North American Ecology (US and Canada)
- Scott, J. A. 1986. The butterflies of North America. Stanford University Press.
The viceroy ranges from central Canada through the eastern United States, into the Cascade Mountains and northern Mexico.
Biogeographic Regions: nearctic (Native )
occurs (regularly, as a native taxon) in multiple nations
Regularity: Regularly occurring
Type of Residency: Year-round
Regularity: Regularly occurring
Type of Residency: Year-round
Global Range: (>2,500,000 square km (greater than 1,000,000 square miles)) Southerb Canada; US except for Pacific coast and higher Rocky Mountains.
In areas of the viceroy's range where monarchs are common, the viceroy tends to mimic the pattern of the monarch (Danaaus plexippus) with black striping and orange areas similar to a monarch. The viceroy can be distinguished from the monarch, however, by one row of white spots within the black fore and hind wing bands. In areas inhabited by the Queen (Danaus glippus), the white spotting of the viceroy becomes less noticeable, and the orange coloration is replaced by a deep mahogany brown.
Other Physical Features: ectothermic ; bilateral symmetry
Viceroys prefer open or slightly shrubby areas that are wet or near water. These include wet meadows, marshes, ponds and lakes, railroad tracks, and roadsides.
Terrestrial Biomes: forest ; mountains
Aquatic Biomes: lakes and ponds; rivers and streams
Comments: Eastward almost any habitat with willows or small aspens which are the main larval foodplants. Habitats include prairies and dry barrens with small willows as well as wetlands. Westward more riparian and only around seeps or watercourses in arid regions.
Non-Migrant: No. All populations of this species make significant seasonal migrations.
Locally Migrant: No. No populations of this species make local extended movements (generally less than 200 km) at particular times of the year (e.g., to breeding or wintering grounds, to hibernation sites).
Locally Migrant: No. No populations of this species make annual migrations of over 200 km.
Larvae feed on various types of willows and poplars. Viceroys produce three generations per year, and the food habits of each generation differs. The first brood consume carrion, decaying fungi, and animal dung. Later generations are more often observed at flowers of plants, such as joe-pye weed, aster, Canada thistle, shepherd's needle, and goldenrod. This difference is likely due to the colder, wetter conditions experienced by the first generation.
Comments: Larvae of all subspecies are virtually resitricted to willows (Salix) an dpoplars (including aspens) (Populus).
Flowering Plants Visited by Limenitis archippus in Illinois
(observations are from Robertson, Graenicher, Betz et al., Fothergill & Vaughn; this butterfly is the Viceroy)
Apiaceae: Cicuta maculata sn (Rb), Eryngium yuccifolium sn (Rb), Oxypolis rigidior sn (Rb); Asclepiadaceae: Asclepias incarnata [plup sn] (Btz), Asclepias syriaca [plab sn] (Rb); Asteraceae: Ageratina altissima sn (Gr), Aster novae-angliae sn (Gr), Bidens aristosa sn (Rb), Bidens cernua sn (Rb), Cirsium altissimum sn (Rb, Gr), Cirsium arvense sn (Gr), Conoclinium coelestinum sn (Rb), Echinacea pallida sn (Rb), Erigeron philadelphicus sn (FV), Eupatoriadelphus purpureus sn (Gr), Eupatorium perfoliatum sn (Gr), Euthamia graminifolia sn (Gr), Heracleum maximum sn (Rb), Liatris pycnostachya sn (Rb), Oligoneuron rigidum sn (Rb), Ratibida pinnata sn (Gr), Rudbeckia hirta sn (Rb), Rudbeckia subtomentosa sn (Rb), Rudbeckia triloba sn (Rb), Silphium perfoliatum sn (Rb); Caprifoliaceae: Symphoricarpos occidentalis sn (Gr); Cornaceae: Cornus obliqua sn (Rb); Lamiaceae: Monarda fistulosa sn (Rb), Pycnanthemum tenuifolium sn (Rb); Rubiaceae: Cephalanthus occidentalis sn (Rb)
Number of Occurrences
Note: For many non-migratory species, occurrences are roughly equivalent to populations.
Estimated Number of Occurrences: 81 to >300
10,000 to >1,000,000 individuals
Life History and Behavior
- Scott, J. A. 1986. The butterflies of North America. Stanford University Press.
