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The Merging of Community Ecology and Phylogenetic Biology

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The Merging of Community Ecology and Phylogenetic Biology Ecology Letters, (2009) 12: 693–715 doi: 10.1111/j.1461-0248.2009.01314.x REVIEW AND SYNTHESIS The merging of community ecology and phylogenetic biology Abstract Jeannine Cavender-Bares,1* The increasing availability of phylogenetic data, computing power and informatics tools Kenneth H. Kozak,2 Paul V. A. has facilitated a rapid expansion of studies that apply phylogenetic data and methods to Fine3 and Steven W. Kembel3† community ecology. Several key areas are reviewed in which phylogenetic information 1 Department of Ecology, helps to resolve long-standing controversies in community ecology, challenges previous Evolution and Behavior, assumptions, and opens new areas of investigation. In particular, studies in phylogenetic University of Minnesota, St. community ecology have helped to reveal the multitude of processes driving community Paul, MN 55108, USA assembly and have demonstrated the importance of evolution in the assembly process. 2Bell Museum of Natural Phylogenetic approaches have also increased understanding of the consequences of History, and Department of Fisheries, Wildlife, and community interactions for speciation, adaptation and extinction. Finally, phylogenetic Conservation Biology, University community structure and composition holds promise for predicting ecosystem processes of Minnesota, St. Paul, MN, and impacts of global change. Major challenges to advancing these areas remain. In 55108, USA particular, determining the extent to which ecologically relevant traits are phylogeneti- 3Department of Integrative cally conserved or convergent, and over what temporal scale, is critical to understanding Biology, University of California, the causes of community phylogenetic structure and its evolutionary and ecosystem Berkeley, CA 94720, USA consequences. Harnessing phylogenetic information to understand and forecast changes † Present address: Center for in diversity and dynamics of communities is a critical step in managing and restoring the Ecology and Evolutionary EarthÕs biota in a time of rapid global change. Biology, University of Oregon, Eugene, OR 97403, USA. Keywords *Correspondence: E-mail: Community assembly, deterministic vs. neutral processes, ecosystem processes, [email protected] experimental approaches, functional traits, phylogenetic community ecology, phylo- genetic diversity, spatial and phylogenetic scale. Ecology Letters (2009) 12: 693–715 Fine et al. 2006; Strauss et al. 2006; Davies et al. 2007; INTRODUCTION Vamosi et al. 2008). Such approaches now allow community Community ecology investigates the nature of organismal ecologists to link short-term local processes to continental interactions, their origins, and their ecological and evolu- and global processes that occur over deep evolutionary time tionary consequences. Community dynamics form the link scales (Losos 1996; Ackerly 2003; Ricklefs 2004; Pennington between uniquely evolved species and ecosystem functions et al. 2006; Mittelbach et al. 2007; Swenson et al. 2007; that affect global processes. In the face of habitat Donoghue 2008; Emerson & Gillespie 2008; Graham & destruction worldwide, understanding how communities Fine 2008). This effort has been facilitated by the rapid rise assemble and the forces that influence their dynamics, in phylogenetic information, computing power and compu- diversity and ecosystem function will prove critical to tational tools. Our goal here is to review how phylogenetic managing and restoring the EarthÕs biota. Consequently, the information contributes to community ecology in terms of study of communities is of paramount importance in the the long-standing questions it helps answer, the assumptions 21st century. it challenges and the new questions it invites. In particular, Recently, there has been a rapidly increasing effort to we focus on the insights gained from applying phylogenetic bring information about the evolutionary history and approaches to explore the ecological and evolutionary genealogical relationships of species to bear on questions factors that underlie the assembly of communities, and of community assembly and diversity (e.g. Webb et al. 2002; how the interactions among species within them ultimately Ackerly 2004; Cavender-Bares et al. 2004a; Gillespie 2004; influence evolutionary and ecosystem processes. Ó 2009 Blackwell Publishing Ltd/CNRS 694 J. Cavender-Bares et al. Review and Synthesis There are three perspectives on the dominant factors both phylogenetic and spatial scale in the interpretation of that influence community assembly, composition and ecological and evolutionary patterns (Box 1, Figs 1 and 2) diversity. First is the classic perspective that communities and cognizance of