Papilio dardanus

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Papilio dardanus
African swallowtail (Papilio dardanus antinorii ) male.jpg
African swallowtail (Papilio dardanus antinorii ) male underside.jpg
both male P. d. antinorii, Ethiopia
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Arthropoda
Class: Insecta
Order: Lepidoptera
Family: Papilionidae
Genus: Papilio
Species:
P. dardanus
Binomial name
Papilio dardanus
Brown, 1776 [1]
Subspecies

See text

Synonyms
List
  • Papilio merope Cramer, 1777
  • Papilio brutus Fabricius, 1781
  • Papilio hippocoon Fabricius, 1793
  • Papilio westermannii Boisduval, 1836
  • Papilio trophonius Westwood, 1842
  • Papilio dionysos Doubleday, 1846
  • Papilio dardanus ab. niobe Aurivillius, 1899
  • Papilio dardanus ab. nioboides Aurivillius, 1899
  • Papilio dardanus heimsi Suffert, 1904
  • Papilio dardanus benio Suffert, 1904
  • Papilio dardanus f. trophonissa Aurivillius, 1907
  • Papilio dardanus polytrophus f. alluaudi Boullet & Le Cerf, 1912
  • Papilio f. latemarginatus Schultze, 1913
  • Papilio dardanus f. sirius Reuss, 1921
  • Papilio dardanus ab. crocotus Poulton, 1923
  • Papilio dardanus f. ceneispila Le Cerf, 1924
  • Papilio dardanus f. fagerskioldi Bryk, 1928
  • Papilio dardanus dardanus ab. punctata Dufrane, 1933
  • Papilio dardanus dardanus ab. divisa Dufrane, 1933
  • Papilio dardanus dardanus ab. bipunctata Dufrane, 1933
  • Papilio dardanus dardanus ab. paradoxa Dufrane, 1946
  • Papilio dardanus dardanus f. completa Dufrane, 1946
  • Papilio dardanus dardanus f. subpunctata Dufrane, 1946
  • Papilio dardanus dardanus f. dawanti Dufrane, 1946
  • Papilio dardanus dardanus f. impunctata Dufrane, 1946
  • Papilio dardanus dardanus f. jottrandi Dufrane, 1946
  • Papilio dardanus dardanus f. extrema Dufrane, 1946
  • Papilio dardanus f. semimelas Basquin & Turlin, 1986
  • Papilio dardanus ab. obscura McLeod & McLeod, 2004
  • Papilio antinorii Oberthür, 1883
  • Papilio antinorii ab. niavioides Kheil, 1890
  • Papilio antinorii ab. ruspinae Kheil, 1890
  • Papilio dardanus antinorii var. niavina Haase, 1891
  • Papilio dardanus antinorii var. alticola Boullet & Le Cerf, 1912
  • Papilio dardanus hodsoni Poulton, 1926
  • Papilio dardanus hodsoni f. weinholti Poulton, 1927
  • Papilio dardanus f. zaoditou Ungemach, 1932
  • Papilio dardanus antinorii f. alameitu Gabriel, 1949
  • Papilio dardanus f. conjunctiflava Stoneham, 1951
  • Papilio dardanus antinorii ab. seriata Storace, 1963
  • Papilio dardanus antinorii morph immaculata Mollet, 1975
  • Papilio dardanus antinorii morph extensinigra Mollet, 1975
  • Papilio dardanus antinorii morph obsoleta Mollet, 1975
  • Papilio dardanus antinorii morph ornata Mollet, 1975
  • Papilio dardanus antinorii morph rufomaculata Mollet, 1975
  • Papilio dardanus antinorii morph aurantiaca Mollet, 1975
  • Papilio dardanus antinorii morph aperta Mollet, 1975
  • Papilio dardanus antinorii morph depuncta Mollet, 1975
  • Papilio dardanus antinorii morph obscura Mollet, 1975
  • Papilio dardanus antinorii morph parvicaudata Mollet, 1975
  • Papilio dardanus antinorii morph cenaeoides Mollet, 1975
  • Papilio dardanus antinorii morph streckerioides Mollet, 1975
  • Papilio dardanus antinorii morph lambornieoides Mollet, 1975
  • Papilio dardanus antinorii morph niobeoides Mollet, 1975
  • Papilio dardanus antinorii morph salaamioides Mollet, 1975
  • Papilio cenea Stoll, 1790
  • Danais rechila Godart, 1819
  • Papilio cephonius Hopffer, 1866
  • Papilio merpoe tibullus var. hippocoonides Haase, 1891
  • Papilio cenea acene Suffert, 1904
