HOME

TheInfoList



OR:

Swallowtail butterflies are large, colorful
butterflies Butterflies are insects in the macrolepidopteran clade Rhopalocera from the Order (biology), order Lepidoptera, which also includes moths. Adult butterflies have large, often brightly coloured wings, and conspicuous, fluttering flight. The ...
in the family Papilionidae, and include over 550
species In biology, a species is the basic unit of classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of the appropriate s ...
. Though the majority are
tropical The tropics are the regions of Earth surrounding the Equator. They are defined in latitude by the Tropic of Cancer in the Northern Hemisphere at N and the Tropic of Capricorn in the Southern Hemisphere at S. The tropics are also referred to ...
, members of the family inhabit every
continent A continent is any of several large landmasses. Generally identified by convention rather than any strict criteria, up to seven geographical regions are commonly regarded as continents. Ordered from largest in area to smallest, these seven ...
except
Antarctica Antarctica () is Earth's southernmost and least-populated continent. Situated almost entirely south of the Antarctic Circle and surrounded by the Southern Ocean, it contains the geographic South Pole. Antarctica is the fifth-largest contine ...
. The family includes the largest butterflies in the world, the birdwing butterflies of the genus ''
Ornithoptera ''Ornithoptera'' is a genus of birdwing butterflies found in the northern portion of the Australasian realm, east of Weber's line; the Moluccas, New Guinea, the Solomon Islands, and northeastern Australia. Except for ''Ornithoptera richmondia'' ...
''. Swallowtails have a number of distinctive features; for example, the papilionid
caterpillar Caterpillars ( ) are the larval stage of members of the order Lepidoptera (the insect order comprising butterflies and moths). As with most common names, the application of the word is arbitrary, since the larvae of sawflies (suborder Sym ...
bears a repugnatorial organ called the
osmeterium The osmeterium is a defensive organ found in all papilionid larvae, in all stages. The organ is situated in the prothoracic segment and can be everted when the larva feels threatened. The everted organ resembles a fleshy forked tongue (not unlik ...
on its prothorax. The osmeterium normally remains hidden, but when threatened, the larva turns it outward through a transverse dorsal groove by inflating it with fluid. The forked appearance in some of the swallowtails' hindwings, which can be seen when the butterfly is resting with its wings spread, gave rise to the common name ''swallowtail''. As for its formal name,
Linnaeus Carl Linnaeus (; 23 May 1707 – 10 January 1778), also known after his ennoblement in 1761 as Carl von Linné Blunt (2004), p. 171. (), was a Swedish botanist, zoologist, taxonomist, and physician who formalised binomial nomenclature, the ...
chose ''
Papilio ''Papilio'' is a genus in the swallowtail butterfly family, Papilionidae, as well as the only representative of the tribe Papilionini. The word ''papilio'' is Latin for butterfly. It includes the common yellow swallowtail (''Papilio machaon''), ...
'' for the
type genus In biological taxonomy, the type genus is the genus which defines a biological family and the root of the family name. Zoological nomenclature According to the International Code of Zoological Nomenclature, "The name-bearing type of a nominal f ...
, as ''papilio'' is Latin for "butterfly". For the specific epithets of the genus, Linnaeus applied the names of Greek figures to the swallowtails. The
type species In zoological nomenclature, a type species (''species typica'') is the species name with which the name of a genus or subgenus is considered to be permanently taxonomically associated, i.e., the species that contains the biological type specimen ...
: ''
Papilio machaon ''Papilio machaon'', the Old World swallowtail, is a butterfly of the family Papilionidae. The butterfly is also known as the common yellow swallowtail or simply the swallowtail (a common name applied to all members of the family, but this speci ...
'' honored Machaon, one of the sons of
Asclepius Asclepius (; grc-gre, Ἀσκληπιός ''Asklēpiós'' ; la, Aesculapius) is a hero and god of medicine in ancient Religion in ancient Greece, Greek religion and Greek mythology, mythology. He is the son of Apollo and Coronis (lover of ...
, mentioned in the ''
Iliad The ''Iliad'' (; grc, Ἰλιάς, Iliás, ; "a poem about Ilium") is one of two major ancient Greek epic poems attributed to Homer. It is one of the oldest extant works of literature still widely read by modern audiences. As with the ''Odysse ...
''. Further, the species ''
Papilio homerus ''Papilio homerus'', commonly known as the Homerus swallowtail or Jamaican swallowtail, is the largest butterfly species in the Western Hemisphere. The species is endangered and faces a potentially bleak future. Only two small populations of the ...
'' is named after the Greek poet,
Homer Homer (; grc, Ὅμηρος , ''Hómēros'') (born ) was a Greek poet who is credited as the author of the ''Iliad'' and the ''Odyssey'', two epic poems that are foundational works of ancient Greek literature. Homer is considered one of the ...
. The Mon of the
Taira clan The Taira was one of the four most important clans that dominated Japanese politics during the Heian, Kamakura and Muromachi Periods of Japanese history – the others being the Fujiwara, the Tachibana, and the Minamoto. The clan is divided ...
of
Japan Japan ( ja, 日本, or , and formally , ''Nihonkoku'') is an island country in East Asia. It is situated in the northwest Pacific Ocean, and is bordered on the west by the Sea of Japan, while extending from the Sea of Okhotsk in the north ...
is an Agehachō (swallowtail butterfly).


