Taxonomic catalogue of Heliconiinae

06 January 2025

Order: Lepidoptera Linnaeus, 1758

Family: Nymphalidae Rafinesque, 1815

Subfamily: Heliconiinae Swainson, 1822

Resources:

• Butterflies of America
• A Catalogue of the Butterflies of the United States and Canada by Jonathan P. Pelham
• Lamas’s Annotated Bibliography of the Neotropical Butterflies and Skippers
• The Global Lepidoptera Names Index @ Natural History Museum, London
• Butterflies and Moths of the World @ Natural History Museum, London
• nymphalidae.net
• Markku Savela’s list
• African Butterfly Database (ABDB) by Szabolcs Sáfián et al.
• Afrotropical butterflies by Mark C. Williams

Tribe Acraeini

Type genus: Acraea Fabricius, 1807

Classification: The tribe Acraeini is divided into eight genera (Williams and Hennig 2023): Acraea, Actinote, Bematistes, Cethosia, Rubraea, Stephenia, Telchinia and Tildia.

References: Penz and Peggie (2003) (phylogeny); Vane-Wright and Jong (2003) (Sulawesi); Silva-Brandão et al. (2008) (phylogeny); Henning and Williams (2010) (taxonomy); Timmermans et al. (2016) (phylogeny); Pierre (2020 a) (Uganda); Lang et al. (2022) (taxonomy; China); Williams and Hennig (2023) (taxonomy)

Genus Acraea

Genus: Acraea Fabricius, 1807

Type species: Acraea horta Linnaeus, 1764; original combination: Papilio horta Linnaeus, 1764

Synonyms:

• Aphanopeltis Mabille, 1887. Type species: Acraea horta Linnaeus, 1764; original combination: Papilio horta Linnaeus, 1764. Refs.: Mabille (1886): 85
• Phanopeltis Mabille, 1887. Type species: Acraea ranavalona Boisduval, 1833; original combination: Acraea ranavalona Boisduval, 1833. Refs.: Mabille (1886): 84
• Solenites Mabille, 1887. Type species: Acraea igati Boisduval, 1833; original combination: Acraea igati Boisduval, 1833. Refs.: Mabille (1886): 82
• Miyana Fruhstorfer, 1914. Type species: Acraea moluccana Felder, 1860; original combination: Acraea moluccana Felder, 1860. Refs.: Fruhstorfer (1912 a): 743

References: Hennig (1993 a); Williams and Hennig (2010); Pierre and Bernaud (2012) (taxonomy, phylogeny); Pierre and Bernaud (2013); Williams and Hennig (2020); Williams and Hennig (2023) (taxonomic revision)

32 species:

• admatha Hewitson [1865]; original combination: Acraea admatha

  • syn. mildbraedi Schultze, 1917; original combination: Acraea admatha ♀ ab. mildbraedi.

• andromacha (Fabricius, 1775)

  • ssp. andromacha (Fabricius, 1775); original combination: Papilio andromacha Distr.: Australia.

    • syn. entoria Godart, 1819

  • ssp. indica Röber, 1885; original combination: Acraea andromacha var. indica

  • ssp. oenone Kirby, 1889; original combination: Acraea oenone

  • ssp. polynesiaca Rebel, 1911; original combination: Acraea andromache[sic] polynesiaca

    • syn. agema Fruhstorfer, 1907

  • ssp. sanderi Rothschild, 1893; original combination: Acraea sanderi

• boopis Wichgraf, 1914

  • ssp. ama Pierre, 1979; original combination: Acraea boopis ama

  • ssp. boopis Wichgraf, 1914; original combination: Acraea boopis

  • ssp. choloui Pierre, 1979; original combination: Acraea boopis choloui

• brainei Henning, 1986; original combination: Acraea brainei

• camaena (Drury, 1773); original combination: Papilio camaena

  • syn. murcia (Fabricius, 1781); original combination: Papilio murcia.

• cuva Grose-Smith, 1889; original combination: Acraea cuva

  • syn. nidama Suffert, 1904; original combination: Acraea dammi nidama.

• dammii Vollenhoven, 1869; original combination: Acraea dammii

  • syn. percussa Keferstein, 1870; original combination: Acraea percussa.

  • syn. masonala Ward, 1872; original combination: Acraea masonala.

  • syn. villettei Oberthür, 1925; original combination: Acraea villettei.

• eltringhami Joicey & Talbot, 1921; original combination: Acraea eltringhami

• endoscota Le Doux, 1928; original combination: Acraea admatha endoscota

  • syn. albanis Pierre, 1979; original combination: Acraea endoscota ♀ f. albanis.

• eugenia Karsch, 1893

  • ssp. eugenia Karsch, 1893; original combination: Acraea eugenia

  • ssp. ochreata Grünberg, 1910; original combination: Acraea eugenia var. ochreata

• hamata Joicey & Talbot, 1922

  • ssp. batangi Bernaud, Ducarme & Pierre, 2017; original combination: Acraea (Acraea) hamata batangi

  • ssp. hamata Joicey & Talbot, 1922; original combination: Acraea hamata

• horta (Linnaeus, 1764); original combination: Papilio horta

  • syn. conjuncta Blachier, 1912; original combination: Acraea horta ab. conjuncta.

• hova Boisduval, 1833; original combination: Acraea hova Refs.: Boisduval (1833 a): 29, pl.4.

• igati Boisduval, 1833; original combination: Acraea igati

• insignis Distant, 1880

  • ssp. gorongozae Van Son, 1963; original combination: Acraea insignis gorongozae

  • ssp. insignis Distant, 1880; original combination: Acraea insignis

    • syn. buxtoni Hewitson, 1877

    • syn. balbina Oberthür, 1888

    • syn. siginna Suffert, 1904

• kappa Pierre, 1979; original combination: Acraea kappa

• kia Pierre, 1990; original combination: Acraea kia

• kinduana Pierre, 1979; original combination: Acraea kinduana

  • syn. kinduana Schouteden, 1919; original combination: Acraea admatha ab. kinduana.

• leucographa Ribbe, 1889

  • ssp. jolyi Pierre, 2009; original combination: Acraea leucographa jolyi. Type loc.: Ghana. Distr.: Ghana; Ivory Coast; Nigeria.

  • ssp. leucographa Ribbe, 1889; original combination: Acraea leucographa

    • syn. gyldenstolpei Aurivillius, 1925

    • syn. sinalba Pierre, 1979

• machequena Grose-Smith, 1887; original combination: Acraea machequena

• magnifica Carpenter & Jackson, 1950; original combination: Acraea chilo magnifica

• mahela Boisduval, 1833; original combination: Acraea mahela

• matuapa Grose-Smith, 1889; original combination: Acraea matuapa

• meyeri Kirsch, 1877; original combination: Acraea meyeri

• moluccana Felder, 1860

  • ssp. buruensis Rothschild, 1899; original combination: Acraea parce buruensis

  • ssp. dohertyi Holland, 1891; original combination: Acraea dohertyi

  • ssp. moluccana Felder, 1860; original combination: Acraea moluccana

    • syn. fumigata Honrath, 1886

  • ssp. nebulosa Hewitson, 1861; original combination: Acraea nebulosa

  • ssp. parce Staudinger, 1896; original combination: Acraea parce

  • ssp. pella Fruhstorfer, 1907; original combination: Acraea moluccana pella

  • ssp. pollonia Godman & Salvin, 1888; original combination: Acraea pollonia

• neobule Doubleday [1847]

  • ssp. legrandi Carcasson, 1964; original combination: Acraea terpsichore legrandi

  • ssp. neobule Doubleday [1847]; original combination: Acraea neobule

    • syn. seis Feisthamel, 1850

    • syn. calyce Godman & Salvin, 1884

    • syn. mhondana Vuillot, 1891

    • syn. arabica Rebel, 1899

    • syn. socotrana Rebel, 1907

    • syn. zambesina Aurivillius, 1909

    • syn. braesioides Wichgraf, 1914

    • syn. guttata Wichgraf, 1914

    • syn. incredibilis Le Doux, 1922

    • syn. camaenopsis Le Doux, 1923

    • syn. cyaniris Le Cerf, 1927

    • syn. kibwezina Le Doux, 1928

    • syn. isabellina Stoneham, 1943

    • syn. didalis Stoneham, 1943

    • syn. montana Stoneham, 1943

    • syn. macra Storace, 1949

    • syn. pallidepicta Storace, 1949

    • syn. sheba Gabriel, 1954

    • syn. melanica Woodhall, 2000

• punctimarginea Pinhey, 1956; original combination: Acraea punctimarginea

• quirina (Fabricius, 1781)

  • ssp. quirina (Fabricius, 1781); original combination: Papilio quirina

    • syn. bourgeoni Schouteden, 1919

  • ssp. rosa Eltringham, 1912; original combination: Acraea quirina rosa

• ranavalona Boisduval, 1833; original combination: Acraea ranavalona

  • syn. maransetra Ward, 1872; original combination: Acraea maransetra. Type loc.: Madagascar.

  • syn. manandaza Ward, 1872; original combination: Acraea manandaza. Type loc.: Madagascar.

• satis Ward, 1871; original combination: Acraea satis

  • syn. corona Staudinger, 1885; original combination: Acraea corona.

  • syn. donatis Woodhall, 2000; original combination: Acraea satis f. donatis.

• terpsicore (Linnaeus, 1758); original combination: Papilio terpsicore

  • syn. violae (Fabricius, 1793 [1775]); original combination: Papilio violae. Refs.: Lang et al. (2022).

  • syn. cephea (Cramer, 1780); original combination: Papilio cephea.

• turlini Pierre, 1979; original combination: Acraea turlini

Genus Actinote

Genus: Actinote Hübner [1819]

Type species: Actinote thalia (Linnaeus, 1758); original combination: Papilio thalia Linnaeus, 1758

Synonyms:

• Calornis Billberg, 1820. Type species: Actinote thalia (Linnaeus, 1758); original combination: Papilio thalia Linnaeus, 1758. Refs.: Billberg (1820): 77
• Abananote Potts, 1943. Type species: Actinote abana (Hewitson, 1868); original combination: Acraea abana Hewitson, 1868. Refs.: Potts (1943): 31 (as subgenus)
• Altinote Potts, 1943. Type species: Actinote neleus (Latreille, [1813]); original combination: Heliconius neleus Latreille, [1813]. Refs.: Potts (1943): 31

References: Hübner ([1819]): 27

59 species:

• abana (Hewitson, 1868)

  • ssp. abana (Hewitson, 1868); original combination: Acraea abana

  • ssp. capnodes Jordan, 1913; original combination: Actinote abana capnodes

• alalia (Felder & Felder, 1860); original combination: Acraea alalia Refs.: Felder 1860:10.5; Kirby 1871- 77:136; Jones 1883:.36; Jones 1883:8; Fritz MUller 1883:216; Mabille 1896:64.

  • syn. eulalia (Fritz Müller, 1878). Refs.: Fritz Müller 1878:296.

  • syn. alalia alalia Eltringham & Jordan, 1913. Refs.: Eltringham and Jordan 1913:10; Jordan 1913b:374.

  • syn. alalia Jordan & Eltringham, 1916. Refs.: Eltringham 1916:19; d’Almeida 193.5a:93; Ebert 1969:3.5; Lewis 197.5:43, 227; Smart 1976:26.5; d’Abrera 1987:44.5; Ackery 1988:136; Francini 1892:16 fig.6-14 [ALA].

  • syn. alalia (Pierre, 1987). Refs.: Pierre 1987:22.

• alberti Neild & Romero, 2008

  • ssp. alberti Neild & Romero, 2008; original combination: Actinote alberti. Type loc.: ~Km 30 entrance to La Mina, Via Barinitas, Sto. Domingo, Barinas, Venezuela. Types: MIZA. Distr.: Cordillera de Mérida. Refs.: Neild (2008): 34. Ety.: Albert Gross (collector)

  • ssp. skinneri Romero & Neild, 2008; original combination: Actinote alberti skinneri. Type loc.: Via Delicia 2000 IV-80. Types: CFR. Distr.: northwestern slope of Sierra de El Tamá. Refs.: Neild (2008): 36. Ety.: Skinner family

• alcione (Hewitson, 1852)

  • ssp. alcione (Hewitson, 1852); original combination: Acraea alcione

  • ssp. corduba (Hewitson, 1874); original combination: Acraea corduba

  • ssp. elatus Druce, 1903; original combination: Actinote elatus

    • syn. sarsanda Druce, 1903

    • syn. alcione subelatus Jordan, 1913

  • ssp. flaugeri Neild & Romero, 2008. Type loc.: Bramón a Delicias, Km 39, Tachira, Venezuela. Types: MIZA. Distr.: Sierra del El Tamá, Venezuela. Refs.: Neild (2008): 43. Ety.: Flauger family

  • ssp. miguelangeli Neild & Romero, 2008. Type loc.: Qda. La Chorrera. Via Merida-Jaji. Estado Merida [Venezuela]. Types: MIZA. Distr.: western Cordillera de Mérida, Venezuela. Refs.: Neild (2008): 42. Ety.: Andrés Orellana

  • ssp. sodalis Butler, 1877; original combination: Actinote sodalis. Type loc.: Peru.

    • syn. corduba messeres Jordan, 1910. Type loc.: Carabaya, Peru.

    • syn. alcyone salmonea Jordan, 1910. Type loc.: Huayabamba River, SE of Chachapoyas, Peru.

    • syn. incarum Oberthür, 1917. Type loc.: Peru.

    • syn. alcione leucothoe (Potts, 1944). Type loc.: Peru.

    • syn. alcione hermosa (Potts, 1944). Type loc.: Peru.

  • ssp. theophila (Dognin, 1887); original combination: Acraea theophila. Type loc.: Ecuador, Zamora.

    • syn. alcyone melina Jordan, 1910. Type loc.: “Brazil(Am)”.

    • syn. alcione mengeli (Potts, 1944). Type loc.: Peru.

    • syn. alcione bassleri (Potts, 1944). Type loc.: Peru.

    • syn. alcione moneta (Potts, 1944). Type loc.: Ecuador, 2-4°S, 78°W.

  • ssp. varians Jordan, 1910; original combination: Actinote alcyone varians. Type loc.: W. Colombia, Rio Dagua.

    • syn. f. cyanea Jordan, 1913

    • syn. f. extensa Jordan, 1913

• anaxo (Hopffer, 1874)

  • ssp. anaxo (Hopffer, 1874); original combination: Acraea anaxo

  • ssp. comta Jordan, 1910; original combination: Actinote anaxo comta

• ballettae Neild & Romero, 2008

  • ssp. ballettae Neild & Romero, 2008; original combination: Actinote ballettae. Type loc.: Río Frío 600m IV-88. Types: CFR. Distr.: eastern slope of Sierra de El Tamá, eastern slope of Cordillera de Mérida, Cordillera de la Costa [Venezuela]. Refs.: Neild (2008): 32. Ety.: Balletta family (contributor of the Romero Family Collection)

  • ssp. bonyae Neild & Romero, 2008; original combination: Actinote ballettae bonyae. Type loc.: La Fría 600m IV-91. Types: CFR. Distr.: western slope of Cordillera de Mérida [Venezuela]. Refs.: Neild (2008): 33. Ety.: Bony (Claire R. Hing King), the late wife of Francisco Fernández Yépez (entomologist)

• bonita Penz, 1996; original combination: Actinote bonita. Type loc.: Minas Gerais, Monteverde, Camanducaia, Brazil. Types: UFPC. Distr.: southeastern Brazil highlands. Host plants: Eupatorium intermedium (Asteraceae) (Penz and Francini 1996). Refs.: Penz and Francini (1996): 315. Ety.: Bonita (a song by Tom Jobim)

• callianthe (Felder & Felder, 1862)

  • ssp. alla Jordan, 1910; original combination: Actinote callianthe alla

  • ssp. amoena Jordan, 1913; original combination: Actinote callianthe amoena

  • ssp. borgesi Neild & Romero, 2008. Type loc.: 1.900M. Via Delicia, P.N., El Tama, Tachira, Venezuela. Types: MIZA. Distr.: Sierra del El Tamá, Venezuela. Refs.: Neild (2008): 40. Ety.: Gustavo Borges (entomologist)

  • ssp. callianthe (Felder & Felder, 1862); original combination: Acraea callianthe

  • ssp. rufa Jordan, 1910; original combination: Actinote callianthe rufa

• canutia (Hopffer, 1874); original combination: Acraea canutia

• carycina Jordan, 1913; original combination: Actinote carycina Host plants: Eupatorium inulaefolium (Asteraceae) (Paluch et al. 2001). Refs.: Paluch et al. (2001) (immature).

  • syn. lorida Oberthür, 1917; original combination: Actinote lorida. Refs.: Paluch (2006) (synonymy).

  • syn. carycinoides d’Almeida, 1935; original combination: Actinote pellenea var. carycinoides. Refs.: Paluch (2006) (synonymy).

  • syn. var. hoffmanni d’Almeida, 1935; original combination: Actinote carycina var. hoffmanni.

  • syn. var. differens d’Almeida, 1935; original combination: Actinote carycina var. differens.

• catarina Penz, 1996; original combination: Actinote catarina. Type loc.: Santa Catarina, Santa Cecilia, Brazil. Distr.: southeastern Brazil highlands. Refs.: Penz and Francini (1996): 314. Ety.: Santa Catarina (a state in Brazil)

• conspicua Jordan, 1913; original combination: Actinote alalia conspicua Host plants: Mikania hirsutissima (Penz and Francini 1996).

  • syn. conspicua Ebert, 1969. Refs.: Ebert 1969:35; Francini 1992:16 fig.6-14 [CON].

• dalmeidai Francini, 1996; original combination: Actinote dalmeidai. Type loc.: Paraná, Sao Jose dos Pinhais, Brazil. Distr.: southeastern Brazil highlands. Refs.: Penz and Francini (1996): 310.

• dicaeus (Latreille [1817])

  • ssp. albofasciata (Hewitson, 1869); original combination: Acraea albofasciata

    • syn. rosaria (Weymer, 1890)

    • syn. crina (Schaus, 1902)

    • syn. dice napensis Jordan, 1910

  • ssp. amida (Hewitson, 1854); original combination: Acraea amida

    • syn. theogonia (Weymer, 1890)

  • ssp. callianira Geyer, 1837; original combination: Actinote callianira

    • syn. callianira stenia Jordan, 1910

  • ssp. carica (Weymer, 1890); original combination: Acraea stratonice var. carica

    • syn. thespias (Weymer, 1890)

    • syn. dicaeus var. pallasia Oberthür, 1917

    • syn. dicaeus var. pallasia subvar. pallasinia Oberthür, 1917

  • ssp. dicaeus (Latreille [1817]); original combination: Heliconius dicaeus

    • syn. dice (Godart, 1819), preocc. (not Drury, 1782), repl. name

    • syn. diceus var. aurantia (Weymer, 1890)

    • syn. olgae Rebel, 1902

    • syn. diceus diceus f. sinefascia Jordan, 1913

    • syn. diceus diceus f. confluens Jordan, 1913

    • syn. diceus diceus f. rubrosticta Jordan, 1913

  • ssp. eulalia Oberthür, 1917; original combination: Actinote flavibasis var. eulalia Refs.: Fritz Miiller 1878:296.

    • syn. flavibasis var. eulogia Oberthür, 1917

  • ssp. euris Jordan, 1910; original combination: Actinote callianira euris

  • ssp. jucunda Jordan, 1910; original combination: Actinote jucunda

  • ssp. laverna (Doubleday [1847]); original combination: Acraea laverna

  • ssp. luisi Neild & Romero, 2008. Type loc.: Monte Zerpa, anove La Hechicera, Merida, Venezuela. Types: MIZA. Distr.: Cordillera de Mérida, Venezuela. Refs.: Neild (2008): 45. Ety.: Luis Colmenares (La Victoria)

  • ssp. ozinta (Schaus, 1902); original combination: Acraea ozinta

    • syn. ozomene var. granadina Rebel, 1902

    • syn. diceus flavibasis Jordan, 1913

    • syn. flavibasis ab. fasciata Jordan, 1913

• discrepans d’Almeida, 1958, ICZN, Art. 45.6.2; original combination: Actinote discrepans

  • syn. parapheles ab. discrepans d’Almeida, 1935; original combination: Actinote parapheles ab. discrepans.

• eberti Francini, Freitas & Penz, 2004; original combination: Actinote eberti. Type loc.: southeastern Brazil (Itatiaia, Rio de Janeiro & Campos do Jordão, São Paulo). Types: Departamento de Zoologia, Universidade Federal do Paraná, Brazil. Distr.: Itatiaia and Campos do Jordão, Brazil. Refs.: Francini et al. (2004): 2. Ety.: Heinz Ebert

• eresia (Felder & Felder, 1862)

  • ssp. albesia Lamas & Willmott, 2024; original combination: Actinote eresia albesia Refs.: Willmott et al. (2024): 8.

  • ssp. canyaris Pyrcz & Lamas, 2024; original combination: Actinote eresia canyaris Refs.: Willmott et al. (2024): 9.

  • ssp. eresia (Felder & Felder, 1862); original combination: Acraea eresia

  • ssp. eresina (Hopffer, 1874); original combination: Acraea eresina

    • syn. binghamae Dyar, 1913 Refs.: Willmott et al. (2024) (synonymy).

  • ssp. lacrymosa Oberthür, 1917; original combination: Actinote lacrymosa

  • ssp. leptogramma Jordan, 1913; original combination: Actinote eresia leptogramma

• erinome (Felder & Felder, 1861)

  • ssp. erinome (Felder & Felder, 1861); original combination: Acraea erinome

    • syn. ab. sciana Jordan, 1910

    • syn. erinome carabaia Jordan, 1910, hyb.

  • ssp. mathani Oberthür, 1917; original combination: Actinote mathani

    • syn. carmentis (Potts, 1944)

    • syn. melini Bryk, 1953

  • ssp. testacea (Salvin & Godman, 1868); original combination: Acraea testacea

    • syn. byzia (Hewitson, 1874)

    • syn. aethilla (Hopffer, 1874)

• euryleuca Jordan, 1910; original combination: Actinote euryleuca

• furtadoi Paluch, Casagrande & Mielke, 2006; original combination: Actinote furtadoi. Type loc.: Alto Rio Arinos, Faz. S. João, Diamantino, MT [Brazil]. Types: UFPC. Distr.: Mato Grosso, Diamantino, Brazil. Refs.: Paluch et al. (2006): 772. Ety.: Eurides Furtado

• genitrix d’Almeida, 1922

  • ssp. genitrix d’Almeida, 1922; original combination: Actinote genitrix

    • syn. ab. subrubicunda d’Almeida, 1925 Refs.: Paluch (2006).

    • syn. ab. moesa d’Almeida, 1925

  • ssp. costae Neild, 2008. Type loc.: La Escalera, Km 123 (1400 m), Estado Bolivar. Types: MIZA. Distr.: Sierra de Lema. Refs.: Neild (2008): 39. Ety.: Mauro Costa

• griseata Butler, 1873; original combination: Actinote griseata

  • syn. aliteria (Hopffer, 1874); original combination: Acraea aliteria.

• guatemalena (Bates, 1864)

  • ssp. guatemalena (Bates, 1864); original combination: Acraea guatemalena

  • ssp. guerrerensis J. Maza, 1982; original combination: Actinote guatemalena guerrerensis

  • ssp. veraecrucis Jordan, 1913; original combination: Actinote guatemalena veraecrucis

• hilaris Jordan, 1910

  • ssp. arcoiris Willmott & Hall, 2024; original combination: Actinote hilaris arcoiris Refs.: Willmott et al. (2024): 9.

  • ssp. desmiala Jordan, 1913; original combination: Actinote desmiala

    • syn. culoti Oberthür, 1916

    • syn. brownorum (Potts, 1943)

  • ssp. hilaris Jordan, 1910; original combination: Actinote hilaris

  • ssp. sourakovi Willmott & Lamas, 2024; original combination: Actinote hilaris sourakovi Refs.: Willmott et al. (2024): 12.

• hylonome (Doubleday, 1844)

  • ssp. hylonome (Doubleday, 1844); original combination: Euterpe hylonome Distr.: Sierras de Perijá and El Tamá, Cordillera de Mérida. Refs.: Neild (2008): 46 (Venezuela).

  • ssp. basilia (Lamas, 1995); original combination: Abananote hylonome basilia Distr.: Cordillera de La Costa, Serranía del Interior. Refs.: Neild (2008): 46 (Venezuela).

• intensa Jordan, 1910; original combination: Actinote radiata intensa Refs.: Willmott et al. (2017): 7 (stat. rest.).

  • syn. - (Hewitson, 1868), preocc.; original combination: Acraea radiata. Refs.: Willmott et al. (2017): 7 (stat. rest.).

  • syn. - (Kemal & Koçak, 2007); original combination: Acraea (Abananote) intensa gerardolamas. Refs.: Willmott et al. (2017): 7 (stat. rest.).

• johncoulsoni Willmott, Lamas & Hall, 2017; original combination: Actinote johncoulsoni. Type loc.: Zamora-Chinchipe, Ecuador. Types: FLMNH. Refs.: Willmott et al. (2017): 9. Ety.: Kenneth John Coulson

• keithbrowni Freitas, Francini & Mielke, 2020. Type loc.: Itatiaia, Rio de Janeiro, Brazil. Types: UFPC; paratypes: ZUEC. Distr.: Itatiaia massif, southeastern Brazil. Refs.: Freitas et al. (2020): 697. Ety.: Keith S. Brown Jr.

• kennethi Freitas, Willmott & Hall, 2009; original combination: Actinote kennethi. Type loc.: Morona-Santiago, Río Abanico, Ecuador. Types: FLMNH; paratypes: MECN, KWJH. Distr.: eastern Ecuador. Refs.: Willmott et al. (2009): 49. Ety.: James Kenneth Robinson

• lapitha (Staudinger, 1885)

  • ssp. calderoni Schaus, 1920; original combination: Actinote calderoni

    • syn. lapitha zilchi Franz & Schröder, 1954

  • ssp. lapitha (Staudinger, 1885); original combination: Acraea lapitha

• latior Jordan, 1913; original combination: Actinote guatemalena latior

  • syn. - Jordan, 1913; original combination: Actinote anteas f. straminosa. Refs.: Paluch (2006) (synonymy).

• mamita (Burmeister, 1861)

  • ssp. elena Hall, 1921; original combination: Actinote elena Refs.: Paluch (2006) (synonymy).

  • ssp. mamita (Burmeister, 1861); original combination: Acrea[sic] mamita

  • ssp. mitama (Schaus, 1902); original combination: Acraea mitama

• mantiqueira Freitas, Francini, Paluch & Barbosa, 2018; original combination: Actinote mantiqueira. Type loc.: Pindamonhangaba, São Paulo, Brazil. Types: ZUEC. Host plants: Eupatorium intermedium (Asteraceae) (Penz and Francini 1996; Freitas et al. 2018). Refs.: Freitas et al. (2018): 136.

• melampeplos Godman & Salvin [1881]; original combination: Actinote melampeplos

• melanisans Oberthür, 1917; original combination: Actinote bubona melanisans

  • syn. travassosi d’Almeida, 1934; original combination: Actinote travassosi. Refs.: Paluch (2006).

• mielkei Paluch & Casagrande, 2006; original combination: Actinote mielkei. Type loc.: Chulumani, Bolivia. Types: UFPC. Distr.: Chulumani, Bolivia. Refs.: Paluch et al. (2006): 770. Ety.: Olaf Hermann Hendrik Mielke

• mirnae Paluch & Mielke, 2006; original combination: Actinote mirnae. Type loc.: Chulumani, Bolivia. Types: UFPC. Distr.: Chulumani, Bolivia. Refs.: Paluch et al. (2006): 767. Ety.: Mirna Martins Casagrande

• momina Jordan, 1910; original combination: Actinote momina

• morio Oberthür, 1917

  • ssp. beckeri Paluch, Casagrande & Mielke, 2006; original combination: Actinote morio beckeri. Type loc.: Planaltina, DF, Brazil. Types: UFPC. Refs.: Paluch et al. (2006): 773. Ety.: Vitor Osmar Becker

  • ssp. morio Oberthür, 1917; original combination: Actinote morio

• negra (Felder & Felder, 1862)

  • ssp. avuncula Oberthür, 1917; original combination: Actinote euclia avuncula

  • ssp. demonica (Hopffer, 1874); original combination: Acraea demonica

    • syn. nicylla (Hopffer, 1874)

    • syn. demonica demonica f. aurantia Jordan, 1913

    • syn. demonica demonica f. intermedia Jordan, 1913

    • syn. demonica var. nicylloides Strand, 1916

    • syn. demonica var. nicylloides ab. flavitincta Strand, 1916

    • syn. subdemonica Strand, 1916

  • ssp. euclia (Dognin, 1887); original combination: Acraea euclia

    • syn. leontine (Weymer, 1890)

    • syn. leontine catochaera Jordan, 1910

    • syn. leontine hypsipetes Jordan, 1910

    • syn. leontine mesia Jordan, 1910

    • syn. leontine bulis Jordan, 1913

  • ssp. menoetes Oberthür, 1917; original combination: Actinote menoetes

  • ssp. negra (Felder & Felder, 1862); original combination: Acraea negra

    • syn. moyobambae Oberthür, 1917

  • ssp. scotosis Jordan, 1910; original combination: Actinote nicylla scotosis

    • syn. amphilecta Jordan, 1913

  • ssp. sobrina Jordan, 1910; original combination: Actinote nicylla sobrina

• neleus (Latreille [1813])

  • ssp. bermudezi Neild & Romero, 2008. Type loc.: Barinitas, Barinas, Venezuela. Types: MIZA. Distr.: Sierra del El Tamá (?), central-eastern and northeastern slopes of Corfillera de Mérida, Venezuela. Refs.: Neild (2008): 44. Ety.: Pedro J. Nermúdez

  • ssp. neleus (Latreille [1813]); original combination: Heliconius neleus

    • syn. ara (Hewitson, 1877)

    • syn. edulis (Weymer, 1890)

    • syn. ab. haemera Jordan, 1913

• ozomene (Godart, 1819)

  • ssp. calimene Rebel, 1902; original combination: Actinote ozomene var. calimene

    • syn. ozomene alicunde (Potts, 1944)

  • ssp. carbonaria (Hopffer, 1874); original combination: Acraea carbonaria

  • ssp. cleasa (Hewitson, 1868); original combination: Acraea cleasa

    • syn. ozomene catopasta Jordan, 1910

  • ssp. gabrielae Rebel, 1902; original combination: Actinote ozomene var. gabrielae

    • syn. ozomene var. cleasa ab. zarayaquilionis Strand, 1916

  • ssp. nox (Bates, 1864); original combination: Acraea nox

    • syn. leucomelas (Bates, 1864)

    • syn. orizava (Reakirt, 1866)

  • ssp. ozomene (Godart, 1819); original combination: Acraea ozomene. Type loc.: S. Amer. [West Colombia].

    • syn. ab. reducta Jordan, 1913

• pallescens Jordan, 1913; original combination: Actinote parapheles f. pallescens Refs.: Williams and Hennig (2023) (to species).

  • syn. brylla Oberthür, 1917; original combination: Actinote brylla. Refs.: Paluch (2006) (synonymy).

  • syn. brasiliensis var. notabilis d’Almeida, 1935; original combination: Actinote brasiliensis var. notabilis. Refs.: Paluch (2006) (synonymy).

• parapheles Jordan, 1913

  • ssp. ochreana Hayward, 1931; original combination: Actinote parapheles f. ochreana Refs.: Paluch (2006) (ssp. rev.).

  • ssp. parapheles Jordan, 1913; original combination: Actinote parapheles

    • syn. distincta d’Almeida, 1922

    • syn. var. fumida d’Almeida, 1925

    • syn. var. hippea d’Almeida, 1925

    • syn. ab. umbrata d’Almeida, 1935

• pellenea Hübner [1819]

  • ssp. adriana (Hopffer, 1874); original combination: Acraea adriana

    • syn. equatoria grammica Jordan, 1913

  • ssp. auloeda Oberthür, 1917; original combination: Actinote auloeda Refs.: Paluch (2006) (ssp. rev.).

    • syn. pellenea nordestina d’Almeida, 1935 Refs.: Paluch (2006) (synonymy).

  • ssp. calymma Jordan, 1913; original combination: Actinote pellenea calymna

    • syn. f. clara Jordan, 1913

    • syn. zaratensis Oberthür, 1917

    • syn. pellenea f. calchaqui Hayward, 1931

    • syn. pellenea f. quasicinerea Hayward, 1931

    • syn. pellenea f. venata Hayward, 1931

    • syn. pellenea f. calymmoides Hayward, 1935

    • syn. pellenea ab. alzaquita Biezanko, Ruffinelli & Carbonell, 1962, nom. nud.

  • ssp. epiphaea Jordan, 1913; original combination: Actinote equatoria epiphaea

  • ssp. equatoria (Bates, 1864); original combination: Acraea equatoria

  • ssp. fernandezi Neild & Romero, 2008; original combination: Actinote pellenea fernandezi. Type loc.: Sta. Elena to Icabaru Road, Bolívar, Venezuela. Types: MIZA. Distr.: Southeastern Bolívar [Venezuela]. Refs.: Neild (2008): 25. Ety.: Francisco Fernández Yépez

  • ssp. giffordi Paluch, Casagrande & Mielke, 2006; original combination: Actinote pellenea giffordi. Type loc.: Planaltina, DF, Brazil. Types: UFPC. Refs.: Paluch et al. (2006): 775. Ety.: David Gifford

  • ssp. hahneli Jordan, 1913; original combination: Actinote equatoria hahneli

    • syn. equatoria hahneli f. semilutea Jordan, 1913

    • syn. hahneli var. meridae Oberthür, 1917

    • syn. brauronia Oberthür, 1917

  • ssp. hyalina Jordan, 1913; original combination: Actinote hyalina

    • syn. pellenea subhyalina f. ferrugata Jordan, 1913

  • ssp. limbata Jordan, 1913; original combination: Actinote equatoria limbata

    • syn. equatoria limbata f. cauca Jordan, 1913

  • ssp. mucia (Hopffer, 1874); original combination: Acraea mucia

  • ssp. lodis Jordan, 1913; original combination: Actinote equatoria lodis. Type loc.: Bolivia.

    • syn. lolia Oberthür, 1917. Type loc.: Bolivia. Refs.: Paluch (2006) (synonymy).

