432:
417:
67:
1080:
1048:
942:
1064:
646:
638:
85:
1032:
410:(spotting) can also, in rare instances, be gray or brown; the spotting pattern, in particularly very rare cases, is sometimes a combination of brown and black/gray. The black speckling varies in amount, in some examples it is almost absent, whilst in others it is so dense that the wings appear to be black sprinkled with white. The antennae of males are strongly bipectinate. Prout (1912–16) gives an account of the forms and congeners.
927:
53:
522:
679:
From their original data, Howlett and
Majerus (1987) concluded that peppered moths generally rest in unexposed positions, using three main types of site. Firstly, a few inches below a branch-trunk joint on a tree trunk where the moth is in shadow; secondly, on the underside of branches and thirdly on
601:
in 1887. Research indicates that the caterpillars can sense the twig's colour with their skin and match their body colour to the background to protect themselves from predators, an ability to camouflage themselves also found in cephalopods, chameleons and some fish, although this colour change is
409:
The wingspan ranges from 45 mm to 62 mm (median 55 mm). It is relatively stout-bodied, with forewings relatively narrow-elongate. The wings are white, "peppered" with black, and with more-or-less distinct cross lines, also black. These transverse wing lines and "peppered" maculation
804:
By contrast, different subspecies of the same species can theoretically interbreed with one another and will produce fully fertile and healthy offspring, but in practice do not, as they live in different regions or reproduce in different seasons. Full-fledged species are either unable to produce
737:
over the seven years 2001–2007 Majerus noted the natural resting positions of peppered moths, and 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.
400:
The caterpillars of the peppered moth not only mimic the form but also the colour of a twig. Recent research indicates that the caterpillars can sense the twig's colour with their skin and match their body colour to the background to protect themselves from predators.
654:
A mating pair or a lone individual will spend the day hiding from predators, particularly birds. In the case of the former, the male stays with the female to ensure paternity. Evidence for resting positions is given by data collected by the peppered moth researcher
980:
Since then, with improved environmental standards, light-coloured peppered moths have again become common, and the dramatic change in the peppered moth's population has remained a subject of much interest and study. This has led to the coining of the term
1624:
Arjen E. van’t Hof, Pascal
Campagne, Daniel J. Rigden, Carl J. Yung, Jessica Lingley, Michael A. Quail, Neil Hall, Alistair C. Darby & Ilik J. Saccheri (2016). "The industrial melanism mutation in British peppered moths is a transposable element".
985:" to refer to the genetic darkening of species in response to pollutants. As a result of the relatively simple and easy-to-understand circumstances of the adaptation, the peppered moth has become a common example used in explaining or demonstrating
659:, and it is given in the accompanying charts. These data were originally published in Howlett and Majerus (1987), and an updated version published in Majerus (1998), who concluded that the moths rest in the upper part of the trees. Majerus notes:
800:
was involved in the observed evolution of the peppered moth. This is not the case; individuals of each morph interbreed and produce fertile offspring with individuals of all other morphs; hence there is only one peppered moth species.
605:
It goes into the soil late in the season, where it pupates in order to spend the winter. The imagines emerge from the pupae between late May and August, the males slightly before the females (this is common and expected from
956:
The evolution of the peppered moth over the last two hundred years has been studied in detail. At the start of this period, the vast majority of peppered moths had light coloured wing patterns which effectively
649:
Figure 2b. Total number of observed moths = 23: Exposed trunk =1; unexposed trunk =1; trunk branch join = 3; branches thicker than 5 centimetres diameter = 10; branches and twigs less thick than 5 centimetres =
674:
on the tree trunks or walls adjacent to our traps, and none elsewhere". The reason now seems obvious. Few people spend their time looking for moths up in the trees. That is where peppered moths rest by
683:
Further support for these resting positions is given from experiments watching captive moths taking up resting positions in both males (Mikkola, 1979; 1984) and females (Liebert and
Brakefield, 1987).
