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The eastern giant swallowtail (''Papilio cresphontes'') is the largest
butterfly Butterflies are winged insects from the lepidopteran superfamily Papilionoidea, characterized by large, often brightly coloured wings that often fold together when at rest, and a conspicuous, fluttering flight. The oldest butterfly fossi ...
in
North America North America is a continent in the Northern Hemisphere, Northern and Western Hemisphere, Western hemispheres. North America is bordered to the north by the Arctic Ocean, to the east by the Atlantic Ocean, to the southeast by South Ameri ...
. It is abundant through many parts of eastern North America; populations from western North America and down into
Panama Panama, officially the Republic of Panama, is a country in Latin America at the southern end of Central America, bordering South America. It is bordered by Costa Rica to the west, Colombia to the southeast, the Caribbean Sea to the north, and ...
are now (as of 2014) considered to belong to a different species, '' Papilio rumiko''.A new Heraclides swallowtail (Lepidoptera, Papilionidae) from North America is recognized by the pattern on its neck
/ref> Though it is often valued in gardens for its striking appearance, its larval stage can be a serious pest to
citrus ''Citrus'' is a genus of flowering trees and shrubs in the family Rutaceae. Plants in the genus produce citrus fruits, including important crops such as oranges, mandarins, lemons, grapefruits, pomelos, and limes. ''Citrus'' is nativ ...
farms, which has earned its caterpillars the names orange dog or orange puppy. The eastern giant swallowtail caterpillars possess remarkable camouflage from predators by closely resembling bird droppings. They use this, along with their osmeteria, to defend against predators such as wasps, flies, and vertebrates.


Geographic range

The eastern giant swallowtail is common across the United States, reaching as far north as southern
New England New England is a region consisting of six states in the Northeastern United States: Connecticut, Maine, Massachusetts, New Hampshire, Rhode Island, and Vermont. It is bordered by the state of New York (state), New York to the west and by the ...
and southern
Canada Canada is a country in North America. Its Provinces and territories of Canada, ten provinces and three territories extend from the Atlantic Ocean to the Pacific Ocean and northward into the Arctic Ocean, making it the world's List of coun ...
. South of the United States, it is found in parts of
Mexico Mexico, officially the United Mexican States, is a country in North America. It is the northernmost country in Latin America, and borders the United States to the north, and Guatemala and Belize to the southeast; while having maritime boundar ...
and also found in
Jamaica Jamaica is an island country in the Caribbean Sea and the West Indies. At , it is the third-largest island—after Cuba and Hispaniola—of the Greater Antilles and the Caribbean. Jamaica lies about south of Cuba, west of Hispaniola (the is ...
and
Cuba Cuba, officially the Republic of Cuba, is an island country, comprising the island of Cuba (largest island), Isla de la Juventud, and List of islands of Cuba, 4,195 islands, islets and cays surrounding the main island. It is located where the ...
. The species was historically considered to occur in the western United States and into South America, but now those populations are treated as a separate species, the western giant swallowtail ('' Papilio rumiko''), based largely on DNA evidence.


Habitat

In the United States, ''P. cresphontes'' mostly inhabit
deciduous forest In the fields of horticulture and botany, the term deciduous () means "falling off at maturity" and "tending to fall off", in reference to trees and shrubs that seasonally shed leaves, usually in the autumn; to the shedding of petals, after flo ...
and citrus orchards. They are only capable of overwintering in
Florida Florida ( ; ) is a U.S. state, state in the Southeastern United States, Southeastern region of the United States. It borders the Gulf of Mexico to the west, Alabama to the northwest, Georgia (U.S. state), Georgia to the north, the Atlantic ...
and the
deep South The Deep South or the Lower South is a cultural and geographic subregion of the Southern United States. The term is used to describe the states which were most economically dependent on Plantation complexes in the Southern United States, plant ...
.


