Papilio dardanus

Papilio dardanus
both male P. d. antinorii, Ethiopia
Scientific classification
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. babingtonia Stoneham, 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]

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:

• Papilio dardanus Brown, 1776
• Papilio constantinus Ward, 1871
• Papilio delalandei Godart, [1824]
• Papilio phorcas Cramer, [1775]
• Papilio rex Oberthür, 1886

Subspecies

Listed alphabetically:^[5]

• P. d. antinorii Oberthür, 1883 ^[6] (highlands of Ethiopia)
• P. d. byatti Poulton, 1926 ^[7] (northern highlands of Somalia)
• P. d. cenea Stoll, [1790] ^[8] (southern Mozambique, eastern Zimbabwe, Botswana, South Africa, Eswatini)
• P. d. dardanus Brown, 1776 (Senegal, Guinea Bissau, Guinea, Sierra Leone, Liberia, Ivory Coast, Ghana, Burkina Faso, Togo, Benin, Nigeria, Cameroon, Equatorial Guinea, Gabon, Congo, Central African Republic, Angola, Democratic Republic of Congo, Uganda, western Kenya, northern Zambia)
• P. d. figinii Storace, 1962^[9] (highlands of Eritrea)
• P. d. flavicornis Carpenter, 1947^[10] (Mt Kulal, north-western Kenya)
• P. d. humbloti Oberthür, 1888^[11] (Comoro Islands)
• P. d. meriones C. & R. Felder, 1865^[12] (Madagascar)
• P. d. meseres Carpenter, 1948 ^[13] (Uganda, south-western Kenya, Tanzania: the western, southern and south-eastern shores of Lake Victoria)
• P. d. ochraceana Vane-Wright 1995 ^[14] (Mt. Marsabit, northern Kenya)
• P. d. polytrophus Rothschild & Jordan, 1903^[15] (Kenya: highlands east of the Rift Valley)
• P. d. sulfurea Palisot de Beauvois, 1806 ^[16] (São Tomé and Príncipe, Bioko)
• P. d. tibullus Kirby, 1880 ^[17] (eastern Kenya, eastern Tanzania, Malawi, Zambia)

Biogeographic realm

Afrotropical realm

Mimicry

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

• Disruptive selection
• Phylogenetics of mimicry
• Supergene
• Cyril Clarke, E. B. Ford and Philip Sheppard (some notable researchers)
• Amauris mimetic model

Gallery

• 
  Chrysalis
• 
  Emerging from chrysalis
• 
  Adult
• 
  In the wild, South Africa
• 
  P. d. antinorii female
• 
  P. d. ochracea female
• 
  P. d. ochracea male
• 
  Ventral view of same male
• Eclosion video
• 
• 
  Papilo dardanus museum specimen

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. 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. 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. 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.

• Carcasson, R.H. (1960). "The Swallowtail Butterflies of East Africa (Lepidoptera, Papilionidae)". Journal of the East Africa Natural History Society pdf Key to East Africa members of the species group, diagnostic and other notes and figures. (Permission to host granted by The East Africa Natural History Society)
• 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.

External links

• Photo showing mimetic morphs and their models
• Photo of caterpillar
• Most Spectacular Batesian Mimicry