780:
These forms are often accidentally elevated to subspecies status when they appear in literature. Not adding the "f." (forma) or morpha implies that the taxon is a subspecies instead of a form, as in
824:. The melanic allele is dominant to the non-melanic allele. This situation is, however, somewhat complicated by the presence of three other alleles that produce indistinguishable morphs of morpha
625:. The males guard the female from other males until she lays the eggs. The female lays about 2,000 pale-green ovoid eggs about 1 mm in length into crevices in bark with her
431:
416:
1670:
765:
of the peppered moth. These are controlled genetically. A particular colour morph can be indicated in a standard way by following the species name in the form "morpha
690:, (2000) have shown that peppered moths are cryptically camouflaged against their backgrounds when they rest in the boughs of trees. It is clear that in human visible
2195:
1794:
2280:
1751:
1902:
457:(Heilongjiang, Jilin, Inner Mongolia, Beijing, Hebei, Shanxi, Shandong, Henan, Shaanxi, Ningxia, Gansu, Qinghai, Xinjiang, Fujian, Sichuan, Yunnan, Tibet),
1063:
730:
common on tree trunks; though they are camouflaged in human wavelengths, in ultraviolet wavelengths, foliose lichens do not reflect ultraviolet light.
1743:
1721:
996:
morph was recorded by
Edleston in Manchester in 1848, and over the subsequent years it increased in frequency. Predation experiments, particularly by
1739:
1411:
2169:
1012:
The evolution of the industrial melanism mutation has been shown to be due to the insertion of a transposable element into the first intron of the
2208:
613:
The males fly every night of their lives in search of females, whereas the females only fly on the first night. Thereafter, the females release
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Figure 2a. Total number of observed moths = 59: Exposed trunk = 7; unexposed trunk = 7; trunk-branch joint = 23; branches = 22
589:, varying in colour between green and brown. On a historical note, it was one of the first animals to be identified as being
2213:
1111:(1999). "Fine tuning the peppered moth paradigm: Reviewed Work: 'Melanism: Evolution in Action' by Michael E. N. Majerus".
2267:
1931:
961:
them against the light-coloured trees and lichens upon which they rested. However, due to widespread pollution during the
2037:
2065:
1266:"A review of Biston Leach, 1815 (Lepidoptera, Geometridae, Ennominae) from China, with description of one new species"
1004:
morph. Subsequent experiments and observations have supported the initial evolutionary explanation of the phenomenon.
2148:
1861:
1821:
1434:
2285:
1698:
2161:
1924:
1177:"Caterpillars of the peppered moth perceive color through their skin to match their body color to the background"
645:
637:
2349:
84:
66:
1892:
1856:
1687:
1108:
921:
386:
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in
England, many of the lichens died out, and the trees which peppered moths rested on became blackened by
2344:
2334:
1985:
1418:
762:
747:
598:
2226:
1197:
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to make it appear flat (shading being the main visual cue that makes things appear solid), in a paper by
2070:
1814:
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560:
179:
2023:
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light that humans cannot see. Using an ultraviolet-sensitive video camera, Majerus et al. showed that
2104:
1800:
1747:
1636:
1328:
1175:
Eacock, A.; Rowland, H. R.; van’t Hof, A. E.; Yung, C.; Edmonds, N.; Saccheri, I. J. (August 2019).
1579:
982:
618:
390:
2259:
1938:
1877:
1496:
1396:
1128:
997:
263:
79:
582:(adults). During the day, the moths typically rest on trees, where they are preyed on by birds.
385:. It is mostly found in the northern hemisphere in places like Asia, Europe and North America.
2200:
2047:
904:) is similarly dominant to the non-melanic allele. There are also some intermediate morphs. In
2306:
2246:
2091:
1963:
1652:
1442:
1356:
1293:
986:
494:
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35:
2236:
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1968:
1945:
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1627:
1527:
1480:
1346:
1336:
1283:
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fertile and healthy offspring, or do not recognize each other's courtship signals, or both.