  • ssp. pellenea Hübner [1819]; original combination: Actinote pellenea

  • ssp. perfulva Jordan, 1913; original combination: Actinote equatoria perfulva

    • syn. equatoria perfulva f. flavifascia Jordan, 1913

    • syn. brychia Oberthür, 1917

  • ssp. subbadia Jordan, 1913; original combination: Actinote pellenea subbadia

    • syn. byzas Oberthür, 1917

  • ssp. subhyalina (Staudinger, 1885); original combination: Acraea subhyalina

  • ssp. trinitatis Jordan, 1913; original combination: Actinote pellenea trinitatis

    • syn. byzas var. insularis Oberthür, 1917

• pratensis Francini, Freitas & Penz, 2004; original combination: Actinote pratensis. Type loc.: Bosque Municipal de Águas da Prata, São Paulo, Brazil. Types: MZUSP; paratypes: ZUEC. Distr.: southeastern Brazil (Águas da Prata, Rio Claro, Campinas, Jundiaí). Refs.: Francini et al. (2004): 5.

• pyrrha (Fabricius, 1775)

  • ssp. crucis Jordan, 1913; original combination: Actinote pellenea crucis

  • ssp. pyrrha (Fabricius, 1775); original combination: Papilio pyrrha Refs.: Francini and Penz (2006) (key).

    • syn. lacaena (Fabricius, 1793), nom. nud.

    • syn. euterpe (Felder, 1862), nom. nud.

    • syn. bubona Oberthür, 1917

    • syn. brasiliensis d’Almeida, 1922

    • syn. brasiliensis ab. magnifica d’Almeida, 1922

    • syn. jordani d’Almeida, 1922 Refs.: Paluch (2006).

    • syn. brasiliensis ab. fuliginosa d’Almeida, 1925 Refs.: Paluch (2006).

    • syn. brasiliensis var. discolora d’Almeida, 1935

• pyrrhosticta Lamas, Willmott & Hall, 2024

  • ssp. alfamayo Lamas & Willmott, 2024; original combination: Actinote pyrrhosticta alfamayo Refs.: Willmott et al. (2024): 8.

  • ssp. apurimac Lamas & Willmott, 2024; original combination: Actinote pyrrhosticta apurimac Refs.: Willmott et al. (2024): 6.

  • ssp. quintecocha Lamas & Willmott, 2024; original combination: Actinote pyrrhosticta quintecocha Refs.: Willmott et al. (2024): 5.

  • ssp. pyrrhosticta Lamas, Willmott & Hall, 2024; original combination: Actinote pyrrhosticta Refs.: Willmott et al. (2024): 2.

• quadra (Schaus, 1902); original combination: Acraea quadra

• rhodope d’Almeida, 1923; original combination: Actinote rhodope

  • syn. f. lynsa d’Almeida, 1923; original combination: Actinote rhodope f. lynsa.

  • syn. ab. tenuilimbata d’Almeida, 1931; original combination: Actinote rhodope ab. tenuilimbata.

  • syn. var. fuscata d’Almeida, 1935; original combination: Actinote rhodope var. fuscata.

• romeroi Neil & Costa, 2008; original combination: Actinote romeroi. Type loc.: Río Surukún 870 V-89. Types: CFR. Distr.: Upper Río Surukún valley in the extreme southeast of Bolívar state (approximately 30–50 km west of Santa Elena on the road to Icabarú). Refs.: Neild (2008): 37. Ety.: Francisco Romero Rodríguez (Venezuela)

• rubrocellulata Hayward, 1960; original combination: Actinote rubrocellulata

• rufina Oberthür, 1917; original combination: Actinote rufina Distr.: east Andean slopes from Ecuador to southern Peru (Freitas et al. 2009). Host plants: Smallanthus sonchifolius (Asteraceae) (Freitas et al. 2009). Refs.: Freitas et al. (2009) (immature).

• stratonice (Latreille [1813])

  • ssp. acipha (Hewitson, 1861); original combination: Acraea acipha

  • ssp. adoxa Jordan, 1910; original combination: Actinote adoxa

  • ssp. aereta Jordan, 1913; original combination: Actinote stratonice aereta

    • syn. stratonice diversa Jordan, 1913

    • syn. laeta Oberthür, 1917

    • syn. stratonice albunea (Potts, 1944)

  • ssp. marthae Jordan, 1910; original combination: Actinote stratonice marthae

  • ssp. meridana Jordan, 1910; original combination: Actinote stratonice meridana

  • ssp. oaxaca Miller & Miller, 1979; original combination: Actinote stratonice oaxaca

  • ssp. stratonice (Latreille [1813]); original combination: Heliconius stratonice Distr.: Sucumbios, Ecuador.

• surima (Schaus, 1902)

  • ssp. perisa Jordan, 1913; original combination: Actinote perisa

  • ssp. surima (Schaus, 1902); original combination: Acraea surima Refs.: Schaus 1902:392; Jordan 1913b:374 pI. 82 f,g; Eltringham and Jordan 1913:13; Jordan and Eltringham 1916:19 Figs. 8, 9; Hayward Hl3l:29,139 pI. 8; Monte 1934:199; Hayward 1935:97; l)’Almeida 1935a:94; d’Almeida 1935b:488; Schweizer and Webster-Kay 1941:14; Biezanko and Pitton 1941:9; Hayward 1950:209; Biezanko, Huflnelli and Carbonell 1957:127; Brown and Mielke 1967:151; D’Araujo e Silva et al. 1968:340; Ebert 1969:35; Brown and Benson 1974:218 pI. 5 fig.]; Smart 1976:191, 265 pI. 5 fig.l; Brown 1984:158; Almeida, Souza and Marques 1986:33; d’Abrera 1987:445; Francini 1989:44 and several other pages, several figs; Brown 1988:36; Francini HJ92:26 flg.6-14 [SUR].

    • syn. f. punctata Hayward, 1935

• tenebrosa (Hewitson, 1868)

  • ssp. erebia Oberthür, 1917; original combination: Actinote erebia

  • ssp. tenebrosa (Hewitson, 1868); original combination: Acraea tenebrosa

    • syn. segesta (Weymer, 1890)

    • syn. dognini (Schaus, 1902)

• thalia (Linnaeus, 1758)

  • ssp. anteas (Doubleday [1847]); original combination: Acraea anteas

    • syn. anteas f. holochroa Jordan, 1913

    • syn. anteas f. ochrotaenia Jordan, 1913

  • ssp. brettia Oberthür, 1917; original combination: Actinote brettia

  • ssp. byssa Oberthür, 1917; original combination: Actinote byssa

  • ssp. cedestis Jordan, 1913; original combination: Actinote cedestis. Type loc.: Zamora, SE. Ecuador.

  • ssp. crassinia (Hopffer, 1874); original combination: Acraea crassinia

    • syn. terpsinoë roqueensis Bryk, 1953

    • syn. terpsinoë roqueensis f. puricella Bryk, 1953

  • ssp. eupelia Jordan, 1913; original combination: Actinote crassinia eupelia. Type loc.: Bolivia.

  • ssp. pierrei Neild, 2008. Type loc.: Sta. Elena to Icabaru Road, Bolívar, Venezuela. Types: MIZA. Distr.: Eastern (and western?) Pantepui, Venezuela. Refs.: Neild (2008): 27. Ety.: Jacques Pierre

  • ssp. sainorum Neild & Romero, 2008. Type loc.: Rio Chucurí, southeast of San Cristobal, Táchira [Venezuela]. Types: MIZA. Distr.: Sierra de El Tamá, Cordilleras de Mérida and la Costa, Serranía del Interior, Sierra de Turimiquire. Refs.: Neild (2008): 30. Ety.: Edwin Saino

  • ssp. suspecta Jordan, 1913; original combination: Actinote cedestes[sic] suspecta. Type loc.: Paramba, Ecuador.

  • ssp. terpsinoe (Felder & Felder, 1862); original combination: Acraea terpsinoe

    • syn. terpsinoë (Felder & Felder, 1862), missp.

  • ssp. thalia (Linnaeus, 1758); original combination: Papilio thalia

    • syn. ixilion (Linnaeus, 1758), nom. nud.

    • syn. acton (Herrich-Schäffer, 1865), nom. nud.

    • syn. ab. idiographa Jordan, 1913

• trinacria (Felder & Felder, 1862)

  • ssp. alegria Boyer & Willmott, 2024; original combination: Actinote trinacria alegria Refs.: Willmott et al. (2024): 16.

  • ssp. naura (Druce, 1875); original combination: Acraea trinacria naura

  • ssp. santamarta (Winhard, 2017); original combination: Altinote santamarta Refs.: Winhard (2017); Willmott et al. (2024) (rev. stat.).

  • ssp. tenebrarum Oberthür, 1917; original combination: Actinote tenebrarum

    • syn. naura unicolor Talbot, 1932

  • ssp. trinacria (Felder & Felder, 1862); original combination: Acraea trinacria

    • syn. iguaquensis Andrade & Restrepo, 1992

  • ssp. vivasi Neild & Romero, 2008. Type loc.: Fundo Piedra Blanca, San Vicente de la Revancha, Sierra de El Tama, Tachira, Venezuela. Types: MIZA. Distr.: N. Colombia (Santander) to Venezuela (Sierra de El Tamá). Refs.: Neild (2008): 41. Ety.: Moises “Chato” Vivas

• zikani d’Almeida, 1951; original combination: Actinote zikani Host plants: Mikania obsoleta (Francini et al. 2011). Refs.: Francini et al. (2011) (immature).

Genus Bematistes

Genus: Bematistes Hemming, 1935

Type species: Bematistes umbra (Drury, 1782); original combination: Papilio umbra Drury, 1782

Synonyms: None

References: Hemming (1935); Son (1963); Timmermans et al. (2016) (phylogeny)

35 species:

• adrasta (Weymer, 1892)

  • ssp. adrasta (Weymer, 1892); original combination: Acraea adrasta

    • syn. machoni (Oberthür, 1893)

    • syn. haydni (Suffert, 1904)

  • ssp. pancalis (Jordan, 1910); original combination: Planema adrasta pancalis

• aganice (Hewitson [1852])

  • ssp. aganice (Hewitson [1852]); original combination: Acraea aganice

  • ssp. montana (Butler, 1888); original combination: Planema montana

    • syn. bertha (Vuillot, 1891)

    • syn. meruana (Rogenhofer, 1891)

    • syn. chanleri (Holland, 1896)

  • ssp. nicega (Suffert, 1904); original combination: Planema aganice nicega

  • ssp. nyassae (Carpenter, 1920); original combination: Planema aganice r. nyasae

  • ssp. orientalis (Ungemach, 1932); original combination: Planema aganice orientalis

  • ssp. ugandae (van Someren, 1936); original combination: craea aganice ugandae

• alcinoe (Felder & Felder [1865])

  • ssp. alcinoe (Felder & Felder [1865]); original combination: Acraea alcinoe

    • syn. godmani (Butler, 1895)

    • syn. timandra (Karsch, 1893)

  • ssp. camerunica (Aurivillius, 1893); original combination: Planema alcinoe var. camerunica

    • syn. salvini (Butler, 1895)

  • ssp. nado (Ungemach, 1932); original combination: Planema nado

  • ssp. racaji Pyrcz, 1991; original combination: Bematistes alcinoe racaji

• bakundu (Bernaud, 2021); original combination: Acraea bakundu

• bana (Pierre & Bernaud, 2012); original combination: Acraea bana

• consanguinea (Aurivillius, 1893)

  • ssp. albicolor (Karsch, 1895); original combination: Planema albicolor

  • ssp. consanguinea (Aurivillius, 1893); original combination: Planema consanguinea

  • ssp. intermedia (Aurivillius, 1899); original combination: Planema consanguinea var. intermedia

• dewitzi (Staudinger, 1896); original combination: Acraea (Planema) dewitzi

• dimonika (Bernaud, 2021); original combination: Acraea dimonika

• ducarmei (Bernaud & Pierre, 2012); original combination: Acraea ducarmei

• elongata (Butler, 1974); original combination: Planema elongata

  • syn. serena (Schultze, 1923); original combination: Planema elongata var. serena.

  • syn. varia (Le Doux, 1937); original combination: Planema elongata f. varia.

• epaea (Cramer [1779]); original combination: Papilio epaea

  • syn. gea (Fabricius, 1781); original combination: Papilio gea.

  • syn. lutosa (Suffert, 1904); original combination: Planema epaea lutosa.

  • syn. sublutosa (Strand, 1914); original combination: Planema epaea ab. sublutosa.

  • syn. insolita (Le Doux, 1937); original combination: Planema epaea f. insolita.

  • syn. cremea (Le Doux, 1937); original combination: Planema epaea f. cremea.

  • syn. aurata (Le Doux, 1937); original combination: Planema epaea f. aurata.

  • syn. bicolorata (Le Doux, 1937); original combination: Planema epaea f. bicolorata.

  • syn. nigrita (Le Doux, 1937); original combination: Planema epaea nigrita.

• epiprotea (Butler, 1874); original combination: Planema epiprotea

  • syn. conformis (Schultze, 1923); original combination: Planema epiprotea ab. conformis.

• epitellus (Staudinger, 1896); original combination: Acraea (Planema) epitellus

• excisa (Butler, 1874); original combination: Planema excisa

• formosa (Butler, 1874); original combination: Planema formosa

• indentata (Butler, 1874); original combination: Planema indentata

• kivuana (Jordan, 1910); original combination: Planema epaea kivuana Refs.: Williams and Hennig (2023) (to species).

• kivuensis (Joicey & Talbot, 1927)

  • ssp. elgonense (Poulton, 1927); original combination: Planema elgonensis Refs.: Williams and Hennig (2023) (transferred from a subspecies of Bematistes obliqua Aurivillius).

    • syn. ela Rumbucher, 1992

  • ssp. kivuensis (Joicey & Talbot, 1927); original combination: Planema obliqua kivuensis

    • syn. elgonense toroense (Poulton, 1929)

• leopoldina (Aurivillius, 1895)

  • ssp. brevimacula (Talbot, 1928); original combination: Planema poggei brevimacula

    • syn. schoutedeni (Le Doux, 1937)

  • ssp. leopoldina (Aurivillius, 1895); original combination: Planema leopoldina

    • syn. albofasciata (Le Doux, 1937)

    • syn. mixta (Le Doux, 1937)

  • ssp. macrosticha (Bethune-Baker, 1908); original combination: Planema macrosticha

    • syn. intermissa (Gaede, 1916)

• lequeuxi (Bernaud & Pierre, 2012); original combination: Acraea lequeuxi

• macaria (Fabricius, 1793)

  • ssp. hemileuca (Jordan, 1914); original combination: Planema macaria hemileuca Refs.: Williams and Hennig (2023) (transferred from a subspecies of Bematistes umbra Drury).

  • ssp. hewitsoni (Aurivillius, 1899); original combination: Planema macarioides var. hewitsoni

  • ssp. macaria (Fabricius, 1793); original combination: Papilio macaria

  • ssp. macaroides (Aurivillius, 1893); original combination: Planema macarioides Refs.: Williams and Hennig (2023) (transferred from a subspecies of Bematistes umbra Drury).

    • syn. rabuma (Suffert, 1904)

• macarista (Sharpe, 1906)

  • ssp. latefasciata (Suffert, 1904); original combination: Planema formosa latefasciata

  • ssp. macarista (Sharpe, 1906); original combination: Planema macarista

• melina (Thurau, 1903); original combination: Planema epaea var. melina Refs.: Williams and Hennig (2023) (to species from a subspecies of Bematistes epaea Cramer).

• obliqua (Aurivillius [1913]); original combination: Planema obliqua

• parageum (Grose-Smith, 1900)

  • ssp. homochroa (Rothscild & Jordan, 1905); original combination: Planema epaea homochroa Refs.: Williams and Hennig (2023) (transferred from a subspecies of Bematistes epaea Cramer).

  • ssp. insulana (Ackery, 1995); original combination: Acraea (Acraea) epaea insulana Refs.: Williams and Hennig (2023) (transferred from a subspecies of Bematistes epaea Cramer).

    • syn. insularis (Aurivillius, 1910) Refs.: Pierre and Bernaud (2012) (synonymy).

  • ssp. parageum (Grose-Smith, 1900); original combination: Planema paragea

    • syn. angustifasciata d’Abrera, 1980

• persanguinea (Rebel, 1914); original combination: Planema persanguinea

  • syn. consanguinoides (Le Doux, 1931); original combination: Planema consanguinoides.

• poggei (Dewitz, 1879)

  • ssp. nelsoni (Grose-Smith & Kirby, 1892); original combination: Acraea nelsoni

    • syn. paragoga (Hering, 1936)

  • ssp. poggei (Dewitz, 1879); original combination: Acraea poggei

    • syn. entalis (Jordan, 1910)

    • syn. amela (Hering, 1936)

  • ssp. poggei (Le Doux, 1937); original combination: Planema poggei nelsoni f. flava

    • syn. entalis (Jordan, 1910)

    • syn. amela (Hering, 1936)

• poggei (Le Doux, 1937)

  • ssp. nelsoni (Grose-Smith & Kirby, 1892); original combination: Acraea nelsoni

    • syn. paragoga (Hering, 1936)

  • ssp. poggei (Dewitz, 1879); original combination: Acraea poggei

    • syn. entalis (Jordan, 1910)

    • syn. amela (Hering, 1936)

  • ssp. poggei (Le Doux, 1937); original combination: Planema poggei nelsoni f. flava

    • syn. entalis (Jordan, 1910)

    • syn. amela (Hering, 1936)

• pseudeuryta (Godman & Salvin, 1890); original combination: Acraea pseudeuryta

  • syn. formosa angulata (Suffert, 1904); original combination: Planema formosa angulata.

• quadricolour (Rogenhöfer, 1891)

  • ssp. itumbana (Jordan, 1910); original combination: Planema quadricolor itumbana

  • ssp. latifasciata (Sharpe, 1892); original combination: Planema latifasciata

  • ssp. leptis (Jordan, 1910); original combination: Planema quadricolor leptis

  • ssp. mahale Kielland, 1990; original combination: Bematistes quadricolor mahale

  • ssp. morogoro Carpenter & Jackson, 1950; original combination: Bematistes quadricolor morogoro

  • ssp. quadricolour (Rogenhöfer, 1891); original combination: Planema quadricolor

  • ssp. uluguru Kielland, 1990; original combination: Bematistes quadricolor uluguru

• sartina (Jordan, 1910); original combination: Planema consanguinea sartina Refs.: Williams and Hennig (2023) (to species).

• scalivittata (Butler, 1896)

  • ssp. kiellandianus (Koçak, 1996); original combination: Acraea scalivittata kiellandianus

    • syn. kiellandi Collins, 1990, preocc.

  • ssp. scalivittata (Butler, 1896); original combination: Planema scalivittata

• schubotzi (Grünberg, 1911); original combination: Planema schubotzi

• tellus (Aurivillius, 1893)

  • ssp. eumelis (Jordan, 1910); original combination: Planema tellus eumelis

  • ssp. tellus (Aurivillius, 1893); original combination: Planema tellus

    • syn. lustella (Suffert, 1904)

    • syn. subapicalis (Strand, 1914)

    • syn. albofasciata (Neustetter, 1916)

    • syn. helichta (Neustetter, 1916)

    • syn. ferruginea (Le Doux, 1937)

    • syn. bernardi Berger, 1981

• umbra (Drury, 1782)

  • ssp. carpenteri (Le Doux, 1937); original combination: Planema umbra carpenteri

  • ssp. umbra (Drury, 1782); original combination: Papilio umbra

• vestalis (Felder & Felder [1865])

  • ssp. congoensis (Le Doux, 1937); original combination: Planema vestalis congoensis

  • ssp. stavelia (Suffert, 1904); original combination: Planema vestalis stavelia

  • ssp. vestalis (Felder & Felder [1865]); original combination: Acraea vestalis

Genus Cethosia

Genus: Cethosia Fabricius, 1807

Type species: Cethosia cydippe (Linnaeus, 1763); original combination: Papilio cydippe Linnaeus, 1763

Synonyms:

• Alazonia Hübner [1819]. Type species: Cethosia cydippe (Linnaeus, 1763); original combination: Papilio cydippe Linnaeus, 1763. Refs.: Hübner ([1819]): 46
• Eugramma Billberg, 1820. Type species: Cethosia cydippe (Linnaeus, 1763); original combination: Papilio cydippe Linnaeus, 1763. Refs.: Billberg (1820): 78

References: Fabricius (1807): 280; Müller and Beheregaray (2010) (phylogeny)

17 species:

• biblis (Drury [1773])

  • ssp. adantonia Fruhstorfer, 1912; original combination: Cethosia biblis adantonia

  • ssp. amboinensis Felder & Felder [1867]; original combination: Cethosia insularis var. amboinensis

    • syn. amboinensis Felder, 1862, nom. nud.

    • syn. ceramensis Fruhstorfer, 1902

  • ssp. biblis (Drury [1773]); original combination: Papilio biblis

  • ssp. buruana Holland, 1900; original combination: Cethosia buruana

  • ssp. hainana Fruhstorfer, 1908; original combination: Cethosia biblis hainana

  • ssp. insularis Felder & Felder, 1861; original combination: Cethosia insularis

    • syn. eurymena Felder & Felder [1867]

  • ssp. javana Felder & Felder [1867]; original combination: Cethosia javana

  • ssp. liacura Fruhstorfer, 1912; original combination: Cethosia biblis liacura

  • ssp. logani Distant, 1881; original combination: Cethosia logani

  • ssp. narmadoides de Nicéville, 1898; original combination: Cethosia narmadoides

  • ssp. nicobarica Felder & Felder, 1862; original combination: Cethosia nikobarica

    • syn. andamanica Stichel, 1908

  • ssp. perakana Fruhstorfer, 1902; original combination: Cethosia biblis perakana

  • ssp. permanggilensis Eliot, 1978; original combination: Cethosia biblis pemanggilensis

  • ssp. phanaroia Fruhstorfer, 1912; original combination: Cethosia biblis phanaroia

  • ssp. picta Felder & Felder [1867]; original combination: Cethosia picta

    • syn. sarsina Fruhstorfer, 1912

    • syn. viridipicta Fruhstorfer, 1912

  • ssp. sandakana Fruhstorfer, 1899; original combination: Cethosia biblis sandakana

  • ssp. tagalorum Fruhstorfer, 1912; original combination: Cethosia biblis tagalorum

  • ssp. tisamena Fruhstorfer, 1912; original combination: Cethosia biblis tisamena

    • syn. thebava Grose-Smith, 1887

    • syn. mixta Fruhstorfer, 1912

  • ssp. togiana Fruhstorfer, 1902; original combination: Cethosia picta togiana

  • ssp. woolletti Hall, 1929; original combination: Cethosia javana woolletti

• cydippe (Linnaeus, 1763)

  • ssp. alkmene Fruhstorfer, 1912; original combination: Cethosia cydippe alkmene

  • ssp. antoni Kawai, 1996; original combination: Cethosia chrysippe antoni

  • ssp. bernsteini Felder & Felder [1867]; original combination: Cethosia bernsteinii

  • ssp. cenchrites Fruhstorfer, 1909; original combination: Cethosia cydippe cenchrites

  • ssp. chrysippe (Fabricius, 1775); original combination: Papilio chrysippe

    • syn. imperialis Butler, 1876

  • ssp. cleanthis Fruhstorfer, 1902; original combination: Cethosia cydippe cleanthis

  • ssp. cydalima Felder & Felder [1867]; original combination: Cethosia cydalima

    • syn. aruensis Talbot, 1932

  • ssp. cydippe (Linnaeus, 1763); original combination: Papilio cydippe Refs.: Linnaeus, 1763, Amoenitates Acad. 6: 409.

    • syn. ino (Cramer [1775])

    • syn. chrysonoe Godart, 1819

    • syn. aurigena Fruhstorfer, 1912

    • syn. tymbrasa Fruhstorfer, 1912

    • syn. ternatensis Hall, 1929

    • syn. koridana Kalis, 1933

    • syn. cydippina Stichel, 1938

  • ssp. cyrene Wallace, 1869; original combination: Cethosia cyrene

  • ssp. damasippe Felder & Felder [1867]; original combination: Cethosia damasippe

    • syn. hermanni Fruhstorfer, 1902

    • syn. lucina Fruhstorfer, 1905

    • syn. coronilla Fruhstorfer, 1909

    • syn. praestabilis Fruhstorfer, 1909

    • syn. claudilla Fruhstorfer, 1912

    • syn. antianeira Fruhstorfer, 1913

    • syn. xerxene Fruhstorfer, 1915

  • ssp. doxata Fruhstorfer, 1913; original combination: Cethosia cydippe doxata

  • ssp. iphigenia Fruhstorfer, 1901; original combination: Cethosia cydippe iphigenia

  • ssp. insulata Butler, 1873; original combination: Cethosia insulata

    • syn. theona Fruhstorfer, 1903

  • ssp. mysolensis Fruhstorfer, 1913; original combination: Cethosia cydippe mysolensis

  • ssp. obiana Fruhstorfer, 1903; original combination: Cethosia cydippe obiana

  • ssp. salwattensis Fruhstorfer, 1913; original combination: Cethosia cydippe salwattensis

  • ssp. sangira Fruhstorfer, 1906; original combination: Cethosia cydippe sangira

  • ssp. schoutensis Joicey & Noakes, 1915; original combination: Cethosia chrysippe schoutensis

  • ssp. woodlarkiana Fruhstorfer, 1912; original combination: Cethosia cydippe woodlarkiana

• gabinia Weymer, 1883; original combination: Cethosia gabinia

• hypsea Doubleday [1847]

  • ssp. aeole Moore [1858]; original combination: Cethosia aeole Distr.: north-eastern Sumatra.

  • ssp. bankana Fruhstorfer, 1912; original combination: Cethosia hypsea bankana Distr.: Bangka Island.

  • ssp. batuensis Stichel, 1908; original combination: Cethosia hypsea batuensis Distr.: Batu Islands.

  • ssp. elioti Okubu, 1983; original combination: Cethosia hypsea elioti

  • ssp. festiva Fruhstorfer, 1909; original combination: Cethosia mindanensis festiva Distr.: Jolo (Sulu Archipelago).

  • ssp. fruhstorferi Stichel, 1908; original combination: Cethosia hypsea fruhstorferi Distr.: eastern Java.

  • ssp. hypsea Doubleday [1847]; original combination: Cethosia hypsea Distr.: Borneo.

  • ssp. hypsina Felder & Felder [1867]; original combination: Cethosia hypsina Distr.: southern Burma to Singapore.

  • ssp. munjava Fruhstorfer, 1912; original combination: Cethosia hypsea munjava Distr.: western Java.

  • ssp. nigrescens van Eecke, 1913; original combination: Cethosia hypsea var. nigrescens

  • ssp. palawana Fruhstorfer, 1900; original combination: Cethosia hypsea palawana Distr.: Palawan.

  • ssp. pallaurea Hagen, 1898; original combination: Cethosia pallaurea Distr.: Mentawai.

  • ssp. triocala Fruhstorfer, 1912; original combination: Cethosia hypsea triocala Distr.: western Sumatra.

• lamarcki Godart, 1819; original combination: Cethosia lamarckii

  • syn. timorensis Stichel, 1908; original combination: Cethosia lamarckii timorensis.

  • syn. justa Fruhstorfer, 1912; original combination: Cethosia lamarcki f. justa.

  • syn. elateia Fruhstorfer, 1912; original combination: Cethosia lamarcki elateia.

  • syn. orientalis Talbot, 1932; original combination: Cethosia lamarckii orientalis.

• lechenault Godart [1824]; original combination: Cethosia leschenault

• luzonica Felder & Felder, 1863

  • ssp. boholica Semper, 1888; original combination: Cethosia luzonica var. boholica

  • ssp. luzonica Felder & Felder, 1863; original combination: Cethosia luzonica

  • ssp. magindanica Semper, 1888; original combination: Cethosia luzonica var. magindanica

  • ssp. pariana Semper, 1888; original combination: Cethosia luzonica var. pariana

• methypsea Butler, 1879

  • ssp. carolinae Forbes, 1885; original combination: Cethosia carolinae

  • ssp. methypsea Butler, 1879; original combination: Cethosia methypsea

• mindanensis Felder & Felder, 1863; original combination: Cethosia mindanensis Distr.: Philippines: Mindanao, Basilan.

• moesta Felder & Felder [1867]; original combination: Cethosia moesta

• myrina Felder & Felder [1867]

  • ssp. melancholica Fruhstorfer, 1912; original combination: Cethosia myrina melancholica

  • ssp. myrina Felder & Felder [1867]; original combination: Cethosia myrina Refs.: Vane-Wright (2012) (taxonomy, Sulawesi).

  • ssp. ribbei Honrath, 1887; original combination: Cethosia myrina var. ribbei

  • ssp. sarnada Fruhstorfer, 1912; original combination: Cethosia myrina sarnada

  • ssp. vanbemmeleni Jurriaanse & Lindemans, 1918, emend. (ICZN, Art. 33.2.3.1); original combination: Cethosia myrina bemmeleni [sic] Refs.: Vane-Wright (2012) (taxonomy, Sulawesi). Ety.: J. F. van Bemmelen

    • syn. vanbemmeleni Jurriaanse & Lindemans, 1920

• nietneri Felder & Felder [1867]

  • ssp. nietneri Felder & Felder [1867]; original combination: Cethosia nietneri

  • ssp. mahratta Moore, 1872; original combination: Cethosia mahratta

• obscura Guérin-Méneville [1830]

  • ssp. antippe Grose-Smith, 1889; original combination: Cethosia antippe

  • ssp. gabrielis Rothschild, 1898; original combination: Cethosia gabrielis

  • ssp. hormisda Fruhstorfer, 1915; original combination: Cethosia obscura hormisda

  • ssp. manusi Rothschild, 1915; original combination: Cethosia obscura manusi

  • ssp. obscura Guérin-Méneville [1830]; original combination: Cethosia obscura

• penthesilea (Cramer [1777])

  • ssp. exsanguis Fruhstorfer, 1897; original combination: Cethosia penthesilea exsanguis

    • syn. filiola Fruhstorfer, 1902

    • syn. diffusa Fruhstorfer, 1905

  • ssp. paksha Fruhstorfer, 1905; original combination: Cethosia penthesilea paksha

  • ssp. penthesilea (Cramer [1777]); original combination: Papilio penthesilea

    • syn. doddi Oberthür, 1911

    • syn. elgitha Fruhstorfer, 1912

    • syn. baweanica Fruhstorfer, 1912

• tambora Doherty, 1891

  • ssp. adonarensis Talbot, 1932; original combination: Cethosia biblis adonarensis

  • ssp. alceste Fruhstorfer, 1905; original combination: Cethosia biblis alceste

  • ssp. atia Fruhstorfer, 1905; original combination: Cethosia biblis (?) atia

  • ssp. floresiana Fruhstorfer, 1902; original combination: Cethosia tambora floresiana

  • ssp. nacoleia Fruhstorfer, 1912; original combination: Cethosia biblis nacoleia

  • ssp. narmada Fruhstorfer, 1896; original combination: Cethosia narmada

  • ssp. narmadoides de Nicéville, 1898; original combination: Cethosia narmadoides

  • ssp. sumbana Pagenstecher, 1894; original combination: Cethosia cyane var. sumbana

  • ssp. tambora Doherty, 1891; original combination: Cethosia tambora

• vasilia Müller, 1999; original combination: Cethosia vasilia

Genus Rubraea

Genus: Rubraea Henning, 1992

Type species: Rubraea egina (Cramer, [1775]); original combination: Papilio egina Cramer, [1775]

Synonyms: None

References: Hennig (1993 b); Williams and Hennig (2023) (to genus)

40 species:

• abdera (Hewitson [1852])

  • ssp. abdera (Hewitson [1852]); original combination: Acraea abdera

  • ssp. eginopsis (Aurivillius, 1899); original combination: Acraea cepheus var. eginopsis

    • syn. nigrescens (Eltringham, 1912)

• acrita (Hewitson [1865])

  • ssp. acrita (Hewitson [1865]); original combination: Acraea acrita

    • syn. pauperata (Thurau, 1903)

    • syn. aquilia (Thurau, 1903)

    • syn. chaeribulula (Strand, 1911)

    • syn. aquilina (Strand, 1911)

    • syn. msamwiae (Strand, 1911)

    • syn. lindica (Strand, 1911)

    • syn. usaramensis (Strand, 1911)

    • syn. nyassicola (Strand, 1911)

    • syn. littoralis (Eltringham, 1912)

    • syn. megaspila (Le Cerf, 1927)

    • syn. albomaculosa (Le Doux, 1931)

    • syn. fusca (Le Doux, 1932)

    • syn. nigromarginata (Le Doux, 1932)

  • ssp. ambigua (Trimen, 1891); original combination: Acraea ambigua

    • syn. bella (Weymer, 1901)

• annonae (Pierre, 1987); original combination: Acraea annonae

  • syn. kapiriensis (Schouteden, 1927); original combination: Acraea lualabae f. kapiriensis.

• asema (Hewitson, 1877); original combination: Acraea asema

  • syn. empusa (Butler, 1894); original combination: Acraea empusa.

  • syn. despecta (Le Doux, 1922); original combination: Acraea asema f. despecta.

  • syn. angustifasciata (Le Doux, 1922); original combination: Acraea asema f. angustifasciata.

  • syn. aspectasemoides (Le Doux, 1922); original combination: Acraea asema f. aspectasemoides.

  • syn. dissimiloides (Le Doux, 1922); original combination: Acraea asema f. dissimiloides.

• atolmis (Westwood, 1881); original combination: Acraea atolmis

  • syn. acontias (Westwood, 1881); original combination: Acraea acontias.

  • syn. luxi (Rogenhofer, 1889); original combination: Acraea luxi.

  • syn. decora (Weymer, 1901); original combination: Acraea acontias ab. decora.

  • syn. nigra (Neustetter, 1916); original combination: Acraea atolmis ab. nigra.

  • syn. westwoodi (van Son, 1963); original combination: Acraea atolmis f. westwoodi.

• bailundensis (Wichgraf, 1918); original combination: Acraea bailunduensis

  • syn. schoutedeni (Overlaet, 1954); original combination: Acraea schoutedeni.

• bellona (Weymer, 1908); original combination: Acraea acrita bellona

• cepheus (Linnaeus, 1758)

  • ssp. bergeriana (Pierre, 1979); original combination: Acraea bergeriana. Type loc.: Uluguru Mts, Kinola. Refs.: Williams and Hennig (2023) (to subspecies).

