs. Identifying the mechanisms underlying the effects of biodiversity on decomposition is critical given the rapid loss of species worldwide and the effects of this loss on human well-being. Yet despite comprehensive syntheses of studies on how biodiversity affects litter decomposition, key questions remain, including when, where and how biodiversity has a role and whether general patterns and mechanisms occur across ecosystems and different functional types of organism. Here, in field experiments across five terrestrial and aquatic locations, ranging from the subarctic to the tropics, we show that reducing the functional diversity of decomposer organisms and plant litter types slowed the cycling of litter carbon and nitrogen. Moreover, we found evidence of nitrogen transfer from the litter of nitrogen-fixing plants to that of rapidly decomposing plants, but not between other plant functional types, highlighting that specific interactions in litter mixtures control carbon and nitrogen cycling during decomposition. The emergence of this general mechanism and the coherence of patterns across contrasting terrestrial and aquatic ecosystems suggest that biodiversity loss has consistent consequences for litter decomposition and the cycling of major elements on broad spatial scales.

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Consequences of biodiversity loss for litter decomposition across biomes

Hill numbers or the effective number of species are increasingly used to quantify species diversity of an assemblage. Hill numbers were recently extended to phylogenetic diversity, which incorporates species evolutionary history, as well as to functional diversity, which considers the differences among species traits. We review these extensions and integrate them into a framework of attribute diversity (the effective number of entities or total attribute value) based on Hill numbers of taxonomic entities (species), phylogenetic entities (branches of unit-length), or functional entities (species-pairs with unit-distance between species). This framework unifies ecologists' measures of species diversity, phylogenetic diversity, and distance-based functional diversity. It also provides a unified method of decomposing these diversities and constructing normalized taxonomic, phylogenetic, and functional similarity and differentiation measures, including N-assemblage phylogenetic or functional generalizations of...

Unifying Species Diversity, Phylogenetic Diversity, Functional Diversity, and Related Similarity and Differentiation Measures Through Hill Numbers

Understanding, modeling, and predicting the impact of global change on ecosystem functioning across biogeographical gradients can benefit from enhanced capacity to represent biota as a continuous distribution of traits. However, this is a challenge for the field of biogeography historically grounded on the species concept. Here we focus on the newly emergent field of functional biogeography: the study of the geographic distribution of trait diversity across organizational levels. We show how functional biogeography bridges species-based biogeography and earth science to provide ideas and tools to help explain gradients in multifaceted diversity (including species, functional, and phylogenetic diversities), predict ecosystem functioning and services worldwide, and infuse regional and global conservation programs with a functional basis. Although much recent progress has been made possible because of the rising of multiple data streams, new developments in ecoinformatics, and new methodological advances, future directions should provide a theoretical and comprehensive framework for the scaling of biotic interactions across trophic levels and its ecological implications.

https://www.pnas.org/content/pnas/111/38/13690.full.pdf

The emergence and promise of functional biogeography

Classifying the biological traits of organisms can test conceptual frameworks of life-history strategies and allow for predictions of how different species may respond to environmental disturbances. We apply a trait-based classification approach to a complex and threatened group of species, scleractinian corals. Using hierarchical clustering and random forests analyses, we identify up to four life-history strategies that appear globally consistent across 143 species of reef corals: competitive, weedy, stress-tolerant and generalist taxa, which are primarily separated by colony morphology, growth rate and reproductive mode. Documented shifts towards stress-tolerant, generalist and weedy species in coral reef communities are consistent with the expected responses of these life-history strategies. Our quantitative trait-based approach to classifying life-history strategies is objective, applicable to any taxa and a powerful tool that can be used to evaluate theories of community ecology and predict the impact of environmental and anthropogenic stressors on species assemblages.

Evaluating life‐history strategies of reef corals from species traits

Summary 1. The goal of conservation and restoration activities is to maintain biological diversity and the ecosystem services that this diversity provides. These activities traditionally focus on the measures of species diversity that include only information on the presence and abundance of species. Yet how diversity influences ecosystem function depends on the traits and niches filled by species. 2. Biological diversity can be quantified in ways that account for functional and phenotypic differences. A number of such measures of functional diversity (FD) have been created, quantifying the distribution of traits in a community or the relative magnitude of species similarities and differences. We review FD measures and why they are intuitively useful for understanding ecological patterns and are important for management. 3. In order for FD to be meaningful and worth measuring, it must be correlated with ecosystem function, and it should provide information above and beyond what species richness or diversity can explain. We review these two propositions, examining whether the strength of the correlation between FD and species richness varies across differing environmental gradients and whether FD offers greater explanatory power of ecosystem function than species richness. 4. Previous research shows that the relationship between FD and richness is complex and context dependent. Different functional traits can show individual responses to different gradients, meaning that important changes in diversity can occur with minimal change in richness. Further, FD can explain variation in ecosystem function even when richness does not. 5. Synthesis and applications. FD measures those aspects of diversity that potentially affect community assembly and function. Given this explanatory power, FD should be incorporated into conservation and restoration decision-making, especially for those efforts attempting to reconstruct or preserve healthy, functioning ecosystems.

