Swallowtail butterfly

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Swallowtail butterfly
Papilio machaon Mitterbach 01.jpg
Papilio machaon
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Arthropoda
Class: Insecta
Order: Lepidoptera
Superfamily: Papilionoidea
Family: Papilionidae
Latreille, [1802]
Type genus
Papilio
Subfamilies and genera

There are 31 genera and about 600 species:

Swallowtail butterfly

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 largest butterflies in the world, the birdwing butterflies of the genus Ornithoptera . [1]

Contents

Swallowtails have a number of distinctive features; for example, the papilionid caterpillar bears a repugnatorial organ called the 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 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 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 figures to the swallowtails. The type species: Papilio machaon honored Machaon, one of the sons of Asclepius, mentioned in the Iliad . [3] Further, the species Papilio homerus is named after the Greek poet, Homer. [4]

The Mon of the Taira clan of Japan is an Agehachō (swallowtail butterfly).

Taxonomy

Subfamilies

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 food plants 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 among papilionidae' as having an Acacia species (family Leguminosae) as its food plant. [5]

Subfamily: Baroniene.

Parnassiinae

The Parnassiinae are a subfamily of essentially Holarctic butterflies. The vast majority of species, mostly Parnassius , 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)". [6] The tribes recognized in the Parnassiinae are Parnassiini, Zerynthiini, and Luehdorfiini.

Tribe Parnassiini contains two 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 genera Archon of Asia minor and the genus Luehdorfia 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 , Bhutanitis , Zerynthia and Allancastria . [5] : 13  [7]

Subfamily: Parnassiinae.

Papilioninae

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

Subfamily: Papilioninae.

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, United States (Durden and Rose, 1978). [8]

Phylogeny

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

Phylogeny of the Papilionidae
  Papilionidae  
after Nazari (2007) [1] [7]

It is now accepted that the subfamily Papilioninae is monophyletic. [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. [9]

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 Baronia brevicornis . They are unique in the family to use the Fabaceae (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. [1]

Distribution

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

Scarce swallowtail, lavender, near Adriatic Swallowtail butterfly and lavender flowers.jpg
Scarce swallowtail, lavender, near Adriatic

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 giant swallowtail (Papilio cresphontes)

Distinguishing characteristics

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

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. [22]

Predators include the red-winged blackbird, Pennsylvania firefly, five-lined skink, green darner, goldenrod spider, Chinese mantis, fiery searcher, and striped skunk. [18]

Biological basis for polymorphisms in mimicry

Not all individuals in some species are identical in appearance. For example, Papilio glaucus (eastern tiger swallowtail), Y-linkage determines whether the females are either wild-type (yellow and black) or melanic (dark melanin replaces the yellow background). [23] This genetic difference stems from the fact that melanism is controlled by a single gene, which controls the level of dopamine in the organism. The 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. [22]

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 mimicry. 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 butterflies. This polymorphism is seen in Papilio 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. [24] Given that the males of the species, which do not have Batesian mimicry, are preyed upon much more frequently by 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 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.

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 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.

Related Research Articles

<i>Papilio glaucus</i> Species of insect

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 many different habitats. It flies from spring until fall, during which it produces two to three broods. Adults feed on the nectar of many species of flowers, mostly from those of the families Apocynaceae, Asteraceae, and Fabaceae. P. glaucus has a wingspan measuring 7.9 to 14 cm. The male is yellow with four black "tiger stripes" on each forewing. Females may be either yellow or black, making them dimorphic. The yellow morph is similar to the male, but with a conspicuous band of blue spots along the hindwing, while the dark morph is almost completely black.

<i>Papilio polyxenes</i> Species of insect

Papilio polyxenes, the (eastern) black swallowtail, American swallowtail or parsnip swallowtail, is a butterfly found throughout much of North America. An extremely similar-appearing species, Papilio joanae, occurs in the Ozark Mountains region, but it appears to be closely related to Papilio machaon, rather than P. polyxenes. The species is named after the figure in Greek mythology, Polyxena, who was the youngest daughter of King Priam of Troy. Its caterpillar is called the parsley worm because the caterpillar feeds on parsley.