Description

One of the eastern giant swallowtail's most notable features is its size. Females have an average wingspan of , and up to , while males' average is , and up to . The wings are black with a horizontal yellow line across the
forewing Insect wings are adult outgrowths of the insect exoskeleton that enable insects to fly. They are found on the second and third thoracic segments (the mesothorax and metathorax), and the two pairs are often referred to as the forewings and hindwi ...
s, and a diagonal yellow line across the hindwing. The underside of the wings is yellow with accents of black. A small patch of red on the ventral wing (within the small blue band) allows for distinction from the similar-looking Schaus' swallowtail. Seitz -"P. cresphontes Cr. (7a). Usually considered a slightly different variety of '' P. thoas''. No cell-spot on the forewing; the fifth discal spot projecting further than the sixth. Claspers of the male separate above, the anal hook quite short, the lower part of the anal segment likewise quite different from that of ''P. thoas''; harpe broad, rounded. A common species in eastern North America, occurring as far as Costa Rica in the south and southern Canada in the north; but the insect is a wanderer, which is found only now and then in the northern districts. In the United States it does not extend westward beyond the Mississippi plain, except in the Southern States. Its true home is the region adjoining the Gulf of Mexico. It occurs also on Cuba. ''Papilio cresphontes'' is a member of the '' Papilio thoas''
species group In biology, a species complex is a group of closely related organisms that are so similar in appearance and other features that the boundaries between them are often unclear. The taxa in the complex may be able to hybridize readily with each oth ...
. '


Food resources


Caterpillars

As well as eating valuable citrus crops, larvae eat a wide variety of plants in the family
Rutaceae The Rutaceae () is a family (biology), family, commonly known as the rueRUTACEAE
in Bo ...
(citrus plants), including different types of prickly ash and common hoptree. They also consume some exotic Rutaceae species that have been introduced to North America, such as gasplant,
rue ''Ruta graveolens'', commonly known as rue, common rue or herb-of-grace, is a species of the genus '' Ruta'' grown as an ornamental plant and herb. It is native to the Mediterranean. It is grown throughout the world in gardens, especially for i ...
and sapote.


Feeding specialization

In specific local subpopulations (studied in Wisconsin, Ohio, and Florida), it was found that eastern giant swallowtail caterpillars do better on their local host plant than on other giant swallowtail host plants. The local host caused the larva to develop faster than other plants that were also edible to the larva. However, this feeding specialization hypothesis has not been tested in the species as a whole beyond these three regions.


Adults

Nectar plants for adults include ''
Lantana ''Lantana'' () is a genus of about 150 species of perennial plant, perennial flowering plants in the verbena family, Verbenaceae. They are native to tropics, tropical regions of the Americas and Africa but exist as an introduced species in num ...
'',
Azalea Azaleas ( ) are flowering shrubs in the genus ''Rhododendron'', particularly the former sections ''Rhododendron sect. Tsutsusi, Tsutsusi'' (evergreen) and ''Pentanthera'' (deciduous). Azaleas bloom in the spring (April and May in the temperate ...
s, '' Bougainvilla, Saponaria officinalis'', '' Hesperis matronalis'', '' Solidago,
Lonicera japonica ''Lonicera japonica'', known as Japanese honeysuckle and golden-and-silver honeysuckle, is a species of honeysuckle native to East Asia, including many parts of China. It is often grown as an ornamental plant, but has become an invasive species i ...
'', and ''
Asclepias incarnata ''Asclepias incarnata'', the swamp milkweed, rose milkweed, rose milkflower, swamp silkweed, or white Indian hemp, is a herbaceous perennial plant species native to North America. It grows in damp through wet soils and also is cultivated as a gar ...
''. Along with flower nectar, adults can also consume liquid from animal waste.


Life history


Eggs

Females lay eggs on the tops of the leaves of one of their preferred host plants. This is because the first larval instars are unable to move from plant to plant, so the mother must select an appropriate plant to support them. One egg is lain at a time, as opposed to in clutches. The small egg (1–1.5 mm) is brownish colored, but appears more orange due to a special orange colored secretion.