Taxonomy


Subfamilies

The genera of extant swallowtails are usually classified into three subfamilies, Baroniinae,
Parnassiinae The Parnassiinae or snow Apollos are a subfamily of the swallowtail butterfly family, Papilionidae. The subfamily includes about 50 medium-sized, white or yellow species. The snow Apollos are high-altitude butterflies and are distributed across ...
, and
Papilioninae Papilioninae is a subfamily of the butterfly family Papilionidae. Papilioninae are found worldwide, but most species are distributed in the tropics. There are roughly 480 species, of which 27 occur in North America. Tribes This subfamily consis ...
, the latter two being further divided into
tribes The term tribe is used in many different contexts to refer to a category of human social group. The predominant worldwide usage of the term in English is in the discipline of anthropology. This definition is contested, in part due to conflic ...
. In swallowtails, besides morphological characteristics, the choice of food plants and ecological lifestyle reflect phylogeny and classification.


Baroniinae

The Baroniinae are a
monotypic In biology, a monotypic taxon is a taxonomic group (taxon) that contains only one immediately subordinate taxon. A monotypic species is one that does not include subspecies or smaller, infraspecific taxa. In the case of genera, the term "unispec ...
subfamily, restricted to a very small region in Mexico and are considered to be the most basal of the subfamilies. ''
Baronia brevicornis ''Baronia brevicornis'', commonly known as the short-horned baronia, is a species of butterfly in the monotypic genus ''Baronia'' and is placed in a subfamily of its own, the Baroniinae, a sister group of the remainder of the swallowtail butterf ...
'' is considered to be a
relict A relict is a surviving remnant of a natural phenomenon. Biology A relict (or relic) is an organism that at an earlier time was abundant in a large area but now occurs at only one or a few small areas. Geology and geomorphology In geology, a r ...
species, and shares features with a fossil taxon ''
Praepapilio ''Praepapilio'' is an extinct genus of swallowtail butterfly from the middle Eocene deposits of Colorado, United States, comparable to the Lutetian epoch in age. The genus is considered to be the only representative of the fossil subfamily Prae ...
''. ''Baronia'' is unique among papilionids as having an ''
Acacia ''Acacia'', commonly known as the wattles or acacias, is a large genus of shrubs and trees in the subfamily Mimosoideae of the pea family Fabaceae. Initially, it comprised a group of plant species native to Africa and Australasia. The genus na ...
'' species (family Leguminosae) as its food plant. Subfamily: Baroniinae. Baronia brevicornis.jpg,


Parnassiinae

The Parnassiinae are a subfamily of essentially
Holarctic The Holarctic realm is a biogeographic realm that comprises the majority of habitats found throughout the continents in the Northern Hemisphere. It corresponds to the floristic Boreal Kingdom. It includes both the Nearctic zoogeographical reg ...
butterflies. The vast majority of species, mostly ''
Parnassius ''Parnassius'' is a genus of northern circumpolar and montane (alpine and Himalayan) butterflies usually known as Apollos or snow Apollos. They can vary in colour and form significantly based on their altitude. They also show an adaptation to h ...
'', can be found in mountain habitats. Parnassiinines can also be found in other habitats such as "arid deserts (''Hypermnestra''), humid forests (''Luehdorfia'') and even lowland meadows (''Zerynthia'')". The tribes recognized in the Parnassiinae are Parnassiini,
Zerynthiini The Zerynthiini are a tribe of swallowtail butterflies. Genera The tribe is thought to consist of four genera: * '' Allancastria'' * '' Bhutanitis'' * ''Sericinus ''Sericinus'' is a genus of swallowtail butterflies placed in the subfamily ...
, and
Luehdorfiini The Luehdorfiini are a tribe of swallowtail butterflies. Genera The tribe is thought to consist of three genera: * '' Archon'' * '' Doritites'' * ''Luehdorfia ''Luehdorfia'' is a genus of butterflies in the family Papilionidae Swallo ...
. Tribe Parnassiini contains two genera, ''
Hypermnestra Hypermnestra ( grc, Ὑπερμνήστρα, ''Hypermnēstra''), in Greek mythology, was a Libyan princess as one of the 50 Danaids the daughter of King Danaus, son of King Belus of Egypt. Her mother was Elephantis and full sister to Gorgophone ...
'', largely confined to
central Asia Central Asia, also known as Middle Asia, is a subregion, region of Asia that stretches from the Caspian Sea in the west to western China and Mongolia in the east, and from Afghanistan and Iran in the south to Russia in the north. It includes t ...
and the genus ''
Parnassius ''Parnassius'' is a genus of northern circumpolar and montane (alpine and Himalayan) butterflies usually known as Apollos or snow Apollos. They can vary in colour and form significantly based on their altitude. They also show an adaptation to h ...
'' (the Apollos), a distinctive group of many species, all of which are alpine and capable of living at high altitudes. Most ''Parnassius'' have two small reddish spots on their
hindwings 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 hindw ...
. The tribe Luehdorfiini contains the genera ''
Archon ''Archon'' ( gr, ἄρχων, árchōn, plural: ἄρχοντες, ''árchontes'') is a Greek word that means "ruler", frequently used as the title of a specific public office. It is the masculine present participle of the verb stem αρχ-, mean ...
'' of
Asia minor Anatolia, tr, Anadolu Yarımadası), and the Anatolian plateau, also known as Asia Minor, is a large peninsula in Western Asia and the westernmost protrusion of the Asian continent. It constitutes the major part of modern-day Turkey. The re ...
and the genus ''
Luehdorfia ''Luehdorfia'' is a genus of butterflies in the family Papilionidae Swallowtail butterflies are large, colorful butterflies in the family Papilionidae, and include over 550 species. Though the majority are tropical, members of the family ...
'' of China and Japan. These two tribes have evolved to change their food plants, while the third tribe, Zerynthiini, has retained the archetypical papilionid food plant, the lowland vine '' Aristolochia''. Zerynthiini comprises four genera – ''
Sericinus ''Sericinus'' is a genus of swallowtail butterflies placed in the subfamily Parnassiinae. The genus has a complex history and a multiplicity of names have been applied to its single species. ''Sericinus montela'', the dragon swallowtail, is the ...
'', '' Bhutanitis'', ''
Zerynthia ''Zerynthia'' is a genus of swallowtail butterflies placed in the subfamily Parnassiinae. The genus has a complex history; a multiplicity of names have been applied to its species. Species ''Zerynthia'' consists of the following species: ...
'' and ''
Allancastria ''Allancastria'' is a genus of Palaearctic swallowtail butterflies in the subfamily Parnassiinae The Parnassiinae or snow Apollos are a subfamily of the swallowtail butterfly family, Papilionidae. The subfamily includes about 50 medium-size ...
''. Subfamily: Parnassiinae. Apollo butterfly.JPG, Archon_apollinus_bellargus_1.jpg, DianeJLH.jpg, BhutliddCRW_4351.jpg,