607:
31:
2293:
1882:
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1725:
1523:
1508:
1457:
719:
656:
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to attract males. Since the pheromone is carried by the wind, males tend to travel up the
528:
caterpillars on birch (left) and willow (right), demonstrating twig mimicry and effective
1640:
1390:
1332:
2156:
2143:
1610:
1595:
1351:
1312:
1288:
1252:
973:, to die off due to predation. At the same time, the dark-coloured, or melanic, moths,
727:
594:
537:
529:
926:
52:
2328:
1907:
1789:
Accusations of data fudging and scientific fraud in the case are found to be vacuous.
707:
553:
510:
276:
195:
156:
146:
1887:
1729:
1706:
1574:
1548:
1374:
Poulton, Edward B. (October 1887). "Notes in 1886 upon
Lepidopterous Larvae, etc".
490:
379:
2096:
1341:
828:. These are of intermediate dominance, but this is not complete (Majerus, 1998).
17:
2272:
2254:
2182:
2017:
1690:
has written several papers on melanism in the peppered moth which are listed on
1265:
722:
common on branches, both in ultraviolet and human-visible wavelengths. However,
711:
663:
571:
557:
474:
470:
136:
521:
1897:
1783:
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958:
797:
691:
666:
critics of the peppered moth have often pointed to a statement made by Clarke
626:
590:
498:
486:
482:
2135:
2008:
1278:
610:). They emerge late in the day and dry their wings before flying that night.
2117:
878:
796:. Either of these two circumstances might lead to the erroneous belief that
734:
718:
reflect ultraviolet light in a speckled fashion and are camouflaged against
614:
545:
116:
96:
1656:
1360:
1297:
621:, i.e., toward the source. During flight, they are subject to predation by
1542:"A modelling exercise for students using the peppered moth as its example"
2078:
2057:
2002:
1795:
An interactive game to simulate how evolution works with Biston betularia
758:
462:
2298:
1648:
2174:
2083:
1132:
586:
541:
502:
2187:
1000:, established that the agent of selection was birds who preyed on the
2032:
813:
699:
567:
506:
458:
126:
106:
1979:
1799:
Wing vein patterns; this information is found in iNaturalist.org at
1124:
2122:
1251:. The Palaearctic Geometridae, 4. 479 pp. Alfred Kernen, Stuttgart.
1596:"Dark moths increased in s. Britain after pollution control began"
905:
644:
636:
579:
564:
520:
478:
466:
454:
1767:: An Excellent Teaching Example of Darwinian Evolution in Action"
1806:
966:
575:
382:
2109:
1983:
1810:
1707:"Moonshine: Why the Peppered Moth remains an Icon of Evolution"
836:
In continental Europe, there are three morphs: the white morph
1691:
622:
777:
in detailing these variations is also a widespread practice.
1611:"In several areas dark moths were more common than expected"
977:, flourished because they could hide on the darkened trees.
908:, no melanic morphs have been recorded; they are all morpha
792:. Rarely, forms have been elevated to species status, as in
680:
foliate twigs. The above data would appear to support this.
1313:"A Reversible Color Polyphenism in American Peppered Moth (
552:, it has one generation per year), whilst in south-eastern
1742:. This is the transcript of Majerus' lecture given at the
1473:"Moth study backs classic 'test case' for Darwin's theory"
1198:"Photos of Peppered Moth (Biston betularia) · iNaturalist"
1311:
Noor MA, Parnell RS, Grant BS (2008). Humphries S (ed.).
989:
to laypeople and classroom students through simulations.
1247:
Prout, L. B. (1912–16). Geometridae. In A. Seitz (ed.)
1016:
gene, resulting in an increase in the abundance of the
670:. (1985): "... In 25 years we have only found two
1236:
1412:"The Peppered Moth: The Proof of Darwinian Evolution"
1020:
transcript, which is expressed in developing wings.
890:
In North
America, the melanic black morph is morpha
1992:
1955:
1916:
1870:
1844:
1740:
The
Peppered Moth: The Proof of Darwinian Evolution
1376:
1722:The Peppered Moth: Decline of a Darwinian Disciple
1709:. Department of Physics, Colorado School of Mines
808:European breeding experiments have shown that in
1699:"The rise and fall of the melanic Peppered Moth"
865:In Britain, the typical white morph is known as
556:it is bivoltine (two generations per year). The
1471:Steve Connor, Science Editor (25 August 2007).
