Morphology and Morphological Diversity 69

co-location implies similarity (with regard to the relevant parameters)

just as spatial co-location implies spatial proximity. Thus by plotting the
location of organisms in these morphospaces, we can reveal patterns in
phenotype evolution. This process can reveal both sparsely and densely
populated regions in a morphospace, both of which need explanation.
We can even map the “directions” organisms take over developmental
time and lineages take over evolutionary time. In the next section, we
discuss a series of examples that illustrate the power of this way of rep-
resenting actual and possible morphological diversity. More generally,
over the next few sections we aim to show that (i) disciplined com-
parison between different phenotypes is indeed possible, and hence the
phenotypic disparity found in a biota is both well-defined and biologi-
cally important; (ii) phenotype diversity is at best imperfectly indexed
by species richness; but (iii) phenotype disparity is not conceptually
independent of taxonomic diversity.

4.4 the power of morphospaces

One of the oldest and still most well known theoretical morphospaces
is David Raup’s cube, developed in 1966 for the analysis of shell coiling
in mollusks. Raup’s model rests on four parameters. Whorl expansion
rate (W) is the rate at which the aperture of the shell opens out. The
distance of the generating curve from the axis (D) is the rate at which
the shell uncoils. The translation of the generating curve (T) is the rate
at which the uncoiling shell is pushed upward from the coiling plane.
The shape of the generating curve (S) is the outline of the growing edge
of the shell (here assumed to be constant). Figure 4.4 represents T, W,
and D in its x, y, and z axes. This is a genuine theoretical morphospace
because, while it is populated by many known morphologies, it places
them within a larger context of biological possibility. The space encom-
passes known forms, but its dimensions are not derived mathemati-
cally from any particular sample of shelled organisms. There are many
potential applications for such morphospaces. The following are a few
representative examples (for a more complete discussion of the value
of this methodology, see Maclaurin 2003).
Morphological biodiversity is immensely important in discovering
evolutionary mechanism. While evolutionary mechanisms generate
both patterns in developmental mechanism and speciation, we often
have access to these other consequences of evolutionary change only via
their effects on morphology. Evolutionary biology is compelled to trade
in inferences from morphological patterns in space and time to the
mechanisms that produce those patterns. These inferences are hard to