Common Buckeye (Junonia Coenia) Anterior View
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''Junonia coenia'', known as the common buckeye or buckeye, is a butterfly in the family Nymphalidae. It is found in the United States east of the Rocky Mountains and in Mexico. Its habitat is open areas with low vegetation and some bare ground. Its original ancestry has been traced to Africa, which then experiences divergence in Asia. The species ''
Junonia grisea ''Junonia grisea'', the gray buckeye or grey buckeye, is a species in the butterfly family Nymphalidae. It is found in North America, west of the Rocky Mountains. Like the common buckeye, the gray buckeye is a brown butterfly with eyespots on it ...
'', the gray buckeye, is found west of the Rocky Mountains and was formerly a subspecies of ''Junonia coenia''. Caterpillars of these butterflies appear to prefer plants that produce iridoid
glycoside In chemistry, a glycoside is a molecule in which a sugar is bound to another functional group via a glycosidic bond. Glycosides play numerous important roles in living organisms. Many plants store chemicals in the form of inactive glycosides. ...
s, which are bitter compounds that release a
hormone A hormone (from the Greek participle , "setting in motion") is a class of signaling molecules in multicellular organisms that are sent to distant organs by complex biological processes to regulate physiology and behavior. Hormones are required ...
called
gastrin Gastrin is a peptide hormone that stimulates secretion of gastric acid (HCl) by the parietal cells of the stomach and aids in gastric motility. It is released by G cells in the pyloric antrum of the stomach, duodenum, and the pancreas. Gastrin ...
that activates the digestive system (i.e. hunger); therefore, iridoid glycoside producing plants stimulate and attract their appetites particularly when found in plants like ''
Plantago lanceolata ''Plantago lanceolata'' is a species of flowering plant in the plantain family Plantaginaceae. It is known by the common names ribwort plantain, narrowleaf plantain, English plantain, ribleaf, lamb's tongue, and buckhorn. It is a common weed on c ...
''. In fact, the presence of these metabolites may trigger
oviposition The ovipositor is a tube-like organ used by some animals, especially insects, for the laying of eggs. In insects, an ovipositor consists of a maximum of three pairs of appendages. The details and morphology of the ovipositor vary, but typical ...
behaviors in female butterflies so that descendant larval bodies may better incorporate iridoid glycosides. Iridoid glycolyside metabolites appear to have a growth-stimulating effect on caterpillars but a growth-reducing effect on predators. Predators like ants, wasps, birds, and small animals prefer to feed on iridoid glycoside poor caterpillars rather than iridoid glycoside rich larvae, potentially due to these effects. Therefore, immunity of ''J. coenia'' larvae to predators like ants appears to be strongly related to the concentration of iridoid glycosides sequestered in their bodies. However, too much iridoid glycosides in the diet can negatively affect the immune response of these larvae and lead to increased susceptibility to parasitism. Adult butterflies feed on flowers with certain pollinator cues: yellow flowers that are 'pre-change', or flowers whose color has not been changed due to insect visitation or other factors. Common buckeye caterpillars feed in isolation rather than relying upon grouping behaviors. Vulnerability to the ''Junonia coenia'' densovirus is another concern for survivorship of common buckeye larvae.


Description


Adult butterfly

Common buckeye butterflies are colored mostly brown with some orange, black, white, blue, and magenta. The forewing features two proximal orange bars and a postmedian white band, which surrounds a prominent black eyespot and borders a smaller, more distal eyespot; both eyespots have a bluish center and each border a distal orange mark. The hindwing is mostly brown with an orange band towards the edge and a brown and white margin. It also has two eyespots, one larger and one smaller, each with a black and white outline, and a center consisting of black, blue, magenta, and orange. They also exhibit blue-green overtones on their front and sometimes hind wings. Their wingspan is typically between 2 and 2.5 inches (5 - 6.5 cm).


