588:"starting block" for the mimic to evolve against. The drive behind this type of evolution must be predation. Eventually, the mimetic population undergoes phenotypic fixation, usually at a point where the wing pattern and colors of the mimic have reached the closest superficial resemblance of its model. As these processes continued, the subspecies divergences began occurring as the mimetic species expanded their geographical range and began mimicking other species of butterfly. Determining what part of the butterfly genome controls wing color and pattern is also a major component that must be taken into account when trying to understand the evolution of mimicry. Each individual stripe or spot on a wing has a distinct identity that can be traced from species to species within a family.
596:
different color and pattern loci with complete suppression of recombination in experimental crosses in a 400,000 base section containing at least 18 genes. This single supergene locus controls differences in a complex phenotype like wing coloration that can involve modifications of wing pattern, shape, and body color. Mimetic patterns have high fitness correlated to locally abundant wing patterns and low fitness when the offspring have recombinant, non-mimetic phenotypes. This tight-linked area of wing pattern genes explains how mimetic phenotypes are not broken up during recombination during sexual reproduction.
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initial mimicry is achieved by a single mutation that has a large effect on the phenotype, which immediately gives the organism some protection, and is then refined by so-called modifier genes with lesser phenotypic effects. Consequently, if the genes for wing pattern and color were normal functioning genes, a single mating would produce several phenotypically different offspring, making the ability for mimicry to evolve very difficult.
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459:(Eurasia and North Africa). Three lineages of mimetic butterflies occur in North America and the evolution of mimicry may have played a large role in the diversification of this group. For butterflies to travel from the Palearctic region to the Nearctic region of the world, the migration must have occurred during a time period when
566:
Some literature suggests that the queen-viceroy may not be a good model-mimic pair for
Batesian mimicry. Experimental evidence has shown that avian predators express aversion to the queen butterfly after being exposed to viceroys. That the avian predators avoided the queen butterfly implies that the
542:
Early experiments suggested that the viceroys use
Batesian mimicry to defend themselves against predators. In these experiments, birds that had not been exposed to monarchs willingly ate viceroys, but those that had tasted the unpalatable monarch refused to touch the mimic. In addition, when given
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feed on members of the willow family as well, suggesting that an (ancestral host plant shift) expansion of a novel host plant across the Bering land bridge could have driven the colonization of the
Nearctic. Species level phylogenies based on the mitochondrial gene COI and the gene EFI-α of Nearctic
591:
A fascinating feature of pattern genetics is that the dramatic phenotypic changes are primarily due to small changes in the gene that determines the sizes and positions of pattern elements. This discovery is in accord with the principal theory for the evolution of mimicry. The theory proposes that
488:
tree confirms that the poplar is the closest existing relative of the
Nearctic taxa and is consistent with the theory that the host plant had a large effect on the evolution of North American admirals. Just like the wing-pattern of the Palearctic butterflies has little evidence of divergence, the
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species. When the monarch's breeding range overlaps with the viceroy, the viceroy will adopt the lighter shades of orange. Towards the south, the viceroy mostly displayed darker orange phenotypes in response to the larger population of queens. It is important to note that the differences between
529:
The viceroy's wing color ranges from tawny orange (resembling monarchs) in the north to dark mahogany (resembling queens) in the south. It has been argued that selective pressures from predators have given rise to "model switching" in the viceroy, with each subspecies being selected to copy the
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Based on phylogenic evidence, it is known that mimicry in the North
American admirals was a driver of speciation. An essential condition for the evolution of mimicry was the presence and abundance of unpalatable models. Mimetic evolution also involved direct selection with the model acting as a
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This unique puzzle led to proposal of a possible supergene. A supergene is a tight cluster of loci that facilitate the co-segregation of adaptive variation, providing integrated control of complex adaptive phenotypes. Different genomic rearrangements have tightened the genetic linkage between
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Viceroys display geographic color polymorphism, which occurs when the viceroy butterflies are observed having different color forms in different regions of their territory. Color polymorphism is hypothesized to be affected by interaction between the viceroy, monarch and queen's overlapping
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When palatability was measured by looking at avian responses to butterfly abdomen, it was found that the viceroy butterfly was significantly more unpalatable than the queen. The queen-viceroy relationship is too asymmetrical for them to be considered real co-mimics of each other. Instead,
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of the three other species, and presumed edible or only mildly unpalatable to predators, but this has since proven not to be true. In an experiment with both the monarch's and viceroy's wings removed, birds were discovered to think the viceroy was just as unpalatable as the monarchs.
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In light of this new interpretation, it has been speculated that different food plants in different geographical locations influence the palatability of the viceroy. Further investigation is needed to clarify the relationship between the viceroy and its purported models.