969:, causing most of the light-coloured moths, or
1822:
1234:Watson, L., and Dallwitz, M.J. 2003 onwards.
563:consists of four stages: ova (eggs), several
8:
1746:meeting on 23 August 2007. The accompanying
1669:: CS1 maint: multiple names: authors list (
1763:"Industrial Melanism in the Peppered Moth,
1392:Concealing-Coloration in the Animal Kingdom
1980:
1829:
1815:
1807:
65:
51:
42:
30:This article is about the insect. For the
1782:
1744:European Society for Evolutionary Biology
1613:. Index to Creationist Claims: CB601.2.2.
1598:. Index to Creationist Claims: CB601.2.3.
1350:
1340:
1287:
1277:
1170:
1168:
940:
925:
1100:
1027:
900:, the melanic allele (producing morpha
710:. However, birds are capable of seeing
412:
324:Amphidasys betularia var. doubledayaria
1662:
1504:
1494:
1452:
1451:
1440:
769:". The use of "form" in the method of
1583:. Index to Creationist Claims: CB601.
1410:Michael E. N. Majerus (August 2007).
1152:National Center for Science Education
7:
2162:7982c8af-0b1c-4356-8caa-b99487f92946
1264:Jiang, N.; Xue, D.; Han, H. (2011).
726:are not as well camouflaged against
578:, which overwinter in the soil, and
1146:Gishlick, Alan (23 November 2006).
602:rather slower in the caterpillars.
1609:David Wilson (10 September 2003).
1594:David Wilson (10 September 2003).
25:
1771:Evolution: Education and Outreach
1249:The Macrolepidoptera of the World
952:, the black-bodied peppered moth.
937:, the white-bodied peppered moth.
1078:
1062:
1046:
1030:
430:
415:
83:
1852:Overview, ecology, and genetics
1801:Wiki - American Peppered Moths
1524:"The peppered moth: an update"
1435:powerpoint presentation as pdf
1223:The Moths of the British Isles
816:for melanism producing morpha
356:Biston cognataria sinitibetica
348:Biston (Eubyjodonta) huberaria
1:
1932:Melanism: Evolution in Action
846:), the dark melanistic morph
340:Biston cognataria alexandrina
273:Phalaena (Geometra) betularia
239:(Barnes & Benjamin, 1923)
1752:also available as a pdf file
1730:British Humanist Association
1724:. This is the transcript of
1342:10.1371/journal.pone.0003142
856:), and an intermediate form
378:) is a temperate species of
2360:Taxa named by Carl Linnaeus
1728:' lecture delivered to the
1701:presented by Laurence Cook.
1315:Biston betularia cognataria
1238:. Version: 29 December 2011
1071:Biston betularia nepalensis
898:Biston betularia cognataria
782:Biston betularia carbonaria
303:Phalaena (Geometra) ulmaria
287:Phalaena (Noctua) p-graecum
2376:
1522:Ken Miller (August 1999).
1389:Thayer, Gerald H. (1909).
946:Biston betularia betularia
931:Biston betularia betularia
919:
820:is controlled by a single
810:Biston betularia betularia
745:
585:The caterpillar is a twig
29:
1925:The Evolution of Melanism
1784:10.1007/s12052-008-0107-y
1575:"The peppered moth story"
1573:Mark Isaak (2 May 2005).
1148:"Icon 6 — Peppered Moths"
1008:Genetic basis of melanism
351:(tienschana Wehrli, 1941)
269:
262:
211:
206:
185:
178:
80:Scientific classification
78:
73:
64:
59:
50:
45:
1279:10.3897/zookeys.139.1308
733:During an experiment in
698:are camouflaged against
2340:Moths described in 1758
1871:Writers and researchers
922:Peppered moth evolution
877:, and the intermediate
871:, the melanic morph is
544:, the peppered moth is
387:Peppered moth evolution
2355:Moths of North America
1759:Majerus, Michael E. N.
1395:. Macmillan. pp.