Caterpillar

These caterpillars have a beautiful and complex color pattern. Their backs are mostly black with light-colored markings (white, gray, beige, or brownish, varying among individuals—see picture below), their sides have white markings and red-orange spots, and they have a brown underside. They have a mostly red-orange head with black markings on the face. They have spines running lengthwise along their back and sides, as many as 7 per segment. There is also a pair of small spines on the top of their head. These spines branch out (have smaller spines coming out of them) and appear a bit shiny-blue at the base. They also have 2 pairs of tiny white spines on each segment, near the bottom. The color patterns and spine forms are less pronounced in earlier instars. There are several other species that resemble these caterpillars in appearance, such as the painted lady, the
Glanville fritillary The Glanville fritillary (''Melitaea cinxia'') is a butterfly of the family Nymphalidae. It is named for the naturalist who discovered it and the checkerboard pattern on its wings. These butterflies live in almost all of Europe, especially Finla ...
, and the red admiral, which can be easily confused by an untrained observer. They can reach 4 cm (1 5/8 inches) in length. File:Common Buckeye variation, Megan McCarty41.JPG, Dorsal variation File:Common Buckeye variation, Megan McCarty75.JPG, Ventral variation File:Common Buckeye larva variation, Megan McCarty42.JPG, Larva variation File:Common Buckeye chrysalis, Megan McCarty43.JPG, Chrysalis File:Hubner1821SammlExotSchmett2Plate32.jpg, Plate accompanying Jacob Hübner's 1822 description


Pupa (chrysalis)

Like the more familiar
monarch A monarch is a head of stateWebster's II New College DictionarMonarch Houghton Mifflin. Boston. 2001. p. 707. Life tenure, for life or until abdication, and therefore the head of state of a monarchy. A monarch may exercise the highest authority ...
s, these caterpillars pupate by spinning a silk pad on a downward facing horizontal surface, attaching themselves with their rear end, and hanging in the form of the letter J for a day or so before shedding their skin to reveal the chrysalis. The chrysalis is mostly brown with beige markings, and has a prickly texture. It has a beige band near the top, another one at the "waist" (border between thorax and abdomen), and 4 pairs of beige dots running down its dorsal side. It is 18 mm long and 8 mm wide. When touched or handled excessively, it has a habit of wiggling its abdomen.


Phylogeny

''Junio'' probably arose from one African colonizing ancestor. In Asia, this diverged into ''J. atlites'', ''J. iphita'', and ''J. hedonia'' that resembled the African ancestor. These species then diverged into ''J. almanac'', ''J. lemonias'', and ''J. villida'' that began to differ morphologically from African ancestors. The Taxon Pulse Hypothesis is invoked to explain the vicariance as well as dispersal exhibited by this species where there is dispersal as the species expands until they find appropriate habitats. This then allows for
speciation Speciation is the evolutionary process by which populations evolve to become distinct species. The biologist Orator F. Cook coined the term in 1906 for cladogenesis, the splitting of lineages, as opposed to anagenesis, phyletic evolution within ...
to occur in several isolated habitats. In ''Junonia'' the species may have expanded from Africa into Asia and then from Asia throughout “Australasia, Africa, and South America.” In the 1930s established populations were recognized in Cuba and the Florida Keys. In the 1940s populations migrated inland from the coast of Florida and further into the southern United States.


Food resources


Caterpillars

Caterpillars feed on a variety of plants including the narrowleaf plantain (''
Plantago lanceolata ''Plantago lanceolata'' is a species of flowering plant in the plantain family Plantaginaceae. It is known by the common names ribwort plantain, narrowleaf plantain, English plantain, ribleaf, lamb's tongue, and buckhorn. It is a common weed on c ...
''), the common greater plantain ('' Plantago major''),
blue toadflax ''Nuttallanthus canadensis'', the blue toadflax, Canada toadflax, or old-field toadflax, is a species of ''Nuttallanthus'' in the family Plantaginaceae, native to eastern North America from Ontario east to Nova Scotia and south to Texas and Flori ...
,
false foxglove False foxglove may refer to: * ''Agalinis'', a genus of plants in the family Orobanchaceae * ''Aureolaria ''Aureolaria'', with the common name false foxgloves, is a genus of 8 species, native to North America. ''Aureolaria'' plants are hemiparas ...
, Mexican petunia (
Ruellia ''Ruellia'' is a genus of flowering plants commonly known as ruellias or wild petunias. They are not closely related to petunias (''Petunia'') although both genera belong to the same euasterid clade. The genus was named in honor of Jean Ruelle ( ...
species), the
firecracker plant ''Russelia'' is a genus of flowering plants in the plantain family, Plantaginaceae. It is sometimes placed in the families Scrophulariaceae or Veronicaceae. The name honours Scottish naturalist Alexander Russell (1715–1768). Members of the gen ...
, and
Cudweed Cudweed is a common name for several species, and may refer to: Plants in the family Asteraceae * '' Gamochaeta'', a plant genus with species in North and South America * ''Gnaphalium'', a plant genus with species in Eurasia and the Americas * ''Fi ...
.


