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Taxonomy Red in Tooth and Claw 25

species richness in phylogenetic context—a good surrogate for bio-


diversity in general? Is this structure a good way of identifying those
patterns that are the signatures of ecological and evolutionary process,
and a good way of specifying the input to further ecological and evo-
lutionary changes? No one thinks that species structure is literally all
there is to biodiversity. If the life sciences had perfect information about
biological systems, and unlimited experimental and computational re-
sources, biologists would not just count species. But the life sciences
need a simple and empirically tractable model of total biodiversity. So
perhaps the number and distribution of species, augmented in various
ways for particular purposes, serves as a good multipurpose measure of
local, regional, and global biodiversity. We begin this project in chapter
2, which seeks to resolve the somewhat puzzling fact that species rich-
ness is often used as a surrogate for overall biodiversity, even though the
nature and identification of species continues to be controversial, and
even though no one thinks that species richness is all there is to biodi-
versity. In chapters 3 and 4 we explore the relationship between species
richness and phenotypic disparity, beginning with Stephen Jay Gould’s
well-known claims that species richness does not track disparity.
We extend the focus on phenotype diversity in chapter 5 by bring-
ing development explicitly into the picture. Genes are paradigmatic
developmental and evolutionary resources; the evolutionary plastic-
ity and resilience of species depends to a considerable extent on the
genetic resources available in species gene pools. So in that chapter,
we discuss the diversity of developmental resources and explore the
extent to which phylogenetically structured species richness is a sur-
rogate for developmental diversity. In the final “filling” chapter of the
conservation biology sandwich, we then turn to ecology. For us, the
crucial question is whether communities or ecosystems function as
biologically important organized systems. If they do not, if species (or
populations) respond to environmental vectors independently of their
neighbors’ response, then species richness captures ecological diversity.
Information about the species present, and the environmental variables
acting on those species, would suffice for understanding ecological out-
comes. That is not true if communities are organized systems. Ecosys-
tem services, for example, would then depend on collective properties
of the community. We then return, rather skeptically, to the problems
of measurement and value as they have been conceived in conservation
biology. Of course, conservation biology can be and is used purely as an
instrumental, applied science to estimate specific dangers from threats
to specific populations and to devise means of defusing those threats.
Conservation biology can, and conservation biologists do, estimate the

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