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Swallowtail butterfly

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Title: Swallowtail butterfly  
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Swallowtail butterfly

Swallowtail butterfly

Swallowtail butterflies are large, colorful [1]

Swallowtails have a number of distinctive features; for example, the papilionid osmeterium 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.[2]

The forked appearance of the swallowtails' hind wings, 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 chose Papilio for the type genus, as Papilio is Latin for 'butterfly'. For the specific epithets of the genus, Linnaeus applied the names of Greek heroes to the swallowtails. The type species: Papilio machaon honoured Machaon, one of the sons of Asclepius, mentioned in the Iliad.[3]


  • Distribution 1
  • Appearance/Morphology 2
    • Distinguishing characteristics 2.1
  • Special adaptations and defense 3
    • Biological basis for polymorphisms in mimicry 3.1
    • Mimicry 3.2
  • Taxonomy 4
    • Subfamilies 4.1
      • Baroniinae 4.1.1
      • Parnassinae 4.1.2
      • Papilioninae 4.1.3
      • Praepapilioninae 4.1.4
    • Phylogeny 4.2
  • Mating and young 5
  • Food 6
  • Swallowtails and humans 7
  • In popular culture 8
  • References 9
  • Further reading 10
  • External links 11


As of 2005, 552 extant species have been identified which are distributed across the tropical and temperate regions.[4] Various species inhabit altitudes ranging from sea level to high mountains, as in the case of most species of Parnassius. The majority of swallowtail species and the greatest diversity are found in the tropics and subtropical regions between 20°N and 20°S,[5]: 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 is found in the British Isles.[6] North America has 40 species, including several tropical species and Parnassius.[7]

The northernmost swallowtail is the Arctic apollo (Parnassius arcticus), found in the Arctic Circle in northeastern Yakutia, at altitudes of 1500 meters above sea level.[8] In the Himalayas, various apollo species such as Parnassius epaphus, have been found at altitudes of 6,000 meters above sea level.[9]:221


The detailed descriptions of morphological characteristics of the Papilionidae, as quoted in Bingham (1905) are as follows:[10]:1,2

Egg. "Dome-shaped, smooth or obscurely facetted, not as high as wide, somewhat leathery, opaque." (Parnassius, the pupa is placed in a loose silken web between leaves. Imago. Wings extraordinarily variable in shape. Hind wing very frequently has a tail, which may be slender, or broad and spatulate, but is always an extension of the termen at vein 4. In one genus, Armandia, the termen of the hind wing is prolonged into tails at the apices of veins 2 and 3 as well as at vein 4. Pore wing (except in the aberrant genera Parnassius and Hypermnestra) with all 12 veins present and in addition a short internal vein, vein 1 a,[11] that invariably terminates on the dorsal margin.

Stages of development of a papilionid – Giant Swallowtail (Papilio cresphontes)

Distinguishing characteristics

The key characteristics that differentiate the Papilionidae from the other butterfly families are:[1]

  • The osmeterium is a forked, fleshy eversible organ found in the prothoracic segment of caterpillars.
  • Venation - 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 sclerites of the cervix (membranous neck between the head and thorax) are fused beneath the neck where the muscles for head movement are anchored.

Special adaptations and defense

Swallowtail butterflies practice Batesian mimicry, a behavior in which the butterflies' appearance closely resemble that of distasteful species that prevents predation. Swallowtails differ from many animals that practice mimicry. The tiger swallowtail butterfly (Papilio glaucus), exhibits a female-limited polymorphism for Batesian mimicry and others, such as the Canadian tiger swallowtail (Papilio canadensis) do not display any form of mimicry.[12]

Predators include the red-winged blackbird, Pennsylvania firefly, five-lined skink, green darner, goldenrod spider, Chinese mantid, fiery searcher, and striped skunk.[13]

Biological basis for polymorphisms in mimicry

Not all individuals in some species are dentical in appearance. For example, Papilio glaucus (Eastern Tiger swallowtail), Y-linkage determines whether the females are either enzyme BAS, which assists dopamine in producing the yellow pigmentation, 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, the pipevine swallowtail. There are also Papilio glaucus that are not wholly black; several possess an intermediate "sooty" color and are sensitive to temperature.[12]

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.[12] 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.


