Disparity and Diversity 53

reply depends on our being able to place our strange-seeming Cambrian

creatures in the tree of animal life, so to that we now turn.
At the time of Wonderful Life, the order of branching of the metazoan
phyla was extremely controversial, in part because their body plans were
so distinct. There was an orthodox view of the history of the Metazoa,
but it was coarse-grained, and many relationships among the phyla were
unresolved or controversial. The fundamental split was between the
basal, embyrologically simple (developing from just two germ layers)
sponges, cnidarians, and ctenophores in one group and the bilaterians
in the other. The bilaterians are morphologically and embryologically
more complex; they have three cell layers in their embryo—they are
tripoblasts. They have front-to-back and up-to-down axes of symmetry;
they have an organized nerve system with a frontal concentration of
nerve cells; most of them have a through-gut with a distinct mouth
and anus; most of them have a true, muscle-walled body cavity. And
(it turned out) they have an enlarged set of Hox genes. The traditional
view saw these morphological innovations as evolving by stages. For not
all bilaterians have a true body cavity. The platyhelminthes—the flat-
worms—do not. Moreover, there is a cluster of poorly known pseudo-
coelomate phyla whose body cavity is partial, and whose development
is embyrologically distinct from that of the true coelomates. So on this
classical view, these groups were evolutionary way stations on the road
to the full-deal bilaterians, which in turn fissured into two main groups
distinguished embryologically: the deuterostomes and protostomes.
The affinities of some phyla remain unclear, but molecular systemat-
ics has led to a consensus about the coarse-grained pattern of metazoan
evolution, and on how to fit much of the eccentric Cambrian fauna into
this picture. This new phylogeny casts doubt on the traditional view of
the stable and gradual development of metazoan body plans.
The picture of the base of the metazoan radiation has not changed.
However, within the bilaterian animals, molecular systematics has led
to a revolutionary change. On previous views of animal evolutionary
history, the evolution of the true body cavity—the “coelom”—was seen
as progressive and gradual. The animal phyla lacking a true coelom
(the acoelomate flatworms, the platyhelminthes) were seen as primi-
tive, resembling the Mother of All Bilaterians in this respect, and those
phyla with partial cavities likewise were seen as more primitive than
the true ceolomates. Molecular data have not supported this view. The
pseudo-coelomates are not a clade; they are secondarily simplified, as
are some of the flatworms. They are not a clade either; some of the
worms without body cavities are indeed primitively simple and are the
most ancient surviving split within the bilaterians, a sister group to all