953:
938:
748:Polymorphism (biology)
677:
651:
642:
619:concentration gradient
599:Edward Bagnall Poulton
533:
517:Ecology and life cycle
963:Industrial Revolution
944:
929:
753:Introduction on forms
746:Further information:
661:
648:
640:
524:
332:Eurbyjodonta concinna
2157:Fauna Europaea (new)
1748:Microsoft PowerPoint
1221:Richard South, 1909
316:Amphidasis huberaria
1761:(6 December 2008).
1649:10.1038/nature17951
1641:2016Natur.534..102H
1580:TalkOrigins Archive
1333:2008PLoSO...3.3142N
983:industrial melanism
391:population genetics
1939:Icons of Evolution
1878:Bernard Kettlewell
1507:has generic name (
998:Bernard Kettlewell
954:
939:
757:There are several
652:
643:
534:
306:(Borkhausen, 1794)
2322:
2321:
2307:Open Tree of Life
1986:Taxon identifiers
1977:
1976:
1964:The Peppered Moth
1635:(7605): 102–105.
1483:on 7 October 2008
1450:External link in
987:natural selection
794:Biston carbonaria
633:Resting behaviour
395:natural selection
389:is an example of
366:
365:
360:
352:
344:
336:
328:
320:
307:
299:
291:
283:
256:
248:
240:
232:
219:
215:B. b. alexandrina
171:B. betularia
36:The Peppered Moth
18:Biston betularius
16:(Redirected from
2367:
2315:
2314:
2302:
2301:
2289:
2288:
2276:
2275:
2273:NBNSYS0000006022
2263:
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2050:
2048:Biston-betularia
2041:
2040:
2038:biston-betularia
2028:
2027:
2026:
2024:Biston betularia
2013:
2012:
2011:
1994:Biston betularia
1981:
1969:Margaret Drabble
1946:Of Moths and Men
1831:
1824:
1817:
1808:
1791:
1786:
1765:Biston betularia
1750:presentation is
1718:
1716:
1714:
1697:Online lecture:
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1615:
1614:
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1600:
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1547:. Archived from
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1532:
1531:
1528:Brown University
1519:
1513:
1512:
1506:
1502:
1500:
1492:
1490:
1488:
1479:. Archived from
1468:
1462:
1461:
1455:
1454:
1448:
1446:
1438:
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1429:
1423:
1417:. Archived from
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1401:
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1308:
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1050:
1034:
842:(syn. morpha/f.
786:Biston betularia
771:Biston betularia
761:and non-melanic
720:crustose lichens
608:sexual selection
526:Biston betularia
451:Biston betularia
438:Biston betularia
434:
423:Biston betularia
419:
375:Biston betularia
358:
350:
342:
334:
327:(Millière, 1870)
326:
318:
311:Eubyja betularia
305:
297:
289:
275:
254:
252:B. b. nepalensis
246:
238:
230:
228:B. b. cognataria
217:
191:
189:Biston betularia
88:
87:
69:
55:
43:
32:Margaret Drabble
21:
2375:
2374:
2370:
2369:
2368:
2366:
2365:
2364:
2350:Moths of Europe
2325:
2324:
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2310:
2305:
2297:
2294:Observation.org
2292:
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2016:
2007:
2006:
2001:
1988:
1978:
1973:
1951:
1912:
1866:
1840:
1835:
1757:
1726:Michael Majerus
1712:
1710:
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1555:
1554:on 3 March 2016
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1484:
1477:The Independent
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1425:
1424:on 15 June 2011
1421:
1414:
1409:
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1368:
1317:) Caterpillars"
1310:
1309:
1305:
1263:
1262:
1258:
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1242:
1233:
1229:
1225:Frederick Warne
1220:
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1883:Mike Majerus
1857:Evolution of
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1711:. Retrieved
1705:Matt Young.
