Perspective: Functional Biodiversity during the Anthropocene
Andrea Belgrano, Ute Jacob, Charles Fowler, and Guy Woodward
We are living through a new temporal epoch, recently described as the Anthropocene (
Latour, 2014), in which human activities are increasingly shaping the biota within and among Earth's ecosystems. These anthropogenic forces bring a variety of major consequences, including the significant loss of biodiversity in all its forms, and not just the more familiar measure of species richness (
Mora et al., 2011;
Cardinale, 2013). Biodiversity, in the broader sense of the term that we use here, includes patterns in the links between species assemblages and their functional organization within a web of interactions under environmental constraints. This shift of focus from the traditional emphasis on the “nodes” to the “links” represents an important philosophical change: it forces us to recognize that multispecies systems are not simply passively mapped onto an environmental template, but that their own internal dynamics influence higher level phenomena. Biodiversity is linked to ecosystem functioning and, by extension, to the socioeconomically valuable services they provide (
Millennium Ecosystem Assessment (MA), 2005), yet the strength and form of these relationships are still surprisingly poorly understood. This emphasizes the urgent need to develop strategies that promote sustainable use of ecosystems at local scales, and of the biosphere at a global scale, in order to preserve a “safe operating space for humanity” (
Naeem et al., 2012;
Perrings et al., 2010;
Dobson, 2009;
Mace et al., 2015).
If we are to do this, first we need to understand how structure and functioning are connected. Functional diversity (FD) is a key component of biodiversity (
Perrings et al., 2010;
Mouillot et al., 2014) and provides a direct link between biodiversity and ecosystem processes (
Naeem, 2006), yet comparisons of FD across systems has largely ignored perspectives that combine ecological and evolutionary principles and understanding (
Perrings et al., 2010). It is imperative that a more comprehensive approach is developed for effectively conserving species and their functional roles that considers both ecological and evolutionary principles (
Stouffer et al., 2012).
Macroecological patterns in the FD can be manifested across a wide range of taxa and systems, as well as across spatial and temporal scales, and organizational levels. This is often achieved by focusing on the explicit links between biodiversity and ecosystem functioning in experimental manipulations, a field that has gained huge traction in the past couple of decades (
Reiss et al., 2009). The (usually positive) relationship between species richness and FD has profound implications for conservation and management (especially of our use of natural resources) and is also increasingly being linked to food web structure and ecosystem services. Combining macroecological studies of system properties with detailed analyses of community functional diversity patterns and food web structure (
Petchey and Gaston, 2007) could provide one means of understanding which ecosystem services (
Dobson, 2009) are particularly important for sustaining overall healthy environmental status, and also which might be most vulnerable to perturbations. This is only one of the many challenges that we need to resolve and understand (
Figure 1), if we are to move toward a functionally based framework that can include the ecological and evolutionary roles that species (including humans) play within the local ecosystems on a global scale.
Figure 1 Challenges and questions at the time of the Anthropocene: linking functional diversity, food web structure, and ecosystem services.
The biodiversity of ecosystems worldwide is undergoing major change, via human-induced extinctions set against gains via intentional and accidental introductions, with the balance increasingly on the red side of the ledger. These combined effects are altering the structure and function of ecosystems on a global scale, and we are now experiencing what has been widely described as the 6th Great Extinction. An especially challenging task is to understand how individual species combine to deliver ecosystem services in a changing world, yet a rigorous, systematic methodology for doing so still eludes us. This is, at least partially, because the dynamics of ecosystem services remain poorly characterized at local-to-regional scales, and the role of species in contributing to services is not fully understood. A priori, we know that functional traits, food web structure, and ecosystem services are interlinked and need to be considered in conjunction with the recognition that ecosystems change and require adaptive governance to ensure their long-term sustainability (
Folke et al., 2005;
Folke, 2007;
Dietz et al., 2003).
Recently there has been an explosion of interest in employing network theory to disentangle and explore the relationships between biodiversity and ecosystem functioning (
Reiss et al., 2009;
Kéfi et al., 2015).
This volume Aquatic Functional Biodiversity: An Ecological and Evolutionary Approach is an attempt by some of the most prominent investigators in the field to provide a more general conceptual framework that can include new ecological and/or evolutionary approaches to the understanding of functional diversity. This is accomplished in a way that holds the promise of leading to effective and realistic conservation, especially in doing our best at ensuring the sustainability of ecosystem services in aquatic systems. The collection of chapters in this book represents a substantive contribution to: (1) defining common ground in terms of terminology and conceptual issues, (2) connecting conceptual frameworks from the ecological and/or evolutionary sciences with those from classical biodiversity theory, to make progress toward better practical application, and (3) providing examples of how biodiversity and ecosystem services might be conserved more effectively in the real world.
Terminology and Conceptual Issues in Ecological and Evolutionary Perspectives
Understanding and predicting interactions between ecological and evolutionary processes are extremely challenging tasks. Pawar and colleagues (
Chapter 1) provide a summary of recent theoretical and empirical advances for developing a mechanistic understanding of trophic interactions, and identify key methods and challenges for understanding and predicting the eco-evolutionary dynamics of aquatic ecosystems. These authors introduce a theoretical approach to understanding how the metabolic and biomechanical bases of trophic interactions can enhance general predictions regarding the eco-evolutionary dynamics and functioning of aquatic ecosystems. Their ideas will be applicable to other types of ecological interactions that involve metabolism and biomechanics (e.g., pollination, parasitism, and competitive interactions). Numerous relationships in which biodiversity contributes positively to ecosystem function have been identified across a wide variety of ecological communities in aquatic and terrestrial ecosystems. Even though these relationships are acknowledged in multiple ways, the underlying mechanisms are poorly understood and are often questioned. In their chapter, Vasseur and Messinger (
Chapter 3) introduce and use two models that depict autotrophic and heterotrophic competitors. Their work includes a case study involving a series of experiments using in silico biodiversity and ecosystem function. They show that the “ancestry” of species, which is defined in terms of whether or not they have coevolved in the presence of competitors or in monocultures, is an important determinant in the extent to which transgressive overyielding can occur. Their research has important implications for the interpretation of previous meta-analyses, and may provide insight of importance to the management of human influence on aquatic communities and their recovery from abnormal human influence.
Borthagaray and coworkers (
Chapter 4) present the state of the art of effects of metacommunity networks on local community structures; their chapter covers theoretical predictions and empirical evaluations. By using metapopulation models they point out the wide range of patterns predicted by different mechanisms. They stress that relative effects of dispersal on dominant and subordinate species determine the weakening or strengthening of patch dynamics important to fully understanding community structure.
Conceptual Frameworks in Ecological and Evolutionary Sciences
In
Chapter 6, Woodward and Perkins focus on the impacts and consequences of different drivers of change on freshwater ecosystems, which are particularly vulnerable to environmental stressors in general, and climate change in particular. A review of the current state of these relationships is presented—as a “jigsaw puzzle of our understanding of ecological and evolutionary responses to climate change.” Freshwater ecology has a long history and is arguably much better understood than that of many marine systems: many of the biotic and abiotic constraints (i.e., chemical or physical properties) on species coexistence and food webs are now relatively well established. Although knowledge about how individuals and ecosystem processes are likely to respond to temperature change has improved over the years, the understanding of the community-level...