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40 chapter two

cycle, the species category is something like a natural kind. In counting


species and comparing the species richness of different regions, we can,
roughly, compare like with like.6

2.4 species and biodiversity

The living world is organized into phenomenological species: recog-


nizable, reidentifiable clusters of organisms. This fact makes the pro-
duction of bird and butterfly field guides, identification keys for inver-
tebrates, regional floras, and the like, all possible. In this chapter we
have embraced the common biological wisdom that phenomenological
species richness captures a crucial dimension of biodiversity. There are
many routes through which one population can become demographi-
cally isolated from, and hence evolutionarily independent of, popula-
tions that were once sources and sinks of its own genes. But the fact of
isolation and evolutionary independence is of immense importance to
the fate of local adaptation in such populations. So the phenomenologi-
cal species richness of a region is, in an importance sense, a catalogue
both of phenotypic variety and of the potential evolutionary resources
available in that region. There is an important difference, on this pic-
ture, between a single widespread and phenotypically variable species
(like the common brushtail) and a set of closely related species. The
available phenotypes, population sizes, and ecological roles might be
exactly the same. But one set of phenotypes will be entrenched by spe-
ciation mechanisms, and hence will survive minor ecological changes
that increase migration rates across the landscape; the other set is much
more fragile in the face of relatively minor ecological change. So it does
not just matter what phenotypes are present; how they are bundled into
species is also important. In effect, we have defended a version of an
evolutionary species concept, and we accept that the collection of in-
dependently evolving lineages in a region is a key component, perhaps
the key component, of that region’s biological diversity.
That said, we need to add some important qualifications. The iden-
tification of phenomenological species with metapopulations in partial
stasis holds good only for some chunks of the tree of life. It may not
fit plants. It clearly does not fit microbes. Frederick Cohan has devel-
oped an account of bacterial species that identifies those species with
the units of bacterial evolution.7 But this involves abandoning the idea
that phenomenological species are typically important units of stasis
and change; in Cohan’s view, phenomenological bacterial species are
amalgams of many evolutionary units (Cohan 2002). Even if we set
that aside, phenomenological species do not represent equal amounts

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