/pdf/beyond-species-functional-diversity-and-the-maintenance-of-1qhn1coswt.pdf

Beyond species: functional diversity and the maintenance of ecological processes and services

For a mutualism to remain evolutionarily stable, theory predicts that mutualists should limit their associations to high-quality partners. However, most mutualists either simultaneously or sequentially associate with multiple partners that confer the same type of reward. By viewing mutualisms through the lens of niche breadth evolution, we outline how the environment shapes partner availability and relative quality, and ultimately a focal mutualist's partner breadth. We argue that mutualists that associate with multiple partners may have a selective advantage compared to specialists for many reasons, including sampling, complementarity, and portfolio effects, as well as the possibility that broad partner breadth increases breadth along other niche axes. Furthermore, selection for narrow partner breadth is unlikely to be strong when the environment erodes variation in partner quality, reduces the costs of interacting with low-quality partners, spatially structures partner communities, or decreases the strength of mutualism. Thus, we should not be surprised that most mutualists have broad partner breadth, even if it allows for ineffective partners to persist.

Using niche breadth theory to explain generalization in mutualisms

The ecological niche is a multi-dimensional concept including aspects of resource use, environmental tolerance, and interspecific interactions, and the degree to which niches overlap is central to many ecological questions. Plant phenotypic traits are increasingly used as surrogates of species niches, but we lack an understanding of how key sampling decisions affect our ability to capture phenotypic differences among species. Using trait data of ecologically distinct monkeyflower (Mimulus) congeners, we employed linear discriminant analysis to determine how (1) dimensionality (the number and type of traits) and (2) variation within species influence how well measured traits reflect phenotypic differences among species. We conducted analyses using vegetative and floral traits in different combinations of up to 13 traits and compared the performance of commonly used functional traits such as specific leaf area against other morphological traits. We tested the importance of intraspecific variation by assessing how population choice changed our ability to discriminate species. Neither using key functional traits nor sampling across plant functions and organs maximized species discrimination. When using few traits, vegetative traits performed better than combinations of vegetative and floral traits or floral traits alone. Overall, including more traits increased our ability to detect phenotypic differences among species. Population choice and the number of traits used had comparable impacts on discriminating species. We addressed methodological challenges that have undermined cross-study comparability of trait-based approaches. Our results emphasize the importance of sampling among-population trait variation and suggest that a high-dimensional approach may best capture phenotypic variation among species with distinct niches.

Trait dimensionality and population choice alter estimates of phenotypic dissimilarity.

Understanding the origins and impacts of novel traits has been a perennial interest in many realms of ecology and evolutionary biology. Here, we build on previous evolutionary and philosophical treatments of this subject to encompass novelties across biological scales and eco-evolutionary perspectives. By defining novelties as new features at one biological scale that have emergent effects at other biological scales, we incorporate many forms of novelty that have previously been treated in isolation (such as novelty from genetic mutations, new developmental pathways, new morphological features, and new species). Our perspective is based on the fundamental idea that the emergence of a novelty, at any biological scale, depends on its environmental and genetic context. Through this lens, we outline a broad array of generative mechanisms underlying novelty and highlight how genomic tools are transforming our understanding of the origins of novelty. Lastly, we present several case studies to illustrate how novelties across biological scales and systems can be understood based on common mechanisms of change and their environmental and genetic contexts. Specifically, we highlight how gene duplication contributes to the evolution of new complex structures in visual systems; how genetic exchange in symbiosis alters functions of both host and symbiont, resulting in a novel organism; and how hybridisation between species can generate new species with new niches.

/pdf/origins-and-evolution-of-biological-novelty-1gr5cwpr.pdf

Origins and evolution of biological novelty.

The contrasting and idiosyncratic changes in biodiversity that have been documented across urbanization gradients call for a more mechanistic understanding of urban community assembly. The reproductive success of organisms in cities should underpin their population persistence and the maintenance of biodiversity in urban landscapes. We propose that exploring individual-level reproductive traits and environmental drivers of reproductive success could provide the necessary links between environmental conditions, offspring production, and biodiversity in urban areas. For 3 years, we studied cavity-nesting solitary bees and wasps in four urban green space types across Toronto, Canada. We measured three reproductive traits of each nest: the total number of brood cells, the proportion of parasite-free cells, and the proportion of non-emerged brood cells that were parasite-free. We determined (a) how reproductive traits, trait diversity and offspring production respond to multiple environmental variables and (b) how well reproductive trait variation explains the offspring production of single nests, by reflecting the different ways organisms navigate trade-offs between gathering of resources and exposure to parasites. Our results showed that environmental variables were poor predictors of mean reproductive trait values, trait diversity, and offspring production. However, offspring production was highly positively correlated with reproductive trait evenness and negatively correlated with trait richness and divergence. This suggests that a narrow range of reproductive traits are optimal for reproduction, and the even distribution of individual reproductive traits across those optimal phenotypes is consistent with the idea that selection could favor diverse reproductive strategies to reduce competition. This study is novel in its exploration of individual-level reproductive traits and its consideration of multiple axes of urbanization. Reproductive trait variation did not follow previously reported biodiversity-urbanization patterns; the insensitivity to urbanization gradients raise questions about the role of the spatial mosaic of habitats in cities and the disconnections between different metrics of biodiversity.

https://onlinelibrary.wiley.com/doi/pdf/10.1002/ece3.7537

Kelly A. Carscadden

Papers

Beyond species: functional diversity and the maintenance of ecological processes and services

Using niche breadth theory to explain generalization in mutualisms

Trait dimensionality and population choice alter estimates of phenotypic dissimilarity.

Origins and evolution of biological novelty.

Reproductive trait differences drive offspring production in urban cavity-nesting bees and wasps.