Principios de Taxonomia
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140j 6 Biological Species as a Gene-Flow Community<br />
Dupre, 1999). All traits of an organism must function in a concerted fashion. This<br />
view consi<strong>de</strong>rs the species as a balanced unit that is resistant to change. Selective<br />
pressure could thus be much more crucial than gene flow (Mallet, 2006).<br />
That selection could be a major factor in maintaining the homogeneity of a<br />
species along large geographical distances already follows from the fact that<br />
uniparental organisms also produce distinct groups of phenotypically similar<br />
individuals. If biparental gene flow were the main factor un<strong>de</strong>rlying the homogeneity<br />
of a species, then uniparental organisms could not generate large<br />
populations of similar individuals in the long term. Therefore, the constancy<br />
in phenotypic appearance of the individuals of a species must not result mainly<br />
from gene flow (Lan<strong>de</strong>, 1980; Mallet, 2006).<br />
Perhaps this phenomenon is comparable to vehicles, which also remain more or<br />
less similarly shaped in the longer term because they cannot freely vary in form.<br />
Trucks, cars and motorcycles have many traits that must be aligned, as they are<br />
nee<strong>de</strong>d to form a functioning whole (Barton, 1993), and this integrity of form is<br />
in<strong>de</strong>pen<strong>de</strong>nt of whether such a vehicle is required to function in the north or the<br />
south of Europe. In roadless terrain, however, other vehicle shapes prove superior,<br />
and four-wheel-drive, off-road vehicles and tracked vehicles are predominant.<br />
However, the assumption that selection is the main factor that maintains species<br />
homogeneity should not be overstressed. The divergent adaptations of organisms<br />
to local environments are evi<strong>de</strong>nt. Many traits are not homogeneously<br />
distributed across the entire range of a species. If this were not the case, there<br />
would be no races (Chapter 5).<br />
6.8<br />
Isolation by Distance<br />
When alleles do not reach geographically distant organisms and thus evolve in<strong>de</strong>pen<strong>de</strong>ntly,<br />
this phenomenon is called isolation by distance. Isolation by distance<br />
must be clearly distinguished from allopatry (see below). Two populations are<br />
consi<strong>de</strong>red allopatric if they are completely separated by external, usually geographical<br />
barriers, so that no gene flow is possible between them. If, however, two<br />
populations were geographically distant but still clinally connected by intermediary<br />
populations, then this situation would qualify as isolation by distance.<br />
Isolation by distance is problematic because the organisms of distant populations<br />
of a species can lose their ability to crossbreed (Sterelny and Griffiths, 1999). If race A<br />
overlaps in range with race B, and race B overlaps with race C, then race C will not<br />
necessarily be cross-fertile with race A (Figure 6.2a). The ability of individuals in<br />
distant populations to crossbreed may become lost <strong>de</strong>spite incremental clinal<br />
transitions along a chain of populations.<br />
An example of the <strong>de</strong>creasing ability of distant organisms of a species to crossbreed<br />
was found in the 1940s and early 1950s by the British evolutionist and lepidopterologist<br />
E. B. Ford (Ford, 1940; 1954). Ford showed by the example of the Satyrid<br />
butterfly Coenonympha tullia (Large Heath) that with an increasing geographical