Limenitis arthemis

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White admiral
White Admiral.jpg
L. a. arthemis
Red Spotted Purple.JPG
L. a. astyanax
Status TNC G5.svg
Secure  (NatureServe) [1]
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Arthropoda
Class: Insecta
Order: Lepidoptera
Family: Nymphalidae
Genus: Limenitis
Species:
L. arthemis
Binomial name
Limenitis arthemis
(Drury, 1773)
Synonyms

Basilarchia arthemis
Papilio arthemis
Papilio lamina
Limenitis proserpina

Contents

Red Spotted Purple Ventral View.jpg

Limenitis arthemis, the red-spotted purple or white admiral, is a North American butterfly species in the cosmopolitan genus Limenitis . It has been studied for its evolution of mimicry, and for the several stable hybrid wing patterns within this nominal species; it is one of the most dramatic examples of hybridization between non-mimetic and mimetic populations.

L. arthemis can be split into two major groups, mainly based on one physical characteristic: the presence of a white band along the wings. Individuals of the northern group, called white admirals, have a conspicuous white band that traverses both the dorsal and ventral surfaces of the wing, while those of the southern group, called red-spotted purples, lack that trait as they have evolved to mimic the poisonous pipevine swallowtail ( Battus philenor ). Due to overlap in distribution among the two major groups, intermediates are numerous as hybridization occurs frequently.

Taxonomy and phylogenetics

Limenitis arthemis is a butterfly species in the tribe Limenitidini of the family Nymphalidae. The Limenitidini are a tribe of the better known "brush-footed butterflies", as they are known to perch on hindlegs, whereas the other two forelegs are positioned curled up. These two forelegs often have brush-like hairs, which is a key identifier of the Nymphalidae. The Limenitidini consist of 25 species grouped primarily by region. The Basilarchia species group, spread in North America, includes the American white admiral species, L. arthemis, as well as L. archippus, L. lorquini, and L. weidemeyerii. [2]

Description and identification

Limenitis arthemis is described to be beautiful and highly active. The butterfly species themselves can be divided into two major groups simply from one main characteristic, the white band on the upper wings. However, besides the look of the butterfly, L. arthemis are in constant motion. Their flights are short in duration and at low altitudes, flying only about 2 to 3 feet off the ground. When not in flight, L. arthemis are constantly walking over leaves and folding their wings. They enjoy the sun as many are found to be resting at the highest points on trees. During the short period they are at rest, L. arthemis keep their wings closed, body at a 45 degree angle upwards, and antennae straight forward. [3]

The two sexes of this species are identical except that the females are slightly larger than the males. [4] The upperside of L. a. arthemis is mostly blackish-blue with white postmedian bands across both wings. Some individuals have a row of red submarginal spots, while others have this area being blue. The underside of the wings is a blackish color with a broad white post-median band. The basal area of both wings contains many red spots. The submarginal area may contain a row of red spots and the marginal area having bluish spots. However, sometimes the submarginal and marginal areas are just a reddish-brown color. [4] [5] [6]

The upperside of L. a. astyanax is very much like L. a. arthemis except it lacks the broad white bands. The forewing submarginal area will sometimes have a row of red spots. The hindwings are either a bright iridescent blue or an iridescent bluish-green. The underside of the wings lacks the white band. The basal area has several red spots. It has a row of red submarginal spots and bluish marginal spots. [4] [5]

Distribution and habitat

Limenitis arthemis are vastly spread out throughout North America. L a. arthemis or the white admiral live on the far north side of the continent, ranging from New England and southern Great Lakes area all the way to various parts of Canada. L. a. astyanax are based further south from the New England and southern Great Lake boundary, and can be found as far south as Florida; [7] the hybridization region is the overlapped region of New England and southern Great Lakes. These butterflies spend their days in deciduous woodlands, along the edges of the forest in shady areas, including roadsides. [8]

