
The evolution of the peppered moth is an
evolutionary
Evolution is the change in the heritable characteristics of biological populations over successive generations. It occurs when evolutionary processes such as natural selection and genetic drift act on genetic variation, resulting in certa ...
instance of directional colour change in the moth population as a consequence of air pollution during the
Industrial Revolution
The Industrial Revolution, sometimes divided into the First Industrial Revolution and Second Industrial Revolution, was a transitional period of the global economy toward more widespread, efficient and stable manufacturing processes, succee ...
. The frequency of dark-coloured moths increased at that time, an example of
industrial melanism
Industrial melanism is an evolutionary effect prominent in several arthropods, where dark pigmentation (melanism) has evolved in an environment affected by industrial pollution, including Sulfur dioxide, sulphur dioxide gas and dark soot deposit ...
. Later, when pollution was reduced in response to clean air legislation, the light-coloured form again predominated. Industrial melanism in the
peppered moth
The peppered moth (''Biston betularia'') is a temperate species of Nocturnality, night-flying moth. It is mostly found in the northern hemisphere in places like Asia, Europe and North America. Peppered moth evolution is an example of populatio ...
was an early test of
Charles Darwin
Charles Robert Darwin ( ; 12 February 1809 – 19 April 1882) was an English Natural history#Before 1900, naturalist, geologist, and biologist, widely known for his contributions to evolutionary biology. His proposition that all speci ...
's
natural selection
Natural selection is the differential survival and reproduction of individuals due to differences in phenotype. It is a key mechanism of evolution, the change in the Heredity, heritable traits characteristic of a population over generation ...
in action, and it remains a classic example in the
teaching of evolution.
In 1978,
Sewall Wright
Sewall Green Wright ForMemRS
HonFRSE (December 21, 1889March 3, 1988) was an American geneticist known for his influential work on evolutionary theory and also for his work on path analysis. He was a founder of population genetics alongside ...
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
Manchester () is a city and the metropolitan borough of Greater Manchester, England. It had an estimated population of in . Greater Manchester is the third-most populous metropolitan area in the United Kingdom, with a population of 2.92&nbs ...
, 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
Camouflage is the use of any combination of materials, coloration, or illumination for concealment, either by making animals or objects hard to see, or by disguising them as something else. Examples include the leopard's spotted coat, the b ...
in a clean environment, such as in rural
Dorset
Dorset ( ; Archaism, archaically: Dorsetshire , ) is a Ceremonial counties of England, ceremonial county in South West England. It is bordered by Somerset to the north-west, Wiltshire to the north and the north-east, Hampshire to the east, t ...
, while the dark colour was beneficial in a polluted environment like industrial
Birmingham
Birmingham ( ) is a City status in the United Kingdom, city and metropolitan borough in the metropolitan county of West Midlands (county), West Midlands, within the wider West Midlands (region), West Midlands region, in England. It is the Lis ...
. 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
''Darwinism'' is a term used to describe a theory of biological evolution developed by the English naturalist Charles Darwin (1809–1882) and others. The theory states that all species of organisms arise and develop through the natural sele ...
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 there principal defender. His seven-year experiment beginning in 2001, the most elaborate of its kind in
population biology
The term population biology has been used with different meanings.
In 1971, Edward O. Wilson ''et al''. used the term in the sense of applying mathematical models to population genetics, community ecology, and population dynamics. Alan Hasting ...
, the results of which were published posthumously in 2012, vindicated Kettlewell's work in great detail. This restored the peppered moth evolution as "the most direct evidence", and "one of the clearest and most easily understood examples of Darwinian evolution in action".
Origin and evolution

Before the
Industrial Revolution
The Industrial Revolution, sometimes divided into the First Industrial Revolution and Second Industrial Revolution, was a transitional period of the global economy toward more widespread, efficient and stable manufacturing processes, succee ...
, 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 University of Oxford is a collegiate university, collegiate research university in Oxford, England. There is evidence of teaching as early as 1096, making it the oldest university in the English-speaking world and the List of oldest un ...
. The first live specimen was caught by R. S. Edleston in
Manchester
Manchester () is a city and the metropolitan borough of Greater Manchester, England. It had an estimated population of in . Greater Manchester is the third-most populous metropolitan area in the United Kingdom, with a population of 2.92&nbs ...
