Editing Peppered moth evolution (section)

{{short description|Significance of the peppered moth in evolutionary biology}}
{{Good article}}
{{About|the peppered moth's significance in [[evolutionary biology]]|its evolutionary ancestry|Insect evolution}}
{{Use dmy dates|date=May 2021}}
{{Use British English|date=August 2022}}
[[File:Biston.betularia.7200.jpg|thumb|upright=1.2|''Biston betularia f. typica'', the white-bodied [[peppered moth]] ]]
[[File:Biston.betularia.f.carbonaria.7209.jpg|thumb|upright=1.2|''Biston betularia f. carbonaria'', the black-bodied peppered moth ]]

The '''evolution of the peppered moth''' is an [[evolutionary biology|evolutionary]] instance of directional colour change in the moth population as a consequence of air pollution during the [[Industrial Revolution]]. The frequency of dark-coloured moths increased at that time, an example of [[industrial melanism]]. Later, when pollution was reduced in response to clean air legislation, the light-coloured form again predominated. Industrial melanism in the [[peppered moth]] was an early test of [[Charles Darwin]]'s [[natural selection]] in action, and it remains a classic example in the [[teaching of evolution]].<ref name="Saccheri">{{cite journal |last1=Cook |first1=L. M. |last2=Saccheri |first2=I J |title=The peppered moth and industrial melanism: evolution of a natural selection case study |journal=Heredity |date=March 2013 |volume=110 |issue=3 |pages=207–212 |pmid=23211788 |doi=10.1038/hdy.2012.92 |pmc=3668657}}</ref><ref name=rudgebio05>{{cite journal |last1=Rudge |first1=David W. |title=The Beauty of Kettlewell's Classic Experimental Demonstration of Natural Selection |journal=[[BioScience]] |year=2005 |volume=55 |issue=4 |pages=369–375 |doi=10.1641/0006-3568(2005)055[0369:TBOKCE]2.0.CO;2 |doi-access=free}}</ref><ref name=majerus2008>{{cite journal |last1=Majerus |first1=Michael E. N. |author-link=Michael Majerus |title=Industrial Melanism in the Peppered Moth, Biston betularia: An Excellent Teaching Example of Darwinian Evolution in Action |journal=Evolution: Education and Outreach |year=2008 |volume=2 |issue=1 |pages=63–74 |doi=10.1007/s12052-008-0107-y |doi-access=free |s2cid=25407417 |url=http://www.docente.unife.it/giorgio.bertorelle/didattica_insegnamenti/biologia-evoluzionistica-1/Bistonbetularia_History.pdf}}</ref> In 1978, [[Sewall Wright]] described it as "the clearest case in which a conspicuous evolutionary process has actually been observed."<ref name=rice>{{cite book |last1=Rice |first1=Stanley A. |title=Encyclopedia of Evolution |year=2007 |publisher=Facts on File |location=New York |isbn=978-1-4381-1005-9 |page=308 |url=https://books.google.com/books?id=YRcAVvmE6eMC}}</ref><ref name="Majerus1998">{{cite book |last=Majerus |first=Michael E. N. |author-link=Michael Majerus |title=Melanism: Evolution in Action |year=1998 |publisher=Oxford University Press |isbn=978-0-19-854983-3 |url=https://books.google.com/books?id=YqgXAQAAIAAJ}}</ref>

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]], 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.<ref>{{cite journal |last1=Clarke |first1=C. A. |last2=Mani |first2=G. S. |last3=Wynne |first3=G. |title=Evolution in reverse: clean air and the peppered moth |journal=Biological Journal of the Linnean Society |date=1985 |volume=26 |issue=2 |pages=189–199 |doi=10.1111/j.1095-8312.1985.tb01555.x}}</ref> 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.<ref>{{cite journal |last=Majerus |first=Michael E. N. |title=Industrial Melanism in the Peppered Moth, ''Biston betularia'': An Excellent Teaching Example of Darwinian Evolution in Action |journal=Evolution: Education and Outreach |date=2008 |volume=2 |issue=1 |pages=63–74 |doi=10.1007/s12052-008-0107-y |doi-access=free}}</ref> 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]] in a clean environment, such as in rural [[Dorset]], while the dark colour was beneficial in a polluted environment like industrial [[Birmingham]]. 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]] evolution and [[coloration evidence for natural selection|evidence for natural selection]] used in standard textbooks.<ref name="Bender">{{cite journal |last=Bender |first=Eric |title=Urban evolution: How species adapt to survive in cities |journal=Knowable Magazine |publisher=Annual Reviews |date=21 March 2022 |doi=10.1146/knowable-031822-1 |doi-access=free |url=https://knowablemagazine.org/article/living-world/2022/urban-evolution-species-adapt-survive-cities}}</ref><ref name="Diamond">{{cite journal |last1=Diamond |first1=Sarah E. |last2=Martin |first2=Ryan A. |title=Evolution in Cities |journal=Annual Review of Ecology, Evolution, and Systematics |date=2 November 2021 |volume=52 |issue=1 |pages=519–540 |doi=10.1146/annurev-ecolsys-012021-021402 |doi-access=free |s2cid=239646134 |url=https://www.annualreviews.org/doi/pdf/10.1146/annurev-ecolsys-012021-021402}}</ref><ref name="Miller">{{cite web |last1=Miller |first1=Ken |title=The Peppered Moth - An Update |date=1999 |website=millerandlevine.com |url=http://www.millerandlevine.com/km/evol/Moths/moths.html |access-date=31 March 2022}}</ref>

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 [[creationism|creationists]]. [[Michael Majerus]] was the principal defender. His seven-year experiment beginning in 2001, the most elaborate of its kind in [[population biology]], the results of which were published posthumously in 2012, vindicated Kettlewell's work in great detail. This restored peppered moth evolution as "the most direct evidence", and "one of the clearest and most easily understood examples of Darwinian evolution in action".<ref name="Cook2012">{{cite journal |last1=Cook |first1=L. M. |last2=Grant |first2=B. S. |last3=Saccheri |first3=I. J. |last4=Mallet |first4=James |author4-link=James Mallet |title=Selective bird predation on the peppered moth: the last experiment of Michael Majerus |journal=[[Biology Letters]] |volume=8 |issue=4 |pages=609–612 |year=2012 |pmid=22319093 |doi=10.1098/rsbl.2011.1136 |pmc=3391436}}</ref>