was speculating that extinction rates might be between 27,000 and
100,000 species per year (280), though in recent times these alarming estimates have come under considerable criticism, particularly in Bjørn Lomborg’s The Skeptical Environmentalist (2001, 249–57).1 Wilson’s was the first of a number of books that have popularized the term “biodiversity.” “Biodiversity” is, of course, a blend of the phrase “biological diversity.” The term was coined in 1985 by Walter G. Rosen for “The National Forum on BioDiversity,” a conference held in Wash- ington DC in 1986 (Harper and Hawksworth 1995). Its proceedings were edited by Wilson (1988) under the title Biodiversity. While Wilson was certainly concerned with species numbers, an important theme of his work, and one that we shall echo, is the idea that diversity cannot be captured by species numbers alone. Species are important, for at least in eukaryotic organisms biological diversity is parceled up into independently evolving lineages. But we cannot assume that we can measure the diversity of a system just by counting species numbers. For the functioning of biological systems—as one quote with which we begin reminds us—depends on the kinds and combinations of organ- isms present, a fact whose importance has recently been reiterated by Kevin McCann (2007). However, if we accept that biodiversity is important, and if we ac- cept that there is more to biodiversity than species number, we must be able to answer the following questions: What is diversity and how do we measure it? What is the appropriate focus for conservation biology? We shall see that, from the beginning, there has been a potentially trou- bling ambiguity in thinking about biodiversity in conservation biology (and hence applied ecology). The ambiguity is between what conserva- tion biology wanted to conserve and the mechanisms of conservation. Biodiversity is sometimes thought of as a measure of what we want to keep, but it is sometimes also thought of as a tool: a measure of an in- strumentally important dimension of biological systems. So in much of this book we will address the purposes to which the idea of biodiversity is put. Disciplines are shaped by their histories, which transmit legacies of tools, assumptions, and projects from one generation to the next. We begin by asking why ecologists and conservation biologists started using the term. Conservation biology is a young discipline because the idea that we ought to put scientific and governmental resources into conserving na- ture is a relatively new one. In the United States in 1873 it was suggested that the government ought to “preserve spots of primitive land-surface of which the vegetation was especially interesting” (Bocking 1997, 21), and indeed there were some limited successes in this respect early in