Papilioninae

The tribes recognized in the Papilioninae are
Leptocircini Leptocircini is a tribe of swallowtail butterflies that includes the genera ''Eurytides'' (kite swallowtails), '' Graphium'' (swordtails), and '' Lamproptera'' (dragontails). Taxonomy The tribe consists of roughly 140 species in nine genera wor ...
, Teinopalpini,
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 ...
, and Papilionini. Subfamily: Papilioninae. Zebra Swallowtail Megan McCarty10.jpg, Teinopalpus imperialis verso.JPG, Trogonoptera trojana male 5.jpg, Papilio troilus01.jpg, Swallowtail butterfly.jpg, Cairns birdwing - melbourne zoo.jpg, Western Tiger Swallowtail (Papilio Rutulus).jpg, Voutyro 2017 822 zoom.jpg,


Praepapilioninae

An additional subfamily, Praepapilioninae, consisting of a single genus ''
Praepapilio ''Praepapilio'' is an extinct genus of swallowtail butterfly from the middle Eocene deposits of Colorado, United States, comparable to the Lutetian epoch in age. The genus is considered to be the only representative of the fossil subfamily Prae ...
'', includes two species of extinct butterflies, each member being described from single
fossil A fossil (from Classical Latin , ) is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserved ...
s found in a middle Eocene deposit in
Colorado Colorado (, other variants) is a state in the Mountain West subregion of the Western United States. It encompasses most of the Southern Rocky Mountains, as well as the northeastern portion of the Colorado Plateau and the western edge of t ...
, United States (Durden and Rose, 1978).


Phylogeny

A
phylogeny A phylogenetic tree (also phylogeny or evolutionary tree Felsenstein J. (2004). ''Inferring Phylogenies'' Sinauer Associates: Sunderland, MA.) is a branching diagram or a tree showing the evolutionary relationships among various biological spe ...
of the Papilionidae based on Nazari (2007) is given: It is now accepted that the subfamily Papilioninae is
monophyletic In cladistics for a group of organisms, monophyly is the condition of being a clade—that is, a group of taxa composed only of a common ancestor (or more precisely an ancestral population) and all of its lineal descendants. Monophyletic gro ...
. The swallowtail butterflies in the nominate tribe Papilionini number about 225 species and studies have been made on their host plant
coevolution In biology, coevolution occurs when two or more species reciprocally affect each other's evolution through the process of natural selection. The term sometimes is used for two traits in the same species affecting each other's evolution, as well ...
and
phylogeny A phylogenetic tree (also phylogeny or evolutionary tree Felsenstein J. (2004). ''Inferring Phylogenies'' Sinauer Associates: Sunderland, MA.) is a branching diagram or a tree showing the evolutionary relationships among various biological spe ...
. Old morphological classifications were also found to be valid in that they formed clusters. Species belonging to the groups that use
Rutaceae The Rutaceae is a family, commonly known as the rueRUTACEAE
in BoDD – Botanical Derm ...
as host plants formed two groups corresponding to
Old World The "Old World" is a term for Afro-Eurasia that originated in Europe , after Europeans became aware of the existence of the Americas. It is used to contrast the continents of Africa, Europe, and Asia, which were previously thought of by the ...
and
American American(s) may refer to: * American, something of, from, or related to the United States of America, commonly known as the "United States" or "America" ** Americans, citizens and nationals of the United States of America ** American ancestry, pe ...
taxa. Those that fed on
Lauraceae Lauraceae, or the laurels, is a plant family that includes the true laurel and its closest relatives. This family comprises about 2850 known species in about 45 genera worldwide (Christenhusz & Byng 2016 ). They are dicotyledons, and occur ma ...
and
Magnoliaceae The Magnoliaceae () are a flowering plant family, the magnolia family, in the order Magnoliales. It consists of two genera: ''Magnolia'' and '' Liriodendron'' (tulip trees). Unlike most angiosperms, whose flower parts are in whorls (rings), t ...
were found to form another cluster which includes both Asian and American taxa. The Parnassinae, like the Papilioninae, were also believed to be monophyletic based on morphological studies but recent studies based on both morphological and molecular characteristics suggest that this is not the case. Of the
Parnassiinae The Parnassiinae or snow Apollos are a subfamily of the swallowtail butterfly family, Papilionidae. The subfamily includes about 50 medium-sized, white or yellow species. The snow Apollos are high-altitude butterflies and are distributed across ...
, the genera ''
Parnassius ''Parnassius'' is a genus of northern circumpolar and montane (alpine and Himalayan) butterflies usually known as Apollos or snow Apollos. They can vary in colour and form significantly based on their altitude. They also show an adaptation to h ...
'' and ''
Hypermnestra Hypermnestra ( grc, Ὑπερμνήστρα, ''Hypermnēstra''), in Greek mythology, was a Libyan princess as one of the 50 Danaids the daughter of King Danaus, son of King Belus of Egypt. Her mother was Elephantis and full sister to Gorgophone ...
'' were found to be extremely close based on molecular studies and are now considered to be part of the tribe Parnassiini. The two taxa, ''
Archon ''Archon'' ( gr, ἄρχων, árchōn, plural: ἄρχοντες, ''árchontes'') is a Greek word that means "ruler", frequently used as the title of a specific public office. It is the masculine present participle of the verb stem αρχ-, mean ...
'' and ''
Luehdorfia ''Luehdorfia'' is a genus of butterflies in the family Papilionidae Swallowtail butterflies are large, colorful butterflies in the family Papilionidae, and include over 550 species. Though the majority are tropical, members of the family ...
'', have been found to be closely related through analysis of nuclear and mitochondrial DNA, and, though they share no morphological similarities, have now been united in the tribe Luehdorfiini. The subfamily Baroniinae is represented by the sole representative species ''
Baronia brevicornis ''Baronia brevicornis'', commonly known as the short-horned baronia, is a species of butterfly in the monotypic genus ''Baronia'' and is placed in a subfamily of its own, the Baroniinae, a sister group of the remainder of the swallowtail butterf ...
''. They are unique in the family to use the
Fabaceae The Fabaceae or Leguminosae,International Code of Nomenc ...
(Leguminosae) as their larval host plants. The Baronninae and the extinct subfamily Praepapilioninae share many external similarities and are traditionally considered to be the most primitive subfamilies and sister to the rest of the swallowtails. Recent research suggests that this may not be the case, the Baroniinae being closely related to only the Parnassiinae, and ''Praepapilio'' to only the Papilionini and neither taxa being sister to the rest of the swallowtails.