,
England
England is a Countries of the United Kingdom, country that is part of the United Kingdom. It is located on the island of Great Britain, of which it covers about 62%, and List of islands of England, more than 100 smaller adjacent islands. It ...
in 1848, but he reported this only 16 years later in 1864, in ''
The Entomologist
The ''Entomological Magazine'' was a publication devoted to entomology.
The ''Entomological Magazine'' was published between September 1832 and October 1838 by the Society of Entomologists of London. The editor was Edward Newman (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
lichen
A lichen ( , ) is a hybrid colony (biology), colony of algae or cyanobacteria living symbiotically among hypha, filaments of multiple fungus species, along with yeasts and bacteria embedded in the cortex or "skin", in a mutualism (biology), m ...
s 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
Soot ( ) is a mass of impure carbon particles resulting from the incomplete combustion of hydrocarbons. Soot is considered a hazardous substance with carcinogenic properties. Most broadly, the term includes all the particulate matter produced b ...
from the new coal-burning factories. Many of the light-bodied lichens died from
sulphur dioxide
Sulfur dioxide (IUPAC-recommended spelling) or sulphur dioxide (traditional Commonwealth English) is the chemical compound with the formula . It is a colorless gas with a pungent smell that is responsible for the odor of burnt matches. It is r ...
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
An ecosystem (or ecological system) is a system formed by Organism, organisms in interaction with their Biophysical environment, environment. The Biotic material, biotic and abiotic components are linked together through nutrient cycles and en ...
: indeed, their bodies now dramatically contrasted with the colour of the bark. Dark-coloured moths, on the other hand, were
camouflage
Camouflage is the use of any combination of materials, coloration, or illumination for concealment, either by making animals or objects hard to see, or by disguising them as something else. Examples include the leopard's spotted coat, the b ...
d very well by the blackened trees.
[Miller, Ken (1999). ]
The Peppered Moth: An Update
' 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%) from the original frequency.
This effect of industrialization in body colour led to the coining of the term "
industrial melanism
Industrial melanism is an evolutionary effect prominent in several arthropods, where dark pigmentation (melanism) has evolved in an environment affected by industrial pollution, including Sulfur dioxide, sulphur dioxide gas and dark soot deposit ...
".
[
The implication that industrial melanism could be evidence supporting ]Charles Darwin
Charles Robert Darwin ( ; 12 February 1809 – 19 April 1882) was an English Natural history#Before 1900, naturalist, geologist, and biologist, widely known for his contributions to evolutionary biology. His proposition that all speci ...
's theory of natural selection
Natural selection is the differential survival and reproduction of individuals due to differences in phenotype. It is a key mechanism of evolution, the change in the Heredity, heritable traits characteristic of a population over generation ...
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
The New Forest is one of the largest remaining tracts of unenclosed pasture land, heathland and forest in Southern England, covering southwest Hampshire and southeast Wiltshire. It was proclaimed a royal forest by William the Conqueror, featu ...
, brown moths on clay and red soil in Herefordshire
Herefordshire ( ) is a Ceremonial counties of England, ceremonial county in the West Midlands (region), West Midlands of England, bordered by Shropshire to the north, Worcestershire to the east, Gloucestershire to the south-east, and the Welsh ...
, and white moths on chalk cliffs in Lewes
Lewes () is the county town of East Sussex, England. The town is the administrative centre of the wider Lewes (district), district of the same name. It lies on the River Ouse, Sussex, River Ouse at the point where the river cuts through the Sou ...
, 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 kiln
A lime kiln is a kiln used for the calcination of limestone (calcium carbonate) to produce the form of lime called ''quicklime'' (calcium oxide). The chemical equation for this reaction is: CaCO3 + heat → CaO + CO2
This reaction can tak ...
s, 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.[
]
Rise and fall of phenotype frequency
Melanism
Melanism is the congenital excess of melanin in an organism resulting in dark pigment.