Home range and territoriality

Male territoriality display

When males are searching for mates, they generally try to defend areas that have high female visitation rates, regardless of the amount of resources. Male L. arthemis are known to be very aggressive when it comes down to defending an area bountiful of female mates. Male residents perch under the sun until another male comes into the vicinity; these engagements generally last approximately 1–5 minutes. Once conflicts comes to an end, males periodically patrol their territory for other outsiders tempted to take over the territory. Males also have high fidelity to the territory they are defending. [9]

Food resources

Caterpillars

Caterpillars of the hybrid region generally feed on tree species in the plant family Salicaceae, including aspen, poplar, and willow trees. Those of the northern region generally feed extensively on yellow birch trees, including Betula aleghaniensis and Betula lenta . Southern caterpillars feed on black cherry ( Prunus serotina ) and other members of the Rosaceae family. [10]

Adults

Red Spotted Purple (L. a. astyanax) RSPTop.png
Red Spotted Purple (L. a. astyanax)

The adult diet generally includes rotting fruit and nectar from small white flowers.

Parental care

Oviposition

Females lay eggs on leaves of food plants (see Food Resources for details), specifically at the very tips of these leaves, to the point where the width of the egg and that of the leaf are the same. These plants are about two to three feet off the ground. Mother undergoes labor for several weeks, laying only a few eggs per day; this is implied as more and more worn-out females are found. [3]

Life history

Life cycle

Larva Limenitis larva.jpg
Larva

Limenitis arthemis have two broods lasting from April to October. Most of the first brood feed and grow until the caterpillar is half-grown. Then they form a hibernaculum and hibernate for the winter until the start of spring. However, some larvae are able to mature during the summer, so they emerge as the second brood early fall. The second brood also mate and lay eggs, but often these larvae are not yet mature enough to undergo hibernation. Ultimately, this could mean death for the larvae. [3]

Caterpillar Limenitis arthemis caterpillar.jpg
Caterpillar

Egg

Lasting about 7 days, the eggs have a grey-green color with kite-shaped cells surrounding a central circular structure.

Caterpillar

Lasting a couple of weeks after hatching, L. arthemis larvae have wood brown heads with dark brown and yellow bordered bodies. Mature larvae are deep brownish-olive color with faint white midsections on their dorsal sides. The legs and prolegs are red-brown. Third stage larvae undergo hibernation at start of winter.

Pupa

Pupa stage last approximately 10–14 days. Chrysalis vary color from a creamy white to silvery gray.

Adult

Red Spotted Purple (L. a. astyanax) RSPFront.png
Red Spotted Purple (L. a. astyanax)

Adult butterflies, if part of first brood, are usually flying by mid-June. Adults are diurnal, meaning they fly from the morning until soon after dusk. [11] Life as an adult lasts approximately 6-14 days.

Sensory biology

Vision

Vision is a dynamic and integral part of an organism. For the L.arthemis it is crucial in order to find food, avoid predators, and engage in social signaling i.e. mating behaviors. Butterfly vision is diverse and unique amongst individual species. This is due to their remarkable range of photoreceptor classes, each with distinct spectral sensitivities. [12] This diversity results not only from the different spectral properties, but also from the alternations in the number and spatial distribution of visual pigments. [13] Vision is a collaborative process between the components of the visual pigment: the opsin protein and the chromophore. The opsin gene encodes opsin proteins, which binds to the chromophore to form a visual pigment. These visual pigments are modulated by specific amino acids which determines the absorbance spectra that allows for animals to detect particular wavelengths of light. In L. arthemis, it has been shown in a study that the diversification of opsins have altered on a longitudinal cline along with natural selection due to environmental factors, unrelated to spectral tuning. [14]

Protective coloration

Batesian mimicry

Red Spotted Purple (L. a. astyanax) Limenitis arthemis arizonensis.jpg
Red Spotted Purple (L. a. astyanax)

This type of mimicry between species of the Limenitis arthemis is a prime example of convergent evolution, in which similar traits are developed among species in common niches or environment. Butterfly wing pattern is a trait often subjected to mimicry among species of different lineages because despite its diversity, the patterns are developed based upon a strict blueprint laid by preceding ground plans that were conserved throughout evolution. [15] Though the exact mechanism by which the wing patterning developed remains unclear, there is evidence of genes associated with the eyespot patterning development. [16] Yet, two conflicting ideas have been proposed to explain the mechanism involved: conserved homology or recent variability. [17]