Host plant preferences

Iridoid
glycoside In chemistry, a glycoside is a molecule in which a sugar is bound to another functional group via a glycosidic bond. Glycosides play numerous important roles in living organisms. Many plants store chemicals in the form of inactive glycosides. ...
s trigger feeding behaviors of ''J. coenia'' larvae in northern California, serving as attractants and as stimulants. It also allows for higher growth and survival rates that appear to be best promoted in the plant ''
Plantago lanceolata ''Plantago lanceolata'' is a species of flowering plant in the plantain family Plantaginaceae. It is known by the common names ribwort plantain, narrowleaf plantain, English plantain, ribleaf, lamb's tongue, and buckhorn. It is a common weed on c ...
'', an invasive plant in California. In fact, leaves from this plant were preferred over artificial diets with just the iridoid glycosides, indicating the presence of additional sources in ''P. lanceolata'' that might play a role in food preferences. Naturally, ''J. coenia'' consider iridoid glycosides in the host plant selection for larvae. This inclination for iridoid glycosides may also reflect its role in
oviposition The ovipositor is a tube-like organ used by some animals, especially insects, for the laying of eggs. In insects, an ovipositor consists of a maximum of three pairs of appendages. The details and morphology of the ovipositor vary, but typical ...
and microhabitat selection. Aucubin and catalpol are two other chemicals that can be stimulants and attractants for larval feeding behaviors.


Pollinator cues

In the case of ''
Lantana camara ''Lantana camara'' (common lantana) is a species of flowering plant within the verbena family (Verbenaceae), native to the American tropics. It is a very adaptable species, which can inhabit a wide variety of ecosystems; once it has been introduc ...
'' flowers, ''J. coenia'' preferentially feed on flowers in the yellow stage at a significantly higher rate that suggests
associative learning Learning is the process of acquiring new understanding, knowledge, behaviors, skills, values, attitudes, and preferences. The ability to learn is possessed by humans, animals, and some machines; there is also evidence for some kind of learnin ...
. During early life stages, the butterfly feeds upon yellow and red flowers almost equally but over time focuses only on those flowers that are yellow and 'pre-change.' This is especially advantageous because the flower's color can be linked to its viability as well as the nectar reward that it can provide the pollinator. With this signal the pollinator can visit the right plant at the right time maximizing both its reproductive advantage as well as that of the flower. Moreover, the amount of nectar found within the flower can also influence the duration of a pollinator's visit to the plant.


Parental Care


Oviposition

In the process of oviposition, contact chemoreception is an important sense used to detect
allelochemicals Allelopathy is a biological phenomenon by which an organism produces one or more biochemicals that influence the germination, growth, survival, and reproduction of other organisms. These biochemicals are known as allelochemicals and can have be ...
on individual leaves that promote or discourage oviposition. Females detect iridoid glycosides found in plants like ''P. lanceolata'' as oviposition cues. Similarly, a mix of catalpol and aucubin, as well as catalpol alone, also promotes oviposition. In addition, the quantity of catalpol also seems to play a role in oviposition because females choose to lay eggs in regions that are most concentrated with catalpol. Just one iridoid glycoside molecule is enough to promote oviposition to the same degree that would be seen on a hostplant leaf. However, with that said, females did prefer plants with more iridoid glycosides. Larvae concentrate large amounts of iridoid glycosides, and theoretical explanations for this include protection from predators. Therefore, evolution would favor female ''J. coenia'' that can detect more iridoid glycosides on host plants so that larvae can incorporate that into their diet and thus gain better protective mechanisms for survival.


Social behavior


Caterpillar sociality

Caterpillars tend to feed individually and in isolation. For example, even in the case that multiple caterpillars are feeding upon the same plant, they will not be doing so cooperatively or based in a group dynamic. Caterpillars are generally not aggressive: they do not mind each other's presence and tend to abstain from fighting when running into one another.


Migration

Common buckeyes move to the south along with tailwinds directed to the north or northwest after the cold fronts from September or October. They are sensitive to the cold and cannot spend the winter in northern regions that will experience extreme cold temperatures. However, they will migrate back from the south during the spring. It was spotted in California in late summer, early fall of 2022.