    • syn. bergeri (Pierre, 1976)

  • ssp. cepheus (Linnaeus, 1758); original combination: Papilio cepheus

    • syn. zosteria (Godart, 1819)

    • syn. baumanni (Rogenhofer, 1889)

    • syn. pheusaca (Suffert, 1904)

    • syn. sucepha (Suffert, 1904)

    • syn. cepheana (Strand, 1914)

    • syn. disjuncta (Dufrane, 1945)

    • syn. addenda (Dufrane, 1945)

    • syn. deficiens (Dufrane, 1945)

• cerasa (Hewitson [1861])

  • ssp. cerasa (Hewitson [1861]); original combination: Acraea cerasa Refs.: Pierre (2020 b) (Acraea); Williams and Hennig (2023) (Rubraea).

  • ssp. cerita (Sharpe, 1906); original combination: Acraea cerita

    • syn. kigezia (Howarth, 1959)

  • ssp. kiellandi (Carcasson, 1964); original combination: Acraea cerasa kiellandi

• chaeribula (Oberthür, 1893); original combination: Acraea chaeribula

• chambezi (Neave, 1910); original combination: Acraea nohara chambezi

• diogenes (Suffert, 1904); original combination: Acraea diogenes

  • syn. acutipennis (Lathy, 1906); original combination: Acraea acutipennis.

  • syn. lactea (Neave, 1910); original combination: Acraea lactea.

• egina (Cramer [1775])

  • ssp. areca (Mabille, 1889); original combination: Acraea areca

  • ssp. egina (Cramer [1775]); original combination: Papilio egina

    • syn. rudolphina (Herbst, 1792)

    • syn. persephone (Fabricius, 1793)

    • syn. zidora (Godart, 1819)

    • syn. khara (Grose-Smith, 1889)

    • syn. contraria (Grünberg, 1910)

    • syn. alba (Eltringham, 1913)

    • syn. intensa (Stoneham, 1937)

    • syn. tenuimarginatus (Stoneham, 1937)

    • syn. rubristriatus (Stoneham, 1937)

    • syn. bellehui (Carcasson, 1961)

  • ssp. harrisoni (Sharpe, 1904); original combination: Acraea harrisoni

  • ssp. pembanus (Kielland, 1990); original combination: Acraea egina pembanus

• eltringhamiana (Le Doux, 1932); original combination: Acraea acrita eltringhamiana

• guillemei (Oberthür, 1893); original combination: Acraea guillemei

• guluensis (Le Doux, 1932); original combination: Acraea guluensis

• kraka (Aurivillius, 1893); original combination: Acraea kraka

  • syn. var. pallida (Carpenter, 1932); original combination: Acraea kraka var. pallida.

  • syn. kibi (Usher, 1986); original combination: Acraea kibi.

• lapidorum (Pierre, 1988); original combination: Acraea lapidorum

• lofua (Eltringham, 1911); original combination: Acraea lofua

• loranae (Pierre, 1987); original combination: Acraea loranae

• lualabae (Neave, 1910); original combination: Acraea lualabae

• manca (Thurau, 1904); original combination: Acraea guillemei var. manca

• mansya (Eltringham, 1911); original combination: Acraea mansya

  • syn. janssensi (Eltringham, 1911); original combination: Acraea mansya janssensi.

• medea (Cramer [1775]); original combination: Papilio medea

  • syn. pasiphae (Fabricius, 1781); original combination: Papilio pasiphae.

  • syn. saronis (Hübner [1819]); original combination: Telchinia saronis.

• niobe (Sharpe, 1893); original combination: Acraea niobe

• nohara (Boisduval, 1847)

  • ssp. dondoensis (Stevenson, 1934); original combination: Acraea dondoensis Refs.: Williams and Hennig (2023) (to subspecies).

    • syn. junodi (Oberthür, 1911)

  • ssp. halali (Marshall, 1896); original combination: Acraea halali

  • ssp. nohara (Boisduval, 1847); original combination: Acraea nohara

    • syn. actiaca (Hewitson, 1852)

• omrora (Trimen, 1894)

  • ssp. omrora (Trimen, 1894); original combination: Acraea omrora

  • ssp. umbraetae (Pierre, 1988); original combination: Acraea omrora umbraetae

• onerata (Trimen, 1891); original combination: Acraea onerata

  • syn. aureola (Eltringham, 1911); original combination: Acraea aureola.

• overlaeti (Pierre, 1988); original combination: Acraea overlaeti

• peetersi (Pierre, 1992); original combination: Acraea peetersi

• periphanes (Oberthür, 1893); original combination: Acraea periphanes

  • syn. beni (Bethune-Baker, 1908); original combination: Acrea[sic] beni.

  • syn. umida (Wichgraf, 1908); original combination: Acraea onerata ♀ f. umida.

  • syn. melaina (Eltringham, 1911); original combination: Acraea periphanes f. melaina.

  • syn. acritoides (Eltringham, 1911); original combination: Acraea periphanes f. acritoides.

  • syn. marginata (Eltringham, 1911); original combination: Acraea periphanes f. marginata.

  • syn. interposita (Wichgraf, 1918); original combination: Acraea periphanes interposita.

  • syn. seitzi (Le Doux, 1922); original combination: Acraea periphanes f. seitzi.

  • syn. bihensis (Le Doux, 1923); original combination: Acraea periphanes f. bihensis.

• petraea (Boisduval, 1847); original combination: Acraea petraea

  • syn. petrina (Suffert, 1904); original combination: Acraea petraea petrina.

  • syn. taborana (Suffert, 1904); original combination: Acraea petraea taborana.

  • syn. pseudacontias (Wichgraf, 1914); original combination: Acraea petraea ab. pseudacontias.

  • syn. grisea (Neustetter, 1916); original combination: Acraea petraea ab. grisea.

• pseudatolmis (Eltringham, 1912); original combination: Acraea nohara pseudatolmis

• pudorina (Staudinger [1885]); original combination: Acraea pudorina

  • syn. emboensis (Gaede, 1915); original combination: Acraea acrita var. emboensis.

  • syn. rubida (Le Doux, 1923); original combination: Acraea acrita pudorina f. rubida.

  • syn. mancamorpha (Le Doux, 1932); original combination: Acraea acrita pudorina f. mancamorpha.

• punctellata (Eltringham, 1912); original combination: Acraea nohara punctellata

  • syn. noharoides (Le Doux, 1923); original combination: Acraea nohara f. noharoides.

• rohlfsi (Suffert, 1904); original combination: Acraea rohlfsi

• severina (Oremans, 2012)

  • ssp. severina (Oremans, 2012); original combination: Acraea severina

  • ssp. terreirovelhoensis (Oremans, 2012); original combination: Acraea severina terreirovelhoensis

• unimaculata (Grose-Smith, 1898); original combination: Acraea unimaculata

• utengulensis (Thurau, 1903); original combination: Acraea acrita var. utengulensis

  • syn. taborensis (Le Doux, 1931); original combination: Acraea (acrita) manca f. taborensis.

• violarum (Boisduval, 1847)

  • ssp. anchietai (Mendes & Bivar-de-Sousa, 2017); original combination: Acraea (Acraea) violarum anchietai

  • ssp. gracilis (Wichgraf, 1909); original combination: Acraea violarum gracilis

  • ssp. violarum (Boisduval, 1847); original combination: Acraea violarum

    • syn. nataliensis (Angas, 1849)

    • syn. assimiliora (Le Doux, 1922)

    • syn. assimilis (Le Doux, 1922)

    • syn. dissimilis (Le Doux, 1922)

    • syn. aspectasema (Le Doux, 1922)

Genus Stephenia

Genus: Stephenia Henning, 1992

Type species: Stephenia caecilia (Fabricius, 1781); original combination: Papilio caecilia Fabricius, 1781

Synonyms: None

References: Hennig (1993 b); Williams and Hennig (2023) (to genus)

30 species:

• aglaonice (Westwood, 1881); original combination: Acraea aglaonice

  • syn. fenestrata (Trimen, 1881); original combination: Acraea fenestrata.

  • syn. albofasciata (Aurivillius, 1913); original combination: Acraea aglaonice ab. albofasciata.

  • syn. leucaspis (Wichgraf, 1914); original combination: Acraea aglaonice f. leucaspis.

  • syn. latimarginata (van Son, 1963); original combination: Acraea aglaonice f. latimarginata.

• asboloplintha (Karsch, 1894)

  • ssp. asboloplintha (Karsch, 1894); original combination: Acraea asboloplintha

    • syn. dissociata (Grose-Smith, 1898)

  • ssp. rubescens (Trimen, 1909); original combination: Acraea asboloplintha rubescens

• atatis (Pierre, 2004); original combination: Acraea atatis

• atergatis (Westwood, 1881); original combination: Acraea atergatis

  • syn. eichleri (van Son, 1963); original combination: Acraea atergatis f. eichleri.

• axina (Westwood, 1881); original combination: Acraea axina

  • syn. illuminata (van Son, 1963); original combination: Acraea axina ♀ f. illuminata.

• braesia (Godman, 1885); original combination: Acraea braesia

  • syn. leucosoma (Staudinger, 1885); original combination: Acraea leucosoma.

  • syn. mystica (Neave, 1904); original combination: Acraea mystica.

  • syn. ochracea (Le Doux, 1931); original combination: Acraea nohara ochracea.

  • syn. lucida (Talbot, 1932); original combination: Acraea bresia[sic] lucida.

  • syn. leucofasciata (Stoneham, 1943); original combination: Acraea braesia leucofasciata.

• buettneri (Rogenhofer, 1890); original combination: Acraea büttneri [= buettneri]

  • syn. felina (Trimen, 1891); original combination: Acraea felina.

  • syn. parapetraea (Schouteden, 1919); original combination: Acraea buettneri parapetraea.

  • syn. contracta (Le Doux, 1923); original combination: Acraea büttneri ♀ f. contracta.

  • syn. nigroapicalis (Overlaet, 1955); original combination: Acraea buettneri f. nigroapicalis.

• caecilia (Fabricius, 1781)

  • ssp. caecilia (Fabricius, 1781); original combination: Papilio caecilia

    • syn. hypatia (Drury, 1782)

    • syn. artemisa (Stoll [1790])

    • syn. bendis (Hübner [1819])

    • syn. varia (Le Doux, 1923)

  • ssp. kulal (van Someren, 1936); original combination: Acraea caecilia kulal

• caldarena (Hewitson, 1877)

  • ssp. caldarena (Hewitson, 1877); original combination: Acraea caldarena

    • syn. amphimalla (Westwood, 1881)

    • syn. dircaea (Westwood, 1881)

    • syn. nero (Butler, 1883)

    • syn. recaldana (Suffert, 1904)

    • syn. mediofasciata (Neustetter, 1916)

    • syn. latiapicalis (Joicey & Talbot, 1921)

    • syn. pallida (Le Doux, 1923)

    • syn. necessaria (Le Doux, 1923)

    • syn. obscuroides (Le Doux, 1923)

    • syn. decepta (Le Doux, 1923)

    • syn. kohambullensis (Le Doux, 1923)

    • syn. marginipunctata (Le Doux, 1931)

  • ssp. neluska (Oberthür, 1878); original combination: Acraea oncaea var. neluska

    • syn. ombria (Weymer, 1892)

• doubledayi (Guérin-Méneville, 1849)

  • ssp. azvaki (d’Abrera, 1980); original combination: Acraea doubledayi azvaki

    • syn. arabica (Eltringham, 1912)

  • ssp. doubledayi (Guérin-Méneville, 1849); original combination: Acraea doubledayi

    • syn. gaekwari (Sharpe, 1901)

    • syn. rileyi (Eltringham, 1913)

• ella (Eltringham, 1911); original combination: Acraea ella

• equatorialis (Neave, 1904)

  • ssp. anaemia (Eltringham, 1912); original combination: Acraea equatorialis anaemia

  • ssp. equatorialis (Neave, 1904); original combination: Acraea doubledayi equatorialis

• intermediodes (Ackery, 1995); original combination: Acraea (Acraea) intermediodes

  • syn. intermedia (Wichgraf, 1908); original combination: Acraea intermedia.

• leucopyga (Aurivillius, 1904); original combination: Acraea leucopyga

  • syn. liszti (Suffert, 1904); original combination: Acraea liszti.

  • syn. propagata (Le Doux, 1923); original combination: Acraea leucopyga ♀ f. propagata.

  • syn. brunnea (Overlaet, 1955); original combination: Acraea leucopyga latiapicalis f. brunnea.

  • syn. albescens (Overlaet, 1955); original combination: Acraea leucopyga latiapicalis f. albescens.

• lyci (Pierre, 2006); original combination: Acraea lyci

• lygus (Druce, 1875); original combination: Acraea lygus

• marnois (Rogenhofer, 1890); original combination: Acraea marnois

• mirabilis (Butler, 1886); original combination: Acraea mirabilis

• miranda (Riley, 1920); original combination: Acraea miranda

  • syn. selousi (Riley, 1920); original combination: Acraea miranda ♀ f. selousi.

• natalica (Boisduval, 1847); original combination: Acraea natalica

  • syn. bellua (Wallengren, 1857); original combination: Acraea bellua.

  • syn. umbrata (Suffert, 1904); original combination: Acraea natalica umbrata.

  • syn. albida (Aurivillius, 1913); original combination: Acraea natalica ♀ ab. albida.

  • syn. mesoleuca (Wichgraf, 1914); original combination: Acraea natalica ♀ f. mesoleuca.

  • syn. dispar (Schouteden, 1919); original combination: Acraea natalica pseudagina ab. dispar.

  • syn. albiventris (Le Doux, 1923); original combination: Acraea natalica natalica f. albiventris.

  • syn. albata (Le Doux, 1923); original combination: Acraea natalica natalica ♂ f. albata.

  • syn. oatesi (van Son, 1936); original combination: Acraea natalica var. oatesi.

• oncaea (Hopffer, 1855)

  • ssp. idjwi (Bernaud & Collins, 2021); original combination: Acraea oncaea idjwi Refs.: Bernaud (2022).

  • ssp. oncaea (Hopffer, 1855); original combination: Acraea oncaea

    • syn. caoncius (Suffert, 1904)

    • syn. modesta (Suffert, 1904)

    • syn. obscura (Suffert, 1904)

    • syn. alboradiata (Suffert, 1904)

    • syn. defasciata (Suffert, 1904)

    • syn. liacea (Suffert, 1904)

    • syn. distincta (Le Doux, 1923)

  • ssp. shimba (Bernaud & Collins, 2021); original combination: Acraea oncaea shimba Refs.: Bernaud (2022).

• pseudegina (Westwood [1852]); original combination: Acraea pseudegina

  • syn. abadima (Ribbe, 1889); original combination: Acraea abadima.

  • syn. clarei (Neave, 1904); original combination: Acraea clarei.

  • syn. stephanophora (Le Cerf, 1927); original combination: Acraea natalica pseudegina f. indiv. stephanophora.

  • syn. inexpectata (Le Doux, 1931); original combination: Acraea natalica abadima ♂ f. inexpectata.

• pudora (Aurivillius, 1910); original combination: Acraea caecilia f. pudora Refs.: Williams and Hennig (2023) (to species from subspecies of caecilia Fabricius).

  • syn. umbrina (Aurivillius, 1910); original combination: Acraea caecilia ab. umbrina.

  • syn. nuda (Wichgraf, 1914); original combination: Acraea caecilia ab. nuda.

• pudorella (Aurivillius [1899])

  • ssp. detecta (Neave, 1010); original combination: Acraea detecta Refs.: Williams and Hennig (2023) (to subspecies from synonym).

  • ssp. pudorella (Aurivillius [1899]); original combination: Acraea caldarena var. pudorella

    • syn. reducta (Wichgraf, 1918)

    • syn. heringi (Le Doux, 1923)

• regalis (Oberthür, 1893); original combination: Acraea regalis

• rhodesiana (Wichgraf, 1909); original combination: Acraea rhodesiana

  • syn. mima (Neave, 1910); original combination: Acraea mima.

  • syn. flaviapicalis (Overlaet, 1955); original combination: Acraea rhodesiana ♀ f. flaviapicalis.

• rogersi (Hewitson, 1873)

  • ssp. lankesteri (Carpenter, 1941); original combination: Acraea rogersi lankesteri

  • ssp. rogersi (Hewitson, 1873); original combination: Acraea rogersi

    • syn. salambo (Grose-Smith, 1887)

    • syn. ehmckei (Dewitz, 1889)

    • syn. lamborni (Eltringham, 1912)

• stenobea (Wallengren, 1860); original combination: Acraea stenobea

  • syn. acronycta (Westwood, 1881); original combination: Acraea acronycta.

  • syn. albomaculata (Weymer, 1892); original combination: Acraea albomaculata.

• sykesi (Sharpe, 1902); original combination: Acraea sykesi

  • syn. candida (Eltringham, 1912); original combination: Acraea doubledayi ♀ f. candida.

• zoumi (Pierre, 1995); original combination: Acraea zoumi

Genus Telchinia

Genus: Telchinia Hübner [1819]

Type species: Telchinia serena (Fabricius, 1775); original combination: Papillio serena Fabricius, 1775

Synonyms:

• Hyalites Doubleday [1848]. Type species: Telchinia encedon (Linnaeus, 1758); original combination: Papilio lycia Fabricius, 1758. Refs.: Doubleday ([1848]): 140
• Gnesia Doubleday [1848]. Type species: Telchinia circeis (Drury, 1782); original combination: Papilio circeis Drury, 1782. Refs.: Doubleday ([1848]): 141
• Pareba Doubleday [1848]. Type species: Telchinia issoria Hübner, [1819]; original combination: Papilio vesta Fabricius, 1787. Refs.: Doubleday ([1848]): 142
• Planema Doubleday [1848]. Type species: Telchinia lycoa (Godart, 1819); original combination: Acraea lycoa Godart, 1819. Refs.: Doubleday ([1848]): 140

References: Hübner ([1819]): 27

108 species:

• acerata (Hewitson, 1874); original combination: Acraea acerata

  • syn. vinidia (Hewitson, 1874); original combination: Acraea vinidia.

  • syn. tenella (Rogenhofer, 1891); original combination: Acraea (Telchinia) tenella.

  • syn. abbottii (Holland, 1892); original combination: Acraea abbottii.

  • syn. abbottii (Holland, 1896); original combination: Acraea abbottii.

  • syn. brahmsi (Suffert, 1904); original combination: Acraea brahmsi.

  • syn. vinidia (Suffert, 1904); original combination: Acraea vinidia diavina.

  • syn. pullula (Grünberg, 1912); original combination: Acraea pullula.

  • syn. vinidia ab. ruandae (Grünberg, 1912); original combination: Acraea vinidia ab. ruandae.

  • syn. vinidia ab. burigensis (Strand, 1913); original combination: Acraea acerata vinidia ab. burigensis.

  • syn. enella f. alluaudi (Le Cerf, 1927); original combination: Acraea acerata tenella f. alluaudi.

  • syn. hoursti (Le Cerf, 1927); original combination: Acraea acerata hoursti.

• actinotina (Lathy, 1903); original combination: Telipna actinotina

• acuta (Howarth, 1969)

  • ssp. acuta (Howarth, 1969); original combination: Acraea ansorgei acuta

    • syn. elgonensis (Howarth, 1969)

    • syn. f. flava (Howarth, 1969)

    • syn. f. alba (Howarth, 1969)

    • syn. f. chrysippina (Howarth, 1969)

    • syn. f. handmani (Howarth, 1969)

    • syn. f. cottrelli (Howarth, 1969)

    • syn. f. australis (Howarth, 1969)

    • syn. f. reducta (Howarth, 1969)

    • syn. f. radiata (Howarth, 1969)

  • ssp. ngorongoro (Kielland, 1990); original combination: Acraea acuta ngorongoro

  • ssp. nigromaculata (Kielland, 1990); original combination: Acraea acuta nigromaculata

  • ssp. rubrobasalis (Kielland, 1990); original combination: Acraea acuta rubrobasalis

    • syn. f. rubrobasalis (Howarth, 1969)

• alalonga (Henning & Henning, 1996); original combination: Hyalites (Auracraea) alalonga

• alberta (Eltringham, 1911); original combination: Acraea cinerea alberta. Type loc.: L. Albert Edward. Refs.: Williams and Hennig (2023) (as species).

• alciope (Hewitson [1852]); original combination: Acraea alciope

  • syn. - (Butler, 1868); original combination: Planema macarina.

  • syn. - (Ward, 1873); original combination: Acraea cydonia.

  • syn. - (Eltringham, 1912); original combination: Acraea alciope ♀ f. cretacea.

  • syn. - (Eltringham, 1912); original combination: Acraea alciope ♀ f. fumida.

  • syn. - (Strand, 1914); original combination: Acraea alciope f. edea.

  • syn. - (Strand, 1914); original combination: Acraea alciope f. lomana.

  • syn. - (Dufrane, 1945); original combination: Acraea alciope alciope ♀ ab. alberici.

  • syn. - (Birket-Smith, 1960); original combination: Acraea alciope alciope f. vulpinopsis.

• alciopoides (Joicey & Talbot, 1921); original combination: Acraea disjuncta f. alciopoides

• alicia Sharpe, 1890

  • ssp. alicia Sharpe, 1890; original combination: Telchinia alicia

    • syn. cappadox (Oberthür, 1893)

    • syn. planesium (Oberthür, 1893)

    • syn. f. cabiroides (Poulton, 1908)

    • syn. f. tenelloides (Poulton, 1908)

    • syn. ab. interruptana (Strand, 1914)

    • syn. ab. ornata (Dufrane, 1945)

    • syn. ab. suffusa (Dufrane, 1945)

  • ssp. mbulu (Kielland, 1990); original combination: Acraea alicia mbulu

  • ssp. uzungwae (Kielland, 1990); original combination: Acraea alicia uzungwae

• althoffi (Dewitz, 1889)

  • ssp. althoffi (Dewitz, 1889); original combination: Acraea althoffi

  • ssp. bitjana (Bethune-Baker, 1926); original combination: Acraea althoffi f. bitjana

  • ssp. neavei (Poulton, 1924); original combination: Acraea althoffi neavei

    • syn. f. telloides (Eltringham, 1912)

    • syn. f. ochreata (Eltringham, 1912)

    • syn. f. drucei (Eltringham, 1912)

    • syn. f. lycioides (Carpenter, 1931)

    • syn. f. budongoensis (Carpenter, 1935)

    • syn. f. albicans (Stoneham, 1936)

    • syn. f. albireducta (Stoneham, 1936)

    • syn. f. ochrafasciata (Stoneham, 1936)

    • syn. f. ochrareducta (Stoneham, 1936)

    • syn. f. ochramaculata (Stoneham, 1936)

    • syn. f. heliconioides (Stoneham, 1936)

  • ssp. rubrofasciata (Aurivillius, 1895); original combination: Acraea althoffi var. rubrofasciata

• amicitiae (Heron, 1909); original combination: Acraea amicitiae

  • syn. polychroma (Rebel, 1911); original combination: Acraea polychroma.

  • syn. f. flavina (Jackson, 1956); original combination: Acrea[sic] amicitiae polychroma ♀ f. flavina.

  • syn. f. lutea (Berger, 1981); original combination: Acraea amicitiae ♀ f. lutea.

  • syn. f. alba (Berger, 1981); original combination: Acraea amicitiae ♀ f. alba.

  • syn. f. intermedia (Berger, 1981); original combination: Acraea amicitiae ♀ f. intermedia.

• anacreon (Trimen, 1868); original combination: Acraea anacreon

• anacreontica (Grose-Smith, 1898)

  • ssp. anacreontica (Grose-Smith, 1898); original combination: Acraea anacreontica

  • ssp. chyulu (van Someren, 1939); original combination: Acraea anacreon chyulu

• ansorgei (Grose-Smith, 1898); original combination: Acraea ansorgei

  • syn. conjuncta (Grose-Smith, 1898); original combination: Acraea conjuncta.

  • syn. nandensis (Sharpe, 1899); original combination: Planema nandensis.

  • syn. f. silacea (Eltringham, 1912); original combination: Acraea conjuncta ♀ f. silacea.

  • syn. f. mutata (Eltringham, 1912); original combination: Acraea conjuncta ♀ f. mutata.

  • syn. f. pica (Eltringham, 1912); original combination: Acraea conjuncta ♀ f. pica.

  • syn. f. lutealba (Eltringham, 1912); original combination: Acraea conjuncta ♀ f. lutealba.

  • syn. f. suffusa (Eltringham, 1912); original combination: Acraea conjuncta ♀ f. suffusa.

  • syn. f. interrupta (Eltringham, 1912); original combination: Acraea conjuncta f. interrupta.

  • syn. f. aurivilliana (Bryk, 1925); original combination: Acrea[sic] ansorgei f. aurivilliana.

  • syn. f. loveni (Bryk, 1925); original combination: Acrea[sic] ansorgei f. loveni.

  • syn. f. jefferyi (Bryk, 1925); original combination: Acrea[sic] ansorgei f. jefferyi.

  • syn. f. aurata (Bryk, 1931); original combination: Acraea ansorgei ♀ f. aurata.

  • syn. f. vansomereni (Bryk, 1931); original combination: Acraea ansorgei ♀ f. vansomereni.

  • syn. bantu (Bryk, 1931); original combination: Acraea ansorgei bantu.

  • syn. f. paulinae (Bryk, 1931); original combination: Acraea ansorgei f. paulinae.

  • syn. f. bryki (Le Doux, 1931); original combination: Acraea ansorgei f. ♀ bryki.

  • syn. f. rufoniger (van Someren, 1936); original combination: Acraea ansorgei f. rufoniger.

  • syn. f. luteflava (van Someren, 1936); original combination: Acraea ansorgei f. luteflava.

  • syn. f. tricolor (van Someren, 1936); original combination: Acraea ansorgei f. tricolor.

  • syn. f. flavipuncta (van Someren, 1936); original combination: Acraea ansorgei f. flavipuncta.

  • syn. f. adaurantica (van Someren, 1936); original combination: Acraea ansorgei f. adaurantica.

  • syn. f. chryssipoides (van Someren, 1936); original combination: Acraea ansorgei f. chryssipoides.

  • syn. f. flaveola (van Someren, 1936); original combination: Acraea ansorgei f. flaveola.

  • syn. f. wickhami (Gabriel, 1939); original combination: Acraea ansorgei f. wickhami.

  • syn. f. uniformis (Gabriel, 1939); original combination: Acraea ansorgei ♀ f. uniformis.

  • syn. f. elgonensis (Howarth, 1969); original combination: Acraea ansorgei ansorgei ♀ f. elgonensis.

  • syn. f. jacksoni (Howarth, 1969); original combination: Acraea ansorgei ansorgei ♀ f. jacksoni.

  • syn. f. gabrieli (Howarth, 1969); original combination: Acraea ansorgei ansorgei ♀ f. gabrieli.

• aubyni (Eltringham, 1912); original combination: Acraea aubyni

• aurivillii (Staudinger, 1896)

  • ssp. aurivillii (Staudinger, 1896); original combination: Acraea (Planema) aurivillii

    • syn. alicia (Grose-Smith, 1900)

    • syn. ab. latifasciata (Grünberg, 1910)

    • syn. ab. bakossua (Strand, 1912)

    • syn. f. tella (Eltringham, 1912)

    • syn. var. smithi (Aurivillius, 1922)

    • syn. bombensis (Stoneham, 1937)

    • syn. ochrextensa (Stoneham, 1937)

    • syn. f. flavifasciata (Stoneham, 1937)

  • ssp. schecana (Rothschild & Jordan, 1905); original combination: Acraea alciope schecana

    • syn. f. vidua (Ungemach, 1932)

• bankoides (Collins & Bernaud, 2021); original combination: Acraea bankoides Refs.: Bernaud (2022): 32.

• baxteri (Sharpe, 1902)

  • ssp. baxteri (Sharpe, 1902); original combination: Acraea baxteri

    • syn. fuelleborni (Thurau, 1903)

    • syn. baxteri subsquamia f. lutea (Carpenter & Jackson, 1950)

    • syn. baxteri philos (Le Cerf, 1933)

  • ssp. oldeani (Kielland, 1990); original combination: Acraea baxteri oldeani

  • ssp. subsquamia (Thurau, 1903); original combination: Acraea fülleborni var. subsquamia

• bergeri (Gaede, 1915); original combination: Acraea bergeri

• bomba (Grose-Smith, 1898); original combination: Acraea bomba

• bonasia (Fabricius, 1775)

  • ssp. banka (Eltringham, 1912); original combination: Acraea bonasia banka

  • ssp. bonasia (Fabricius, 1775); original combination: Papilio bonasia

    • syn. cynthius (Drury, 1782)

    • syn. praeponina (Staudinger, 1896)

    • syn. siabona (Suffert, 1904)

    • syn. ab. implicata (Schultze, 1923)

    • syn. ab. tristis (Schultze, 1923)

    • syn. f. flavistrigata (Le Cerf, 1927)

    • syn. bonasia ab. obscura (Dufrane, 1945)

• burgessi (Jackson, 1956); original combination: Acraea burgessi

• burni (Butler, 1896); original combination: Acraea burni

  • syn. meyeri (van Son, 1963); original combination: Acraea obeira meyeri.

  • syn. meyerana (Ackery, 1995); original combination: Acraea meyerana.

• buschbecki (Dewitz, 1889); original combination: Acraea buschbecki

  • syn. zaire (Rogenhofer, 1889); original combination: Acraea zaire.

• cabira (Hopffer, 1855); original combination: Acraea cabira

  • syn. apecida (Oberthür, 1893); original combination: Acraea apecida.

  • syn. var. flavomaculatus (Lanz, 1896); original combination: Acraea apecida var. flavomaculatus.

  • syn. var. natalensis (Staudinger, 1896); original combination: Acraea cabira var. natalensis.

  • syn. biraca (Suffert, 1904); original combination: Acraea cabira biraca.

  • syn. var. abrupta (Grünberg, 1910); original combination: Acraea apecida var. abrupta.

  • syn. var. swinburnei (Stevenson, 1940); original combination: Acraea cabira apecida var. swinburnei.

• calida (Butler, 1878); original combination: Acraea calida

• cinerea (Neave, 1904); original combination: Acraea cinerea

  • syn. luluae (Berger, 1981); original combination: Acraea cinerea luluae.

• circeis (Drury, 1782); original combination: Papilio circeis

  • syn. mandane (Fabricius, 1793); original combination: Papilio mandane.

  • syn. opis (Herbst, 1793); original combination: Papilio opis.

  • syn. leona (Staudinger, 1896); original combination: Acraea leona.

  • syn. igola leonina (Bethune-Baker,, 1903); original combination: Acraea igola leonina.

  • syn. servona f. reversa (Eltringham, 1912); original combination: Acraea servona f. reversa.

• comor (Pierre, 1992); original combination: Acraea comor

• conradti (Oberthür, 1893)

  • ssp. conradti (Oberthür, 1893); original combination: Acraea conradti

    • syn. ab. flavescens (Blachier, 1912)

    • syn. salmonea (Le Doux, 1922)

    • syn. f. maculosa (Le Doux, 1928)

    • syn. f. usambarensis (Le Doux, 1928)

    • syn. f. vossleri (Le Doux, 1928)

  • ssp. kuekenthalie Le Doux, 1922; original combination: Acraea kükenthali [= kuekenthali]

• disjuncta (Grose-Smith, 1898); original combination: Acraea disjuncta

  • syn. kigeziensis (Jackson, 1956); original combination: Acraea disjuncta kigeziensis.

• encedana (Pierre, 1976); original combination: Acraea encedana

  • syn. ab. alcippina (Aurivillius, 1898); original combination: Acraea encedon ab. alcippina.

  • syn. ab. radiata (Aurivillius, 1905); original combination: Acraea encedon ab. radiata.

  • syn. f. dairalcippa (Le Doux, 1923); original combination: Acraea encedon ♀ f. dairalcippa.

  • syn. f. micropunctata (Le Doux, 1931); original combination: Acraea encedon f. micropunctata.

  • syn. f. macropunctata (Le Doux, 1931); original combination: Acraea encedon f. macropunctata.

  • syn. f. radiofasciata (Stoneham, 1943); original combination: Acraea encedon f. radiofasciata.

  • syn. f. dairana (Pierre, 1976); original combination: Acraea encedana ♀ f. dairana.

• encedon (Linnaeus, 1758)

  • ssp. encedon (Linnaeus, 1758); original combination: Papilio encedon

    • syn. lycia (Fabricius, 1775)

    • syn. sganzini (Boisduval, 1833)

    • syn. lycia var. fulva (Doubleday [1848])

    • syn. daira (Godman & Salvin, 1884)

    • syn. braunei (Staudinger, 1885)

    • syn. lycia ab. infuscata (Staudinger, 1885)

    • syn. usagarae (Vuillot, 1891)

    • syn. lycia ab. ascrepticia (Strand, 1912)

    • syn. ab. fumosa (Aurivillius, 1913)

    • syn. f. commixta (Poulton, 1913)

    • syn. f. alcippoides (Le Doux, 1923)

    • syn. var. nigeriae (Schultze, 1923)

    • syn. f. perradiata (Le Cerf, 1927)

    • syn. f. carbonaria (Le Cerf, 1927)

    • syn. f. poultoni (Le Doux, 1931)

    • syn. f. infuscatoides (Le Doux, 1931)

    • syn. f. umbratalcippina (Le Doux, 1931)

    • syn. f. lycoides (Le Doux, 1931)

    • syn. f. fulvoides (Le Doux, 1931)

    • syn. encedonoides (Le Doux, 1931)

    • syn. f. aurantiaca (Le Doux, 1931)

  • ssp. rathjensi (Le Doux, 1933); original combination: Acraea encedon rathjensi

• encoda (Pierre, 1981); original combination: Acraea encoda

• esebria (Hewitson [1861]); original combination: Acraea esebria

  • syn. protea (Trimen, 1866); original combination: Acraea protea.

  • syn. arctifascia (Butler, 1874); original combination: Planema arctifascia.

  • syn. pseudoprotea (Butler, 1874); original combination: Planema pseudoprotea.

  • syn. amphiprotea (Butler, 1874); original combination: Planema amphiprotea.

  • syn. metaprotea (Butler, 1874); original combination: Planema metaprotea.

  • syn. monteironis (Butler, 1874); original combination: Plenama monteironis.

  • syn. jacksoni (Sharpe, 1890); original combination: Planema jacksoni.

  • syn. ertli (Aurivillius, 1904); original combination: Acraea ertli.

  • syn. f. nubilata (Eltringham, 1912); original combination: Acraea esebria ♀ f. nubilata.

  • syn. jodutta f. inaureata (Eltringham, 1912); original combination: Acraea jodutta ♀ f. inaureata.

  • syn. f. kibwezia (Strand, 1913); original combination: Acraea esebria f. kibwezia.

  • syn. var. actinotis (Neustetter, 1916); original combination: Acraea esebria var. actinotis.