The red-spotted purple butterfly, also known as the subspecies L. arthemis astyanax, is a well known Batesian mimic. [2] The palatable red-spotted purple mimics the unpalatable pipevine swallowtail ( Battus philenor ) with its overall dark wings with bright iridescent blue along the hind wings. [2]

The two subspecies differ in the presence of a bright white band on both upper wings in the L. arthemis arthemis (non-mimic form).

Pipevine swallowtail Battus philenor - dorsal.jpg
Pipevine swallowtail

A study hypothesized that the higher density of either model or mimic would cause a decrease in the effect of the mimicry in evading predators; however, this was not the case, since they showed that even with the lowest density of the model, the effectiveness of evading predators by the mimics did not decline. [18]

A further study indicates that this phenomenon may be regulated by positionally orthologous nucleotide variants in the genome of these butterflies. Specifically, the gene WntA was found to be responsible for mimicry between L. arthemis and the pipevine swallowtail (Battus philenor), species that diverged more than 65 million years ago. [19]

Another study ruled out a major hypothesis stating that the reappearance of the non-mimic form was due to the evolutionary loss of mimicry and a reversion to the ancestral phenotype. [2] It showed that this hypothesis is most likely incorrect. These reasons included such significant levels of both contemporary and historical gene flow between the two phenotypic species that individual gene trees would not be accurate. [2] In addition, past work on this hypothesis utilized mitochondrial DNA, which is an unreliable guide to phenotype. [2]

Unlike previous studies which suggested that melanin pathway genes were responsible for variation in patterning and pigment expression, [20] the WntA pathway is a crucial part of the early development of embryos and therefore highly conserved between species; thus making it the most likely candidate responsible for the evolution of mimicry in L. arthemis. [17]

Genetics

Subspecies

L. arthemis Distribution Map:
Red: L. a. arthemis
Orange: L. arthemis astyanax
Yellow: Hybridization region between L. a. arthemis and L. a. astyanax
Green: L. a. arizonensis Limenitis arthemis range map.PNG
L. arthemis Distribution Map:
Red: L. a. arthemis
Orange: L. arthemis astyanax
Yellow: Hybridization region between L. a. arthemis and L. a. astyanax
Green: L. a. arizonensis

Hybridization

Limenitis arthemis is one of the most dramatic examples of hybridization within a nominal species. The two major subspecies that dominate the hybridization are L. a. arthemis (white admiral) and L. a. astyanax (red-spotted purple). The white admiral is characterized by a bright, white band on its wings, while the red-spotted purple lacks the white band, but has cooler blue-green shades on its wings. Hybridization occurs in the area across New England, southern Ontario, and the Great Lakes, also known as the "suture region". It is at these regions that the subspecies were formed. [10] [18]

According to one study, these hybrid zones were of secondary origin, meaning that both the white admiral and the red-spotted purple are already genetically distinct and the two diverged lineages reconnect at this hybrid area. [10] They were able to support their claim by examining various of mitochondria DNA of the population to determine the similarities and differences of the origins of the hybrid zones and the evolution of mimicry. Another study suggests that hybridization is highly frequent in this species because similar species mate together regardless of being the same butterfly or not. For instance, the L. arthemis was found to be mating with a L. archippus butterfly in New England. [21]

Mating

Mate searching behavior

Mississauga Ontario Canada L Arthemis Mississauga Ontario Canada - Top view.jpg
Mississauga Ontario Canada

When males are searching for female mates, they tend to seek territories with plentiful visits from females. Mate-locating behaviors for males tended to start in the afternoon, between 11 am and 4 pm. Most males perched as a sign of defense of the territory. Males tend not to favor perching on host trees, but rather on various kinds of foliage like maple, elm trees, or raspberry bushes, generally about 1–2 meters off the ground. This particular mating behavior seems ironic because males should want to perch on host trees as females tend to seek mates in areas where resources like food or host trees are bountiful. Therefore, many times, males are defending areas that do not attract the most females. [9]

