664:(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.
749:, 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."
183:
50:
35:
668:
382:, 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
302:, 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
742:. 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."
260:
very well by the blackened trees. 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
237:
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
624:
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
729:
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
710:
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." However,
386:
process of genetic drift, because the changes were too fast. 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
174:, 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".
1793:
van't Hof, Arjen E.; Campagne, Pascal; Rigden, Daniel J.; Yung, Carl J.; Lingley, Jessica; Quail, Michael A.; Hall, Neil; Darby, Alistair; Saccheri, Ilik J. (2 June 2016). "The industrial melanism mutation in
British peppered moths is a transposable element".
433:
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
486:
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.
347:
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,
610:, 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.
546:
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
605:
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
1963:
Hasebroek, K. (1925). "Die prinzipielle
Loesung des Problems des Grossstadt- und Industriemelanismus der Schmetterlinge" [The principal solution of problems of city- and industrial-melanism of butterflies].
734:, 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
111:, 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.
693:
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
2891:
450:
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
1740:
van't Hof, Arjen E.; Edmonds, Nicola; Dalikova, Martina; Marec, Frantisek; Saccheri, Ilik J. (20 May 2011). "Industrial
Melanism in British Peppered Moths Has a Singular and Recent Mutational Origin".
276:
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
229:
298:, 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
459:
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.
1645:
355:
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.
1343:
499:
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
2282:
Sargent, T. D.; Millar, C.D.; Lambert, D. M. (1988). "Chapter 9: The 'classical' explanation of industrial melanism: Assessing the evidence". In Hecht, Max K.; Wallace, Bruce (eds.).
730:
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
2044:
115:), 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
317:
has appeared 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
652:
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."
125:
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
233:. 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
371:
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.
1456:
906:
137:. 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
2843:
1609:
704:, 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
152:
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
2997:
695:
538:
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
249:
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
551:
showed that Heslop-Harrison's controls were inadequate, and that Hughes's findings made the 6% mutation rate required by Heslop-Harrison "improbable".
2835:
2813:
146:
2831:
2440:
1369:
644:, discussed studies which raised questions about Kettlewell's original experimental methods, and called for further research. Reviewing the book,
2923:
2763:
2551:
993:
963:
119:
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.
590:
215:, the black form of the peppered moth was rare. The first black specimen (of unknown origin) was collected before 1811, and kept in the
2660:
1924:"Genetical studies in the moths of the geometrid genus Oporabia (Oporinia) with a special consideration of melanism in the lepidoptera"
2082:
1074:
577:
to perform the experiments. In 1953, Kettlewell started a preliminary experiment in which moths were released into a large (18m × 6m)
241:
During the early decades of the Industrial Revolution in England, the countryside between London and Manchester became blanketed with
162:
was published in 2002, Kettlewell's story was more sternly attacked, and accused of fraud. The criticism became a major argument for
2568:
1867:
1681:
1326:
745:
Majerus died before he could complete the writing up of his experiments, so the work was carried on by Cook, Grant, Saccheri and
503:
of the organism. In 1925, K. Hasebroek made an early attempt to prove this hypothesis, exposing pupae to pollutant gases, namely
467:
Several alternative hypotheses to natural selection as the driving force of evolution were proposed during the 1920s and 1930s.
442:. Through elimination of candidates within the region based on rarity, a 21,925 base pair insert remained. The insert, labelled
392:
2291:
1669:
256:: indeed, their bodies now dramatically contrasted with the colour of the bark. Dark-coloured moths, on the other hand, were
907:"Industrial Melanism in the Peppered Moth, Biston betularia: An Excellent Teaching Example of Darwinian Evolution in Action"
2237:
Sargent, T. D. (1969). "Background Selections of the Pale and Melanic Forms of the Cryptic Moth, Phigalia titea (Cramer)".
3094:
3026:
634:
2374:
3089:
2821:
2035:
1983:
1919:
496:
2720:
2540:(2005). "The peppered moth: decline of a Darwinian disciple". In Fellowes, Mark; Holloway, Graham; Rolf, Jens (eds.).
2695:
526:
In 1926 and 1928, Heslop-Harrison suggested that the increase of melanic moths in industrialised regions was due to "
2956:
2916:
1616:
619:
560:
138:
2790:
1884:
3019:
2401:
1859:
640:
523:". He used eight species in his studies, four of which were species of butterfly that did not exhibit melanism.
882:
863:
414:) as they are indistinguishable in appearance. Genetic analysis indicates that both phenotypes are inherited as
2898:
and record their observations of local moths, in an effort to help increase the available data for researchers.
700:
2186:
Sargent, T. D. (1968). "Cryptic moths: effects on background selections of painting the circumocular scales".
1103:
2596:"Second Thoughts about Peppered Moths; This classical story of evolution by natural selection needs revising"
2347:
2987:
2779:
1521:
752:
Coyne said he was "delighted to agree with this conclusion , which answers my previous criticisms about the
379:
349:
96:
described it as "the clearest case in which a conspicuous evolutionary process has actually been observed."
76:. Later, when pollution was reduced, the light-coloured form again predominated. Industrial melanism in the
367:. The melanic morphs were better camouflaged against the bark of trees without foliose lichen, whereas the
182:
170:
was the principal defender. His seven-year experiment beginning in 2001, the most elaborate of its kind in
3084:
2040:"A further induction of melanism in the lepidopterous insect, Selenia bilunaria Esp., and its inheritance"
765:
68:
instance of directional colour change in the moth population as a consequence of air pollution during the
2887:
2429:
2164:
2909:
1847:
724:
212:
89:
69:
1009:
Clarke, C. A.; Mani, G. S.; Wynne, G. (1985). "Evolution in reverse: clean air and the peppered moth".
