Morphology and Morphological Diversity 79

The Library of Mendel is bad enough; the Library of D’Arcy Thomp-

son (the space of all possible morphologies) is worse. The Library of
Mendel is at least anchored by the base sequences. We might stipulate
that all possible genomes are sequences of the four bases, even though
we cannot specify in advance the possible ways those sequences can be
used in the generation of phenotypes, or how sequences can be grouped
into genes. The morphology of organisms (or even that of animals) does
not have this common currency. We cannot even specify a catalogue
of all possible cell types out of which organisms could be built. Any
theoretical morphospace must be anchored in actual organisms and be
underpinned by some scientific goal, be it mapping the evolution of a
clade, the effects of an extinction event, or the innovations in an eco-
logical guild. It is these purposes that give morphospaces their dimen-
sions. It is tempting to think that the availability and utility of these
partial morphospaces—morphospaces that represent some aspects of
the morphology of a particular group of organisms—implies that there
is a global morphospace that represents all the ways organisms might
have been, and locates each actual organism in that space. We think that
temptation should be resisted. That space is as ill defined as the library
of all possible books.

4.6 morphological biodiversity

Over the last few chapters we have been exploring the relationship be-
tween evolutionary differentiation expressed in the speciation pattern
of clades and morphological differentiation. We saw in chapters 2 and 3
that these patterns are at least partly independent: it is possible for
a clade to be species rich without extensive differentiation, and it is
possible for a clade to be species poor, but with species markedly dif-
ferentiated from their (surviving) sister taxa. For that reason, species
richness is not invariably a good surrogate for morphological disparity.
Even so, there is an intimate relationship between phylogenetic and
morphological biodiversity. In part, that relationship is causal. We can
have speciation without differentiation, and that is one reason why a
clade can be species rich without being morphologically diverse. But
on the Futuyma-Eldredge model (Futuyma 1987; Eldredge 1995, 2003),
speciation is often necessary for morphological differentiation.
There is a second relationship as well. In 1.2, in discussing phenet-
ics, we pointed out that similarity and difference are undefined; we
must talk, instead, of similarity with respect to one or more character
states. In discussing morphospace, this issue reappears as the choice of
dimensions. A global morphospace is as undefined as the idea of overall