Caterpillars

Once emerged from the eggs, there are five larval instars, wherein the larva grow to about 2 inches before pupating. Larvae are mostly
nocturnal Nocturnality is a ethology, behavior in some non-human animals characterized by being active during the night and sleeping during the day. The common adjective is "nocturnal", versus diurnality, diurnal meaning the opposite. Nocturnal creatur ...
, feeding at night. Their appearance changes slightly throughout the
instar An instar (, from the Latin '' īnstar'' 'form, likeness') is a developmental stage of arthropods, such as insects, which occurs between each moult (''ecdysis'') until sexual maturity is reached. Arthropods must shed the exoskeleton in order to ...
s, with the younger ones having
seta In biology, setae (; seta ; ) are any of a number of different bristle- or hair-like structures on living organisms. Animal setae Protostomes Depending partly on their form and function, protostome setae may be called macrotrichia, chaetae, ...
e and the older instars lacking setae. The caterpillars have remarkable
camouflage Camouflage is the use of any combination of materials, coloration, or illumination for concealment, either by making animals or objects hard to see, or by disguising them as something else. Examples include the leopard's spotted coat, the b ...
patterns.


Pupation

Larvae must find a vertical plant, or sometimes a man-made object, to form their chrysalis on, and often they choose the host plant they already occupy. They attach themselves to their substrate of choice and molt to reveal a brown, mottled chrysalis (resembling a bit of dead twig) in which they remain for approximately 10–12 days.


Mating

Males search for females along set flight paths and near host plants, and mate with females in the afternoon. Male and female then copulate facing away from each other.


Enemies


Parasites

Eastern giant swallowtails are most vulnerable to
parasites Parasitism is a close relationship between species, where one organism, the parasite, lives (at least some of the time) on or inside another organism, the host, causing it some harm, and is adapted structurally to this way of life. The en ...
when they are in their
chrysalis A pupa (; : pupae) is the life stage of some insects undergoing transformation between immature and mature stages. Insects that go through a pupal stage are holometabolous: they go through four distinct stages in their life cycle, the stages the ...
. Common parasites include flies and wasps such as '' Brachymeria robusta'', '' Pteromalus cassotis'', ''Pteromalus vanessae'', and '' Lespesia rileyi''.


Protective coloration and behavior

Larvae have many adaptations to protect themselves from predators.


Coloration

The caterpillars' intricate coloring patterns are an effective
camouflage Camouflage is the use of any combination of materials, coloration, or illumination for concealment, either by making animals or objects hard to see, or by disguising them as something else. Examples include the leopard's spotted coat, the b ...
and defense against both vertebrate and invertebrate predators and parasites. It is thought that the coloration was naturally selected for because of its imitation of bird and lizard droppings. The caterpillar mimics certain droppings based on its habitat and which instar it is. The caterpillars' coloration, particularly the saddle pattern, is also thought to be
disruptive coloration Disruptive coloration (also known as disruptive camouflage or disruptive patterning) is a form of camouflage that works by breaking up the outlines of an animal, soldier or military hardware with a strongly contrasting pattern. It is often com ...
. This means the coloration makes it harder for a predator to distinguish the shape of the camouflaged prey, which explains the continuation of this coloration pattern in larger instars that are too large to be mistaken for bird droppings. It has also been hypothesized that the older instars' pattern is meant to be reminiscent of a snake.


Other defensive measures

When camouflage is not enough, larvae will employ their osmeteria when they are threatened. The osmeterium is an organ behind the head that “inflates” into an orange/red Y-shaped growth that resembles a snake's forked tongue. By the fourth instar, the osmeterium is more than just a startle mechanism and also has bad-smelling and toxic mixture of acidic chemicals. This is only effective on small invertebrate predators, and the caterpillar will try to rub its osmeterium on the predator to deter it. It has been experimentally tested and found that osmeteria are ineffective at deterring predation from birds.


