Danaus (butterfly)

Danaus, commonly called tigers, milkweeds, monarchs, wanderers, and queens, is a genus of butterflies in the tiger butterfly tribe. They are found worldwide, including North America, South America, Africa, Asia, Indonesia and Australia. For other tigers see the genus, Parantica.

Danaus
Male Danaus chrysippus
Danaus plexippus and Danaus gilippus
Scientific classification
Domain:	Eukaryota
Kingdom:	Animalia
Phylum:	Arthropoda
Class:	Insecta
Order:	Lepidoptera
Family:	Nymphalidae
Subtribe:	Danaina
Genus:	Danaus Kluk, 1802
Type species
Papilio plexippus Linnaeus, 1758
Species
12–13 species, see text

Taxonomy

Following the review of Smith et al. (2005), 12 species are provisionally accepted based on morphological, mtDNA 12S rRNA and cytochrome c oxidase subunit I, and nuclear DNA 18S rRNA and EF1 subunit α sequence data:

Butterfly	Caterpillar	Name	Common name	Distribution
		Danaus affinis (Fabricius, 1775)	Malay tiger, mangrove tiger, or swamp tiger	from Thailand to the Philippines and southwards through Indonesia to Melanesia and northeastern Australia
		Danaus chrysippus (Linnaeus, 1758)	plain tiger, common tiger, African monarch, lesser wanderer, or African queen	Asia, Australia, and Africa
		Danaus cleophile (Godart, 1819)	Jamaican monarch	Dominican Republic, Haiti, and Jamaica.
		Danaus dorippus (Klug, 1845)	dorippus tiger (formerly included in D. chrysippus)	eastern and southern Africa (mainly in Kenya, Uganda, Erythrea, Oman, Tanzania) and sporadically in India
		Danaus eresimus (Cramer, [1777])	soldier or tropical queen, includes D. plexaure	America: southern Florida, southern Texas, Mexico, the Caribbean, Central America, and South America.
		Danaus erippus (Cramer, [1775])	southern monarch	South America, mainly in Brazil, Uruguay, Paraguay, Argentina, Bolivia, Chile, and southern Peru.
		Danaus genutia (Cramer, [1779])	common tiger, Indian monarch, or orange tiger	Sri Lanka, Myanmar and extending to South-East Asia and Australia (except New Guinea).
		Danaus gilippus (Cramer, [1775])	queen	America: from central United States (Kansas, Colorado, and Utah) to Argentina
		Danaus ismare (Cramer, [1780])	Ismare tiger	South East Asia
		Danaus melanippus (Cramer, [1777])	white tiger, common tiger, or eastern common tiger	Asia: Assam in eastern India through South-East Asia south to Indonesia, and eastwards to the Philippines and through southern China to Taiwan.
		Danaus petilia (Stoll, 1790)	lesser wanderer	Australia
		Danaus plexippus (Linnaeus, 1758)	monarch or wanderer	from southern Canada through northern South America. Bermuda, Cook Islands, Hawaii, Cuba, and other Caribbean islands, the Solomons, New Caledonia, New Zealand, Papua New Guinea, Australia, the Azores, the Canary Islands, Gibraltar, the Philippines, and North Africa.

This genus was formerly split into the subgenera Danaus, Salatura, and Anosia, but this arrangement has been abolished. While the first (the 2–3 monarch butterflies) and Salatura (species ismare, genutia, affinis, and melanippus) do indeed seem to be clades, the relationship of these to the other species, especially the puzzling D. dorippus, is not clear.

Hybridization producing fertile offspring is known to occur between some species, confounding mtDNA data; this seems to be especially true in the case of D. dorippus (Smith et al. 2005). In addition, male-killing Spiroplasma infection has been shown in D. chrysippus and probably also occurs in other species (Jiggins et al. 2000); the consequences for speciation and evolution are probably similar to those observed in infection with male-killing strains of the better-researched Wolbachia bacteria.

Phylogeny

Plain tiger (Danaus chrysippus) caterpillar on a milkweed (Calotropis) species in Hyderabad, India

	Tirumala septentrionis
	Tirumala limniace
	Danaus plexippus Danaus erippus Danaus genutia Danaus melanippus Danaus affinis Danaus ismare Danaus eresimus Danaus gilippus Danaus petilia Danaus dorippus dorippus-2 Danaus d. bataviana Danaus chrysippus chrysippus Danaus c. orientis Danaus c. alcippus Danaus d. dorippus-1

Phylogeny of the genus (without D. cleophile).