  • syn. f. swynnertoni (O’Neal, 1919); original combination: Acraea esebria f. swynnertoni.

  • syn. f. victoris (Poulton, 1927); original combination: Acraea esebria ♀ f. victoris.

  • syn. f. flavibrunnea (Stoneham, 1943); original combination: Acraea esebria f. flavibrunnea.

  • syn. f. pallidibrunnea (Stoneham, 1943); original combination: Acraea esebria f. pallidibrunnea.

• excelsior (Sharpe, 1891)

  • ssp. excelsior (Sharpe, 1891); original combination: Acraea excelsior

  • ssp. usambarae (Jackson, 1951); original combination: Acraea excelsior usambarae

• fornax (Butler, 1879); original combination: Acraea fornax

  • syn. smithii (Mabille [1880]); original combination: Acraea smithii.

  • syn. var. blachieri (Oberthür, 1916); original combination: Acraea fornax var. blachieri.

• goetzei (Thurau, 1903); original combination: Acraea goetzei

  • syn. byatti (Neave, 1904); original combination: Acraea byatti.

• grosvenori (Eltringham, 1912); original combination: Acraea grosvenori

• guichardi (Gabriel, 1949); original combination: Acraea guichardi

• hecqui (Berger, 1981); original combination: Acraea hecqui

• humilis (Sharpe, 1897); original combination: Acraea humilis

• igola (Trimen, 1889); original combination: Acraea igola

  • syn. maculiventris (Grose-Smith & Kirby, 1894); original combination: Acraea maculiventris.

  • syn. f. turbata (Le Doux, 1923); original combination: Acraea igola f. turbata.

  • syn. f. fasciola (Le Doux, 1923); original combination: Acraea igola f. fasciola.

• induna (Trimen, 1895)

  • ssp. induna (Trimen, 1895); original combination: Acraea induna

  • ssp. salmontana (Henning & Henning, 1996); original combination: Hyalites (Auracraea) induna salmontana

• insularis (Sharpe, 1893); original combination: Acraea insularis

• issoria Hübner [1819]

  • ssp. issoria Hübner [1819]; original combination: Telchinia issoria

    • syn. vesta (Fabricius, 1787)

    • syn. anomala (Kollar, 1848)

    • syn. vesta f. le-douxi (Bryk, 1928)

  • ssp. formosana (Fruhstorfer, 1914); original combination: Pareba vesta formosana

  • ssp. sordice (Fruhstorfer, 1914); original combination: Pareba vesta sordice

  • ssp. vestalina (Fruhstorfer, 1906); original combination: Pareba vesta f. vestalina

  • ssp. vestita (de Nicéville, 1895); original combination: Pareba vestita

    • syn. alticola (Fruhstorfer, 1906)

  • ssp. vestoides (Moore, 1901); original combination: Pareba vestoides

    • syn. vesta narona (Fruhstorfer, 1914)

    • syn. vesta f. arsa (Fruhstorfer, 1914)

• iturina (Grose-Smith, 1890); original combination: Acraea iturina

  • syn. iturinoides (Stoneham, 1936); original combination: Acraea iturinoides.

  • syn. f. sinepeduncles (Stoneham, 1936); original combination: Acraea iturina f. sinepeduncles.

  • syn. f. albicolor (Stoneham, 1943); original combination: Acraea iturina f. albicolor.

• jodutta (Fabricius, 1793)

  • ssp. aethiops (Rothschild & Jordan, 1905); original combination: Acraea jodutta aethiops

    • syn. jodutta ab. clara (Ungemach, 1932)

  • ssp. jodutta (Fabricius, 1793); original combination: Papilio jodutta

    • syn. carmentis (Doubleday [1848])

    • syn. flava (Dewitz, 1879)

    • syn. dorotheae (Sharpe, 1902)

    • syn. f. interjecta (Eltringham, 1912)

    • syn. f. subfulva (Eltringham, 1912)

    • syn. f. castanea (Eltringham, 1912)

    • syn. f. joduttana (Strand, 1914)

    • syn. ab. integra (Schultze, 1923)

• johnstoni (Godman, 1885); original combination: Acraea johnstoni

  • syn. confusa (Rogenhofer, 1891); original combination: Planema confusa.

  • syn. telekiana (Rogenhofer, 1891); original combination: Planema telekiana.

  • syn. proteina (Oberthür, 1893); original combination: Acraea proteina.

  • syn. proteina var. semifulvescens (Oberthür, 1893); original combination: Acraea proteina var. semifulvescens.

  • syn. proteina var. flavescens (Oberthür, 1893); original combination: Acraea proteina var. flavescens.

  • syn. proteina var. semialbescens (Oberthür, 1893); original combination: Acraea proteina var. semialbescens.

  • syn. proteina fulvescens (Oberthür, 1893); original combination: Acraea proteina fulvescens.

  • syn. octobalia (Karsch, 1894); original combination: Acraea octobalia.

  • syn. silvicola (Richelmann, 1913); original combination: Acraea silvicola.

  • syn. f. valdemaculosa (Carpenter, 1932); original combination: Acraea johnstoni f. valdemaculosa.

  • syn. f. pretiosa (Carpenter, 1932); original combination: Acraea johnstoni f. pretiosa.

• kaduna (Pierre, 1993)

• kakana (Eltringham, 1911); original combination: Acraea iturina kakana

  • syn. ab. infernalis (Ungemach, 1932); original combination: Acraea kakana ab. infernalis.

• kalinzu (Carpenter, 1936); original combination: Acraea kalinzu

  • syn. var. albipicta (Carpenter, 1936); original combination: Acraea kalinzu var. albipicta.

• lia (Mabille, 1879); original combination: Acraea lia

• lumiri (Bethune-Baker, 1908); original combination: Acraea lumiri

  • syn. var. camerunica (Strand, 1914); original combination: Acraea lumiri var. camerunica.

• lusinga (Overlaet, 1955); original combination: Acraea anacreon lusinga

• lycoa (Godart [1819])

  • ssp. aequalis (Rothschild & Jordan, 1905); original combination: Acraea lycoa aequalis

  • ssp. fallax (Rogenhofer, 1891); original combination: Planema fallax

  • ssp. kenia (Eltringhsm, 1911); original combination: Acraea lycoa kenia

• masamba (Ward, 1872); original combination: Acraea masamba

  • syn. ruepelli (Saalmüller, 1878); original combination: Acraea rüppelli.

  • syn. boseae (Saalmüller, 1880); original combination: Acraea boseae.

  • syn. vulgaris (Oberthür, 1916); original combination: Acraea masamba r.? vulgaris.

  • syn. torquata (Le Cerf, 1927); original combination: Acraea masamba f. torquata.

• masaris (Oberthür, 1893)

  • ssp. jodina (Pierre, 1992); original combination: Acraea masaris jodina

  • ssp. masaris (Oberthür, 1893); original combination: Acraea masaris

• melanoxantha (Sharpe, 1891); original combination: Acraea melanoxantha

• mirifica (Lathy, 1906); original combination: Acraea mirifica

• necoda (Hewitson [1861]); original combination: Acraea necoda

• newtoni (Sharpe, 1893); original combination: Acraea newtoni

• ntebiae (Sharpe, 1897)

  • ssp. dewitzi (Carcasson, 1981); original combination: Acraea ntebiae dewitzi

    • syn. dewitzi (Aurivillius, 1904)

  • ssp. kigoma (Kielland, 1978); original combination: Acraea ntebiae kigoma

  • ssp. ntebiae (Sharpe, 1897); original combination: Acraea ntebiae

    • syn. mairessei (Aurivillius, 1904)

    • syn. melanosticta (Sharpe, 1904)

  • ssp. nyongana (d’Abrera, 1980); original combination: Acraea ntebiae nyongana

    • syn. mairessei ab. nyongana (Strand, 1914)

    • syn. mairessei var. alticola (Schultze, 1923)

• obeira (Hewitson, 1863); original combination: Acraea obeira

  • syn. piva (Guenée, 1864); original combination: Acrea[sic] piva.

  • syn. andromba (Grose-Smith, 1891); original combination: Acraea andromba.

• oberthueri (Butler, 1895), missp.

  • ssp. laetopicta (Rebel, 1914); original combination: Acraea oberthüri ab. laetopicta

  • ssp. oberthueri (Butler, 1895), missp.; original combination: Acraea oberthürii [= oberthuerii]. Type loc.: “Johann-Albrechthöhe, Nord-Camerun”.

    • syn. - (Suffert, 1904)

    • syn. - (Schultze, 1923)

    • syn. - (Le Doux, 1923)

• ochrascens (Sharpe, 1902); original combination: Acraea ochrascens Refs.: Williams and Hennig (2023) (as species).

  • syn. terpsichore var. bukoba (Weymer, 1903); original combination: Acraea terpsichore var. bukoba.

• odzalae (Collins, 1997); original combination: Acraea odzalae

• oreas (Sharpe, 1891)

  • ssp. oboti (Collins & Larsen, 2000); original combination: Acraea oreas oboti

  • ssp. oreas (Sharpe, 1891); original combination: Acraea oreas

    • syn. f. albimaculata (Neave, 1904)

    • syn. angolanus (Lathy, 1906)

    • syn. ab. radians (Aurivillius, 1913)

• orestia (Hewitson, 1874)

  • ssp. orestia (Hewitson, 1874); original combination: Acraea orestia

    • syn. orestina (Plötz, 1880)

    • syn. f. transita (Eltringham, 1912)

    • syn. f. carpenteri (Eltringham, 1913)

    • syn. ab. moliwensis (Strand, 1914)

    • syn. acutipennis (Schultze, 1923)

    • syn. f. clara (Stoneham, 1943)

    • syn. f. pseudohumilis (Stoneham, 1943)

  • ssp. sambar (Stoneham, 1943); original combination: Acraea sambar

• orina (Hewitson, 1874); original combination: Acraea orina

  • syn. oreta (Hewitson, 1874); original combination: Acraea oreta.

  • syn. var. nigroapicalis (Aurivillius, 1893); original combination: Acraea orina var. nigroapicalis.

• orinata (Oberthür, 1893); original combination: Acraea orinata

  • syn. orineta (Eltringham, 1912); original combination: Acraea orina orineta.

• parei (Henning & Henning, 1996)

  • ssp. orangica (Henning & Henning, 1996); original combination: Hyalites (Auracraea) parei orangica

  • ssp. parei (Henning & Henning, 1996); original combination: Hyalites (Auracraea) parei

• parrhasia (Fabricius, 1793)

  • ssp. kenya (van Someren & Rogers, 1926); original combination: Acraea parrhasia kenya

  • ssp. limonata (Eltringham, 1912); original combination: Acraea servona limonata

  • ssp. orientis (Aurivillius, 1904); original combination: Acraea circeis var. orientis

    • syn. ab. depunctella (Strand, 1911)

    • syn. ab. unipunctella (Strand, 1911)

    • syn. ab. semipunctella (Strand, 1911)

    • syn. ab. transienda (Strand, 1911)

  • ssp. parrhasia (Fabricius, 1793); original combination: Papilio parrhasia

    • syn. leona var. parrhoppidia (Staudinger, 1896)

    • syn. f. pseudoppidia (Strand, 1914)

  • ssp. servona (Godart [1819]); original combination: Acraea servona

    • syn. lycoides (Boisduval, 1836)

    • syn. dejana (Godman & Salvin, 1890)

    • syn. rhodina (Rothschild & Jordan, 1905)

    • syn. var. subochreata (Grünberg, 1910)

    • syn. f. rubra (Eltringham, 1912)

    • syn. tenebrosa (Eltringham, 1912)

    • syn. f. digitata (Carpenter, 1935)

• pelopeia (Staudinger, 1896); original combination: Acraea penelope var. pelopeia

  • syn. tropicalis (Blachier, 1912); original combination: Acraea tropicalis.

  • syn. var. lineata (Aurivillius, 1925); original combination: Acraea pelopeia var. lineata.

• peneleos (Ward, 1871)

  • ssp. gelonica (Rothschild & Jordan, 1905); original combination: Acraea peneleos gelonica

  • ssp. pelasgia (Grose-Smith, 1900); original combination: Acraea pelasgius

  • ssp. peneleos (Ward, 1871); original combination: Acraea peneleos

    • syn. fenelos (Aurivillius, 1893)

    • syn. f. helvimaculata (Eltringham, 1912)

    • syn. f. lactimaculata (Eltringham, 1912)

    • syn. f. sepia (Eltringham, 1912)

    • syn. f. pseudopelasgius (Strand, 1914)

    • syn. f. peneloides (Strand, 1914)

    • syn. ab. castanea (Schultze, 1923)

    • syn. ab. strigipygida (Birket-Smith, 1960)

• penelope (Staudinger, 1896)

  • ssp. derubescens (Eltringham, 1912); original combination: Acraea penelope derubescens

    • syn. translucida (Eltringham, 1912)

  • ssp. penelope (Staudinger, 1896); original combination: Acraea penelope

    • syn. pomponia (Grose-Smith, 1900)

    • syn. f. argentea (Eltringham, 1912)

    • syn. f. exalbescens (Eltringham, 1912)

    • syn. f. penella (Eltringham, 1912)

    • syn. ab. sulphurescens (Aurivillius, 1913)

    • syn. f. laeticolor (Le Doux, 1928)

    • syn. f. josetta (Jackson, 1951)

    • syn. vitrea (Eltringham, 1912)

• pentapolis (Ward, 1871)

  • ssp. epidica (Oberthür, 1893); original combination: Acraea epidica

  • ssp. pentapolis (Ward, 1871); original combination: Acraea pentapolis

    • syn. thelestis (Oberthür, 1893)

    • syn. ab. sjoestedti (Bryk, 1917)

• perenna (Doubleday [1847])

  • ssp. kaffana (Rothschild, 1902); original combination: Acraea perenna kaffana

  • ssp. perenna (Doubleday [1847]); original combination: Acraea perenna

    • syn. polydectes (Ward, 1871)

    • syn. thesprio (Oberthür, 1893)

    • syn. ab. usagara (Strand, 1913)

• pharsalus (Ward, 1871)

  • ssp. carmen (Pyrcz, 1991); original combination: Acraea pharsalus carmen

  • ssp. pharsalus (Ward, 1871); original combination: Acraea pharsalus

    • syn. pharsaloides (Holland, 1892)

    • syn. pharsaloides (Holland, 1896)

    • syn. saluspha (Suffert, 1904)

    • syn. var. pharsaloides ab. pallidepicta (Strand, 1911)

    • syn. var. pharsaloides ab. nia (Strand, 1911)

    • syn. f. ruperti (O’Neal, 1919)

    • syn. f. duplicatus (Stoneham, 1943)

  • ssp. rhodina (Rothschild, 1902); original combination: Acraea pharsalus rhodina

• pierrei (Berger, 1981); original combination: Acraea pierrei

• polis (Pierre, 1999); original combination: Acraea polis

• pseudepaea (Dudgeon, 1909)

  • ssp. pseudepaea (Dudgeon, 1909); original combination: Acraea pseudepaea

  • ssp. ziama Belcastro, Boireau & Safian, 2020; original combination: Telchinia pseudepaea ziama. Type loc.: Forêt de Ziama, Guinea. Types: NHMUK. Refs.: Sáfián et al. (2020): 46. Ety.: Ziama Massif

• quirinalis (Grose-Smith, 1900); original combination: Acraea quirinalis

• rahira (Boisduval, 1833)

  • ssp. mufindi (Kielland, 1990); original combination: Acraea rahira mufindi Distr.: Tanzania.

  • ssp. rahira (Boisduval, 1833); original combination: Acraea rahira Distr.: southern Africa, Zimbabwe, Mozambique to Malawi, southern Tanzania, Zambia, southern Zaire (Shaba), Angola, western Kenya, Uganda.

    • syn. rahira f. melanoradiata (Stoneham, 1943)

    • syn. rahira f. crippsi (Stoneham, 1943)

  • ssp. uasingishuensis (Stoneham, 1943); original combination: Acraea rahira uasingishuensis Refs.: Williams and Hennig (2023) (as subspecies).

• rangatana (Eltringham, 1912)

  • ssp. basilewskyi (Berger, 1956); original combination: Acraea ventura basilewskyi

  • ssp. bettiana (Joicey & Talbot, 1921); original combination: Acraea bettiana

    • syn. ab. kissejensis (Joicey & Talbot, 1921)

    • syn. ab. hades (Joicey & Talbot, 1927)

  • ssp. ecketti (Jackson, 1951); original combination: Acraea rangatana ecketti

  • ssp. maji (Carpenter, 1935); original combination: Acraea maji

  • ssp. rangatana (Eltringham, 1912); original combination: Acraea terpsichore f. rangatana

• rileyi (Le Doux, 1931); original combination: Acraea rileyi

• rupicola (Schultze, 1912); original combination: Acraea rupicola

• safie (Felder & Felder [1865])

  • ssp. antinorii (Oberthür, 1880); original combination: Acraea antinorii

    • syn. f. neumanni (Le Doux, 1928)

    • syn. ab. albipuncta (Ungemach, 1932)

    • syn. ab. oumbiana (Ungemach, 1932)

  • ssp. safie (Felder & Felder [1865]); original combination: Acraea safie

    • syn. f. tillini (Gabriel, 1949)

• sambavae (Ward, 1873); original combination: Acraea sambavae

• semivitrea (Aurivillius, 1895); original combination: Acraea semivitrea

  • syn. pervia (Sharpe, 1897); original combination: Acraea pervia.

• serena (Fabricius, 1775); original combination: Papilio serena

  • syn. eponina (Cramer [1780]); original combination: Papilio eponina.

  • syn. janisca (Godart, 1819); original combination: Acraea janisca.

  • syn. manjaca (Boisduval, 1833); original combination: Acraea manjaca.

  • syn. var. rougetii (Guérin-Méneville, 1849); original combination: Acraea serena var. rougetii.

  • syn. buxtoni Butler, 1875; original combination: Telchinia buxtoni.

  • syn. perrupta Butler, 1883; original combination: Telchinia perrupta.

  • syn. subserena (Grose-Smith, 1900); original combination: Acraea subserena.

  • syn. terpsichore ab. venturina (Thurau, 1903); original combination: Acraea terpsichore ab. venturina.

  • syn. terpsichore ab. connexa (Thurau, 1903); original combination: Acraea terpsichore ab. connexa.

  • syn. terpsichore ab. excentrica (Thurau, 1903); original combination: Acraea terpsichore ab. excentrica.

  • syn. intermediana (Strand, 1911); original combination: Acraea intermediana.

  • syn. terpsichore rougeti ab. contraria (Strand, 1912); original combination: Acraea terpsichore rougeti ab. contraria.

  • syn. eliana (Strand, 1912); original combination: Acraea eliana.

  • syn. eliana ab. toka (Strand, 1912); original combination: Acraea eliana ab. toka.

  • syn. terpsichore ab. janiscella (Strand, 1913); original combination: Acraea terpsichore ab. janiscella.

  • syn. eponina f. akoafima (Le Doux, 1928); original combination: Acraea eponina eponina ♀ f. akoafima.

  • syn. eponina f. latifasciata (Le Doux, 1928); original combination: Acraea eponina eponina ♀ f. latifasciata.

  • syn. eponina manjaca f. reducta (Le Doux, 1928); original combination: Acraea eponina manjaca f. reducta.

  • syn. jordani (Le Doux, 1928); original combination: Acraea jordani.

  • syn. terpsichore ab. bankoides (Carpenter, 1935); original combination: Acraea terpsichore ab. bankoides.

  • syn. terpsichore buxtoni ab. subjanisca (Dufrane, 1945); original combination: Acraea terpsichore buxtoni ab. subjanisca.

• silia (Mabille [1885]); original combination: Acraea masamba var. silia

  • syn. r. debilis (Oberthür, 1916); original combination: Acraea masamba r. debilis.

• siliana (Oberthür, 1916); original combination: Acraea siliana

• simulata (Le Doux, 1923); original combination: Acraea simulata

• simulator (Ackery, 1995); original combination: Acraea simulator

  • syn. simulata (Le Doux, 1932), preocc.; original combination: Planema simulata.

• sotikensis (Sharpe, 1892)

  • ssp. karschi (Aurivillius [1899]); original combination: Acraea viviana var. karschi

    • syn. supponina ninapo (Suffert, 1904)

    • syn. karscheni var. cruentata (Gaede, 1916)

    • syn. ab. vulcanica (Schultze, 1923)

  • ssp. sotikensis (Sharpe, 1892); original combination: Acraea sotikensis

    • syn. f. katana (Eltringham, 1912)

    • syn. rowena (Eltringham, 1912)

    • syn. var. hansmeyeri (Strand, 1913)

    • syn. var. rowenina (Gaede, 1915)

    • syn. f.? bayeri (Schouteden, 1919)

    • syn. katerensis (Stoneham, 1943)

    • syn. f. exclamationis (Stoneham, 1943)

    • syn. f. albisubapex (Stoneham, 1943)

    • syn. f. rowena ab. mariae (Dufrane, 1945)

    • syn. f. rowena ab. alberici (Dufrane, 1945)

• speciosa (Wichgraf, 1909); original combination: Acraea anacreon speciosa

• strattipocles (Oberthür, 1893); original combination: Acraea strattipocles

  • syn. var. albescens (Oberthür, 1916); original combination: Acraea strattipocles var. albescens.

  • syn. var. cervina (Oberthür, 1916); original combination: Acraea strattipocles var. cervina.

• supponina (Staudinger, 1896); original combination: Acraea supponina

• toruna (Grose-Smith, 1900); original combination: Acraea toruna

  • syn. lycoa ab. butleri (Aurivillius, 1898); original combination: Acraea lycoa ab. butleri.

• ungemachi (Le Cerf, 1927); original combination: Acraea ungemachi

  • syn. f. nuda (Ungemach, 1932); original combination: Acraea ungemachi f. nuda.

• uvui (Grose-Smith, 1890)

  • ssp. balina (Karsch, 1892); original combination: Acraea balina

  • ssp. uvui (Grose-Smith, 1890); original combination: Acraea uvui

    • syn. minima (Holland, 1892)

    • syn. minima (Holland, 1896)

    • syn. ab. interruptella (Strand, 1909)

    • syn. f. subsuffusa (Stoneham, 1936)

• ventura (Hewitson, 1877); original combination: Acraea ventura

  • syn. serena ab. melas (Sharpe, 1902); original combination: Acraea serena ab. melas.

• vesperalis (Grose-Smith, 1890)

  • ssp. catori (Bethune-Baker, 1904); original combination: Acraea catori

  • ssp. vesperalis (Grose-Smith, 1890); original combination: Acraea vesperalis

    • syn. ab. punctula (Strand, 1914)

    • syn. ab. picta (Schouteden, 1919)

• viviana (Staudinger, 1896); original combination: Acraea viviana

• vuilloti (Mabille, 1888); original combination: Acraea vuilloti Refs.: Mabille (1888): CLXX. Ety.: M. P. Vuillot

• vumbui (Stevenson, 1934); original combination: Acraea conradti r. vumbui

  • syn. vumbui ab. barnesi (Stevenson, 1940); original combination: Acraea conradti vumbui ab. barnesi.

  • syn. vumbui f. cooksoni (van Son, 1963); original combination: Acraea conradti vumbui f. cooksoni.

• wigginsi (Neave, 1904)

  • ssp. occidentalis (Bethune-Baker, 1926); original combination: Acraea wigginsi r. occidentalis

  • ssp. wigginsi (Neave, 1904); original combination: Acraea wigginsi

• zitja (Boisduval, 1833); original combination: Acraea zitja

  • syn. rakeli (Boisduval, 1833); original combination: Acraea rakeli. Refs.: Boisduval (1833 a): 32, pl.4.

  • syn. zitja var. radiata (Guenée, 1864); original combination: Acraea zitja var. radiata.

  • syn. zitja var. fumida (Mabille, 1880); original combination: Acrooea[sic] zitja var. fumida.

Genus Tildia

Genus: Tildia Williams & Henning, 2023

Type species: Tildia zetes (Linnaeus, 1758); original combination: Papilio zetes Linnaeus, 1758

Synonyms: None

References: Williams and Hennig (2023)

13 species:

• acara (Hewitson, 1865)

  • ssp. acara (Hewitson, 1865); original combination: Acraea acara

    • syn. - (Felder & Felder [1865])

    • syn. - (Suffert, 1904)

    • syn. - (Le Doux, 1923)

    • syn. - (Le Cerf, 1927)

    • syn. - (van Son, 1963)

  • ssp. melanophanes (Le Cerf, 1927); original combination: Acraea zetes sufferti f. melanophanes

• anemosa (Hewitson, 1865); original combination: Acraea anemosa

  • syn. arcticincta (Butler, 1883); original combination: Acraea arcticincta.

  • syn. alboradiata (Aurivillius, 1898); original combination: Acraea anemosa ab. alboradiata.

  • syn. interrupta (Thurau, 1903); original combination: Acraea anemosa ab. interrupta.

  • syn. mosana (Suffert, 1904); original combination: Acraea anemosa mosana.

  • syn. dubiosa (Suffert, 1904); original combination: Acraea anemosa dubiosa.

  • syn. discoguttata (Strand, 1909); original combination: Acraea anemosa ab. discoguttata.

  • syn. ufipana (Strand, 1911); original combination: Acraea anemosa ab. ufipana.

  • syn. urungensis (Strand, 1911); original combination: Acraea anemosa ab. urungensis.

  • syn. conjuncta (Niepelt, 1937); original combination: Acraea anemosa f. conjuncta.

  • syn. macrosticta (Storace, 1949); original combination: Acraea anemosa macrosticta.

• barberi (Trimen, 1881); original combination: Acraea barberi

• chilo (Godman, 1880)

  • ssp. chilo (Godman, 1880); original combination: Acraea chilo

    • syn. crystallina (Grose-Smith, 1890)

    • syn. rosina (Rogenhofer, 1891)

    • syn. hoehneli (Holland, 1896)

    • syn. wissmanni (Weymer, 1903)

  • ssp. yemenensis (Le Doux, 1931); original combination: Acraea chilo yemensis

• hypoleuca (Trimen, 1898); original combination: Acraea hypoleuca

• oscari (Rothschild, 1902); original combination: Acraea oscari. Type loc.: Abyssinia.

• pseudolycia (Butler, 1874)

  • ssp. astrigera (Butler, 1899); original combination: Acraea astrigera

    • syn. emini (Weymer, 1903)

    • syn. auasa (Gabriel, 1949)

  • ssp. pseudolycia (Butler, 1874); original combination: Acraea pseudolycia. Type loc.: Angola, Quanza.

    • syn. brunnea (Eltringham, 1911). Type loc.: Angola; Unyoro and Masindi; Entebbe.

• rabbaiae (Ward, 1873)

  • ssp. perlucida (Henning & Henning, 1996); original combination: Acraea (Acraea) rabbaiae perlucida

  • ssp. rabbaiae (Ward, 1873); original combination: Acraea rabbaiae

    • syn. mombasae (Grose-Smith, 1889)

• trimeni (Aurivillius, 1898); original combination: Acraea barberi ab. (et var.?) trimeni

  • syn. eros (Le Doux, 1923); original combination: Acraea zetes barberi f. eros.

  • syn. nigromacula (Le Doux, 1931); original combination: Acraea zetes trimeni f. nigromacula.

• turna (Mabille, 1877); original combination: Acraea turna

  • syn. marmorata (Grose-Smith & Kirby, 1892); original combination: Acraea marmorata.

  • syn. lacteata (Le Cerf, 1927); original combination: Acraea turna lacteata.

  • syn. scioptera (Le Cerf, 1927); original combination: Acraea turna lacteata f. scioptera.

• welwitschii (Rogenhofer, 1892); original combination: Acraea (Telchinia) welwitschii

  • syn. lobemba (Eltringham, 1912); original combination: Acraea welwitschii lobemba.

  • syn. lutea (Riley, 1921); original combination: Acraea welwitschi lutea.

  • syn. nivea (Riley, 1921); original combination: Acraea welwitschi nivea.

• zetes (Linnaeus, 1758)

  • ssp. annobona (d’Abrera, 1980); original combination: Acraea zetes annobona

  • ssp. rudolfi (Eltringham, 1929); original combination: Acraea zetes rudolfi

  • ssp. sidamona (Rothschild & Jordan, 1905); original combination: Acraea zetes sidamona

  • ssp. zetes (Linnaeus, 1758); original combination: Papilio zetes

    • syn. menippe (Drury, 1782)

    • syn. mycenaea (Hübner [1819])

    • syn. jalema (Godart, 1819)

• zonata (Hewitson, 1877); original combination: Acraea zonata

  • syn. makupa (Grose-Smith, 1889); original combination: Acraea makupa.

Tribe Argynnini

Type genus: Argynnis [Fabricius], 1807

Classification: The tribe Argynnini is divided into four subtribes and eight genera (Simonsen 2006 a; Simonsen et al. 2006):

• Subtribe Argynnina: Argynnis, Brenthis, Issoria

• Subtribe Boloriina: Boloria (monobasic)

• Subtribe Euptoietina: Euptoieta (monobasic)

• Subtribe Yrameina: Yramea (monobasic)

• Genus Pardopsis (no subtribe assigned).

References: Warren (1944) (taxonomy); dos Passos and Grey (1945 a) (taxonomy); Warren et al. (1946) (taxonomy); Warren (1955) (taxonomy); Lamas (2004) (Neotropics); Penz and Peggie (2003) (phylogeny); Simonsen (2006 a) (taxonomy, phylogeny); Simonsen et al. (2006) (phylogeny)

Genus Argynnis

Genus: Argynnis Fabricius, 1807

Type species: Argynnis paphia (Linnaeus, 1758); original combination: Papilio paphia Linnaeus, 1758

Synonyms: None

References: Fabricius (1807): 283; De Moya et al. (2017) (phylogeny); Zhang et al. (2020) (genome phylogeny)

Subgenus Argynnis

Subgenus: Argynnis Fabricius, 1807

Type species: Argynnis paphia (Linnaeus, 1758); original combination: Papilio paphia Linnaeus, 1758

Synonyms:

• Argyreus Scopoli, 1777. Type species: Argynnis hyperbius Linnaeus, 1763; original combination: Papilio niphe Linnaeus, 1767 (repl. name). Refs.: Scopoli (1777): 431; ICZN (1945)
• Acidalia Hübner [1819]. Type species: Argynnis hyperbius Linnaeus, 1763; original combination: Papilio niphe Linnaeus, 1767 (repl. name). Refs.: Hübner ([1819]): 31
• Argyronome Hübner [1819]. Type species: Argynnis laodice (Pallas, 1771); original combination: Papilio laodice Pallas, 1771. Refs.: Hübner ([1819]): 32
• Argyrea Billberg, 1820. Type species: Argynnis hyperbius Linnaeus, 1763; original combination: Papilio niphe Linnaeus, 1767 (repl. name). Refs.: Billberg (1820): 77
• Damora Nordmann, 1851. Type species: Argynnis sagana paulina (Nordmann, 1851); original combination: Damora paulina Nordmann, 1851. Refs.: Nordmann (1851): 439, pl.II tab.XII
• Eudaphne Reuss, 1922. Type species: Argynnis laodice (Pallas, 1771); original combination: Papilio laodice Pallas, 1771. Refs.: Reuss (1922): 221 (as subgenus)
• Mimargyra Reuss, 1922. Type species: Argynnis hyperbius (Linnaeus, 1763); original combination: Papilio hyperbius Linnaeus, 1763. Refs.: Reuss (1922): 221 (as subgenus)
• Mesodryas Reuss, 1926. Type species: Argynnis paphia (Linnaeus, 1758); original combination: Papilio paphia Linnaeus, 1758. Refs.: Reuss (1926 a): 435
• Eudryas Reuss, 1926. Type species: Argynnis childreni Gray, 1831; original combination: Argynnis childreni Gray, 1831. Refs.: Reuss (1926): 253; Hemming (1943): 30; Hemming (1967): 173
• Pandoriana Warren, 1942. Type species: Argynnis pandora ([Denis & Schiffermüller], 1775); original combination: Papilio maja Cramer, [1775]. Refs.: Warren (1942): 245-246
• Childrena Hemming, 1943. Type species: Argynnis childreni Gray, 1831; original combination: Argynnis childreni Gray, 1831. Refs.: Hemming (1943): 30
• Nephargynnis Shirôzu & Saigusa, 1973. Type species: Argynnis anadyomene Felder & Felder, 1862; original combination: Argynnis anadyomene Felder & Felder, 1862. Refs.: Shirôzu and Saigusa (1973): 111

References: Fabricius (1807): 283; De Moya et al. (2017) (phylogeny); Zhang et al. (2020) (genome phylogeny)

10 species:

• anadyomene Felder & Felder, 1862

  • ssp. anadyomene Felder & Felder, 1862; original combination: Argynnis anadyomene Distr.: Southeast Russia, China, Korea, Japan (De Moya et al. 2017).

  • ssp. ella Bremer, 1864; original combination: Argynnis ella

    • syn. obliterata Kardakoff, 1928

  • ssp. midas Butler, 1866; original combination: Argynnis midas

  • ssp. prasoides Fruhstorfer, 1907; original combination: Argynnis anadyomene prasoides

• childreni Gray, 1831

  • ssp. caesarea Fruhstorfer, 1912; original combination: Argynnis childreni caesarea

  • ssp. childreni Gray, 1831; original combination: Argynnis childreni

  • ssp. sakontala Kollar, 1848; original combination: Argynnis sakontala

• hyperbius (Linnaeus, 1763)

  • ssp. castetsi Oberthür, 1891; original combination: Argynnis castetsi

  • ssp. centralis Martin, 1913; original combination: Argynnis hyperbius centralis

  • ssp. hyperbius (Linnaeus, 1763); original combination: Papilio hyperbius Refs.: Linnaeus, 1763; Amoenitates Acad. 6: 408.

  • ssp. inconstans Butler, 1873; original combination: Argynnis inconstans

  • ssp. javanica Oberthür, 1889; original combination: Argynnis niphe javanica

  • ssp. neumanni Rothschild, 1902; original combination: Argynnis hyperbius neumanni

  • ssp. niugini Samson, 1976; original combination: Argyreus hyperbius niugini

  • ssp. sagada Fruhstorfer, 1912; original combination: Argynnis hyperbius sagada

  • ssp. sumatrensis Fruhstorfer, 1903; original combination: Argynnis hyperbius sumatrensis

  • ssp. taprobana (Moore, 1900); original combination: Acidalia taprobana

• kuniga (Chou & Tong, 1994); original combination: Argyronome kuniga. Type loc.: Linan, Zhejiang.