Only certain subsets of swallowtail females practice mimicry. This polymorphism is seen in Papilo dardanus, the African swallowtail butterfly, whose females have three different morphs for wing color pattern: a black-and-white pattern for Batesian mimicry, 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.[15] Given that the males of the species, which do not have Batesian mimicry, are preyed upon much more frequently by predators than the females, scientists initially questioned why females would choose an andromorph wing pattern, which would seemingly lower their fitness compared to the mimicry form.

Several hypotheses for this phenomena 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.[16] 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.[17]

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.[15] The scientists concluded that frequency-dependent selection did lead to equal success for all three alternative strategies: the Batesian females suffered the least number of 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.



Short-horned baronia
(Baronia brevicornis),
subfamily Baroniinae.

The genera of extant swallowtails are usually classified into three subfamilies, Baroniinae, Parnassiinae, and Papilioninae, the latter two being further divided into tribes. In swallowtails, besides morphological characteristics, the choice of foodplants and ecological lifestyle reflect phylogeny and classification.


The Baroniinae are a monotypic subfamily, restricted to a very small region in Mexico and are considered to be the most basal of the subfamilies. Baronia brevicornis is considered to be a relict species, and shares features with a fossil taxon Praepapilio. Baronia is unique amongst papilionids as having an Acacia species (family Leguminosae) as its foodplant.[5]


The Parnassinae are a subfamily of essentially [18] The tribes recognized in the Parnassinae are Parnassiini, Zerynthiini, and Luehdorfiini.

Tribe Parnassiini contains two monotypic genera, [19]

Subfamily: Parnassiinae.


The tribes recognised in the Papilioninae are Leptocircini, Teinopalpini, Troidini, and Papilionini.

Subfamily: Papilioninae.


An additional subfamily, Praepapilioninae, consisting of a single genus Praepapilio, includes two species of extinct butterflies, each member being described from single fossils found in a middle Eocene deposit in Colorado, U.S.A. (Durden and Rose, 1978).[20]


A [19]


Praepapilioninae (†)











Phylogeny of the Papilionidae
(after Nazari, 2007)[1][19]

It is now accepted that the subfamily Papilioninae is [1] The Swallowtail butterflies in the nominate tribe Papilionini number about 225 species and studies have been made on their host-plant coevolution and phylogeny. Old morphological classifications were also found to be valid in that they formed clusters. Species belonging to the groups that use Rutaceae as host plants formed two groups corresponding to Old World and American taxa. Those that fed on Lauraceae and Magnoliaceae were found to form another cluster which includes both Asian and American taxa.[21]

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.[1] Of the [19]

The subfamily Baroniinae is represented by the sole representative species [1]

Mating and young

After mating, the male Parnassines produce a glue like substance that is used to seal the female genital opening and prevent other males from mating.[23] They lay individual eggs on the underside of the leaves of their food plants.[24] There is no parental investment once the eggs have been laid.

The pupae are typically attached to the substrate by the cremaster but with head up held by a silk girdle. The Apollos, however, pupate in debris on the ground and also build a loose cocoon. In the temperate regions, the winters are passed in a pupal diapause stage.


Scarce swallowtail butterfly, Iphiclides podalirius on lavender flowers, near Adriatic coast

The caterpillars of various swallowtail butterfly species feed on a wide range of different plants, most depending on only one of five families: Aristolochiaceae, Annonaceae, Lauraceae, Umbelliferae (Apiaceae) and Rutaceae. 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.[25] Swallowtail tribes Zerynthiini (Parnassiinae), Luehdorfiini (Parnassiinae) and Troidini (Papilioninae) almost exclusively use the Aristolochiaceae family as their host plants.

For example, the eastern black swallowtail's main host plant in the wild is Queen Anne's lace, but they also eat garden plants in the carrot family, including carrots, parsley, dill, and fennel.[13]

Adult swallowtails sip nectar, but also mud and sometimes manure.[24]

Swallowtails and humans

As swallowtail butterflies are large, colourful, and attractive, they were once targeteted by butterfly collectors. The largest of these, the birdwing butterflies are particularly sought after and are cultured in butterfly farms for the purpose of collectors.

Many members of the family feed as pests in citrus orchards.

The Delaware, and South Carolina. The black swallowtail is the state butterfly of Oklahoma.

