74 chapter four

fitness than our current location.) But as the number of demands on our
plant morphologies expands, the range of equally fit forms also expands.
Hence our walks can end in more and more locations. All are more-
or-less equally fit, but each makes different compromises (none of these
compromise morphologies are as good at any one of these tasks as a de-
sign optimal for just that task). The more demands on the morphology,
the greater the distance between the all-rounder’s performance and the
specialist performance for each particular task (Niklas 1994; 2002).
Niklas is careful to emphasize the preliminary nature of these re-
sults. The fitness assumptions are idealized. Nutrient acquisition is left
out of the model, and the assumption that each functional requirement
contributes equally and independently to overall fitness is obviously
unrealistic. In some environments, light energy is readily available but
water is not. In others, the reverse is true. Even so, the morphologies
found in these multiply constrained walks—walks in which plant mor-
phology must compromise between all these functional demands—are
similar to the morphologies found by evolution in the radiation of
the land plants. This is an intriguing model, opening up the possibil-
ity of a coevolutionary interaction between ecological complexity and
morphological disparity. Niklas’s point about the expansion of equi-fit
morphologies as distinct adaptive demands expands seems to be quite
general (Marshall 2006). Nonetheless, we are agnostic about the extent
to which this model explains plant diversification.
While we are agnostic about the extent to which this model explains
plant diversity, we do want to underline an important element of it: the
use of phylogenetic information to control our choice of dimensions for
the morphospace and its point of origin. We think this constraint on the
choice of dimensions is crucial to the use of morphospaces to generate
testable ideas about the biological world. Likewise, it is crucial to have
a principled point of origin for those models—like Nilsson and Pelger’s
model of eye evolution—that explore specific evolutionary trajectories.
This in turn has implications for the relationship between phylogenetic
diversity and morphological diversity. So far in this chapter, and in the
last, we have been emphasizing the imperfect correlation between mor-
phological disparity and species richness, even when we add in how
species are sorted into clades, as in studies of mass extinction and adap-
tive radiation. That idea stands. But the dimensions of similarity and
difference that are biologically relevant are clade-specific. In discuss-
ing phenetics in 1.2, we argued that questions of phenotype similar-
ity and difference are ill defined unless we can answer, in a principled
way: similar or different in what ways? Our measures of disparity are
conceptually and theoretically dependent on phylogentically organized