Pseudomelanism, also called abundism, is another variant of pigmentation, identifiable by dark spots or enlarged stripes, which cover a large part of the bod ...
has been observed in both European and North American peppered moth populations. Information about the rise in frequency is scarce. Much more is known about the subsequent fall in phenotype
In genetics, the phenotype () is the set of observable characteristics or traits of an organism. The term covers the organism's morphology (physical form and structure), its developmental processes, its biochemical and physiological propert ...
frequency, as it has been measured by lepidopterists using moth traps.
Steward compiled data for the first recordings of the peppered moth by locality, and deduced that the ''carbonaria'' morph was the result of a single mutation
In biology, a mutation is an alteration in the nucleic acid sequence of the genome of an organism, virus, or extrachromosomal DNA. Viral genomes contain either DNA or RNA. Mutations result from errors during DNA or viral replication, ...
that subsequently spread. By 1895, it had reached a reported frequency of 98% in Manchester.
From around 1962 to the present, the phenotype frequency of ''carbonaria'' has steadily fallen in line with cleaner air around industrial cities. Its decline has been measured more accurately than its rise, through more rigorous scientific studies. Notably, Kettlewell conducted a national survey in 1956, Bruce Grant conducted a similar one in early 1996, and L.M. Cook in 2003.
Similar results were found in North America. Melanic forms have not been found in Japan. It is believed that this is because peppered moths in Japan do not inhabit industrialised regions.[
]
Genetics
Tutt was the first to propose the "differential bird predation hypothesis" in 1896, as a mechanism of natural selection
Natural selection is the differential survival and reproduction of individuals due to differences in phenotype. It is a key mechanism of evolution, the change in the Heredity, heritable traits characteristic of a population over generation ...
. The melanic morphs were better camouflaged against the bark of trees without foliose lichen, whereas the ''typica'' morphs were better camouflaged against trees with lichens. As a result, birds would find and eat those morphs that were not camouflaged with increased frequency.
In 1924, J.B.S. Haldane calculated, using a simple general selection model, the selective advantage necessary for the recorded natural evolution of peppered moths, based on the assumption that in 1848 the frequency of dark-coloured moths was 2%, and by 1895 it was 95%. The dark-coloured, or melanic, form would have had to be 50% more fit than the typical, light-coloured form. Even taking into consideration possible errors in the model, this reasonably excluded the stochastic Stochastic (; ) is the property of being well-described by a random probability distribution. ''Stochasticity'' and ''randomness'' are technically distinct concepts: the former refers to a modeling approach, while the latter describes phenomena; i ...
process of genetic drift, because the changes were too fast.[ Haldane, J.B.S. (1924). '' A Mathematical Theory of Natural and Artificial Selection''.] Haldane's statistical analysis of selection for the melanic variant in peppered moths became a well known part of his effort to demonstrate that mathematical models that combined natural selection with Mendelian genetics could explain evolution – an effort that played a key role in the foundation of the discipline of population genetics
Population genetics is a subfield of genetics that deals with genetic differences within and among populations, and is a part of evolutionary biology. Studies in this branch of biology examine such phenomena as Adaptation (biology), adaptation, s ...
, and the beginnings of the modern synthesis of evolutionary theory with genetics.
The peppered moth ''Biston betularia'' is also a model of parallel evolution in the incidence of melanism in the British form (''f. carbonaria'') and the American form (''f. swettaria'') as they are indistinguishable in appearance. Genetic analysis indicates that both phenotypes are inherited as autosomal dominant
In genetics, dominance is the phenomenon of one variant (allele) of a gene on a chromosome masking or overriding the Phenotype, effect of a different variant of the same gene on Homologous chromosome, the other copy of the chromosome. The firs ...
s. Cross hybridizations indicate that the phenotypes are produced by alleles at a single locus.
The gene for ''carbonaria'' in ''B. betularia'' was thought to be in a region of chromosome
A chromosome is a package of DNA containing part or all of the genetic material of an organism. In most chromosomes, the very long thin DNA fibers are coated with nucleosome-forming packaging proteins; in eukaryotic cells, the most import ...
17. It was later concluded that the gene could not be in that region, because none of the genes in the chromosome coded for either wing pattern or melanisation. The region that was used to find it was the first intron of the orthologue of the ''cortex'' gene in ''Drosophila''. Through elimination of candidates within the region based on rarity, a 21,925 base pair insert remained. The insert, labelled ''carb''-TE, is a class II transposable element
A transposable element (TE), also transposon, or jumping gene, is a type of mobile genetic element, a nucleic acid sequence in DNA that can change its position within a genome.