Synonyms of Danaus species and subspecies

• Source: The higher classification of Nymphalidae, at Nymphalidae.net
• Note: Names preceded by an equal sign ( = ) are synonyms, homonyms, rejected names, or invalid names. Synonymy, given below in abbreviated form, is from Ackery & Vane-Wright (1984).

Tribe Danaini Boisduval, 1833

• Danaus Kluk, 1802 ( = Danaida Latreille, 1804; = Limnas Hübner, 1806; = Danais Latreille, 1807; = Danaus Latreille, 1809; = Anosia Hübner, 1816; = Festivus Crotch, 1872; = Salatura Moore, 1880; = Nasuma Moore, 1883; = Tasitia Moore, 1883; = Danaomorpha Kremky, 1925; = Panlymnas Bryk, 1937; = Diogas d'Almeida, 1938)
  • Danaus cleophile (Godart, 1819) ( original name = Danais cleophile Godart, 1819)
  • Danaus plexippus (Linnaeus, 1758) ( original name = Papilio plexippus Linnaeus, 1758; = misippiformis Meuschen, 1781; = archippus Fabricius, 1793; = menippe Hübner, 1816; = megalippe Hübner, 1826; = leucogyne Butler, 1884; = fumosus Hulstaert, 1886; = pulchra Strecker, 1900; = nigrippus Haensch, 1909; = americanus Gunder, 1927; = nivosus Gunder, 1927; = curassavicae Fabricius, 1938; = portoriciensis Clark, 1941; = tobagi Clark, 1941; = disjuncta Dufrane, 1948; = bipunctata Dufrane, 1948; = obliterata Dufrane, 1948)
  • Danaus erippus (Cramer, 1775) ( original name = Papilio erippus Cramer, 1775 ( = larensis Köhler, 1929; = asclepiadis Fabricius, 1938)
  • Danaus ismare (Cramer, 1780) ( original name = Papilio ismare Cramer, 1780; = ismareola Butler, 1866; = fulvus Ribbe, 1890; = celebensis Rothschild, 1892; = felicia Fruhstorfer, 1907; = goramica Fruhstorfer, 1907; = alba Morishita, 1981)
  • Danaus genutia (Cramer, 1779) ( original name = Papilio genutia Cramer, 1779; = adnana Swinhoe, 1917; = albipars Talbot, 1943; = alexis Waterhouse & Lyell, 1914; = bandjira Martin, 1911; = bimana Martin, 1911; = connectens Moulton, 1921; = conspicua Butler, 1866; = grynion Fruhstorfer, 1907; = intensa Moore, 1883; = intermedia Moore, 1883; = kyllene Fruhstorfer, 1910; = laratensis Butler, 1883; = leucoglene Felder & Felder, 1865; = niasicus Fruhstorfer, 1899; = nipalensis Moore, 1877; = partita Fruhstorfer, 1897; = plexippus auctt. nec Linnaeus, 1758; = sumatrana Moore, 1883; = sumbana Talbot, 1943; = telmissus Fruhstorfer, 1910; = tuak Pryer & Cator, 1894; = tychius Fruhstorfer, 1910; = uniens Martin, 1911; = wetterensis Fruhstorfer, 1899; = yuchingkini Murayama & Shimonoya, 1960 (hybrid chrysippus x genutia, Kurosawa, 1973)
  • Danaus affinis (Fabricius, 1775) ( original name = Papilio affinis Fabricius, 1775; = abigar Eschscholtz, 1821; = adustus Godman & Salvin, 1882; = affinoides Fruhstorfer, 1899; = albistriga Talbot, 1943; = albonotata Howarth, 1962; = artenice Cramer, 1781; = aruana Moore, 1883; = astakos Fruhstorfer, 1906; = batjana Fruhstorfer, 1899; = bipuncta Talbot, 1943; = biseriata Butler, 1882; = bonguensis Fruhstorfer, 1899; = cecilia Bouganville, 1837; = chionippe Hübner, 1823; = cometho Godman & Salvin, 1888; = coriacea Fruhstorfer, 1906; = decentralis Fruhstorfer, 1899; = decipiens Butler, 1882; = decipientis Strand, 1914; = djampeana Van Eecke, 1915; = fergussonia Fruhstorfer, 1907; = ferruginea Butler, 1876; = fuliginosa Hagen, 1894; = fulgurata Butler, 1866; = fuscata Talbot, 1943; = galacterion Fruhstorfer, 1906; = gelanor Waterhouse & Lyell, 1914; = hegesippinus Röber, 1891; = insolata Butler, 1870; = jimiensis Miller & Miller, 1978; = jobiensis Grose-Smith, 1894; = kapaura Talbot, 1943; = kawiensis Fruhstorfer, 1899; = kiriwina Fruhstorfer, 1907; = leucippis Röber, 1891; = litoralis Doherty, 1891; = luxurians Fruhstorfer, 1907; = malayana Fruhstorfer, 1899; = molyssa Fruhstorfer, 1907; = mysolica Moore, 1883; = mytilene Felder & Felder, 1860; = nigrita Moore, 1883; = nore Swinhoe, 1917; = nubila Butler, 1866; = obscura Capronnier, 1886; = oesypera Talbot, 1943; = olga Swinhoe, 1917; = oros Fruhstorfer, 1907; = paradoxa Röber, 1939; = philene Stoll 1782; = piepersi Kalis, 1933; = pittakus Fruhstorfer, 1907; = pleistarchus Fruhstorfer, 1912; = pseudophilene Fruhstorfer, 1907; = pullata Butler, 1866; = rubrica Fruhstorfer, 1906; = sabrona Talbot, 1943; = sangira Fruhstorfer, 1899; = signata Talbot, 1943; = snelleni Kalis, 1933; = strephon Fruhstorfer, 1907; = subnigra Joicey & Talbot, 1922; = subnubila Fruhstorfer, 1907; = tambora Fruhstorfer, 1899; = taroena Van Eecke, 1915; = taruna Fruhstorfer, 1899; = transfuga Fruhstorfer, 1907; = tualana Talbot, 1943; = vandeldeni Kalis, 1933; = vorkeinus Röber, 1866; = wentholti Martin, 1914; = woodlarkiana Fruhstorfer, 1907)
