Swallowtail butterfly

This article is about the insect family. For other uses, see Swallowtail.

Swallowtail butterflies
Cairns Birdwing, Ornithoptera euphorion (G.R. Gray 1852)
Scientific classification
Kingdom:	Animalia
Phylum:	Arthropoda
Class:	Insecta
Order:	Lepidoptera
Suborder:	Ditrysia
Superfamily:	Papilionoidea
Family:	Papilionidae Latreille, [1802]
Type species
Papilio machaon (Old World Swallowtail)
Subfamilies and genera
There are 27 genera and about 600 species: Subfamily Baroniinae Baronia Subfamily Parnassiinae Allancastria Archon Bhutanitis Hypermnestra Luehdorfia Parnassius Sericinus Subfamily Papilioninae Atrophaneura Battus Cressida Euryades Eurytides Graphium Iphiclides Lamproptera Meandrusa Mimoides Ornithoptera Papilio Parides Pharmacophagus Protesilaus Protographium Teinopalpus Trogonoptera Troides

Swallowtail butterflies are large, colorful butterflies that form the family Papilionidae. There are over 550 species,^[1] and though the majority are tropical, members of the family are found on all continents except Antarctica. The family includes the largest butterflies in the world, the birdwing butterflies of Australia (genus Ornithoptera).^[2]

Swallowtails differ from all other butterflies in a number of anatomical traits. Most notably, their caterpillars possess a unique organ behind their heads, called the osmeterium. Normally hidden, this forked structure can be everted when the caterpillar is threatened, or forced out with a gentle squeeze, and emits smelly secretions containing terpenes. The adults are often tailed like the forked tail of some swallows, giving the insect its name.

Distribution

As of 2005, 552 extant species have been identified^[1] which are distributed across the entire world (except Antarctica) and occur from sea level to high mountains, as in the case of most species of Parnassius. The majority of swallowtail species and greatest diversity in form and lifestyle are found in the tropics between 20°N and 20°S,^[3]: particularly Southeast Asia, and between 20°N and 40°N in East Asia. Only 12 species are found in Europe^[4] and only one species, Papilio machaon is found in the British Isles.^[5] North America records 40 species which include tropical species and Parnassius.^[6]

The northern-most swallowtail is the Arctic Apollo (Parnassius arcticus) which is found in the Arctic Circle in northeastern Yakutia, at an altitude of 1500 meters above sea level.^[7] In the Himalayas, various Apollo species such as Parnassius epaphus, besides others, have been recorded to occur up to an altitude of 6,000 meters above sea level.^[8]:221

Morphology

The detailed descriptions of morphological characteristics of the Papilionidae, as quoted in Bingham (1905) are as follows:^[9]:1,2

Egg. "Dome-shaped, smooth or obscurely facetted, not as high as wide, somewhat leathery, opaque." (Doherty.) Larva. Stout, smooth or with a series of fleshy tubercles on the dorsum : sometimes with a raised fleshy protuberance (the so-called hood or crest) on the fourth segment which is also generally thickened above. The second segment has a transverse opening, out of which the larva can protrude at will an erect, forked, glandular fleshy organ that emits a strong, somewhat pleasant, but always penetrating odour. Pupa. Variable in form but superiorly most often curved backwards, sometimes very strongly so ; angulate, with the head truncate or rounded, often bifid ; back of abdomen smooth or tuberculate. Attached by the tail, normally in a perpendicular position, and further secured by a silken girth round the middle. In Parnassius strangely enough the pupa is placed in a loose silken web between leaves. Imago. Wings extraordinarily variable in shape. Hind wing very frequently with 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,^[10] that invariably terminates on the dorsal margin. There is also a short transverse vein present at base of wing between the median vein and vein 1a in all genera except Leptocircus, Armandia, Parnassius, and Hypermnestra. Hind wing : vein 1a absent; precostal vein and precostal cell both present; dorsal margin not excavated so as to receive the abdomen, but in the male frequently folded over and studded within the fold with specialized scales (androconia) or hairs that are often strongly scented. Antennae comparatively short, with generally a distinct club; "the distal joints mostly more expanded ventrally than dorsally, so that the club is curved dorsad" (Jordan). The scaling most extended in Leptocircus, but in Papilio confined to the basal joints. Body stout; claspers at apex of abdomen in the male generally well-developed, absent in a few forms. Six walking legs; the fore tibiae with a medial pad; claws simple except in one form of Leptocircus, which has them bifid.

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

• 

  Egg

• 

  Caterpillar

• 

  Chrysalis

• 

  Adult

Distinguishing characteristics

The key characteristics that differentiate the Papilionidae from the other butterfly families are:^[2]

• The presence of the osmeterium, a forked, fleshy eversible organ found in the prothoracic segment of papilionid caterpillars.
• Venation in the adult forewing of papilionids is characteristic. 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.