Comments: In most of the range two to four broods, the first in late spring (usualy May or June) the others following at about two month intervals but with stragglers often flying essentially all summer. Third instar larvae overwinter in shelters on the foodplant.
Mating occurs in the afternoon, and the female is the egg carrier. She deposits one egg onto the tip of a leaf and chooses only leaves that have not been eaten by other insects. She deposits about three eggs per sapling.
Molecular Biology and Genetics
Barcode data: Limenitis archippus
Below is a sequence of the barcode region Cytochrome oxidase subunit 1 (COI or COX1) from a member of the species.
See the BOLD taxonomy browser for more complete information about this specimen and other sequences.
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Download FASTA File
Statistics of barcoding coverage: Limenitis archippus
Public Records: 4
Specimens with Barcodes: 13
Species With Barcodes: 1
The viceroy has a wide range and is not threatened.
US Federal List: no special status
CITES: no special status
State of Michigan List: no special status
National NatureServe Conservation Status
Rounded National Status Rank: N5 - Secure
Rounded National Status Rank: N5 - Secure
NatureServe Conservation Status
Rounded Global Status Rank: G5 - Secure
Reasons: Widespread and common; adapts to some disturbance.
Degree of Threat: D : Unthreatened throughout its range, communities may be threatened in minor portions of the range or degree of variation falls within natural variation
Global Protection: Many to very many (13 to >40) occurrences appropriately protected and managed
The Viceroy (Limenitis archippus) is a North American butterfly that 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 Northwest Territories along the eastern edges of the Cascade Range and Sierra Nevada mountains, southwards into central Mexico. Its easternmost range extends along the Atlantic and Gulf Coasts of North America from Nova Scotia into Texas.
Its wings feature an orange and black pattern, and over most of its range it is a Müllerian mimic with the Monarch butterfly (Danaus plexippus). The viceroy's wingspan is between 53 and 81 mm. It can be distinguished from the Monarch by its smaller size and the postmedian black line that runs across the veins on the hindwing.
In Florida, Georgia, and the Southwest, Viceroys share the pattern of the Queen (Danaus gilippus) and in Mexico they share the pattern of the Soldier (Danaus eresimus). In all three areas, the local Danaus population mimic the coloration of the Viceroy species. It was originally believed that the Viceroy was a Batesian mimic of the three other species, and presumed edible or only mildly unpalatable to predators, but this has since proven not to be true.
The caterpillar feeds on trees in the willow family Salicaceae, including willows (Salix), and poplars and cottonwoods (Populus). The caterpillars sequester the salicylic acid in their bodies, which makes them bitter, and upsets predators' stomachs. As further protection, the caterpillars, as well as their chrysalis stage, resemble bird droppings.
Evolution of admiral butterflies (Limenitis)
The world is divided into eight biogeographic areas called ecozones: Palearctic, Nearctic, Afrotropic, Neotropic, Australasia, Indo-Malaya, Oceania, and Antarctica. Palearctic includes most of Eurasia and North Africa while Nearctic includes most of North America. Limenitis butterfly wing patterns are much more diverse in the Nearctic than the Palearctic. 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 Beringia, the land bridge between Eurasia and North America, was still above water. Based on crude divergence rate calculations, the colonization of the Nearctic Leminitis 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 speciation occurred before the evolution of mimicry, meaning mimicry was the result of speciation instead of the driver of speciation.
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 larvae host plant shift. The position of the Poplar admiral (L. populi), a Palearctic species, in a phylogenetic 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 host plant use of these species also shows no sign of divergence. These species only feed on different species of honeysuckle (Lonicera ssp.) The exception is the Poplar that feeds exclusively on aspen (Populus tremulus of the willow family). All North American Limenitis 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 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 L. arthemis. established itself in North America and resulted in several major lineages, three of which involved mimicry independently of each other. Given the present monophyly of the Nearctic species, it is likely that a single migration and subsequent expansion of the population was the foundation of the Nearctic butterflies.
Predators and avoidance
The Viceroy’s main predators – like many other butterflies – consist mostly of birds.
Mimicry as defense
The Viceroy’s major defense against predators is purported to be mimicry. 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. As such, the Viceroy wing color ranges from tawny orange (resembling Monarchs) in the north to dark mahogany (resembling Queens) in the south.
