Ecology Community Ecology
by
Herman A. Verhoef
  • LAST MODIFIED: 23 May 2012
  • DOI: 10.1093/obo/9780199830060-0042

Introduction

At the beginning of the 20th century there was much debate about the “nature” of communities. The driving question was whether the community was a self-organized system of co-occurring species or simply a haphazard collection of populations with minimal functional integration. At that time, two extreme views dominated the discussion: one view considered a community as a superorganism, the member species of which were tightly bound together by interactions that contributed to repeatable patterns of species abundance in space and time. This concept led to the assumption that communities are fundamental entities, to be classified as the Linnaean taxonomy of species. Frederick E. Clements was one of the leading proponents of this approach, and his view became known as the organismic concept of communities. This assumes a common evolutionary history for the integrated species. The opposite view was the individualistic continuum concept, advocated by H. A. Gleason. His focus was on the traits of individual species that allow each to live within specific habitats or geographical ranges. In this view a community is an assemblage of populations of different species whose traits allow persisting in a prescribed area. The spatial boundaries are not sharp, and the species composition can change considerably. Consequently, it was discussed whether ecological communities were sufficiently coherent entities to be considered appropriate study objects. Later, consensus was reached: that properties of communities are of central interest in ecology, regardless of their integrity and coherence. From the 1950s and 1960s onward, the discussion was dominated by the deterministic outcome of local interactions between species and their environments and the building of this into models of communities. This approach, indicated as “traditional community ecology,” led to a morass of theoretical models, without being able to provide general principles about many-species communities. Early-21st-century approaches to bringing general patterns into community ecology concern (1) the metacommunity approach, (2) the functional trait approach, (3) evolutionary community ecology, and (4) the four fundamental processes. The metacommunity approach implicitly recognizes and studies the important role of spatiotemporal dynamics. In the functional trait approach, four themes are focused upon: traits, environmental gradients, the interaction milieu, and performance currencies. This functional, trait-focused approach should have a better prospect of understanding the effects of global changes. Evolutionary community ecology is an approach in which the combination of community ecology and evolutionary biology will lead to a better understanding of the complexity of communities and populations. The four fundamental processes are selection, drift, speciation, and dispersal. This approach concerns an organizational scheme for community ecology, based on these four processes to describe all existing specific models and frameworks, in order to make general statements about process–pattern connections.

General Overviews

The now-classic discussion on the nature of communities dates from the early 1920s, the two opponents of which have already been mentioned in the Introduction: F. E. Clements with his organismic concept (Clements 1916) and H. A. Gleason with his individualistic continuum concept (Gleason 1926). Especially in the former concept, the community was thought to live as a superorganism, as a fundamental unit, in a specific area. As “the most general definition one can give,” C. J. Krebs (Krebs 1972) describes a community in his classic textbook as “an assemblage of populations of living organisms in a prescribed area or habitat.” In that same year, MacArthur 1972, a book on geographical ecology, stated that “the goal of community ecology is to find general rules.” It is this quest to find general rules in community ecology that has caused much debate in the field of (theoretical) community ecology until the present. From the 1960s onward the focus within community ecology was on the study of the population dynamics of pairs of species and the building of models. Important publications concerning this subject are May 2001 and Tilman 1976. These publications were very important to explain one- to few-species systems but hardly provided general rules for communities existing of many species, as is mentioned in Putman 1994, which states that “. . . it is difficult to keep pace and retain a comprehensive overview of the entire discipline.” More recently, approaches have been presented that try to handle the current explosion of massive, high-resolution data based on individual-level sampling and satellite images, which attempt to bring this together into general patterns in community ecology. The following four books should also be mentioned in this section: Macroecology (Brown 1995, cited under Historical Background), Ecological Niches (Chase and Leibold 2003), Community Ecology (Verhoef and Morin 2010, cited under Applications), and Community Ecology (Morin 2011). Brown 1995 integrates data from ecology, systematics, evolutionary biology, paleobiology, and biogeography to investigate problems, providing a rich, more complete understanding of how patterns of life have moved across the earth over time. It also demonstrates the advantages of macroecology for conservation, showing how it allows scientists to look beyond endangered species and ecological communities to consider the long history and large geographic scale of human impacts. In Chase and Leibold 2003, questions such as “Why do species live where they live?”; “What determines the abundance and diversity of species in a given area?”; and “What role do species play in the functioning of entire ecosystems?” share a single core concept: the ecological niche. The authors define the niche as including both what an organism needs from its environment and how that organism’s activities shape its environment. Verhoef and Morin 2010 provides a survey of the state of the art in theory and applications of community ecology, with special attention to topology, dynamics, and the importance of spatial and temporal scale, as well as applications to emerging problems in human-dominated ecosystems, including the restoration and reconstruction of viable communities (see Applications). Morin 2011 gives an introduction to a coverage of concepts and theories central to community ecology, using examples drawn from terrestrial, freshwater, and marine systems, and focusing on animal, plant, and microbial species. Throughout, there is an emphasis on the crucial interplay among observations, experiments, and mathematical models.

