Disparity and Diversity 59

richness together with species genealogy would keep track of dispar-

ity for free. But phenotype disparity is not well-captured by traditional
higher taxa, as their presentist bias makes old taxa look stranger than
they are.
Budd’s argument is powerful. But it is also important to be clear
about what it does not show: the fundamental distinction between
crown and stem taxa is neutral on the rate of evolutionary change, and
neutrality cuts both ways. Our ability to explain the systematics of the
metazoan radiation using that distinction is compatible with a highly
disparate Cambrian fauna. We could and should make the distinction
between stem and crown arthropods, even if stem arthropods are as
disparate as Gould, and Mark and Dianna McMenamin suppose. The
stem/crown distinction makes no special assumptions about the nature
of Cambrian evolution. As we have just noted, it is compatible with
conservative assumptions about Cambrian differentiation. But so long
as the Metazoa are a monophyletic clade, it is equally compatible with
the idea that this differentiation was unique. Even if the Burgess fauna
were as rich and weird as Gould suggests, the strange and problematic
Cambrian fossils would still be members of stem groups of extant meta-
zoan lineages: they are bilaterian branches.
The Budd-Jensen argument suggests that, in thinking about the
disparity of animal life, we need to guard against taxonomic illusion:
over-estimating early disparity because early fossils are hard to fit into
taxonomic schemes designed to fit extant organisms. If the Cambrian
fauna were indeed highly disparate, we could still construct a well-
confirmed phylogeny with a stem/crown distinction, one showing (for
example) where Opabinia and Anomalocaris fit into the stem arthropods.
Equally, we can construct a phylogeny if Cambrian disparity slowly
increases over time. But is there any reason independent of taxonomic
awkwardness to suppose that Cambrian fauna were unusually disparate?
This takes us to the problem of morphospace: the idea that we can rep-
resent morphology as a multidimensional space, with each dimension
of that space corresponding to a variable morphological feature. If there
is a space of animal form, the disparity of life at a time is the volume of
that space occupied by life at that time. There would then be an open
question about the covariation between species richness and the
occupation of morphospace. While conceding that we are yet to de-
velop an explicit characterization of morphospace, Gould suggests that
the Burgess arthropods are highly disparate in just this sense (Gould
1991). In the next chapter we explore the idea of defining phenotype
biodiversity via the occupation of morphospace, and the connection
between morphospace and species richness.