49:
363:
Tutt was the first to propose the "differential bird predation hypothesis" in 1896, as a mechanism of
332:
Steward compiled data for the first recordings of the peppered moth by locality, and deduced that the
2248:
2195:
2091:
1805:
1751:
1673:
1663:
1269:
447:
216:
65:
22:
2131:"A survey of the frequencies of Biston betularia (L.) (Lep.) and its melanic forms in Great Britain"
601:, involved marking, releasing and recapturing marked moths. He found that in this polluted woodland
261:
original frequency. This effect of industrialization in body colour led to the coining of the term "
1928:
1703:
574:
388:
262:
206:
73:
3099:
3033:
2972:
2618:
2520:
2264:
2219:
2135:
2017:
1945:
1829:
1775:
1580:
1503:
1402:
1125:
931:
706:
698:
wrote an essay on the subject, a shortened version of which appeared in the 24 May 1999 issue of
690:
686:
566:
415:
403:
282:
171:
122:
1923:
418:. Cross hybridizations indicate that the phenotypes are produced by alleles at a single locus.
3058:
2759:
2691:
2547:
2287:
2211:
2009:
1863:
1821:
1767:
1742:
1722:
1677:
1572:
1394:
1322:
1297:
1209:
989:
959:
841:
770:
364:
273:
107:) was rare, though a specimen had been collected by 1811. After field collection in 1848 from
85:
1525:
340:
that subsequently spread. By 1895, it had reached a reported frequency of 98% in Manchester.
3063:
3040:
2870:
2668:
2512:
2483:
2475:
2326:
2317:
2256:
2239:
2203:
2144:
2109:
2099:
2053:
1999:
1937:
1899:
1852:
1813:
1796:
1759:
1712:
1640:
1564:
1495:
1445:. Manuscripts Room, Cambridge University Library, West Road, Cambridge, England. DAR 164:26.
1386:
1287:
1277:
1236:
1199:
1191:
1115:
1082:
1053:
1018:
921:
877:
831:
823:
679:
660:
504:
472:
158:
26:
2977:
2850:
2817:
2749:
2645:
2537:
2435:
2352:
1659:
1605:
1424:
1377:
1182:
1144:
1120:
979:
902:
629:
468:
167:
2572:
2252:
2199:
2095:
1809:
1755:
1273:
812:"The peppered moth and industrial melanism: evolution of a natural selection case study"
715:
states that there is little difference between the 'staged' photos and 'unstaged' ones.
2692:"Selective bird predation on the peppered moth: the last experiment of Michael Majerus"
2488:
2463:
1499:
1292:
1255:
1240:
1227:
Berry, R. J. (1990). "Industrial melanism and peppered moths (Biston betularia (L.))".
1204:
1178:"Selective bird predation on the peppered moth: the last experiment of Michael Majerus"
1177:
1022:
836:
811:
375:
269:
246:
81:
34:
3078:
3002:
2946:
2932:
2881:
Accusations of data fudging and scientific fraud in the case are found to be vacuous.
2624:
2524:
1885:"A proposed selective mechanism based on metal chelation in industrial melanic moths"
1555:
Cook, L. M. (2003). "The rise and fall of the Carbonaria form of the peppered moth".
1129:
655:
548:
476:
322:
291:
250:
153:
93:
77:
42:
2595:
2378:
2223:
2021:
1949:
1779:
1584:
1507:
1482:
Steward, R. C. (1977). "Industrial and non-industrial melanism in the peppered moth
1406:
935:
864:"The Beauty of Kettlewell's Classic Experimental Demonstration of Natural Selection"
387:
evolution – an effort that played a key role in the foundation of the discipline of
2982:
2798:
2406:
2268:
2073:
1833:
1254:
Saccheri, I. J.; Rousset, F.; Watts, P. C.; Brakefield, P. M.; Cook, L. M. (2008).
1173:
746:
2753:
2541:
2207:
1316:
983:
953:
133:, while the dark colour was beneficial in a polluted environment like industrial
2782:
has written several papers on melanism in the peppered moth which are listed on
2716:
2641:
1368:
Hart, Adam G.; Stafford, Richard; Smith, Angela L.; Goodenough, Anne E. (2010).
712:
671:
645:
527:
326:
245:
from the new coal-burning factories. Many of the light-bodied lichens died from
163:
789:
Sargent (1936–2018) was a biologist at the University of Massachusetts Amherst.
667:
2992:
2875:
2854:
2839:
2825:
2479:
1390:
1087:
1058:
1037:
926:
868:
594:
570:
500:
430:
383:
303:
287:
257:
220:
198:
134:
126:
116:
108:
2013:
1988:"The induction of melanism in the lepidoptera and its subsequent inheritance"
1438:
72:. The frequency of dark-coloured moths increased at that time, an example of
2516:
1763:
1717:
1698:
1457:"How an extraordinary letter to Darwin spotted industrial melanism in moths"
1282:
582:
535:
483:
318:
299:
253:
142:
2104:
2078:"On the Evidence Against the Chemical Induction of Melanism in Lepidoptera"
2077:
2058:
2039:
2004:
1987:
1825:
1771:
1726:
1576:
1398:
1301:
1213:
1195:
845:
648:
noted these points, and concluded that "for the time being we must discard
530:", not to selection by predators which he regarded as negligible. Salts of
2215:
1256:"Selection and gene flow on a diminishing cline of melanic peppered moths"
565:
The first important experiments on the peppered moth were carried out by
520:
337:
314:
197:(below the bark's scar) is nearly invisible on this pollution-free tree,
1817:
1321:(Reprint ed.). Cambridge: Cambridge University Press. p. 171.
827:
2149:
2130:
1941:
1439:"Farn, A.B. to Darwin C.R., Darwin Correspondence Project Letter 11747"
598:
512:
482:
P. A. Riley proposed an additional selective factor, where heavy metal
234:
224:
2114:
1904:
268:
The implication that industrial melanism could be evidence supporting
2894:
on this evolutionary phenomenon and implored UK readers to visit the
2621:: Science or Myth? Why Much of What We Teach About Evolution is Wrong
2260:
607:
578:
130:
2628:
1568:
2331:
2308:
666:
295:
2859:: An Excellent Teaching Example of Darwinian Evolution in Action"
1042:: An Excellent Teaching Example of Darwinian Evolution in Action"
2901:
1260:
731:
531:
242:
2905:
2799:"Moonshine: Why the Peppered Moth remains an Icon of Evolution"
306:
explicitly linked peppered moth melanism to natural selection.
2783:
1370:"Evidence for contemporary evolution during Darwin's lifetime"
352:
conducted a similar one in early 1996, and L.M. Cook in 2003.
