Battus philenor
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''Battus philenor'', the pipevine swallowtail or blue swallowtail, Retrieved April 19, 2018. is a swallowtail butterfly found in North America and
Central America Central America ( es, América Central or ) is a subregion of the Americas. Its boundaries are defined as bordering the United States to the north, Colombia to the south, the Caribbean Sea to the east, and the Pacific Ocean to the west. ...
. This butterfly is black with iridescent-blue hindwings. They are found in many different habitats, but are most commonly found in
forest A forest is an area of land dominated by trees. Hundreds of definitions of forest are used throughout the world, incorporating factors such as tree density, tree height, land use, legal standing, and ecological function. The United Nations' ...
s. 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.


Nomenclature

The pipevine swallowtail was first identified in 1771 by Linnaeus and originally was placed in the genus '' Papilo'', as was typical for butterflies. In 1777, it was moved to the genus '' Battus'' by
Scopoli Giovanni Antonio Scopoli (sometimes Latinized as Johannes Antonius Scopolius) (3 June 1723 – 8 May 1788) was an Italian physician and naturalist. His biographer Otto Guglia named him the "first anational European" and the "Linnaeus of the Au ...
. The name ''Battus'' comes from the founder of the
Greek Greek may refer to: Greece Anything of, from, or related to Greece, a country in Southern Europe: *Greeks, an ethnic group. *Greek language, a branch of the Indo-European language family. **Proto-Greek language, the assumed last common ancestor ...
colony of
Cyrenaica Cyrenaica ( ) or Kyrenaika ( ar, برقة, Barqah, grc-koi, Κυρηναϊκή παρχίαKurēnaïkḗ parkhíā}, after the city of Cyrene), is the eastern region of Libya. Cyrenaica includes all of the eastern part of Libya between ...
, Battus I, and the species name ''philenor'' comes from the Greek word meaning ‘fond of husband’. The pipevine swallowtail also belongs to the tribe
Troidini Troidini is a tribe of swallowtail butterflies that consists of some 135 species in 12 genera. Members of this tribe are superlatively large among butterflies (in terms of both wingspan and surface area) and are often strikingly coloured. Gener ...
, a group of butterflies whose larvae all feed on plants of the genus '' Aristolochia.'' They are referred to as the ''Aristolochia'' butterflies.


