Taxonomy Red in Tooth and Claw 7

over time, thus alerting us to troubling changes, and enabling us to

evaluate the success or failure of protection. But these surrogates are
supposed to be measures of overall richness or variety.
For example, in their influential overview of conservation planning,
C. R. Margules and R. L. Pressey write:

Biological systems are organized hierarchically from the molecular to

the ecosystem level. Logical classes such as individuals, populations,
species, communities and ecosystems are heterogeneous. Each member
of each class can be distinguished from every other member. It is not
even possible to enumerate all of the species of any one area, let alone
the members of logical classes at lower levels such as populations of
individuals. Yet this is biodiversity, and maintaining that complexity is
the goal of conservation planning. (Margules and Pressey 2000, 245)

They are not alone. In a similarly wide-ranging and much cited over-
view, Craig Groves and his colleagues define biodiversity as “the variety
of living organisms; the biological complexes in which they occur, and
the ways in which they interact with each other and the physical envi-
ronment . . . this definition . . . characterises biodiversity as having three
primary components, composition, structure and function” (Groves
et al. 2002, 500).
We will be interested in this idea of biodiversity as a natural feature
of biological systems, though like Kevin Gaston and John Spicer (2004),
we will reject the idea that there is a single measure of the diversity of
a biological system. We doubt, in fact, that anyone really thinks there is
a single natural property of a biological system that captures all its bio-
logically relevant diversity, though perhaps Daniel Brooks and Deborah
McLennan come close, suggesting that diversity is essentially species
in their phylogenetic structure. They begin their 1991 monograph with
a thought experiment about a tidal pool, inviting their readers to com-
pare how much they know about an organism in the pool if given eco-
logical information (the organism is a predator) or if given phylogenetic
information (the organism is a fish). A predator, after all, might be an
octopus, a starfish, a crab, or a fish, yet a starfish and an octopus differ
far more than any two fishes (Brooks and McLennan 1991; 2002).
We will not find much reason to accept the idea that diversity is es-
sentially captured by species and their phylogeny. But we shall see that
a somewhat more modest view deserves to be taken seriously: that a
phylogenetically informed species count is a good general purpose in-
dicator or surrogate for total biodiversity (see, for example, Forest et al.
2007). We discuss a number of proposals for meshing species richness