Morphology and Morphological Diversity 63

in comparisons, for example, between protostomes (such as Opabinia)

and deuterostomes (such as human beings). Furthermore, reliance on
homology makes landmark analysis inherently historical. This makes it
a powerful tool for the analysis of change within a taxon over time (for
example, Eble 2000; Wagner 1995b; Vizcaíno and De Iuliis 2003) as
well as between geographically isolated populations of the same species
(for example, Hoffmann and Shirriffs 2002). Tools of this kind are quite
powerful, as Dan McShea’s work on the evolution of complexity shows.
There is a consensus in evolutionary biology that the complexity of life
has increased over time, though there is no consensus on the nature or
cause of that trend. McShea has made this question empirically trac-
table, by relativizing it to clades, and by developing an operational mea-
sure of complexity for homologous organ systems inherited throughout
a clade. The vertebrate system, for example, becomes more complex on
McShea’s operationalization if the number of individual parts (verte-
brae) increase, or if the parts become more differentiated, one from an-
other (McShea 1992; 1996). By tracing changes in such systems through
a lineage, and by developing these measures of complexity, McShea can
then ask whether, on average, organizational complexity in the skeletal
systems of vertebrates tends to increase over time. If in most clades and
most major organizational systems, complexity tends to increase, we
could then conclude that there really was an overall trend to increasing
complexity.
However, the same historical foundations render landmark analysis
less useful in a comparison of, for example, wing structure between
bees, birds, bats, and pterodactyls; their wings are not homologous,
each having evolved from a different morphological precursor (see ta-
ble 4.1). But perhaps the greatest drawback for those who would map
large-scale changes in disparity is the fact that the study of landmark
data is bound by actual form. It is necessarily derived from existing
morphology interpreted in light of known phylogeny. But shouldn’t
we be able to represent possible as well as actual morphologies? It
seems important to be able to represent morphologies we might have,
but do not, if we are to identify “gaps in nature.” Such gaps are organ-
ta b l e 4 . 1 : Wing Precursors in Four Distantly Related Taxa.

Taxon Wings evolved from

Insects Gill branches

Birds Arms
Bats Hands
Pterodactyls A single elongated digit