Disparity and Diversity 43

will depend both on the number of distinct elements in it and the

extent of their differentiation.
In this chapter and the next, we will consider the idea that tracking
species number does not track a second important dimension of biodi-
versity: phenotypic richness. We will focus on the claim that diversity
(= species number) does not track disparity (= variation across pheno-
types). A biota can be species rich but not very disparate, if the species
composing the biota are rather similar. Arguably, many island faunas are
more diverse than disparate, for they often derive from a few founder
species, and this constrains the variation that evolves. Despite their dif-
ferent beaks, Darwin’s finches really are pretty similar birds. Stephen
Jay Gould made this diversity-disparity distinction famous in his 1989
classic Wonderful Life, and it has generated ongoing controversy.
The phenotypic spread of a biota is important to its evolutionary
and ecological future; a phenotypically richer biota is more apt to be
biologically prepared for change of various kinds. It can recruit and
amplify existing variation to meet change, rather than having to wait
on migration or evolution to create new variation. And the phenotypic
spread of a biota is also a signal of its ecological and evolutionary past,
of the processes that have been important in its making. In this chap-
ter our main interest will be in the idea that high phenotypic biodi-
versity is a signal of distinctive mechanisms (we discuss the ecological
consequences of phenotypic diversity in chapter 6). But whether we
are interested in diversity as a signal or as an input to further change,
the spread of phenotype is important. What is not clear is whether we
have to count it separately (and if so, how we should count) or whether
spread is indexed by species-level diversity. While there is more to the
phenotype of an organism than its morphology, we will use morphology
as our surrogate for phenotype diversity. For we have temporal depth
in morphological information. Behavior and physiology leave less of
a paleobiological signal. There is no doubt that phenotypes vary from
one another in objective and important ways. There is, however, con-
siderable doubt that there is a single metric or framework into which
these variations can be placed; a decent multipurpose measure of the
phenotypic spread of a biota. As we shall see, a wide range of biological
projects suggest the existence of such a framework. But we are skeptics;
there is no overall, objective measure of phenotype difference. If there
were, the phenetic program in systematics could be revived.
Gould himself explored this distinction in the context of paleobiol-
ogy, of long-term trends in biodiversity. There has been considerable
debate about the impact of biases in the fossil record on our ability to re-
liably estimate changes in species number over time (Alroy 2000; Foote