  • Chase, J. M., and M. A. Leibold. 2003. Ecological niches: Linking classical and contemporary approaches. Interspecific Interactions. Chicago: Univ. of Chicago Press.

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    Although recently the niche concept has had a negative connotation among ecologists, Chase and Leibold argue that the niche is an ideal tool for unifying disparate research and theoretical approaches in contemporary ecology. Their niche concept is flexible enough to include a variety of small- and large-scale processes, from resource competition, predation, and stress to community structure, biodiversity, and ecosystem function.

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    • Clements, F. E. 1916. Plant succession: An analysis of the development of vegetation. Carnegie Institution of Washington Publication 242. Washington, DC: Carnegie Institution of Washington.

      DOI: 10.5962/bhl.title.56234E-mail Citation »

      Clements’s magnus opus, in which he conceived of the community as a superorganism whose member species are tightly bound together both now and in their common evolutionary history. His ideas are out of date but are referred to in a historical context.

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      • Gleason, H. A. 1926. The individualistic concept of the plant association. Bulletin of the Torrey Botanical Club 53.1: 7–26.

        DOI: 10.2307/2479933E-mail Citation »

        In this individualistic concept, which contrasts with Clements 1916, Gleason saw the relationship of coexisting species as simply the result of similarities in their requirements and tolerances. Though the current view is close to the individualistic concept, this article is mentioned only in a historical context.

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        • Krebs, C. J. 1972. Ecology: The experimental analysis of distribution and abundance. New York: Harper and Row.

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          A much-used textbook on ecology during the 1970s. As mentioned by the author, “This book is not an encyclopedia of ecology, but an introduction to its problems.” It approaches ecology in a general way, and not as the ecology per biotope.

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          • MacArthur, R. H. 1972. Geographical ecology: Patterns in the distribution of species. New York: Harper & Row.

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            This book forms the summation of the life work of one of the most influential scientists of our time. According to MacArthur the structure of the environment, the morphology of the species, the economics of species behavior, and the dynamics of population changes are the four essential ingredients of biogeographic patterns.

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            • May, R. M. 2001. Stability and complexity in model ecosystems. 2d ed. Princeton Landmarks in Biology. Princeton, NJ: Princeton Univ. Press.

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              Originally published in 1973. Robert May addressed in this classic book the following questions: What makes populations stabilize? What makes them fluctuate? Are populations in complex ecosystems more stable than populations in simple ecosystems? This book played a key role in introducing nonlinear mathematical models and the study of deterministic chaos into ecology.

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              • Morin, P. J. 2011. Community ecology. 2d ed. Oxford: Wiley-Blackwell.

                DOI: 10.1002/9781444341966E-mail Citation »

                This second, updated edition gives an introduction to a balanced coverage of concepts and theories central to community ecology, using examples drawn from terrestrial, freshwater, and marine systems, and focusing on animal, plant, and microbial species. It offers a valuable resource for advanced undergraduates and graduate students.

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                • Putman, R. J. 1994. Community ecology. London and New York: Chapman & Hall.

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                  This is a typical, standard textbook for advanced undergraduates in ecology, with a chapter division following the basic elements of community ecology (definitions, population interactions, food webs, niche theory, etc.).