the multiplicity of processes that assemble according to niche-related processes, following underlie patterns. Observational, experimental and theo- fundamental ÔrulesÕ dictated by local environmental retical studies aimed at deciphering the mechanisms filters and the principle of competitive exclusion (e.g. involved in community assembly and how they shift with Diamond 1975; Tilman 1982; Bazzaz 1991; Weiher & scale are paving the way for phylogenetic approaches to Keddy 1999). An alternative perspective is that commu- large-scale prediction of ecosystem dynamics in response nity assembly is largely a neutral process in which species to global change. are ecologically equivalent (e.g. Hubbell 2001). A third We first discuss the historical origins of the classic perspective emphasizes the role of historical factors in debates in community ecology that phylogenetics helps to dictating how communities assemble (Ricklefs 1987; address. We then turn to specific examples in the general Ricklefs & Schluter 1993). In the latter view, the starting areas highlighted above and review contributions made conditions and historical patterns of speciation and possible by integrating community ecology and phyloge- dispersal matter more than local processes. The relative netic biology. In doing so, we discuss the challenges influence of niche-related, neutral and historical processes involved in further progress. We close with a summary of is at the core of current debates on the assembly of the major advances, challenges and prospects for the communities and the coexistence of species (Hubbell emerging field of phylogenetic community ecology. We 2001; Chase & Leibold 2003; Fargione et al. 2004; include illustrative examples from animals, plants and Ricklefs 2004; Tilman 2004). This debate falls within other organisms in discussing the contributions of the larger historic controversy about the nature of phylogenetic information to understanding community communities and the extent to which they represent assembly and the feedbacks to evolutionary processes. associations of tightly interconnected species shaped over However, we focus largely on the plant literature in long periods of interaction or are the result of chance co- discussing the ecosystem and global consequences of occurrences of individually dispersed and distributed community assembly, reflecting the plant orientation of organisms (Clements 1916; Gleason 1926; Davis 1981; much of the relevant literature. Brooks & McLennan 1991; Callaway 1997; DiMichele et al. 2004; Ricklefs 2008). Here we review how the merging of community HISTORICAL OVERVIEW ecology and phylogenetic biology advances these debates Niche-related processes and assembly rules and allows new areas of enquiry to be addressed. First, phylogenetics helps to resolve the long-standing contro- Early ecologists, including Darwin, recognized that specific versy about the relative roles of neutral vs. niche-related attributes of species could influence their interactions with processes in community assembly and facilitates identifi- other species and with the environment in predictable ways. cation of the kinds of processes that underlie community In particular, Darwin noted a paradox inherent in assembly. Second, insights from phylogenetic approaches phenotypic similarity of species with shared ancestry. On present strong challenges to the classical idea that the the one hand, if closely related species are ecologically species pool (and the traits of species within it) is static similar, they should share similar environmental on the time scale over which communities are assembled. requirements and may thus be expected to co-occur. On These approaches are also beginning to demonstrate that the other hand, closely related species should experience community interactions might strongly influence how the strong competitive interactions due to their ecological pool itself evolves and changes across space and time. similarity, thereby limiting coexistence and thus driving Finally, phylogenetic diversity and composition is relevant selection for divergent traits. to predicting ecosystem properties that impact global The idea that similar phenotypes should share habitat processes. affinities was championed by the Danish plant ecologist, We argue that ongoing efforts to integrate knowledge Eugenius Warming (1895), who emphasized differences in of phylogenetic relationships of organisms with their the physiological abilities of plants to adjust to some functional attributes will enhance understanding of the environments but not others. The core idea was that similar distribution and function of the EarthÕs biota at multiple physiological attributes would be selected for by similar scales, increasing our ability to predict outcomes of environments in different regions and that plant pheno- species interactions
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