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Kettlewell's experiment

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460:, said Kettlewell's experiments had appeared to be "the slam-dunk of natural selection", but argued that the cause of the dark forms appearing was still an "irreducible mystery". Although not a creationist herself, Hooper argued that the peppered moth experiments failed to represent evolution. She claimed 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. She then alleged that scientists in general showed "credulous and biased" acceptance of evolution. The book's reception led to claims that the peppered moth evolution story ought to be deleted from textbooks. 440:, 14 March 1999, claiming that "the rise and fall of the peppered moth, is based on a series of scientific blunders. Experiments using the moth in the Fifties and long believed to prove the truth of natural selection are now thought to be worthless, having been designed to come up with the 'right' answer." Majerus regarded this view as surprising, and not one that would be shared by those involved in the field. He noted numerous scientific inaccuracies, misquotations and misrepresentations in the article, but thought this was common in press reports. He stated that he had spoken to Matthews for over half an hour and had to explain many details as Matthews hadn't read the book, but "Even then, he got nearly everything wrong." 411:, he stated that the most serious problem was that only two peppered moths had been found on tree trunks. He also wrote that the white moths had increased in numbers before the lichen had returned and that Kettlewell's findings of moths choosing matching backgrounds had not been replicated in later experiments. Coyne compared his reaction to "the dismay attending my discovery, at the age of 6, that it was my father and not Santa who brought the presents on Christmas Eve". He concluded that "for the time being we must discard 173: 20: 473: 420:
on tree trunks was incorrect, as the book gives the resting positions of 47 peppered moths Majerus had found in the wild between 1964 and 1996; twelve were on tree trunks (six exposed, six unexposed), twenty were at the trunk/branch joint, and fifteen resting on branches. Majerus found that the review did not reflect the factual content of the book or his own views, and cited an assessment by the
249:) to prey upon. He found that the initial procedure failed, as the birds actively looked for any moth, regardless of their colour or background. He succeeded only by using freshly captured moths consisting of 9 black and 8 white types, which he released separately. He found that the birds preferentially caught the moths according to the background colour on which the moths were present. 152:. He estimated that for the peppered moth having reproductive cycle in a year, it would take 48 generations to produce the dominant (melanic or black) forms, and the melanic population could dominate the entire moth population after 13 generations. He concluded that "the only probable explanation is the not very intense degree of natural selection". University of Oxford zoologist 385:, published a critique of Kettlewell's work. Based on his experiments between 1965 and 1969, he concluded that it was not possible to reproduce Kettlewell's results, and said that birds showed no preference on moth on either black or white tree trunks. He suggested that Kettlewell had trained the birds to pick moths on tree trunks to obtain desired results. 524:, in which he claims, "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." The arguments were dismissed by Majerus, Cook and 533:
are unstaged pictures of live moths in the wild, and the photographs of moths on tree-trunks, apart from some slight blurring, look little different from the "staged" photographs. While an experiment did involve the gluing of dead moths to trees, this practice was just one of many different ways used
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The experiment found that birds selectively prey on peppered moths depending on their body colour in relation to their environmental background. Thus, the evolution of a dark-coloured body provided a survival advantage in a polluted locality. The study concluded that "industrial melanism in moths is
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Contrary to this review, Majerus had stressed that the basic findings from that work were correct, and that differential bird predation of polluted environment "is the primary influence of the evolution of melanism in the peppered moth". Coyne's statement that only two peppered moths had been found
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Kettlewell first devised a standard procedure for scoring the moths. It was necessary to determine how far apart the moths should be placed so they were indistinguishable from their backgrounds. Correct separation would result in effective and selective predation by birds, because if the moths were
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type. After several days, he found that out the total 190 moths eaten by birds in the wild, 86% were the black type while only 14% were the white type. In his release-and-recapture experiment for 11 days, he used 799 moths, and his recapture rate was 13.7% for the white type, but only 4.7% for the
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17%. (The total capture after release was 770, but 621 of them were non-experimental moths, i.e. not bearing the paint marks.) Thus their survival values were 5.72%, 1.48%, and 4.32% respectively. This shows that black moths had the best survival advantage in a darkened and polluted environment.