• laodice (Pallas, 1771)

  • ssp. ariana Fruhstorfer, 1907; original combination: Argynnis laodice ariana

  • ssp. ferruginea Watkins, 1924; original combination: Argynnis laodice ferruginea Refs.: Watkins (1924): 456.

  • ssp. fletcheri Watkins, 1924; original combination: Argynnis laodice fletcheri Refs.: Watkins (1924): 456.

  • ssp. huochengice (Huang & Murayama, 1992); original combination: Argyronome laodice huochengice

  • ssp. indroides Tytler, 1940; original combination: Argynnis laodice indroides

  • ssp. japonica Ménétriés, 1857; original combination: Argynnis laodice japonica. Type loc.: Japan.

  • ssp. laodice (Pallas, 1771); original combination: Papilio laodice

  • ssp. melloides (Draeseke, 1925); original combination: Dryas laodice f. geogr. melloides

  • ssp. rudra Moore, 1857; original combination: Argynnis rudra

  • ssp. samana Fruhstorfer, 1902; original combination: Argynnis laodice samana

  • ssp. tomaridice Bryk, 1942; original combination: Argynnis laodice tomaridice

• pandora ([Denis & Schiffermüller], 1775)

  • ssp. argentifasciata Kotzsch, 1937; original combination: Argynnis pandora argentifasciata

  • ssp. pandora ([Denis & Schiffermüller], 1775); original combination: Papilio pandora

    • syn. transcaucasica (Moucha, 1967)

  • ssp. pasargades Fruhstorfer, 1908; original combination: Argynnis pandora pasargades

  • ssp. seitzi Fruhstorfer, 1908; original combination: Argynnis maja seitzi

    • syn. chrysobarylla Fruhstorfer, 1909

• paphia (Linnaeus, 1758)

  • ssp. angustia Churkin & Pletnev, 2012; original combination: Argynnis paphia angustia. Type loc.: Kyrgyzstan, Ferghansky Mts. (west. edges), W. Urum-Bash river. Types: DSM. Refs.: Churkin and Pletnev (2012): 114. Ety.: Angustia (L. - a narrow gorge, narrow limits)

  • ssp. argyrophontes Oberthür, 1923; original combination: Argynnis paphia argyrophontes

  • ssp. argyrorrhytes Alphéraky, 1908; original combination: Argynnis paphia ab. argyrorrhytes

  • ssp. delila Röber, 1896; original combination: Argynnis paphia var. delila

  • ssp. dives Oberthür, 1908; original combination: Argynnis paphia var. dives

  • ssp. formosicola Matsumura, 1927; original combination: Argynnis paphia formosicola. Type loc.: Formosa.

  • ssp. geisha Hemming, 1941; original combination: Argynnis paphia geisha

  • ssp. masandarensis Gross & Ebert, 1975; original combination: Argynnis (Argynnis) paphia masandarensis

  • ssp. megalegoria Fruhstorfer, 1907; original combination: Argynnis paphia megalegoria. Type loc.: Szechuan, Yunnan.

  • ssp. neopaphia Fruhstorfer, 1907; original combination: Argynnis paphia neopaphia. Type loc.: Amur, Ussuri, Sakhalin, Kuriles, Transbaikal, Kamchatka.

  • ssp. paphia (Linnaeus, 1758); original combination: Papilio paphia

    • syn. valesina (Esper, 1798)

  • ssp. pusilla Wnukowsky, 1927; original combination: Argynnis paphia pusilla

  • ssp. thalassata Fruhstorfer, 1908; original combination: Argynnis paphia thalassata

  • ssp. tsushimana Fruhstorfer, 1906; original combination: Argynnis paphia tsushimana

  • ssp. virescens Nakahara, 1926; original combination: Argynnis paphia virescens

• ruslana Motschulsky, 1866

  • ssp. lysippe Janson, 1877; original combination: Argynnis lysippe. Type loc.: Japan. Refs.: Janson (1877): 154.

  • ssp. ruslana Motschulsky, 1866; original combination: Argynnis ruslana. Type loc.: Amur. Distr.: Southeast Russia, eastern China, Korea, Japan (De Moya et al. 2017).

• sagana Doubleday [1847]

  • ssp. liane Fruhstorfer, 1907; original combination: Argynnis sagana liane

  • ssp. paulina (Nordman, 1851); original combination: Damora paulina

  • ssp. relicta Korshunov, 1984; original combination: Argynnis sagana relicta

  • ssp. sagana Doubleday [1847]; original combination: Argynnis sagana

• zenobia Leech, 1890

  • ssp. penelope Staudinger [1892]; original combination: Argynnis penelope

  • ssp. zenobia Leech, 1890; original combination: Argynnis zenobia Distr.: Southeast Russia, China, Tibet, Korea (De Moya et al. 2017).

Subgenus Fabriciana

Subgenus: Fabriciana Reuss, 1920

Type species: Fabriciana niobe (Linnaeus, 1758); original combination: Papilio niobe Linnaeus, 1758

Synonyms: None

References: Reuss (1920): 192 (footnote); Reuss (1922): 197

12 species:

• adippe ([Denis & Schiffermüller], 1775)

  • ssp. adippe ([Denis & Schiffermüller], 1775); original combination: Papilio adippe

  • ssp. alaiensis Reuss, 1921; original combination: Argynnis alaiensis. Type loc.: “Alai Tal, Koksu”.

  • ssp. bischoffi (Reuss, 1922); original combination: Fabriciana cydippe f. bischoffi. Type loc.: “Turkestan”.

  • ssp. chalciope Jachontov [1909]; original combination: Argynnis adippe f. chalciope. Type loc.: “Borzhom”.

  • ssp. chayuensis (Huang, 2001); original combination: Fabriciana adippe chayuensis. Type loc.: Chayu, Chayu area Tibet.

  • ssp. chinensis (Belter, 1931); original combination: Fabriciana niobe chinensis. Type loc.: “Kunkalashan, Szechuan”.

  • ssp. chrysodippe Staudinger, 1892; original combination: Argynnis adippe var. chrysodippe. Type loc.: “Amur”. Distr.: Amur region.

  • ssp. elwesi (Reuss, 1921); original combination: Fabriciana niobe f. elwesi

  • ssp. kurosawae Matsumura, 1915; original combination: Argynnis adippe var. kurosawae. Type loc.: “Sapporo”.

  • ssp. garcila Fruhstorfer, 1910; original combination: Argynnis adippe garcila. Type loc.: “Saratow”. Distr.: Southern Europe.

  • ssp. locuples Butler, 1881; original combination: Argynnis locuples. Type loc.: “Nikko, C. Japan”.

  • ssp. ornatissima Leech, 1892; original combination: Argynnis ornatissima. Type loc.: “Ta-chien-lu”.

  • ssp. martini (Reuss, 1922); original combination: Fabriciana cydippe f. martini. Type loc.: “Turkestan”.

  • ssp. milina (Lang, 2009); original combination: Fabriciana adippe niilina. Type loc.: Tibet: Milin County, Namula Pass, 3800 m,. Types: IZCAS. Refs.: Lang and Han (2010): 493. Ety.: Milin (type locality)

  • ssp. norvegica Schultz, 1904; original combination: Argynnis adippe ab. (var?) norvegica. Type loc.: “Norwegen”. Distr.: Northern Europe, north and central Siberia, Far East.

  • ssp. satakei Nakahara, 1926, repl. name; original combination: Argynnis adippe satakei. Type loc.: “Sakhalin”.

    • syn. sachalinensis Satake, 1916, preocc.

  • ssp. taliana (Reuss, 1922); original combination: Fabriciana cydippe f. taliana. Type loc.: “Yunnan, Tali”.

  • ssp. taurica Staudinger, 1878; original combination: Argynnis adippe var. taurica. Type loc.: “Taurus”. Distr.: Armenian Highland and Kopet-Dagh.

  • ssp. thalestria Jachontov [1909]; original combination: Argynnis adippe var. thalestria. Type loc.: “Caucasus centralis; Transcaucasia: Borzhom”. Distr.: Caucasus Major and Minor.

  • ssp. tianshanica Alphéraky, 1881; original combination: Argynnis adippe var. tianschanica. Type loc.: “.sur le Jouldousse”. Distr.: Tian-Shan, Tarbagatai and Saur.

    • syn. juldana (Reuss, 1922). Type loc.: “Yuldus, Tienschan”.

  • ssp. zarewna Fruhstorfer, 1912; original combination: Argynnis adippe zarewna. Type loc.: “Irkutsk”. Distr.: Sayan, Transbaikalia.

• argyrospilata Kotsch, 1938; original combination: Argynnis argyrospilata Distr.: Afghanistan and Tajikistan (De Moya et al. 2017).

• auresiana Fruhstorfer, 1908

  • ssp. astrifera (Higgins, 1965); original combination: Fabriciana niobe astrifera

  • ssp. auresiana Fruhstorfer, 1908; original combination: Argynnis adippe auresiana

    • syn. buckwelli (Varin, 1961)

  • ssp. hassani (Weiss, 1979); original combination: Fabriciana niobe hassani

  • ssp. maroccana (Belter, 1935); original combination: Profabriciana auresiana maroccana

• elisa Godart [1824]; original combination: Argynnis elisa Distr.: Corsica, Sardinia (De Moya et al. 2017).

• hallasanensis (Okano, 1998); original combination: Fabriciana coredippe hallasanensis

• jainadeva Moore, 1864; original combination: Argynnis jainadeva Distr.: Pakistan and northern India, western Himalayas (De Moya et al. 2017).

• kamala Moore, 1857; original combination: Argynnis kamala Distr.: Western Himalayas (De Moya et al. 2017).

• nerippe Felder & Felder, 1862

  • ssp. coreana Butler, 1882; original combination: Argynnis coreana

  • ssp. mumon Matsumura, 1929; original combination: Argynnis adippe mumon

  • ssp. nerippe Felder & Felder, 1862; original combination: Argynnis nerippe

  • ssp. nerippina Fruhstorfer, 1907; original combination: Argynnis nerippe nerippina

• niobe (Linnaeus, 1758)

  • ssp. barkhatovi (Gorbunov, 2001); original combination: Fabriciana niobe barkhatovi

  • ssp. changaica (Reuss, 1922); original combination: Fabriciana niobe changaica

  • ssp. demavendis Gross & Ebert, 1975; original combination: Argynnis (Fabriciana) niobe demavendis

  • ssp. gigantea Staudinger, 1871; original combination: Argynnis gigantea

  • ssp. intermedian (Reuss, 1925); original combination: Farbriciana f. xanthodippoides subf. intermedian

  • ssp. khusestana Gross & Ebert, 1975; original combination: Argynnis (Fabriciana) niobe khusestana

  • ssp. kunlunensis (Lang, 2010); original combination: Fabriciana niobe kunlunensis. Type loc.: Xinjiang Uygur Autonomous Region: Kunlun Mts., 3300 m. Types: IZCAS. Distr.: South Xinjiang, China. Refs.: Lang and Wang (2010): 222. Ety.: Kunlun Mountains

  • ssp. kurana Wyatt & Omoto, 1966; original combination: Argynnis (Fabriciana) niobe kurana

  • ssp. mohmandorum Fruhstorfer, 1912; original combination: Argynnis adippe mohmandorum

  • ssp. niobe (Linnaeus, 1758); original combination: Papilio niobe

  • ssp. orientalis Alphéraky, 1881; original combination: Argynnis niobe var. orientalis

  • ssp. ornata Staudinger, 1901; original combination: Argynnis niobe v. ornata

  • ssp. shiva Wyatt & Omoto, 1966; original combination: Argynnis (Fabriciana) niobe shiva

  • ssp. tekkensis Christoph, 1893; original combination: Argynnis niobe var. tekkensis

  • ssp. valesinoides (Reuss, 1926); original combination: Fabriciana coredippe valesinoides

  • ssp. voraxides Reuss, 1921; original combination: Argynnis voraxides

• pallescens Butler, 1873; original combination: Argynnis pallescens. Type loc.: “Hakodadi” [Hokkaido, Japan]. Distr.: Hokkaido, Honshu and the Kuril Islands. Refs.: Spitsyna et al. (2024): 186 (to species).

  • syn. taigetana (Reuss, 1922); original combination: Fabriciana taigetana.

  • syn. microvorax (Belter, 1935); original combination: Fabriciana vorax microvorax.

  • syn. doii Matsumura, 1928; original combination: Argynnis adippe doii.

• vorax Butler, 1871; original combination: Argynnis vorax Distr.: Southeast Russia, China, Korea, Tibet, Japan (De Moya et al. 2017).

  • syn. xanthodippoides Reuss, 1922; original combination: Argynnis vorax xanthodippoides.

  • syn. paki (Okano, 1998); original combination: Fabriciana vorax paki.

• xipe Groum-Grshimailo, 1891

  • ssp. aglaiaeformis Watkins, 1924; original combination: Arggnnis cydippe aglaiaeformis. Type loc.: Korniloff Bay, 40-70 km S of Vladivostok. Refs.: Watkins (1924): 455.

    • syn. mandschurica (Belter, 1935). Type loc.: Ussuri.

  • ssp. gyala Tytler, 1940; original combination: Argynnis adippe gyala

  • ssp. niraea Oberthür, 1913; original combination: Argynnis niraea

    • syn. anargyra (Draeseke & Reuss, 1925)

  • ssp. rueckerti Fruhstorfer, 1911; original combination: Argynnis adippe rückerti. Type loc.: “Tschifu, N.O. China”.

    • syn. subornatissima (Reuss, 1922)

    • syn. transbaicalensis (Belter, 1935)

    • syn. subtransbaicalensis (Belter, 1935)

  • ssp. xipe Groum-Grshimailo, 1891; original combination: Argynnis adippe var. xipe

    • syn. xanthodippe Fixsen, 1887

    • syn. coredippe Leech, 1892 Refs.: Lee (2005) (synonymy).

    • syn. musanna Mori, Doi & Cho, 1934

    • syn. souyoensis Mori, Doi & Cho, 1934

Subgenus Speyeria

Subgenus: Speyeria Scudder, 1872

Type species: Speyeria idalia (Drury, 1773); original combination: Papilio idalia Drury, 1773

Synonyms:

• Semnopsyche Scudder, 1875. Type species: Speyeria diana (Cramer, [1777]); original combination: Papilio diana Cramer, [1777]. Refs.: Scudder (1875): 238, 258
• Neoacidalia Reuss, 1926. Type species: Speyeria cybele (Fabricius, 1775); original combination: Papilio cybele Fabricius, 1775. Refs.: Reuss (1926 b): 69
• Mesoacidalia Reuss, 1926. Type species: Speyeria aglaja (Linnaeus, 1758); original combination: Papilio aglaja Linnaeus, 1758. Refs.: Reuss (1926 b): 69
• Proacidalia Reuss, 1926. Type species: Speyeria clara (Blanchard, 1844); original combination: Argynnis clara Blanchard, 1844. Refs.: Reuss (1926 b): 69

Distribution: North America, Europe, Asia

References: Scudder (1872): 23; Dunford (2009) (taxonomy); Hammond et al. (2013) (crossing exp.); Zhang et al. (2020): 17 (as subgenus of Argynnis); Hill et al. (2018) (DNA barcode); Thompson et al. (2019) (phylogeny)

22 species:

• adiaste Edwards, 1864

  • ssp. adiaste Edwards, 1864; original combination: Argynnis Adiaste. Type loc.: California, USA. Types: lectotype in CMNH. Refs.: W. H. Edwards, 1864, Proc. Entomol. Soc. Philad. 3(3): 436, no. 7; figured by W. H. Edwards (1890), Butterflies N. Am. 3(10): pl. Argynnis 8, figs. 1 ♂ D, 2 ♂ V (lectotype), 3 ♀ D (as adiante); Brown (1965): 291, fig. 17 ♂ D&V (lectotype); Dunford (2009).

    • syn. adiante Boisduval, 1869. Type loc.: “au bord des bois, dans la partie orientale de la Californie”; defined as “Santa Cruz Mountains, California” by dos Passos and Grey (1947), Am. Mus. Novit. (1370): 16; further defined as “2 miles southeast of Summit Road along Highland Way, Santa Cruz Mountains, Santa Cruz County, California” by J. Emmel et al. (1998), Syst. W. N. Am. Butterflies: 34. Types: lectotype in CMNH. Refs.: Boisduval, 1869, Annls. Soc. Entomol. Belg. 12: 61, no. 57.

  • ssp. clemencei Comstock, 1925; original combination: Argynnis adiaste race clemencei. Type loc.: “Atascadero, Calif” [San Luis Obispo County, California]. Types: LACM. Refs.: Comstock, 1925, Bull. South. Calif. Acad. Sci. 24(3): 67; figured by J. A. Comstock (1927), Butterflies Calif.: pl. 28, figs. 7 ♂ D.

• aglaja (Linnaeus, 1758)

  • ssp. aglaja (Linnaeus, 1758); original combination: Papilio aglaia. Type loc.: Sweden. Refs.: Linnaeus (1758): 481; Hemming (1942) (taxonomy); dos Passos and Grey (1945 a).

  • ssp. auxo Yakhontov, 1909; original combination: Argynnis aglaja var. auxo. Type loc.: Caucasus. Refs.: Yakhontov (1909): 287.

  • ssp. bessa Fruhstorfer, 1907; original combination: Argynnis aglaja bessa. Type loc.: Tse-Kou. Refs.: Fruhstorfer (1907): 257.

  • ssp. boreas (Hemming, 1942), repl. name; original combination: Mesoacidalia charlotta boreas. Type loc.: see aglaja borealis Strand. Refs.: Hemming (1942): 160.

    • syn. aglaja v. borealis Strand, 1901, preocc. by Argynnis thore borealis Staudinger, 1861. Type loc.: “Arktisch, Norwegen”. Refs.: Strand (1901): 49.

  • ssp. clavimacula Matsumura, 1929; original combination: Argynnis aglaia clavimacula. Type loc.: Korea. Refs.: Matsumura (1929): 154.

  • ssp. excelsior Rothschild, 1933; original combination: Argynnis aglaia excelsior. Type loc.: Ketama, Morocco. Refs.: Rothschild (1933): 317.

  • ssp. fortuna Janson, 1877; original combination: Argynnis fortuna. Type loc.: Japan. Refs.: Janson (1877): 154.

  • ssp. gigasvitatha Verity, 1935; original combination: Argynnis aglaja gigasvitatha. Type loc.: Issyk kul. Refs.: Verity (1935): 202.

  • ssp. kenteana Stichel, 1901; original combination: Argynnis aglaja kenteana Refs.: Stichel (1901): 23.

    • syn. graeseri Kardakoff, 1928. Type loc.: Ussuri. Refs.: Kardakoff (1928): 270.

  • ssp. matsumurai Nakahara, 1926; original combination: Argynnis aglaja matsumurai Refs.: Nakahara (1926).

    • syn. chishimensis Matsumura, 1928 Refs.: Matsumura (1928): 192.

  • ssp. ottomana Röber, 1896; original combination: Argynnis aglaja var. ottomana Refs.: Röber (1896): 82.

  • ssp. lyauteyi Oberthür, 1920; original combination: Argynnis lyauteyi. Type loc.: Azrou, Morocco. Refs.: Oberthür (1920): 48, pl.D.

  • ssp. tonnai Matsumura, 1928; original combination: Argynnis aglaia tonnai. Type loc.: Saghalien. Refs.: Matsumura (1928): 192.

    • syn. sachalinensis Matsumura, 1911, preocc. Type loc.: Saghalien. Refs.: Matsumura (1911): 43.

  • ssp. vitatha Moore [1875]; original combination: Argynnis vitatha. Type loc.: “Cashmere, north side of Rajdiangan Pass and Gurais”. Refs.: Moore ([1875]): 568.

• alexandra Ménétriés, 1832; original combination: Argynnis alexandra. Type loc.: Talysh Mountains, Zuvand, Azerbaijan. Distr.: Azerbaijan, Turkmenistan, Iran (De Moya et al. 2017). Refs.: Ménétriés (1832): 246; dos Passos and Grey (1945 a).

• aphrodite (Fabricius, 1787)

  • ssp. alcestis Edwards [1877]; original combination: Argynnis Alcestis. Type loc.: “Northern Illinois, Iowa and Colorado”, interpreted as “Galena, Jo Davies County, Illinois” (dos Passos and Grey 1947). Types: lectotype in CMNH. Refs.: dos Passos and Grey (1947) (lectotype).

    • syn. suffusa Wolcott, 1916. Type loc.: “bog south of Grand Rapids, Michigan” [Kent County]. Types: “Holotype” in OSU. Refs.: Wolcott, 1916, Entomol. News 27(3): 97-98, pl. 4, fig. 5 ♂ D (“holotype”).

  • ssp. aphrodite (Fabricius, 1787); original combination: P[apilio]. N[ymphalis]. P[halerata]. Aphrodite. Type loc.: “America meridionali”; neotype from “New York City, New York County, New York”. Types: neotype in AMNH. Refs.: Fabricius, 1787, Mantissa Ins. 2: 62, no. 590; dos Passos and Grey (1947) (neotype).

    • syn. bartschi Reiff, 1910. Type loc.: in West Roxbury, Mass” [Suffolk County, Massachusetts]. Types: “Holotype” in MCZ. Refs.: Reiff, 1910, Psyche 17(6): 252-255, pl. 13, fig. 1 ♀ D&V.

    • syn. bakeri Clark, 1932. Type loc.: “Waynesburg, Ohio” [Stark County]. Types: “Holotype” in NMNH. Refs.: Clark, 1932, Bull. U. S. Nat. Mus. (157): 276, pl. 19, figs. 1 ♂ D, 2 ♂ V (“holotype”).

    • syn. hughi (Clark & Clark, 1951). Type loc.: not stated in the original description; defined as “from a single bog in Essex, Mass” by A. Clark (1927), Ann. Assoc. Am. Geogr. 17(3): 126, where the name is a nomen nudum. [Essex County, Massachusetts]. Types: “holotype” probably in NMNH. Refs.: Clark & Clark, 1951, Smithsonian Misc. Coll. 116(7): 60-61.

  • ssp. byblis (Barnes & Benjamin, 1926); original combination: Dryas aphrodite race byblis. Type loc.: “White Mts., Ariz” [Arizona]. Types: NMNH. Refs.: Barnes & Benjamin, 1926, Bull. South. Calif. Acad. Sci. 25(3): 91, no. 157d.

  • ssp. columbia Edwards, 1877; original combination: Argynnis Columbia. Type loc.: “Lahache, near the Alaskan border of British Columbia”; further defined as “Lakes Lahache and Quesnelle, British Columbia” by W. H. Edwards [1878], Field & Forest 3(6): 102-103. [Lac La Hache]. Types: lectotype in CMNH. Refs.: Hy. Edwards, 1877, Pac. Coast Lepid. (26): [1].

    • syn. columbia Edwards, 1878, preocc. Type loc.: “Lakes Lahache and Quesnelle, British Columbia” [Lac La Hache]. Types: Type(s) in CMNH. Refs.: W. H. Edwards, 1878, Field & Forest 3(6): 102-103.

  • ssp. cullasaja (Gatrelle, 1998); original combination: Speyeria aphrodite cullasaja. Type loc.: “trail to Scaly Mountain, 4200’, Macon Co., North Carolina”. Types: FLMNH.

  • ssp. ethne Hemming, 1933; original combination: Argynnis ethne. Type loc.: “Arizona to Montana.Colorado.So. Colorado.Arizona.Colorado”, interpreted as “Bighorn, Treasure County, Montana” (dos Passos and Grey 1947). Types: lectotype in CMNH. Refs.: Hemming, 1933, Entomol. 66(847): 276, no. 10; dos Passos and Grey (1947) (lectotype).

    • syn. cypris Edwards, 1886, preocc. by Argynnis cypris Meigen, [1828], Syst. Beschr. Europ. Schmett. 1(2): 59-60, no. 13. Type loc.: “Arizona to Montana.Colorado.So. Colorado.Arizona.Colorado”; defined as “Bighorn, Treasure County, Montana” by dos Passos and Grey (1947), Am. Mus. Novit. (1370): 9; suggested to be “Big Horn Mountains, near Sheridan, Wyoming” by F. Brown (1965), Trans. Am. Entomol. Soc. 91(3): 245. [Sheridan County]. Types: Lectotype in CMNH, designated by dos Passos and Grey (1947), Am. Mus. Novit. (1370): 9; figured by W. Holland (1898), Butterfly Book: pl. 12, fig. 3 ♂ D, and F. Brown (1965), Trans. Am. Entomol. Soc. 91(3): 244, fig. 3 ♂ D&V.

  • ssp. manitoba Chermock & Chermock, 1940; original combination: Argynnis manitoba. Type loc.: “Sand Ridge, Manitoba”. Types: CNC. Refs.: Chermock & Chermock, 1940, Can. Entomol. 72(4): 83.

    • syn. mayae (dos Passos & Grey, 1947) Refs.: dos Passos and Grey (1947): 8 (adopted Argynnis aphrodite cypris tr. fm. mayae Gunder, 1932, to represent a subspecies-level taxon, with the same data; Code Articles 10.2, 45.5.1).

    • syn. mayae Gunder, 1932. Type loc.: “Sand Ridge, Bener Dam Lake, near Kelwood, Man., Can” [Manitoba]. Types: “Holotype” in AMNH. Refs.: Gunder, 1932, Can. Entomol. 64(12): 279.

  • ssp. whitehousei Gunder, 1932; original combination: Argynnis whitehousei. Type loc.: “Jaffray, B. C” [British Columbia]. Types: AMNH. Refs.: Gunder, 1932, Can. Entomol. 64(12): 279-280.

  • ssp. winni Gunder, 1932; original combination: Argynnis aphrodite race winni. Type loc.: “St. Calixte de Kilkenny, Que. (about 40 miles from Montreal)” [Quebec]. Types: AMNH. Refs.: Gunder, 1932, Can. Entomol. 64(12): 278-279.

• atlantis Edwards, 1862

  • ssp. atlantis Edwards, 1862; original combination: Argynnis atlantis. Type loc.: “mountainous districts of the northern states and to parts of British America.Catskill Mountains, near the Mountain House.White Mountains.Williamstown, Mass.Lake Winnepeg.near Hudson’s Bay.also.north side of Lake Superior”; defined as “Hunter, Green County, New York” by dos Passos (1935); lectotype labeled “Catskills”. Types: lectotype in CMNH. Refs.: W. H. Edwards, 1862, Proc. Acad. Nat. Sci. Philad. 14(1/2): 54-55, no. 1; figured by W. H. Edwards (1868), Butterflies N. Am. 1: pl. Argynnis 5, figs. 1 ♂ D, 2 ♂ V, 3 ♂ V; dos Passos (1935) (lectotype).

    • syn. chemo Scudder, 1889. Type loc.: “Lake Chemo, ten miles northeast of Bangor, Me” [Penobscot County, Maine]. Types: “Lectotype” (#25828) in MCZ. Refs.: Scudder, 1889, Butterflies East. U. S. and Can. 1(4): 573-574; dos Passos (1935).

  • ssp. brittanae Pavulaan, 2022; original combination: Argynnis (Speyeria) atlantis brittanae. Type loc.: “West Virginia, Randolph County, Spruce Knob Lake”. Types: FLMNH. Refs.: Pavulaan (2022): 4-6, figs. 7 ♀ D, 9 ♀ V (holotype), 8 ♂ D, 10 ♂ V (allotype).

  • ssp. canadensis dos Passos, 1935; original combination: Argynnis atlantis canadensis. Type loc.: “Doyles Station, Newfoundland” [Newfoundland and Labrador]. Types: AMNH. Refs.: dos Passos (1935): 85-87; Dunford (2009): 90, fig. 21 ♂ D&V.

  • ssp. hollandi Chermock & Chermock, 1940; original combination: Argynnis atlantis hollandi. Type loc.: “Riding Mountains, Manitoba”. Types: CNC. Refs.: F. Chermock & R. Chermock, 1940, Can. Entomol. 72(4): 82-83; Dunford (2009) (fig. 22).

  • ssp. pahasapa (Spomer, Scott & Kondla, 1998); original combination: Speyeria atlantis pahasapa. Type loc.: “Deerfield Reservoir, Pennington Co., S. D” [South Dakota]. Types: AMNH. Refs.: Spomer, J. Scott & Kondla, 1998, Papilio (n.s.) 8: 13, figs. 1 ♂ D, 2 ♂ V (holotype); 3 ♀ D, 4 ♀ V, 5 ♂ D, 6 ♂ D, 7 ♂ V, 8 ♂ V, 9 ♂ V; Dunford (2009) (fig. 24).

  • ssp. sorocko (Scott, Kondla & Spomer, 1998); original combination: Speyeria atlantis sorocko. Type loc.: “several km NE of Mt. Judge, 9100 feet altitude, Clear Creek Co., Colo” [Colorado]. Types: AMNH. Refs.: J. Scott, Kondla & Spomer, 1998, Papilio (n.s.) 8: 13, figs. 10 ♂ V (holotype), 11 ♀ V, 12 ♂ D, 13 ♂ V, 14 ♀ D, 15 ♀ V; Dunford (2009) (fig. 23).

• bischoffii Edwards, 1870

  • ssp. artonis Edwards, 1881; original combination: Argynnis Artonis. Type loc.: Nevada, Elko Co., USA. Types: lectotype in CMNH. Refs.: Edwards, 1881, Trans. Am. Entomol. Soc. 9(1): 1-2; Brown (1965) (lectotype); Zhang et al. (2022) (genome; transfer from mormonia Boisduval).

    • syn. artonis form rubyensis Gunder, 1932, nom. nud. Type loc.: “Franklin Slough, near Wright’s Ranch, Ruby Valley, Elko Co., Nevada”. Types: AMNH. Refs.: Gunder, 1932, Can. Entomol. 64(12): 282-283.

  • ssp. bischoffii Edwards, 1870; original combination: Argynnis Bischoffii. Type loc.: Kodiak, Alaska, USA. Types: neotype in CMNH. Refs.: W. H. Edwards, 1870, Trans. Am. Entomol. Soc. 3(2): 189-190, no. 2; figured by W. H. Edwards (1875), Butterflies N. Am. 2(4): pl. Argynnis 3, figs. 1 ♂ D, 2 ♂ V, 3 ♀ D, 4 ♀ V; dos Passos and Grey (1947); Brown (1965) (neotype); Zhang et al. (2022) (genome; transfer from mormonia Boisduval).

  • ssp. erinna Edwards, 1883; original combination: Argynnis Eurynome var. Erinna. Type loc.: USA: Washington, Spokane Co. Types: lectotype in CMNH. Refs.: W. H. Edwards, 1883, Can. Entomol. 15(2): 33-34; dos Passos and Grey (1947) (lectotype); Zhang et al. (2022) (genome; transfer from mormonia Boisduval).

    • syn. aberrant form cunninghami Owen, 1893, nom. nud. Type loc.: “Klamath County, Oregon”. Types: NMNH. Refs.: Owen, 1893, Entomol. News 4(7): 246.

  • ssp. eurynome Edwards, 1872; original combination: Argynnis Eurynome. Type loc.: USA: Colorado, Park Co. Types: lectotype in CMNH. Refs.: Edwards, 1872, Trans. Am. Entomol. Soc. 4(1/2): 61, 66-67, no. 6; figured by W. H. Edwards (1875), Butterflies N. Am. 2(4): pl. Argynnis 1, figs. 1 ♂ D, 2 ♂ V, 3 ♀ D, 4 ♀ V; Zhang et al. (2022) (genome; transfer from mormonia Boisduval).

    • syn. clio Edwards, 1874. Type loc.: Teton County, Wyoming, USA. Types: neotype in AMNH. Refs.: Edwards, 1874, Trans. Am. Entomol. Soc. 5(2): 106-107; dos Passos and Grey (1947) (neotype); Brown (1965) (fig. 33).

    • syn. eurynome ab. eris Igel, 1922, nom. nud. Type loc.: “der Umgebung von Calgary (Alberta).aus Montana”. Types: Igel collection. Refs.: Igel, 1922, Entomol. Z. 36(5): 20.

    • syn. mormonia tr. f. benjamini Gunder, 1927, nom. nud. Type loc.: “Arangie, Idaho” [Arangee precinct, or Arange Peak, Fremont County]. Types: NMNH. Refs.: Gunder, 1927, Can. Entomol. 59(12): 287, no. 22, pl. B, figs. 22 ♂ D, 22a ♂ V (holotype).

    • syn. eurynome tr. f. brucei Gunder, 1927, nom. nud. Type loc.: Colorado, USA. Types: FMNH. Refs.: Gunder, 1927, Can. Entomol. 59(12): 287, no. 23, pl. B, figs. 23 ♀ D, 23a ♀ V (“holotype”).

    • syn. eurynome race clio tr. f. gunderi Field, 1936, nom. nud. Type loc.: Broadwater Co., Montana. Types: NMNH. Refs.: Field, 1936, J. Entomol. & Zool. 28(2): 23.

    • syn. eurynome ab. igeli (dos Passos & Grey, 1947), repl. name of eurynome ab. eris Igel, 1922 Refs.: dos Passos and Grey (1947): 23-24.

    • syn. eurynome ab. fieldi (dos Passos & Grey, 1947), repl. name of eurynome race clio tr. f. gunderi Field, 1936 Refs.: dos Passos and Grey (1947): 23-24.

  • ssp. kimimela Marrone, Spomer & Scott, 2008; original combination: Argynnis (Speyeria) mormonia kimimela. Type loc.: USA: South Dakota, Lawrence Co. Types: SDSU. Refs.: Scott et al. (2008): 31; Zhang et al. (2022) (genome; transfer from mormonia Boisduval).

  • ssp. luski Barnes & McDunnough, 1913; original combination: A[rgynnis]. eurynome luski. Type loc.: “White Mts., Arizona”. Types: lectotype in NMNH. Refs.: Barnes & McDunnough, 1913, Contr. Nat. Hist. Lepid. N. Am. 2(3): 95-96, pl. 1, figs. 1 ♂ D, 2 ♀ D, 3 ♂ V, 4 ♀ V; dos Passos and Grey (1947) (lectotype); Zhang et al. (2022) (genome; transfer from mormonia Boisduval).

  • ssp. opis Edwards, 1874; original combination: Argynnis Opis. Type loc.: Canada: “Bald Mountain, Cariboo, British Columbia”. Types: lectotype in CMNH. Refs.: W. H. Edwards, 1874, Trans. Am. Entomol. Soc. 5(2): 105-106; figured by W. H. Edwards (1875), Butterflies N. Am. 2(4): pl. Argynnis 3, figs. 5 ♂ D, 6 ♂ V, 7 ♀ D, 8 ♀ V; dos Passos and Grey (1947) (lectotype); Zhang et al. (2022) (genome; transfer from mormonia Boisduval).

