Swallowtail butterfly

Praepapilioninae

An additional subfamily,

A phylogeny of the Papilionidae based on Nazari (2007) is given:^[1][7]

It is now accepted that the subfamily Papilioninae is

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 Parnassiinae, the genera Parnassius and Hypermnestra were found to be extremely close based on molecular studies^[10] and are now considered to be part of the tribe Parnassiini.^[7] 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.^[7]

The subfamily Baroniinae is represented by the sole representative species

As of 2005, 552 extant species have been identified which are distributed across the tropical and temperate regions.[11] 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.^[12] North America has 40 species, including several tropical species and Parnassius.^[13]

The northernmost swallowtail is the Siberian Apollo (Parnassius arcticus), found in the Arctic Circle in northeastern Yakutia, at altitudes of 1500 meters above sea level.^[14] In the Himalayas, various Apollo species such as Parnassius epaphus, have been found at altitudes of 6,000 meters above sea level.^[15]: 221

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, 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.^[16] Similarly, the Parnassius smintheus sequesters sarmentosin from its host plant Sedum lanceolatum for protection from predators.^[17] Swallowtail tribes Zerynthiini (Parnassiinae), Luehdorfiini (Parnassiinae) and Troidini (Papilioninae), almost exclusively use the family Aristolochiaceae as their host plants.

For example, the eastern black swallowtail's (Papilio polyxenes) 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.^[18]

Adult swallowtails sip nectar, but also mud and sometimes manure.^[19]

Life cycle

The detailed descriptions of morphological characteristics of the Papilionidae, as quoted in Bingham (1905) are as follows:^{[20]: 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. The second segment has a transverse opening, out of which the larva protrudes at will and an erect, forked, glandular fleshy organ that emits a strong, penetrating, and somewhat unpleasant odor.

Pupa. Variable in form but most often curved backwards. It is angulate, with the head truncate or rounded and the back of abdomen is smooth or tuberculate. It is attached by the tail, normally in a perpendicular position, and further secured by a silken girth round the middle. In Parnassius, the pupa is placed in a loose silken web between leaves.

Imago. Wings extraordinarily variable in shape. Hindwing 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 hindwing is prolonged into tails at the apices of veins 2 and 3 as well as at vein 4. Forewing (except in the aberrant genera Parnassius and Hypermnestra) with all 12 veins present and in addition a short internal vein, vein 1 a,^[21] that invariably terminates on the dorsal margin.

Stages of development of a papilionid, the

• 
  Egg
• 
  Larva
• 
  Pupa
• 
  Imago

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.

• 
  Old World swallowtail caterpillar everting its osmeterium

  in defense.
• 
  Papilio demodocus larva applying one horn of the evaginated osmeterium to an offending finger.
• 
  Adult forewing of Graphium agetes. Second anal vein, 2A, extends up to 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.

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.^[22]

Predators include the

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.[22] 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

predators 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

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.^[25] 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.^[26]

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.[24] The scientists concluded that frequency-dependent selection 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, the male Parnassines produce a glue-like substance that is used to seal the female genital opening and prevent other males from mating.^[27] They lay individual eggs on the underside of the leaves of their food plants.^[19] There is no parental investment once the eggs have been laid.

The

temperate regions, the winters are passed in a pupal diapause

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, feed on plants of the citrus family Rutaceae, making some of these attractive butterflies pests in citrus orchards.

The

black swallowtail is the state butterfly of Oklahoma

.

References

1. ^ ^{a b c d e f g} Reed, Robert D.; Sperling, Felix A.H. (2006). "Papilionidae – The Swallowtail Butterflies". [Tree of Life Web Project]. Retrieved 7 November 2010.
2. ISBN 0-412-61390-5
   .
3. ISBN 978-0-520-22963-1
4. PMID 28698508
   .
5. ^
   ISBN 978-2-88032-603-6
   – via Biodiversity Heritage Library.
6. ^ Nazari, Vazrick (2006). "Parnassius Latreille 1804". [Tree of Life Web Project]. Retrieved 7 November 2010.
7. ^ ^{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.
8. ^ 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..
9. ^ Aubert, J.; Legal, L.; Descimon, H.; Michel, F. (1999). "Molecular phylogeny of swallowtail butterflies of the tribe Papilionini (Papilionidae, Lepidoptera)".
   PMID 10381318
   .
10. ^ 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" (PDF).
    S2CID 23898737
    .
11. ^ 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)". Archived from the original on 9 September 2010. Retrieved 8 November 2010.
12. ^ Coombs, Simon (30 September 2010). "European Butterfly checklist". butterfly-guide.co.uk. Archived from the original on 16 March 2017. Retrieved 11 November 2010.
13. ISBN 0-618-15312-8
    .
14. ^ Stumpe, Felix. "Parnassius arctica Eisner, 1968". Russian-Insects.com. Retrieved 9 November 2010.
15. ISBN 978-90-6193-114-0
    . Retrieved 9 November 2010.
16. doi:10.1002/ijch.196800084
    ..
17. ^ Doyle, Amanda. "The roles of temperature and host plant interactions in larval development and population ecology of Parnassius smintheus Doubleday, the Rocky Mountain Apollo butterfly" (PDF). University of Alberta. Retrieved 13 November 2017.
18. ^ ^{a b} Moran, Mark. "Eastern Black Swallowtail". Archived from the original on 1 September 2016. Retrieved 19 October 2013.
19. ^ ^{a b} "Swallowtail Butterflies". University of Michigan. Retrieved 19 October 2013.
20. ^ Bingham, C. T. (1905). The Fauna of British India, Including Ceylon and Burma: Butterflies Volume I. London: Taylor and Francis. p. 519. Retrieved 7 November 2010.
21. ^ The vein is since named 2A or second anal vein in modern venation systems.
22. ^
    PMID 28568864
    .
23. S2CID 21556770
    .
24. ^
    S2CID 53186308
    . Retrieved 15 September 2013.
25. S2CID 85401171
    .
26. doi:10.1139/z89-044
    .
27. ^ 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.

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
  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. Novitates Zoologicae 2(3):167–463 and plates 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 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 (
  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

• Media related to Papilionidae at Wikimedia Commons
• Data related to Papilionidae at Wikispecies

New International Encyclopedia article "Swallowtail

".

• "Papilionidae" on Tree of Life web project.
• "Family Papilionidae". Insecta.pro. Retrieved 7 February 2020.
• Revised GloBIS/GART Papilionidae species checklist. Archived 9 September 2010 at the Wayback Machine
• GloBIS Database Archived 5 November 2013 at the Wayback Machine Includes type images
• Natural History Museum, London web site on "Afrotropical Kite Swallowtails".
• Collins, N. Mark; Morris, Michael G. (1985). Threatened Swallowtail Butterflies of the World: The IUCN Red Data Book. Gland & Cambridge:
  ISBN 978-2-88032-603-6
  – via Biodiversity Heritage Library.
• Photoset and description of lesser known species of swallowtails
• Swallowtails Papilionidae of the World: a pictorial summary. Comprehensive.