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who describes Wells as distorting the picture by selectively omitting or scrambling references in a way that is dishonest. Professional photography to illustrate textbooks uses dead insects because of the considerable difficulty in getting good images of both forms of moth in the same shot. The
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Scientists have examined the allegations made by Hooper, and found them to be without merit. Majerus described the book as "littered with errors, misrepresentations, misinterpretations and falsehoods". David W. Rudge, after critical analyses of Kettlewell' works, declared that "none of Hooper's
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arguments is found to withstand careful scrutiny", and that all "these charges are baseless and stem from a fundamental misunderstanding of the nature of science as a process." He concluded "that Hooper does not provide one shred of evidence to support this serious allegation."
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to study different individual elements of the overall hypothesis. This particular experiment was not meant to exactly reproduce natural conditions but instead was used to assess how the numbers of moths available (their density) affected the foraging practices of birds.
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of 24 May 1999, claiming that "In 25 years of fieldwork, C.A. Clarke and his colleagues found only one peppered moth on a tree trunk", and concluding that "The fact that peppered moths do not normally rest on tree trunks invalidates Kettlewell's experiments".
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paint, so that he would be able to identify them later from non-experimental individuals after recapture. He started capturing the moths on the night of 26 June 1953, and lasted till 5 July. Out of his total capture, he selected 630 (447
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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" and that further studies of the animal's habits were needed.
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said that the moths "do not sit on tree trunks", "moths had to be glued to the trunks" for pictures and that the experiments were "fraudulent" and a "scam." This led Frack to exchange with intelligent design proponent
508:, who conceded that Majerus listed six moths on exposed tree trunks (out of 47), but argued that this was "an insignificant proportion". Wells wrote an essay on the subject, a shortened version of which appeared in 299:
To compare the conditions of predation in different environments Kettlewell planned to perform similar experiment in both polluted and clean environments. In 1954 he surveyed several woodlands including
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all too close then birds would be able to differentiate even well camouflaged individuals. He tested his scoring method in the woodlands near Birmingham by releasing 651 peppered moths (consisting of
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were covered with black soot. R.S. Edleston was the first to identify the unusual black peppered moth in 1848 in Manchester. By the end of the century, it was recorded that the black moth, the
148: 261:, Birmingham, because it was heavily polluted, but still inhabited by a number of bird species. He caught all three types of peppered moth and marked them underneath their wings with 1197:
Sargent, T.D.; Millar, C.D.; Lambert, D.M. (1988). "Ch. 9: The 'classical' explanation of industrial melanism: Assessing the evidence". In Hecht, Max K.; Wallace, Bruce (eds.).
335:. Tinbergen was responsible for filming the experiment, particularly to verify whether or not birds were the main predators. Out of 227 moths they released, 154 were 138:
was the first to come up with natural selection as an explanation, and stated in 1894 that the phenomenon was due to selective predation by birds. With the rise of
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designed by Wells and other ID scholars. (In the book Wells accused Kettlewell's experiment as "fraudulent" and "staged".) Coyne and Grant wrote a letter to
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In clean and lichened area, dark moths remained scarce and were rapidly eliminated because of their conspicuousness even when experimentally introduced.
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the most striking evolutionary phenomenon ever actually witnessed in any organism, animal or plant." It is now regarded as the classic demonstration of
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The main experiment, called mark-release-recapture, started in the summer of 1953 and lasted for three years. It consists of two continuous phases.
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Moth were eaten by birds selectively in both polluted and clean forests, indicating camouflage efficiency of the different varieties of moths.
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The more conspicuous form of moth was always less in number after recapture; i.e. the white type in Birmingham, and the black type in Dorset.
382: 1348: 1206: 188:, is whitish-grey in colour with dark speckles on the wings. The colour was a perfect camouflage on light-coloured trees covered with 1866: 1836: 1435: 1368: 635: 1751: 139: 347:. The recapture rate within two days was 82%, 16%, and 2% respectively. It was another success, with Tinbergen capturing live 529:
scientific studies actually consisted of observational data rather than using such photographs. The photographs in Majerus's
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that there was essentially no resemblance between the book and Coyne's review, which appeared to be a summary of the Sargent
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Sargent, T. D. (1969). "Background Selections of the Pale and Melanic Forms of the Cryptic Moth, Phigalia titea (Cramer)".
