The evolution of the peppered moth is an evolutionary instance of directional colour change in the moth population as a consequence of air pollution during the Industrial Revolution in England in the 19th century. The frequency of dark-coloured moths increased at that time, an example of industrial melanism. Later, when pollution was reduced in response to clean air legislation, the light-coloured form again predominated. Industrial melanism in the peppered moth was an early test of Charles Darwin's natural selection in action, and it remains a classic example in the teaching of evolution. In 1978, Sewall Wright described it as "the clearest case in which a conspicuous evolutionary process has actually been observed." The dark-coloured or melanic form of the peppered moth (var. carbonaria) was rare, though a specimen had been collected by 1811. After field collection in 1848 from Manchester, an industrial city in England, the frequency of the variety was found to have increased drastically. By the end of the 19th century it almost completely outnumbered the original light-coloured type (var. typica), with a record of 98% in 1895. The evolutionary importance of the moth was only speculated upon during Darwin's lifetime. It was 14 years after Darwin's death, in 1896, that J. W. Tutt presented it as a case of natural selection. Because of this, the idea spread widely, and more people came to believe in Darwin's theory. Bernard Kettlewell was the first to investigate the evolutionary mechanism behind peppered moth adaptation, between 1953 and 1956. He found that a light-coloured body was an effective camouflage in a clean environment, such as in rural Dorset, while the dark colour was beneficial in a polluted environment like industrial Birmingham. This selective survival was due to birds, which easily caught dark moths on clean trees and white moths on trees darkened with soot. The story, supported by Kettlewell's experiment, became the canonical example of Darwinian evolution and evidence for natural selection used in standard textbooks. However, failure to replicate the experiment and Theodore David Sargent's criticism of Kettlewell's methods in the late 1960s led to general skepticism. When Judith Hooper's Of Moths and Men was published in 2002, Kettlewell's story was more sternly attacked, and accused of fraud. The criticism became a major argument for creationists. Michael Majerus was the principal defender of Kettlewell's original experiment. His seven-year experiment beginning in 2001, the most elaborate of its kind in population biology, the results of which were published posthumously in 2012, vindicated Kettlewell's work in great detail. This restored peppered moth evolution as "the most direct evidence [of evolution]", and "one of the clearest and most easily understood examples of Darwinian evolution in action".
Origin and evolution
Before the Industrial Revolution, the black form of the peppered moth was rare. The first black specimen (of unknown origin) was collected before 1811, and kept in the University of Oxford. The first live specimen was caught by R. S. Edleston in Manchester, England in 1848, but he reported this only 16 years later in 1864, in The Entomologist. Edleston notes that by 1864 it was the more common type of moth in his garden in Manchester. The light-bodied moths were able to blend in with the light-coloured lichens and tree bark, and the less common black moths were more likely to be eaten by birds. As a result of the common light-coloured lichens and English trees, therefore, the light-coloured moths were much more effective at hiding from predators, and the frequency of the dark allele was very low, at about 0.01%. During the early decades of the Industrial Revolution in England, the countryside between London and Manchester became blanketed with soot from the new coal-burning factories. Many of the light-bodied lichens died from sulphur dioxide emissions, and the trees became darkened. This led to an increase in bird predation for light-coloured moths, as they no longer blended in as well in their polluted ecosystem: indeed, their bodies now dramatically contrasted with the colour of the bark. Dark-coloured moths, on the other hand, were camouflaged very well by the blackened trees. The population of dark-coloured moth rapidly increased. By the mid-19th century, the number of dark-coloured moths had risen noticeably, and by 1895, the percentage of dark-coloured moths in Manchester was reported at 98%, a dramatic change (of almost 100 times) from the original frequency. This effect of industrialization in body colour led to the coining of the term "industrial melanism". The implication that industrial melanism could be evidence supporting Charles Darwin's theory of natural selection was noticed during his lifetime. Albert Brydges Farn (1841–1921), a British entomologist, wrote to Darwin on 18 November 1878 to discuss his observation of colour variations in the Annulet moth (then Gnophos obscurata, now Charissa obscurata). He noted the existence of dark moths in peat in the New Forest, brown moths on clay and red soil in Herefordshire, and white moths on chalk cliffs in Lewes, and suggested that this variation was an example of "survival of the fittest". He told Darwin that he had found dark moths on a chalk slope where the foliage had been blackened by smoke from lime kilns, and he had also heard that white moths had become less common at Lewes after lime kilns had been in operation for a few years. Darwin does not seem to have responded to this information, possibly because he thought natural selection would be a much slower process. A scientific explanation of moth coloration was only published in 1896, 14 years after Darwin's death, when J. W. Tutt explicitly linked peppered moth melanism to natural selection.
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