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Joron, M; Frezal, L; Jones, R; Chamberlain, N; Lee, S; Haag, C; Whibley, A; Becuwe, M; Baxter, S; Ferguson, L; Wilkinson, P; Salazar, C; Davidson, C; Clark, R; Quail, M; Beasley, H; Glithero, R; Lloyd, C; Sims, S; Jones, M; Rogers, J; Jiggins, C; Constant, R (2011).
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It had been long accepted that the viceroy practiced
Batesian mimicry, with the monarch and the queen serving as models. Batesian mimicry is a type of defensive behavior in which a palatable species closely resembles unpalatable or toxic species to avoid predation.
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When avian predators were exposed to butterfly abdomina without the wings, many avian predators rejected the viceroy after a single peck. Furthermore, they exhibited distress behavior similar to that displayed when eating other, known, unpalatable species.
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dates back approximately four million years. Whether the migration event was a single or multiple occurrence event has a significant effect on how we look at the evolution of mimicry. A history of multiple migrations would suggest that
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Color warnings in viceroy butterflies have been shaped by natural selection in an evolutionary relationship between prey and predator. The viceroy's main predators – like many other butterflies – consist mostly of birds.
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However, much evidence supports that a single event colonization is the best explanation. One theory of
Nearctic colonization states that the reason for the colonization was a larva host plant shift. The position of the
286:). The viceroy's wingspan is between 53 and 81 mm (2.1 and 3.2 in). It can be distinguished from the monarch by its smaller size and the postmedian black line that runs across the veins on the hindwing.
1282:
Ritland, David B. (June 1991). "Revising a
Classic Butterfly Mimicry Scenario: Demonstration of MĂĽllerian Mimicry between Florida Viceroys (Limenitis archippus floridensis) and queens (Danaus gilippus berenice)".
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Research has argued that the viceroy may be unpalatable to avian predators. If that is the case, then the viceroy butterfly displays MĂĽllerian mimicry, and both viceroy and monarch are co-mimics of each other.
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environments. In the northern areas of their region, where monarchs predominate, viceroys are lighter, while in southern
Florida, they are darker due to queens being more abundant than monarchs.
1141:
van Zandt Brower, Jane (March 1958). "Experimental
Studies of Mimicry in Some North American Butterflies: Part I. The Monarch, Danaus plexippus, and Viceroy, Limenitis archippus archippus".
1176:
Platt, Austin P.; Raymond P. Coppinger; Lincoln P. Brower (December 1971). "Demonstration of the Selective Advantage of Mimetic Limenitis Butterflies Presented to Caged Avian Predators".
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Mallet, James and Joron, Mathieu (1999). Evolution of Diversity in Warning Color and Mimicry: Polymorphism, Shifting Balance and Speciation. Pub Annu Rev Ecol Syst 200x 30:201-33
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these two morphs is only the color of the wings and the line drawn through the viceroy's lower wings; other features, such as body size and wing-pattern elements, are identical.
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mathematical models have suggested that the queen enjoys the benefits of mimicry at the viceroy's expense, and that the model-mimic dynamic between the two should be switched.
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mountains, southward into central Mexico. Its easternmost range extends along the Atlantic and Gulf coasts of North America from Nova Scotia into Texas. It has been possibly
504:
and Palearctic species also indicate a single colonization of the Nearctic species. The phylogenies produced indicate that a white-banded ancestor similar to the species
1226:
Ritland, David B. (August 1995). "Comparative unpalatability of mimetic viceroy butterflies (Limenitis archippus) from four south-eastern United States populations".
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1041:"Hybrid zone origins, species boundaries, and the evolution of wing-pattern diversity in a polytypic species complex of North American butterflies (Nymphalidae:
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The viceroy ranges through most of the contiguous United States as well as parts of Canada and Mexico. The westernmost portion of its range extends from the
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established itself in North America and resulted in several major lineages, three of which involved mimicry independently of each other. Given the present
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of the Nearctic species, it is likely that a single migration and subsequent expansion of the population was the foundation of the Nearctic butterflies.
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in their bodies, which makes them bitter, and upsets predators' stomachs. As further protection, the caterpillars, as well as their
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the choice between a mimic and non-mimic after being exposed to an unpalatable model, avian predators never ate the viceroy mimic.
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host plant use of these species also shows no sign of divergence. These species only feed on different species of honeysuckle (
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1932:
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Mullen, S P (2006). "Wing pattern evolution and the origins of mimicry among North American admiral butterflies (Nymphalide:
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occurred before the evolution of mimicry, meaning mimicry was the result of speciation instead of the driver of speciation.