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                  • Tilman, D. 1976. Ecological competition between algae: Experimental confirmation of resource-based competition theory. Science 192.4238: 463–465.

                    DOI: 10.1126/science.192.4238.463E-mail Citation »

                    In laboratory studies of two species of freshwater diatoms potentially limited by phosphate and silicate, all possible outcomes of ecological competition, including stable coexistence, were found, with the relative abundance of these nutrients determining the outcome of competition. This important article is recurring in many textbooks in which examples are given of competition between two species.

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                    Journals

                    Most ecological journals publish articles that include the topic of community ecology, including Trends in Ecology & Evolution, Ecology, Journal of Ecology, Oikos, Functional Ecology, and The American Naturalist. However, opinion journals, also used for this bibliography, are included in this section, such as The Quarterly Review of Biology and Proceedings of the National Academy of Sciences.

                    • The American Naturalist.

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                      First published in 1867, The American Naturalist is the oldest scientific journal in the world, dedicated to the study of ecology, evolution, and behavior. It addresses topics in community and ecosystem dynamics, evolution of sex and mating systems, organismal adaptation, and genetic aspects of evolution.

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                      • Ecology.

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                        Established in 1920, Ecology publishes research and synthesis papers on all aspects of ecology, with particular emphasis on papers that develop new concepts in ecology that test ecological theory, or that lead to an increased appreciation of the diversity of ecological phenomena. Preference is given to research and synthesis that lead to generalizations potentially applicable to other species, populations, communities, or ecosystems.

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                        • Ecology Letters.

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                          Ecology Letters is a forum for the very rapid publication of the most novel research in ecology. Manuscripts relating to the ecology of all taxa, in any biome and geographic area, will be considered, and priority will be given to those papers exploring or testing clearly stated hypotheses.

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                          • Functional Ecology.

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                            Functional Ecology publishes high-impact papers on organismal ecology, including physiological, behavioral, and evolutionary ecology. Further, it publishes organism-level studies that have clear and important implications for community and ecosystem processes. As a more recent development, it has been including studies on behavioral, physiological, or life history traits at the genomic level.

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                            • Journal of Ecology.

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                              Journal of Ecology publishes both experimental and theoretical papers on all aspects of plant-related ecology, covering terrestrial and aquatic ecosystems; studies of plant communities, populations, or individuals; and interactions between plants and their environment or plants and other organisms.

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                              • Oikos.

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                                Oikos publishes original and innovative research on all aspects of ecology. Emphasis is on theoretical and empirical work aimed at generalization and synthesis across taxa, systems, and ecological disciplines. Papers should be well founded in ecological theory and contribute to new developments in ecology by reporting novel theory or critical experimental results.

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                                • Proceedings of the National Academy of Sciences USA.

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                                  PNAS, established in 1914, is one of the world’s most cited multidisciplinary scientific serials. It publishes research reports, commentaries, reviews, perspectives, colloquium papers, and actions of the Academy. It covers the biological-biomedical, physical, and social sciences. PNAS is published weekly.

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                                  • The Quarterly Review of Biology.

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                                    This review journal in biology dates back to 1926 and publishes articles in all areas of biology, but with a traditional emphasis on evolution, ecology, and organismal biology. QRB papers offer new ideas, concepts, and syntheses. In addition, the book review section of QRB is the most comprehensive in biology.

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                                    • Trends in Ecology & Evolution.

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                                      Trends in Ecology & Evolution keeps scientists informed of new developments and ideas across the full range of ecology and evolutionary biology—from the pure to the applied and from molecular to global. It appears in monthly issues.

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                                      Defining Community Ecology

                                      The definition of community refers to the discourse mentioned under General Overviews on the degree to which communities are “sufficiently coherent entities to be considered appropriate objects of study” (reviewed in Ricklefs 2008). The modern definition of community (see, e.g., Vellend 2010, cited under The Four Fundamental Processes), which is “a group of organisms representing multiple species living in a specified place and time, including all scales of space and time,” stipulates the properties of the community, regardless of their integrity and possible coherence. This makes the definition of community ecology the following: “the study of patterns in the diversity, abundance, and composition of species in communities, and the processes underlying these patterns” (Vellend 2010). Johnson and Stinchcombe 2007 accentuates the study of the interactions between species and their environment, affecting abundance, distribution, and diversity of species within communities. Because these authors are evolutionary biologists, they stipulate that within these species genetic variation and evolutionary change take place, with great implications for these interactions.