Distribution

As of 2005, 552 extant species have been identified which are distributed across the tropical and temperate regions. Various species inhabit altitudes ranging from sea level to high mountains, as in the case of most species of ''
Parnassius ''Parnassius'' is a genus of northern circumpolar and montane (alpine and Himalayan) butterflies usually known as Apollos or snow Apollos. They can vary in colour and form significantly based on their altitude. They also show an adaptation to h ...
''. The majority of swallowtail species and the greatest diversity are found in the tropics and subtropical regions between 20°N and 20°S, particularly Southeast Asia, and between 20°N and 40°N in East Asia. Only 12 species are found in Europe and only one species, ''
Papilio machaon ''Papilio machaon'', the Old World swallowtail, is a butterfly of the family Papilionidae. The butterfly is also known as the common yellow swallowtail or simply the swallowtail (a common name applied to all members of the family, but this speci ...
'' is found in the
British Isles The British Isles are a group of islands in the North Atlantic Ocean off the north-western coast of continental Europe, consisting of the islands of Great Britain, Ireland, the Isle of Man, the Inner and Outer Hebrides, the Northern Isles, ...
. North America has 40 species, including several tropical species and ''Parnassius''. The northernmost swallowtail is the Siberian Apollo (''
Parnassius arcticus ''Parnassius arcticus'', the Siberian Apollo, is a high-altitude butterfly which is found in Northeastern Yakutia, Russia. It is a member of the snow Apollo genus ('' Parnassius'') of the swallowtail family, Papilionidae. The taxonomic status of ...
''), found in the Arctic Circle in northeastern
Yakutia Sakha, officially the Republic of Sakha (Yakutia),, is the largest republic of Russia, located in the Russian Far East, along the Arctic Ocean, with a population of roughly 1 million. Sakha comprises half of the area of its governing Far Eas ...
, at altitudes of 1500 meters above sea level. In the
Himalayas The Himalayas, or Himalaya (; ; ), is a mountain range in Asia, separating the plains of the Indian subcontinent from the Tibetan Plateau. The range has some of the planet's highest peaks, including the very highest, Mount Everest. Over 100 ...
, various Apollo species such as '' Parnassius epaphus'', have been found at altitudes of 6,000 meters above sea level.


