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58 3 Reframing Urban Ecology In this chapter, I examine the origin of urban ecology and its evolution toward a science of coupled human-natural systems. During the past hundred years, advances in the scientific understanding of ecological systems have opened up new opportunities to integrate humans into ecology. Yet despite increasing human domination of Earth’s ecosystems, ecological theories are still conceptualized primarily in terms of biophysical, ecological, and evolutionary processes that are substantially unaffected by human agency. I discuss observations of “anomalies” in urban ecosystems that are difficult to reconcile within ecological theory and the emergence of a co-evolutionary paradigm that better explains patterns observed in urbanizing regions. I conclude the chapter by highlighting emerging elements for reframing the science and practice of urban ecology. Ecology for an Urban Planet The science of ecology has evolved through many stages and definitions (Likens 1992; O’Neill 2001; Marquet et al. 2014). What characterizes its evolution is the dynamic interplay between changes that have occurred in Earth’s ecosystems over time and parallel shifts in our scientific understanding. It is through such interplay, over the past century, that new ideas have developed (Graham and Dayton 2002). The emergence and evolution of the ecosystem concept provided a framework for studying ecological interactions among individuals, populations, and communities and their abiotic environments, and for studying the changes in these relationships over time (Likens 1992). Although the term ecosystem Reframing Urban Ecology 59 was coined by Tansley in 1935 and the underlying idea can be traced back to Marsh (1864), the concept of ecosystem became a standard paradigm for studying ecological systems only when systems analysis emerged in the second half of the past century (Holling 1973; Odum 1971). Since then, ecological scholars have revised the concept to acknowledge multiple equilibria and the open, hierarchical, spatially heterogeneous, and scaled nature of ecosystems (Levin 1999; O’Neill et al. 1982; Pickett, Parker, and Fiedler 1992). Still, the term essentially excludes humans. The ecosystem concept, which is at the core of ecological thinking, emerged from a view of systems theory that is difficult to reconcile with our current understanding of dynamic ecological systems that may operate far from equilibrium (O’Neill 2001, 3275). In his MacArthur Lecture in Ecology, Robert O’Neill (ibid., 3276) proposed that in ecology the “ecosystem concept” has constituted a paradigm in the Kuhnian sense, an “a priori intellectual structure” rather than an “a posteriori empirical observation ” about how nature works. While the concept offers a practical approach to the study of ecological systems—one that has proved instrumental in ecology’s progress—it has some drawbacks. O’Neill argued powerfully that the ecosystem paradigm accepts a set of assumptions that limits our thinking and the questions we ask because it emphasizes some properties of nature while ignoring others. For example, emphasis on the self-regulating nature of ecosystems has biased ecosystem scientists to see disturbance as interference from outside. Yet we know that disturbance regimes are critical to understanding stability and ecosystem function. A key property of ecosystems is their ability to change state in response to a spectrum of variable conditions (Holling 1973). Ecosystems have evolved over millions of years through changes in biotic-abiotic interactions. Since the Industrial Revolution, humans have increasingly dominated such interactions. But in ecology, humans are the only species that is considered to be external to ecosystems. They are seen as consumers of ecosystem services rather than active participants in eco­ system processes. Yet today humans are creating novel ecosystem functions well outside the range of values and conditions that Earth’s ecosystems have experienced throughout their evolution (O’Neill 2001; Tilman and Lehman 2001). If ecology is the study of organisms’ interactions with one another 60 Chapter 3 and with the environment, and of the transformations of matter, energy, and information, then the advent of the Anthropocene implies a fundamental shift in ecology as a science. Earth’s atmosphere, on which we all depend, emerged from the metabolic processes of vast numbers of singlecelled algae and bacteria living in the seas 2.3 billion years ago. These organisms transformed the environment into a place where human life could develop. From a planetary perspective, the emergence and rapid expansion of cities across the globe may be a turning point in the life of our planet, one on the scale of the Great Oxidation. Recent increases in positive feedback (e.g., climate change), along with the emergence of evolutionary innovations (e.g., novel metabolisms...


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