In popular culture

The American TV show Gilligan's Island had an episode[26] where an explorer[27] came to the island seeking a rare "pussycat swallowtail."

Swallowtails have appeared in various forms of Japanese entertainment, such as [28] In the manga and anime Bleach, Shinigami use Hell Butterflies to send messages and travel between Soul Society and the Living World (Earth); the same butterflies also guide souls during soul burials. These swallowtails are entirely black except for a few red markings on the wings, making them resemble Papilio protenor.[29] The butterfly-based entity Beautifly from the Pokémon anime series is pictured as a swallowtail. Swallowtail butterflies also appear in the manga xxxHolic which resemble yuuko ichihara. The antagonist Koushaku Chono (Papillon) in the manga/anime Busou Renkin features swallowtail-like imagery.

The band Rudolf Steiner, later known as Schwarz Stein, recorded a song entitled 黒揚羽 (lit. Black Swallowtail) on an early demotape. It is a track on the 2006 collaboration album Another Cell. The band Elvenking recorded a song titled "Swallowtail", which was released on their 2006 album, The Winter Wake. Paramore's 2009 release "Brand New Eyes" features a dissected swallowtail butterfly on its cover.

In the Japanese video game Bayonetta the main character Bayonetta is a witch who is in a pact with a demon(s) who takes on a swallowtail form.


  1. ^ a b c d e f g Reed, Robert D.; Sperling, Felix A.H. (2006). – The Swallowtail Butterflies"Papilionidae". Tree of Life Web Project. Retrieved 7 November 2010. 
  2. ^ Richards, O. W.; Davies, R.G. (1977). Imms' General Textbook of Entomology: Volume 1: Structure, Physiology and Development Volume 2: Classification and Biology. Berlin: Springer.  
  3. ^ Salmon, Michael A., Marren, Peter, Harley, Basil. The Aurelian Legacy: British Butterflies and Their Collectors. page 252. Publisher: University of California Press. 2001. ISBN 978-0-520-22963-1
  4. ^ Häuser, Christoph L.; de Jong, Rienk; Lamas, Gerardo; Robbins, Robert K.; Smith, Campbell; Vane-Wright, Richard I. (28 July 2005). "Papilionidae – revised GloBIS/GART species checklist (2nd draft)". Retrieved 8 November 2010. 
  5. ^ a b c Collins, N. Mark; Collins, Michael G. (1985). Threatened Swallowtails of the World: the IUCN red data book. IUCN Protected Area Programme Series. Gland, Switzerland and Cambridge, U.K.: IUCN. pp. 401 & 8 plates.  
  6. ^ Coombs, Simon (30 September 2010). "European Butterfly checklist". Retrieved 11 November 2010. 
  7. ^ Brock, Jim P.; Kaufman, Kenn (2003). Butterflies of North America. Boston: Houghton Mifflin.  
  8. ^ Stumpe, Felix. "Parnassius arctica Eisner, 1968". Retrieved 9 November 2010. 
  9. ^ Mani, M. S. (1968). Ecology and Biogeography of High Altitude Insects. Volume 4 of Series entomologica. Springer. p. 530.  
  10. ^  
  11. ^ The vein is since named 2A or second anal vein in modern venation systems.
  12. ^ a b c Scriber, Mark; Hagen, Robert; Lederhouse, Robert (February 1996). "Genetics of Mimicry in the Tiger Swallowtail Butterflies, Papilio glaucus and P. canadensis (Lepidoptera: Papilionidae)". Evolution 50 (1): 222.  
  13. ^ a b Moran, Mark. "Eastern Black Swallowtail". Retrieved 19 October 2013. 
  14. ^ Koch, Bernhardt; Behnecke, Bettina; ffrench-Constant, Richard H. (May 2000). "The molecuar basis of melanism and mimicry in a swallowtail butterfly". Current Biology 10 (10): 591–4.  
  15. ^ a b Cook, S. E.; Jennifer G. Vernon; Melissa Bateson; Tim Guilford (1994). "Mate Choice in the Polymorphic African Swallowtail Butterfly, Papilio dardanus: male-like females may avoid sexual harassment" (PDF). Animal Behavior 47 (2): 389–397.  
  16. ^ Vane-Wright, R.; C.R. Smith (1991). "Phylogenetic Relationships of Three African Swallowtail Butterflies, Papilio dardanus, P. phorcas, and P. constantinus: a cladistic analysis (Lepidoptera: Papilionidae)". Systematic Entomology. Biology of Butterflies 16 (3): 275–291.  
  17. ^ Conrad, K.F.; G Pritchard (1989). "Female Dimorphism and Physiological Colour Change in the Damselfly Argia vivida Hagen Odonata: Coenagrionidae)". Canadian Journal of Zoology 67 (2): 298–304.  
  18. ^ Nazari, Vazrick (2006). Latreille 1804"Parnassius". Tree of Life Web Project. Retrieved 7 November 2010. 
  19. ^ a b c d e Nazari, Vazrick; Sperling, Felix A.H. (2006). "Parnassiinae Duponchel, [1835]". Tree of Life. Tree of Life Web Project. Retrieved 7 November 2010. 
  20. ^ Durden, C. J.; Rose, H. & Rothschild, Miriam (1978). "Butterflies from the middle Eocene: the earliest occurrence of fossil Papilionidae (Lepidoptera)". Pearce-Sellards Ser. Tex. Mem. Mus. 29 (5): 1–25 .
  21. ^ Aubert, J.; Legal, L; Descimon, H.; Michel, F. (1999). "Molecular phylogeny of swallowtail butterflies of the tribe Papilionini (Papilionidae, Lepidoptera)". Mol Phylogenet Evol. 12 (2): 156–167.  .
  22. ^ Katoh, T.; Chichvarkhin, A.; Yagi, T.; Omoto, K. (2005). "Phylogeny and evolution of butterflies of the genus Parnassius: inferences from mitochondrial 16S and ND1 sequences". Zoolog Sci. 22 (3): 343–351.  .
  23. ^ Ramel, Alain. "Les Papilionides, une famille en beauté". Les Insectes – Petit cours illustré d'entomologie(The Insects – A short illustrated course in Entomology). Retrieved 8 November 2010.  English translation.
  24. ^ a b "Swallowtail Butterflies". University of Michigan. Retrieved 19 October 2013. 
  25. ^ von Euw, J.; Reichstein, T. & Rothschild, M. (1968). "Aristolochic acid in the swallowtail butterfly Pachlioptera aristolochiae". Isr. J. Chem. 6: 659–670.  .
  26. ^ """Gilligan's Island Script Episode #75, "Man With A Net. episode #75 script. 
  27. ^ "Gilligan's Island Season 3, Episode 7 summary". Man with a Net. 
  28. ^ Though her insect designation was never announced in Beetleborgs Metallix, her name being Ladyborg, the astral coin that was used to summon her has the illustration of a swallowtail.
  29. ^ "Bleach Soul Reaper Guide". Retrieved 30 November 2009. 