Food

The caterpillars of various swallowtail butterfly species feed on a wide range of different plants, most depending on only one of five families:
Aristolochiaceae The Aristolochiaceae () are a family, the birthwort family, of flowering plants with seven genera and about 400 known species belonging to the order Piperales. The type genus is ''Aristolochia'' L. Description They are mostly perennial, he ...
,
Annonaceae The Annonaceae are a Family (biology), family of flowering plants consisting of trees, shrubs, or rarely lianas commonly known as the custard apple family or soursop family. With 108 accepted genera and about 2400 known species, it is the largest ...
,
Lauraceae Lauraceae, or the laurels, is a plant family that includes the true laurel and its closest relatives. This family comprises about 2850 known species in about 45 genera worldwide (Christenhusz & Byng 2016 ). They are dicotyledons, and occur ma ...
, Umbelliferae (
Apiaceae Apiaceae or Umbelliferae is a family of mostly aromatic flowering plants named after the type genus ''Apium'' and commonly known as the celery, carrot or parsley family, or simply as umbellifers. It is the 16th-largest family of flowering plants ...
) and
Rutaceae The Rutaceae is a family, commonly known as the rueRUTACEAE
in BoDD – Botanical Derm ...
. By eating some of these toxic plants, the caterpillars sequester aristolochic acid which renders both the caterpillars and the butterflies of some of these as toxic, thus protecting them from predators. Similarly, the ''
Parnassius smintheus ''Parnassius smintheus'', the Rocky Mountain parnassianJim P. Brock and K. Kaufman. Kaufman Field Guide to Butterflies of North America, New York, NY:Houghton Mifflin, 2003. or Rocky Mountain apollo, is a high-altitude butterfly found in the Rock ...
'' sequesters sarmentosin from its host plant ''
Sedum lanceolatum ''Sedum lanceolatum'' is a species of flowering plant in the family Crassulaceae known by the common names spearleaf stonecrop and lanceleaf stonecrop. It is native to western North America and occurs in western Canada and the United States. It ...
'' for protection from predators. Swallowtail tribes Zerynthiini (Parnassiinae), Luehdorfiini (Parnassiinae) and Troidini (Papilioninae), almost exclusively use the family
Aristolochiaceae The Aristolochiaceae () are a family, the birthwort family, of flowering plants with seven genera and about 400 known species belonging to the order Piperales. The type genus is ''Aristolochia'' L. Description They are mostly perennial, he ...
as their host plants. For example, the eastern black swallowtail's main host plant in the wild is
Queen Anne's lace Queen Anne's lace is a common name for a number of plants in the family Apiaceae. including: * ''Ammi majus'', native in the Nile River Valley * '' Anthriscus sylvestris'', a herbaceous biennial or short-lived perennial plant * '' Daucus carota'', ...
, but they also eat garden plants in the carrot family, including carrots, parsley, dill, and fennel. Adult swallowtails sip nectar, but also mud and sometimes manure.


Life cycle

The detailed descriptions of morphological characteristics of the Papilionidae, as quoted in Bingham (1905) are as follows: Stages of development of a papilionid, the
giant swallowtail The giant swallowtail (''Papilio cresphontes'') is the largest butterfly in North America. It is abundant through many parts of eastern North America; populations from western North America and down into Panama are now (as of 2014) considered t ...
(''Papilio cresphontes'') Giant Swallowtail Egg.jpg, Egg Giant Swallowtail Larva.jpg, Larva Papilio cresphontes chrysalis.JPG, Pupa Butterfly 8731-2.jpg, Imago


Distinguishing characteristics

The key characteristics that differentiate the Papilionidae from the other butterfly families are: * The
osmeterium The osmeterium is a defensive organ found in all papilionid larvae, in all stages. The organ is situated in the prothoracic segment and can be everted when the larva feels threatened. The everted organ resembles a fleshy forked tongue (not unlik ...
is a forked, fleshy eversible organ found in the prothoracic segment of caterpillars. *
Venation Venation may refer to: * Venation (botany), the arrangement of veins in leaves * Wing venation, the arrangement of veins in insect wings See also * * Vernation Vernation (from ''vernal'' meaning ''spring'', since that is when leaves spring fort ...
– in swallowtails, the second anal vein, 2A, extends up to the wing margin and does not link with the first anal vein, 1A. These veins are fused in other butterfly families and 2A does not reach the wing margin. * The
sclerite A sclerite (Greek , ', meaning "hard") is a hardened body part. In various branches of biology the term is applied to various structures, but not as a rule to vertebrate anatomical features such as bones and teeth. Instead it refers most commonly ...
s of the cervix (membranous neck between the head and thorax) are fused beneath the neck where the muscles for head movement are anchored. 01 chenille machaon (detail).jpg,
Old World swallowtail ''Papilio machaon'', the Old World swallowtail, is a butterfly of the family Papilionidae. The butterfly is also known as the common yellow swallowtail or simply the swallowtail (a common name applied to all members of the family, but this speci ...
caterpillar everting its
osmeterium The osmeterium is a defensive organ found in all papilionid larvae, in all stages. The organ is situated in the prothoracic segment and can be everted when the larva feels threatened. The everted organ resembles a fleshy forked tongue (not unlik ...
in defense. Papilio demodocus larva IMG_8543s.jpg, ''
Papilio demodocus ''Papilio demodocus'', the citrus swallowtail or Christmas butterfly, is a swallowtail butterfly which commonly occurs over the entirety of sub-Saharan Africa, including Madagascar, besides the southern Arabian Peninsula. The caterpillars feed ...
'' larva applying one horn of the evaginated osmeterium to an offending finger. Papilionidae id char venation.png, Adult forewing of ''
Graphium agetes ''Graphium agetes'', the fourbar swordtail, is a widespread species of swallowtail butterfly found in tropical Asia. It is common and not threatened. Description Males and females upperside white. Forewing: cell crossed by three comparatively ...
''. Second anal vein, 2A, extends up to the wing margin and does not link with the first anal vein, 1A. Black Swallowtail Chrysalis Megan McCarty33.jpg, Papilionid chrysalis are typically attached to a substrate by the cremaster and with head up held by a
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 coc ...
girdle.


