Industrial Melanism: The Case of the Peppered Moth

The idea of natural selection is that organisms in a species that have characteristics favoring survival are most likely to survive and produce offspring with the same characteristics. Because the survival of organisms with particular characteristics is favored over the survival of other organisms in the same species that lack these characteristics, future generations of the species are likely to include more organisms with the favorable characteristics.

One of the most thoroughly analyzed examples of natural selection in operation is the change in color that has occurred in certain populations of the peppered moth, Biston betularia, in industrial regions of Europe during the past 100 years. Originally moths were uniformly pale gray or whitish in color; dark-colored (melanic)individuals were rare and made up less than 2 percent of the population. Over a period of decades, dark-colored forms became an increasingly large fraction of some populations and eventually came to dominate peppered moth populations in certain areas-especially those of extreme industrialization such as the Ruhr Valley of Germany and the Midlands of England. Coal from industry released large amounts of black soot into the environment, but the increase of the dark-colored forms was not due to genetic mutations caused by industrial pollution. For example, caterpillars that feed on soot-covered leaves did not give rise to dark-colored adults. Rather, pollution promoted the survival of dark forms on soot-covered trees. Melanics were normally quickly eliminated in nonindustrial areas by adverse selection; birds spotted them easily. This phenomenon, an increase in the frequency of dark-colored mutants in polluted areas, is known as industrial melanism. The North American equivalent of this story is another moth, the swettaria form of Biston cognataria, first noticed in industrialized areas such as Chicago and New York City in the early 1900s. By 1961 it constituted over 90 percent of the population in parts of Michigan.

The idea that natural selection was responsible for the changing ratio of dark- to light-colored peppered moths was developed in the 1950s by H.B.D.Kettlewell of Oxford University. If natural selection was the explanation, then there should be different survival rates for dark- and light-colored moths. To determine whether this was true, Kettlewell released thousands of light and dark moths (each marked with a paint spot) into rural and industrialized areas. In the nonindustrial area of Dorset, he recaptured 14.6 percent of the pale forms but only 4.7 percent of the dark forms. In the industrial area of Birmingham, the situation was reversed: 13 percent of pale forms but 27.5 percent of dark forms were recaptured.

Clearly some environmental factor was responsible for the greater survival rates of dark moths. Birds were predators of peppered moths. Kettlewell hypothesized that the normal pale forms are difficult to see when resting on lichen-covered trees, whereas dark forms are conspicuous. In industrialized areas, lichens are destroyed by pollution, tree barks become darker, and dark moths are the ones birds have difficulty detecting. As a test, Kettlewell set up hidden observation positions and watched birds voraciously eat moths placed on tree trunks of a contrasting color. The action of natural selection in producing a small but highly significant step of evolution was seemingly demonstrated, with birds as the selecting force.

Not every researcher has been convinced that natural selection by birds is the only explanation of the observed frequencies of dark and light peppered moths. More recent data, however, provide additional support for Kettlewell's ideas about natural selection. The light-colored form of the peppered moth is making a strong comeback. In Britain, a Clean Air Act was passed in 1965. Sir Cyril Clarke has been trapping moths at his home in Liverpool, Merseyside, since 1959. Before about 1975, 90 percent of the moths were dark, but since then there has been a steep decline in melanic forms, and in 1989 only 29.6 percent of the moths caught were melanic. The mean concentration of sulphur dioxide pollution fell from about 300 micrograms per cubic meter in 1970 to less than 50 micrograms per cubic meter in 1975 and has remained fairly constant since then. If the spread of the light-colored form of the moth continues at the same speed as the melanic form spread in the last century, soon the melanic form will again be only an occasional resident of the Liverpool area.