  • Papilio dardanus f. leighi Poulton, 1912
  • Papilio dardanus cenea f. radiata Reuss, 1921
  • Papilio dardanus cenea f. natalica Le Cerf, 1924
  • Papilio dardanus cenea f. hypolimnides Le Cerf, 1924
  • Papilio dardanus cenea f. extensiflava Le Cerf, 1924
  • Papilio dardanus cenea f. sylvicola van Son, 1949
  • Papilio dardanus cenea f. neocenea Stoneham, 1951
  • Papilio dardanus cenea f. nigricans Storace, 1955
  • Papilio dardanus cenea f. transiens Storace, 1955
  • Papilio dardanus cenea f. aikeni van Son, 1956
  • Papilio dardanus cenea f. infuscata van Son, 1956
  • Papilio dardanus antinorii f. vaccaroi Storace, 1947
  • Papilio dardanus antinorii f. protoniavioides Storace, 1962
  • Papilio dardanus figinii f. protomima Storace, 1962
  • Papilio humbloti Oberthür, 1888
  • Papilio meriones Felder & Felder, 1865
  • Papilio dardanus-meriones f. palaeotypus Le Cerf, 1924
  • Papilio merope f. planemoides Trimen, 1904
  • Papilio dardanus f. dionysoides Aurivillius, 1907
  • Papilio dardanus f. pemtolipus Aurivillius, 1907
  • Papilio dardanus f. mixtus Aurivillius, 1907
  • Papilio dardanus f. acenides Le Cerf, 1924
  • Papilio dardanus f. swynnertoni Poulton, 1929
  • Papilio dardanus f. carpenteri Poulton, 1929
  • Papilio dardanus f. xanthocaudatus Stoneham, 1932
  • Papilio dardanus f. hippocoonatus Stoneham, 1933
  • Papilio dardanus f. dominicanoides Stoneham, 1933
  • Papilio dardanus f. planematus Stoneham, 1934
  • Papilio dardanus f. epiplanemoides Stoneham, 1934
  • Papilio dardanus xanthocaudatus f. briani Stoneham, 1944
  • Papilio dardanus ochracea Carpenter, 1948
  • Papilio dardanus cenea ochracea f. ochracea Poulton, 1924
  • Papilio dardanus ochracea f. atavica Storace, 1955
  • Papilio nandina Rothschild & Jordan, 1901
  • Papilio dardanus tibullus f. dorippoides Trimen, 1909
  • Papilio dardanus f. speciosa Le Cerf, 1912
  • Papilio dardanus f. jeanneli Le Cerf, 1912
  • Papilio dardanus var. polytrophus f. punctimargo Le Cerf, 1912
  • Papilio dardanus polytrophus f. hippocooninus Reuss, 1921
  • Papilio dardanus polytrophus f. albescens Reuss, 1921
  • Papilio dardanus polytrophus f. nigrescens Reuss, 1921
  • Papilio dardanus polytrophus f. acenoides Reuss, 1921
  • Papilio dardanus polytrophus f. trophonoides Reuss, 1921
  • Papilio dardanus polytrophus f. mixtoides Reuss, 1921
  • Papilio dardanus var. polytrophus f. protocenea Bryk & Peebles, 1932
  • Papilio dardanus var. polytrophus f. astarte Bryk & Peebles, 1932
  • Papilio dardanus f. nairobianus Stoneham, 1932
  • Papilio dardanus f. poultoni Ford, 1936
  • Papilio dardanus nairobianus f. akechia Stoneham, 1951
  • Papilio dardanus nairobianus f. neria Stoneham, 1951
  • Papilio dardanus f. millari Stoneham, 1951
  • Papilio dardanus f. babingtoniaStoneham, 1951
  • Papilio dardanus nairobianus f. akechiana Stoneham, 1951
  • Papilio dardanus nairobianus f. neriana Stoneham, 1951
  • Papilio dardanus f. harmonia Stoneham, 1951
  • Papilio dardanus f. ariadne Stoneham, 1951
  • Papilio sulfurea Palisot de Beauvois, 1806
  • Papilio dardanus storacei Gauthier, 1984
  • Papilio dardanus sulphureus Bernardi, Pierre & Nguyen, 1985
  • Papilio tibullus Kirby, 1880
  • Papilio cenea maculatus Suffert, 1904
  • Papilio cenea discopunctatus Suffert, 1904
  • Papilio cenea salaami Suffert, 1904
  • Papilio boosi Suffert, 1904
  • Papilio dardanus f. trimeni Poulton, 1906
  • Papilio dardanus var. tibullus ab. gomia Strand, 1911
  • Papilio dardanus tibullus f. lamborni Poulton, 1918
  • Papilio dardanus f. mombasicus Stoneham, 1951
  • Papilio dardanus f. melanoleuca Stoneham, 1951