Further reading

  • Chattopadhyay, J. (2007). Swallowtail Butterflies, Biology & Ecology of a few Indian Species. Desh Prashan, Kolkata, India. - 134 pp. ISBN 978-81-905719-1-3.
  • Glassberg, J. (2001). Butterflies through Binoculars. The West.
  • Guppy, C.S. and Shepard, J.H. (2001). Butterflies of British Columbia.
  • Igarashi, S. (1979). Papilionidae and their early stages [in Japanese]. 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. Part II. Papilionidae. - Moscow, 387 p., 20 color tabs, ISBN 978-5-00077-163-1 [1]
  • 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. Novit. Zool. 2(3):167-463 and plates pdf
  • 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 and Pieridae. 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, Rhopalocera). 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. [2]

External links

  • web project.Tree of Life"Papilionidae" on
  • Family Papilionidae at
  • Revised GloBIS/GART Papilionidae species checklist.
  • GloBIS Database Includes type images
  • Natural History Museum, London web site on "Afrotropical Kite Swallowtails".
  • Collins, N.M., Morris, M.G. (1985) Threatened Swallowtail Butterflies of the World. IUCN. ISBN 2-88032-603-6
  • Photoset and description of lesser known species of Swallowtails
  • Swallowtails Papilionidae of the World: a pictorial summary . Comprehensive.
  • Butterflies and Moths of North America
  • Butterflies of America

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