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951:"Once a Batesian mimic, not always a Batesian mimiic: Mimoic reverts back to ancestral phetype wen the modpel is absent"
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queen does not serve as a model and the viceroy as a parasitic mimic; rather, they may be MĂĽllerian co-mimics.
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Ritland, David (1998). "Mimicry-Related Predation on Two Viceroy Butterfly (Limenitis Archippus) Phenotypes".
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Ritland, David (1998). "Mimicry-related Predation on Two Viceroy Butterfly (Limenitis archippus) Phenotypes".
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population mimic the coloration of the viceroy species. It was originally believed that the viceroy was a
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Nijhout, H F (1994). "Developmental perspectives on evolution of butterfly mimicry".
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Its wings feature an orange and black pattern, and over most of its range it is a
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Platt, A P (1983). "Evolution of North American admiral butterflies".
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ssp.) The exception is the poplar that feeds exclusively on aspen (
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Layberry, Ross A.; Hall, Peter W.; J. Donald, Lafontaine. (1998).
703:(11 April 1991). "The viceroy butterfly is not a batesian mimic".
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Catalogue of the Butterflies of the United States and Canada
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10.1674/0003-0031(1998)140[0001:mrpotv]2.0.co;2
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10.1674/0003-0031(1998)140[0001:mrpotv]2.0.co;2
818:. Toronto, Canada: Royal Ontario Museum. pp. 244–245.
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Monarch (left) and viceroy (right) butterflies exhibiting
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Description of Viceroy on Butterflies and Moths website
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791:"Viceroy Butterflies, Caterpillars, Chrysalis Photos"
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butterfly wing patterns are much more diverse in the
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839:Fullard, James H.; Nadia Napoleone (August 2001).
816:The ROM Field Guide to the Butterflies of Ontario
809:
807:
1331:Bulletin of the Entomological Society of America
1039:Mullen, S P; Dopman, E B; Harrison, R G (2008).
770:"Viceroy (Limenitis archippus) (Cramer, 1776)"
305:) and in Mexico they share the pattern of the
650:10.2305/IUCN.UK.2020-3.RLTS.T160397A831906.en
262:The viceroy was named the state butterfly of
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1504:Guppy, Crispin S. and Shepard, Jon H.
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530:color pattern of the locally dominant
1511:James, David G. and Nunnallee, David
1004:Molecular Phylogenetics and Evolution
908:Molecular Phylogenetics and Evolution
7:
360:feeds on trees in the willow family
297:, viceroys share the pattern of the
1903:IUCN Red List least concern species
636:IUCN Red List of Threatened Species
955:Proceedings of the Royal Society B
440:Evolution of admiral butterflies (
378:). The caterpillars sequester the
25:
949:Prudic, K L; Oliver, J C (2008).
313:). In all three areas, the local
49:Typical viceroy, co-mimic of the
1943:Lepidoptera of the United States
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1062:10.1111/j.1558-5646.2008.00366.x
427:
415:
403:
386:stage, resemble bird droppings.
372:), and poplars and cottonwoods (
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1506:Butterflies of British Columbia
1461:The American Midland Naturalist
1108:The American Midland Naturalist
755:. University of Toronto Press.
336:along the eastern edges of the
484:), a Palearctic species, in a
247:. It was long thought to be a
1:
1923:Butterflies described in 1776
558:The MĂĽllerian mimicry dispute
1918:Butterflies of North America
583:Evolution of viceroy mimicry
455:(North America) than in the
1571:Featured Creatures Web site
1527:The Butterflies of Cascadia
928:10.1016/j.ympev.2006.01.021
1959:
1908:NatureServe secure species
753:The Butterflies of Canada
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109:Scientific classification
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682:Kentucky State Butterfly
328:Distribution and habitat
1577:, Butterflies of Canada
516:Predators and avoidance
1017:10.1006/mpev.1994.1018
998:Brower, A V Z (1994).
967:10.1098/rspb.2007.1766
860:10.1006/anbe.2001.1753
672:, NatureServe Explorer
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243:) is a North American
1933:Insects of the Arctic
1928:Lepidoptera of Mexico
1525:Pyle, Robert Michael
549:
334:Northwest Territories
1343:10.1093/besa/29.3.10
793:. Gardens With Wings
643:: e.T160397A831906.
389:Adults are strictly
27:Species of butterfly
1938:Symbols of Kentucky
1649:Limenitis-archippus
1639:Limenitis_archippus
1625:Limenitis archippus
1595:Limenitis archippus
1497:Glassberg, Jeffrey
1428:10.1038/nature10341
1420:2011Natur.477..203J
1240:1995Oecol.103..327R
961:(1639): 1125–1132.
