Peppered moth
A moth studied as an example of natural selection.
The peppered moth (Biston betularia) belongs to the Geometridae family. It flies at night and lives mostly in the temperate parts of the northern hemisphere. Its evolution is a well-known case study in population genetics and natural selection.
The caterpillars of this moth mimic both the shape and color of twigs. Recent research shows they can detect a twig’s color through their skin and adjust their own body color to match the background, helping them avoid predators.
**Description** The moth’s wingspan is 45 to 62 mm, with an average of 55 mm. It has a fairly stout body and narrow, elongated forewings. The wings are white with black speckling, plus more or less distinct black cross lines. Rarely, the speckling and lines can be gray or brown, and in very rare cases, a mix of brown and black or gray. The amount of black speckling varies widely—some individuals have almost none, while others are so heavily marked the wings look black with white spots. Males have strongly comb-like antennae. Prout (1912–16) described the various forms and related species.
**Distribution** This moth occurs in China (many provinces including Heilongjiang, Jilin, Inner Mongolia, Beijing, Hebei, Shanxi, Shandong, Henan, Shaanxi, Ningxia, Gansu, Qinghai, Xinjiang, Fujian, Sichuan, Yunnan, Tibet), Russia, Mongolia, Japan, North and South Korea, Nepal, Kazakhstan, Kyrgyzstan, Turkmenistan, Georgia, Azerbaijan, Armenia, Europe, and North America.
**Ecology and Life Cycle** In Great Britain and Ireland, the peppered moth has one generation per year. In southeastern North America, it has two. Its life cycle has four stages: eggs, several caterpillar stages, pupae (which overwinter in the soil), and adults. During the day, the moths rest on trees, where birds prey on them. The caterpillar mimics a twig and can be green or brown. In 1887, Edward Bagnall Poulton noted it as one of the first animals known to use countershading for camouflage, making it appear flat. The caterpillar senses a twig’s color with its skin and changes its own color to match—a slower version of the camouflage seen in cephalopods, chameleons, and some fish. Late in the season, it enters the soil to pupate for winter. Adults emerge from pupae between late May and August, with males emerging slightly before females. They emerge late in the day, dry their wings, and fly that night. Males fly every night searching for females, while females fly only on their first night. After that, females release pheromones, which males follow upwind. During flight, bats prey on them. Males guard females from other males until eggs are laid. Females lay about 2,000 pale-green, oval eggs, each about 1 mm long, into bark crevices using their ovipositor.
**Resting Behavior** A mating pair or a lone moth spends the day hiding from birds. Males stay with females to ensure paternity. Peppered moths usually rest in the upper parts of trees in sheltered spots: a few inches below a branch-trunk joint on a trunk (where it’s shadowed), on the underside of branches, or on leafy twigs. Researcher Michael Majerus noted that creationist critics often cite a 1985 statement by Clarke et al. that in 25 years they found only two moths on tree trunks or walls near traps. Majerus explained that few people look for moths high in trees, where peppered moths actually rest by day. This is supported by experiments with captive moths (Mikkola, 1979, 1984; Liebert and Brakefield, 1987). Majerus et al. (2000) showed that moths are cryptically camouflaged against their backgrounds when resting on tree boughs. In human-visible light, the typical light form (typica) blends with lichens, and the dark form (carbonaria) blends with plain bark. Birds see ultraviolet light, which humans cannot. Using an ultraviolet-sensitive video camera, Majerus et al. found that typica reflects UV light in a speckled pattern, matching crustose lichens on branches in both UV and human light. However, typica is less well camouflaged against foliose lichens on trunks—though it matches in human light, foliose lichens do not reflect UV. In a seven-year Cambridge experiment (2001–2007), Majerus recorded natural resting spots of 135 moths: over half were on tree branches (mostly on the lower half), 37% on trunks (mostly on the north side), and 12.6% on or under twigs.
**Polymorphism** The peppered moth has several melanic and non-melanic forms, controlled genetically. A particular color form is indicated in a standard way by following...
- wingspan
- 45 to 62 mm (median 55 mm)
- distribution
- China, Russia, Mongolia, Japan, North Korea, South Korea, Nepal, Kazakhstan, Kyrgyzstan, Turkmenistan, Georgia, Azerbaijan, Armenia, Europe, North America
- life cycle
- univoltine in Great Britain and Ireland; bivoltine in southeastern North America
- known for
- example of population genetics and natural selection; industrial melanism
Lore & Background
The peppered moth has a wingspan ranging from 45 to 62 mm, with a median of 55 mm. Its wings are white, 'peppered' with black, and have more-or-less distinct black cross lines. The black speckling varies; in some examples it is almost absent, while in others it is so dense that the wings appear black sprinkled with white. The antennae of males are strongly bipectinate. The caterpillar is a twig mimic, varying in color between green and brown, and can sense the twig's color with its skin to match its body color to the background, an ability also found in cephalopods, chameleons, and some fish. The peppered moth is found across a wide range including China, Russia, Mongolia, Japan, North Korea, South Korea, Nepal, Kazakhstan, Kyrgyzstan, Turkmenistan, Georgia, Azerbaijan, Armenia, Europe, and North America. In Great Britain and Ireland it is univoltine (one generation per year), while in southeastern North America it is bivoltine (two generations per year). The life cycle consists of eggs, several larval instars (caterpillars), pupae that overwinter in the soil, and adults. Adults emerge between late May and August, with males slightly before females. Males fly every night in search of females, while females fly only on the first night and then release pheromones to attract males. The female lays about 2,000 pale-green ovoid eggs about 1 mm in length into crevices in bark. Peppered moths rest by day in the upper part of trees at unexposed positions, such as below branch-trunk joints, on the underside of branches, or on foliate twigs. The moths are cryptically camouflaged against their backgrounds; typica are camouflaged against lichens and carbonaria against plain bark. Birds can see ultraviolet light, and typica reflect ultraviolet light in a speckled fashion, camouflaging them against crustose lichens common on branches.
Reader's Guide
The peppered moth is a key example in the study of population genetics and natural selection. Its evolution over the last two hundred years has been studied in detail, particularly in relation to the Industrial Revolution in England. At the start of this period, most peppered moths had light-colored wing patterns that camouflaged them against light-colored trees and lichens. Due to widespread pollution, many lichens died out and trees became blackened by soot, causing most light-colored moths (typica) to die off due to predation, while dark-colored melanic moths (carbonaria) flourished. This shift illustrates how environmental changes can drive natural selection. The peppered moth exhibits polymorphism, with several melanic and non-melanic morphs controlled genetically. In Europe, there are three morphs: the white typica, the dark melanistic carbonaria, and an intermediate medionigra. In Britain, the intermediate is named insularia. In North America, the melanic black morph is morpha swettaria. In Japan, no melanic morphs have been recorded. The melanic allele is dominant to the non-melanic allele in European and North American populations. Importantly, individuals of each morph interbreed and produce fertile offspring, so there is only one species. The peppered moth's resting behavior and camouflage have been studied using ultraviolet-sensitive video cameras, showing that typica are camouflaged against crustose lichens in both human-visible and ultraviolet wavelengths.
Did You Know?
- The caterpillars can sense the twig's color with their skin and match their body color to the background.
- In Japan, no melanic morphs have been recorded; all are morpha typica.
- The female lays about 2,000 pale-green ovoid eggs about 1 mm in length.
More in Arthropods 1-24
Spotted an error? Know more?
This is a living reference — every entry is fact-audited, and reader corrections feed straight into our audit queue. Suggest an edit · See this site's audit record