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Hagen, Joel B. (1999). "Retelling experiments: H.B.D. Kettlewell's studies of industrial melanism in peppered moths".
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When serious criticism and controversy arose, the story was picked up by creationists. Coyne's review was taken up by
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in that it was not whitish. The moths were active at night, and rested on tree trunks and boughs during the day.
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By the time of Kettlewell, it was known in England that there were three varieties of peppered moth. The normal,
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Sargent, T. D. (1968). "Cryptic moths: effects on background selections of painting the circumocular scales".
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continued efforts to favor intelligent design teaching by requiring the use of alternative resources, such as
1897:"Second Thoughts about Peppered Moths; This classical story of evolution by natural selection needs revising" 1494: 2132: 2096: 1942: 525: 172: 19: 2229: 1571: 146:
estimated in 1924 the rate of evolution by natural selection in the peppered moth in his first series of
2054: 1946: 278:) male moths and released them into the woods. Within two days, 149 moths were recaptured, out of which 67: 39: 320:
form, indicating clean environment. In mid-June 1955 he started the experiment. He brought along 3,000
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in England in the 19th century. He conducted his first experiment in 1953 in the polluted woodland of
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The review was subsequently picked up by the journalist Robert Matthews, who wrote an article for
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Jonathan Wells's book Icons of Evolution and why most of what it teaches about evolution is wrong
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supported the bird-predation hypothesis. To experimentally investigate the issue he recruited
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Immediately after completion, he headed back to Birmingham, now accompanied by the renowned
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Cook, Laurence M. (2003). "The rise and fall of the Carbonaria form of the peppered moth".
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Evolution and the Nature of Science Institutes for High School Biology Teachers: Resources
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in which they defended the moth experiments and revealed the misrepresentations by Wells.
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For his first experimental site, Kettlewell chose Christopher Cadbury Bird Reserve, near
1140: 1089: 999: 574:"The peppered moth and industrial melanism: evolution of a natural selection case study" 2026: 1623: 1598: 824: 598: 573: 496: 332: 122:. There were conflicting ideas as to the biological basis of this industrial melanism. 83: 2223: 2147: 2091: 2077: 1925: 1702: 1638:
Cook, L. M. (2003). "The rise and fall of the Carbonaria form of the peppered moth".
1466: 877: 457: 99: 51: 1896: 1813: 1786: 1667: 1523: 1306: 1113: 793: 750: 698:"The Beauty of Kettlewell's Classic Experimental Demonstration of Natural Selection" 2127: 1156: 1023: 74:, and his second experiment in 1955 in Birmingham as well as in the clean woods of 1719: 1171: 1097: 957: 625: 316:, which was well covered with lichens. He found only the white type, including 1 1302: 476: 404: 472: 312:
forms, which indicated unclean environment. Finally he went to Deanend Wood in
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in action and one of the most beautiful experiments in evolutionary biology.
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The conclusion of Kettlewell's experiment can be summarised as follows:
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in Cambridge. In the aviary he released 69 moths, which he allowed two
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Darwin, Then and Now: The Most Amazing Story in the History of Science
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type, outnumbered (90% in some regions) the natural white ones, named
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Rudge, David W. (2006). "Myths about Moths: a Study in Contrasts".