Papilio dardanus , the Saharan swallowtail,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. [2] The British entomologist E. B. Poulton described it as "the most interesting butterfly in the world". [3]

Contents

Classification

Molecular studies have provided evidence that this species' closest relative is Papilio phorcas , with Papilio constantinus being the next closest (see images below). [4] It is a member of the Papilio genus of which Papilio appalachiensis and Papilio xuthus are also members.

Papilio dardanus is the nominal member of the dardanus species group. The members of the clade are:

Subspecies

Listed alphabetically: [5]

Biogeographic realm

Afrotropical realm

Mimicry

Mating Papilio dardanus mating.jpg
Mating

The species shows polymorphism in wing appearance, though this is limited to females, which are often given as an example of Batesian mimicry in insects. This female-limited mimicry was first described in 1869 by Roland Trimen. [18] Males have a more or less uniform appearance throughout the species' range, but females come in at least 14 varieties or morphs. [2]

Some female morphs share a very similar pattern of colouration with various species of distasteful butterfly (e.g. from the Danainae, a subfamily of nymphalids), [19] while others have been found that mimic male appearance (andromorphs). The persistence of these various morphs or different types of females may be explained by frequency-dependent selection. Cook et al. suggest that Batesian mimics gain a fitness advantage by avoiding predators, but suffer harassment from males (see sexual conflict), whereas andromorphs (male mimics) are vulnerable to predation but are not harassed by male mating attempts. [20]

Morphs are divided into three general groups based on patterning: the hippocoon group, the cenea group, and the planemoides group. The hippocoon group holds the largest amount of morphs; phenotypes within this group are characterized by four bands of alternating black and color patterns. Within the cenea group patterns are greatly dominated by black coloration and contain small splotches of color. The planemoides group has black bands surrounding the outside of the wing with a large splotch of color through the middle of the wing. This group also contains the female forms that are male-like mimics. Diversity in the wing patterns of each group is seen mostly in the coloration of each organism, while black patterns are generally consistent in each morph. [2]

Phenotypic variation within the female morphs of Papilio dardanus has been found to be controlled at one locus named H that contains at least 11 different alleles. [2] Recent studies have narrowed down the region of H to approximately 24 genes that is centered around the engrailed (en) gene which codes for specific transcription factors. [21] The engrailed site has been found to have non-synonymous mutations throughout individuals in the species which would allow the divergence of each morph. [21] Studies support that the engrailed gene in Papilio dardanus is monophyletic and has only evolved once within the species. [22] Findings also suggest that the many different mimetic alleles in the Papilio dardanus genome are solely from mutations in the species. In other words, alleles did not enter into the genome from genetic transfer from other species. [22]

Different combinations of the alleles at H lead to the variety of forms seen within the species. Genetic crosses of individuals found a general dominance hierarchy within the alleles. [2] Allele combinations also determine not only which morph will be expressed but the actual size of the patterns shown. Each allele is able to either influence a larger or smaller mimetic pattern in an organism. [2]

Such female-limited Batesian mimicry is not unique to this species, even in the genus Papilio . For instance Papilio memnon shows a similar case of polymorphism in females. [23] Similarly, male mimicry has been observed in another insect, a damselfly Ischnura ramburii which also appears to have evolved camouflage to avoid sexual coercion by males. [24]

See also

Related Research Articles

<span class="mw-page-title-main">Polymorphism (biology)</span> Occurrence of two or more clearly different morphs or forms in the population of a species

In biology, polymorphism is the occurrence of two or more clearly different morphs or forms, also referred to as alternative phenotypes, in the population of a species. To be classified as such, morphs must occupy the same habitat at the same time and belong to a panmictic population.

<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</i> Genus of butterflies

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.

<span class="mw-page-title-main">Swallowtail butterfly</span> Butterflies of 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 inhabit every continent except Antarctica. The family includes the largest butterflies in the world, the birdwing butterflies of the genus Ornithoptera.

<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.

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.

<span class="mw-page-title-main">Batesian mimicry</span> Bluffing imitation of a strongly defended species

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, who worked on butterflies in the rainforests of Brazil.

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

The viceroy is a North American butterfly. It was long thought to be a Batesian mimic of the monarch butterfly, but since the viceroy is also distasteful to predators, it is now considered a Müllerian mimic instead.