                                      • Johnson, M. T. J., and J. R. Stinchcombe. 2007. An emerging synthesis between community ecology and evolutionary biology. Trends in Ecology & Evolution 22.5: 250–257.

                                        DOI: 10.1016/j.tree.2007.01.014E-mail Citation »

                                        The authors suggest a synthesis between community ecology and evolutionary biology, identifying how genetic variation and evolution within one species can shape the ecological properties of entire communities and, in turn, how community context can govern evolutionary processes and patterns.

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                                        • Ricklefs, R. E. 2008. Disintegration of the ecological community. American Naturalist 172.6: 741–750.

                                          DOI: 10.1086/593002E-mail Citation »

                                          In his essay, Ricklefs argues that “the seemingly indestructible concept of the community as a local, interacting assemblage of species has hindered progress toward understanding species richness at local to regional scales.” He advocates a community concept based on interactions between populations over a continuum of spatial and temporal scales within entire regions, including the population and evolutionary processes that produce new species.

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                                          Historical Background

                                          The now-classic debate about the nature of communities, whether they are integrated fundamental units or arbitrary assemblages of populations whose traits allow them to exist in a certain area, is personalized in, respectively, Clements 1916 and Gleason 1926 (both cited under General Overviews). Focus on the mathematical theory in community ecology was developed in the same period. Lotka 1925 and Volterra 1926 independently modeled two-species community models, which are still used as the point of departure for many models of modern population dynamics. The 1960s were characterized as the period of the “traditional community ecology,” during which time a vast body of theoretical and empirical research was developed (see Brown 1995). More recently there have been successful attempts within the vast field of community ecology on approaches that make it possible to bring structure in this vast body of data and models (Chase and Leibold 2003, under General Overviews; McGill, et al. 2006; Johnson and Stinchcombe 2007, under Defining Community Ecology; Ellers 2010, under Evolutionary Community Ecology; Vellend 2010, under The Four Fundamental Processes). The following section, New Approaches, concentrates on four recent, promising approaches that may lead to a more structurized discipline.

                                          • Brown, J. H. 1995. Macroecology. Chicago: Univ. of Chicago Press.

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                                            In this book Brown presents the rationale for a macroscopic, statistical, nonmanipulative approach that can overcome some of the inherent limitations of traditional “microscopic” experimental ecology and provide new insights into the structure and dynamics of complex ecological systems.

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                                            • Lotka, A. J. 1925. Elements of physical biology. Baltimore: Williams & Williams.

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                                              Alfred Lotka wrote this book on theoretical biology in 1925. He is best known for the predator-prey model he proposed, at the same time but independent from Volterra 1926 (the Lotka-Volterra model), which still forms the basis of many models used in the analysis of population dynamics. Reprinted in 1956 as Elements of Mathematical Biology (New York: Dover).

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                                              • McGill, B. J., B. J. Enquist, E. Weiher, and M. Westoby. 2006. Rebuilding community ecology from functional traits. Trends in Ecology & Evolution 21.4: 178–185.

                                                DOI: 10.1016/j.tree.2006.02.002E-mail Citation »

                                                In their article the authors state that community ecology has lost its way by focusing on pairwise species interactions, independent of the environment. They assert that community ecology should return to an emphasis on four interlinked themes: functional traits, environmental gradients, the interaction milieu, and performance currencies.

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                                                • Volterra, V. 1926. Fluctuations in the abundance of a species considered mathematically. Nature 118.2972: 558–560.

                                                  DOI: 10.1038/118558a0E-mail Citation »

                                                  In 1926, the Italian mathematician Vito Volterra proposed a differential equation model to explain the observed increase in predator fish (and corresponding decrease in prey fish) in the Adriatic Sea. At the same time in the United States, the equations studied by Volterra were derived independently by Alfred Lotka (see Lotka 1925). The Lotka-Volterra model is the simplest model of predator–prey interactions.

