Species: A Modest Propasal 39

these source populations are not under selection to acquire or retain

these new genes. Furthermore, the demographic center of gravity usu-
ally consists of populations with the original phenotype of the species.
Thus, to the extent that an adapting population is well connected de-
mographically to the rest of the species, the local adaptations it acquires
are liable to be lost.
That said, demographic connections between metapopulations do
not invariably stabilize phenotypes. Some selective impacts will be of
the right spatial scale to generate change. Climatic and other changes in
the physical environment might well generate coarse-grained selective
forces, affecting all or most of the species’ populations. Tectonic and
other geological forces that alter the basic structure of the landscape
(uplift, erosion and deposition, sea level changes) may well exert con-
sistent effects over large areas. So changes that are adaptive throughout
the species’ range can become established, but as Futuyma notes (1987,
468), these changes are likely to be uncommon. Hence, the connection
between phenotype change and speciation.5 Speciation is not required
for phenotype change that adapts a population to specific local condi-
tions, but it is often required to make such changes permanent. The dis-
tribution of a species through an ecological mosaic, together with gene
flow between the fragments, acts as a brake on evolutionary change.
Stasis is not permanent, however, in part because environmental
change has the potential to release the evolutionary brake imposed by
metapopulation dynamics. Climate change and other large-scale physi-
cal changes can turn species mosaics into patchworks of isolated popu-
lations (see Bennett 1997; Eldredge 1995; Vrba 1993 and 1995). Some-
times the effects of environmental change will not be dramatic: the
potential space available to a species might shrink a bit, expand a bit, or
shift latitude. If physical barriers do not intervene, the species can shift
with it. However, stasis breaks down when environments both change
(creating new selection pressures) and a species’ range fragments, dis-
solving the metapopulation by chopping it into its component popula-
tions. A local, isolated population is not ecologically fragmented. The
vast majority of such small populations will go extinct. But if in these
fragments the population is not so small that genetic variation is sharply
reduced, selection can act, and act without counterbalance, from ho-
mogenizing gene flow from neighboring populations. For there are no
neighboring populations. While many fragments go extinct, a few will
survive as new species. Many species emerge through a life cycle: from
population ⇒ metapopulation ⇒ isolated population ⇒ incipient spe-
cies. This life cycle is itself one important mechanism of evolutionary
change. As many phenomenological species are the product of this life