The discovery of mobile genetic elements earned Barbara McClinto ...
that has an approximately 9-kb non-repetitive sequence tandemly repeated two and one third times. There are 6 base pairs of inverted repeats and duplicated 4 base pairs at the target site not present in ''typica'' moths. ''Carb-''TE has higher expression during the stage of rapid wing disc morphogenesis. The mechanism of how the gene increases expression, and whether it is the only gene involved, is still not known.
Alternative hypotheses
Several alternative hypotheses to natural selection as the driving force of evolution were proposed during the 1920s and 1930s. Random mutation, migration
Migration, migratory, or migrate may refer to: Human migration
* Human migration, physical movement by humans from one region to another
** International migration, when peoples cross state boundaries and stay in the host state for some minimum le ...
, and genetic drift
Genetic drift, also known as random genetic drift, allelic drift or the Wright effect, is the change in the Allele frequency, frequency of an existing gene variant (allele) in a population due to random chance.
Genetic drift may cause gene va ...
were also seen as major forces of evolution.
P. A. Riley proposed an additional selective factor, where heavy metal chelation
Chelation () is a type of bonding of ions and their molecules to metal ions. It involves the formation or presence of two or more separate coordinate bonds between a polydentate (multiple bonded) ligand and a single central metal atom. These l ...
by melanin would supposedly protect peppered moths against the toxic effects of heavy metals associated with industrialisation. This selective advantage would supplement the major selective mechanism of differential bird predation.
Phenotypic induction
In 1920, John William Heslop-Harrison rejected Tutt's differential bird predation hypothesis, on the basis that he did not believe that birds ate moths. Instead he proposed that pollutants could cause changes to the soma and germ plasm
Germ plasm () is a biological concept developed in the 19th century by the German biologist August Weismann. It states that heritable information is transmitted only by germ cells in the gonads (ovaries and testes), not by somatic cells. The ...
of the organism. In 1925, K. Hasebroek made an early attempt to prove this hypothesis, exposing pupae to pollutant gases, namely hydrogen sulfide
Hydrogen sulfide is a chemical compound with the formula . It is a colorless chalcogen-hydride gas, and is toxic, corrosive, and flammable. Trace amounts in ambient atmosphere have a characteristic foul odor of rotten eggs. Swedish chemist ...
(H2S), ammonia
Ammonia is an inorganic chemical compound of nitrogen and hydrogen with the chemical formula, formula . A Binary compounds of hydrogen, stable binary hydride and the simplest pnictogen hydride, ammonia is a colourless gas with a distinctive pu ...
(NH3), and "pyredin". He used eight species in his studies, four of which were species of butterfly that did not exhibit melanism.
In 1926 and 1928, Heslop-Harrison suggested that the increase of melanic moths in industrialised regions was due to " mutation pressure", not to selection by predators which he regarded as negligible. Salts of lead
Lead () is a chemical element; it has Chemical symbol, symbol Pb (from Latin ) and atomic number 82. It is a Heavy metal (elements), heavy metal that is density, denser than most common materials. Lead is Mohs scale, soft and Ductility, malleabl ...
and manganese
Manganese is a chemical element; it has Symbol (chemistry), symbol Mn and atomic number 25. It is a hard, brittle, silvery metal, often found in minerals in combination with iron. Manganese was first isolated in the 1770s. It is a transition m ...
were present in the airborne pollutant particles, and he suggested that these caused the mutation of genes for melanin production but of no others. He used ''Selenia bilunaria'' and ''Tephrosia bistortata'' as material. The larvae were fed with leaves that had incorporated these salts: melanics subsequently appeared. A similar experiment in 1932 by McKenney Hughes failed to replicate these results; the statistician and geneticist Ronald Fisher
Sir Ronald Aylmer Fisher (17 February 1890 – 29 July 1962) was a British polymath who was active as a mathematician, statistician, biologist, geneticist, and academic. For his work in statistics, he has been described as "a genius who a ...
showed that Heslop-Harrison's controls were inadequate, and that Hughes's findings made the 6% mutation rate required by Heslop-Harrison "improbable".