1665:cite journal
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1549:the original
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2255:NatureServe
2183:iNaturalist
2018:Wikispecies
1893:Bruce Grant
1688:Bruce Grant
1487:9 September
1428:21 February
1207:16 November
1202:iNaturalist
1157:19 December
1085:Subspecies
1069:Subspecies
1053:Subspecies
1037:Subspecies
959:camouflaged
784:instead of
712:ultraviolet
692:wavelengths
664:Creationist
591:camouflaged
475:South Korea
471:North Korea
405:Description
244:B. b. parva
207:Subspecies
147:Geometridae
137:Lepidoptera
34:novel, see
2329:Categories
2247:BistoBetul
1898:E. B. Ford
1734:Darwin Day
1095:References
1087:nepalensis
1002:carbonaria
994:carbonaria
992:The first
975:carbonaria
950:carbonaria
874:carbonaria
859:medionigra
849:carbonaria
832:Form names
826:medionigra
818:carbonaria
798:speciation
790:carbonaria
767:morph name
704:carbonaria
627:ovipositor
615:pheromones
561:life cycle
546:univoltine
499:Azerbaijan
487:Kyrgyzstan
483:Kazakhstan
295:marmoraria
117:Arthropoda
1713:3 January
1497:cite news
1113:Evolution
916:Evolution
902:swettaria
893:swettaria
884:insularia
881:is named
879:phenotype
844:betularia
735:Cambridge
686:Majerus,
672:betularia
165:Species:
103:Kingdom:
97:Eukaryota
2260:2.118961
2244:MaBENA:
2222:LepIndex
2201:10299269
2079:BugGuide
2055:BioLib:
2045:BAMONA:
2003:Wikidata
1657:27251284
1558:27 April
1443:cite web
1361:18769543
1321:PLOS ONE
1298:22259309
1182:2 August
1089:(female)
1057:(female)
775:formname
580:imagines
463:Mongolia
277:Linnaeus
264:Synonyms
196:Linnaeus
143:Family:
113:Phylum:
107:Animalia
93:Domain:
2175:1975897
2009:Q865651
1845:Biology
1637:Bibcode
1352:2518955
1329:Bibcode
1289:3260909
1270:ZooKeys
1133:2640740
1024:Gallery
948:morpha
933:morpha
759:melanic
700:lichens
568:instars
542:Ireland
503:Armenia
495:Georgia
153:Genus:
133:Order:
127:Insecta
123:Class:
74:Female
2312:968114
2227:231130
2214:117560
2188:143517
2149:445615
2136:308438
2123:BISTBE
2110:298821
2033:ARKive
1655:
1628:Nature
1378:: 294.
1359:
1349:
1296:
1286:
1131:
1073:(male)
1041:(male)
1018:cortex
1014:cortex
971:typica
935:typica
910:typica
868:typica
852:(syn.
839:typica
814:allele
812:, the
763:morphs
724:typica
716:typica
696:typica
688:et al.
565:larval
507:Europe
459:Russia
158:Biston
2286:82595
2234:LoB:
2196:IRMNG
2131:EUNIS
2084:19107
2071:15515
2058:52017
1917:Works
1736:2004.
1552:(PDF)
1545:(PDF)
1422:(PDF)
1415:(PDF)
1129:JSTOR
1055:parva
1039:parva
906:Japan
896:. In
822:locus
668:et al
593:with
587:mimic
576:pupae
479:Nepal
467:Japan
455:China
60:Male
2299:8396
2281:NCBI
2237:5604
2209:ITIS
2170:GBIF
2118:EPPO
2097:LX46
2066:BOLD
1715:2009
1671:link
1653:PMID
1560:2011
1509:help
1489:2007
1458:help
1430:2011
1357:PMID
1294:PMID
1209:2020
1184:2019
1159:2009
967:soot
708:bark
702:and
675:day.
623:bats
550:i.e.
540:and
509:and
440:♂ △
393:and
383:moth
368:The
281:1758
200:1758
2268:NBN
2105:EoL
2092:CoL
1967:by
1779:doi
1732:on
1645:doi
1633:534
1347:PMC
1337:doi
1284:PMC
1274:doi
1253:pdf
1121:doi
788:f.
773:f.
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Text is available under the Creative Commons Attribution-ShareAlike License. Additional terms may apply.
