70 chapter four

assess, even if we represent the data optimally. Hence it is important to

get the patterns right: no pseudo-patterns; no pattern blindness. Hence
models like Raup’s are of great importance. In Raup’s space, the white
areas are those not populated by extant or extinct mollusks. We might
therefore naturally ask (as did Raup) why these regions of possibility
space have remained unpopulated. This in turn leads to an explora-
tion of selection pressures and developmental constraints that might
be responsible for the observed distributions of molluscan form. Such
studies have been remarkably productive both in the discovery of mor-
phological regularities and in the generation of novel hypotheses. We
discuss one case below: the interaction of ammonite and nautilus evo-
lution. A second common use of theoretical morphology is to plot the
occupation of morphospace over time to generate and test hypotheses
about the way in which clades move in response to long-term environ-
mental change. This gives us a sense of the rates and modes of change
of which different clades are capable. Mike Foote refers to this as inves-
tigating the “morphological exuberance” of clades (1997, 133). In The
Geometry of Evolution (2006), George McGhee makes a strong case for
the idea that such analysis will finally allow us to make operational the
idea of the adaptive landscape.
Crucial in the success of theoretical morphology has been the contin-
gent connection between morphological diversity and taxonomic diver-
sity; it allows us to investigate the relationship between these two forms
of biodiversity rather than assuming that species richness covaries with
phenotypic variation. This is especially important in the study of major
extinction events. By definition these extinguish large numbers of taxa,
but employing theoretical morphology, we can investigate the relation-
ship between taxonomic loss and morphological loss. In some cases
extinction events “merely thin the number of species present, without
having much effect on the total range of morphologies” (McGhee 1999,
190). In others, in cases of what David Raup calls “wanton” and “fair
game” extinctions4 (Raup 1991, 185–89), particular morphologies may
be targeted in a way that will lead to loss of morphological diversity as
well as taxonomic diversity.
The Cretaceous/Tertiary extinction event was both morphologically
and taxonomically selective. It extinguished all the existing species of
ammonites while taking a less severe toll on other similar marine mol-
lusks, including the nautilids. This is a tale that could have been told
by taxonomy. Theoretical morphology is not needed to identify the
differential impact of extinction on clades, but what happened next
we know about only through the efforts of theoretical morphologists.
Ammonites and nautilids are both univalve marine mollusks, and so