Distribution and habitat

The pipevine swallowtail has a wide distribution across the Northern
Americas The Americas, which are sometimes collectively called America, are a landmass comprising the totality of North and South America. The Americas make up most of the land in Earth's Western Hemisphere and comprise the New World. Along with th ...
. In the
United States The United States of America (U.S.A. or USA), commonly known as the United States (U.S. or US) or America, is a country primarily located in North America. It consists of 50 states, a federal district, five major unincorporated territori ...
, the butterfly is found in
New England New England is a region comprising 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 Can ...
down to
Florida Florida is a state located in the Southeastern region of the United States. Florida is bordered to the west by the Gulf of Mexico, to the northwest by Alabama, to the north by Georgia, to the east by the Bahamas and Atlantic Ocean, and to ...
west to
Nebraska Nebraska () is a state in the Midwestern region of the United States. It is bordered by South Dakota to the north; Iowa to the east and Missouri to the southeast, both across the Missouri River; Kansas to the south; Colorado to the sout ...
,
Texas Texas (, ; Spanish: ''Texas'', ''Tejas'') is a state in the South Central region of the United States. At 268,596 square miles (695,662 km2), and with more than 29.1 million residents in 2020, it is the second-largest U.S. state by ...
,
New Mexico ) , population_demonym = New Mexican ( es, Neomexicano, Neomejicano, Nuevo Mexicano) , seat = Santa Fe , LargestCity = Albuquerque , LargestMetro = Tiguex , OfficialLang = None , Languages = English, Spanish ( New Mexican), Navajo, Ke ...
,
Arizona Arizona ( ; nv, Hoozdo Hahoodzo ; ood, Alĭ ṣonak ) is a state in the Southwestern United States. It is the 6th largest and the 14th most populous of the 50 states. Its capital and largest city is Phoenix. Arizona is part of the Fou ...
,
California California is a state in the Western United States, located along the Pacific Coast. With nearly 39.2million residents across a total area of approximately , it is the most populous U.S. state and the 3rd largest by area. It is also the m ...
, and
Oregon Oregon () is a state in the Pacific Northwest region of the Western United States. The Columbia River delineates much of Oregon's northern boundary with Washington, while the Snake River delineates much of its eastern boundary with Idaho. T ...
. There is also an isolated population in
central California Central California is generally thought of as the middle third of the state, north of Southern California, which includes Los Angeles, and south of Northern California, which includes San Francisco. It includes the northern portion of the S ...
. They have been observed as far south as
Mexico Mexico (Spanish: México), officially the United Mexican States, is a country in the southern portion of North America. It is bordered to the north by the United States; to the south and west by the Pacific Ocean; to the southeast by Guatema ...
and as far north as
Ontario Ontario ( ; ) is one of the thirteen provinces and territories of Canada.Ontario is located in the geographic eastern half of Canada, but it has historically and politically been considered to be part of Central Canada. Located in Central C ...
, although these sightings are inconsistent. The butterfly frequents warmer, more
temperate In geography, the temperate climates of Earth occur in the middle latitudes (23.5° to 66.5° N/S of Equator), which span between the tropics and the polar regions of Earth. These zones generally have wider temperature ranges throughout ...
environments, and can be found mostly in open
grassland A grassland is an area where the vegetation is dominated by grasses ( Poaceae). However, sedge ( Cyperaceae) and rush ( Juncaceae) can also be found along with variable proportions of legumes, like clover, and other herbs. Grasslands occur na ...
s,
woodland A woodland () is, in the broad sense, land covered with trees, or in a narrow sense, synonymous with wood (or in the U.S., the ''plurale tantum'' woods), a low-density forest forming open habitats with plenty of sunlight and limited shade (se ...
s,
meadow A meadow ( ) is an open habitat, or field, vegetated by grasses, herbs, and other non- woody plants. Trees or shrubs may sparsely populate meadows, as long as these areas maintain an open character. Meadows may be naturally occurring or arti ...
s, and backyard gardens. They favor anywhere pipevine (''Aristolochia)'' can grow in abundance.


Host plants and food resources


Larvae

Pipevine swallowtail larvae feed on plants within the genus '' Aristolochia.'' These are commonly known as pipevine plants, which is where the butterfly gets its common name. This plant genus is known to have active aristolochic acids, which pipevine swallowtail larvae sequester and use for defensive properties. A common host plant is the
Virginia snakeroot ''Aristolochia serpentaria'' is a species of perennial flowering plant in the Aristolochiaceae (birthwort) family. The species is commonly known as Virginia snakeroot and is native to eastern North America, from Connecticut to southern Michigan ...
, (''Aristolochia serpentaria''), which can be found in the Eastern United States and in Florida. It has broad-leaved and narrow-leaved forms, and both act as hosts for swallowtail larvae. Other host plants include Pipevine ('' Aristolochia macrophylla'')'','' Woolly Dutchman’s pipe ''(Aristolochia tomentosa),'' Texas Dutchman’s pipe'' (Aristolochia reticulate),'' Watson's Dutchman’s pipe ''(Aristolochia watsonii),'' and California Dutchman’s pipe (''Aristolochia californica).'' Some ''Aristolochia'' species are actually toxic or extremely distasteful to larvae and thus may act as death traps for the butterflies. This is especially true of exotic variants.