Kettlewell's experiment
The first important experiments on the peppered moth were carried out by Bernard Kettlewell at Oxford University, under the supervision of E. B. Ford, who helped him gain a grant from the Nuffield Foundation
The Nuffield Foundation is a charitable trust established in 1943 by William Morris, Lord Nuffield, the founder of Morris Motors Ltd. It aims to improve social well-being by funding research and innovation projects in education and social pol ...
to perform the experiments. In 1953, Kettlewell started a preliminary experiment in which moths were released into a large (18m × 6m) aviary
An aviary is a large enclosure for confining birds, although bats may also be considered for display. Unlike birdcages, aviaries allow birds a larger living space where Bird flight, they can fly; hence, aviaries are also sometimes known as flig ...
, where they were fed on by great tit
The great tit (''Parus major'') is a small passerine bird in the tit family Paridae. It is a widespread and common species throughout Europe, the Middle East, Central Asia and east across the Palearctic to the Amur River, south to parts of No ...
s (''Parus major''). His main experiment, at Christopher Cadbury Wetland Reserve in Birmingham
Birmingham ( ) is a City status in the United Kingdom, city and metropolitan borough in the metropolitan county of West Midlands (county), West Midlands, within the wider West Midlands (region), West Midlands region, in England. It is the Lis ...
, England
England is a Countries of the United Kingdom, country that is part of the United Kingdom. It is located on the island of Great Britain, of which it covers about 62%, and List of islands of England, more than 100 smaller adjacent islands. It ...
, involved marking, releasing, and recapturing marked moths. He found that in this polluted woodland ''typica'' moths were preferentially preyed upon. He thus showed that the melanic phenotype was important to the survival of peppered moths in such a habitat. Kettlewell repeated the experiment in 1955 in unpolluted woodlands in Dorset
Dorset ( ; Archaism, archaically: Dorsetshire , ) is a Ceremonial counties of England, ceremonial county in South West England. It is bordered by Somerset to the north-west, Wiltshire to the north and the north-east, Hampshire to the east, t ...
, and again in the polluted woods in Birmingham.[ In 1956 he repeated the experiments and found similar results; in Birmingham, birds ate most of the white moths (75%), whereas in Dorset, most of the dark moths (86%) were eaten.][
]
Criticisms
Theodore David Sargent performed experiments between 1965 and 1969, from which he concluded that it was not possible to reproduce Kettlewell's results, and said that birds showed no preference for moths on either black or white tree trunks. He suggested that Kettlewell had trained the birds to pick moths on tree trunks to obtain the desired results.
Two chapters in Michael Majerus's 1998 book '' Melanism: Evolution in Action'' critiqued the research in Kettlewell's '' The Evolution of Melanism'', discussed studies which raised questions about Kettlewell's original experimental methods, and called for further research.[ Reviewing the book, Jerry Coyne noted these points, and concluded that "for the time being we must discard ''Biston'' as a well-understood example of natural selection in action, although it is clearly a case of evolution. There are many studies more appropriate for use in the classroom."]
Judith Hooper's book '' Of Moths and Men'' (2002) severely criticised Kettlewell's experiment. Hooper argued that Kettlewell's field notes could not be found and suggested that his experiment was fraudulent, on the basis of Sargent's criticisms alleging that the photographs of the moths were taken of dead moths placed on a log. She said that E. B. Ford was a "Darwinian zealot", and claimed that he exploited the scientifically naive Kettlewell to obtain the desired experimental results. The book's reception led to demands that the peppered moth evolution story be deleted from textbooks. Scientists have examined the allegations made by Hooper, and found them to be without merit.[
Phillip E. Johnson, a co-founder of the creationist ]intelligent design
Intelligent design (ID) is a pseudoscientific argument for the existence of God, presented by its proponents as "an evidence-based scientific theory about life's origins".#Numbers 2006, Numbers 2006, p. 373; " Dcaptured headlines for it ...
