Industrial melanism

Industrial melanism was first noticed in 1900 by the geneticist William Bateson; he observed that the colour morphs were inherited, but did not suggest an explanation for the polymorphism.^[1][6]

In 1906, the geneticist Leonard Doncaster described the increase in frequency of the melanic forms of several moth species from about 1800 to 1850 in the heavily industrialised north-west region of England.^[7]

In 1924, the evolutionary biologist J. B. S. Haldane constructed a mathematical argument showing that the rapid growth in frequency of the carbonaria form of the peppered moth, Biston betularia, implied selective pressure.^[8][9]

From 1955 onwards, the geneticist Bernard Kettlewell conducted a series of experiments exploring the evolution of melanism in the peppered moth. He used a capture-mark-recapture technique to show that dark forms survived better than light ones.^{[5][10][11][12][13]}

By 1973, pollution in England had begun to decrease, and the dark carbonaria form had declined in frequency. This provided convincing evidence, gathered and analysed by Kettlewell and others such as the entomologist and geneticist Michael Majerus and the population geneticist Laurence M. Cook, that its rise and fall had been caused by natural selection in response to the changing pollution of the landscape.^[14][15][16]

Industrial melanism is known from over 70 species of moth that Kettlewell found in England, and many others from Europe and North America.^[17] Among these, Apamea crenata (clouded border brindle moth) and Acronicta rumicis (knot grass moth) are always polymorphic, though the melanic forms are more common in cities and (like those of the peppered moth) are declining in frequency as those cities become less polluted.^[1]

Among other insects, industrial melanism has been observed in a beetle (Adalia bipunctata, the two-spot ladybird^[18]) and a barklouse (Mesopsocus unipunctatus^[19]).

In the vertebrates, industrial melanism is known from the turtle-headed seasnake Emydocephalus annulatus,^[20] and may be present in urban feral pigeons.^[21]

Biston betularia, the peppered moth, in typical and melanic forms

Originally, peppered moths lived where light-colored lichens covered the trees. For camouflage from predators against that clean background, they had generally light coloration.^[18] During the Industrial Revolution in England, sulphur dioxide pollution in the atmosphere reduced the lichen cover, while soot blackened the bark of urban trees, making the light-colored moths more vulnerable to predation. This provided a selective advantage to the gene responsible for melanism, and the darker-colored moths increased in frequency.^[22][23] The melanic phenotype of Biston betularia has been calculated to give a fitness advantage as great as 30 per cent.^[24] By the end of the 19th century it almost completely replaced the original light-coloured type (var. typica), forming a peak of 98% of the population in 1895.^[25]

Melanic B. betularia have been widely observed in North America. In 1959, 90% of B. betularia in Michigan and Pennsylvania were melanic. By 2001, melanism dropped to 6% of the population, following clean air legislation.^[26] The drop in melanism was correlated with an increase in species diversity of lichens, a decrease in the atmospheric pollutant sulphur dioxide, and an increase in the pale phenotype. The return of lichens is in turn directly correlated with the reduction in atmospheric sulphur dioxide.^[27]

An additional study in 2018 further quantified survivability by looking at color and luminance camouflage and avian artificial predation models. For color camouflage, typica moths blended better under lichen bark than carbonaria, but when placed under plain bark, there was no significant difference. However, in luminance camouflage, carbonaria moths blended better compared to typica on a plain bark tree. When both variants were placed on an unpolluted lichen covered tree, typica moths had a 21% better survival rate.^[28]

Kettlewell's experiments were criticised by the zoologist Theodore David Sargent, who failed to reproduce Kettlewell's results between 1965 and 1969, and argued that Kettlewell had specially trained his birds to give the desired results.^{[29][30][31][32]} Michael Majerus however found that Kettlewell was basically correct in concluding that differential bird predation in a polluted environment was the primary cause of industrial melanism in the peppered moth.^[33] The story was in turn taken up in a 2002 book Of Moths and Men, by the journalist Judith Hooper, asserting that Kettlewell's findings were fraudulent.^[34] The story was picked up by creationists who repeated the assertions of fraudulence.^[35] Zoologists including L. M. Cook, B. S. Grant, Majerus and David Rudge however all upheld Kettlewell's account, finding that each of Hooper's and the creationists' claims collapsed when the facts were examined.^{[3][33][36][37][38][39]}

It has been suggested that the demonstrated relationship between melanism and pollution can not be fully proven because the exact reason for increase in survivability can not be tracked and pin-pointed. However, as air quality has improved in industrial areas of America and Britain, through improved regulation, offering the conditions for a natural experiment, melanism has sharply declined in moths including B. betularia and Odontopera bidentata. Cook and J. R. G. Turner have concluded that "natural selection is the only credible explanation for the overall decline",^[1] and other biologists working in the area concur with this judgement.^[40]