    • syn. jesmondensis (dos Passos & Grey, 1947). Type loc.: see jesmondensis McDunnough, 1940. Refs.: dos Passos and Grey (1947): 22; Grey (1989) (synonymy).

    • syn. opis form jesmondensis McDunnough, 1940, nom. nud. Type loc.: “Jesmond, B. C.(3500 ft.)” [British Columbia]. Types: CNC. Refs.: McDunnough, 1940, Can. Entomol. 72(2): 25.

  • ssp. washingtonia Barnes & McDunnough, 1913; original combination: A[rgynnis]. bischoffi [sic] washingtonia. Type loc.: USA: Washington, Pierce Co. Types: lectotype in NMNH. Refs.: Barnes & McDunnough, 1913, Contr. Nat. Hist. Lepid. N. Am. 2(3): 95, pl. 1, figs. 5 ♂ D (lectotype), 6 ♀ D, 7 ♂ V, 8 ♀ V; dos Passos and Grey (1947) (lectotype); Zhang et al. (2022) (genome; transfer from mormonia Boisduval).

    • syn. washingtonia form ranierensis Gunder, 1932, nom. nud. Type loc.: “Paradise Valley, Mount Ranier [sic], Pierce Co., Washington” [Mt. Rainier]. Types: AMNH. Refs.: Gunder, 1932, Can. Entomol. 64(12): 282.

• callippe Boisduval, 1852

  • ssp. adiasteoides (Emmel, Emmel & Mattoon, 1998); original combination: Speyeria callippe adiasteoides. Type loc.: “California: Mendocino County, Cahto Peak summit, 4000-4200 feet elevation”. Types: LACM.

  • ssp. calgariana McDunnough, 1924; original combination: [Argynnis nevadensis] race calgariana. Type loc.: “Calgary.Alta”; suggested to be “the head of Pine Creek, Alberta (Township 22, Range 2, West of the 5th meridian)” by Kondla (1996), Alberta Naturalist 26(2): 39. Types: CNC. Refs.: McDunnough, 1924, Can. Entomol. 56(2): 42.

  • ssp. callippe Boisduval, 1852; original combination: Argynnis callippe. Type loc.: “au bord des forets”, interpreted as “San Francisco, San Francisco County, California” (dos Passos and Grey 1947). Types: Lectotype in NMNH; figured by J. Emmel et al. (1998), Syst. W. N. Am. Butterflies (2): 66, figs. 90 ♂ D, 91 ♂ V, and Dunford (2009), Insecta Mundi 90: fig. 16 ♂ D&V. Refs.: Boisduval, 1852, Annls. Soc. Entomol. Fr. (2)10(2): 302, no. 43; figured by Oberthür (1914), Étud. Lépid. Comp. 9(2): pl. 258, fig. 2166 ♂ D&V, 2167 ♀ D&V; dos Passos and Grey (1947) (lectotype).

  • ssp. chilcotinensis (Guppy & Shepard, 2001); original combination: Speyeria callippe chilcotinensis. Type loc.: “BC, Riske Creek” [British Columbia]. Types: RBCM. Refs.: C. Guppy & J. Shepard, 2001, Butterflies B. C.: 276-277, figs. ♂ D&V, ♀ D&V (holotype).

  • ssp. comstocki Gunder, 1925; original combination: Argynnis calippe [sic] var. comstocki. Type loc.: “Los Angeles, Los Angeles County, California”. Types: AMNH. Refs.: Gunder, 1925, Entomol. News 36(1): 8-9, no. 13, pl. 1, figs. 6 ♂ D, 7 ♀ D.

    • syn. comstocki tr.f. creelmani Gunder, 1934. Type loc.: “East San Diego, California” [San Diego County]. Types: AMNH. Refs.: Gunder, 1934, Can. Entomol. 66(6): 127.

  • ssp. elaine (dos Passos & Grey, 1945); original combination: Speyeria callippe elaine. Type loc.: “Butte Falls, Oregon” [Jackson County]. Types: AMNH. Refs.: dos Passos and Grey (1945 b).

  • ssp. gallatini McDunnough, 1929; original combination: Argynnis nevadensis var. gallatini. Type loc.: “Elkhorn Ranch, Upper Gallatin Canyon, Mont” [Gallatin County, Montana]. Types: CNC. Refs.: McDunnough, 1929, Can. Entomol. 61(5): 106-107.

  • ssp. hagemani (Emmel, Emmel & Mattoon, 1998); original combination: Speyeria callippe hagemani. Type loc.: “California: Shasta County; Shingletown, 3500’ elevation”. Types: LACM. Refs.: J. Emmel, T. Emmel & Mattoon, 1998, Syst. W. N. Am. Butterflies (12): 150, figs. 39 ♂ D, 40 ♂ V (holotype); 41 ♀ D, 42 ♀ V.

  • ssp. harmonia (dos Passos & Grey, 1945); original combination: Speyeria callippe harmonia. Type loc.: “Mount Wheeler (Snake Range near Utah border), Nevada” [White Pine County]. Types: AMNH. Refs.: dos Passos and Grey (1945 b).

  • ssp. inornata Edwards, 1972; original combination: Argynnis Inornata. Type loc.: California, USA. Types: neotype in CMNH. Refs.: Edwards, 1872, Trans. Am. Entomol. Soc. 4(1/2): 61, 64-65, no. 5; figured by W. H. Edwards (1876), Butterflies N. Am. 2(4): pl. Argynnis 5, figs. 1 ♂ D, 2 ♂ V, 3 ♀ D, 4 ♀ V; dos Passos and Grey (1947) (neotype); Brown (1965): 274, fig. 12 ♂ D&V; Grey (1989).

  • ssp. juba Boisduval, 1869; original combination: Argynnis Juba. Type loc.: “les prairies élevées de la Juba”, interpreted as “Downieville, Sierra County, California” (dos Passos and Grey 1947). Types: lectotype in NMNH. Refs.: Boisduval, 1869, Annls. Soc. Entomol. Belg. 12: 60, no. 54; figured by Oberthür (1914), Étud. Lépid. Comp. 9(2): pl. 262, fig. 2197 ♂ D&V, 2198 ♀ D&V; dos Passos and Grey (1947) (lectotype); Emmel et al. (1998), Syst. W. N. Am. Butterflies (2): 76, figs. 255 ♂ D, 256 ♂ V.

    • syn. laura Edwards, 1879. Type loc.: “Nevada” implied from the title of paper; suggested to be “the region mountain-wards from Virginia City” by Brown (1965); defined as “vicinity of Downieville, canyon of the North Fork of the Yuba River, Sierra County, California” by Austin (1998), Syst. W. N. Am. Butterflies (50): 634. Types: lectotype in CMNH. Refs.: W. H. Edwards, 1879, Can. Entomol. 11(3): 49-50; dos Passos and Grey (1947) (lectotype).

    • syn. sierra (dos Passos & Grey, 1945). Type loc.: “Gold Lake, Sierra County, California”. Types: AMNH. Refs.: dos Passos and Grey (1945 b): 5-6, figs. 15 ♂ D, 16 ♂ V (holotype).

  • ssp. laurina Wright, 1905; original combination: [Argynnis] Laurina; Argynnis Laurina n. v.. Type loc.: “Southern California.flies with laura”, interpreted as “Greenhorn Mountains, Kern County, California” (dos Passos and Grey 1947). Types: Lectotype (#4290) in CAS, designated by dos Passos and Grey (1947): 16. Refs.: Wright, 1905, Butterflies W. Coast: 53, no. 134; 138-139, no. 134, pl. 15, fig. 134 ♂ V (lectotype); dos Passos and Grey (1947) (lectotype).

    • syn. wrighti Wright, 1905, nom. nud. Refs.: Wright, 1905, Butterflies W. Coast: 139, no. 134.

  • ssp. liliana Edwards, 1877; original combination: Argynnis Liliana. Type loc.: “near St. Helena, Napa Co”, interpreted as “[Bale Station], near [Mount] St. Helena, Napa Co[unty]., California” (dos Passos and Grey 1947). Types: AMNH.

    • syn. baroni Edwards, 1881. Type loc.: Northern California”, interpreted as “Bale Station, near Mount St. Helena, Napa County, California” (dos Passos and Grey 1947) but Brown (1965) suggests “Mendocino, Lake or Sonoma counties”. Types: NMNH.

  • ssp. macaria Edwards, 1877; original combination: Argynnis Macaria. Type loc.: “Havilah, California” [Kern County]. Types: Lectotype in CMNH; figured by W. Holland (1931), Butterfly Book (rev. ed.): 96, pl. 13, fig. 9 ♂ D and Brown (1965): 283, fig. 15 ♂ D&V. Refs.: Edwards, 1877, Field & Forest 3(5): 86-87; dos Passos and Grey (1947) (lectotype).

  • ssp. meadii Edwards, 1872; original combination: Argynnis Meadii. Type loc.: “Turkey Creek Junction, in Colorado”; defined as “Turkey Creek Junction, Colorado.Turkey Creek just south of Morrison, near Denver” probably “the junction of South Turkey Creek with Turkey Creek, a few miles up in the foothills” by F. Brown (1934), J. N. Y. Entomol. Soc. 42(2): 159; also see F. Brown (1956), Lepid. News 9(6): 185-190. Summarized as “within 9.7 km (6 mi) northeast of Bradford Junction (“Turkey Creek Junction,” within the present-day town of Conifer), along South Turkey Creek, Jefferson County, Colorado” by Calhoun (2015), J. Lepid. Soc. 69(1): 26. Types: lectotype in CMNH. Refs.: W. H. Edwards, 1872, Trans. Am. Entomol. Soc. 4(1/2): 61, 67-68, no. 7; figured by W. H. Edwards (1875), Butterflies N. Am. 2(3): pl. Argynnis 2, figs. 1 ♂ D, 2 ♂ V, 3 ♀ D, 4 ♀ V; dos Passos and Grey (1947) (lectotype).

    • syn. gerhardi Gunder, 1927. Type loc.: “Colorado”; defined as “Turkey Creek Junction, Colorado” by dos Passos and Grey (1947). Types: “Holotype” in FMNH. Refs.: Gunder, 1927, Can. Entomol. 59(12): 287, no. 24, pl. B, figs. 24 ♂ D, 24a ♂ V (“holotype”).

  • ssp. nevadensis Edwards, 1870; original combination: Argynnis Nevadensis. Type loc.: “valleys of the Sierra, near Virginia City”; defined as “immediate neighborhood of Virginia.especially abundant near Washoe Lake” by W. H. Edwards (1871), Butterflies N. Am. 1: [94]; neotype from “Lake Washoe, Nevada” [Washoe County]. Types: neotype in CMNH. Refs.: W. H. Edwards, 1870, Trans. Am. Entomol. Soc. 3(1): 10, 14-15, no. 6; figured by W. H. Edwards (1890), Butterflies N. Am. 3(9): pl. Argynnis 5, figs. 1 ♂ D (neotype), 2 ♂ V; dos Passos and Grey (1965) (neotype); Brown (1965).

  • ssp. rupestris Behr, 1863; original combination: Argynnis rupestris. Type loc.: “California” implied from the title of the paper, “the steep rocky declivities characterizing the lower part of the Sierra at moderate elevation above the sea” from the description; defined as “Trinity Center, Trinity County, California” by dos Passos and Grey (1947), Am. Mus. Novit. (1370): 15; neotype from “California: Mariposa County; Moore Creek Road 1.5 to 2 road miles south of Highway 120, elevation 3200’; S.35, T.1S., R.17E”. Types: Neotype in CAS, designated by J. Emmel et al. (1998), Syst. W. N. Am. Butterflies (12): 148-149, figs. 33 ♂ D, 34 ♂ V; figured by J. Emmel et al. (1998), Syst. W. N. Am. Butterflies (6): 109, figs. 10 ♂ D, 11 ♂ V. dos Passos and Grey (1947), Am. Mus. Novit. (1370): 15, designated a “lectotype” (in MGCL), but this is invalid (Code Article 74.2). Refs.: Behr, 1863, Proc. Calif. Acad. Sci. [1]3(6): 84, no. 3.

  • ssp. semivirida McDunnough, 1924; original combination: [Argynnis callippe] race semivirida. Type loc.: “Aspen Grove, B. C” [British Columbia]. Types: CNC. Refs.: McDunnough, 1924, Can. Entomol. 56(2): 42-43.

  • ssp. shasta (Emmel, Emmel & Mattoon, 1998); original combination: Speyeria callippe shasta. Type loc.: “California: Shasta County; 8 road miles southeast of McCloud Arm bridge, elevation 1500’, Lake Shasta”. Types: LACM. Refs.: J. Emmel, T. Emmel & Mattoon, 1998, Syst. W. N. Am. Butterflies (12): 149, figs. 35 ♂ D, 36 ♂ V (holotype); 37 ♀ D, 38 ♀ V.

• clara Blanchard, 1844

  • ssp. clara Blanchard, 1844; original combination: Argynnis clara. Type loc.: Tihri Garhwal, Western Himalayas. Distr.: Northern India, Tibet, West China (De Moya et al. 2017). Refs.: de Nicéville (1889): 165, pl.A fig.6; dos Passos and Grey (1945 a).

  • ssp. clarina Staudinger, 1901; original combination: Argynnis clara v. clarina. Type loc.: “Amdo”. Refs.: Staudinger and Rebel (1901): 38.

  • ssp. manis Fruhstorfer, 1903; original combination: Argynnis clara manis. Type loc.: Sikkim. Refs.: Fruhstorfer (1903): 41.

    • syn. claudia Fawcett, 1904. Type loc.: Khamba Jong, Thibet. Refs.: Fawcett (1904): 136.

  • ssp. menba (Lang, 2009); original combination: Mesoacidalia clara menba. Type loc.: Tibet, Milin Co., Namula Pass, 3800m. Types: IZCAS. Refs.: Lang and Han (2010): 494. Ety.: Menba (a tribe in Motuo County, Southeast Tibet)

  • ssp. tongtianensis (Lang, 2009); original combination: Mesocacidalia clara tongtianensis. Type loc.: Qinghai, Qumalai Co., Chuhuier River, 4500m. Types: IZCAS. Refs.: Lang and Han (2010): 495. Ety.: basin of Tongtian River (type locality)

  • ssp. kanga (Lang, 2010); original combination: Mesocacidalia clara kanga. Type loc.: Sichuan, Kangding, Muge Cuo Lake. Types: IZCAS. Distr.: West Sichuan and East Tibet. Refs.: Lang and Wang (2010): 221.

• coronis Behr, 1864

  • ssp. carolae dos Passos & Grey, 1942; original combination: Argynnis coronis carolae. Type loc.: “Charleston Park, Clark County, Nevada”. Types: AMNH. Distr.: Spring Mountains, Nevada, U.S.A. (De Moya et al. 2017). Refs.: dos Passos and Grey (1942); Emmel and Austin (1998) (as species); Zhang et al. (2020) (genome, as subspecies).

  • ssp. coronis Behr, 1864; original combination: Argynnis Coronis. Type loc.: California, USA. Types: neotype in CMNH. Refs.: Behr, 1864, In: W. H. Edwards, Proc. Entomol. Soc. Philad. 3(3): 435, no. 2; figured by W. H. Edwards (1868), Butterflies N. Am. 1: pl. Argynnis 5, figs. 3 ♀ D, 4 ♀ V (as nevadensis), and (1887), Butterflies N. Am. 3(2): pl. Argynnis 14, figs. 1 ♂ D, 2 ♂ V, 3 ♀ D, 4 ♀ V (as nevadensis).

    • syn. californica Skinner, 1917. Type loc.: California, USA. Types: Holotype (#7034) in CMNH; a paralectotype figured by W. Holland (1931), Butterfly Book (rev. ed.): pl. 56, fig. 5 ♂ V. Refs.: Skinner, 1917, Entomol. News 28(7): 328; cited W. H. Edwards (1887), Butterflies N. Am. 3(2): pl. Argynnis 4, figs. 1 ♂ D, 2 ♂ V, 3 ♀ D, 4 ♀ V (as coronis).

  • ssp. halcyone Edwards [1869]; original combination: Argynnis Halcyone. Type loc.: “Colorado”; defined as “Southern border of South Park, Park Co.along the road from Hartzel east to Howbert. The altitude varies a few feet above 7600 feet” by F. Brown (1934), J. N. Y. Entomol. Soc. 42(2): 159, suggested to be “Estes Park, Larimer County, Colorado” by dos Passos and Grey (1947), Am. Mus. Novit. (1370): 12; neotype labeled “Colo”. Types: neotype in CMNH. Refs.: Edwards, [1869], Butterflies N. Am. 1: [81], pl. Argynnis 9, figs. 5 ♂ D, 6 ♂ V (1869), 3(9): pl. Argynnis 5, figs. 3 ♀ D, 4 ♀ V (1890); dos Passos and Grey (1947) (neotype).

  • ssp. hennei Gunder, 1934; original combination: Argynnis hennei. Type loc.: “near summit, elevation 7700 ft., Mt. Pinos, Frazier Mountain Park, Santa Barbara Nat. Forest, Ventura Co., Calif” [California]. Types: AMNH. Refs.: Gunder, 1934, Can. Entomol. 66(6): 126-127.

  • ssp. semiramis Edwards, 1886; original combination: Argynnis Semiramis. Type loc.: “San Bernardino, California” [San Bernardino County]. Types: lectotype in CMNH. Refs.: Edwards, 1886, Can. Entomol. 18(4): 61-62, no. 1; dos Passos and Grey (1947) (lectotype).

    • syn. semiramis tr.f. bernardensis Gunder, 1933. Type loc.: “Camp Wasew[a]gan for Boy Scouts, San Bernardino Mts., San Bernardino Co., Calif” [California]. Types: AMNH. Refs.: Gunder, 1933, Can. Entomol. 65(8): 172-173.

  • ssp. simaetha (dos Passos & Grey, 1945); original combination: Speyeria coronis simaetha. Type loc.: “Black Canyon, Cascade Mountains, near Brewster, Washington” [Okanogan County]. Types: AMNH. Refs.: dos Passos and Grey (1945 b).

  • ssp. snyderi Skinner, 1897; original combination: Argynnis Snyderi. Type loc.: “Salt Lake City, Utah.Ogden”, interpreted as “City Creek Canyon, Salt Lake City, Salt Lake County, Utah” (dos Passos and Grey 1947). Types: Lectotype (#7033) in CMNH, designated by dos Passos and Grey (1947), ibid.; paralectotype figured by W. Holland (1898), Butterfly Book: pl. 18, fig. 6 ♂ D. Refs.: dos Passos and Grey (1947) (lectotype).

• cybele (Fabricius, 1775)

  • ssp. carpenterii Edwards, 1876; original combination: Argynnis Carpenterii. Type loc.: “New Mexico, above timber line”; defined as “Taos Peak, about 12,000 to 13,000 feet elevation” by W. H. Edwards (1889), Butterflies N. Am. 3(8): [138]; lectotype from “Taos Mtn N. M” [Taos Peak, Taos County, New Mexico]. Types: lectotype in CMNH. Refs.: dos Passos and Grey (1947) (lectotype).

  • ssp. charlottii Barnes, 1897; original combination: Argynnis Charlottii. Type loc.: “Glenwood Springs, Colo” [Garfield County, Colorado]. Types: NMNH.

  • ssp. cybele (Fabricius, 1775); original combination: P[apilio]. N[ymphalis]. P[halerata]. Cybele. Type loc.: “America”, interpreted as “New York City, New York County, New York” (dos Passos and Grey 1947). Types: NHMUK. Refs.: Fabricius (1775): 516.

    • syn. daphnis (Cramer [1775]). Type loc.: “ze te Nieuw-York en te Jamaika” “Nouvelle York & la Jamaïque”; neotype from “New York City, New York County, New York”. Types: neotype in NHMUK. Refs.: dos Passos and Grey (1947) (neotype; syntype of Papilio cybele).

    • syn. baal Strecker, 1878. Type loc.: “Ohio”; defined as “Toledo, Lucas County, Ohio” by dos Passos and Grey (1947). Types: “holotype” in FMNH.

  • ssp. krautwurmi Holland, 1931; original combination: Argynnis cybele var. krautwurmi. Type loc.: “Les Cheneaux, Upper Peninsula of Michigan” [Mackinac County]. Types: CMNH.

  • ssp. neomexicana Grishin, 2022; original combination: Argynnis cybele neomexicana Refs.: Zhang et al. (2022).

  • ssp. novascotiae McDunnough, 1935; original combination: Argynnis cybele novascotiae. Type loc.: “White Point Beach, Queens Co., N. S” [Nova Scotia, Canada]. Types: CNC.

  • ssp. pseudocarpenteri Chermock & Chermock, 1940; original combination: Argynnis cybele pseudocarpenteri. Type loc.: Sand Ridge, Manitoba. Types: CNC.

• diana (Cramer [1777])

  • ssp. diana (Cramer [1777]); original combination: Pap[ilio]. Nymph[alis]. Phaler[ata]. Diana. Type loc.: “Virginien”, “Virginie” on p.3, interpreted as “Jamestown, James City County, Virginia” (dos Passos and Grey 1947). Types: NHMUK. Distr.: East coast U.S.A. extending west to Tennessee (De Moya et al. 2017). Refs.: Cramer ([1777]): 4, pl.XCVIII, fig.D,E.

  • ssp. arkansana Pavulaan, 2021; original combination: Argynnis (Speyeria) diana. Type loc.: “County Road 30, 0.64 miles west of U.S. Route 71, Needmore, Scott County, Arkansas, USA”. Types: FLMNH. Refs.: Pavulaan (2021): 4. Ety.: Arkansas (range)

• edwardsii Reakirt, 1866; original combination: Argynnis Edwardsii. Type loc.: “California; Pikes Peak, Colorado Territory.Washington, D. C.?.Rocky Mountains.near Empire City, Colorado Territory, in the heart of the mountains only”; defined as “Pike’s Peak, Teller County, Colorado” by dos Passos and Grey (1947), Am. Mus. Novit. (1370): 10; subsequently defined as “Empire, Clear Creek County, Colorado” by Brown (1965). Types: lectotype in FLMNH. Distr.: Primarily in north central U.S.A. (De Moya et al. 2017). Refs.: Reakirt, 1866, Proc. Entomol. Soc. Philad. 6(1): 137-139; dos Passos and Grey (1947) (lectotype).

  • syn. - (Reuss, 1927); original combination: A[cidalia]. edwardsi montana. Type loc.: Montana, USA. Types: ZMHU. Refs.: Reuss, 1927, Dtsche. Entomol. Z. 1926(5): 439; cited Reuss (1922), Arch. Naturgesch. 87A(11): 205.

  • syn. - Gunder, 1934; original combination: Argynnis edwardsii tr. f. edonis. Type loc.: “Beaver City, Larimer County, Colorado (10-12,000 ft.)”. Types: AMNH. Refs.: Gunder, 1934, Can. Entomol. 66(6): 125.

• egleis Behr, 1862

  • ssp. albrighti Gunder, 1932; original combination: Argynnis albrighti. Type loc.: “Highwood Mts., Chouteau County, Montana”. Types: AMNH. Refs.: Gunder, 1932, Can. Entomol. 64(12): 281-282.

  • ssp. egleis Behr, 1862; original combination: A[rgynnis]. Egleis Boisd.?. Type loc.: “California” implied from the title of the paper, “high up in the mountains” from the description; defined as “vicinity of Gold Lake, Sierra County, California” (dos Passos and Grey 1947). Types: neotype in NMNH. Refs.: Behr, 1862, Proc. Calif. Acad. Sci. [1]2(12): 174, no. 4.

    • syn. montivaga Behr, 1863. Type loc.: “California” implied from the title of the paper, “widely spread through the Sierra.at different localities.from an elevation of 10,500 feet” from the description; defined as “vicinity of Gold Lake, Sierra County, California” (dos Passos and Grey 1947). Types: neotype in NMNH. Refs.: Behr, 1863, Proc. Calif. Acad. Sci. [1]3(6): 84, no. 2.

    • syn. egleis Boisduval, 1869, preocc. by Argynnis egleis Behr, 1862. Type loc.: “les lieux incultes et aux borde des forets, dans l’intérieur de la Californie”. Types: lectotype in NMNH. Refs.: Boisduval, 1869, Annls. Soc. Entomol. Belg. 12: 59-60, no. 53; figured by Oberthür (1914), Étud. Lépid. Comp. 9(2): pl. 261, fig. 2194 ♂ D&V, pl. 262, fig. 2195 ♀ D&V; dos Passos and Grey (1947) (lectotype).

    • syn. astarte Edwards, 1864, preocc. by Argynnis astarte E. Doubleday 1848. Type loc.: “Mountainous regions of California”; defined as “Gold Lake, Sierra County, California” by dos Passos and Grey (1947). Types: neotype in FMNH. Refs.: W. H. Edwards, 1864, Proc. Entomol. Soc. Philad. 3(3): 434, 435, no. 4.

    • syn. mammothi Gunder, 1924. Type loc.: “Mammoth Camp, Mono County, California”. Types: “Holotype” possibly in LACM. Refs.: Gunder, 1924, Entomol. News 35(5): 157, pl. 2, fig. H ♂ D (“holotype”).

    • syn. boharti Gunder, 1930. Type loc.: “Mono Pass Trail (Yosemite to Mono Lake), Calif” [California]. Types: “Holotype” in AMNH. Refs.: Gunder, 1930, Bull. Brooklyn Entomol. Soc. 24(5): 326-327, pl. 31, fig. 5 ♀ D&V (“holotype”).

  • ssp. kutoyisiks (Kohler, 2020); original combination: Speyeria egleis kutoyisiks. Type loc.: “Montana: Toole Co.: Pratt Canyon, 4470-5000’, West Butte Sweet Grass Hills”. Types: FLMNH. Refs.: Kohler (2020): 6, figs. 93 ♂ D, 94 ♂ V (holotype), 109 ♀ D, 110 ♀ V (allotype), 95-108, 121, 122, 125-126 ♂ D&V, 111-120, 123, 124 ♀ D&V.

  • ssp. linda dos Passos & Grey, 1942; original combination: Argynnis utahensis linda. Type loc.: “Heyburn Peak, Sawtooth-Boise, Idaho, 9500-10,000 feet” [Custer County]. Types: AMNH. Refs.: dos Passos and Grey (1942).

  • ssp. macdunnoughi Gunder, 1932; original combination: Argynnis macdunnoughi. Type loc.: “Elkhorn Ranch Resort, Gallatin County, Mont.up what they call (at the Resort) Spring Creek for about half a mile, the path branches to the right about where the water ceases to flow and rises sharply over a ridge, ending in an open meadow”; “7000 feet, Upper Gallatin Canyon” added by dos Passos and Grey (1947). Types: CNC. Refs.: Gunder, 1932, Can. Entomol. 64(12): 280-281.

  • ssp. mattooni (Emmel & Emmel, 1998); original combination: Speyeria egleis mattooni. Type loc.: “California: Siskiyou County; Dry Lake Mountain, U. S. Forest Service Road 40S01, Siskiyou Mountains”. Types: LACM. Refs.: J. Emmel & T. Emmel, 1998, Syst. W. N. Am. Butterflies (33): 438, figs. 1 ♂ D, 2 ♂ V (holotype); 3 ♀ D, 4 ♀ V.

  • ssp. moecki (Hammond & Dornfeld, 1983); original combination: Speyeria egleis moecki. Type loc.: “Skookum Mdw., Walker Rim, Klamath Co., Ore” [Oregon]. Types: AMNH. Refs.: Hammond & Dornfeld, 1983, J. Lepid. Soc. 37(2): 115-117, fig. 1 ♂ D&V (holotype) ♀ D&V.

  • ssp. morrellensis (Kohler, 2020); original combination: Speyeria egleis morrellensis. Type loc.: “Montana: Powell Co.: vic. Morrell Lookout, 7600’, 10 mi. E. Seeley Lake”. Types: FLMNH. Refs.: Kohler (2020): 5, figs. 45 ♂ D, 46 ♂ V (holotype), 65 ♀ D, 66 ♀ V (allotype), 47-64, 85-90 ♂ D&V, 67-84, 91-92 ♀ D&V.

  • ssp. oweni Edwards, 1892; original combination: Argynnis Oweni. Type loc.: “Mt. Shasta, elevation 7,500 feet” [Siskiyou County, California]. Types: lectotype in CMNH. Refs.: W. H. Edwards, 1892, Can. Entomol. 24(5): 105-106, no. [1] (not 2 as given); dos Passos and Grey (1947) (lectotype).

  • ssp. reidi (Austin, 1998); original combination: Speyeria egleis reidi. Type loc.: “Nevada: White Pine County; Snake Range, Wheeler Peak Campground, 3033 m, T13N R68E (unsectioned) on USGS Sacramento Pass, Nev. 15’ quadrangle”. Types: FLMNH. Refs.: Austin, 1998, Syst. W. N. Am. Butterflies (46): 574-575, figs. 5 ♂ D, 6 ♂ V (holotype); 7 ♀ D, 8 ♀ V.

  • ssp. secreta (dos Passos & Grey, 1945); original combination: Speyeria montivaga secreta. Type loc.: “Rocky Mountain National Park, Colorado, 8000 feet.in the Estes Park area” [Larimer County]; more likely to be from Rabbit Ears Pass, Routt-Grand counties, Colorado (A. Warren pers. comm. 2006). Types: AMNH. Refs.: dos Passos and Grey (1945 b): 7-8, figs. 19 ♂ D, 20 ♂ V (holotype).

  • ssp. tehachapina Comstock, 1920; original combination: Argynnis montivaga var. tehachapina. Type loc.: “highest peak in Tehachapi Mts., California” [Kern County]. Types: LACM. Refs.: J. A. Comstock, 1920, Southwest Sci. Bull: 6-9, pl. [1], figs. 7 ♂ V, 8 ♂ D (holotype); 9 ♀ D.

  • ssp. toiyabe (Howe, 1975); original combination: Speyeria egleis toiyabe. Type loc.: “Kingston Canyon, Toiyabe Range, Lander County, Nevada”. Types: LACM. Refs.: Howe, 1975, Butterflies N. Am.: 237, pl. 31, fig. 14 ♂ D&V (holotype).

  • ssp. utahensis Skinner, 1919; original combination: Argynnis utahensis. Type loc.: “City Creek Canyon, Salt Lake City, Utah” [Salt Lake County]. Types: CMNH. Refs.: Skinner, 1919, Entomol. News 30(8): 216.

  • ssp. yolaboli (Emmel & Emmel, 1998); original combination: Speyeria egleis yolaboli. Type loc.: “California: Tehama County; South Yolla Bolly Mountain”. Types: LACM. Refs.: J. Emmel & T. Emmel, 1998, Syst. W. N. Am. Butterflies (33): 439, figs. 5 ♂ D, 6 ♂ V (holotype); 7 ♀ D, 8 ♀ V.

• hesperis Edwards, 1864

  • ssp. beani (Barnes & Benjamin, 1926); original combination: Dryas atlantis race beani. Type loc.: “Banff, Alta” [Alberta]. Types: NMNH. Refs.: Barnes & Benjamin, 1926, Bull. South. Calif. Acad. Sci. 25(3): 92, no. 160b; Dunford (2009) (fig. 28).

  • ssp. brico (Kondla, Scott & Spomer, 1998); original combination: Speyeria hesperis brico. Type loc.: “Castle Creek Forest Service Rd. Km 23.5, Cariboo Mts. (25 air km due S of McBride), B.C” [British Columbia]. Types: CNC. Refs.: Kondla, J. Scott & Spomer, 1998, Papilio (n.s.) 8: 14, figs. 24 ♂ V, 25 ♀ V; Dunford (2009) (fig. 29).

  • ssp. capitanensis (Holland, 1988); original combination: Speyeria atlantis capitanensis. Type loc.: “Padilla Point, crest of Capitan Ridge, Capitan Mts., Lincoln Co., NM” [New Mexico]. Types: AMNH. Refs.: R. Holland, 1988, Bull. Allyn. Mus. (115): 2-3, fig. 4 ♂&♀ D&V (top row holotype); Dunford (2009) (fig. 46).

  • ssp. dennisi (dos Passos & Grey, 1945); original combination: Speyeria atlantis dennisi. Type loc.: see dennisi Gunder, 1927. Refs.: dos Passos and Grey (1945 b): 20.

    • syn. lais tr. f. dennisi Gunder, 1927. Type loc.: “Beulah, Man., Can” [Manitoba]. Types: AMNH. Refs.: Gunder, 1927, Can. Entomol. 59(12): 287, no. 21, pl. B, figs. 21 ♂ D, 21a ♂ V (holotype); dos Passos and Grey (1945 b); Dunford (2009) (fig. 27).

    • syn. helena (dos Passos & Grey, 1957), Proposed to replace Argynnis lais W. H. Edwards, 1884, Can. Entomol. 15(11): 209, with the same data. See Boloria eunomia triclaris synonym lais (Scudder, 1875). Type loc.: see lais Edwards [1884]. Refs.: dos Passos & Grey, 1957, J. N. Y. Entomol. Soc. 63: 95-96.

    • syn. lais Edwards [1884], preocc. by Argynnis lais Scudder (1875), Proc. Boston Soc. Nat. Hist. 17(3): [294]-296. Type loc.: “Red Deer River”; corrected to “Fort Edmonton” by Geddes [1884], Can. Entomol. 15(12): 222. [Alberta]. Types: lectotype in CMNH. Refs.: Edwards, [1884], Can. Entomol. 15(11): 209; figured by W. H. Edwards (1887), Butterflies N. Am. 3(1): pl. Argynnis 2, figs. 1 ♂ D, 2 ♂ V, 3 ♀ D, 4 ♀ V; dos Passos and Grey (1947) (lectotype); Brown (1965) (fig. 26); Dunford (2009) (fig. 26).

  • ssp. dorothea (Moeck, 1947); original combination: Speyeria atlantis dorothea. Type loc.: “Sandia Peak” [Sandia Mountains, Sandoval County, New Mexico]. Types: AMNH. Refs.: Moeck, 1947, Entomol. News 58(3): 73-75, figs. 1 ♂ D, 2 ♂ V (holotype); 3 ♀ D, 4 ♀ V; Dunford (2009) (fig. 45).

  • ssp. electa Edwards, 1878; original combination: Argynnis Electa. Type loc.: Colorado, USA. Types: lectotype in CMNH. Refs.: W. H. Edwards, 1878, Field & Forest 3(9): 143-144; dos Passos and Grey (1947).