• 

  Old World Swallowtail caterpillar everting its osmeterium in defense.

• 

  Adult forewing of Graphium agetes. Second anal vein, 2A, extends upto the wing margin and does not link with the first anal vein, 1A.

• 

  Papilionid chrysalis are typically attached to a substrate by the cremaster and with head up held by a silk girdle.

Taxonomy

Subfamilies

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.

Baroniinae

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.^[3]

Parnassinae

The Parnassinae are a subfamily of essentially holarctic butterflies. The vast majority of species, mostly Parnassius, can be found in mountain habitats. Parnassinines can also be found in other habitats such as "arid deserts (Hypermnestra), humid forests (Luehdorfia) and even lowland meadows (Zerynthia)" (Nazari, 2006).^[11] The tribes recognized in the Parnassinae are Parnassiini, Zerynthiini, and Luehdorfiini.

Tribe Parnassiini contains two monotypic genera, Hypermnestra, largely confined to central Asia and the genus Parnassius (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. The tribe Luehdorfiini contains the monotypic genera Archon of Asia minor and the genus Luehdorfia of China and Japan. These two tribes have evolved to change their foodplants, while the third tribe, Zerynthiini, has retained the archetypical papilionid foodplant, the lowland vine Aristolochia. Zerynthhini comprises four genera – Sericinus, Bhutanitis, Zerynthia and Allancastria.^[3]:13[12]

Subfamily : Parnassiinae.

• 

  Mountain Apollo
  (Parnassius apollo),
  tribe Parnassiini.
• 

  False Apollo
  (Archon apollinus),
  tribe Luehdorfiini.
• 

  Southern Festoon
  (Zerynthia polyxena),
  tribe Zerythiini.
• 

  Bhutan Glory
  (Bhutanitis lidderdalii),
  tribe Zerythiini.

Papilioninae

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

Subfamily : Papilioninae.

• 

  Zebra Swallowtail
  (Eurytides marcellus)
  Tribe Leptocircini.
• 

  Kaiser-i-hind
  (Teinopalpus imperialis)
  Tribe Teinopalpini.
• 

  Palawan Birdwing
  (Trogonoptera trojana)
  Tribe Troidini.
• 

  Spicebush Swallowtail
  (Papilio trolius)
  Tribe Papilionini.
• 

  Papilio protenor Cramer

Praepapilioninae

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).^[13]

Phylogeny

A phylogeny of the Papilionidae based on Nazari (2007) is given:^{[2] [12]}

Phylogeny

Praepapilioninae (†) Baroniinae Parnassiinae Parnassiini Zerynthiini Luehdorfiini Papilioninae Leptocircini Teinopalpini Papilionini Troidinini

Phylogeny of the Papilionidae (after Nazari, 2007)[2][12]

It is now accepted that the subfamily Papilioninae is monophyletic.^[2] 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.^[14]

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.^[2] Of the Parnassiinae, the genera Parnassius and Hypermnestra were found to be extremely close based on molecular studies^[15] and are now considered to be part of the tribe Parnassiini.^[12] The two taxa, Archon and Luehdorfia, 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.^[12]

The subfamily Baroniinae is represented by the sole representative species Baronia brevicornis. They are unique in the family to use the Fabaceae (Leguminosae) as their larval host plants. The Baronninae and the extinct family Praepapilioninae share many external similarities and are traditionally considered to be the most primitive families 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.^[2]

Natural history

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.^[16]

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.

Foodplants

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.^[17] Swallowtail tribes Zerynthiini (Parnassiinae), Luehdorfiini (Parnassiinae) and Troidini (Papilioninae) almost exclusively use the Aristolochiaceae family as their host plants.

Swallowtails and humans

Swallowtail butterflies, being large, colourful, and attractive, have been the target of butterfly collectors in earlier times. 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 larvae on plants of the citrus family, Rutaceae. Some of these attractive butterflies are therefore considered pests in citrus orchards.

The Oregon Swallowtail is the state insect of Oregon. The Eastern Tiger Swallowtail is the state insect of Virginia and the state butterfly of Georgia, 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^[18] where an explorer^[19] came to the island seeking a rare "pussycat swallowtail."

Swallowtails have appeared in various forms of Japanese entertainment, such as tokusatsu, manga and anime. In the 1996 Season of the popular Japanese tokusatsu Metal Hero Series B-Fighter Kabuto and the 1997 American show Beetleborgs Metallix, one of the B-fighters/Astral Borgs motifs was a swallowtail hence her Japanese designated name "B-Fighter Ageha".^[20] 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.^{[citation needed]} These swallowtails are entirely black except for a few red markings on the wings, making them resemble Papilio protenor.^[21] The butterfly-based entity Beautifly from the Pokémon anime series is pictured as a swallowtail.^[22] Swallowtail butterflies also appear in the manga xxxHolic.^{[citation needed]}