Local or regional dispersal

''J. coenia'' appears to readily migrate on a local scale between patches that are connected by corridors over those that are not connected by corridors. They could also affect large scale immigration by making patches more accessible to immigrating butterflies if they are connected by corridors as opposed to if they are unconnected. As a result, this would play a role in increasing the population densities of these habitats, which would then affect resource availability for affected individuals.


Enemies


Predators

Predators for these organisms include ants, wasps, birds, and small animals. Predators appear to select larvae that feed on artificial diets deprived of iridoid glycoside (iridoid glycoside poor) rather than those that consume organic leaf material (iridoid glycoside rich larvae), which may indicate the role of iridoid glycoside consumption. This selection then favors larvae that have diets including iridoid glycosides as well as females who oviposition on those sites by providing offspring with necessary defense mechanisms.


Defense effect on predators

Prey which contain iridoid glycosides can affect invertebrate predators differently. Chemical differences as a result of leaf age can affect the development of these butterflies, which can then affect the quality of these butterflies as food sources for spiders. One such observation indicated that iridoid glycosides may diminish growth of predators so that predators select for caterpillars that do not feed on iridoid glycoside-rich substrates. Iridoid glycosides have been shown to stimulate growth in prey while decreasing growth in predators, which may then be responsible for its role in providing chemical protection to butterflies.


Virus

The densovirus ''Junonia coenia'' infects the common buckeye larvae by focusing on targeting dividing cells in order to propagate. It consists of single stranded DNA genomes with a smooth, sphere-shaped capsid. It is from the family Parvoviridae and has been considered a potential
insecticide Insecticides are substances used to kill insects. They include ovicides and larvicides used against insect eggs and larvae, respectively. Insecticides are used in agriculture, medicine, industry and by consumers. Insecticides are claimed to b ...
, as is hypothesized for other one-species densoviruses. The ''Junonia coenia'' densovirus is unique in its separation of DNA strands for structural proteins versus nonstructural proteins.


Parasites

High intake of iridoid glycosides by ''J. coenia'' can have a negative effect on their immune response. This can then lead to higher susceptibility to parasitoids as well as a decreased ability to resist parasitism. Therefore, it is necessary to balance the chemical defense provided by these secondary metabolites with their immunological cost that correlates with increased risk of parasitism. Specifically, secondary metabolites may be hindering the immune response by operating through decreased melanization. This becomes even more problematic since parasitism is a prominent cause of mortality for caterpillars. This relationship exemplifies the vulnerable host hypothesis.


Immunity

Common predators of ''J. coenia'' larvae are ants, and chemical defense appears to be a strong survival mechanism for these organisms against predation. The concentration of iridoid glycoside sequestered by larvae seems to be a strong predictor for the decreased likelihood that they will be predated.


Physiology


Gustation (taste)

In order to find a host plant for oviposition, females search for appropriate concentrations of catalpol or iridoid glycosides. Therefore, they practice drumming, which allows them to pierce the plant exterior and ‘taste’ the variety of allelochemicals that are present within its structure. This can be detected through the female ''J. coenia'' foretarsal chemoreceptors.


Protective coloration and behavior


Automimicry

Larvae also appear to weakly adopt automimicry as a survival strategy. In the presence of caterpillars that have sequestered iridoids (making them unpalatable to predators) and those that have not sequestered iridoids (making them palatable to predators), unpalatable caterpillars are able to escape predation. However, this trend is a weak example because it also may be the consequence of predators entirely avoiding this species of caterpillars rather than predators rejecting caterpillars.


In popular culture

The common buckeye was featured on the 2006 United States Postal Service 24-cent postage stamp.


References


Citations


Literature

* * * * * * * *


External links


common buckeye
on the UF /
IFAS IFAS may refer: * Institute of Food and Agricultural Sciences * Integrated Fixed-Film Activated Sludge, a sewage treatment process * International French adjectival system In rock climbing, mountaineering, and other climbing disciplines, clim ...
Featured Creatures Web site
Common Buckeye
Butterflies of Canada {{Taxonbar, from=Q1257151 Butterflies of Central America Butterflies of Cuba Butterflies of North America Butterflies of the Caribbean coenia Nymphalidae of South America Taxa named by Jacob Hübner