227:
in 1848, but he reported this only 16 years later in 1864, in
2402:"Of Moths and Men: Intrigue, Tragedy & the Peppered Moth"
2895:
674:
have disputed the occurrence or significance of the melanic
343:
From around 1962 to the present, the phenotype frequency of
2165:"Obituary: Theodore Sargent, Professor Emeritus of Biology"
883:
10.1641/0006-3568(2005)055[0369:TBOKCE]2.0.CO;2
2834:. This is the transcript of Majerus' lecture given at the
2661:"Moth study backs classic 'test case' for Darwin's theory"
219:. The first live specimen was caught by R. S. Edleston in
21:
This article is about the peppered moth's significance in
2569:"Evolution – April 1999: Peppered Moths and Creationists"
1699:"Allelic melanism in American and British peppered moths"
1646:
A Mathematical Theory of Natural and Artificial Selection
1075:"Urban evolution: How species adapt to survive in cities"
18:
Significance of the peppered moth in evolutionary biology
2546:. Wallingford, Oxon: CABI Publishing. pp. 375–377.
2167:. University of Massachusetts Amherst. 4 September 2018
711:
peppered moths do rest on tree trunks on occasion, and
286:). He noted the existence of dark moths in peat in the
1102:
Diamond, Sarah E.; Martin, Ryan A. (2 November 2021).
1610:"The Peppered Moth: The Proof of Darwinian Evolution"
2045:
Proceedings of the Royal Society of London. Series B
1992:
Proceedings of the Royal Society of London. Series B
1108:
Annual Review of Ecology, Evolution, and Systematics
625:
moths on tree trunks to obtain the desired results.
3050:
3011:
2965:
2939:
2832:
The Peppered Moth: The Proof of Darwinian Evolution
88:in action, and it remains a classic example in the
2814:The Peppered Moth: Decline of a Darwinian Disciple
2428:
1851:
406:in the incidence of melanism in the British form (
2801:. Department of Physics, Colorado School of Mines
2791:"The rise and fall of the melanic Peppered Moth"
2646:"Icon of Obfuscation: Chapter 7: Peppered Moths"
2627:, Washington, D.C., p. 138 (book available from
569:at Oxford University, under the supervision of
1665:Evolution: The history of an idea, 3rd edition
479:were also seen as major forces of evolution.
2917:
2503:Grant, B. S. (2002), "Sour grapes of wrath",
1600:
1598:
1596:
1594:
897:
895:
893:
8:
2838:meeting on 23 August 2007. The accompanying
2377:. W. W. Norton & Company. Archived from
1172:Cook, L. M.; Grant, B. S.; Saccheri, I. J.;
857:
855:
193:morphs on the same tree. The light-coloured
2855:"Industrial Melanism in the Peppered Moth,
1550:
1548:
1546:
1167:
1165:
1038:"Industrial Melanism in the Peppered Moth,
2924:
2910:
2902:
2874:
2836:European Society for Evolutionary Biology
2487:
2330:
2148:
2113:
2103:
2057:
2003:
1966:Internationale entomologische Zeitschrift
1903:
1892:Biological Journal of the Linnean Society
1716:
1418:
1416:
1291:
1281:
1229:Biological Journal of the Linnean Society
1203:
1119:
1086:
1057:
1011:Biological Journal of the Linnean Society
925:
881:
835:
810:Cook, L. M.; Saccheri, I J (March 2013).
620:Kettlewell's experiment § Criticisms
129:in a clean environment, such as in rural
958:. New York: Facts on File. p. 308.
181:
48:
33:
1363:
1361:
802:
782:
638:critiqued the research in Kettlewell's
573:, who helped him gain a grant from the
947:
945:
395:of evolutionary theory with genetics.
321:frequency, as it has been measured by
290:, brown moths on clay and red soil in
1854:Genetics and the Evolutionary Process
1526:"Distribution of melanism in Britain"
1121:10.1146/annurev-ecolsys-012021-021402
294:, and white moths on chalk cliffs in
25:. For its evolutionary ancestry, see
7:
310:Rise and fall of phenotype frequency
2758:. Oxford: Oxford University Press.
2346:Kenney, Michael (22 October 2002).
591:Christopher Cadbury Wetland Reserve
2721:"The peppered moth story is solid"
2348:"Of Dark Moths, Men and Evolution"
2083:Proceedings of the Royal Society B
1500:10.1111/j.1365-2311.1977.tb00886.x
1318:Introduction to Population Biology
1241:10.1111/j.1095-8312.1990.tb00518.x
1023:10.1111/j.1095-8312.1985.tb01555.x
689:, a co-founder of the creationist
141:, became the canonical example of
14:
2863:Evolution: Education and Outreach
1046:Evolution: Education and Outreach
914:Evolution: Education and Outreach
429:was thought to be in a region of
336:morph was the result of a single
56:, the black-bodied peppered moth.
2690:Matzke, Nick (8 February 2012).
2659:Connor, Steve (25 August 2007).
2443:from the original on 25 May 2022
2309:"Not Black and White. Review of
1998:(696): 241–263 /rspb.1926.0012.
1437:Farn, A. B. (18 November 1878).
103:form of the peppered moth (var.
2947:Overview, ecology, and genetics
2567:Frack, Donald (16 April 1999).
2400:Smith, Peter D. (11 May 2002).
1557:The Quarterly Review of Biology
1145:"The Peppered Moth - An Update"
1036:Majerus, Michael E. N. (2008).
2286:. Vol. 23. Plenum Press.
1670:University of California Press
1073:Bender, Eric (21 March 2022).
147:evidence for natural selection
62:evolution of the peppered moth
54:Biston betularia f. carbonaria
1:
3027:Melanism: Evolution in Action
2755:Melanism: Evolution in Action
2311:Melanism: Evolution in Action
2129:Kettlewell, H. B. D. (1958).
985:Melanism: Evolution in Action
635:Melanism: Evolution in Action
2822:British Humanist Association
2816:. This is the transcript of
2208:10.1126/science.159.3810.100
1425:The Peppered Moth: An Update
581:, where they were fed on by
497:John William Heslop-Harrison
391:, and the beginnings of the
149:used in standard textbooks.