<i>Limenitis arthemis</i> Species of butterfly

Limenitis arthemis, the red-spotted purple or white admiral, is a North American butterfly species in the cosmopolitan genus Limenitis. It has been studied for its evolution of mimicry, and for the several stable hybrid wing patterns within this nominal species; it is one of the most dramatic examples of hybridization between non-mimetic and mimetic populations.

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

Papilio polytes, the common Mormon, is a common species of swallowtail butterfly widely distributed across Asia.

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

Papilio memnon, the great Mormon, is a large butterfly native to southern Asia that belongs to the swallowtail family. It is widely distributed and has thirteen subspecies. The female is polymorphic and with mimetic forms.

<i>Callosamia promethea</i> Species of moth

Callosamia promethea, commonly known as the promethea silkmoth, is a member of the family Saturniidae, which contains approximately 1,300 species. It is also known as the spicebush silkmoth, which refers to one of the promethea silkmoth's common host plants, spicebush. C. promethea is classified as a silk moth, which stems from its ability to produce silk, which it does in the formation of its cocoon. C. promethea lives in forests in the eastern U.S. and does not damage the trees on which it lives. The species was first described by Dru Drury in 1773.

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

Papilio constantinus, the Constantine's swallowtail, is a butterfly of the family Papilionidae. It is found in Sub-Saharan Africa.

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

Papilio cynorta, the mimetic swallowtail or common white banded papilio, is a butterfly of the family Papilionidae. It is found in Africa, including Sierra Leone, Liberia, Ivory Coast, Ghana, Togo, southern Nigeria, Cameroon, Equatorial Guinea, the Democratic Republic of the Congo, Angola, the Republic of the Congo, Uganda, Kenya and Tanzania.

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

Papilio rex, the regal swallowtail or king papilio, is a butterfly of the family Papilionidae. It is found in Africa. It is a semi-montane and montane forest species. The larvae feed on Teclea tricocarpa, Teclea stuhlmanni, Calodendrum, Citrus, Clausena, Fagara and Toddalia species. In the early morning and late afternoon adults of both sexes descend from the forest canopy to feed from the flowers of Lantana, Impatiens and Bougainvillea. It hilltops on granite outcrops and mud puddles.

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

Papilio delalandei is a butterfly of the family Papilionidae. It is endemic to Madagascar.

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

Papilio phorcas, the apple-green swallowtail or green-banded swallowtail, is a butterfly of the family Papilionidae. It is found in Africa.

Many types of polymorphism can be seen in the insect order Lepidoptera. Polymorphism is the appearance of forms or "morphs" differing in color and number of attributes within a single species. In Lepidoptera, polymorphism can be seen not only between individuals in a population but also between the sexes as sexual dimorphism, between geographically separated populations in geographical polymorphism and also between generations flying at different seasons of the year. It also includes the phenomenon of mimicry when mimetic morphs fly alongside non-mimetic morphs in a population of a particular species. Polymorphism occurs both at a specific level with heritable variation in the overall morphological design of individuals as well as in certain specific morphological or physiological traits within a species.

<i>Dazzled and Deceived</i> Camouflage book by Peter Forbes

Dazzled and Deceived: Mimicry and Camouflage is a 2009 book on camouflage and mimicry, in nature and military usage, by the science writer and journalist Peter Forbes. It covers the history of these topics from the 19th century onwards, describing the discoveries of Henry Walter Bates, Alfred Russel Wallace and Fritz Müller, especially their studies of butterflies in the Amazon. The narrative also covers 20th-century military camouflage, begun by the painter Abbot Thayer who advocated disruptive coloration and countershading and continued in the First World War by the zoologist John Graham Kerr and the marine artist Norman Wilkinson, who developed dazzle camouflage. In the Second World War, the leading expert was Hugh Cott, who advised the British army on camouflage in the Western Desert.