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                                                  New Approaches

                                                  This section contains a selection of four new approaches in this broad field: The Metacommunity Approach, The Functional Trait Approach, Evolutionary Community Ecology, and The Four Fundamental Processes. They were chosen because of their promising outlook and the fact that all four generated considerable scientific interest, resulting in subsequent articles referring to these approaches.

                                                  The Metacommunity Approach

                                                  While community ecology in the 1980s and 1990s focused on how species interactions within localities influence patterns and abundances, assuming that local communities are closed and isolated, more recently the importance of space and time has been recognized, suggesting that different principles might apply at different times and scales. Recent syntheses, such as in Leibold, et al. 2004, have suggested that metacommunity theory can be categorized into four general frameworks: neutral, patch dynamics, species sorting, and mass effects. Neutral models emphasize how far one can go by assuming that communities are structured by stochastic processes only. Patch dynamic models assume that species may display trade-offs in their abilities to compete and colonize. Species sorting focuses on deterministic processes resulting from differential responses of species to heterogeneous environments. Mass effects combine aspects of patch dynamics and species sorting. This framework assumes that species have differential responses to environmental conditions and differential extinction, colonization, and dispersal rates. This metacommunity approach has presented new insight in several aspects of community ecology, showing that, concerning the hypothesis of “community-wide character displacement” (locally coexisting species are less similar than random draws from a larger regional pool), a metacommunity view results in the reverse pattern: locally coexisting species should be more similar than random draws. Other examples concern the inference of regional influences on local communities, based on nonasymptotic relationships between local and regional diversity and consistent patterns in the distribution of local versus regional biodiversity across different gradients. Underlining the importance of upscaling for the discipline of community ecology, Leibold, et al. 2004 concludes that biodiversity at larger spatial scales may regulate the dynamic behavior of ecosystems in ways strongly differing from currently documented effects of local diversity. In their broad overview of metacommunity ecology (Chase and Bengtsson 2010), Jonathan Chase and Janne Bengtsson conclude that, while metacommunity ecology is a rapidly growing field, considerable new insights, synthesis, theories, and empirical studies are needed.

                                                  • Chase, J. M., and J. Bengtsson. 2010. Increasing spatio-temporal scales: Metacommunity ecology. In Community ecology, processes, models, and applications. Edited by H. A. Verhoef and P. J. Morin, 57–68. Oxford Biology. Oxford: Oxford Univ. Press.

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                                                    In their review, the authors discuss a number of recent advances in metacommunity ecology, including diversity partitioning at different spatial scales, the interaction between stochastic and deterministic factors, food web interactions, and cross-ecosystem boundaries.

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                                                    • Leibold, M. A., M. Holyoak, N. Mouquet, et al. 2004. The metacommunity concept: A framework for multi-scale community ecology. Ecology Letters 7.7: 601–613.

                                                      DOI: 10.1111/j.1461-0248.2004.00608.xE-mail Citation »

                                                      The authors review current understanding about the metacommunity concept, which is an important way to think about linkages among different spatial scales in ecology.

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                                                      The Functional Trait Approach