Adults

Pipevine swallowtail adults use nectar-producing plants as hosts; there are many plants that fit this criterion. These butterflies frequent
thistle Thistle is the common name of a group of flowering plants characterised by leaves with sharp prickles on the margins, mostly in the family Asteraceae. Prickles can also occur all over the planton the stem and on the flat parts of the leaves ...
(''
Cirsium ''Cirsium'' is a genus of perennial and biennial flowering plants in the Asteraceae, one of several genera known commonly as thistles. They are more precisely known as plume thistles. These differ from other thistle genera (''Carduus'', '' Sily ...
'') flowers, the pink and purple flowers of the ''
Phlox ''Phlox'' (; Greek φλόξ "flame"; plural "phlox" or "phloxes", Greek φλόγες ''phlóges'') is a genus of 67 species of perennial and annual plants in the family Polemoniaceae. They are found mostly in North America (one in Siberia) in di ...
'' species, and ironweed of ''
Vernonia ''Vernonia'' is a genus of about 350 species of forbs and shrubs in the Daisy family Asteraceae. Some species are known as ironweed. Some species are edible and of economic value. They are known for having intense purple flowers. There have been ...
'' species.


Life cycle


Eggs

The eggs of the pipevine swallowtail vary in color from red to orange. They are small and are laid on host plant stems. A distinctive feature of these eggs which is shared by all ''Aristolochia'' butterflies is that the outer layer of the egg is covered by a firm and nourishing
excretion Excretion is a process in which metabolic waste is eliminated from an organism. In vertebrates this is primarily carried out by the lungs, kidneys, and skin. This is in contrast with secretion, where the substance may have specific tasks after ...
laid in vertical stripes along the outside. The egg appears bumpy due to the excretion forming large beads along the bands. The female pipevine swallowtail deposits this excretion from a gland above the ovipositor during the egg laying process. These butterflies lay their eggs on the leaves of host plants in clusters with access to sunlight. The larvae hatch after a few weeks and immediately eat the remnants of the egg from which they emerged.


Larvae

Pipevine swallowtail larvae are around 5 cm in length and vary in color from dark brown to black. In areas of higher temperature such as
Texas Texas (, ; Spanish: ''Texas'', ''Tejas'') is a state in the South Central region of the United States. At 268,596 square miles (695,662 km2), and with more than 29.1 million residents in 2020, it is the second-largest U.S. state by ...
and
Arizona Arizona ( ; nv, Hoozdo Hahoodzo ; ood, Alĭ ṣonak ) is a state in the Southwestern United States. It is the 6th largest and the 14th most populous of the 50 states. Its capital and largest city is Phoenix. Arizona is part of the Fou ...
, a red coloring dominates. Larvae have bright orange spots on the ends of
tubercle In anatomy, a tubercle (literally 'small tuber', Latin for 'lump') is any round nodule, small eminence, or warty outgrowth found on external or internal organs of a plant or an animal. In plants A tubercle is generally a wart-like projection ...
s in rows along their body, and at either end of the body the tubercles are elongated into filaments. Full grown larvae have a glossy or velvet appearance due to many fine hairs. Larvae sex determination can be done by looking at pits along the
ventral Standard anatomical terms of location are used to unambiguously describe the anatomy of animals, including humans. The terms, typically derived from Latin or Greek roots, describe something in its standard anatomical position. This position prov ...
surface at the ends of the
abdominal The abdomen (colloquially called the belly, tummy, midriff, tucky or stomach) is the part of the body between the thorax (chest) and pelvis, in humans and in other vertebrates. The abdomen is the front part of the abdominal segment of the torso ...
sections. Larvae spend almost all their time eating the leaves of their host plants. Once they completely remove edible matter from one plant they move to the next one. They eat in groups at first but as food sources deplete they become more solitary as they move on in search of fresh host plants. Larvae filaments help them identify vertical plants, which are then determined to be host or non-host by using their mouthparts. The larvae feed and wander for several weeks before finding an isolated spot to
pupa A pupa ( la, pupa, "doll"; plural: ''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 thei ...
te.


