Unit Four. The Evolution and Diversity of Life
14.11. Peppered Moths and Industrial Melanism
The peppered moth, Biston betularia, is a European moth that rests on tree trunks during the day. Until the mid-19th century, almost every captured individual of this species had light-colored wings. From that time on, individuals with dark-colored wings increased in frequency in the moth populations near industrialized centers, until they made up almost 100% of these populations. Dark individuals had a dominant allele that was present but very rare in populations before 1850. Biologists soon noticed that in industrialized regions where the dark moths were common, the tree trunks were darkened almost black by the soot of pollution. Dark moths were much less conspicuous resting on them than light moths were. In addition, air pollution that was spreading in the industrialized regions had killed many of the light-colored lichens on tree trunks, making the trunks darker.
Selection for Melanism
Can Darwin’s theory explain the increase in the frequency of the dark allele? Why did dark moths gain a survival advantage around 1850? An amateur moth collector named J. W. Tutt proposed in 1896 what became the most commonly accepted hypothesis explaining the decline of the light-colored moths. He suggested that light forms were more visible to predators on sooty trees that have lost their lichens. Consequently, birds ate the peppered moths resting on the trunks of trees during the day. The dark forms, in contrast, were at an advantage because they were camouflaged (figure 14.28). Although Tutt initially had no evidence, British ecologist Bernard Kettlewell tested the hypothesis in the 1950s by rearing populations of peppered moths with equal numbers of dark and light individuals. Kettlewell then released these populations into two sets of woods: one, near heavily polluted Birmingham, the other, in unpolluted Dorset. Kettlewell set up traps in the woods to see how many of both kinds of moths survived. To evaluate his results, he had marked the released moths with a dot of paint on the underside of their wings, where birds could not see it.
Figure 14.28. Tutt's hypothesis explaining industrial melanism.
Color variants of the peppered moth (Biston betularia). Tutt proposed that the dark moth is more visible to predators on unpolluted trees (top), while the light moth is more visible to predators on bark blackened by industrial pollution (bottom).
In the polluted area near Birmingham, Kettlewell trapped 19% of the light moths, but 40% of the dark ones. This indicated that dark moths had a far better chance of surviving in these polluted woods where the tree trunks were dark. In the relatively unpolluted Dorset woods, Kettlewell recovered 12.5% of the light moths but only 6% of the dark ones. This indicated that where the tree trunks were still light-colored, light moths had a much better chance of survival. Kettlewell later solidified his argument by placing dead moths on trees and filming birds looking for food. Sometimes the birds actually passed right over a moth that was the same color as its background.
Industrial melanism is a term used to describe the evolutionary process in which darker individuals come to predominate over lighter individuals since the industrial revolution as a result of natural selection. The process is widely believed to have taken place because the dark organisms are better concealed from their predators in habitats that have been darkened by soot and other forms of industrial pollution, as suggested by Kettlewell.
Dozens of other species of moths have changed in the same way as the peppered moth in industrialized areas throughout Eurasia and North America, with dark forms becoming more common from the mid-19th century onward as industrialization spread.
Selection Against Melanism
As of the second half of the 20th century, with the widespread implementation of pollution controls, these trends are reversing, not only for the peppered moth in many areas in England but also for many other species of moths throughout the northern continents. These examples provide some of the best-documented instances of changes in allelic frequencies of natural populations as a result of natural selection due to specific factors in the environment.
In England, the air pollution promoting industrial melanism began to reverse following enactment of Clean Air legislation in 1956. Beginning in 1959, the Biston population at Caldy Common outside Liverpool has been sampled each year. The frequency of the melanic (dark) form dropped from a high of 94% in 1960 to a low of 19% in 1995 (figure 14.29). Similar reversals have been documented at numerous other locations throughout England. The drop correlates well with a drop in air pollution, particularly with tree-darkening sulfur dioxide and suspended particulates.
Figure 14.29 Selection against melanism.
The circles indicate the frequency of melanic Biston betularia moths at Caldy Common in England, sampled continuously from 1959 to 1995. Red diamonds indicate frequencies of melanic B. betularia in Michigan from 1959 to 1962 and from 1994 to 1995.
Interestingly, the same reversal of industrial melanism appears to have occurred in America during the same time that it was happening in England. Industrial melanism in the American subspecies of the peppered moth was not as widespread as in England, but it has been well documented at a rural field station near Detroit. Of 576 peppered moths collected there from 1959 to 1961, 515 were melanic, a frequency of 89%. The American Clean Air Act, passed in 1963, led to significant reductions in air pollution. Resampled in 1994, the Detroit field station peppered moth population had only 15% melanic moths! The moths in Liverpool and Detroit, both part of the same natural experiment, exhibit strong evidence of natural selection.
Reconsidering the Target of Natural Selection
Tutt’s hypothesis, widely accepted in the light of Kettlewell’s studies, is currently being reevaluated. The problem is that the recent selection against melanism does not appear to correlate with changes in tree lichens. At Caldy Common, the light form of the peppered moth began its increase in frequency long before lichens began to reappear on the trees. At the Detroit field station, the lichens never changed significantly as the dark moths first became dominant and then declined over the last 30 years. In fact, investigators have not been able to find peppered moths on Detroit trees at all, whether covered with lichens or not. Wherever the moths rest during the day, it does not appear to be on tree bark. Some evidence suggests they rest on leaves on the treetops, but no one is sure.
The action of selection may depend on other differences between light and dark forms of the peppered moth as well as their wing coloration. Researchers report, for example, a clear difference in their ability to survive as caterpillars under a variety of conditions. Perhaps natural selection is also targeting the caterpillars rather than the adults. While we can’t yet say exactly what the targets of selection are, researchers are actively investigating this, one of the best-documented instances of natural selection in action.
Natural Selection for Melanism in Mice
Melanism is not restricted to insects. Cats and many other mammals have melanic forms that are subject to natural selection in much the same way as moths. The coat color of desert pocket mice that live on differently colored rock habitats provides a clear-cut example of natural selection acting on melanism. In Arizona and New Mexico, these small, wild pocket mice live in isolated black volcanic lava beds and the pale soils between them. Melanin synthesis during hair development of pocket mice is regulated by the receptor gene MC1R. Mutations that disable MC1R lead to melanism. Such mutations are dominant alleles, so whenever they are present in a population, dark pocket mice are seen. When wild populations of pocket mice were surveyed by biologists from the University of Arizona, there was a striking correlation between coat color and the color of the rock on which the population of pocket mice lived.
As you can see in the two upper photographs, the close match between coat color and background color gives the mice cryptic protection from avian predators, particularly owls. These mice are very visible when placed in the opposite habitats (lower photos).
Key Learning Outcome 14.11. Natural selection has favored the dark form of the peppered moth in areas subject to severe air pollution, perhaps because on darkened trees they are less easily seen by motheating birds. Selection has in turn favored the light form as pollution has abated.