1651: 777: 1280: 1259: 301: 200:, which was light-coloured with speckled wings, but distinct from 171: 18: 1468:
Of Moths and Men : Intrigue, Tragedy & the Peppered Moth
658:"Selection experiments on industrial melanism in the Lepidoptera" 2046: 348: 2050: 196:, was completely black. There was an intermediate form, called 1547:"Of Moths and Men: Intrigue, Tragedy & the Peppered Moth" 807:
Berry, R.J. (1990). "Industrial melanism and peppered moths (
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have disputed the occurrence or significance of the melanic
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10.1641/0006-3568(2005)055[0369:TBOKCE]2.0.CO;2
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Theodore David Sargent, professor of zoology at the
2195: 2156: 2110: 2084: 1369:"Peppered Moths – in black and white (part 2 of 2)" 925:(L.) (Lep.) and its melanic forms in Great Britain" 499:on 13 March 1999, creationist and professor of law 142:, the theoretical background was set. For example, 1465: 1928:, Washington, D.C., p. 138 (book available from 130:, disease, temperature and protection (such as 495:creationists, and at a seminar presenting the 2062: 1681:Grant, B. S. (2002), "Sour grapes of wrath", 8: 1969:"Intelligent Design in Pratt County, Kansas" 1745: 1743: 1522:. W. W. Norton & Company. Archived from 2033:. Department of Biology, Indiana University 2069: 2055: 2047: 651: 649: 647: 16:Biological experiment on the peppered moth 1992:Padian, Kevin; Gishlick, Alan D. (2002). 1622: 1279: 1060: 1035: 1033: 962:. New York: Academic Press. p. 173. 940: 911: 909: 813:Biological Journal of the Linnean Society 713: 691: 689: 687: 685: 673: 597: 537:On 27 November 2000, the school board of 1362: 1360: 1176:. 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Archived from 1403:Majerus, Michael 1399: 1388: 1387: 1385: 1384: 1375:. Archived from 1364: 1355: 1354: 1331: 1325: 1324: 1322: 1321: 1299: 1286: 1285: 1283: 1255: 1249: 1248: 1246: 1244: 1238: 1232:. Archived from 1231: 1219: 1213: 1212: 1194: 1188: 1187: 1167: 1161: 1160: 1149:10.1038/222585b0 1124: 1118: 1117: 1073: 1067: 1066: 1064: 1037: 1028: 1027: 1008:10.1038/175943a0 980: 974: 973: 953: 947: 946: 944: 923:Biston betularia 913: 904: 903: 901: 899: 888: 882: 881: 856:(1–2): 209–240. 847: 835: 829: 828: 809:Biston betularia 804: 798: 797: 761: 755: 754: 726: 720: 719: 717: 693: 680: 679: 677: 653: 642: 641: 621: 612: 611: 601: 569: 453:Of Moths and Men 445:Of Moths and Men 192:. 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Haldane 95: 92: 84:Charles Darwin 15: 13: 10: 9: 6: 4: 3: 2: 2253: 2252: 2241: 2238: 2236: 2233: 2231: 2230:Peppered moth 2228: 2227: 2225: 2210: 2206: 2205: 2201: 2200: 2198: 2194: 2188: 2187: 2183: 2181: 2180: 2176: 2174: 2173: 2169: 2167: 2166: 2162: 2161: 2159: 2155: 2149: 2148:Judith Hooper 2146: 2144: 2141: 2139: 2136: 2134: 2131: 2129: 2126: 2124: 2121: 2119: 2116: 2115: 2113: 2109: 2103: 2100: 2098: 2095: 2093: 2090: 2089: 2087: 2083: 2079: 2078:Peppered moth 2072: 2067: 2065: 2060: 2058: 2053: 2052: 2049: 2032: 2028: 2022: 2019: 2014: 2010: 2006: 2002: 1995: 1988: 1985: 1974: 1970: 1963: 1960: 1948: 1944: 1938: 1935: 