Monarch
Female Male
Scientific classification
Kingdom: Animalia Phylum: Arthropoda Class: Insecta Order: Lepidoptera Superfamily: Papilionoidea Family: Nymphalidae Subfamily: Danainae Tribe: Danaini Genus: Danaus Kluk, 1780 Species: D. plexippus
Binomial name
Danaus plexippus (Linnaeus, 1758)

The monarch (Danaus plexippus) is a milkweed butterfly (subfamily Danainae), in the family Nymphalidae. It is perhaps the best known of all North American butterflies. Since the 19th century, it is also found in New Zealand, and has been known in Australia since 1871. [1] [2] In Australia it is also known as the wanderer butterfly. [3] In Europe it is resident in the Canary Islands and Madeira, and is found as a migrant in Russia, Azores, Sweden and Spain. Its wings feature an easily recognizable orange and black pattern, with a wingspan of 8.5–12.5 cm (3.34 in–4.92in). (The viceroy butterfly has a similar size, color, and pattern, but can be distinguished by an extra black stripe across the hindwing.) Female monarchs have darker veins on their wings, and the males have a spot in the center of each hindwing [4] from which pheromones are released. Males are also slightly larger.

Taxonomy

The common name “monarch” was coined in 1874 by Samuel H. Scudder because “it is one of the largest of our butterflies, and rules a vast domain”.^[1] The monarch was one of the many species originally described by Linnaeus in his Systema Naturae of 1758. It was first placed in the genus Papilio.^[2] In 1780, Jan Krzysztof Kluk used the monarch as the type species for a new genus; Danaus. Since Latin grammar requires that the specific epithet and gender names agree it is unclear if the genus Danaus is drawn from Danaus (Greek Δαναός), a mythical king of Egypt and great-grandson of Zeus or is a masculinised version of Danae (Greek Δανάη), Danaus’ great-great-granddaughter. The species name, plexippus, refers to Plexippus, one of the 50 sons of Aegyptus, Danaus’ twin brother.^[1][3]

Underside of a Danaus plexippus plexippus feeding on nectar from an Echinacea purpurea flower

This butterfly is closely related to one or two very similar species which were formerly considered to form the Danaus (Danaus) subgenus (see Smith et al. 2005). One is the Jamaican monarch, Danaus cleophile, of Jamaica and Hispaniola.

The other is a butterfly which is almost indistinguishable from the monarch as an adult (the pupae are somewhat more different) and occurs south of the Amazonas river. This taxon, Danaus erippus ("Southern monarch"), is often considered a subspecies of the monarch butterfly proper. Analysis of morphological, mtDNA 12S rRNA and cytochrome c oxidase subunit I, and nuclear DNA 18S rRNA and EF1 subunit α sequence data (Smith et al. 2005) indicates that it is better considered a distinct species.

However, as a species the southern monarch is only comparatively recently evolved. In all likelihood, its ancestors separated from the monarch's population some 2 mya, at the end of the Pliocene, when sea levels were higher and the entire Amazonas lowland was a vast expanse of brackish swamp that offered hardly any butterfly habitat (Smith et al. 2005).

Following the review of Smith et al. (2005), two subspecies are nowadays recognized:

• Danaus plexippus plexippus

  The migratory subspecies known from most of the USA

• Danaus plexippus megalippe

  The non-migratory subspecies which is found from Florida and Georgia southwards, throughout the Caribbean and Central America to the Amazon River. This has several local forms:

  • Danaus plexippus megalippe forma leucogyne
  • Danaus plexippus megalippe forma portoricensis
  • Danaus plexippus megalippe forma tobagi

Migration

Monarchs are especially noted for their lengthy annual migration. In North America they make massive southward migrations starting in August until the first frost. A northward migration takes place in the spring. (In Australia, they make limited migrations in cooler areas, [5] but the blue tiger butterfly is better known in Australia for its lengthy migration. [6]) Female monarchs deposit eggs for the next generation during these migrations. By the end of October, the population of the Rocky Mountains migrates to the sanctuaries in the area of Angangueo, Ocampo, Zitácuaro and El Rosario in the Mexican state of Michoacán. The western population overwinters in various sites in central coastal California, United States, notably in Pacific Grove and Santa Cruz.