<i>Papilio troilus</i> Species of butterfly

Papilio troilus, the spicebush swallowtail or green-clouded butterfly, is a common black swallowtail butterfly found in North America. It has two subspecies, Papilio troilus troilus and Papilio troilus ilioneus, the latter found mainly in the Florida peninsula. The spicebush swallowtail derives its name from its most common host plant, the spicebush, members of the genus Lindera.

<span class="mw-page-title-main">Osmeterium</span> Odoriferous defensive structure

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, and this along with the large eye-like spots on the body might be used to startle birds and small reptiles. The osmeterial organ remains inside the body in the thoracic region in an inverted position and is everted when the larva is disturbed in any way emitting a foul, disagreeable odor which serves to repel ants, small spiders and mantids. To humans, this odour is rather strong but not unpleasant, usually smelling like a concentrated scent of the caterpillar's food plant and pineapple.

<span class="mw-page-title-main">Apollo (butterfly)</span> Species of butterfly

The Apollo, also called the mountain Apollo, is a butterfly of the family Papilionidae.

<span class="mw-page-title-main">Parnassiinae</span> Subfamily of butterflies

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 Asia, Europe and North America.

<i>Battus philenor</i> Species of butterfly

Battus philenor, the pipevine swallowtail or blue swallowtail, is a swallowtail butterfly found in North America and Central America. This butterfly is black with iridescent-blue hindwings. They are found in many different habitats, but are most commonly found in forests. Caterpillars are often black or red, and feed on compatible plants of the genus Aristolochia. They are known for sequestering acids from the plants they feed on in order to defend themselves from predators by being poisonous when consumed. The adults feed on the nectar of a variety of flowers. Some species of Aristolochia are toxic to the larvae, typically tropical varieties. While enthusiasts have led citizen efforts to conserve pipevine swallowtails in their neighborhoods on the West coast, the butterfly has not been the subject of a formal program in conservation or protected in legislation. The butterfly is however of "Special Concern" in Michigan, which is on the Northern limit of its range.

<i>Papilio demodocus</i> Species of butterfly

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 on various native plants of especially the family Rutaceae, but have also taken to the leaves of cultivated citrus trees.

<i>Parnassius</i> Genus of insects

Parnassius is a genus of northern circumpolar and montane butterflies usually known as Apollos or snow Apollos. They can vary in colour and form significantly based on their altitude. They also exhibit altitudinal melananism, a high-altitude adaptation. They have dark bodies and darker coloring at the base of their wings, which allows them to absorb solar energy more quickly.

<i>Baronia</i> Species of butterfly

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 butterflies. It is endemic to a very small area of Mexico, where the distribution is patchy and restricted.

<i>Papilio aristophontes</i> Species of butterfly

Papilio aristophontes is a species of butterfly in the family Papilionidae. It is endemic to Comoros.

<i>Papilio xuthus</i> Species of butterfly

Papilio xuthus, the Asian swallowtail, Chinese yellow swallowtail, Japanese Swallowtail, or Xuthus swallowtail, is a yellow-colored, medium to large sized swallowtail butterfly found in northeast Asia, northern Myanmar, southern China, Taiwan, the Korean Peninsula, Japan, Siberia and the Hawaiian Islands. The butterfly has been observed thrice in New Zealand. Once in Dunedin in 1996 emerging from a chrysalis in a car yard specializing in Japanese used cars; it is thought the chrysalis arrived through one of the cars. and later in Auckland in 2011 and 2016. It was also recorded in the state of Arunachal Pradesh, India, in 2014.

<i>Papilio appalachiensis</i> Species of butterfly

Papilio appalachiensis, the Appalachian tiger swallowtail, is a species of swallowtail butterfly found in eastern North America, particularly in the Appalachian Mountains. It is a hybrid of another two Papilio species, Papilio canadensis and Papilio glaucus, with which it shares many characteristics. The butterflies are normally yellow and contain black patterns in their wings. Their wingspans range from 86 to 115 mm. The caterpillars range in color from green and yellow to orange and are ornamented with black specks that give them the appearance of a bird dropping, which is useful for camouflage, or a large eye, a form of mimicry that is also efficient for protection. This species is univoltine. Females lay their eggs in May.