2820:' lecture delivered to the
2793:presented by Laurence Cook.
2543:Insect Evolutionary Ecology
988:. Oxford University Press.
589:). His main experiment, at
378:calculated, using a simple
3116:
722:
718:
617:
558:
554:
204:
39:Biston betularia f. typica
20:
3020:The Evolution of Melanism
2876:10.1007/s12052-008-0107-y
2594:Wells, J. (24 May 1999).
2480:10.1038/sj.embor.embor778
1986:; Garrett, F. C. (1926).
1860:Columbia University Press
1391:10.1016/j.cub.2009.12.010
1088:10.1146/knowable-031822-1
1059:10.1007/s12052-008-0107-y
955:Encyclopedia of Evolution
952:Rice, Stanley A. (2007).
927:10.1007/s12052-008-0107-y
641:The Evolution of Melanism
410:) and the American form (
2313:by Michael E.N. Majerus"
2307:Coyne, Jerry A. (1998).
1342:Edleston, R. S. (1864).
862:Rudge, David W. (2005).
2966:Writers and researchers
2517:10.1126/science.1073593
1764:10.1126/science.1203043
1283:10.1073/pnas.0803785105
1149:www.millerandlevine.com
561:Kettlewell's experiment
555:Kettlewell's experiment
380:general selection model
139:Kettlewell's experiment
2851:Majerus, Michael E. N.
2750:Majerus, Michael E. N.
2538:Majerus, Michael E. N.
2105:10.1098/rspb.1933.0018
2059:10.1098/rspb.1928.0009
2036:Heslop-Harrison, J. W.
2005:10.1098/rspb.1926.0012
1984:Heslop-Harrison, J. W.
1920:Heslop Harrison, J. W.
1606:Majerus, Michael E. N.
1196:10.1098/rsbl.2011.1136
980:Majerus, Michael E. N.
903:Majerus, Michael E. N.
738:frequency compared to
683:
463:Alternative hypotheses
202:
57:
46:
2725:Why Evolution Is True
2462:Dover, Gabby (2003).
1883:Riley, P. A. (2013).
1718:10.1093/jhered/esh022
1697:Grant, B. S. (2004).
1488:Ecological Entomology
1104:"Evolution in Cities"
725:Michael E. N. Majerus
723:Further information:
670:
618:Further information:
213:Industrial Revolution
205:Further information:
185:
99:The dark-coloured or
90:teaching of evolution
80:was an early test of
70:Industrial Revolution
52:
37:
3095:Evolution of insects
2719:(12 February 2012).
2629:Iconsofevolution.com
2439:. 4 September 2003.
2284:Evolutionary Biology
1422:Miller, Ken (1999).
1344:"[No title]"
1143:Miller, Ken (1999).
719:Majerus's experiment
678:morph's increase in
544:Tephrosia bistortata
491:Phenotypic induction
448:transposable element
217:University of Oxford
178:Origin and evolution
23:evolutionary biology
3090:Biology experiments
2896:Moths Count website
2381:on 10 December 2014
2253:1969Natur.222..585S
2200:1968Sci...159..100S
2096:1933RSPSB.112..407F
1929:Journal of Genetics
1818:10.1038/nature17951
1810:2016Natur.534..102H
1756:2011Sci...332..958V
1704:Journal of Heredity
1315:Neal, Dick (2004).
1274:2008PNAS..10516212S
1268:(42): 16212–16217.
828:10.1038/hdy.2012.92
575:Nuffield Foundation
446:-TE, is a class II
416:autosomal dominants
402:is also a model of
389:population genetics
263:industrial melanism
207:Industrial melanism
74:industrial melanism
41:, the white-bodied
3034:Icons of Evolution
2973:Bernard Kettlewell
2619:Icons of Evolution
2430:"Of moths and men"
2375:"Of Moths and Men"
2150:10.1038/hdy.1958.4
1942:10.1007/BF02983273
1782:– via JSTOR.
1463:. 27 February 2018
1081:. Annual Reviews.
707:Icons of Evolution
691:intelligent design
687:Phillip E. Johnson
684:
567:Bernard Kettlewell
404:parallel evolution
398:The peppered moth
283:Charissa obscurata
203:
201:it from predators.
172:population biology
123:Bernard Kettlewell
58:
47:
3072:
3071:
3059:The Peppered Moth
2886:On 19 June 2009,
2765:978-0-198-54983-3
2696:The Panda's Thumb
2671:on 7 October 2008
2616:Wells J. (2000).
2575:on 26 August 2007
2553:978-1-84593-140-7
2511:(5583): 940–941,
2247:(5193): 585–586.
2194:(3810): 100–101.
1905:10.1111/bij.12062
1848:Dobzhansky, T. G.
1804:(7605): 102–117.
1750:(6032): 958–960.
1443:The Darwin Papers
1079:Knowable Magazine
995:978-0-19-854983-3
965:978-1-4381-1005-9
771:Scottish red deer
540:Selenia bilunaria
528:mutation pressure
365:natural selection
278:Gnophos obscurata
274:natural selection
86:natural selection
3107:
3064:Margaret Drabble
3041:Of Moths and Men
2926:
2919:
2912:
2903:
2883:
2878:
2857:Biston betularia
2842:presentation is
2810:
2808:
2806:
2789:Online lecture:
2769:
2736:
2735:
2733:
2731:
2713:
2707:
2706:
2704:
2702:
2687:
2681:
2680:
2678:
2676:
2667:. Archived from
2656:
2650:
2649:
2642:Matzke, Nicholas
2638:
2632:
2614:
2608:
2607:
2591:
2585:
2584:
2582:
2580:
2571:. Archived from
2564:
2558:
2557:
2534:
2528:
2527:
2500:
2494:
2493:
2491:
2459:
2453:
2452:
2450:
2448:
2432:
2425:
2419:
2418:
2416:
2414:
2397:
2391:
2390:
2388:
2386:
2371:
2365:
2364:
2362:
2360:
2343:
2337:
2336:
2334:
2304:
2298:
2297:
2279:
2273:
2272:
2261:10.1038/222585b0
2234:
2228:
2227:
2183:
2177:
2176:
2174:
2172:
2161:
2155:
2154:
2152:
2126:
2120:
2119:
2117:
2107:
2090:(778): 407–416.