The length of these journeys exceeds the normal lifespan of most monarchs, which is less than two months for butterflies born in early summer. The last generation of the summer enters into a non-reproductive phase known as diapause and may live seven months or more. [7] During diapause, butterflies fly to one of many overwintering sites. The generation that overwinters generally does not reproduce until it leaves the overwintering site sometime in February and March. It is thought that the overwinter population may reach as far north as Texas and Oklahoma during the spring migration. It is the second, third and fourth generations that return to their northern locations in the United States and Canada in the spring. How the species manages to return to the same overwintering spots over a gap of several generations is still a subject of research; the flight patterns appear to be inherited, based on a combination of circadian rhythm and the position of the sun in the sky.^[4]

Monarch butterflies are one of the few insects capable of making transatlantic crossings. They are becoming more common in Bermuda due to increased usage of milkweed as an ornamental plant in flower gardens. Monarch butterflies born in Bermuda remain year round due to the island's mild climate.

A few monarchs turn up in the far southwest of Great Britain in years when the wind conditions are right, and have been sighted as far east as Long Bennington. Monarchs can also be found in New Zealand. On the islands of Hawaii no migrations have been noted.

Reproduction

The mating period for the overwinter population occurs in the spring, just prior to migration from the overwintering sites. The courtship is fairly simple and less dependent on chemical pheromones in comparison with other species in its genus. [8] Courtship is composed of two distinct stages, the aerial phase and the ground phase. During the aerial phase, the male pursues, nudges, and eventually takes down the female. Copulation occurs during the ground phase and involves the transfer of a spermatophore from the male to the female. Along with sperm, the spermatophore is thought to provide the female with energy resources that aid her in carrying out reproduction and remigration. The overwinter population returns only as far north as they need to go to find the early milkweed growth; in the case of the eastern butterflies that is commonly southern Texas. The life cycle of a monarch includes a change of form called complete metamorphosis. The monarch goes through four radically different stages:

1. The eggs are laid by the females during spring and summer breeding months.
2. The eggs hatch, revealing worm-like larva, the caterpillars. The caterpillars consume their egg cases, then feed on milkweed, and sequester substances called cardenolides, a type of cardiac glycosides. During the caterpillar stage, monarchs store energy in the form of fat and nutrients to carry them through the non-feeding pupa stage.
3. In the pupa or chrysalis stage, the caterpillar spins a silk pad on a twig, leaf, etc, and hangs from this pad by its last pair of prolegs. It hangs upside down in the shape of a 'J', and then molts, leaving itself encased in an articulated green exoskeleton. At this point, hormonal changes occur, leading to the development of a butterfly. The chrysalis darkens (actually becomes transparent) a day before it emerges, and its orange and black wings can be seen.
4. The mature butterfly emerges after about two pupal weeks and hangs from the split chrysalis for several hours until its wings are dry (often in the morning). Meanwhile fluids are pumped into the crinkled wings until they become full and stiff. Some of this orangy fluid drips from the wings. Finally (usually in the afternoon) the monarch spreads its wings, quivers them to be sure they are stiff, and then flies in a circle and away, to feed on a variety of flowers, including milkweed flowers, red clover, and goldenrod.

Monarchs can live a life of twenty to eighty weeks in a garden having their host Asclepias plants and sufficient flowers for nectar. This is especially true if the flower garden happens to be surrounded by native forest that seems to be lacking in flowers.

Pictorial lifecycle

Monarch male showing its wings to attract a mate	Monarch butterflies mating	Monarch butterfly laying eggs on Asclepias curassavica	Monarch eggs on milkweed
An early instar monarch caterpillar	A late instar caterpillar feeding	Monarch caterpillar beginning pupation	Monarch butterfly chrysalis
Butterfly emerging from chrysalis	Adult monarch butterfly feeding on a Zinnia	Wintering monarch butterflies

Defense against predators

Swamp milkweed, one of many species of Asclepias milkweeds used by the monarch

Monarchs are foul-tasting and poisonous due to the presence of cardenolide aglycones in their bodies, which the caterpillars ingest as they feed on milkweed. [9] Both forms advertise their unpalatability with bright colors and areas of high contrast on the skin or wings. This phenomenon is known as aposematism.

Monarchs share this defense with the even more unpleasant-tasting and similar-appearing viceroy butterfly, in an example of Müllerian mimicry. (Viceroys were at one time believed to be Batesian mimics of monarchs.)

The monarch (left) and viceroy (right) butterflies exhibiting Müllerian mimicry

Relationship with humans

A monarch butterfly in a butterfly garden

The monarch is the state insect of Alabama,^[5] Idaho,^[6] Illinois,^[7] Minnesota,^[8] Texas,^[9] and the state butterfly of Vermont^[10] and West Virginia.^[11] It was nominated in 1990 as the national insect of the United States of America, along with the honeybee (Apis mellifera),^[12] but the legislation did not pass.^[13]

Many people like to attract monarchs by growing a butterfly garden. Others enjoy raising them for pleasure or for educational purposes. For migrating flocks, sanctuaries have been created at favorite wintering locations; Even tourism revenue is thus generated.