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.^{[citation needed]} The band Elvenking recorded a song titled Swallowtail, which was released on their 2006 album, The Winter Wake.^{[citation needed]} Paramore's 2009 release "Brand New Eyes" features a dissected swallowtail butterfly on its cover.^{[citation needed]}

References

1. ^ ^{a b} 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)". http://www.insects-online.de/frames/papilio.htm. Retrieved 8 November 2010. 
2. ^ ^{a b c d e f g} Reed, Robert D.; Sperling, Felix A.H. (2006). "Papilionidae – The Swallowtail Butterflies". Tree of Life Web Project. http://tolweb.org/Papilionidae/12177. Retrieved 7 November 2010. 
3. ^ ^{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. ISBN 9782880326036. http://books.google.co.in/?id=RomV7uO_t9YC. Retrieved 22 October 2010. 
4. ^ Coombs, Simon (30 September 2010). "European Butterfly checklist". http://www.butterfly-guide.co.uk/. http://www.butterfly-guide.co.uk/help/euro_chklist.htm. Retrieved 11 November 2010. 
5. ^ Coombs, Simon (30 September 2010). "UK Butterfly checklist". http://www.butterfly-guide.co.uk/. http://www.butterfly-guide.co.uk/help/uk_chklist.htm. Retrieved 11 November 2010. 
6. ^ Brock, Jim P.; Kaufman, Kenn (2003). Butterflies of North America. Boston: Houghton Mifflin. ISBN 0-618-15312-8. 
7. ^ Stumpe, Felix. "Parnassius arctica Eisner, 1968". Russian-Insects.com. http://rusinsects.com/p-arct.htm. Retrieved 9 November 2010. 
8. ^ Mani, M. S. (1968). Ecology and Biogeography of High Altitude Insects. Volume 4 of Series entomologica. Springer. pp. 530. ISBN 9789061931140. http://books.google.co.in/?id=n4qSTCkniZoC&lpg=PP1&pg=PA221. Retrieved 9 November 2010. 
9. ^ Bingham, C.T. (1905). The Fauna of British India including Ceylon and Burma – Butterflies (Vol 1). London: Taylor and Francis. pp. 519. http://www.archive.org/details/TheFunaOfBritishIndiaButterfliesVolI. Retrieved 7 November 2010. 
10. ^ The vein is since named 2A or second anal vein in modern venation systems.
11. ^ Nazari, Vazrick (2006). "Parnassius Latreille 1804". Tree of Life Web Project. http://tolweb.org/Parnassius/65393. Retrieved 7 November 2010. 
12. ^ ^{a b c d e} Nazari, Vazrick; Sperling, Felix A.H. (2006). "Parnassiinae Duponchel, [1835"]. Tree of Life. Tree of Life Web Project. http://tolweb.org/Parnassiinae. Retrieved 7 November 2010. 
13. ^ Durden, C. J. & Rose, H. (1978). "Butterflies from the middle Eocene: the earliest occurrence of fossil Papilionidae (Lepidoptera)". Pearce-Sellards Ser. Tex. Mem. Mus. 29: 1–25 .
14. ^ 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. doi:10.1006/mpev.1998.0605. PMID 10381318 .
15. ^ 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. doi:10.2108/zsj.22.343. PMID 15795497 .
16. ^ Ramel, Alain. "Les Papilionides, une famille en beauté". Les Insectes – Petit cours illustré d'entomologie(The Insects – A short illustrated course in Entomology). http://aramel.free.fr/INSECTES13-5.shtml. Retrieved 8 November 2010.  English translation.
17. ^ von Euw, J.; Reichstein, T. & Rothschild, M. (1968). "Aristolochic acid in the swallowtail butterfly Pachlioptera aristolochiae". Isr. J. Chem. 6: 659–670 .
18. ^ "Gilligan's Island Script Episode #75, "Man With A Net"". episode #75 script. gilligansisle.com. http://www.gilligansisle.com/scripts/script75.html. 
19. ^ "Gilligan's Island Season 3, Episode 7 summary". Man with a Net. tv.com. http://www.tv.com/gilligans-island/man-with-a-net/episode/10185/summary.html. 
20. ^ 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.
21. ^ "Bleach Soul Reaper Guide". MyFavoriteGames.com. http://www.myfavoritegames.com/bleach/Shinigami_Guide/Shinigami-About.aspx. Retrieved 30 November 2009. 
22. ^ "Beautifly (Pokémon)". Bulbapedia. Bulbasaur's Mysterious Garden (an English-language Pokémon community). http://bulbapedia.bulbagarden.net/wiki/Beautifly_%28Pok%C3%A9mon%29. Retrieved 9 November 2010. 

External links

• "Papilionidae" on Tree of Life web project.
• Family Papilionidae at Lepidoptera.pro
• Revised GloBIS/GART Papilionidae species checklist.
• Natural History Museum, London web site on "Afrotropical Kite Swallowtails".