Morphology and Morphological Diversity 75

species richness. Diversity understood through phylogeny enables us to

identify the similarities and differences to count. We develop this point
further in the next section.
There is no doubt that theoretical morphology provides us with a
new and powerful tool for the analysis of evolutionary change. The
discoveries just listed would have been impossible without it. But in
the final sections of this chapter we explore the power of theoretical
morphology when we extend its scope beyond that of tracking closely
related taxa sharing small numbers of distinct parameters. We run into
its limits, for we no longer have a principled account of the dimensions
of similarity and difference.

4.5 here there be no dragons: the limits of theoretical

morphology

In 4.4, we illustrated the use of morphospace to represent the morpho-

logical diversity of clades at and over time. These representations are
mathematically tractable, theoretically principled, and biologically im-
portant. Thus in a famous series of studies, Joan Roughgarden studied
the morphology of anolis lizards on Caribbean islands, comparing liz-
ards that were alone on an island with those that shared an island with
congeneric species. The idea was to test a hypothesis about character
displacement. Namely, when congeneric lizards share an island they
evolve away from one another, to minimize comparative interactions.
Studies of this kind involve what are known as empirical morphospaces,
and they form part of the wider study of actual (as opposed to possible)
biological form known as morphometrics. Their dimensions are derived
from observations of actual organisms. Sometimes the derivation is not
direct. The dimensions of Raup’s cube might not be “derived” from any
particular mollusks. But clearly the power of the model stems precisely
from the fact that it has been developed with molluscan architecture in
mind. So these dimensions, too, have been determined by organisms.
The same is true of the space of possible skeletal form developed by
Thomas and Reif (1993). Here too the possibility space is based on the
range of forms found, without it being closely tied to a particular clade
or a particular statistical technique for detecting variation within the
clade. As with Raup’s shell space, this skeletal space represents plenty
of merely possible morphological forms (fig. 4.6).
For reasons of both theoretical coherence and empirical tractability,
it is important to have a constrained and principled choice of dimensions.
More than anything else, the development of theoretical morphospaces
has been a means of investigating adaptationist hypotheses. These are