  • Danaus melanippus (Cramer, 1777) ( original name = Papilio melanippus Cramer, 1777; = hegesippus Cramer, 1777; = lotis Cramer, 1779; = thoe Hübner, 1816; = edmondii Bougainville, 1837; = nesippus C. Felder, 1862; = eurydice Butler, 1884; = celebensis Staudinger, 1889; = erebus, Röber, 1891 ; = haruhasa Doherty, 1891; = pietersii Doherty, 1891; = taimanu Doherty, 1891; = fruhstorferi Röber, 1897; = keteus Hagen, 1898; = indicus Fruhstorfer, 1899; = malossona Fruhstorfer, 1899; = lotina Fruhstorfer, 1904; = umbrosus Fruhstorfer, 1906; = philozigetes Fruhstorfer, 1907; = mezentius Fruhstorfer, 1910; = meridionigra Martin, 1913; = edwardi Van Eecke, 1914; = insularis Moulton, 1921; = albescens Röber, 1927; = kotoshonis Matsumara, 1929; = camorta Evans, 1932)
  • Danaus eresimus (Cramer, 1777) ( original name = Papilio eresimus Cramer, 1777; = ares d'Almeida, 1944; = asclepidea Fabricius, 1938; = dilucida Forbes, 1939; = erginus Godman & Salvin, 1897; = estevana Talbot, 1943; = icensis Fuchs, 1954; = montezuma Talbot, 1943; = tethys Forbes, 1943)
  • Danaus plexaure (Godart, 1819) ( original name = Danais plexaure Godart, 1819; = brasiliensis Capronnier, 1874)
  • Danaus gilippus (Cramer, 1775) ( original name = Papilio gilippus Cramer, 1775; = berenice Cramer, 1779; = vincedoxici Hübner, 1816; = vincetoxici Hübner, 1816; = cleothera Godart, 1819; = manuja Eschscholtz, 1821; = xanthippus Felder & Felder, 1860; = thersippus Bates, 1863; = jamaicensis Bates, 1864; = strigosa Bates, 1864; = hermippus Felder & Felder, 1865; = nivosus Godman & Salvin, 1897; = esperanza Hoffman, 1924; = centralis Joicey & Talbot, 1925; = kempfferi Hall, 1925; = kerri Comstock, 1925; = vincetoxici Fabricius, 1938; = candidus Clark, 1941; = gilippina Hoffman, 1940; = wheeleri Talbot, 1943; = roedingeri Fuchs, 1954)
  • Danaus chrysippus (Linnaeus, 1758) ( original name = Papilio chrysippus Linnaeus, 1758; = albinus Lanz, 1896; = amplifascia Talbot, 1943; = anomala Dufrane, 1948; = asclepiadis Gagliardi, 1811; = auriflava Van Eecke, 1914; = axantha Hayward, 1922; = bataviana Moore, 1883; = bipunctata Dufrane, 1948; = bowringi Moore, 1883; = candidata Hayward, 1922; = chrysipellus Strand, 1910; = clarippus Weymer, 1884; = completa Dufrane, 1948; = cratippus Felder, 1860; = deficiens Dufrane, 1948; = duplicata Dufrane, 1948; = duponti Dufrane, 1948; = evanescens Storace, 1949; = fuscippus Van Eecke, 1915; = gelderi Snellen, 1891; = hanoiensis Dufrane, 1948; = hypermnestra Stoneham, 1958; = impunctata Dufrane, 1948; = infumata Aurivillius, 1898; = joannisi Dufrane, 1948; = kanariensis Fruhstorfer, 1898; = lemeemagdalenae Lemeé, 1950; = limbata Matsumura, 1929; = luxurians Dufrane, 1948; = margarita Röber, 1926; = orientis Aurivillius, 1909; = ornata Dufrane, 1948; = petilia Stoll, 1790; = praealbata Froreich, 1928; = pseudopetilea Kalis, 1933; = radiata Dufrane, 1948; = reducta Dufrane, 1948; = rubra Van Eecke, 1915; = semialbinus Strand, 1910; = subpurpurea Matsumura, 1929; = subreducta Dufrane, 1948; = vigelii Heylaerts, 1884; = witteellus Overlaet, 1955; = yuchingkini Murayama & Shimonoya, 1960 (hybrid D. chrysippus x D. genutia)
    • Danaus chrysippus chrysippus (Linnaeus, 1758)
    • Danaus chrysippus aegyptius (Schreber, 1759) ( original name = Papilio aegyptius Schreber, 1759; = Danaus chrysippus klugii Butler, 1886; = Papilio alcippus Cramer, 1777; = Danaus chrysippus liboria Hulstaert, 1931; = Danaus chrysippus alcippoides Moore, 1883; = Danaus chrysippus dorippus Klug, 1845; = Danaus chrysippus transiens Suffert, 1900)