920:2006MolPE..39..747M
719:1991Natur.350..497R
699:Ritland, David B.;
629:Limenitis archippus
625:Walker, A. (2020).
241:Limenitis archippus
218:Limenitis archippus
59:Conservation status
18:Limenitis archippus
1248:10.1007/bf00328621
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295:American Southwest
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1875:Open Tree of Life
1587:Taxon identifiers
1561:viceroy butterfly
1518:Pelham, Jonathan
1414:(7363): 203–207.
825:978-0-88854-497-1
713:(6318): 497–498.
701:Lincoln P. Brower
552:MĂĽllerian mimicry
348:from California.
280:monarch butterfly
253:monarch butterfly
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74:Least Concern
64:
60:
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52:
47:
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38:
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872:the original
851:
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752:
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640:
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600:Polymorphism
594:
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494:
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486:phylogenetic
481:
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369:
364:, including
355:
352:Food sources
331:
314:
310:
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289:In Florida,
288:
283:
273:
261:
240:
236:
234:
217:
215:
199:
198:
186:
29:
1823:NatureServe
1745:iNaturalist
1619:Wikispecies
506:L. arthemis
422:Caterpillar
397:Life stages
358:caterpillar
270:Description
176:Nymphalidae
166:Lepidoptera
99:NatureServe
1897:Categories
1363:BioScience
881:2007-06-15
797:2018-01-23
609:References
470:speciation
457:Palearctic
362:Salicaceae
346:extirpated
293:, and the
146:Arthropoda
1913:Limenitis
1285:Evolution
1228:Oecologia
1178:Evolution
1143:Evolution
1049:Evolution
1043:Limenitis
904:Limenitis
767:cited at
510:monophyly
501:Limenitis
482:L. populi
465:Leminitis
449:Limenitis
442:Limenitis
384:chrysalis
278:with the
266:in 1990.
259:instead.
245:butterfly
194:Species:
187:Limenitis
132:Kingdom:
126:Eukaryota
1828:2.107284
1797:LepIndex
1763:10775497
1680:BugGuide
1656:BioLib:
1646:BAMONA:
1604:Wikidata
1481:86071506
1446:21841803
1313:28564042
1264:13436225
1256:28306826
1206:28564787
1128:86071506
1114:: 1–20.
1079:38769164
1071:18331459
985:18285285
936:16500119
868:53182157
735:28667520
491:Lonicera
461:Beringia
264:Kentucky
172:Family:
142:Phylum:
136:Animalia
122:Domain:
79:IUCN 3.1
35:Viceroy
1867:viceroy
1737:5132398
1610:Q310832
1575:Viceroy
1563:on the
1437:3717454
1416:Bibcode
1383:1312251
1305:2409699
1236:Bibcode
1198:2406950
1163:2405902
1026:8075834
976:2602694
916:Bibcode
775:4 April
715:Bibcode
656:21 June
391:diurnal
375:Populus
366:willows
307:soldier
291:Georgia
251:of the
237:viceroy
228:, 1776)
182:Genus:
162:Order:
156:Insecta
152:Class:
97: (
95:Secure
77: (
51:monarch
1880:581024
1802:160351
1789:160397
1776:777969
1724:LIMEAC
1711:159832
1659:523658
1529:(2002)
1522:(2008)
1515:(2011)
1508:(2001)
1501:(2001)
1479:
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1408:Nature
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706:Nature
532:danaid
316:Danaus
226:Cramer
1854:79907
1841:42270
1758:IRMNG
1750:58586
1698:6QB84
1672:79910
1477:S2CID
1467:: 1.
1379:JSTOR
1301:JSTOR
1260:S2CID
1194:JSTOR
1159:JSTOR
1124:S2CID
1075:S2CID
875:(PDF)
864:S2CID
844:(PDF)
731:S2CID
410:Adult
370:Salix
299:queen
1862:ODNR
1836:NCBI
1815:4523
1810:MONA
1784:IUCN
1771:ITIS
1732:GBIF
1719:EPPO
1667:BOLD
1569:IFAS
1442:PMID
1309:PMID
1252:PMID
1202:PMID
1067:PMID
1022:PMID
981:PMID
932:PMID
906:)".
820:ISBN
777:2015
757:ISBN
658:2021
641:2020
434:Pupa
356:The
340:and
235:The
1706:EoL
1693:CoL
1685:548
1634:ADW
1469:doi
1432:PMC
1424:doi
1412:477
1371:doi
1339:doi
1293:doi
1244:doi
1232:103
1186:doi
1151:doi
1116:doi
1112:140
1057:doi
1012:doi
971:PMC
963:doi
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