200 CHAPTER 8
A very different outcome occurs when migration onto an island is much stron-
ger than selection (s << m). The frequency of the locally adapted allele evolves to
0, and the allele is lost entirely. This outcome, in which gene flow overwhelms
local adaptation, is called gene swamping. Like many insects, populations of the
mosquito Culex pipiens have evolved resistance to insecticides. On the island of
Corsica, however, resistance has not evolved, despite the presence of resistance
alleles at low frequency. The best explanation is gene swamping. Patches of habitat
on Corsica where insecticide is applied are small. Mosquitoes disperse into those
patches from neighboring populations that are not resistant, swamping the resis-
tance alleles that are favored inside the patches [19, 29].
The evolutionary tension between selection and gene flow plays out in continu-
ous habitats as well as on islands. Consider a grass growing in a prairie that has
two kinds of soil that meet at a sharp boundary. The soil to the west of the bound-
ary favors allele A 1 , while the soil to the east favors allele A 2. To be specific, say that
allele A 1 has a relative fitness advantage of s in the west, while A 2 has the same
advantage in the east, and there is no dominance. What happens? Without gene
f low, A 1 will become fixed everywhere to the west of the boundary, and A 2 will be
fixed everywhere to the east. But with gene flow, alleles move across the boundary.
This introduces into each habitat the allele that is not favored there, and a cline
develops (FIGURE 8.11). The cline can be short or long, depending on the relative
strengths of gene flow and selection. The width of the cline is
(8.4)
This is the width of region over which allele A 2 increases from a low frequency
(p = 0.1) to a high frequency (p = 0.9).
wc=2.5 σ^2 ms
Futuyma Kirkpatrick Evolution, 4e
Sinauer Associates
Troutt Visual Services
Evolution4e_08.10 .ai Date 11-17-2016 01-24-17
Lava
O’Neill
Xmas Tule West Mid East O’Neill
Arizona,
USA
Sonora, Mexico
50
km
Frequency of melanic allele^0
0.5
1
FIGURE 8.10 The rock pocket mouse (Chaeto- Light rock Dark lava Light rock
dipus intermedius) typically has light-colored fur
where it lives on light-colored granite, but is dark
colored where it lives on dark lava flows in Arizona.
The dark coloration, which results from a melanic
allele at the Mc1r locus, camouflages mice on the
lava from predators. Selection favoring the melanic
allele on the lava is intense, with a selection coef-
ficient estimated to be as high as s = 0.4. (After [12];
photos from [24].)
08_EVOL4E_CH08.indd 200 3/23/17 9:12 AM