References

• Ackery, P. & Vane-Wright, R. (1984). Milkweed Butterflies: Their Cladistics and Biology. London: British Museum (Natural History.
• Jiggins, F. M.; Hurst, G. D. D.; Jiggins, C. D.; Schulenburg, J. H. G. v. D. & Majerus, M. E. N. (2000). The butterfly Danaus chrysippus is infected by a male-killing Spiroplasma bacterium. Parasitology 120(5): 439–446. doi:10.1017/S0031182099005867 (HTML abstract)
• Lushai, G., Goulson, D., Allen, J. A., Maclean, N., Smith, D. A. S., & Gordon, I. J. (2003). Incomplete sexual isolation in sympatry between subspecies of the butterfly Danaus chrysippus (L.) and the creation of a hybrid zone. Heredity, 90(3), 236-246.
• Smith, D. A. S., Gordon, I. J., Lushai, G., Goulson, D., Allen, J. A., & Maclean, N. (2002). Hybrid queen butterflies from the cross Danaus chrysippus (L.) x D. gilippus (Cramer): Confirmation of species status for the parents and further support for Haldane's Rule. Biological Journal of the Linnean Society, 76(4), 535-544.
• Smith, D. A. S., Lushai, G., & Allen, J. A. A. (2005). A classification of Danaus butterflies (Lepidoptera: Nymphalidae) based upon data from morphology and DNA. Zoological Journal of the Linnean Society, 144(2), 191-212.