Male-male behavior

Males tend to display aggressive behavior with each other during territorial fights. Fights have been classified into two types: 1) jostling, which are contests over eclosed or eclosing females and 2) aerial maneuvers, which are fought over ownership of mating territories in a specific breeding ground. [22] These fights are significant due to the ability of female butterflies to "double-mate" which introduce the idea of sperm precedence and competition to reproduce between males. A study showed that within female butterflies that "double-mate" a small percent of the progeny resulted from the mixing of sperm, which illustrate the individual fitness of the male. Therefore, there is an incentive for males to claim their mate prior to other males, which can also affect male mate-seeking strategies as well. [23] Generally, the fights engaged by L. arthemis are aerial maneuvers which are initiated once a nonresident male files into his peripheral vision. After which a series of flight patterns such as circling flights, vertical and/or horizontal chases, and hovering flights ensue. It is established that the "competition" component of the fight is the circling flight, often followed by the horizontal chase in which the 'loser' is chased from the disputed territory. Territorial fights are not limited to mating, but it is a method to optimize their mate-seeking behavior. [22]

Female-male behavior

Once a female lands on a leaf, the male lands behind her quite shortly after. If the female does not want to copulate, she closes her dorsal wings.

Ecology

Preferred host plants: birches, including Betula lenta ; Salicaceae, including Salix bebbiana and Populus tremuloides , and Prunus virginiana (Rosaceae).

Also but not as often: Crataegus , Amelanchier , Malus pumila , Prunus pensylvanica and Prunus serotina (Rosaceae), Populus deltoides , P. grandidentata and P. balsamifera (Salicaceae), Alnus rugosa , Betula alleghaniensis and Carpinus caroliniana (Betulaceae), Ulmus americana (Ulmaceae), Tilia americana (Malvaceae) and Fagus grandifolia (Fagaceae).

Caterpillar feeding on a willow tree Limenitis arthemis caterpillar feeding on a willow tree.jpg
Caterpillar feeding on a willow tree

Etymology

Limenitis (Neo-Latin "of harbours", from Ancient Greek Λιμενιτις (from λιμήν, a harbour, haven) – an epithet of Artemis, goddess of the hunt and the wild) – arthemis, from Artemis. [24]

The white admiral is the official state butterfly of New York. Limenitis arthemis arthemis is also considered by some to be the unofficial insect emblem of Quebec.

See also

Related Research Articles

<span class="mw-page-title-main">Mimicry</span> Imitation of another species for selective advantage

In evolutionary biology, mimicry is an evolved resemblance between an organism and another object, often an organism of another species. Mimicry may evolve between different species, or between individuals of the same species. Often, mimicry functions to protect a species from predators, making it an anti-predator adaptation. Mimicry evolves if a receiver perceives the similarity between a mimic and a model and as a result changes its behaviour in a way that provides a selective advantage to the mimic. The resemblances that evolve in mimicry can be visual, acoustic, chemical, tactile, or electric, or combinations of these sensory modalities. Mimicry may be to the advantage of both organisms that share a resemblance, in which case it is a form of mutualism; or mimicry can be to the detriment of one, making it parasitic or competitive. The evolutionary convergence between groups is driven by the selective action of a signal-receiver or dupe. Birds, for example, use sight to identify palatable insects and butterflies, whilst avoiding the noxious ones. Over time, palatable insects may evolve to resemble noxious ones, making them mimics and the noxious ones models. In the case of mutualism, sometimes both groups are referred to as "co-mimics". It is often thought that models must be more abundant than mimics, but this is not so. Mimicry may involve numerous species; many harmless species such as hoverflies are Batesian mimics of strongly defended species such as wasps, while many such well-defended species form Müllerian mimicry rings, all resembling each other. Mimicry between prey species and their predators often involves three or more species.