                                                      In McGill, et al. 2006 (cited under Historical Background), the authors state that for years in community ecology, the main focus has been the study and modeling of species interactions. They call for a return to the trait- and environment-focused route: “The time is ripe to get back to species, characterized by their traits, located in a heterogeneous environment” (McGill, et al. 2006). They give two reasons to follow this route: (1) the availability of remote sensing data, geographical information, and the accumulation of worldwide data sets, and (2), the need to predict the effects of global warming. In this approach, four themes play a role in the functional-traits research program: traits, environmental gradients, the interaction milieu, and performance currencies. Examples of functional traits include basal metabolic rate, beak size, seed or egg size, nutrient concentrations and stoichiometries, adult body mass, frost tolerance, potential photosynthetic rate, and leaf mass per area. According to the authors, statements about traits lead to generality and predictability, contrary to the standard nomenclatural approach. Natural communities occur on gradients of temperature, moisture, and soil chemistry, and notwithstanding the importance of species interactions, predicting the impact of global warming, for example, can be done based only on knowledge on the effect of environmental gradients on communities. Biotic interactions are best modeled as a milieu or biotic background with which an organism interacts. Frequency distributions of traits (e.g., a histogram of heights) that are important for a given type of interaction give an operational definition of the interaction milieu (e.g., the milieu of plant light competition). Finally, in this approach, performance currencies derive from the processes of acquiring, allocating, and spending energy and mineral nutrients, because they are linked to the physical environment and to interactions in the interaction milieu. The above-mentioned four themes are tied together by the fundamental-realized niche framework (Hutchinson 1957). This approach differs strongly from three common approaches in community ecology: past empirical studies of species interactions, community matrices, and neutral theory. It has some overlap with common resource models and macroecology. If we want to get a more mechanistic understanding of multispecies communities in relation to physical geography, this functional-traits-on-gradients approach may be successful.

                                                      • Hutchinson, G. E. 1957. Concluding remarks. Cold Spring Harbor Symposia on Quantitative Biology 22:415–427.

                                                        DOI: 10.1101/SQB.1957.022.01.039E-mail Citation »

                                                        In these concluding remarks at a Cold Spring Harbor Symposium, Hutchinson presents a detailed analysis of the nature of the ecological niche and the validity of the principle of niche specificity. His new concept of the niche has become known as the Hutchinsonian Niche and was considered by the author as his best contribution to science.

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                                                        Evolutionary Community Ecology

                                                        In this approach, bridging the gap between community ecology and evolutionary biology may lead to “new insight if evolution in one species affects some ecological property of a community or, alternatively, the ecological properties of a community influence the evolutionary dynamics within one or more populations” (Johnson and Stinchcombe 2007), cited under Defining Community Ecology) It has been found that evolutionary processes over long as well as short time scales influence community dynamics and patterns, whereas, vice versa, interactions among species in communities can influence evolutionary processes and patterns. In Johnson and Stinchcombe 2007, four conditions that affect support for the hypotheses that bridge community ecology and evolutionary biology are presented: genetic variation and microevolution affect community properties, evolutionary history influences community assembly, community context affects the direction and rate of evolution, and multiple species within communities codiversify. However, despite the convincing results presented in this article, the importance of bridging community ecology and evolutionary biology has, according to these authors, not yet been convincingly demonstrated, due to a lack of evidence. A similar but further-reaching opinion is presented in Ellers 2010.This article, titled “Evolutionary Processes in Community Ecology,” expresses the need for a greater number of field studies, the expansion of the range of taxa (mainly plants), a careful choice of the study system in relation to the extensive work to measure both the genetic and phenotypic diversity of all species. It is the author’s opinion that a true integrative approach can address a wider range of questions than just the reciprocal effects of genetic and species diversity. One of those questions is if, and to what extent, genetically diverse species persist longer in a community than genetically uniform species—or whether species at particular trophic positions are more likely to be genetically diverse. If this promising new field of evolutionary community ecology will flourish depends on the researchers of both disciplines extending their original niches.

                                                        • Ellers, J. 2010. Evolutionary processes in community ecology. In Community ecology, processes, models and applications. Edited by H. A. Verhoef and P. J. Morin, 151–161. Oxford Biology. Oxford: Oxford Univ. Press.

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                                                          Ellers states that bridging the gaps between community and evolutionary ecology can be mutually beneficial. “In fact, we may even need an integrative approach in order to face fast changing environmental conditions such as global warming and urbanization, which pose ecological as well as evolutionary challenges” (p. 152).

