Disparity and Diversity 55

bilaterian body plan is not fixed once in place. We will see in chapter 5
that there might be something to the idea that body plans, once they
evolve, are stabilized and become difficult to change. This idea is an
important theme of Bill Wimsatt’s work (see Wimsatt 2007). But this
idea of stabilization does not show there is anything special about phyla,
about, say, the arthropod rather than the trilobite body plan. The body
organizations we take to be distinctive of the metazoan phyla are not
especially, uniquely stabilized. A phylum is a large monophyletic chunk
of the tree of animal life, and the organisms in a phylum will resemble
one another in various ways due to their shared deep ancestry. To be
told that a biota includes representatives from, say, the arthropods, mol-
lusks, and bivalves is to be given useful information (in contrast, say,
to being told that it contains organisms used in Wiccan spells). But
phyla are not objectively countable units. After all, the idea of a body
plan is fundamentally hierarchical. Cephalopods are mollusks. There is
a cephalopod body plan, and a mollusk body plan, and the first is a ver-
sion of the second. But there is nothing objective that determines which
of these organizations, if either, characterizes a phylum.
So we should be cautious about inferring phenotypic disparity from
traditional taxonomy. We should be especially cautious if the animals
are ancient. This new phylogeny shows how the Cambrian fauna can be
integrated within the tree of life, and this integration predicts that the
Cambrian fauna would seem to be very disparate, even if it were not.
The crucial distinction is between the stem and the crown members of
a lineage. This distinction is best explained through an example, and
we will borrow Andrew Knoll’s example of the divergence between the
arthropods and their (possible) sister phylum, nematodes (Knoll 2003,
187–90). These are both members of the Ecdysozoa, so they share the
molting cuticle characteristic of that clade as well as its genetic and
developmental signatures, but apparently not much else. Arthropods
are segmented, with jointed appendages and an external skeleton made
from chitin. Nematodes are a species-rich clade of tiny worms, tapered
at both ends. The joint ancestor of this clade—their last common
ancestor—would have resembled neither. It would not have possessed
the distinctive suite of arthropod characteristics, but neither would it
have had the radically simplified anatomy (compared to many bilateria)
of the nematodes. So consider the evolutionary history of the arthropod
lineage leading from this last common ancestor to crustaceans, insects,
and spiders. On this lineage the distinctive characteristics that unite the
arthropods—segmentation, chitinous skeleton, jointed appendages—
would have evolved. But not all at once. Perhaps the order was chitin,
then segmented body, then jointed appendages.