    • syn. cornelia Edwards, 1892. Type loc.: “Ouray, and Manitou, Colorado, and in the adjacent districts.Crested Butte in the Ouray region”; defined as “Ouray, Ouray County, Colorado” by dos Passos and Grey (1947). Types: lectotype in CMNH. Refs.: W. H. Edwards, 1892, Can. Entomol. 24(5): 106-107, no. 2; dos Passos and Grey (1947) (lectotype); Brown (1965) (fig. 25).

    • syn. nikias Ehrmann, 1917. Type loc.: “Temez [sic] Springs, New Mexico” [Jemez Springs, Sandoval County]. Types: lectotype in CMNH. Refs.: Ehrmann, 1917, Lepidopterist 1(7): 55-56; dos Passos and Grey (1947) (lectotype); Dunford (2009) (fig. 42).

  • ssp. hesperis Edwards, 1864; original combination: Argynnis Hesperis. Type loc.: “Rocky Mountains”; defined as “Turkey Creek Junction, Jefferson County, Colorado” by dos Passos and Grey (1947). Types: neotype in CMNH. Refs.: Edwards, 1864, Proc. Entomol. Soc. Philad. 2(4): 501, 502-503, no. 3; figured by W. H. Edwards (1868), Butterflies N. Am. 1: pl. Argynnis 7, figs. 1 ♂ D, 2 ♂ V, 3 ♀ D, 4 ♀ V; dos Passos and Grey (1947) (lectotype); dos Passos and Grey (1965) (neotype); Brown (1965) (fig. 21); Dunford (2009) (fig. 25).

  • ssp. hutchinsi (dos Passos & Grey, 1947); original combination: Speyeria atlantis hutchinsi. Type loc.: see hutchinsi Gunder. Refs.: dos Passos and Grey (1947): 20 (adopted Argynnis atlantis beani tr. f. hutchinsi Gunder, 1932, to represent a subspecies-level taxon, with the same data; Code Articles 10.2, 45.5.1).

    • syn. beani tr. f. hutchinsi Gunder, 1932. Type loc.: “Twp 2N Range 1W, Jefferson County, Mont” [Montana]. Types: AMNH. Refs.: Gunder, 1932, Can. Entomol. 64(12): 280; dos Passos and Grey (1947).

  • ssp. lurana (dos Passos & Grey, 1945); original combination: Speyeria atlantis lurana. Type loc.: “Black Hills of South Dakota.Harney Peak” [Custer County]. Types: AMNH. Refs.: dos Passos and Grey (1945 b): 8-9, figs. 23 ♂ D, 24 ♂ V (holotype); Dunford (2009) (fig. 32).

  • ssp. ratonensis (Scott, 1981); original combination: Speyeria atlantis ratonensis. Type loc.: “Raton Mesa, Colfax Co., New Mexico”. Types: LACM. Refs.: Scott, 1981, Papilio (n.s.) 1: 4; figured by J. Scott (1998), Papilio (n.s.) 8: figs. 30 ♂ D, 31 ♂ V, 32 ♀ D, 33 ♀ V; Dunford (2009) (fig. 30).

• hydaspe Boisduval, 1869

  • ssp. davisi (Emmel, Emmel & Mattoon, 1998); original combination: Speyeria hydaspe davisi. Type loc.: “California: Humboldt County; McClellan Mountain peat bog”. Types: LACM. Refs.: J. Emmel, T. Emmel & Mattoon, 1998, Syst. W. N. Am. Butterflies (12): 147, figs. 29 ♂ D, 30 ♂ V (holotype); 31 ♀ D, 32 ♀ V.

  • ssp. hydaspe Boisduval, 1869; original combination: Argynnis Hydaspe. Type loc.: “sud de la Californie; suggested to be “Yosemite Valley, Mariposa County, California” by dos Passos and Grey (1947); defined as “Gold Lake, Sierra County, California” by J. Emmel et al. (1998), Syst. W. N. Am. Butterflies (2): 34. Types: lectotype in NMNH. Refs.: Boisduval, 1869, Annls. Soc. Entomol. Belg. 12: 60, no. 55; figured by Oberthür (1914), Étud. Lépid. Comp. 9(2): pl. 263, fig. 2200 ♂ D&V (lectotype), 2201 ♂ D&V, 2202 ♀ D&V; dos Passos and Grey (1947) (lectotype).

    • syn. purpurascens Edwards, 1877. Type loc.: “Siskiyou County, in the region surrounding Mt. Shasta, as well as in some parts of Oregon.at the Dalles, Oregon.at Soda Springs, Siskiyou County”; defined as “Soda Springs, Siskiyou County, California” by dos Passos and Grey (1947). Types: lectotype in AMNH. Refs.: Edwards, 1877, Proc. Calif. Acad. Sci. [1]7(13): 170-171.

    • syn. hydaspe ab. caliginosa Comstock, 1925. Type loc.: “Modoc Co., Calif” [California]. Types: AMNH. Refs.: Comstock, 1925, Bull. South. Calif. Acad. Sci. 24(3): 66; figured by J. A. Comstock (1927), Butterflies Calif.: pl. 26, fig. 3 ♂ D (holotype).

  • ssp. minor (dos Passos & Grey, 1947); original combination: S[peyeria]. hydaspe minor. Type loc.: see hydaspe minor McDunnough, 1927. Refs.: dos Passos and Grey (1947): 21.

    • syn. rhodope form alt. minor McDunnough, 1927. Type loc.: “Mt. McLean, B. C., (5000 feet)” [near Lillooet, British Columbia]. Types: CNC. Refs.: McDunnough, 1927, Can. Entomol. 59(7): 155.

    • syn. rhodope tr. f. gregsoni Gunder, 1932. Type loc.: “Mount Washington (6000 ft.), Forbidden Plateau, V. I., B. C., Can” [Vancouver Island, British Columbia]. Types: AMNH. Refs.: Gunder, 1932, Can. Entomol. 64(12): 281.

  • ssp. rhodope Edwards, 1874; original combination: Argynnis Rhodope. Type loc.: British Columbia, Canada. Types: lectotype in CMNH. Refs.: W. H. Edwards, 1874, Trans. Am. Entomol. Soc. 5(1): 13-14; figured by W. H. Edwards (1875), Butterflies N. Am. 2(2): pl. Argynnis 6, figs. 1 ♂ D, 2 ♂ V, 3 ♀ D, 4 ♀ V; dos Passos and Grey (1947) (lectotype); Brown (1965) (fig. 27).

    • syn. sakuntala Skinner, 1911. Type loc.: “Kaslo, B. C” [British Columbia]. Types: lectotype in CMNH. Refs.: Skinner, 1911, Entomol. News 22(3): 108; dos Passos and Grey (1947) (lectotype); Kondla (2001) (synonymy).

    • syn. skinneri Holland, 1931, Proposed to replace Argynnis sakuntala Skinner, 1911, erroneously though to be preoccupied; a junior objective synonym of sakuntala Refs.: W. Holland, 1931, Butterfly Book (rev. ed.): 95, pl. 56, fig. 4 ♀ V.

  • ssp. tehama (Emmel, Emmel & Mattoon, 1998); original combination: Speyeria hydaspe tehama. Type loc.: “California: Tehama County; Covelo Road, at Government Flat, 5900’ elevation”. Types: LACM. Refs.: J. Emmel, T. Emmel & Mattoon, 1998, Syst. W. N. Am. Butterflies (12): 146, figs. 25 ♂ D, 26 ♂ V (holotype); 27 ♀ V, 28 ♀ V.

  • ssp. viridicornis Comstock, 1925; original combination: Argynnis hydaspe form viridicornis. Type loc.: “Greenhorn Mountains, Calif” [Kern County]. Types: LACM. Refs.: Comstock, 1925, Bull. South. Calif. Acad. Sci. 24(3): 63-64; figured by J. A. Comstock (1927), Butterflies Calif.: pl. 25, figs. 6 ♂ V (holotype).

• idalia (Drury [1773])

  • ssp. idalia (Drury [1773]); original combination: Nym[phalis]. Phal[eratus]. Idalia. Type loc.: “New York”; neotype from “New York City, New York County, New York”, labeled “West Farms, New York City”. Types: AMNH. Refs.: Drury (1970): 25, pl.XIII, figs. 1 ♀ D, 2 ♂ D, 3 ♀ V, index (1770); 2: index (1773, name given); dos Passos and Grey (1947) (neotype); B. Williams (2001).

    • syn. astarte Fisher [1859], preocc. by Argynnis astarte Doubleday, 1848, in: Gen. diurn. Lepid. 1(21): 181. Type loc.: “Nova Caesarea.New Jersey.Succasunna Plains near Schooley’s Mountain, in Morris Co”. Types: FMNH. Refs.: Fisher, [1859], Proc. Acad. Nat. Sci. Philad. 10(13): 179-180, pl. 2 ♀ D&V.

    • syn. ashtaroth Fisher [1860], proposed to replace Argynnis astarte J. Fisher, [1859], with the same data (Code Article 60.3). Types: FMNH. Refs.: Fisher, [1860], Proc. Acad. Nat. Sci. Philad. 11(27): 352.

    • syn. infumata Oberthür, 1912. Type loc.: “Newark, N. J” [Essex County, New Jersey]. Types: “holotype” in NMNH. Refs.: Oberthür, 1912, Études Lépid. Comp. 6: 315, pl. 104, fig. 969 ♂ D&V.

    • syn. dolli Gunder, 1927. Type loc.: “Richmond Hill, Long Island, N. Y” [Queens County, New York]. Types: “holotype” in NMNH. Refs.: Gunder, 1927, Can. Entomol. 59(12): 286, no. 20, pl. B, figs. 20 ♀ D, 20b ♀ V (“holotype”).

  • ssp. occidentalis (Williams, 2001); original combination: Speyeria idalia occidentalis. Type loc.: “1 mile S.E. of Crete, IL” [Will County, Illinois]. Types: INHS. Refs.: B.E. Williams (2001).

    • syn. pallida Eisner, 1942. Type loc.: “Decorah, Iowa” [Winneshiek County]. Types: “Holotype” in NCBN. Refs.: Eisner, 1942, Zool. Meded. 24(1/2): 124.

• irene Boisduval, 1869

  • ssp. cottlei Comstock, 1925; original combination: Argynnis cottlei. Type loc.: “near Alturas, Modoc Co., Calif” [California]. Types: LACM. Refs.: Comstock, 1925, Bull. South. Calif. Acad. Sci. 24(3): 64-65; figured by J. A. Comstock (1927), Butterflies Calif.: pl. 26, fig. 1 ♀ V (holotype); Zhang et al. (2020) (as subspecies of irene).

  • ssp. dodgei Gunder, 1931; original combination: Argynnis dodgei. Type loc.: “Diamond Lake, Douglas Co., Oregon”. Types: AMNH. Refs.: Gunder, 1931, Bull. South. Calif. Acad. Sci. 30(2): 46; Dunford (2009) (fig. 35); Zhang et al. (2020) (as subspecies of irene).

  • ssp. hanseni Emmel, Emmel & Mattoon, 1998; original combination: Speyeria atlantis hanseni. Type loc.: “California: Tehama County; Anthony Peak on Covelo Road”. Types: LACM. Refs.: J. Emmel, T. Emmel & Mattoon, 1998, Syst. W. N. Am. Butterflies (12): 152, figs. 47 ♂ D, 48 ♂ D (holotype); 49 ♀ D, 50 ♀ V; Dunford (2009) (fig. 34); Zhang et al. (2020) (as subspecies of irene).

  • ssp. irene Boisduval, 1869; original combination: [Argynnis Egleis variété] Irene. Type loc.: “les lieux incultes et aux borde des forets, dans l’intérieur de la Californie”; defined as “Massack, Plumas County, California” by dos Passos and Grey (1947), Am. Mus. Novit. (1370): 20; suggested to be “Gold Lake, Sierra County, California” by J. Emmel et al. (1998), Syst. W. N. Am. Butterflies (2): 33. Types: lectotype in NMNH. Refs.: Boisduval, 1869, Annls. Soc. Entomol. Belg. 12: 59-60, under no. 53; figured by Oberthür (1914), Étud. Lépid. Comp. 9(2): pl. 262, fig. 2196 ♀ D&V; dos Passos and Grey (1947) (lectotype); Zhang et al. (2020) (to species).

• leto Behr, 1862

  • ssp. eileenae (Emmel, Emmel & Mattoon, 1998); original combination: Speyeria cybele eileenae. Type loc.: “California: Humboldt County; Bear River Ridge Road, 0 to 5 road miles E of junction of Wildcat Road, 7 air miles SE of Ferndale, 2300’ elevation”. Types: LACM.

  • ssp. leto Behr, 1862; original combination: Argynnis leto. Type loc.: “California” implied from title of paper, “peculiar to the western slope” from the description; neotype from “Near Carson City, Ormsby County, Nevada” [Ormsby County no longer exists]. Types: neotype in CMNH. Refs.: dos Passos and Grey (1947) (neotype); Pohl et al. (2010) (listed as Speyeria leto); Zhang et al. (2022) (to species).

    • syn. letis Wright, 1905. Type loc.: “Northern California” “Sisson, Cal” [Mount Shasta City, Siskiyou County, California]. Types: CAS.

    • syn. valesinoidesalba (Reuss [1926]), emend. Type loc.: not stated. Types: ZMHU.

    • syn. lethe Gunder, 1934. Type loc.: “Big Horn Mts., Wyoming”. Types: “Holotype” in AMNH. Refs.: Gunder, 1934, Can. Entomol. 66(6): 125.

  • ssp. letona (dos Passos & Grey, 1945); original combination: Speyeria cybele letona. Type loc.: “City Creek Canyon, Salt Lake City, Utah, 4500 feet” [Salt Lake County]. Types: AMNH. Refs.: dos Passos and Grey (1945 b).

  • ssp. pugetensis (Chermock & Frechin, 1947); original combination: Speyeria cybele pugetensis. Type loc.: “Stimson Creek, near Belfair, Mason County, Wash” [Washington]. Types: AMNH.

• mormonia Boisduval, 1869

  • ssp. mormonia Boisduval, 1869; original combination: Argynnis Mormonia. Type loc.: USA: Arizona, White Mts. Refs.: Boisduval, 1869, Annls. Entomol. Soc. Belg. 12: 58-59, no. 51; figured by Oberthür (1914), Étud. Lépid. Comp. 9(2): pl. 261, figs. 2192 ♂ D&V (lectotype), 2193 ♀ D&V.

    • syn. astarte Edwards, 1862, preocc. by Argynnis astarte Doubleday, 1848, in: Gen. diurn. Lepid. 1(21): 180. Type loc.: “Oregon; California”; defined as “Monache Meadows, Tulare County, California” by dos Passos and Grey (1947). Types: “neotype” in FMNH. Refs.: W. H. Edwards, 1862, Proc. Entomol. Soc. Philad. 1(7): 221-222, no. 1; dos Passos and Grey (1947) (neotype, which is the lectotype of arge Strecker).

    • syn. montivaga Behr, 1864, preocc. by Argynnis montivaga Behr, 1863, Proc. Calif. Acad. Sci. [1]3(6): 84, no. 2. Type loc.: “always found in mountainous regions”; defined as “Monache Meadows, Tulare County, California” by dos Passos and Grey (1947). Types: “neotype” in FMNH. Refs.: Behr, 1864, In: W. H. Edwards, Proc. Entomol. Soc. Philad. 3(3): 435, nos. 4 and 5.

    • syn. arge Strecker, 1878. Type loc.: “California”; defined as “Monache Meadows, Tulare County, California” by dos Passos and Grey (1947). Types: lectotype in FMNH. Refs.: Strecker, 1878, Butterflies & Moths N. Am., Comp. Syn. Cat.: 114, no. 210; dos Passos and Grey (1947) (lectotype).

  • ssp. obsidiana (Emmel, Emmel & Mattoon, 1998); original combination: Speyeria mormonia obsidiana. Type loc.: USA: California, Mono Co. Types: LACM. Refs.: J. Emmel, T. Emmel & Mattoon, 1998, Syst. W. N. Am. Butterflies (12): 145-146, figs. 21 ♂ D, 22 ♂ V (holotype); 23 ♀ D, 24 ♀ V.

• nausicaa Edwards, 1874

  • ssp. chitone Edwards, 1879; original combination: Argynnis chitone. Type loc.: “Southern Utah and Arizona”, interpreted as “Cedar Breaks National Monument, Iron County, Utah” (dos Passos and Grey 1947). Types: lectotype in CMNH. Refs.: W. H. Edwards, 1879, Can. Entomol. 11(5): 82; Brown (1965) (fig. 23); Dunford (2009) (fig. 40); Zhang et al. (2020) (as subspecies of nausicaa).

    • syn. wasatchia (dos Passos & Grey, 1945), “considered to be a synonym of S. h. chitone by Hammond (1991), J. Res. Lepid. 29(1/2): 59 (as S. atlantis chitone), and a synonym of S. h. tetonia by J. Scott et al. (1998), Papilio (n.s.) 8: 6, where tetonia was given priority over wasatchia”. Type loc.: “Payson Canyon, Payson, Utah” [Utah County]. Types: AMNH. Refs.: dos Passos and Grey (1945 b): 8, figs. 25 ♂ D, 26 ♂ V (holotype); Dunford (2009) (fig. 40).

  • ssp. elko (Austin, 1984); original combination: Speyeria atlantis elko. Type loc.: “Nev(ada): Elko Co(unty); Owyhee R(iver). Valley, Wildhorse Creek Campg(round)., ca. 10 mi(les). S(outh). (of) Mountain City”. Types: NVMC. Refs.: Austin, 1984, J. Lepid. Soc. 37(3): 244-247, figs. 1 upper left ♂ D, 2 upper right ♂ V (holotype), 1 lower left ♀ D, 2 lower left ♀ V; Dunford (2009) (fig. 37); Zhang et al. (2020) (as subspecies of nausicaa).

  • ssp. greyi (Moeck, 1950); original combination: Speyeria atlantis greyi. Type loc.: “Lamoille Canyon, Ruby Mountains, Elko County, Nevada.at an elevation of about 8,500 feet”. Types: AMNH. Refs.: Moeck, 1950, Entomol. News 61(3): 61-62, figs. 1 ♂ D, 2 ♂ V (holotype); 3 ♀ D, 4 ♀ V; Dunford (2009) (fig. 31); Zhang et al. (2020) (as subspecies of nausicaa).

  • ssp. nausicaa Edwards, 1874; original combination: Argynnis Nausicaa. Type loc.: “Rocky Canon, Arizona”; further defined as “Graham County” by Brown (1965). Types: lectotype in CMNH. Refs.: W. H. Edwards, 1874, Trans. Am. Entomol. Soc. 5(2): 104-105; figured by W. H. Edwards (1888), Butterflies N. Am. 3(4): pl. Argynnis 10, figs. 1 ♂ D, 2 ♂ V, 3 ♀ D, 4 ♀ V; dos Passos and Grey (1947) (lectotype); Brown (1965) (fig. 22); Dunford (2009) (fig. 44); Zhang et al. (2020) (to species); Campbell et al. (2022) (population genomics).

    • syn. arizonensis Elwes, 1889. Type loc.: “Arizona (Wheeler Expedition).also in Colorado”; suggested to be “Rocky Can[y]on, Arizona” by dos Passos and Grey (1947), Am. Mus. Novit. (1370): 19; defined as “Rock Canyon, Graham County, Arizona” by F. Brown (1965), Trans. Am. Entomol. Soc. 91(3): 301. Types: neotype in CMNH. Refs.: Elwes, 1889, Trans. Entomol. Soc. Lond. (4)1889(4): 546; cited Mead [1876], Rep. Coll. diurn. Lepid. 5(Zool.): 752; dos Passos and Grey (1947) (neotype; same as lectotype of Argynnis nausicaa Edwards, 1874).

  • ssp. schellbachi (Garth, 1949); original combination: Speyeria atlantis schellbachi. Type loc.: “Neal Springs, North Rim, Grand Canyon National Park, Coconino County, Arizona, 8,175 feet”. Types: LACM. Refs.: Garth, 1949, Bull. South. Calif. Acad. Sci. 48(1): 1-2, figs. 1 ♂ D, 2 ♂ V (holotype); 3 ♀ D, 4 ♀ V; Dunford (2009) (fig. 43); Zhang et al. (2020) (as subspecies of nausicaa).

  • ssp. tetonia (dos Passos & Grey, 1945); original combination: Speyeria atlantis tetonia. Type loc.: “Teton Mountains region, Wyoming.Teton Mountains” [Teton County]. Types: AMNH. Refs.: dos Passos and Grey (1945 b): 9-10, figs. 27 ♂ D, 28 ♂ V (holotype); Dunford (2009) (fig. 38); Zhang et al. (2020) (as subspecies of nausicaa).

    • syn. conquista (dos Passos & Grey, 1945), assumed mislabeled specimens of hydaspe rhodope. Type loc.: “Little Tesuque Canyon, near Santa Fe, New Mexico, 8000 feet”, almost certainly in error [Santa Fe County]. Types: AMNH. Refs.: dos Passos and Grey (1945 b): 8, figs. 21 ♂ D, 22 ♂ V (holotype); Dunford (2009) (fig. 39). As noted by Ferris (1981), in: Ferris and F. Brown (1981), Butterflies Rocky Mtn. States: 311-312, S. hydaspe has not been recorded from New Mexico since the description of this taxon; the types are therefore assumed to be mislabeled. Zhang et al. (2020): 19, fig. 14, determine that the holotype ‘clusters closely’ with the holotype of tetonia.

  • ssp. viola (dos Passos & Grey, 1945); original combination: Speyeria atlantis viola. Type loc.: “Sawtooth Mountains, Idaho.Trail Creek, 7400 feet” [Custer County]. Types: AMNH. Refs.: dos Passos and Grey (1945 b): 10, figs. 29 ♂ D, 30 ♂ V (holotype); Dunford (2009) (fig. 36); Zhang et al. (2020) (as subspecies of nausicaa).

• nokomis Edwards, 1862

  • ssp. apacheana Skinner, 1918; original combination: Argynnis apacheana. Type loc.:  “Arizona”; suggested to be “mountains west of Camp Independence” by F. Brown [1958], J. N. Y. Entomol. Soc. 65(3/4): 228-229; defined as “vicinity of Independance, Inyo Co., California” by F. Brown (1983), J. Lepid. Soc. 37(1): 79-80. [Inyo County]. Types: lectotype in CMNH. Refs.: Skinner, 1918, Entomol. News 29(2): 67-68; cited W. H. Edwards (1868), Butterflies N. Am. 1: pl. Argynnis 4, figs. 1 ♂ D, 2 ♂ V (lectotype), 3 ♀ D, 4 ♀ V (as Argynnis nokomis).

    • syn. hermosa Comstock, 1925. Type loc.: “Round Valley, Inyo County, California”. Types: “Holotype” in LACM. Refs.: Comstock, 1925, Bull. South. Calif. Acad. Sci. 24(1): 3-4; figured by J. A. Comstock (1927), Butterflies Calif.: pl. 23, fig. 4 ♂ D (holotype).

  • ssp. carsonensis (Austin, 1998); original combination: Speyeria nokomis carsonensis. Type loc.: “Nevada: Douglas County; Carson Valley, Scossa Ranch, Nevada state route 206, 3.6 miles south of Nevada state route 207, 1463 m, T12N R19E S26 on USGS Freel Peak, Calif.-Nev. 15’ quadrangle”. Types: FLMNH. Refs.: Austin, 1998, Syst. W. N. Am. Butterflies (46): 573-574, figs. 1 ♂ D, 2 ♂ V (holotype); 3 ♀ D, 4 ♀ V.

  • ssp. coerulescens Holland, 1900; original combination: Argynnis nitocris var. coerulescens. Type loc.: “State of Chihuahua, Mexico.at the head of the Rio Piedras Verdes, in Chihuahua, at an elevation of from 7,100 to 7,300 feet above sea level in the Sierra Madre region”. Types: lectotype in CMNH. Refs.: Holland, 1900, Entomol. News 10(11): 332-333; figured by W. Holland (1931), Butterfly Book (rev. ed.): pl. 55, fig. 2 ♀ D (paralectotype), 3 ♂ D (lectotype), 4 ♀ V (paralectotype); dos Passos and Grey (1947) (lectotype).

  • ssp. melaena (Mooser & García, 1980); original combination: Speyeria nokomis melaena

  • ssp. nitocris Edwards, 1874; original combination: Argynnis Nitocris. Type loc.: “White Mountains, Arizona”. Types: CMNH. Refs.: Edwards, 1874, Trans. Am. Entomol. Soc. 5(1): 15-16; figured by W. H. Edwards (1887), Butterflies N. Am. 3(1): pl. Argynnis 1, figs. 1 ♂ D, 2 ♂ V (holotype); 3 ♀ D, 4 ♀ V; Ferris and Fisher (1971).

  • ssp. nokomis Edwards, 1862; original combination: Argynnis Nokomis. Type loc.: Beulah, New Mexico, USA. Types: AMNH. Refs.: Edwards, 1862, Proc. Acad. Nat. Sci. Philad. 14(5): 221-222, no. 1; Holland (2008); Scott and Fisher (2014) (taxonomy).

    • syn. nigrocaerulea Cockerell & Cockerell, 1900. Type loc.: “Sapello Canon, Beulah, New Mexico” [San Miguel County]. Types: NMNH. Refs.: Cockerell & Cockerell, 1900, Entomol. News 11(10): 622; figured by W. Cockerell and T. Cockerell (1902), Birds & Nature: unnumbered plate.

    • syn. valesinoidesalba (Reuss [1926]), emend. Type loc.: not stated. Types: ZMHU. Refs.: Reuss, [1926], Dtsche. Entomol. Z. 1926(1): 69.

    • syn. rufescens Cockerell, 1909. Type loc.: “At Beulah, in the Sapello Cañon” [San Miguel County, New Mexico]. Types: NMNH. Refs.: Cockerell, 1909, Entomol. Rec. & J. Var. 21(7/8): 186.

    • syn. tularosa Holland, 2010. Type loc.: “Mescalero, Tularosa River, Otero County, New Mexico, ca. 7000’”. Types: CMNH. Refs.: Holland, 2010, J. Res. Lepid. 42: 79, 79, figs. 1, ♂ D, 2 ♂ V, 3 ♂ D, 4 ♂ V, 5 ♀ D, 6 ♀ V (holotype), 7 ♀ D, 8 ♀ V.

  • ssp. wenona (dos Passos & Grey, 1945); original combination: Speyeria wenona Refs.: dos Passos and Grey (1945 b).

• zerene Boisduval, 1852

  • ssp. atossa Edwards, 1890; original combination: Argynnis Atossa. Type loc.: “North American.Tehachipe [sic], south California.in a little valley about four miles from town, by a small stream” [Tehachapi, Kern County]. Types: lectotype in CMNH. Refs.: Edwards, 1890, Butterflies N. Am. 3(10): [127]-[128], pl. Argynnis 8, figs. 4 ♂ D, 5 ♂ V, 6 ♀ D; Zhang et al. (2020) (transferred from adiaste).

    • syn. tejonica Comstock, 1925. Type loc.: “Tejon region (Collins Ranch), California” [Los Angeles County]. Types: LACM. Refs.: Comstock, 1925, Bull. South. Calif. Acad. Sci. 24(3): 68; figured by J. A. Comstock (1927), Butterflies Calif.: pl. 28, fig. 6 ♂ V; Zhang et al. (2020) (transferred from adiaste).

    • syn. sineargentatus Gunder, 1927. Type loc.: “Mammoth, Mono Co., Calif” [California]. Types: “Holotype” in AMNH. Refs.: Gunder, 1927, Can. Entomol. 59(12): 286, no. 19, pl. B, fig. 19 ♂ V (“holotype”).

  • ssp. behrensii Edwards, 1869; original combination: Argynnis Behrensii. Type loc.: “Mendocino, California” [Mendocino County]. Types: Lectotype in CMNH, designated by dos Passos and Grey (1947), Am. Mus. Novit. (1370): 13. F. Brown (1965), Trans. Am. Entomol. Soc. 91(3): 269-271, fig. 9 ♂ D&V, noted that this specimen may not be one of the original types; therefore the “lectotype” may need to be redesignated as neotype. Refs.: Edwards, 1869, Trans. Am. Entomol. Soc. 2(4): 370-371; figured by W. H. Edwards (1868), Butterflies N. Am. 1: pl. Argynnis 12, figs. 1 ♂ D, 2 ♂ V, 3 ♀ D, 4 ♀ V; dos Passos and Grey (1947) (lectotype, but see Brown (1965)).

  • ssp. bremnerii Edwards, 1872; original combination: Argynnis Bremnerii. Type loc.: “San Juan Island” [San Juan County, Washington]. Types: CMNH. Refs.: Edwards, 1872, Trans. Am. Entomol. Soc. 4(1/2): 61, 63-64, no. 4; figured by W. H. Edwards (1874), Butterflies N. Am. 2(1): pl. Argynnis 4, figs. 1 ♂ D, 2 ♂ V, 3 ♀ D, 4 ♀ V; dos Passos and Grey (1947) (lectotype).

    • syn. sordida Wright, 1905. Type loc.: “Puget Sound, Washington” implied from the text. Types: “Holotype” (#4326) in CAS; figured by Howe (1975), Butterflies N. Am.: pl. 31, fig. 5 ♂ D&V. The text clearly indicates that this taxon was described as an individual variant. Gunder (1932), Can. Entomol. 64(12): 281, determined that the holotype is a worn specimen of bremnerii.

  • ssp. gloriosa (Moeck, 1957); original combination: Speyeria zerene gloriosa. Type loc.: “Illinois River Valley, Josephine County, southwestern Oregon, on the road leading west from Selma, along the river, from about 1,600 feet elevation down to about 1,100 feet”. Types: AMNH. Refs.: Moeck, 1957, Geogr. var. Speyeria: 21, 24, unnumbered fig. ♂ D&V (holotype), ♀ D&V; fig. p. 23, 9 ♂ V, 6 ♀ V.

  • ssp. gunderi Comstock, 1925; original combination: Argynnis gunderi. Type loc.: “Modoc Co., Cal”, interpreted as “Mountains above Davis Creek ranger station, north of Alturas” (dos Passos and Grey 1947) [California, USA]. Types: LACM. Refs.: Comstock, 1925, Bull. South. Calif. Acad. Sci. 24(3): 67-68; figured by J. A. Comstock (1927), Butterflies Calif.: pl. 27, fig. 4 ♂ D (holotype).

    • syn. cynna (dos Passos & Grey, 1945). Type loc.: “Ruby Valley, Elko County, Nevada.Humboldt National Forest”. Types: AMNH. Refs.: dos Passos and Grey (1945 b); Grey (1989): 3 (synonymy).

  • ssp. hippolyta Edwards, 1879; original combination: Argynnis Hippolyta. Type loc.: “Oregon [.] Northern California”; interpreted as “Oceanside, Tillamook County, Oregon [,USA]” (dos Passos and Grey 1947). Types: Lectotype in CMNH, designated by dos Passos and Grey (1947), Am. Mus. Novit. (1370): 13; figured by W. Holland (1931), Butterfly Book (rev. ed.): 93, pl. 12, fig. 10 ♂ D, F. Brown (1965), Trans. Am. Entomol. Soc. 91(3): 266, fig. 8 ♂ D&V, and Howe (1975), Butterflies N. Am.: pl. 22, figs. 9 ♂ D, 10 ♂ V. Refs.: Edwards, 1879, Can. Entomol. 11(5): 81; dos Passos and Grey (1947) (lectotype).

  • ssp. malcolmi Comstock, 1920; original combination: Argynnis Malcolmi. Type loc.: “Mammoth, Cal” [Mono County, California, USA]. Types: LACM. Refs.: Comstock, 1920, Southwest Sci. Bull. [1]: 4-5, pl. [1], figs. 4 ♂ D (holotype), 5 ♀ D, 6 ♂ V.

  • ssp. monticola Behr, 1863; original combination: Argynnis monticola. Type loc.: “California” implied from the title of the paper; “Yosemite Valley and some other localities of the Sierra Nevada” in the descripton; suggested to be “several localities in the Sierra Nevada” by Behr (1862), Proc. Calif. Acad. Sci. [1]2(12): 175, no. 8 [unnamed]; defined as “Yosemite Valley, Mariposa County, California” by dos Passos and Grey (1947), Am. Mus. Novit. (1370): 12; neotype from “Mather, ca. 15 air miles NW of Yosemite Valley, Tuolumne County, California”. Types: Neotype in CAS, designated by J. Emmel et al. (1998), Syst. W. N. Am. Butterflies (6): 97, figs. 7 ♂ D, 8 ♂ V. dos Passos and Grey (1947), Am. Mus. Novit. (1370): 12, designated a “lectotype” in the USNM, but J. Emmel et al. (1990), suggested this is not a syntype, and that a neotype was required. Refs.: Behr, 1863, Proc. Calif. Acad. Sci. [1]3(6): 84, no. 4.

  • ssp. myrtleae (dos Passos & Grey, 1945); original combination: Speyeria zerene myrtleae. Type loc.: San Mateo, California, USA. Types: AMNH. Refs.: dos Passos and Grey (1945 b).

  • ssp. picta McDunnough, 1924; original combination: [Argynnis halcyone] race picta. Type loc.: “Aspen Grove, B. C” [British Columbia, Canada]. Types: CNC. Refs.: McDunnough, 1924, Can. Entomol. 56(2): 43.

    • syn. garretti Gunder, 1932. Type loc.: “Cranbrook, B. C., Can” [British Columbia, Canada]. Types: AMNH. Refs.: Gunder, 1932, Can. Entomol. 64(12): 282.

  • ssp. platina Skinner, 1897; original combination: Argynnis platina. Type loc.: “Ogden, Utah.Beaver Canon, Idaho”, interpreted as “Ogden, Weber County, Utah” (dos Passos and Grey 1947). Types: CMNH. Refs.: Skinner, 1897, Can. Entomol. 29(6): 154; dos Passos and Grey (1947) (lectotype).

    • syn. pfoutsi Gunder, 1933. Type loc.: “Mt. Loafer, nr. Payson, Utah” [Utah County]. Types: AMNH. Refs.: Gunder, 1933, Can. Entomol. 65(8): 171-172; Grey (1989): 3 (synonymy).

  • ssp. puntareyes (Emmel, Emmel & Mattoon, 1998); original combination: Speyeria zerene puntareyes. Type loc.: “California: Marin County; Point Reyes Peninsula; stabilized dunes 2.0 air miles SW of RCA radio towers (0.4 air mi. WSW of BM 157 on 1954 USGS 15’ Pt. Reyes quadrangle)”. Types: LACM. Refs.: J. Emmel & T. Emmel, 1998, Syst. W. N. Am. Butterflies (35): 451-452, figs. 1 ♂ D, 2 ♂ V (holotype); 3 ♀ D, 4 ♀ V.