Special adaptations and defense

Swallowtail butterflies practice
Batesian mimicry 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 bu ...
, a
behavior Behavior (American English) or behaviour (British English) is the range of actions and mannerisms made by individuals, organisms, systems or artificial entities in some environment. These systems can include other systems or organisms as wel ...
in which the butterflies' appearance closely resemble that of distasteful species that prevents predation. Swallowtails differ from many animals that practice
mimicry 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 f ...
. The tiger swallowtail butterfly (''
Papilio glaucus ''Papilio glaucus'', the eastern tiger swallowtail, is a species of butterfly native to eastern North America. It is one of the most familiar butterflies in the eastern United States, ranging north to southern Ontario, Canada, and is common in m ...
''), exhibits a female-limited polymorphism for Batesian mimicry and others, such as the Canadian tiger swallowtail (''
Papilio canadensis ''Papilio canadensis'', the Canadian tiger swallowtail, is a species of butterfly in the family Papilionidae. It was once classified as a subspecies of ''Papilio glaucus''. Description Adult The wingspan of this species is 67 to 80 mm. It ...
'') do not display any form of mimicry. Predators include the
red-winged blackbird The red-winged blackbird (''Agelaius phoeniceus'') is a passerine bird of the family Icteridae found in most of North America and much of Central America. It breeds from Alaska and Newfoundland south to Florida, the Gulf of Mexico, Mexico, and G ...
,
Pennsylvania firefly ''Photuris pensylvanica'', known by the common names Pennsylvania firefly, lightning bug, dot-dash firefly and (in its larval state) glowworm, is a species of firefly from the United States and Canada. It is also widely known under the Latin nam ...
, five-lined skink,
green darner The green darner or common green darner (''Anax junius''), after its resemblance to a darning needle, is a species of dragonfly in the family Aeshnidae. One of the most common and abundant species throughout North America, it also ranges south ...
,
goldenrod spider Goldenrod is a common name for many species of flowering plants in the sunflower family, Asteraceae, commonly in reference to the genus ''Solidago''. Several genera, such as ''Euthamia'', were formerly included in a broader concept of the genus ...
,
Chinese mantis The Chinese mantis (''Tenodera sinensis'') is a species of mantis native to Asia and the nearby islands. In 1896 this species was accidentally introduced by a nursery tender at Mt. Airy near Philadelphia, United States. ''Tenodera sinensis'' oft ...
,
fiery searcher ''Calosoma scrutator'', also known as the fiery searcher and caterpillar hunter, is a species of ground beetle belonging to the genus ''Calosoma'', subgenus '' Acalosoma''. This beetle can be as large as long. The distribution of this species is ...
, and striped skunk.


Biological basis for polymorphisms in mimicry

Not all individuals in some species are identical in appearance. For example, ''
Papilio glaucus ''Papilio glaucus'', the eastern tiger swallowtail, is a species of butterfly native to eastern North America. It is one of the most familiar butterflies in the eastern United States, ranging north to southern Ontario, Canada, and is common in m ...
'' (eastern tiger swallowtail), Y-linkage determines whether the females are either
wild-type The wild type (WT) is the phenotype of the typical form of a species as it occurs in nature. Originally, the wild type was conceptualized as a product of the standard "normal" allele at a locus, in contrast to that produced by a non-standard, "m ...
(yellow and black) or melanic (dark
melanin Melanin (; from el, μέλας, melas, black, dark) is a broad term for a group of natural pigments found in most organisms. Eumelanin is produced through a multistage chemical process known as melanogenesis, where the oxidation of the amino ...
replaces the yellow background). This genetic difference stems from the fact that
melanism The term melanism refers to black pigment and is derived from the gr, μελανός. Melanism is the increased development of the dark-colored pigment melanin in the skin or hair. Pseudomelanism, also called abundism, is another variant of ...
is controlled by a single
gene In biology, the word gene (from , ; "...Wilhelm Johannsen coined the word gene to describe the Mendelian units of heredity..." meaning ''generation'' or ''birth'' or ''gender'') can have several different meanings. The Mendelian gene is a ba ...
, which controls the level of dopamine in the organism. The
enzyme Enzymes () are proteins that act as biological catalysts by accelerating chemical reactions. The molecules upon which enzymes may act are called substrates, and the enzyme converts the substrates into different molecules known as products. A ...
BAS, which assists dopamine in producing the yellow
pigmentation A pigment is a colored material that is completely or nearly insoluble in water. In contrast, dyes are typically soluble, at least at some stage in their use. Generally dyes are often organic compounds whereas pigments are often inorganic compoun ...
, normally found on the wings' background, is suppressed. Without the pigmentation, the butterfly appears mostly black (the melanic form) and is a Batesian mimic of ''
Battus philenor ''Battus philenor'', the pipevine swallowtail or blue swallowtail, Retrieved April 19, 2018. is a swallowtail butterfly found in North America and Central America. This butterfly is black with iridescent-blue hindwings. They are found in many d ...
'', the pipevine swallowtail. There are also ''Papilio glaucus'' that are not wholly black; several possess an intermediate "sooty" color and are sensitive to
temperature Temperature is a physical quantity that expresses quantitatively the perceptions of hotness and coldness. Temperature is measured with a thermometer. Thermometers are calibrated in various temperature scales that historically have relied o ...
. The different polymorphisms (wild-type, melanic, and the 'sooty' intermediate) depend upon the geographical distribution and abundance of its mimic, the ''Battus philenor'', whose wing color varies depending on its geographical location. In order to be successfully confused for the ''B. philenor'' by predators, the ''Papilio glaucus's'' background wing color matches that of the ''B. philenor'' residing in the same regional area. Studies support this theory; in the southeastern United States, the relative abundance of melanic females has been found to geographically correlate with ''B. philenor''.