2070:
2064:
2063:
2061:
2052:(718): 338–347.
2032:
2026:
2025:
2007:
1980:
1974:
1973:
1960:
1954:
1953:
1916:
1910:
1909:
1907:
1889:
1880:
1874:
1873:
1857:
1844:
1838:
1837:
1790:
1784:
1783:
1737:
1731:
1730:
1720:
1694:
1688:
1687:
1660:Bowler, Peter J.
1656:
1650:
1638:
1632:
1631:
1629:
1627:
1621:
1615:. Archived from
1614:
1602:
1589:
1588:
1552:
1541:
1540:
1538:
1536:
1518:
1512:
1511:
1484:Biston betularia
1479:
1473:
1472:
1470:
1468:
1453:
1447:
1446:
1434:
1428:
1420:
1411:
1410:
1374:
1365:
1356:
1355:
1339:
1333:
1332:
1312:
1306:
1305:
1295:
1285:
1251:
1245:
1244:
1224:
1218:
1217:
1207:
1169:
1160:
1159:
1157:
1155:
1140:
1134:
1133:
1123:
1099:
1093:
1092:
1090:
1070:
1064:
1063:
1061:
1040:Biston betularia
1033:
1027:
1026:
1006:
1000:
999:
976:
970:
969:
949:
940:
939:
929:
911:
899:
888:
887:
885:
859:
850:
849:
839:
807:
790:
787:
661:Of Moths and Men
628:Two chapters in
505:hydrogen sulfide
400:Biston betularia
393:modern synthesis
230:The Entomologist
159:Of Moths and Men
27:Insect evolution
3115:
3114:
3110:
3109:
3108:
3106:
3105:
3104:
3075:
3074:
3073:
3068:
3046:
3007:
2961:
2935:
2930:
2888:Telegraph.co.uk
2849:
2818:Michael Majerus
2804:
2802:
2796:
2776:
2766:
2748:
2745:
2743:Further reading
2740:
2739:
2729:
2727:
2715:
2714:
2710:
2700:
2698:
2689:
2688:
2684:
2674:
2672:
2665:The Independent
2658:
2657:
2653:
2640:
2639:
2635:
2615:
2611:
2593:
2592:
2588:
2578:
2576:
2566:
2565:
2561:
2554:
2536:
2535:
2531:
2502:
2501:
2497:
2461:
2460:
2456:
2446:
2444:
2436:The Independent
2427:
2426:
2422:
2412:
2410:
2399:
2398:
2394:
2384:
2382:
2373:
2372:
2368:
2358:
2356:
2353:Chicago Tribune
2345:
2344:
2340:
2325:(6706): 35–36.
2306:
2305:
2301:
2294:
2281:
2280:
2276:
2236:
2235:
2231:
2185:
2184:
2180:
2170:
2168:
2163:
2162:
2158:
2128:
2127:
2123:
2072:
2071:
2067:
2034:
2033:
2029:
1982:
1981:
1977:
1962:
1961:
1957:
1918:
1917:
1913:
1887:
1882:
1881:
1877:
1870:
1846:
1845:
1841:
1792:
1791:
1787:
1739:
1738:
1734:
1696:
1695:
1691:
1684:
1658:
1657:
1653:
1641:Haldane, J.B.S.
1639:
1635:
1625:
1623:
1622:on 15 June 2011
1619:
1612:
1608:(August 2007).
1604:
1603:
1592:
1554:
1553:
1544:
1534:
1532:
1520:
1519:
1515:
1481:
1480:
1476:
1466:
1464:
1455:
1454:
1450:
1436:
1435:
1431:
1421:
1414:
1378:Current Biology
1372:
1367:
1366:
1359:
1341:
1340:
1336:
1329:
1314:
1313:
1309:
1253:
1252:
1248:
1226:
1225:
1221:
1183:Biology Letters
1171:
1170:
1163:
1153:
1151:
1142:
1141:
1137:
1101:
1100:
1096:
1072:
1071:
1067:
1035:
1034:
1030:
1008:
1007:
1003:
996:
978:
977:
973:
966:
951:
950:
943:
909:
901:
900:
891:
861:
860:
853:
809:
808:
804:
799:
794:
793:
788:
784:
779:
762:
727:
721:
630:Michael Majerus
622:
616:
563:
557:
518:
510:
493:
469:Random mutation
465:
361:
312:
247:sulphur dioxide
209:
180:
168:Michael Majerus
30:
19:
12:
11:
5:
3113:
3111:
3103:
3102:
3097:
3092:
3087:
3077:
3076:
3070:
3069:
3067:
3066:
3054:
3052:
3051:Creative works
3048:
3047:
3045:
3044:
3037:
3030:
3023:
3015:
3013:
3009:
3008:
3006:
3005:
3000:
2998:Jonathan Wells
2995:
2990:
2985:
2980:
2975:
2969:
2967:
2963:
2962:
2960:
2959:
2957:Experiments on
2954:
2949:
2943:
2941:
2937:
2936:
2931:
2929:
2928:
2921:
2914:
2906:
2900:
2899:
2884:
2847:
2844:also available
2829:
2811:
2794:
2787:
2775:
2774:External links
2772:
2771:
2770:
2764:
2744:
2741:
2738:
2737:
2708:
2682:
2651:
2633:
2609:
2586:
2559:
2552:
2529:
2495:
2454:
2420:
2392:
2366:
2338:
2299:
2292:
2274:
2229:
2178:
2156:
2121:
2065:
2027:
1975:
1955:
1936:(3): 195–280.
1911:
1898:(2): 298–301.
1875:
1868:
1839:
1785:
1732:
1689:
1682:
1651:
1633:
1590:
1569:10.1086/378925
1563:(4): 399–417.
1542:
1513:
1494:(3): 231–243.
1474:
1448:
1429:
1412:
1357:
1334:
1327:
1307:
1246:
1235:(4): 301–322.
1219:
1190:(4): 609–612.
1161:
1135:
1114:(1): 519–540.
1094:
1065:
1028:
1017:(2): 189–199.
1001:
994:
971:
964:
941:
889:
876:(4): 369–375.