1931: 1927: 1926:Regnery Press 1923: 1921: 1914: 1911: 1906: 1902: 1901:The Scientist 1898: 1891: 1888: 1877:on 2007-09-27 1876: 1872: 1868: 1861: 1858: 1847:on 2007-08-26 1846: 1842: 1838: 1831: 1828: 1823: 1819: 1815: 1811: 1807: 1803: 1796: 1793: 1788: 1784: 1780: 1776: 1772: 1768: 1764: 1760: 1753: 1746: 1744: 1740: 1735: 1729: 1725: 1721: 1717: 1711: 1708: 1704: 1700: 1696: 1692: 1688: 1684: 1677: 1674: 1669: 1665: 1661: 1657: 1653: 1649: 1645: 1641: 1634: 1631: 1625: 1620: 1616: 1612: 1608: 1604: 1600: 1599:"Mothbusters" 1593: 1590: 1577: 1573: 1567: 1564: 1552: 1548: 1541: 1538: 1525: 1521: 1515: 1512: 1500: 1496: 1489: 1486: 1481: 1475: 1470: 1469: 1460: 1457: 1446:on 2007-09-27 1445: 1441: 1437: 1430: 1427: 1416:on 2007-09-26 1412: 1408: 1404: 1398: 1396: 1394: 1390: 1379:on 2007-09-30 1378: 1374: 1370: 1363: 1361: 1357: 1352: 1346: 1342: 1341: 1336: 1330: 1327: 1316: 1312: 1308: 1305:(2002–2004). 1304: 1298: 1296: 1294: 1292: 1288: 1282: 1281:10.1038/23856 1277: 1273: 1269: 1265: 1263: 1254: 1251: 1235: 1228: 1224: 1218: 1215: 1210: 1204: 1200: 1193: 1190: 1185: 1179: 1175: 1174: 1166: 1163: 1158: 1154: 1150: 1146: 1142: 1138: 1134: 1130: 1123: 1120: 1115: 1111: 1107: 1103: 1099: 1095: 1091: 1087: 1083: 1079: 1072: 1069: 1063: 1058: 1054: 1050: 1046: 1042: 1036: 1034: 1030: 1025: 1021: 1017: 1013: 1009: 1005: 1001: 997: 993: 989: 985: 979: 976: 971: 965: 961: 960: 952: 949: 943: 938: 934: 930: 926: 924: 918: 912: 910: 906: 893: 887: 884: 879: 875: 871: 867: 863: 859: 855: 851: 844: 840: 834: 831: 826: 822: 818: 814: 810: 803: 800: 795: 791: 787: 783: 779: 775: 771: 767: 760: 757: 752: 748: 744: 740: 736: 732: 725: 722: 716: 711: 707: 703: 699: 692: 690: 688: 686: 682: 676: 671: 667: 663: 659: 652: 650: 648: 644: 639: 637:9780595515752 633: 629: 628: 620: 618: 614: 609: 605: 600: 595: 591: 587: 583: 579: 575: 568: 566: 562: 556: 554: 552: 551: 546: 545: 540: 535: 532: 527: 523: 522: 516: 513: 512: 511:The Scientist 507: 502: 498: 494: 486: 482: 478: 474: 467: 465: 461: 459: 458:Judith Hooper 455: 454: 446: 443: 441: 439: 438: 432: 430: 426: 423: 417: 414: 410: 406: 402: 401: 396: 395: 391:'s 1998 book 390: 386: 384: 376: 371: 368: 365: 362: 361: 360: 354: 352: 350: 346: 342: 338: 334: 331: 326: 323: 319: 315: 311: 307: 303: 294: 292: 289: 285: 281: 277: 273: 269: 264: 260: 252: 250: 248: 244: 240: 236: 232: 228: 224: 215: 213: 207: 205: 203: 199: 195: 191: 187: 179: 174: 167: 165: 163: 159: 155: 151: 150: 145: 141: 137: 133: 129: 125: 121: 117: 113: 109: 105: 101: 100:Great Britain 93: 91: 89: 85: 79: 77: 73: 69: 65: 61: 57: 53: 52:peppered moth 49: 46:mechanism of 45: 41: 37: 30: 26: 21: 2202: 2184: 2177: 2170: 2163: 2128:Cyril Clarke 2123:Mike Majerus 2101: 2097:Evolution of 2035:. 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Tutt 132:camouflage 116:carbonaria 112:Birmingham 108:Manchester 94:Background 72:Birmingham 25:carbonaria 1907:(11): 13. 1871:Evolution 1841:Evolution 1802:Endeavour 1703:161367302 1440:Evolution 1373:Evolution 878:189884360 841:(1990) . 485:frequency 345:insularia 318:insularia 288:insularia 286:13%, and 276:insularia 274:, and 46 263:cellulose 239:Madingley 231:insularia 198:insularia 178:insularia 104:pollution 1822:16549216 1787:25525719 1779:16240545 1718:(2005). 