''Papilio dardanus'', the African swallowtail, mocker swallowtail or flying handkerchief, is a species of

butterfly Butterflies are insects in the macrolepidopteran clade Rhopalocera from the order Lepidoptera, which also includes moths. Adult butterflies have large, often brightly coloured wings, and conspicuous, fluttering flight. The group comprise ...

in the 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 larg ...

(the swallowtails). 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".

Classification

Molecular studies have provided evidence that this species' closest relative is '' Papilio phorcas'', with '' Papilio constantinus'' being the next closest (see images below). It is a member of the ''

Papilio ''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. It includes the common yellow swallowtail (''Papilio machaon''), ...

genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of living and fossil organisms as well as viruses. In the hierarchy of biological classification, genus comes above species and below family. In binomial nom ...

of which '' Papilio appalachiensis'' and ''

Papilio xuthus ''Papilio xuthus'', the Asian swallowtail, Chinese yellow 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 Penins ...

'' are also members. ''Papilio dardanus'' is the nominal member of the ''dardanus''

species group In biology, a species complex is a group of closely related organisms that are so similar in appearance and other features that the boundaries between them are often unclear. The taxa in the complex may be able to hybridize readily with each oth ...

. The members of the clade are: *''Papilio dardanus'' Brown, 1776 *'' Papilio constantinus'' Ward, 1871 *'' Papilio delalandei'' Godart, 824/small> *'' Papilio phorcas'' Cramer, 775/small> *'' Papilio rex'' Oberthür, 1886

Subspecies

Listed alphabetically: *''P. d. antinorii'' Oberthür, 1883 (highlands of Ethiopia) *''P. d. byatti'' Poulton, 1926 (northern highlands of Somalia) *''P. d. cenea'' Stoll,

790 __NOTOC__ Year 790 ( DCCXC) was a common year starting on Friday (link will display the full calendar) of the Julian calendar. The denomination 790 for this year has been used since the early medieval period, when the Anno Domini calendar era ...

/small> (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 (highlands of Eritrea) *''P. d. flavicornis'' Carpenter, 1947 (Mt Kulal, north-western Kenya) *''P. d. humbloti'' Oberthür, 1888 (Comoro Islands) *''P. d. meriones'' C. & R. Felder, 1865 (Madagascar) *''P. d. meseres'' Carpenter, 1948 (Uganda, south-western Kenya, Tanzania: the western, southern and south-eastern shores of Lake Victoria) *''P. d. ochraceana'' Vane-Wright 1995 (Mt. Marsabit, northern Kenya) *''P. d. polytrophus'' Rothschild & Jordan, 1903 (Kenya: highlands east of the Rift Valley) *''P. d. sulfurea'' Palisot de Beauvois, 1806 (São Tomé and Príncipe, Bioko) *''P. d. tibullus'' Kirby, 1880 (eastern Kenya, eastern Tanzania, Malawi, Zambia)

Biogeographic realm

Afrotropical realm

Mimicry

The species shows polymorphism in

wing A wing is a type of fin that produces lift while moving through air or some other fluid. Accordingly, wings have streamlined cross-sections that are subject to aerodynamic forces and act as airfoils. A wing's aerodynamic efficiency is e ...

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 Roland Trimen FRS (29 October 1840 in London – 25 July 1916 in London) was a British-South African naturalist, best known for ''South African Butterflies'' (1887–89), a collaborative work with Colonel James Henry Bowker. He was among ...

. Males have a more or less uniform appearance throughout the species' range, but females come in at least 14 varieties or morphs. Some female morphs share a very similar pattern of colouration with various species of distasteful butterfly (e.g. from the

Danainae Danainae is a subfamily of the family Nymphalidae, the brush-footed butterflies. It includes the Daniadae, or milkweed butterflies, who lay their eggs on various milkweeds on which their larvae (caterpillars) feed, as well as the clearwing butt ...

, a subfamily of nymphalids), 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 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. * In positive frequency-dependent selection, the fitness of a phenotyp ...

. Cook ''et al.'' suggest that Batesian mimics gain a fitness advantage by avoiding predators, but suffer harassment from males (see

sexual conflict Sexual conflict or sexual antagonism occurs when the two sexes have conflicting optimal fitness strategies concerning reproduction, particularly over the mode and frequency of mating, potentially leading to an evolutionary arms race between ma ...

), whereas andromorphs (male mimics) are vulnerable to

predation Predation is a biological interaction where one organism, the predator, kills and eats another organism, its prey. It is one of a family of common feeding behaviours that includes parasitism and micropredation (which usually do not kill th ...

but are not harassed by male mating attempts. 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. 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. 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. The engrailed site has been found to have non-synonymous mutations throughout individuals in the species which would allow the divergence of each morph. Studies support that the engrailed gene in ''Papilio dardanus'' is monophyletic and has only evolved once within the species. 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. 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. 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. Such female-limited Batesian mimicry is not unique to this species, even in the genus ''

''. For instance ''

Papilio memnon ''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. Range Its rang ...

'' shows a similar case of polymorphism in females. Similarly, male mimicry has been observed in another insect, a

damselfly Damselflies are flying insects of the suborder Zygoptera in the order Odonata. They are similar to dragonflies, which constitute the other odonatan suborder, Anisoptera, but are smaller and have slimmer bodies. Most species fold the wings along ...

'' Ischnura ramburii'' which also appears to have evolved camouflage to avoid sexual coercion by males.Lessells, K. (2005). "Sexual Conflict". In '' Encyclopedia of Life Sciences''.

Gallery

File:Papilio dardanus chrysalis.JPG, Chrysalis File:Papilio dardanus emerging 2.JPG, Emerging from chrysalis File:Papilio dardanus v.JPG, Adult File:Papilio dardanus on flower (cropped).jpg, In the wild, South Africa File:Papilio dardanus antinorii female.jpg, ''P. d. antinorii'' female File:Papilio dardanus ochracea female.jpg, ''P. d. ochracea'' female File:Papilio dardanus ochracea male.jpg, ''P. d. ochracea'' male File:Papilio dardanus male ventral view.jpg, Ventral view of same male File:Papilio dardanus emerging.ogv, Eclosion video File:Illustrations of new species of exotic butterflies Papilio XII.jpg, File:Specimen at Natural History Museum, Gothenburg 48.jpg, Papilo dardanus museum specimen

References

*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) *

External links

Photo of caterpillarMost Spectacular Batesian Mimicry
{{Taxonbar, from=Q1315393 dardanus Butterflies of Africa Butterflies described in 1776 Articles containing video clips Taxa named by Peter Brown (naturalist)