Each level requires different methods of analysis, different modes of understanding, and, ultimately, different approaches to management. For managers, it is not just a matter of counting species or individuals. Managers must consider the role of biodiversity in the functioning of ecosystems and the effects of management and use of resources on ecosystem processes (chapter 3).

George Evelyn Hutchison (1965), one of the founders of modern ecology, wrote about the "evolutionary play in the ecological theater". This multilayered drama generates, sustains, shapes, and sometimes even diminishes biodiversity. Charles Darwin's reflections on species diversity underlay one of the most far-reaching theories in the history of ideas: the theory of evolution by natural selection. His travels from England to the strikingly different landscapes of the New World left him awestruck and inspired. Whatever constitutes biodiversity, Darwin recognized that Brazil had a lot of it and certainly more than he left behind in an English midwinter. No modern biologist would disagree. Like Darwin, we often equate biodiversity with the number and novelty of the species present.

Species, Populations, and Genes

There is genetic diversity within species. If each species were reduced to one small population of genetically similar individuals, we would lose much biodiversity. As we move across a region, the species change, even if the numbers of species in different places might not; a forest and an adjacent grassland might contain almost entirely different assemblages of species, for instance. Moreover, the ecosystem processes in a grassland differ from those in the forest nearby.

A population consists of individuals of the same species that live in the same place and interact in various ways, including interbreeding. Populations of the same species living in different places can exchange members, but they often are genetically differentiated to some degree and the further they are separated from each other, the more distinctive they become. Metapopulations are groups of spatially separated populations that occur in patches of habitat across a landscape. Populations can become locally extinct in different habitat patches across a landscape; they infrequently exchange members, and when they do, the passage between local populations is generally hazardous and entails movement across inhospitable habitat. Local populations that make up a metapopulation experience extinction, and habitat left open is recolonized at some finite probability by other local populations within the metapopulation.

The genetic variability among individuals within a species can result from gene recombination or mutation, genetic polymorphism (the presence of different forms of the same gene), isolation of gene pools, local selection pressures, habitat (environmental) complexity, landscape mosaics, and environmental gradients. Specific genetic combinations in populations result from natural selection acting



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