Physiology


Identifying host plants

Eastern giant swallowtail butterflies must correctly identify their host plants by antennal sensitivity to the specific volatile compounds in the plants. A study found that antennal response to these volatiles depends upon the concentration of the volatiles, the host plant of origin (whether it is a primary or secondary host), and the sex of the butterfly. This last dependency is thought to be because the females, not the males, must identify the correct host plant for egg laying.


Flight

Because of the butterfly's notably large wingspan (14 to 18 cm), eastern giant swallowtails are very strong fliers and are able to glide long distances with very few wing beats.


Interactions with humans

Sometimes referred to as "orange dogs", "orange pups", or "bird poop caterpillars" by farmers, the larva targets all varieties of citrus plants, often causing significant damage to new foliage and younger trees, which they can more thoroughly defoliate. Large mature citrus trees are not generally significantly impacted. Outside of farms, the species is valued for its aesthetic appeal and can also be raised to butterflies successfully at home.


Control

Biological insecticides, such as ''
Bacillus thuringiensis ''Bacillus thuringiensis'' (or Bt) is a gram-positive bacteria, gram-positive, soil-dwelling bacterium, the most commonly used biological pesticide worldwide. ''B. thuringiensis'' also occurs naturally in the gut of caterpillars of various types ...
'', as well as chemical insecticides, are used to protect trees against larvae.
Methionine Methionine (symbol Met or M) () is an essential amino acid in humans. As the precursor of other non-essential amino acids such as cysteine and taurine, versatile compounds such as SAM-e, and the important antioxidant glutathione, methionine play ...
, an
essential amino acid An essential amino acid, or indispensable amino acid, is an amino acid that cannot be synthesized from scratch by the organism fast enough to supply its demand, and must therefore come from the diet. Of the 21 amino acids common to all life forms ...
in humans, has also been found to be an effective killer of caterpillars, with possible use as a nontoxic pesticide against eastern giant swallowtail larvae.


Effects of climate change

There has been a northern expansion of the range of the eastern giant swallowtail in recent years, which has been linked to increasingly warm temperatures, and particularly to a lack of September frosts in regions of expansion starting in 2001. Larvae were then able to withstand a few frosts before they pupated. The immediate effects of this warming, as well as their effect on host plants and predators, can explain the giant swallowtail's range expansion.


Gallery

File:Giant swallowtail egg.jpg, Egg File:Papilio cresphontes larva defensive.JPG, ''P. cresphontes'' caterpillar showing defensive posture File:Butterfly House 1.JPG, Adult, underside File: Papilio cresphontes - The giant swallowtail.webm, In flight. Video clip


References


Further reading

* * * * * * *


External links


Giant swallowtail butterfly life cycle photographs
on the UF / IFAS Featured Creatures Web site
Butterflycorner
Images from
Naturhistorisches Museum Wien The Natural History Museum Vienna () is a large natural history museum located in Vienna, Austria. The NHM Vienna is one of the largest museums and non-university research institutions in Austria and an important center of excellence for all matt ...

Giant Swallowtail
Butterflies and Moths of North America

funet.fi
Giant Swallowtail (''Papilio cresphontes'')
CBIF
Insecta pro
{{Taxonbar, from=Q950614
cresphontes In Greek mythology, Cresphontes (; ) was a son of Aristomachus, husband of Merope, father of Aepytus and brother of Temenus and Aristodemus. He was a great-great-grandson of Heracles and helped lead the fifth and final attack on Mycenae in t ...
Butterflies described in 1777 Butterflies of Central America Lepidoptera of the United States Taxa named by Pieter Cramer Articles containing video clips Lepidoptera of Canada Lepidoptera of Mexico Fauna of Costa Rica Fauna of Panama