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                                                          The Four Fundamental Processes

                                                          Under the title “Conceptual Synthesis in Community Ecology,” Vellend 2010 presents a framework 1) to clarify the essential similarities and differences among the many conceptual and theoretical approaches and 2) to articulate a very general theory of community dynamics. This framework is based on four distinct kinds of processes: selection, drift, speciation, and dispersal. These processes are close analogues of the “big four” in population genetics: selection, drift, mutation, and gene flow. In the author’s opinion, the addition of speciation, drift, and dispersal to the deterministic interactions among species and between species and their environments (i.e., selection) gives a “logically complete set of process categories within which all other more specific processes can be placed.” In a comprehensive table, Vellend relates many of the influential theories, models, and frameworks in community ecology to their emphasis on selection, drift, speciation, and dispersal, and the essential differences and similarities are presented clearly. The author hopes that this conceptual framework will be useful to practicing community ecologists to place their work in a process-based context. Furthermore, this framework is supposed to clarify the diffuse way in which some textbooks arrange their chapters. In his general theory of community dynamics, he incorporates the four fundamental processes in the following way: “species are added to communities via speciation and dispersal, and the relative abundances of these species are then shaped by drift and selection, as well as ongoing dispersal, to drive community dynamics.”

                                                          • Vellend, M. 2010. Conceptual synthesis in community ecology. Quarterly Review of Biology 85.2: 183–206.

                                                            DOI: 10.1086/652373E-mail Citation »

                                                            In his generalizing framework, Vellend states that community ecology is influenced by only four processes: selection, drift, speciation, and dispersal. In his view, all theoretical and conceptual models in community ecology can be understood with respect to their emphasis on these four processes. Organizing the material of community ecology according to this framework clarifies similarities and differences among the many conceptual and theoretical approaches to the discipline.

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                                                            Applications

                                                            Community ecology has many real-world applications, deriving from the fact that the abundance of species strongly depends on competitive and predatory interactions. These interactions are major drivers of ecosystem processes, and they are key to the delivery of ecosystem services. In Verhoef and Morin 2010, the authors give four examples of applications in the context of community ecology: applications of community ecology approaches in terrestrial ecosystems, structure and functioning of emerging marine communities, diversity and ecosystem services at the intersection of local and regional processes, and management of salt marshes. Regarding applications of community ecology approaches in terrestrial ecosystems, there are linkages between belowground and aboveground subsystems that influence many major processes, such as ecosystem restoration following land abandonment or the fate of introduced exotic alien species. In terms of structure and functioning of emerging marine communities, fishing pressures can ultimately affect average food chain length, which can initiate regime shifts to alternate semistable states and affect the provision of marine ecosystem services to society. Under diversity and ecosystem services at the intersection of local and regional processes, examples drawn from research on organic farming and biological control illustrate that (meta)community theory combined with good knowledge of the system under management is useful to understand how human-dominated ecosystems can be managed at spatial scales relevant to managers. Finally, regarding the management of salt marshes, natural selection in these systems results in the dominance of a single tall-grass species, to the detriment of the natural herbivores such as geese and hares. The introduction of large herbivores can reverse succession, facilitating geese and hares. Without this management, these marshes would develop into low-diversity systems of reduced value. Rohr, et al. 2006 suggests community ecology as a framework for predicting contaminant effects. The authors present an approach that offers predictions of the strength and direction of indirect effects, which species are crucial for propagating these effects, which communities will be sensitive to contaminants, and which contaminants will be most insidious to communities. They discuss its value for integrated pest management, ecological risk assessment, and the development of remediation and ecosystem management strategies.

                                                            • Rohr, J. R., J. L. Kerby, and A. Sih. 2006. Community ecology as a framework for predicting contaminant effects. Trends in Ecology & Evolution 21.11: 606–613.

                                                              DOI: 10.1016/j.tree.2006.07.002E-mail Citation »

                                                              In their article the authors state that is it important to identify a predictive theory for the direct and indirect effects of the multitude of anthropogenic chemicals entering most ecosystems. They propose that the impacts of contaminants can be simplified and unified under the framework of community ecology.

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                                                              • Verhoef, H. A., and P. J. Morin, eds. 2010. Community ecology, processes, models, and applications. Oxford Biology. Oxford: Oxford Univ. Press.

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                                                                This book provides a survey of the state of the art in theory and applications of community ecology, with special attention to topology, dynamics, and the importance of spatial and temporal scale, as well as applications to emerging problems in human-dominated ecosystems. This book contributes to a better understanding of the mechanisms that shape ecological communities. It provides interesting course material for graduate seminars.

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