<i>Papilio dardanus</i> Species of butterfly

Papilio dardanus, the Saharan swallowtail,African swallowtail, mocker swallowtail or flying handkerchief, is a species of butterfly in the family Papilionidae. The species is broadly distributed throughout Sub-Saharan Africa. The British entomologist E. B. Poulton described it as "the most interesting butterfly in the world".

<i>Papilio noblei</i> Species of butterfly

Papilio noblei is a butterfly of the family Papilionidae. It is found in Southeast Asia, from Burma to Vietnam. Recently, this species was also sighted in Namdapha National Park, India.

<i>Papilio gallienus</i> Species of butterfly

Papilio gallienus, the narrow-banded swallowtail, is a butterfly of the family Papilionidae. It is found in Nigeria, Cameroon, the Democratic Republic of the Congo and the central part of the Republic of the Congo.

<span class="mw-page-title-main">Parnassiini</span> Tribe of butterflies

The Parnassiini are a tribe of swallowtail butterflies.

References

  1. 1 2 3 4 5 6 7 Reed, Robert D.; Sperling, Felix A.H. (2006). "Papilionidae – The Swallowtail Butterflies". [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. ISBN   0-412-61390-5.
  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. ↑ Lehnert, Matthew S.; Kramer, Valerie R.; Rawlins, John E.; Verdecia, Vanessa; Daniels, Jaret C. (10 July 2017). "Jamaica's Critically Endangered Butterfly: A Review of the Biology and Conservation Status of the Homerus Swallowtail (Papilio (Pterourus) homerus Fabricius)". Insects. 8 (3): 68. doi: 10.3390/insects8030068 . PMC   5620688 . PMID   28698508.
  5. 1 2 3 Collins, N. Mark; Morris, Michael G. (1985). Threatened Swallowtail Butterflies of the World: The IUCN Red Data Book. Gland & Cambridge: IUCN. 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. 1 2 3 4 5 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 Series, Texas Memorial Museum. 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)". Molecular Phylogenetics and Evolution . 12 (2): 156–167. Bibcode:1999MolPE..12..156A. doi:10.1006/mpev.1998.0605. 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). Zoological Science . 22 (3): 343–351. doi:10.2108/zsj.22.343. hdl: 2115/14485 . PMID   15795497. 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. ↑ Brock, Jim P.; Kaufman, Kenn (2003). Butterflies of North America. Boston: Houghton Mifflin. ISBN   0-618-15312-8.
  14. ↑ Stumpe, Felix. "Parnassius arctica Eisner, 1968". Russian-Insects.com. Retrieved 9 November 2010.
  15. ↑ Mani, M. S. (1968). Ecology and Biogeography of High Altitude Insects. Volume 4 of Series entomologica. Springer. p. 530. ISBN   978-90-6193-114-0 . Retrieved 9 November 2010.
  16. ↑ von Euw, J.; Reichstein, T. & Rothschild, M. (1968). "Aristolochic acid in the swallowtail butterfly Pachlioptera aristolochiae". Israel Journal of Chemistry . 6: 659–670. 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. 1 2 Moran, Mark. "Eastern Black Swallowtail". Archived from the original on 1 September 2016. Retrieved 19 October 2013.
  19. 1 2 "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. 1 2 3 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): 222–236. doi:10.2307/2410795. JSTOR   2410795. PMID   28568864.
  23. ↑ 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. Bibcode:2000CBio...10..591K. doi: 10.1016/s0960-9822(00)00494-2 . PMID   10837223. S2CID   21556770.
  24. 1 2 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 Behaviour. 47 (2): 389–397. doi:10.1006/anbe.1994.1053. S2CID   53186308 . Retrieved 15 September 2013.
  25. ↑ 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. 16 (3): 275–291. Bibcode:1991SysEn..16..275V. doi:10.1111/j.1365-3113.1991.tb00689.x. S2CID   85401171.
  26. ↑ 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. 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.

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