References

  1. ↑ Peter Brown, 1776 . Nouvelles illustrations de zoologie, contenant cinquante planches enluminées d'oiseaux curieux, et qui non etés jamais descrits, et quelques de quadrupedes, de reptiles et d'insectes, avec de courtes descriptions systematiques. - New illustrations of zoology, containing fifty coloured plates of new, curious, and non-descript birds, with a few quadrupeds, reptiles and insects. Together with a short and scientific description of the same London. (White).
  2. 1 2 3 4 5 6 Nijhout, H. F. (2003). "Polymorphic mimicry in Papilio dardanus: mosaic dominance, big effects, and origins" (PDF). Evolution and Development. 5 (6): 579–592. doi:10.1046/j.1525-142X.2003.03063.x. PMID   14984040. S2CID   25478508 . Retrieved 2009-08-22.
  3. ↑ Poulton, E. B. (1924). "Papilio dardanus. The most interesting butterfly in the world". Journal of the East African and Ugandan Natural History Society. 20: 4–22.
  4. ↑ Vane-Wright, R. I.; Raheem, D. C.; Cieslak, A.; Vogler, A. P. (1999). "Evolution of the mimetic African swallowtail butterfly Papilio dardanus: molecular data confirm relationships with P. phorcas and P. constantinus". Biological Journal of the Linnean Society . 99 (2): 215–229. doi: 10.1111/j.1095-8312.1999.tb01885.x .
  5. ↑ Papilio dardanus, funet.fi
  6. ↑ Oberthür, C., 1883 Spedezione Italiana nell Africa Equatoriale risulati zoologicae Lepidotteri:Rhopalocera Parte Seconda Ann. Mus. Stor. nat. Genova 18: 711 Full text
  7. ↑ Poulton, E.B. 1926 Proceedings of the Royal Entomological Society of London. 1: xlv
  8. ↑ Stoll, C [1790] Die Uitlandsche Kapellen voorkomende in de drie waerrelddeelen Asia, Africa en America Amsterdam (2-5): 134 Full text
  9. ↑ Storace, L., 1962; Doriana 3 (129): 2
  10. ↑ Carpenter, G.D.H. 1947 Proceedings of the Royal Entomological Society of London (B) 16: 55 (55-56)
  11. ↑ Oberthür, C. 1888 Bulletin de la Société Entomologique de France (6) 8: XL (40-42)Full text
  12. ↑ C. & R. Felder, 1865 . Reise der Österreichischen Fregatte Novara Wien, Bd 2 (Abth. 2) (1): 95 Full text
  13. ↑ Carpenter, G. D. H., 1948 Proceedings of the Royal Entomological Society of London. (B) 17 (1-2): 14
  14. ↑ Vane-Wright, R. I., 1995 Carcasson's African Butterflies 144
  15. ↑ Rothschild, W. & Jordan, K., 1903 Novitates Zoologicae 10 (3): 488 Full text
  16. ↑ Palisot de Beauvois, 1806 Insectes receueillis en Afrique et en Amérique 40 (276 pp.) Paris 46, pl.
  17. ↑ Kirby, W.F., 1880 Scientific Proceedings of the Royal Dublin Society 2: 339 (292-340).
  18. ↑ Trimen, R. (1869). "On some remarkable mimetic analogies among African butterflies". Transactions of the Linnean Society of London . 26 (3): 497–522. doi:10.1111/j.1096-3642.1869.tb00538.x.
  19. ↑ Mallet, J. (2001). "Batesian mimicry in Papilio dardanus" . Retrieved 2009-08-23.
  20. ↑ Cook, S. E.; Vernon, J. G.; Bateson, M.; Guilford, T. (1994). "Mate choice in the polymorphic African swallowtail butterfly, Papilio dardanus: Male-like females may avoid sexual harassment". Animal Behaviour . 47 (2): 389–397. doi:10.1006/anbe.1994.1053. S2CID   53186308.
  21. 1 2 Timmermans, M. J. T. M.; Baxter, S. W.; Clark, R.; Heckel, D. G.; Vogel, H.; Collins, S.; Papanicolaou, A.; Fukova, I.; Joron, M.; Thompson, M. J.; Jiggins, C. D.; ffrench-Constant, R. H.; Vogler, A. P. (2014). "Comparative genomics of the mimicry switch in Papilio dardanus". Proceedings of the Royal Society B: Biological Sciences. 281 (1787): 20140465. doi:10.1098/rspb.2014.0465. PMC   4071540 . PMID   24920480.
  22. 1 2 Thompson, M. J.; Timmermans, M. J. T. N.; Jiggins, C. D.; Vogler, A. P. (2014). "The evolutionary genetics of highly divergent alleles of the mimicry locus in Papilio dardanus". BMC Evolutionary Biology. 14 (1): 140. Bibcode:2014BMCEE..14..140T. doi: 10.1186/1471-2148-14-140 . PMC   4262259 . PMID   25081189.
  23. ↑ Mallet, James (2001). "Batesian mimicry in Papilio memnon" . Retrieved 2009-09-09.
  24. ↑ Lessells, K. (2005). "Sexual Conflict". In Encyclopedia of Life Sciences .
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