Mimicry

Only certain subsets of swallowtails practice
mimicry 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 f ...
. Species differ in whether one or both sexes is mimetic, and whether the mimicry is monomorphic or polymorphic. A phenomenon which has received particular attention is female-limited polymorphism, in which only the females of a species are mimetic and polymorphic, often mimicking different, distantly-related
aposematic Aposematism is the advertising by an animal to potential predators that it is not worth attacking or eating. This unprofitability may consist of any defences which make the prey difficult to kill and eat, such as toxicity, venom, foul taste o ...
butterflies. This polymorphism is seen in ''
Papilio dardanus ''Papilio dardanus'', the African swallowtail, mocker swallowtail or flying handkerchief, is a species of butterfly in the family Papilionidae (the swallowtails). The species is broadly distributed throughout Sub-Saharan Africa. The British e ...
'', the African swallowtail butterfly, whose females have three different morphs for
wing A wing is a type of fin that produces lift while moving through air or some other fluid. Accordingly, wings have streamlined cross-sections that are subject to aerodynamic forces and act as airfoils. A wing's aerodynamic efficiency is expres ...
color pattern: a black-and-white pattern for
Batesian mimicry 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 bu ...
, a black-and-yellow pattern that resembles the males of the species, and a pattern with orange patches that resembles the elderly males of the species. Given that the males of the species, which do not have Batesian mimicry, are preyed upon much more frequently by
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 than the females, an ongoing question is why females would exhibit the non-mimetic wing pattern, which would seemingly lower their fitness compared to the mimicry form. The
pipevine swallowtail ''Battus philenor'', the pipevine swallowtail or blue swallowtail, Retrieved April 19, 2018. is a swallowtail butterfly found in North America and Central America. This butterfly is black with iridescent-blue hindwings. They are found in many di ...
exhibits Batesian mimicry as well. Several hypotheses for this phenomenon were made, the two noteworthy being the pseudosexual selection hypothesis and the male avoidance hypothesis. In the pseudosexual hypothesis, male butterflies aggressively approached the ''male''-looking females and then mellowed their behavior into sexual behavior when they were close enough to identify them as females. In the male avoidance hypothesis, female butterflies disguise themselves in an attempt to evade male harassment, as courtship can be harmful, time-consuming, and attract predators. One study recorded male responses to females of each morphs and found that the males consistently favored the Batesian mimics, then the black and yellow, and then the morph with orange patches. The scientists concluded that
frequency-dependent selection Frequency-dependent selection is an evolutionary process by which the fitness of a phenotype or genotype depends on the phenotype or genotype composition of a given population. * In positive frequency-dependent selection, the fitness of a phenotyp ...
did lead to equal success for all three alternative strategies: the Batesian females suffered the fewest predators but their fitness was reduced the most by sexual harassment, while the other two faced lower sexual harassment but also lost fitness from predators' attacks.


Mating and young

After
mating In biology, mating is the pairing of either opposite-sex or hermaphroditic organisms for the purposes of sexual reproduction. ''Fertilization'' is the fusion of two gametes. ''Copulation'' is the union of the sex organs of two sexually reproduc ...
, the male ''Parnassines'' produce a glue-like substance that is used to seal the female genital opening and prevent other males from mating.English translation
They lay individual eggs on the underside of the leaves of their food plants. There is no parental investment once the eggs have been laid. The
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 typically attached to the substrate by the cremaster but with head up held by a
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 coc ...
girdle. The Apollos, however, pupate in debris on the ground and also build a loose cocoon. In the
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 t ...
regions, the winters are passed in a pupal
diapause In animal dormancy, diapause is the delay in development in response to regular and recurring periods of adverse environmental conditions.Tauber, M.J., Tauber, C.A., Masaki, S. (1986) ''Seasonal Adaptations of Insects''. Oxford University Press I ...
stage.


In culture

Since swallowtail butterflies are large, colorful, and attractive, they have been targeted by butterfly collectors. The largest of these, the birdwing butterflies are particularly sought after and are cultured in butterfly farms to supply collectors. Many members of the family, as
larvae A larva (; plural larvae ) is a distinct juvenile form many animals undergo before metamorphosis into adults. Animals with indirect development such as insects, amphibians, or cnidarians typically have a larval phase of their life cycle. The ...
, feed on plants of the citrus family
Rutaceae The Rutaceae is a family, commonly known as the rueRUTACEAE
in BoDD – Botanical Derm ...
, making some of these attractive butterflies
pests PESTS was an anonymous American activist group formed in 1986 to critique racism, tokenism, and exclusion in the art world. PESTS produced newsletters, posters, and other print material highlighting examples of discrimination in gallery represent ...
in
citrus ''Citrus'' is a genus of flowering plant, flowering trees and shrubs in the rue family, Rutaceae. Plants in the genus produce citrus fruits, including important crops such as Orange (fruit), oranges, Lemon, lemons, grapefruits, pomelos, and lim ...
orchards. The
Oregon swallowtail The Oregon swallowtail (''Papilio machaon oregonius'', sometimes classified as ''Papilio oregonius'' or ''Papilio bairdii oregonius'') is a subspecies of swallowtail butterfly native to the United States of America, Oregon, Washington, and Id ...
is the state insect of
Oregon Oregon () is a U.S. state, state in the Pacific Northwest region of the Western United States. The Columbia River delineates much of Oregon's northern boundary with Washington (state), Washington, while the Snake River delineates much of it ...
. The eastern tiger swallowtail is the state insect of
Virginia Virginia, officially the Commonwealth of Virginia, is a state in the Mid-Atlantic and Southeastern regions of the United States, between the Atlantic Coast and the Appalachian Mountains. The geography and climate of the Commonwealth ar ...
and the
state butterfly State insects are designated by 48 individual states of the fifty United States. Some states have more than one designated insect, or have multiple categories (e.g., state insect and state butterfly, etc.). Iowa and Michigan are the two states with ...
of
Georgia Georgia most commonly refers to: * Georgia (country), a country in the Caucasus region of Eurasia * Georgia (U.S. state), a state in the Southeast United States Georgia may also refer to: Places Historical states and entities * Related to the ...
,
Delaware Delaware ( ) is a state in the Mid-Atlantic region of the United States, bordering Maryland to its south and west; Pennsylvania to its north; and New Jersey and the Atlantic Ocean to its east. The state takes its name from the adjacent Del ...
, and
South Carolina )''Animis opibusque parati'' ( for, , Latin, Prepared in mind and resources, links=no) , anthem = " Carolina";" South Carolina On My Mind" , Former = Province of South Carolina , seat = Columbia , LargestCity = Charleston , LargestMetro = ...
. The
black swallowtail ''Papilio polyxenes'', the (eastern) black swallowtail, American swallowtail or parsnip swallowtail, is a butterfly found throughout much of North America. It is the state butterfly of Oklahoma and New Jersey New Jersey is a state in ...
is the state butterfly of
Oklahoma Oklahoma (; Choctaw language, Choctaw: ; chr, ᎣᎧᎳᎰᎹ, ''Okalahoma'' ) is a U.S. state, state in the South Central United States, South Central region of the United States, bordered by Texas on the south and west, Kansas on the nor ...
.