<span class="mw-page-title-main">Nymphalidae</span> Largest butterfly family

The Nymphalidae are the largest family of butterflies, with more than 6,000 species distributed throughout most of the world. Belonging to the superfamily Papilionoidea, they are usually medium-sized to large butterflies. Most species have a reduced pair of forelegs and many hold their colourful wings flat when resting. They are also called brush-footed butterflies or four-footed butterflies, because they are known to stand on only four legs while the other two are curled up; in some species, these forelegs have a brush-like set of hairs, which gives this family its other common name. Many species are brightly coloured and include popular species such as the emperors, monarch butterfly, admirals, tortoiseshells, and fritillaries. However, the under wings are, in contrast, often dull and in some species look remarkably like dead leaves, or are much paler, producing a cryptic effect that helps the butterflies blend into their surroundings.

<span class="mw-page-title-main">Batesian mimicry</span> Bluffing imitation of a strongly defended species

Batesian mimicry is a form of mimicry where a harmless species has evolved to imitate the warning signals of a harmful species directed at a predator of them both. It is named after the English naturalist Henry Walter Bates, after his work on butterflies in the rainforests of Brazil.

<span class="mw-page-title-main">Viceroy (butterfly)</span> Species of butterfly

The viceroy is a North American butterfly. It was long thought to be a Batesian mimic of the monarch butterfly, but since the viceroy is also distasteful to predators, it is now considered a Müllerian mimic instead.

<span class="mw-page-title-main">Müllerian mimicry</span> Mutually beneficial mimicry of strongly defended species

Müllerian mimicry is a natural phenomenon in which two or more well-defended species, often foul-tasting and sharing common predators, have come to mimic each other's honest warning signals, to their mutual benefit. The benefit to Müllerian mimics is that predators only need one unpleasant encounter with one member of a set of Müllerian mimics, and thereafter avoid all similar coloration, whether or not it belongs to the same species as the initial encounter. It is named after the German naturalist Fritz Müller, who first proposed the concept in 1878, supporting his theory with the first mathematical model of frequency-dependent selection, one of the first such models anywhere in biology.

<i>Danaus chrysippus</i> Species of butterfly

Danaus chrysippus, also known as the plain tiger, African queen, or African monarch, is a medium-sized butterfly widespread in Asia, Australia and Africa. It belongs to the Danainae subfamily of the brush-footed butterfly family Nymphalidae. Danainae primarily consume plants in the genus Asclepias, more commonly called milkweed. Milkweed contains toxic compounds, cardenolides, which are often consumed and stored by many butterflies. Because of their emetic properties, the plain tiger is unpalatable to most predators. As a result, its coloration is widely mimicked by other species of butterflies. The plain tiger inhabits a wide variety of habitats, although it is less likely to thrive in jungle-like conditions and is most often found in drier, wide-open areas.

<span class="mw-page-title-main">Queen (butterfly)</span> Species of butterfly

The queen butterfly is a North and South American butterfly in the family Nymphalidae with a wingspan of 80–85 mm. It is orange or brown with black wing borders and small white forewing spots on its dorsal wing surface, and reddish ventral wing surface fairly similar to the dorsal surface. The ventral hindwings have black veins and small white spots in a black border. The male has a black androconial scent patch on its dorsal hindwings. It can be found in meadows, fields, marshes, deserts, and at the edges of forests.

<i>Heliconius cydno</i> Species of butterfly

Heliconius cydno, the cydno longwing, is a nymphalid butterfly that ranges from Mexico to northern South America. It is typically found in the forest understory and deposits its eggs on a variety of plants of the genus Passiflora. It is a member of the Heliconiinae subfamily of Central and South America, and it is the only heliconiine that can be considered oligophagous. H. cydno is also characterized by hybridization and Müllerian mimicry. Wing coloration plays a key role in mate choice and has further implications in regards to sympatric speciation. Macrolide scent gland extracts and wing-clicking behavior further characterize this species.