Early experiments suggested that the Viceroy use Batesian mimicry to defend themselves against predators. Birds that had not been exposed to Monarchs willingly ate Viceroy, but those who had tasted the unpalatable Monarch refused to touch the mimic. In addition, when given the choice between a mimic and non-mimic after being exposed to an unpalatable model, avian predators never ate the Viceroy mimic.
It has been argued that selective pressures from predators have given rise to “model switching” in the Viceroy, with each subspecies choosing to copy the color pattern of the locally dominant Danaine subspecies. 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 difference between these two morphs is only the color of the wings; other features, such as body size and wing-pattern elements, are identical.
Recent research has argued that the Viceroy may be unpalatable to avian predators. If that is the case, then the Viceroy butterfly displays Müllerian mimicry, and both Viceroy and Monarch are co-mimics of each other.
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 Viceroy. That the avian predators avoided the Queen butterfly implies that the Queen does not serve as a model and the Viceroy as a parasitic mimic; rather, they may be Müllerian co-mimics.
When avian predators were exposed to butterfly abdomen 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.
Interestingly, 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, 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.
In light of this new interpretation, it has been speculated that different food plants in different geographical locations influence the palatability of Viceroy. Further investigation is needed to clarify the relationship between the Viceroy and its purported models.
Evolution of Viceroy mimicry
Based on phylogenic evidence we know 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 “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. 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 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 phenyotypically different offspring, making the ability for mimicry to evolve very difficult.
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 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. Color warnings in Viceroy butterfly have been shaped by natural selection in an evolutionary relationship between prey and predator.
Polymorphism in Viceroy Butterflies
There is color polymorphism in Viceroy butterfly. This polymorphism shows geographic variation. This polymorphism has the advantage of enabling the viceroy to evade predation in the different environments. Color polymorphism is where the Viceroy butterflies display different color forms. Color polymorphism has been used by Viceroy butterflies by having different patterns of color mimicry for different environments and predator relationships. The viceroy has a lighter color pattern in Georgia in the United States, and it changes to a darker color pattern in the southern part of Florida as they switched from one model to another. The Viceroy butterfly is a perfect mimic of its model the monarch (Danaus plexippus) that has chemical defenses. The viceroy butterflies in different parts of the US have switched from mimicking the lighter orange monarch to the darker mahogany-purple monarch (D. gilippus). A kind of “model-switching”. This switching is regionally tuned based on the abundance of the models on different geographical latitudes. Natural selection based on the relative abundance of the models in a particular region has resulted in the viceroy butterfly “model-switching” in different regions to escape prey. This was variation with latitude. The darker colored butterflies were more abundant in the south, and the lighter colored butterflies were more abundant in the north. The evolutionary relationship of prey and predator has resulted in color mimicry. Closely related species tend to differ in their color mimicry, and mimicry can serve as a barrier to species isolation. Mimicry is variable geographically within species, and between sister species. Mimicry systems tend to evolve. This can be seen in insects and specifically butterflies. The butterflies that were being mimicked were unpalatable. It is expensive for the palatable butterflies to produce distasteful secondary chemicals, and it is cheaper to mimic the unpalatable butterflies. The unpalatable species Zygaena spp and Heliconius spp feed on cyanogenic plants, but are capable of producing their own toxins too. The bad taste has also evolved to protect potential eaters from toxic substances. Some species also show sex-linked mimicry where female butterflies are mimetic while males are not, and this is thought to be due to sexual selection. Visual mimicry shows natural selection for many organisms. It also shows parallel evolution. Butterflies display many color patterns and show a lot of mimicry, but there has been debate as to the explanations for this.
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Names and Taxonomy
Comments: A widespread species with some geographical and much individual variation. Two subspecies are almost universally recognized: FLORIDENSIS (mainly Florida) and OBSOLETA (southwestern USA and adjacent Mexico). Subspecies LAHONTANI (Nevada) is more or less transitional to OBSOLETA. Subspecies IDAHO needs to be reevaluated. It is close to typical ARCHIPPUS but probably shows some tendancy towards LAHONTANI. Based on the illustrations in its description and examination of additional published illustrations and specimens of eastern viceroys, dorsally IDAHO seems to fall mostly within the range of variation of northern transcontinental viceroys but the ventral tan areas are paler than at least eastern populations (D. Schweitzer).
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