References


Further reading

* Chattopadhyay, J. (2007). ''Swallowtail Butterflies, Biology & Ecology of a few Indian Species.'' Desh Prashan, Kolkata, India. – 134 pp. . * Glassberg, J. (2001). ''Butterflies through Binoculars.'' The West. * Guppy, C.S. and Shepard, J.H. (2001). ''Butterflies of British Columbia.'' * Igarashi, S. (1979). ''
Papilionidae Swallowtail butterflies are large, colorful butterflies in the family Papilionidae, and include over 550 species. Though the majority are tropical, members of the family inhabit every continent except Antarctica. The family includes the larg ...
'' and their early stages
n Japanese N, or n, is the fourteenth letter in the Latin alphabet, used in the modern English alphabet, the alphabets of other western European languages and others worldwide. Its name in English is ''en'' (pronounced ), plural ''ens''. History ...
2 vols. – Tokyo, Kodansha, 218 pp., 357 pls. * James, D.G. & Nunnallee, D. (2011). Life Histories of Cascadia Butterflies. * Korolev V.A. (2014). Cataloges on the collection of ''
Lepidoptera Lepidoptera ( ) is an order (biology), order of insects that includes butterfly, butterflies and moths (both are called lepidopterans). About 180,000 species of the Lepidoptera are described, in 126 Family (biology), families and 46 Taxonomic r ...
''. Part II. ''
Papilionidae Swallowtail butterflies are large, colorful butterflies in the family Papilionidae, and include over 550 species. Though the majority are tropical, members of the family inhabit every continent except Antarctica. The family includes the larg ...
''. – Moscow, 387 p., 20 color tabs,

* Pelham, J. (2008). ''Catalogue of the Butterflies of the United States and Canada.'' * Pyle, R.M. (2002). ''The Butterflies of Cascadia.'' * Rothschild, L.W. (1895). A revision of the Papilios of the Eastern Hemisphere, exclusive of Africa. ''Novitates Zoologicae'' 2(3):167–463 and plate
pdf
*Rothschild, W. and Jordan, K. (1906). A revision of the American Papilios. ''Novitates Zoologicae'' 13: 411–752
online
(and as pdf) (Facsimile edition ed. P.H. Arnaud, 1967). * Seitz, A. (1907). 1. ''Gattung Papilio, Schwalbenschwänze.'' In: Seitz, A. (ed). ''Die Groß-Schmetterlinger der Erde. I. Abteilung (Die Großschmetterlinge des Palaeakrtischen Faunengebietes). 1. Band: Tagfalter.'' pp. 8–15. F. Lehmann, Stuttgart. * Talbot, G. (1939). ''The Fauna of British India. Butterflies. Volume 1.
Papilionidae Swallowtail butterflies are large, colorful butterflies in the family Papilionidae, and include over 550 species. Though the majority are tropical, members of the family inhabit every continent except Antarctica. The family includes the larg ...
and
Pieridae The Pieridae are a large family of butterflies with about 76 genera containing about 1,100 species, mostly from tropical Africa and tropical Asia with some varieties in the more northern regions of North America and Eurasia.DeVries P. J. in Le ...
.'' Taylor & Francis, London; xxix, 600 pp., 3 pls, 1 map. * Tuzov V., Bogdanov P., Devyatkin A., Kaabak L., Korolev V., Murzin V., Samodurov G., Tarassov E. (1997). ''Guide to the butterflies of Russia and adjacent territories (
Lepidoptera Lepidoptera ( ) is an order (biology), order of insects that includes butterfly, butterflies and moths (both are called lepidopterans). About 180,000 species of the Lepidoptera are described, in 126 Family (biology), families and 46 Taxonomic r ...
,
Rhopalocera Butterflies are insects in the macrolepidopteran clade Rhopalocera from the order Lepidoptera, which also includes moths. Adult butterflies have large, often brightly coloured wings, and conspicuous, fluttering flight. The group comprises ...
). Vol. 1.'' Sofia-Moscow: Pensoft-Press, 480 p., 79 col. plates. * Warren, A.D., Davis, K.J., Grishin, N.V., Pelham, J.P., Stangeland, E.M. (2012). Interactive Listing of American Butterflies


External links

* *
"Papilionidae" on ''Tree of Life '' web project.
*


GloBIS Database
Includes type images

* * ttp://www.arkinspace.com/2012/04/sublime-swallowtail.html Photoset and description of lesser known species of swallowtails
Swallowtails
Papilionidae of the World: a pictorial summary. Comprehensive. {{Authority control Papilionoidea Articles containing video clips