22 chapter one

more stable, perhaps even more productive. It might even be that di-
verse global biotas hedge our bets against an uncertain future. But not
all descriptions of biological systems are forward-looking in this way.
In evolutionary theory, our interest in patterns in diversity is often mo-
tivated by the thought that differences in pattern are symptoms of dif-
ferences in process: biodiversity patterns are informative signals of the
processes that caused them. A good example of this is recent work on
phenotypic diversity in evolutionary biology. Within microevolutionary
studies, there is a long tradition that attempts to measure the strength
of competition for resources between similar species by seeing whether
competing species exhibit character displacement. For example, such
studies measure whether two species of anolis lizards that live together
on the same island are phenotypically different from populations of
those same lizards when they are not in contact.12 Divergence, if found,
is a trace of competition. The same is true of macroevolutionary studies.
For example, phenotype conservatism—no change over long periods of
time—is often taken to be a signal of constraint on the power of selec-
tion to shape new forms of life. We will meet this interpretation of the
evolution of the animals in 3.1.
Cladistic methodology is supposed to allow us to estimate the tree of
life13 while making only uncontentious assumptions about evolutionary
mechanisms. Even so, the whole point of identifying genealogical rela-
tionships is to zero in on evolutionary mechanisms. We cannot estimate
the extent to which Australian eucalypt phenotypes are adaptations to
their current environment without a phylogeny that tells us which of
their traits are shared derived inheritances from a pre-arid Australia,
and which are convergent or parallel adaptations to their new and
harder world. If eucalypts’ characteristically hard, waxy leaves evolved
before the great Australian drying, they cannot be an adaptation to that
drying. Identifying phylogeny is essential to understanding phenotypic
change.
Patterns in speciation are also signals of evolutionary process. Why
are there so many beetle species? How was it possible for the cichlids
in the east African lake systems to evolve so many species so fast? Pat-
terns in the overall shape of the tree of life are signals of the processes
that produce those patterns, and that is one reason why it’s important to
have a principled and objective characterization of those patterns. We
need a well-established phylogeny to show a clade is unusually species
rich or unusually morphologically diverse. Consider, for example, one
of Wallace Arthur’s striking examples of developmental constraint. As
a group, centipedes vary considerably in segment number. But among
the Lithobiomorpha centipedes there is no variation at all. All thousand