1668:26831926 1660:14737825 1405:(2004). 1337:(1998). 1114:32124765 1049:Heredity 1043:(1956). 1016:14383775 929:Heredity 919:(1958). 811:(L.))". 794:26831926 786:14737825 751:81591392 662:Heredity 608:23211788 578:Heredity 343:, and 9 306:Cornwall 128:heredity 124:Humidity 2085:Biology 1683:Science 1624:1315906 1157:4202131 1137:Bibcode 1106:5634373 1086:Bibcode 1078:Science 1024:4169783 996:Bibcode 870:2185859 599:3668657 190:lichens 50:in the 1820:  1785:  1777:  1730:  1701:  1666:  1658:  1621:  1476:  1414:(.doc) 1347:  1268:Nature 1205:  1180:  1155:  1129:Nature 1112:  1104:  1022:  1014:  988:Nature 966:  876:  868:  792:  784:  749:  634:  606:  596:  413:Biston 409:Nature 341:typica 314:Dorset 284:typica 272:typica 270:, 137 259:Rubery 235:aviary 223:typica 202:typica 186:typica 120:typica 76:Dorset 38:was a 29:typica 2157:Works 1997:(PDF) 1783:S2CID 1755:(PDF) 1699:S2CID 1664:S2CID 1237:(PDF) 1230:(PDF) 1153:S2CID 1110:S2CID 1020:S2CID 874:S2CID 846:(PDF) 790:S2CID 747:S2CID 429:et al 349:movie 339:, 64 302:Devon 1818:PMID 1775:PMID 1728:ISBN 1656:PMID 1584:2014 1558:2014 1532:2014 1506:2014 1474:ISBN 1345:ISBN 1245:2007 1203:ISBN 1178:ISBN 1102:PMID 1012:PMID 964:ISBN 900:2014 866:PMID 782:PMID 632:ISBN 604:PMID 304:and 229:and 110:and 2207:by 2009:doi 1810:doi 1767:doi 1691:doi 1687:297 1648:doi 1619:PMC 1611:doi 1276:doi 1272:396 1145:doi 1133:222 1094:doi 1082:159 1057:doi 1004:doi 992:175 937:doi 858:doi 821:doi 774:doi 739:doi 710:doi 670:doi 594:PMC 586:doi 582:110 86:'s 2226:: 2029:. 2005:77 2003:. 1999:. 1971:. 1924:. 1905:13 1903:. 1899:. 1869:. 1839:. 1816:. 1806:30 1804:. 1781:. 1773:. 1763:14 1761:. 1757:. 1742:^ 1697:, 1685:, 1662:. 1654:. 1644:78 1642:. 1617:. 1605:. 1601:. 1574:. 1549:. 1497:. 1438:. 1392:^ 1371:. 1359:^ 1313:. 1309:. 1290:^ 1270:. 1266:. 1151:. 1143:. 1131:. 1108:. 1100:. 1092:. 1080:. 1053:10 1051:. 1047:. 1032:^ 1018:. 1010:. 1002:. 990:. 933:12 931:. 927:. 908:^ 872:. 864:. 854:52 852:. 848:. 817:39 815:. 788:. 780:. 770:78 768:. 745:. 735:14 733:. 706:55 704:. 700:. 684:^ 664:. 660:. 646:^ 616:^ 602:. 592:. 580:. 576:. 564:^ 225:, 164:. 78:. 2070:e 2063:t 2056:v 2040:. 2015:. 2011:: 1981:. 1956:. 1932:) 1884:. 1854:. 1824:. 1812:: 1789:. 1769:: 1736:. 1693:: 1670:. 1650:: 1627:. 1613:: 1607:4 1586:. 1560:. 1534:. 1508:. 1482:. 1453:. 1423:. 1386:. 1353:. 1323:. 1284:. 1278:: 1247:. 1211:. 1186:. 1159:. 1147:: 1139:: 1116:. 1096:: 1088:: 1065:. 1059:: 1026:. 1006:: 998:: 972:. 945:. 939:: 902:. 880:. 860:: 827:. 823:: 796:. 776:: 753:. 741:: 718:. 712:: 678:. 672:: 666:9 640:. 610:. 588:: 487:. 245:( 54:(

Index


biological experiment
evolutionary
industrial melanism
peppered moth
Bernard Kettlewell
University of Oxford
Industrial Revolution
Birmingham
Dorset
Charles Darwin
natural selection
Great Britain
pollution
Manchester
Birmingham
Humidity
heredity
camouflage
J. W. Tutt
evolutionary statistics
J.B.S. Haldane
A Mathematical Theory of Natural and Artificial Selection
E. B. Ford
Bernard Kettlewell
Nuffield Foundation

lichens
aviary
Madingley

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