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Scientists have long recognized the incredible diversity within a species. But they thought it reflected evolutionary changes that unfolded imperceptibly, over millions of years. That divergence between populations within a species was enforced, according to Ernst Mayr, the great evolutionary biologist of the $1940$s, when a population was separated from the rest of the species by a mountain range or a desert, preventing breeding across the divide over geologic scales of time. Without the separation, gene flow was relentless. But as the separation persisted, the isolated population grew apart and speciation occurred

In the mid-$1960$s, the biologist Paul Ehrlich- author of The Population Bomb $(1968)$ - and his Stanford University colleague Peter Raven challenged Mary’s idea about speciation. They had studied checkerspot butterflies living in the Jasper Ridge Biological Preserve in California, and it soon became clear that they were not examining a single population. Through years of capturing, marking and then recapturing the butterflies, they were able to prove that within the population, spread over just $50$ acres of suitable checkerspot habitat, there were three groups that rarely interacted despite their very close proximity.

Among other ideas, Ehrlich and Raven argued in a now classic paper from $1969$ that gene flow was not as predictable and ubiquitous as Mayr and his cohort maintained, and thus evolutionary divergence between neighbouring groups in a population was probably common. They also asserted that isolation and gene flow were less important to evolutionary divergence than natural selection (when factors such as mate choice, weather, diseases or predation cause better-adapted individuals to survive and pass on their successful genetic traits). For example, Ehrlich and Raven suggested  that, without the force of natural selection, an isolated population would remain unchanged and that,  in other scenarios, natural selection could be strong enough to overpower gene flow.

Which of the following best sums up Ehrlich and Raven’s argument in their classic $1969$ paper?

1. Ernst Mayr was wrong in identifying physical separation as the cause of species diversity
2. Checkerspot butterflies in the $50$-acre Jasper Ridge Preserve formed three groups that rarely interacted with each other
3. While a factor, isolation was not as important to speciation as natural selection
4. Gene flow is less common and more erratic than Mayr and his colleagues claimed.

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