851:
822:(3): 207–212.
801:
800:
798:
795:
792:
791:
781:
780:
778:
775:
774:
773:
768:
761:
758:
720:
717:
696:Jonathan Wells
615:
612:
559:Main article:
556:
553:
516:
508:
492:
489:
464:
461:
376:J.B.S. Haldane
360:
357:
323:lepidopterists
311:
308:
270:Charles Darwin
179:
176:
145:evolution and
82:Charles Darwin
17:
13:
10:
9:
6:
4:
3:
2:
3112:
3101:
3098:
3096:
3093:
3091:
3088:
3086:
3085:Peppered moth
3083:
3082:
3080:
3065:
3061:
3060:
3056:
3055:
3053:
3049:
3043:
3042:
3038:
3036:
3035:
3031:
3029:
3028:
3024:
3022:
3021:
3017:
3016:
3014:
3010:
3004:
3003:Judith Hooper
3001:
2999:
2996:
2994:
2991:
2989:
2986:
2984:
2981:
2979:
2976:
2974:
2971:
2970:
2968:
2964:
2958:
2955:
2953:
2950:
2948:
2945:
2944:
2942:
2938:
2934:
2933:Peppered moth
2927:
2922:
2920:
2915:
2913:
2908:
2907:
2904:
2897:
2893:
2889:
2885:
2882:
2877:
2872:
2868:
2864:
2860:
2858:
2852:
2848:
2845:
2841:
2837:
2833:
2830:
2827:
2823:
2819:
2815:
2812:
2800:
2797:Young, Matt.
2795:
2792:
2788:
2785:
2784:his home page
2781:
2778:
2777:
2773:
2767:
2761:
2757:
2756:
2751:
2747:
2746:
2742:
2726:
2722:
2718:
2712:
2709:
2697:
2693:
2686:
2683:
2670:
2666:
2662:
2655:
2652:
2647:
2643:
2637:
2634:
2630:
2626:
2625:Regnery Press
2622:
2620:
2613:
2610:
2605:
2601:
2600:The Scientist
2597:
2590:
2587:
2574:
2570:
2563:
2560:
2555:
2549:
2545:
2544:
2539:
2533:
2530:
2526:
2522:
2518:
2514:
2510:
2506:
2499:
2496:
2490:
2485:
2481:
2477:
2473:
2469:
2465:
2464:"Mothbusters"
2458:
2455:
2442:
2438:
2437:
2431:
2424:
2421:
2409:
2408:
2403:
2396:
2393:
2380:
2376:
2370:
2367:
2355:
2354:
2349:
2342:
2339:
2333:
2332:10.1038/23856
2328:
2324:
2320:
2319:
2314:
2312:
2303:
2300:
2295:
2289:
2285:
2278:
2275:
2270:
2266:
2262:
2258:
2254:
2250:
2246:
2242:
2241:
2233:
2230:
2225:
2221:
2217:
2213:
2209:
2205:
2201:
2197:
2193:
2189:
2182:
2179:
2166:
2160:
2157:
2151:
2146:
2142:
2138:
2137:
2132:
2125:
2122:
2116:
2111:
2106:
2101:
2097:
2093:
2089:
2085:
2084:
2079:
2075:
2074:Fisher, R. A.
2069:
2066:
2060:
2055:
2051:
2047:
2046:
2041:
2037:
2031:
2028:
2023:
2019:
2015:
2011:
2006:
2001:
1997:
1993:
1989:
1985:
1979:
1976:
1971:
1968:(in German).
1967:
1959:
1956:
1951:
1947:
1943:
1939:
1935:
1931:
1930:
1925:
1921:
1915:
1912:
1906:
1901:
1897:
1893:
1886:
1879:
1876:
1871:
1869:0-231-08306-8
1865:
1861:
1856:
1855:
1849:
1843:
1840:
1835:
1831:
1827:
1823:
1819:
1815:
1811:
1807:
1803:
1799:
1798:
1789:
1786:
1781:
1777:
1773:
1769:
1765:
1761:
1757:
1753:
1749:
1745:
1744:
1736:
1733:
1728:
1724:
1719:
1714:
1711:(2): 97–102.
1710:
1706:
1705:
1700:
1693:
1690:
1685:
1683:0-520-23693-9
1679:
1675:
1671:
1667:
1666:
1661:
1655:
1652:
1648:
1647:
1642:
1637:
1634:
1618:
1611:
1607:
1601:
1599:
1597:
1595:
1591:
1586:
1582:
1578:
1574:
1570:
1566:
1562:
1558:
1551:
1549:
1547:
1543:
1531:
1527:
1523:
1517:
1514:
1509:
1505:
1501:
1497:
1493:
1489:
1485:
1478:
1475:
1462:
1461:Science Focus
1458:
1452:
1449:
1444:
1440:
1433:
1430:
1427:
1426:
1419:
1417:
1413:
1408:
1404:
1400:
1396:
1392:
1388:
1384:
1380:
1379:
1371:
1364:
1362:
1358:
1353:
1349:
1345:
1338:
1335:
1330:
1328:9780521532235
1324:
1320:
1319:
1311:
1308:
1303:
1299:
1294:
1289:
1284:
1279:
1275:
1271:
1267:
1263:
1262:
1257:
1250:
1247:
1242:
1238:
1234:
1230:
1223:
1220:
1215:
1211:
1206:
1201:
1197:
1193:
1189:
1185:
1184:
1179:
1175:
1174:Mallet, James
1168:
1166:
1162:
1150:
1146:
1139:
1136:
1131:
1127:
1122:
1117:
1113:
1109:
1105:
1098:
1095:
1089:
1084:
1080:
1076:
1069:
1066:
1060:
1055:
1051:
1047:
1043:
1041:
1032:
1029:
1024:
1020:
1016:
1012:
1005:
1002:
997:
991:
987:
986:
981:
975:
972:
967:
961:
957:
956:
948:
946:
942:
937:
933:
928:
923:
919:
915:
908:
904:
898:
896:
894:
890:
884:
879:
875:
871:
870:
865:
858:
856:
852:
847:
843:
838:
833:
829:
825:
821:
817:
813:
806:
803:
796:
786:
783:
776:
772:
769:
767:
764:
763:
759:
757:
755:
750:
748:
743:
741:
737:
733:
726:
716:
714:
709:
708:
703:
702:
701:The Scientist
697:
692:
688:
681:
677:
673:
669:
665:
663:
662:
657:
656:Judith Hooper
653:
651:
647:
643:
642:
637:
636:
632:'s 1998 book
631:
626:
621:
613:
611:
609:
604:
600:
596:
592:
588:
584:
580:
576:
572:
568:
562:
552:
550:
549:Ronald Fisher
545:
541:
537:
533:
529:
524:
522:
514:
506:
502:
498:
490:
488:
485:
480:
478:
477:genetic drift
474:
470:
462:
460:
458:
454:
449:
445:
441:
437:
432:
428:
424:
421:The gene for
419:
417:
413:
409:
408:f. carbonaria
405:
401:
396:
394:
390:
385:
381:
377:
372:
370:
366:
358:
356:
353:
351:
346:
341:
339:
335:
330:
328:
324:
320:
316:
309:
307:
305:
301:
297:
293:
292:Herefordshire
289:
285:
284:
279:
275:
272:'s theory of
271:
266:
264:
259:
255:
252:
248:
244:
239:
236:
232:
231:
226:
222:
218:
214:
208:
200:
196:
192:
188:
184:
177:
175:
173:
169:
165:
161:
160:
155:
154:Judith Hooper
150:
148:
144:
140:
136:
132:
128:
124:
120:
118:
114:
110:
106:
102:
97:
95:
94:Sewall Wright
91:
87:
83:
79:
78:peppered moth
75:
71:
67:
63:
55:
51:
44:
43:peppered moth
40:
36:
32:
28:
24:
16:
3057:
3039:
3032:
3025:
3018:
2983:Cyril Clarke
2978:Mike Majerus
2952:Evolution of
2951:
2880:
2869:(1): 63–74.
2866:
2862:
2856:
2803:. Retrieved
2754:
2728:. Retrieved
2724:
2717:Coyne, Jerry
2711:
2699:. Retrieved
2685:
2673:. Retrieved
2669:the original
2664:
2654:
2636:
2617:
2612:
2603:
2599:
2589:
2577:. Retrieved
2573:the original
2562:
2542:
2532:
2508:
2504:
2498:
2471:
2468:EMBO Reports
2467:
2457:
2445:. Retrieved
2434:
2423:
2411:. Retrieved
2407:The Guardian
2405:
2395:
2383:. Retrieved
2379:the original
2369:
2357:. Retrieved
2351:
2341:
2322:
2316:
2310:
2302:
2283:
2277:
2244:
2238:
2232:
2191:
2187:
2181:
2169:. Retrieved
2159:
2143:(1): 51–72.
2140:
2134:
2124:
2087:
2081:
2068:
2049:
2043:
2030:
1995:
1991:
1978:
1969:
1965:
1958:
1933:
1927:
1914:
1895:
1891:
1878:
1853:
1842:
1801:
1795:
1788:
1747:
1741:
1735:
1708:
1702:
1692:
1664:
1654:
1644:
1636:
1624:. Retrieved
1617:the original
1560:
1556:
1533:. Retrieved
1530:Talk Origins
1529:
1522:Grant, Bruce
1516:
1491:
1487:
1483:
1477:
1465:. Retrieved
1460:
1451:
1442:
1432:
1423:
1382:
1376:
1351:
1348:Entomologist
1347:
1337:
1317:
1310:
1265:
1259:
1249:
1232:
1228:
1222:
1187:
1181:
1152:. Retrieved
1148:
1138:
1111:
1107:
1097:
1078:
1068:
1052:(1): 63–74.
1049:
1045:
1039:
1031:
1014:
1010:
1004:
984:
974:
954:
920:(1): 63–74.
917:
913:
873:
867:
819:
815:
805:
785:
766:Polymorphism
753:
751:
747:James Mallet
744:
739:
735:
728:
705:
699:
685:
675:
672:Creationists
659:
654:
649:
639:
633:
627:
623:
602:
586:
564:
543:
539:
525:
494:
481:
466:
456:
452:
443:
439:
435:
427:B. betularia
426:
422:
420:
412:f. swettaria
411:
407:
399:
397:
373:
368:
362:
354:
344:
342:
333:
331:
313:
281:
277:
267:
240:
228:
210:
199:camouflaging
194:
190:
186:
164:creationists
157:
151:
121:
112:
104:
100:
98:
66:evolutionary
61:
59:
53:
38:
31:
15:
2988:Bruce Grant
2780:Bruce Grant
2675:9 September
2447:10 December
2413:10 December
2385:10 December
2359:10 December
1672:. pp.
1626:9 September
713:Nick Matzke
646:Jerry Coyne
587:Parus major
350:Bruce Grant
258:camouflaged
211:Before the
92:. In 1978,
3079:Categories
2993:E. B. Ford
2892:an article
2890:published
2840:PowerPoint
2826:Darwin Day
2474:(3): 235.
2293:0306429772
2115:2440/15114
1385:(3): R95.
869:BioScience
797:References
736:carbonaria
676:carbonaria
614:Criticisms
595:Birmingham
583:great tits
571:E. B. Ford
501:germ plasm
440:Drosophila
431:chromosome
423:carbonaria
384:stochastic
345:carbonaria
334:carbonaria
327:moth traps
304:J. W. Tutt
300:lime kilns
288:New Forest
221:Manchester
191:carbonaria
135:Birmingham
127:camouflage
117:J. W. Tutt
109:Manchester
105:carbonaria
3100:Selection
2805:3 January
2606:(11): 13.
2579:26 August
2525:161367302
2014:0950-1193
1130:239646134
680:frequency
536:manganese
495:In 1920,
484:chelation
473:migration
374:In 1924,
319:phenotype
254:ecosystem
143:Darwinian
2853:(2009).
2752:(1998).
2441:Archived
2224:32124765
2136:Heredity
2076:(1933).
2038:(1928).
2022:84987348
1950:38996034
1922:(1920).
1850:(1937).
1826:27251284
1780:24400858
1772:21493823
1727:15073224
1662:(2003).
1643:(1924).