Pupae

The pipevine swallowtail
pupa A pupa ( la, pupa, "doll"; plural: ''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 thei ...
e are colored green or brown. They are about 6 to 7 cm in length. These pupae differ from other swallowtail butterfly pupae in that the sides of their pupae bodies widen into a sort of a winged appearance, with distinctive purplish edges along the sides of these extensions. Pipevine swallowtail pupation takes place many feet off the ground on tree trunks or other suitable spots. Pupation is rare on green surfaces. Pupation begins when the larva releases
silk Silk is a natural protein fiber, some forms of which can be woven into textiles. The protein fiber of silk is composed mainly of fibroin and is produced by certain insect larvae to form cocoons. The best-known silk is obtained from the ...
to form a support structure so that the chrysalis can hang safely. In colder climates, it will spend the winter as a pupa, but in warmer
climate Climate is the long-term weather pattern in an area, typically averaged over 30 years. More rigorously, it is the mean and variability of meteorological variables over a time spanning from months to millions of years. Some of the meteorologi ...
s, pupation only lasts a few weeks. Prior to emerging as an adult, the wing markings of the butterfly can be seen through the chrysalis.


Adults

Pipevine swallowtail adults have a
wingspan The wingspan (or just span) of a bird or an airplane is the distance from one wingtip to the other wingtip. For example, the Boeing 777–200 has a wingspan of , and a wandering albatross (''Diomedea exulans'') caught in 1965 had a wingspan o ...
from 7 to 13 cm. The dorsal wings of an adult male are black with vibrant blue iridescence on the hind wings. Female iridescence is duller in appearance. Bright orange spots are visible on the back end of the
ventral Standard anatomical terms of location are used to unambiguously describe the anatomy of animals, including humans. The terms, typically derived from Latin or Greek roots, describe something in its standard anatomical position. This position prov ...
wings. The adults from Californian populations of these butterflies have smaller and hairier bodies and are thought to be a subspecies '' Battus philenor hirsute.'' The adult pipevine swallowtail male spends most of its time feeding and searching for mates. Males have also been observed to take moisture and nutrients from mud, a behavior that is motivated by the presence of other males. Females spend their time feeding, being courted by males, and reproducing. Immediately after emerging from the chrysalis, adult butterflies spend time near the pupa case and dry their wings and remove pupal waste products from their bodies. The butterflies are seen during the spring and summer months. File:Battus philenor (18466985321).jpg, Pipevine swallowtail eggs with vertical stripes File:Battus philenor Caterpillar.jpg, Red pipevine swallowtail larva File:Pipevine Swallowtail larva, Megan McCarty52.jpg, Dark pipevine swallowtail larva File:Pipevine Swallowtail chrysalis, Megan McCarty53.jpg, Pipevine swallowtail chrysalis File:Battus philenor - dorsal.jpg, Pipevine swallowtail dorsal view


Mating


Mating process

Males frequently visit host plants to find females. Once found, males spend a long time hovering above the female, and
courtship Courtship is the period wherein some couples get to know each other prior to a possible marriage. Courtship traditionally may begin after a betrothal and may conclude with the celebration of marriage. A courtship may be an informal and private m ...
occurs as the male fans the female with
pheromone A pheromone () is a secreted or excreted chemical factor that triggers a social response in members of the same species. Pheromones are chemicals capable of acting like hormones outside the body of the secreting individual, to affect the behavio ...
s from above. The males’ iridescent hind wings are also believed to be involved in attracting females. Males may also use
sodium Sodium is a chemical element with the symbol Na (from Latin ''natrium'') and atomic number 11. It is a soft, silvery-white, highly reactive metal. Sodium is an alkali metal, being in group 1 of the periodic table. Its only stable ...
taken up from mud as a nuptial gift during mating.


Oviposition

Females identify host plants on which to lay their eggs by leaf shape, which can lead to mistakes in egg laying and thus compromising the eggs' survival rates. Aristolochic acid presence may also act as an oviposition chemical stimulant.