movement, said that the moths "do not sit on tree trunks", that "moths had to be glued to the trunks" for pictures, and that the experiments were "fraudulent" and a "scam." The intelligent design advocate Jonathan Wells wrote an essay on the subject, a shortened version of which appeared in the 24 May 1999 issue of '' The Scientist'', claiming that "The fact that peppered moths do not normally rest on tree trunks invalidates Kettlewell's experiments". Wells further wrote in his 2000 book '' Icons of Evolution'' that "What the textbooks don't explain, however, is that biologists have known since the 1980s that the classical story has some serious flaws. The most serious is that peppered moths in the wild don't even rest on tree trunks. The textbook photographs, it turns out, have been staged."[Wells J. (2000). '' Icons of Evolution: Science or Myth? Why Much of What We Teach About Evolution is Wrong''. Regnery Press, Washington, D.C., p. 138 (book available fro]
Iconsofevolution.com
However, peppered moths do rest on tree trunks on occasion, and Nick Matzke states that there is little difference between the 'staged' photos and 'unstaged' ones.
Majerus's experiment
From 2001 to 2007, Majerus carried out experiments in Cambridge to resolve the various criticisms of Kettlewell's experiment. During his experiment, he noted the natural resting positions of peppered moths. Of the 135 moths examined, over half were on tree branches, mostly on the lower half of the branch; 37% were on tree trunks, mostly on the north side; and only 12.6% were resting on or under twigs. Following correspondence with Hooper, he added an experiment to find if bats
Bats are flying mammals of the order Chiroptera (). With their forelimbs adapted as wings, they are the only mammals capable of true and sustained flight. Bats are more agile in flight than most birds, flying with their very long spread-out ...
, not birds, could be the main predators. He observed a number of species of bird actually preying on the moths, and found that differential bird predation was a major factor responsible for the decline in ''carbonaria'' frequency compared to ''typica''.[ He described his results as a complete vindication of the natural selection theory of peppered moth evolution, and said "If the rise and fall of the peppered moth is one of the most visually impacting and easily understood examples of Darwinian evolution in action, it should be taught. It provides after all the proof of evolution."]
Majerus died before he could complete the writing up of his experiments, so the work was carried on by Cook, Grant, Saccheri, and James Mallet, and published on 8 February 2012 as "Selective bird predation on the peppered moth: the last experiment of Michael Majerus." The experiment became the largest ever in the study of industrial melanism, involving 4,864 individuals in a six-year investigation, and it confirmed that melanism in moths is a genuine example of natural selection involving camouflage and predation. Their concluding remark runs: "These data provide the most direct evidence yet to implicate camouflage and bird predation as the overriding explanation for the rise and fall of melanism in moths."
Coyne said he was "delighted to agree with this conclusion f Majerus's experiment which answers my previous criticisms about the ''Biston'' story."
See also
* Polymorphism
* Scottish red deer
Notes
References
Further reading
*
External links
* Bruce Grant has written several papers on melanism in the peppered moth which are listed o
his home page
* Online lecture
"The rise and fall of the melanic Peppered Moth"
presented by Laurence Cook.
*
The Peppered Moth: Decline of a Darwinian Disciple
This is the transcript of Michael Majerus' lecture delivered to the British Humanist Association
Humanists UK, known from 1967 until May 2017 as the British Humanist Association (BHA), is a charitable organisation which promotes secular humanism and aims to represent non-religious people in the UK through a mixture of charitable servic ...
on Darwin Day
Darwin Day is a celebration to commemorate the birthday of Charles Darwin on 12 February 1809. The day is used to highlight Darwin's contributions to science and to promote science in general. Darwin Day is celebrated around the world.
Histo ...
2004.
The Peppered Moth: The Proof of Darwinian Evolution
This is the transcript of Majerus' lecture given at the European Society for Evolutionary Biology meeting on 23 August 2007. The accompanying PowerPoint presentation i
also available
*
* On 19 June 2009
Telegraph.co.uk
publishe
on this evolutionary phenomenon and implored UK readers to visit th
Moths Count website
and record their observations of local moths, in an effort to help increase the available data for researchers.
{{Pepperedmoth
Biology experiments
Evolution of insects
Evolution
Evolution is the change in the heritable Phenotypic trait, characteristics of biological populations over successive generations. It occurs when evolutionary processes such as natural selection and genetic drift act on genetic variation, re ...
Selection