1585:26831926
1577:14737825
1508:85624115
1407:31093691
1399:20144776
1302:18854412
1214:22319093
1176:(2012).
1154:31 March
982:(1998).
936:25407417
905:(2008).
846:23211788
816:Heredity
760:See also
756:story."
658:'s book
438:gene in
359:Genetics
338:mutation
315:Melanism
251:polluted
2940:Biology
2730:7 March
2701:7 March
2505:Science
2489:1315906
2269:4202131
2249:Bibcode
2216:5634373
2196:Bibcode
2188:Science
2092:Bibcode
1834:3989607
1806:Bibcode
1752:Bibcode
1743:Science
1674:331–332
1486:(L.)".
1467:16 June
1293:2571026
1270:Bibcode
1205:3391436
837:3668657
599:England
521:pyredin
519:) and "
513:ammonia
455:moths.
238:0.01%.
235:lichens
225:England
101:melanic
2762:
2550:
2523:
2486:
2318:Nature
2290:
2267:
2240:Nature
2222:
2214:
2171:31 May
2020:
2012:
1948:
1866:
1832:
1824:
1797:Nature
1778:
1770:
1725:
1680:
1583:
1575:
1535:2 June
1506:
1405:
1397:
1354:: 150.
1325:
1300:
1290:
1212:
1202:
1128:
992:
962:
934:
844:
834:
754:Biston
740:typica
650:Biston
608:Dorset
603:typica
579:aviary
453:typica
436:cortex
369:typica
325:using
280:, now
195:typica
187:Typica
131:Dorset
113:typica
64:is an
3012:Works
2828:2004.
2521:S2CID
2265:S2CID
2220:S2CID
2018:S2CID
1946:S2CID
1888:(PDF)
1830:S2CID
1776:S2CID
1620:(PDF)
1613:(PDF)
1581:S2CID
1504:S2CID
1403:S2CID
1373:(PDF)
1126:S2CID
932:S2CID
910:(PDF)
777:Notes
457:Carb-
296:Lewes
2807:2009
2760:ISBN
2732:2012
2703:2012
2677:2007
2581:2007
2548:ISBN
2449:2014
2415:2014
2387:2014
2361:2014
2288:ISBN
2212:PMID
2173:2022
2010:ISSN
1864:ISBN
1822:PMID
1768:PMID
1723:PMID
1678:ISBN
1628:2007
1573:PMID
1537:2022
1469:2018
1395:PMID
1323:ISBN
1298:PMID
1261:PNAS
1210:PMID
1156:2022
990:ISBN
960:ISBN
842:PMID
732:bats
542:and
534:and
532:lead
511:S),
444:carb
243:soot
189:and
60:The
3062:by
2871:doi
2824:on
2513:doi
2509:297
2484:PMC
2476:doi
2327:doi
2323:396
2257:doi
2245:222
2204:doi
2192:159
2145:doi
2110:hdl
2100:doi
2088:112
2054:doi
2050:102
2000:doi
1938:doi
1900:doi
1896:109
1814:doi
1802:534
1760:doi
1748:332
1713:doi
1565:doi
1496:doi
1387:doi
1288:PMC
1278:doi
1266:105
1237:doi
1200:PMC
1192:doi
1116:doi
1083:doi
1054:doi
1019:doi
922:doi
878:doi
832:PMC
824:doi
820:110
593:in
515:(NH
475:or
425:in
265:".
156:'s
84:'s
3081::
2879:.
2865:.
2861:.
2723:.
2694:.
2663:.
2644:.
2623:.
2604:13
2602:.
2598:.
2519:,
2507:,
2482:.
2470:.
2466:.
2433:.
2404:.
2350:.
2321:.
2315:.
2263:.
2255:.
2243:.
2218:.
2210:.
2202:.
2190:.
2141:12
2139:.
2133:.
2108:.
2098:.
2086:.
2080:.
2048:.
2042:.
2016:.
2008:.
1996:99
1994:.
1990:.
1970:19
1944:.
1932:.
1926:.
1894:.
1890:.
1862:.
1858:.
1828:.
1820:.
1812:.
1800:.
1774:.
1766:.
1758:.
1746:.
1721:.
1709:95
1707:.
1701:.
1676:.
1668:.
1593:^
1579:.
1571:.
1561:78
1559:.
1545:^
1528:.
1524:.
1502:.
1490:.
1459:.
1441:.
1415:^
1401:.
1393:.
1383:20
1381:.
1375:.
1360:^
1350:.
1346:.
1296:.
1286:.
1276:.
1264:.
1258:.
1233:39
1231:.
1208:.
1198:.
1186:.
1180:.
1164:^
1147:.
1124:.
1112:52
1110:.
1106:.
1077:.
1048:.
1044:.
1015:26
1013:.
944:^
930:.
916:.
912:.
892:^
874:55
872:.
866:.
854:^
840:.
830:.
818:.
814:.
682:.
597:,
507:(H
471:,
329:.
223:,
166:.
2925:e
2918:t
2911:v
2873::
2867:2
2846:.
2809:.
2786:.
2768:.
2734:.
2705:.
2679:.
2648:.
2631:)
2583:.
2556:.
2515::
2492:.
2478::
2472:4
2451:.
2417:.
2389:.
2363:.
2335:.
2329::
2296:.
2271:.
2259::
2251::
2226:.
2206::
2198::
2175:.
2153:.
2147::
2118:.
2112::
2102::
2094::
2062:.
2056::
2024:.
2002::
1972:.
1952:.
1940::
1934:9
1908:.
1902::
1872:.
1836:.
1816::
1808::
1762::
1754::
1729:.
1715::
1686:.
1649:.
1630:.
1587:.
1567::
1539:.
1510:.
1498::
1492:2
1471:.
1409:.
1389::
1352:2
1331:.
1304:.
1280::
1272::
1243:.
1239::
1216:.
1194::
1188:8
1158:.
1132:.
1118::
1091:.
1085::
1062:.
1056::
1050:2
1025:.
1021::
998:.
968:.
938:.
924::
918:2
886:.
880::
848:.
826::
585:(
517:3
509:2
45:.
29:.
Text is available under the Creative Commons Attribution-ShareAlike License. Additional terms may apply.