Natural enemies

The pipevine swallowtail larva has few natural
predator Predation is a biological interaction where one organism, the predator, kills and eats another organism, its prey. It is one of a family of common feeding behaviours that includes parasitism and micropredation (which usually do not kill th ...
s but there have been observed cases of other butterfly larvae feeding on swallowtail larva.
Parasites Parasitism is a close relationship between species, where one organism, the parasite, lives on or inside another organism, the host, causing it some harm, and is adapted structurally to this way of life. The entomologist E. O. Wilson ha ...
can also threaten larvae, with certain fly and
wasp A wasp is any insect of the narrow-waisted suborder Apocrita of the order Hymenoptera which is neither a bee nor an ant; this excludes the broad-waisted sawflies (Symphyta), which look somewhat like wasps, but are in a separate suborder. ...
species being the most dangerous.
Bird Birds are a group of warm-blooded vertebrates constituting the class Aves (), characterised by feathers, toothless beaked jaws, the laying of hard-shelled eggs, a high metabolic rate, a four-chambered heart, and a strong yet lightweig ...
s are the greatest threat to larvae, as many species will eat them whenever they can. As a result of bird predation, this butterfly has evolved a chemical defense using the aristolochic acids found in their host plants.


Defense

All host plants for the pipevine swallowtail have some form of aristolochic acids, which the larva sequesters while they feed. These acids are passed on to future eggs, pupa, and adult butterflies, so over
generation A generation refers to all of the people born and living at about the same time, regarded collectively. It can also be described as, "the average period, generally considered to be about 20–⁠30 years, during which children are born and gr ...
s, the level of acid in the body of the butterflies rises as the acid accumulates. High levels of acid make the larvae and adults unpalatable to bird predation. The bright orange spots present on larvae and adult butterflies are thought to serve as warnings to predators, alerting them to the potential bad taste, should the predator decide to make a meal out of the swallowtail.


Mimicry

As a result of the pipevine swallowtails' natural defense through acid sequestering, many other species of butterflies, like the red-spotted purple butterfly, female eastern tiger swallowtail and eastern black swallowtail butterflies, promothea silkmoth males and the spicebush swallowtail use the pipevine swallowtail as a template for mimicry. Several other swallowtail subspecies have similar coloring to the pipevine swallowtail so that predators may also associate them with distaste. This is called Batesian mimicry. Pipevine swallowtails may also be involved with
Müllerian mimicry 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 Mülleria ...
, in which two distasteful species resemble each other, and thus act as a mimic and model; certain millipede species look like pipevine larvae and release hydrogen cyanide when in danger. File:Pipevine Swallowtail - Battus philenor, Great Smoky Mountains National Park, North Carolina.jpg, Pipevine swallowtail UP (model) File:Red Spotted Purple (2455520620).jpg, Red spotted purple butterfly UP (mimic) File:Eastern Tiger Swallowtail (Papilio glaucus) (6016411244).jpg, Female eastern tiger swallowtail UP (mimic) File:Pipevine Swallowtail (2661168441).jpg, Pipevine swallowtail UN (model) File:Spicebush Swallowtail (20304827146).jpg, Spicebush swallowtail UN (mimic)


References


Further reading

*Edwin Möhn, 1999 ''Schmetterlinge der Erde'', Butterflies of the World Part VIII (8), Papilionidae V. ''New and rare Neotropical Papilionidae''. Edited by Erich Bauer and Thomas Frankenbach Keltern : Goecke & Evers ; Canterbury : Hillside Books. , plate 2, f. 3-4, pl. 16, f. 1-2


External links


Pipevine Swallowtail, Butterflies of North Carolina
Cirrus Digital Imaging
Pipevine Swallowtail, Butterflies and Moths of North America
at Animal Diversity Web
Symbiosis: The Pipevine Swallowtail and the Dutchman's Pipevine
by David W. Klemp {{Taxonbar, from=Q1758032 philenor Butterflies of North America Butterflies of Central America Fauna of the San Francisco Bay Area Butterflies described in 1771 Taxa named by Carl Linnaeus