<i>Battus philenor</i> Species of butterfly

Battus philenor, the pipevine swallowtail or blue swallowtail, is a swallowtail butterfly found in North America and Central America. This butterfly is black with iridescent-blue hindwings. They are found in many different habitats, but are most commonly found in forests. Caterpillars are often black or red, and feed on compatible plants of the genus Aristolochia. They are known for sequestering acids from the plants they feed on in order to defend themselves from predators by being poisonous when consumed. The adults feed on the nectar of a variety of flowers. Some species of Aristolochia are toxic to the larvae, typically tropical varieties. While enthusiasts have led citizen efforts to conserve pipevine swallowtails in their neighborhoods on the West coast, the butterfly has not been the subject of a formal program in conservation or protected in legislation. The butterfly is however of "Special Concern" in Michigan, which is on the Northern limit of its range.

<i>Heliconius</i> Genus of brush-footed butterflies

Heliconius comprises a colorful and widespread genus of brush-footed butterflies commonly known as the longwings or heliconians. This genus is distributed throughout the tropical and subtropical regions of the New World, from South America as far north as the southern United States. The larvae of these butterflies eat passion flower vines (Passifloraceae). Adults exhibit bright wing color patterns which signal their distastefulness to potential predators.

<i>Heliconius melpomene</i> Species of butterfly

Heliconius melpomene, the postman butterfly, common postman or simply postman, is a brightly colored butterfly found throughout Central and South America. It was first described by Carl Linnaeus in his 1758 10th edition of Systema Naturae. Its coloration coevolved with a sister species H. erato as a warning to predators of its inedibility; this is an example of Müllerian mimicry. H. melpomene was one of the first butterfly species observed to forage for pollen, a behavior that is common in other groups but rare in butterflies. Because of the recent rapid evolutionary radiation of the genus Heliconius and overlapping of its habitat with other related species, H. melpomene has been the subject of extensive study on speciation and hybridization. These hybrids tend to have low fitness as they look different from the original species and no longer exhibit Müllerian mimicry.

<span class="mw-page-title-main">Eyespot (mimicry)</span> Eye-like marking used for mimicry or distraction

An eyespot is an eye-like marking. They are found in butterflies, reptiles, cats, birds and fish.

<i>Callosamia promethea</i> Species of moth

Callosamia promethea, commonly known as the promethea silkmoth, is a member of the family Saturniidae, which contains approximately 1,300 species. It is also known as the spicebush silkmoth, which refers to is one of the promethea silkmoth's common host plants, spicebush. C. promethea is classified as a silk moth, which stems from its ability to produce silk, which it does in the formation of its cocoon. C. promethea lives in forests in the eastern U.S. and does not damage the trees on which it lives. The species was first described by Dru Drury in 1773.

<i>Papilio dardanus</i> Species of butterfly

Papilio dardanus, the African swallowtail, mocker swallowtail or flying handkerchief, is a species of butterfly in the family Papilionidae. The species is broadly distributed throughout Sub-Saharan Africa. The British entomologist E. B. Poulton described it as "the most interesting butterfly in the world".

Many types of polymorphism can be seen in the insect order Lepidoptera. Polymorphism is the appearance of forms or "morphs" differing in color and number of attributes within a single species. In Lepidoptera, polymorphism can be seen not only between individuals in a population but also between the sexes as sexual dimorphism, between geographically separated populations in geographical polymorphism and also between generations flying at different seasons of the year. It also includes the phenomenon of mimicry when mimetic morphs fly alongside non-mimetic morphs in a population of a particular species. Polymorphism occurs both at a specific level with heritable variation in the overall morphological design of individuals as well as in certain specific morphological or physiological traits within a species.

<i>Danaus dorippus</i> Species of butterfly

Danaus dorippus, the dorippus tiger, is a butterfly belonging to the danaine group of the brush-footed butterflies family.

Deception in animals is the transmission of misinformation by one animal to another, of the same or different species, in a way that propagates beliefs that are not true.

In evolutionary biology, mimicry in vertebrates is mimicry by a vertebrate of some model, deceiving some other animal, the dupe. Mimicry differs from camouflage as it is meant to be seen, while animals use camouflage to remain hidden. Visual, olfactory, auditory, biochemical, and behavioral modalities of mimicry have been documented in vertebrates.

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