Many schools also enjoy growing, and attending to monarch butterflies, starting with the caterpillar form. When the butterflies reach adulthood they are then released into the wild.

Threats

Recent illegal deforestation of the monarch's overwintering grounds have led to a drastic reduction in the butterfly's population. Efforts to classify it as a protected species and to restore its habitat are under way. Mexican authorities expected a significant increase in the monarch population in the 2005-2006 season.

Predators

Although monarchs feed on milkweed, variations in the quantity of cardiac glycosides exist between species, individuals and even parts of the host plant. The levels of toxins in adult monarchs reflect the levels in their host plants. This means some monarchs are not foul-tasting but are Batesian or auto-mimics. Some species of predators have learnt to measure the toxins by taste and reject butterflies with high cardiac glycosides contents, eating only the ones with low cardiac glycosides contents. In the butterfly, the cardiac glycosides are concentrated in the abdomen and wings. Some species of predators differenciate these parts and consume only the most palatable ones.^[14] Bird predators include Brown Thrashers, Grackles, Robins, Cardinals, Sparrows, Scrub Jays and Pinyon Jays.^[14]

In North America, eggs and first instar larvae of the monarch are eaten by larvae and adults of the introduced Asian lady beetle (Harmonia axyridis).^[15]

White morph of the monarch in Hawaii

On Oahu, a white morph of the monarch has emerged. This is because of the introduction, in 1965 and 1966, of two bulbul species, Pycnonotus cafer and Pycnonotus jacosus. They are now the most common insectivore birds, and probably the only ones preying on insects as big as the monarch. Monarchs in Hawaii are known to have low cardiac glycoside levels, but the birds may also be tolerant for the chemical. The two species hunt the larvae and some pupae from the branches and underside of leaves in milkweed bushes. The bulbuls also eat resting and ovipositing adults, but rarely flying ones. Because of its colour the white morph has a higher survival rate than the orange one. This is either because of apostatic selection (i.e. the birds have learnt the orange monarchs can be eaten) or because of camouflage; the white morph matches the white pubescence of milkweed, or the patches of light shining through foliage.^[16]

Parasites

Parasites include the tachinid fly Lesperia archippivora. Parasitized larvae complete their moult, suspend, but die before pupation. At that time one white maggot comes out of the larvae, suspended by a silken thread. The maggot then forms a brown pupa on the ground.^[17]

The bacteria Micrococcus flacidifex danai also infects the larvae and causes “black death”. As usual, just before pupation the larvae migrate to a horizontal surface. They die a few hours later, attached only by one pair of prolegs, thorax and abdomen hanging limp. The body turns black shortly after. The bacteria Pseudomonas aeriginosa has no invasive powers, but causes secondary infections in weakened insects. It is a common cause of death in laboratory reared insects.^[17]

The protozoan Ophryocystis elektroscirrha is another parasite of the monarch. It infects the subcutaneous tissues and propagates by spores formed during the pupal stage. The spores are found over all of the body of infected butterflies, with the greatest number on the abdomen. These spores are passed, from female to caterpillar, when spores rub off during egg-laying and are then ingested by caterpillars. Severely infected individuals are weak, unable to expand their wings, or unable to eclose, and have a shortened lifespan but probably occur a low frequencies in nature. This is not the case in laboratory or commercial rearing, where after a few generations all individuals can be infected.^[18]

References

• Smith, David A. S.; Lushai, Gugs & Allen, John A. (2005): A classification of Danaus butterflies (Lepidoptera: Nymphalidae) based upon data from morphology and DNA. Zool. J. Linn. Soc. 144(2): 191–212. doi:10.1111/j.1096-3642.2005.00169.x (HTML abstract)

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External links

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Danaus plexippus

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Danaus plexippus

• Monarch butterfly facts and photos - NatureMapping Program
• Video of a monarch butterfly emerging from chrysalis
• Australian Museum fact sheet on monarch butterflies
• Critters of Calamvale Creek — a monarch tells its story
• Monarch Migration Maps
• Monarch Watch of the University of Kansas Entomology Department
• Michoacan Reforestation Fund
• USGS description of monarch butterfly
• MonarchHealth!
• Monarch Butterfly Trust NZ
• Monarch butterfly life cycle photographs
• More Than Monarchs