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Evolution__3rd_Edition

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..<br />

Could postzygotic isolation evolve<br />

by changes at one locus?<br />

rarely arranged in a ring. But several examples do exist (Irwin et al. 2001b). The herring<br />

gull (Larus argentatus) and lesser black-backed gull (L. fuscus) of northern Europe are a<br />

ring species, connected by a ring of populations around the North Pole. In central Asia,<br />

the warbler Phylloscopus trochiloides is distributed in a ring around the treeless Tibetan<br />

Plateau. The species appears to have originated in the south, in India. It then colonized<br />

northwards, either side of the Tibetan Plateau. The colonizing populations evolved<br />

slight genetic differences, and the songs diverged between the eastern and western<br />

populations. Eventually, these two lines of populations meet up again in central<br />

Siberia, but here the songs are so different that the members of the east population do<br />

not interbreed with members of the western population (Irwin et al. 2001a). And on<br />

the tiny Pacific island of Moorea, certain species of snail in the genus Partula had<br />

evolved ring species around individual small mountains (Murray & Clarke 1980). Alas,<br />

the Partula of Moorea are extinct. They all succumbed in the 1980s and 1990s, in yet<br />

another unnecessary manmade ecological disaster (Tudge 1992).<br />

14.3.4 Speciation as a by-product of divergence is well documented<br />

In summary, we have abundant evidence that reproductive isolation evolves as a byproduct<br />

of divergence between geographically separate populations. If populations are<br />

experimentally kept in different conditions in the lab, those populations evolve adaptations<br />

to their conditions a and reproductive isolation also evolves as a consequence. If<br />

separate populations adapt to different local conditions in nature, they also evolve<br />

reproductive isolation as a consequence. For prezygotic isolation, we saw that the genetic<br />

basis is probably pleiotropy or hitch-hiking. However, that explanation is mainly hypothetical<br />

at present, because little genetic research has been done on prezygotic isolation.<br />

We can turn now to the genetics of postzygotic isolation. It has been the subject of<br />

much genetic research, and is well understood both empirically and theoretically.<br />

14.4 The Dobzhansky–Muller theory of postzygotic isolation<br />

14.4.1 The Dobzhansky–Muller theory is a genetic theory of postzygotic<br />

isolation, explaining it by interactions among many gene loci<br />

What kind of genetic changes give rise to postzygotic isolation? Postzygotic isolation<br />

means that hybrid offspring are produced but they either die before breeding or live<br />

and are sterile. In this section, we shall see that postzygotic isolation is likely to be<br />

caused, genetically, by an interaction between the genotypes at multiple loci, rather<br />

than by genotypes at a single locus. The simplest hypothetical genetic control would<br />

have one genetic locus, and the fitnesses of the genotypes would be as follows:<br />

Species 1 Hybrid Species 2<br />

One-locus genotype AA Aa aa<br />

Fitness High Zero High<br />

CHAPTER 14 / Speciation 389

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