34 chapter two

ideas are neutral on the mechanisms of speciation, they could be tied

to further claims about the process or processes that give rise to that
structure. However, there seems not to be a single mechanism respon-
sible for lineage bifurcation. Indeed, one natural interpretation of much
of the species debate is that it reflects our increasing knowledge of the
many mechanisms underlying diversity and differentiation.
John Wilkins has developed a helpful way of thinking about this
diversity of mechanisms and the relationship between them: a three-
dimensional conceptual space (2007). One dimension represents the
role of chance. Sir Ronald Fisher and Sewall Wright famously debated
the role of genetic drift and other chance factors in generating the di-
vergence between populations in a sundering lineage. For example, in
vicariant models of speciation a widely distributed ancestral popula-
tion is divided into fragments by geological changes. These fragments
then diverge, and chance is important as they wander morphologically
away from one another. So if this model is important, chance plays an
important role in much speciation. A second dimension concerns the
relative role of intrinsic and external factors when selection does drive
differentiation. For example, if hybrids between two subpopulations
are less fit, then there will be selection of traits that cause like to mate
with like. Features of the evolving population itself shape the selective
environment. In contrast, on Mayr’s peripheral isolate model, selection
will drive differentiation due to external environmental differences be-
tween the center of the species’ range and the periphery. Wilkins’s third
dimension focuses on the role of gene flow and barriers to that flow.
Mayr, famously, argued for the importance of geographic isolation in
the evolution of differentiation. But there are many models of specia-
tion that allow speciation to take place without geographic isolation;
for example, speciation that involves host switching by parasites, and
speciation that involves hybridization or chromosomal reorganization
(a mechanism quite common in plants).
We will illustrate these points about the diversity of mechanism
through a brief discussion of ecological and biological species concepts.
As usual, the picture is complex. Some ecological species concepts are
deliberately agnostic as to the details of the processes that give rise to
speciation. For example, Alan Templeton’s cohesion species concept
takes cohesion to be crucial in the production and maintenance of spe-
cies, but he accepts that there are many biological processes that gener-
ate cohesion. Leigh van Valen’s ecological species concept ties species
to niche occupation. However, the relationship between species and
niches is very complex. It was once supposed that communities were
organized in ways that made a variety of roles or occupations available