  • ssp. sinope (dos Passos & Grey, 1945); original combination: Speyeria zerene sinope. Type loc.: Rocky Mountain National Park, Colorado, USA. Types: AMNH. Refs.: dos Passos and Grey (1945 b).

  • ssp. sitka (Hammond, Harry & McCorkle, 2001); original combination: Speyeria zerene sitka. Type loc.: “AK, Haines” [Alaska]. Types: CAS. Refs.: Hammond, Harry & McCorkle, 2001, Butterflies B. C.: 274-276, figs. ♂ D&V (holotype), ♀ D&V.

  • ssp. sonomensis (Emmel, Emmel & Mattoon, 1998); original combination: Speyeria zerene sonomensis. Type loc.: “California: Sonoma County; south end of Sonoma Mountains, 1.5 air mi. N. of intersection of Lakeville Hwy. & Rt. 37, 175’ elev”. Types: LACM. Refs.: J. Emmel, T. Emmel & Mattoon, 1998, Syst. W. N. Am. Butterflies (36): 455-456, figs. 1 ♂ D, 2 ♂ V (holotype); 3 ♀ D, 4 ♀ V.

  • ssp. zerene Boisduval, 1852; original combination: Argynnis Zerene. Type loc.: California, USA. Types: lectotype in NMNH. Refs.: Boisduval, 1852, Annls. Soc. Entomol. Fr. (2)10(2): 303, no. 44; figured by Oberthür (1914), Étud. Lépid. Comp. 9(2): pl. 258, figs. 2168 ♂ D&V (lectotype), 2169 ♀ D&V; dos Passos and Grey (1947) (lectotype).

    • syn. conchyliatus Comstock, 1925. Type loc.: “Mt. Shasta, Calif” [Siskiyou County, California]. Refs.: Comstock, 1925, Bull. South. Calif. Acad. Sci. 24(3): 63; figured by J. A. Comstock (1927), Butterflies Calif.: pl. 25, figs. 1 ♂ V (holotype).

    • syn. shastaensis Comstock, 1925. Type loc.: “Mt. Shasta, Siskiyou Co” [California]. Refs.: Comstock, 1925, Bull. South. Calif. Acad. Sci. 24(3): 65-66 NHMLA; figured by J. A. Comstock (1927), Butterflies Calif.: pl. 26, fig. 1 ♂ D (“holotype”).

Genus Brenthis

Genus: Brenthis Hübner [1819]

Type species: Brenthis hecate ([Denis & Schiffermüller], 1775); original combination: Papilio hecate [Denis & Schiffermüller], 1775

Synonyms:

• Neobrenthis Koçak, 1980. Type species: Brenthis hecate ([Denis & Schiffermüller], 1775); original combination: Papilio hecate [Denis & Schiffermüller], 1775. Refs.: Koçak (1980): 30

References: Hübner ([1819]): 30

4 species:

• daphne (Bergsträsser, 1780)

  • ssp. daphne (Bergsträsser, 1780); original combination: Papilio daphne

    • syn. daphne ([Denis & Schiffermüller], 1775), nom. nud.

    • syn. chloris (Esper, 1778), preocc.

    • syn. epidaphne (Fruhstorfer, 1907). Type loc.: Saratow.

    • syn. syriaca Belter, 1935. Type loc.: Syria, Akbes.

    • syn. anatolica Belter, 1935. Type loc.: Aksehir, Anatolia.

  • ssp. fumida (Butler, 1882); original combination: Argynnis daphne var. fumida. Type loc.: Posiette Bay, NE.Korea.

    • syn. mediofusca (Matsumura, 1929). Type loc.: Korea.

  • ssp. iwatensis Okano, 1951

  • ssp. japygia (Stauder, 1921); original combination: Argynnis daphne japygia

  • ssp. nikator (Fruhstorfer, 1909); original combination: Argynnis daphne nikator

  • ssp. ochroleuca (Fruhstorfer, 1907); original combination: Argynnis daphne ochroleuca. Type loc.: Amur.

  • ssp. rabdia (Butler, 1877); original combination: Argynnis rabdia

• hecate ([Denis & Schiffermüller], 1775)

  • ssp. aigina (Fruhstorfer, 1908); original combination: Argynnis hecate aigina

  • ssp. alaica (Staudinger, 1886); original combination: Argynnis hecate var. alaica

  • ssp. duxtina Klazlauskas, 1984

  • ssp. hecate ([Denis & Schiffermüller], 1775); original combination: Papilio hecate

  • ssp. latericia Tuzov, 1999; original combination: Brenthis hecate latericia. Type loc.: Tajikistan, Peter I Mts.

  • ssp. tergemina Churkin & Zhdanko, 2002; original combination: Brenthis hecate tergemina

  • ssp. transcaucasica (Wnukowsky, 1929); original combination: Argynnis hecate transcaucasica

    • syn. caucasica (Staudinger, 1871)

  • ssp. triburniana (Fruhstorfer, 1908); original combination: Argynnis hecate triburniana

  • ssp. warren Kudrna, 1974

• ino (Rottemburg, 1775)

  • ssp. achasis (Fruhstorfer, 1907), repl. name; original combination: Argynnis ino achasis

    • syn. clara (Staudinger, 1892), preocc.

  • ssp. acrita (Fruhstorfer, 1907); original combination: Argynnis ino acrita

    • syn. discolus (Fruhstorfer, 1907)

    • syn. borealis (Seitz [1909]), preocc.

    • syn. vitimensis (Wnukowsky, 1929), repl. name

  • ssp. adula (Fruhstorfer, 1910); original combination: Argynnis ino adula

  • ssp. amurensis (Staudinger, 1887); original combination: Argynnis ino var. amurensis

  • ssp. eporedia (Fruhstorfer, 1910); original combination: Argynnis ino eporedia

  • ssp. ino (Rottemburg, 1775); original combination: Papilio ino

    • syn. tergesta (Fruhstorfer, 1910)

  • ssp. magadanica (Kurentzov, 1970); original combination: Argynnis ino magadanica

  • ssp. paidicus (Fruhstorfer, 1907); original combination: Argynnis ino paidicus

  • ssp. parvimarginalis (Nakahara, 1926), repl. name; original combination: Argynnis ino parvimarginalis

    • syn. NA (Matsumura, 1925)

  • ssp. schmitzi Wagener, 1983; original combination: Brenthis ino schmitzi

  • ssp. siopelus (Fruhstorfer, 1907); original combination: Argynnis ino siopelus

    • syn. herzi (Seitz [1909])

  • ssp. tigroides (Fruhstorfer, 1907); original combination: Argynnis ino tigroides

  • ssp. trachalus (Fruhstorfer, 1917); original combination: Argynnis ino trachalus

• mofidii Wyatt, 1969

  • ssp. mofidii Wyatt, 1969; original combination: Brenthis hecate mofidii

  • ssp. zabensis Leestmans & Carbonell, 1993; original combination: Brenthis mofidii zabensis

Genus Issoria

Genus: Issoria Hübner [1819]

Type species: Issoria lathonia (Linnaeus, 1758); original combination: Papilio lathonia Linnaeus, 1758

Synonyms:

• Rathora Moore, 1900. Type species: Issoria lathonia (Linnaeus, 1758); original combination: Papilio lathonia Linnaeus, 1758. Refs.: Moore (1900): 241
• Kuekenthaliella Reuss, 1921. Type species: Issoria gemmata (Butler, 1881); original combination: Argynnis gemmata Butler, 1881. Refs.: Reuss (1921 a): 16
• Pseudorathora Reuss, 1926. Type species: Issoria lathonia (Linnaeus, 1758); original combination: Rathora isaeae [sic] f.geogr. isaeoides Reuss, 1925. Refs.: Reuss (1926 b): 68
• Prokuekenthaliella Reuss, 1926. Type species: Issoria baumanni (Rebel & Rogenhofer, 1894); original combination: Argynnis excelsior Butler, [1896]. Refs.: Reuss (1926 a): 435

References: Hübner ([1819]): 31

9 species:

• altissima (Elwes, 1882); original combination: Argynnis altissima

• baileyi Huang, 1998; original combination: Kuekenthaliella baileyi

• baumanni (Rebel & Rogenhofer, 1894)

  • ssp. baumanni (Rebel & Rogenhofer, 1894); original combination: Argynnis baumanni

  • ssp. excelsior (Butler [1896]); original combination: Argynnis excelsior. Type loc.: Ruwenzori.

    • syn. uganda (Röber, 1937). Type loc.: Uganda.

    • syn. alberici (Dufrane, 1945). Type loc.: Bilati.

  • ssp. katangae (Neave, 1910); original combination: Brenthis excelsior katangae

  • ssp. orientalis Kielland, 1990; original combination: Issoria baumanni orientalis. Type loc.: Tanzania, Mufindi, Luisenga, 1750m.

• eugenia (Eversmann, 1847)

  • ssp. eugenia (Eversmann, 1847); original combination: Argynnis eugenia

  • ssp. genia (Fruhstorfer, 1903); original combination: Argynnis eugenia genia

    • syn. rheaoides Draeseke & Reuss, 1925

    • syn. anargyron (Oberthür, 1914)

  • ssp. montana (Bang-Haas, 1906); original combination: Argynnis eugenia var. montana

  • ssp. rhea (Groum-Grshimailo, 1891); original combination: Argynnis eugenia var. rhea

    • syn. fulgens (Bang-Haas, 1927)

  • ssp. pulchella Huang, 2001; original combination: Kuekenthaliella eugenia pulchella

  • ssp. tibetana Huang, 1998; original combination: Kuekenthaliella eugenia tibetana

  • ssp. vega (Christoph, 1889); original combination: Argynnis vega

• gemmata (Butler, 1881)

  • ssp. gemmata (Butler, 1881); original combination: Argynnis gemmata. Type loc.: Darjeeling, India. Types: NHMUK. Refs.: Butler (1881): 32.

  • ssp. wui Huang, 1998; original combination: Kuekenthaliella gemmata wui

• hanningtoni (Elwes, 1889)

  • ssp. hanningtoni (Elwes, 1889); original combination: Argynnis hanningtoni

  • ssp. imatonga (Riley, 1932); original combination: Argynnis hanningtoni imatonga

  • ssp. jeanneli Bernardi, 1968; original combination: Issoria hanningtoni jeanneli

• lathonia (Linnaeus, 1758)

  • ssp. isaea (Gray, 1846); original combination: Argynnis isaea

  • ssp. isaeoides Reuss, 1925; original combination: Rathora isaeae f. geogr. isaeoides

  • ssp. lathonia (Linnaeus, 1758); original combination: Papilio lathonia Refs.: Simonsen (2006 b) (taxonomy).

    • syn. valdensis (Esper [1804])

  • ssp. messoa (Fruhstorfer, 1912); original combination: Argynnis lathonia messoa

• mackinnonii (de Nicéville, 1891); original combination: Argynnis mackinnonii

• smaragdifera (Butler, 1895)

  • ssp. reducta Carcasson, 1961; original combination: Issoria smaragdifera reducta

  • ssp. smaragdifera (Butler, 1895); original combination: Argynnis smaragdifera

Genus Boloria

Genus: Boloria Moore, 1900

Type species: Boloria pales ([Denis & Schiffermüller], 1775); original combination: Papilio pales [Denis & Schiffermüller], 1775

Synonyms: None

References: Moore (1900): 243; Simonsen (2005) (phylogeny); Pelham (2008); Simonsen et al. (2010) (phylogeny)

Subgenus Boloria

Subgenus: Boloria Moore, 1900

Type species: Boloria pales ([Denis & Schiffermüller], 1775); original combination: Papilio pales [Denis & Schiffermüller], 1775

Synonyms:

• Smoljana Slivov, 1995. Type species: Boloria graeca (Staudinger, 1870); original combination: Boloria (Smoljana) rhodopensis Slivov, 1995. Refs.: Slivov (1995): 62

References: Warren (1944); Simonsen (2005); Simonsen et al. (2010)

13 species:

• alaskensis (Holland, 1900)

  • ssp. alaskensis (Holland, 1900); original combination: B[renthis] pales var. alaskensis. Type loc.: “mountains between Forty-Mile and Mission Creeks” [Alaska]. Types: CMNH. Refs.: Holland (1900) Entomol. News 11(3): 383, no. 6; Crosson du Cormier (1977), Alexanor 10(1): 35-47, detailed the separation of this species from B. napaea (Hoffmansegg, 1804) in: Illiger, Mag. f. Insektenk. 3(3): 196. This arrangement was corroborated by Dubatolov (1992), Vestnik Zoologii 1992(6): 40-41, Kosterin (2000), Far East Entomol. 86(1): 1-10, and Gorbunov (2001), Butterflies Russia Class.: 201-202.

    • syn. reiffi Reuss, 1925. Type loc.: “Britisch-Columbien” [British Columbia]. Types: syntypes in Reuss collection. Refs.: Reuss (1925) Int. Entomol. Z. 19(36): cols. 279-280; Pike (1981), J. Res. Lepid. 18(3): 157, suggested that this may be a subspecies-level taxon.

    • syn. nearctica Verity, 1932. Type loc.: “north-eastern Alaska (69o 40’ N. Lat. and 141oW. Long.)”. Types: type location unknown. Refs.: Verity (1932) Dtsche. Entomol. Z. “Iris” 46(3): 104, 108; cited Lehmann (1913), in: Seitz, Grossschmett. Erde 5(173): 424.

  • ssp. bato Churkin, 1999; original combination: Boloria alaskensis bato

  • ssp. halli Klots, 1940; original combination: Boloria pales halli. Type loc.: “vicinity of Green River Pass, Wind River Range, Sublette Co., Wyoming”. Types: AMNH. Refs.: Klots (1940).

  • ssp. nikolajewski (Heydemann, 1920); original combination: Argynnis arsilache nikolajewski

  • ssp. sedykhi Crosson du Cormier, 1977; original combination: Boloria alaskensis sedykhi

• aquilonaris (Stichel, 1980)

  • ssp. aquilonaris (Stichel, 1980); original combination: Argynnis pales aquilonaris

  • ssp. huerteri Churkin & Bogdanov, 2003; original combination: Boloria aquilonaris huerteri

  • ssp. sima Churkin, 2000; original combination: Boloria aquilonaris sima

• banghaasi Seitz, 1909

  • ssp. banghaasi Seitz, 1909; original combination: Boloria pales f. banghaasi

  • ssp. infans Churkin, 2000; original combination: Boloria banghaasi infans Refs.: Churkin, 2000, Helios 1: 109.

  • ssp. jakubovi Kosterin & Gorbunov, 2007; original combination: Boloria aquilonaris jakubovi Refs.: Kosterin & Gorbunov, 2007, Butt. North. Asia in nat. 2: 197.

  • ssp. neopales (Nakahara, 1926); original combination: Argynnis pales neopales

• caucasica (Lederer, 1852)

  • ssp. caucasica (Lederer, 1852); original combination: Argynnis arsilache v. caucasica

  • ssp. petrovi Churkin, 2002; original combination: Boloria caucasica petrovi. Type loc.: Daghestan. Refs.: Churkin, 2002; Helios 3: 109.

• frigidalis Warren, 1944; original combination: Boloria napaea altaica f. frigidalis

• generator (Staudinger, 1886)

  • ssp. generator (Staudinger, 1886); original combination: Argynnis pales var. generator

    • syn. juldussica (Wagner, 1913)

    • syn. extrema (Wnukowsky, 1927)

  • ssp. klimenkoi Churkin & Zhdanko, 2001; original combination: Boloria sipora klimenkoi

• graeca (Staudinger, 1870)

  • ssp. balcanica (Rebel, 1903); original combination: Argynnis pales var. balcanica

    • syn. rhodopensis Slivov, 1955

  • ssp. brogotarus (Fruhstorfer, 1909); original combination: Argynnis pales brogotarus

  • ssp. graeca (Staudinger, 1870); original combination: Argynnis pales var. graeca

  • ssp. karina Oorschot & Wagener, 1990; original combination: Boloria graeca karina

• napaea (Hoffmannsegg, 1804)

  • ssp. altaica (Groum-Grshimailo, 1893); original combination: Argynnis pales var. altaica

    • syn. isisoides (Heydemann, 1920)

  • ssp. contaminata Gorbunov, 2007; original combination: Boloria napaea contaminata

  • ssp. napaea (Hoffmannsegg, 1804); original combination: Papilio napaea

  • ssp. pustagi Korshunov & Ivonin, 1955; original combination: Boloria altaica pustagi

  • ssp. shanxiana Murayama, 1983; original combination: Boloria napaea shanxiana

  • ssp. vinokurovi Dubatolov, 1992; original combination: Boloria napaea vinokurovi

• pales ([Denis & Schiffermüller], 1775)

  • ssp. eupales (Fruhstorfer, 1903); original combination: Argynnis pales eupales

  • ssp. nirvana Kocman, 1999; original combination: Boloria pales nirvana

  • ssp. pales ([Denis & Schiffermüller], 1775); original combination: Papilio pales

  • ssp. palina (Fruhstorfer, 1903); original combination: Argynnis pales palina

    • syn. palinoides Reuss, 1925

  • ssp. palustris (Fruhstorfer, 1909); original combination: Argynnis pales palustris

  • ssp. pyrenesmiscens Verity, 1932; original combination: Boloria pales r. pyrenesmiscens

  • ssp. rilaensis Varga, 1971; original combination: Boloria pales rilaensis

  • ssp. shambhala Kocman, 1999; original combination: Boloria pales shambhala

  • ssp. yangi Hsu & Yen, 1997; original combination: Boloria pales yangi

• purpurea Churkin, 1999; original combination: Boloria purpurea. Type loc.: “Buryatia, Barguzin Mts., Nesterikha River, Kedrovoe Lake, 1900-2000m”. Refs.: Churkin, 1999; Atalanta 29 (1/4): 118.

• roddi Kosterin, 2000; original combination: Boloria pales roddi

  • syn. sanguinea Churkin & Bogdanov, 2002; original combination: Boloria sanguinea.

• sifanica (Grum-Grshimailo, 1891); original combination: Argynnis pales var. sifanica

  • syn. mingi Kocman, 1995; original combination: Boloria pales mingi.

• sipora (Moore [1875])

  • ssp. korla (Fruhstorfer, 1904); original combination: Argynnis pales korla

  • ssp. sipora (Moore [1875]); original combination: Argynnis sipora. Type loc.: Cashmere, near Gungabul Lake, at foot of Haramook Peak. Refs.: Moore ([1875]): 568.

Subgenus Clossiana

Subgenus: Clossiana Reuss, 1920

Type species: Boloria selene ([Denis & Schiffermüller], 1775); original combination: Papilio selene [Denis & Schiffermüller], 1775

Synonyms: None

References: Reuss (1920): 192 (footnote); Reuss (1921 b): 190

26 species:

• alberta (Edwards, 1890); original combination: Argynnis Alberta. Type loc.: “Laggan, Alberta.strictly above timberline, and not very near it either, though not at the highest points in the mountains”; defined as “vicinity of.Lake Louise, Banff National Park, Alberta” by Kondla (1996), Alberta Naturalist 26(2): 40. Types: Lectotype in CMNH, designated by F. Brown (1965), Trans. Am. Entomol. Soc. 91(3): 339, fig. 35 ♂ D&V; figured by W. Holland (1898), Butterfly Book: pl. 15, fig. 15 ♂ D. Refs.: Edwards, 1890, Can. Entomol. 22(6): 113-114; figured by W. H. Edwards (1894), Butterflies N. Am. 3(15): pl. Argynnis 7, figs. 5 ♂ D, 6 ♂ V, 7 ♀ D, 8 ♀ V.

  • syn. banffensis (Gunder, 1932); original combination: Brenthis alberta tr. f. banffensis. Type loc.: “Banff, Alberta, Canada”. Types: “Holotype” in AMNH. Refs.: Gunder, 1932, Pan-Pac. Entomol. 8(3): 125.

• angarensis (Erschoff, 1870)

  • ssp. alticoa (Sushkin & Tshetverikov, 1907); original combination: Argynnis angarensis alticola

    • syn. shelkovnikovi Korb & Yakovlev, 1997

  • ssp. angarensis (Erschoff, 1870); original combination: Argynnis angarensis

  • ssp. hakutozana (Matsumura, 1927); original combination: Argynnis hakutozana

  • ssp. herzi (Wnukowsky, 1927); original combination: Argynnis (Brentis[sic]) angarensis herzi

  • ssp. miyakei (Matsumura, 1919); original combination: Argynnis amathusia var. miyakei

  • ssp. samkoi (Sheljuzhko, 1931); original combination: Brenthis selenis samkoi

  • ssp. sedychi (Weiss, 1964); original combination: Clossiana angarensis sedychi

• astarte (Doubleday [1847])

  • ssp. astarte (Doubleday [1847]); original combination: Melitaea Astarte. Type loc.: “Rocky Mountains, North America”; defined as “Rock Lake, Alberta (53o 27’N, 118o 16’W)” by J. Shepard (1984), Quaest. Entomol. 20(1): 40-41. Types: Syntype (#45-136) in NHMUK. Refs.: Doubleday, 1847, In: Gen. diurn. Lepid. (1)(12): pl. 23, fig. 5; (1)(21): 181, footnote (1848), name corrected to Argynnis astarte; C. Guppy and J. Shepard (2001), Butterflies B. C.: 295-296, suggested that astarte is conspecific with B. tritonia (Böber, 1812), Mém. Soc. Moscou 3: 20-21, no. 6, pl. 1, figs. 1 ♂ D, 2 ♂ V; index: 320, an arrangement adopted by Gorbunov (2001), Butterflies Russia Class.: 195-196. However, Pike (1981), J. Res. Lepid. 18(3): 162, and Bogdanov (2009), Proc. State Darwin Mus. 13: 68-99, among others, have questioned whether distincta, and by implication tschukotkensis, should be considered subspecies of B. astarte.

    • syn. victoria (Edwards, 1891). Type loc.: “Laggan, Alberta”; defined as “two mountains near Laggan, one of them three miles southwest, in the main range, the other a low smooth mountain directly north of Laggan” by W. H. Edwards (1894), Butterflies N. Am. 3(15): [115-118]; subsequently interpreted to be “outliers of Mt. Temple and Mt. Richardson” by F. Brown (1965), Trans. Am. Entomol. Soc. 91(3): 341. Types: Holotype in CMNH, figured by F. Brown (1965), Trans. Am. Entomol. Soc. 91(3): 340, fig. 36 ♂ D&V; figured by W. Holland (1898), Butterfly Book: pl. 18, fig. 15 ♂ V. Refs.: Edwards, 1891, Can. Entomol. 23(9): 198-199; figured by W. H. Edwards (1894), Butterflies N. Am. 3(15): pl. Argynnis 7, figs. 1 ♂ D, 2 ♂ V, 3 ♀ D, 4 ♀ V (as astarte).

  • ssp. alaskia Grishin, 2024; original combination: Boloria astarte alaskia. Type loc.: USA: Alaska, Seward Peninsula, Darby Mts., Omilak. Types: FLMNH. Distr.: From the Seward Peninsula to the Brooks Range in Alaska, USA. Refs.: Zhang et al. (2024): 8 (taxonomy, genome). Ety.: Alaska

  • ssp. distincta (Gibson, 1920); original combination: Brenthis distincta. Type loc.: “Harrington Creek, Yukon Territory, lat. 65o 05’”. Types: CNC. Refs.: Gibson, 1920, Rept. Can. Arct. Exped. 3(i): 25i, 54i, pl. 4, fig. 12 ♀ V (holotype).

  • ssp. tschukotkensis (Wyatt, 1961); original combination: Clossiana distincta tschukotkensis. Type loc.: “Berge von Tschukotka, N. O. Sibirien, Geröll-zone” [Magadan Oblast, Chukotka Autonomous Okrug, Russia]. Types: LNKD. Refs.: Wyatt, 1961, Zeit. Weiner Entomol. Ges. 4672: 99, pl. 5, fig. 3 ♀ D, 4 ♀ V (holotype).

    • syn. ershovi (Korshunov & Gorbunov, 1995). Type loc.: “Khaptagai on the Lena River” [Sakha Republic, Russia]. Types: SZMN. Refs.: Korshunov & Gorbunov, 1995, Butterflies Asiatic Russia: 108, under no. 211.

    • syn. suntara (Korshunov, 1998). Type loc.: “Suntar-Khayata Range, 180 km ESE Khandyga, upper Eastern Khandyga River, 233 km on the Khandyga to Magadan line, mountains at 1300-1500 m, Yakutia” [Sakha Republic, Russia]. Types: Syntypes in SZMN. Refs.: Korshunov, 1998, New descr. specif. Butterflies Asiatic Russia 21-22.

• bellona (Fabricius, 1775)

  • ssp. bellona (Fabricius, 1775); original combination: P[apilio]. N[ymphalis]. P[halerata]. Bellona. Type loc.: “America boreali”. Types: Syntype in ZMUC. Refs.: Fabricius, 1775, Syst. Entomol.: 517-518, no. 317.

    • syn. fasciata (Cockerell, 1889). Type loc.: Not stated; “Massachusetts” implied from reference to Maynard. Types: “Type(s)” probably lost. The text clearly indicates that this taxon was described as an individual variant. Refs.: Cockerell, 1889, Entomol. 22(311): 99; cited Maynard (1886), Butterflies New England: 27-28 (in part), under no. 31.

    • syn. kleenei (Watson, 1921). Type loc.: “West Hartford, Connecticut” [Hartford County]. Types: “Holotype” in AMNH. Refs.: Watson, 1921, J. N. Y. Entomol. Soc. 28(3/4): 171-172.

    • syn. pardopsis (Holland, 1928). Type loc.: “Crafton, Allegheny County, Pa” [Pennsylvania]. Types: “Holotype” in CMNH. Refs.: Holland (1928): 45; figured by W. Holland (1931), Butterfly Book (rev. ed.): pl. 55, fig. 26 ♀ D (“holotype”).

    • syn. ammiralis (Hemming, 1933), Proposed to replace Papilio bellona Fabricius, 1775, erroneously thought preoccupied; a junior objective synonym of bellona. Refs.: Hemming, 1933, Entomol. 66(847): 276, no. 7.

  • ssp. jenistae Stallings & Turner, 1947; original combination: Boloria toddi jenistai. Type loc.: “Lloydminster, Sask”; suggested to be “the vicinity of Rivercourse, Alberta” by Kondla (1996), Alberta Naturalist 26(2): 39. Types: NMNH. Refs.: Stallings & Turner, 1947, Can. Entomol. 78(7/8): 135-136.

  • ssp. toddi (Holland, 1928); original combination: B[renthis]. bellona toddi. Type loc.: St. Margarets River.near Sept-Iles along a river that was called Riviere Sainte-Marguerite; Quebec” This river was dammed near its confluence with the St. Laurence River, and the resulting reservoir is called Lac-Walker. Types: Holotype (#5269) in CMNH. Refs.: Holland (1928): 45; figured by W. Holland (1931), Butterfly Book (rev. ed.): pl. 55, fig. 14 ♂ D (holotype), 15 ♀ D (paratype).

• chariclea (Schneider, 1794)

  • ssp. arctica (Zetterstedt [1839]); original combination: A[rgynnis]. arctica. Type loc.: “Groenlandia”. Types: Location of type(s) not known, possibly lost. Refs.: Zetterstedt, [1839], Ins. Lappon. (5): col. 899, no. 18.

    • syn. tullia (Fabricius, 1780), preocc. by Papilio tullia Müller, 1764, Fauna Ins. Fridrichsdal.: 36. Type loc.: “Groenl. Sullernek.Kugluiak” [Greenland]. Types: Location of type(s) not known, possibly lost. Refs.: Fabricius, 1780, Fauna Groenl.: 192-193, no. 143.

    • syn. obscurata (M’Lachlan, 1878). Type loc.: “81°41’N”; holotype from “Grinnell Land, W. side of Smith Sound 81o45’ N” [Ellesmere Island, Nunavut]. Types: NHMUK. Refs.: M’Lachlan, 1878, J. Linn. Soc. Lond., Zool. 14(74): 110.

  • ssp. boisduvalii (Duponchel [1832]); original combination: Argynnis Boisduvalii. Type loc.: “les contrées les plus septentrionales de l’Europe”; suggested to be “probably Labrador” by Klots (1951), Field Guide Butterflies N. Am.: 90. Types: Type(s) may be in MNHN or NHMUK. Refs.: Duponchel, [1832], In: Godart, Hist. Nat. Lepid. 1(Diurnes), Suppl. 1(6): 127-128, no. 45, pl. 20, no. 4 ♂ V. “Both Duponchel [1832] ibid., and Boisduval [1833], Icon. Hist. Lépid. Europ. 1(10): 98, pl. 20, figs. 5 ♂ D, 6 ♂ V, cited “M. Sommer” as the source of the specimens described. Sommer has been noted as a source of many butterflies from Labrador (e.g., Boisduval and Le Conte [1830], Hist. Lépid. Amér. sept. (8): 66-67, Staudinger (1857), Stettiner Entomol. Ztg. 18(7-9): 300-301, and Hemming (1937), Hübner 2: 123). Scudder (1868), Proc. Boston Soc. Nat. Hist. 11: 33, and Aurivillius (1889), Bih. K. svenska VetenskAkad. Handl. 15(4)(1): 9-10 , fig. 1 VHW, referred to samples from Labrador as boisduvalii. Examination of the type(s), if extant, is needed; if there are no syntypes, the designation of a neotype may be required”.

    • syn. labradorensis (Gunder, 1928). Type loc.: “Labrador” [Newfoundland and Labrador]. Types: “holotype” in NMNH. Refs.: Gunder, 1928, Can. Entomol. 60(7): 163, pl. A, figs. 5, 5a (“holotype”).

    • syn. oenone (Scudder, 1875), nom. nud. Refs.: Scudder, 1875, Proc. Boston Soc. Nat. Hist. 17(3): [297]-299 (listed as a synonymy of chariclea Schneider, 1794 without a description or indication).

  • ssp. butleri (Edwards, 1883); original combination: Argynnis Butleri. Type loc.: “Cape Thomson, North-west America.Kotzebue Sound”; lectotype from “Cape Thompson” [Alaska]. Distr.: Lectotype in CMNH designated by F. Brown (1965), Trans. Am. Entomol. Soc. 91(3): 346, fig. 38 ♂ D&V; figured by W. Holland (1931), Butterfly Book (rev. ed.): pl. 55, fig. 18 ♂ V. Refs.: Edwards, 1883, Can. Entomol. 15(2): 32-33.

    • syn. tshuktsha (Korshunov, 1998). Type loc.: “mouth of the Chaun River, 100 km south of Pevek, Chukotka” [Magadan Oblast, Chukotka Autonomous Okrug, Russia]. Types: SZMN. Refs.: Korshunov, 1998, New descr. specif. Butterflies Asiatic Russia: 21.

  • ssp. chariclea (Schneider, 1794); original combination: Papilio Chariclea. Type loc.: “Lappland” implied from the title of paper. Types: Location of type(s) not known, probably lost. Refs.: Schneider, 1794, Neu. Mag. f. Leibhaber Entomol. 5: 588-589; Pike (1981), J. Res. Lepid. 18(3): 156, and J. Shepard (1998), Syst. W. N. Am. Butterflies (59): 727-730. It is probable that this assemblage actually consists of two species.

  • ssp. grandis (Barnes & McDunnough, 1916); original combination: Brenthis chariclea grandis. Type loc.: “Hymers, Ont” [Ontario]. Types: syntypes in NMNH; figured by W. Holland (1931), Butterfly Book (rev. ed.): pl. 60, figs. 7 ♂ D, 8 ♀ V. Refs.: Barnes & McDunnough, 1916, Can. Entomol. 48(7): 223.

  • ssp. helena (Edwards, 1871); original combination: Argynnis Helena. Type loc.: “Colorado”; suggested to be “region of the range north and west of Fairplay and about Twin Lakes”, probably “Mt. Elbert, Mosquito Pass, Mt. Lincoln and Hoosier Pass” by F. Brown (1934), J. N. Y. Entomol. Soc. 42(2): 160; defined as “Mosquito Pass, Lake-Park Counties, Colorado” by Brown (1965): 342-343. Types: Lectotype in CMNH, designated by F. Brown (1965), Trans. Am. Entomol. Soc. 91(3): 344, fig. 37 ♂ D&V. Refs.: Edwards, 1871, Trans. Am. Entomol. Soc. 3(3/4): 266, 268, no. 3.

    • syn. sangredecristo Scott, 1981. Type loc.: “Hermit Pass, Sangre de Cristo Mts., Custer Co., Colorado”. Types: LACM. Refs.: Scott, 1981, Papilio (n.s.) 1: 5.

  • ssp. ingens (Barnes & McDunnough, 1918); original combination: Brenthis helena ingens. Type loc.: “Yellowstone Park” [Wyoming]. Types: NMNH. Refs.: Barnes & McDunnough, 1918, Contr. Nat. Hist. Lepid. N. Am. 4(2): 71, 180, pl. 11, figs. 5 ♂ D (holotype), 6 ♂ V.

    • syn. martini (Gunder, 1934). Type loc.: “Teton trail, Teton Mts., Teton County, Wyoming (just south of the ingens type locality)”. Types: “Holotype” in AMNH. Refs.: Gunder, 1934, Can. Entomol. 66(6): 127-128.

  • ssp. montinus (Scudder, 1863); original combination: Argynnis Montinus. Type loc.: “the lower half of barren summits of the White Mountains, N[ew]. H[ampshire]”; defined as “summit of Mount Madison, the head of Tuckerman’s Ravine, and a short distance above the Ledge, (all, it will be noticed, within the subalpine region)” by Scudder (1863), Boston J. Nat. Hist. 7(4): 618, 628. Types: Syntypes probably in MCZ. Refs.: Scudder, 1863, Proc. Essex Inst. 3: 166, no. 32; redescribed and figured by Scudder (1863), Boston J. Nat. Hist. 7(4): 626-628, pl. 14, fig. 1 ♂ D&V.

  • ssp. rainieri (Barnes & McDunnough, 1913); original combination: B[renthis]. chariclea rainieri. Type loc.: “Mt. Ranier [sic!], Wash.(6-7000 ft)”; the figure captions added “Paradise Valley” [Mt. Rainier, Pierce County]. Types: Syntype in NMNH and LACM; figured by W. Holland (1931), Butterfly Book (rev. ed.): pl. 60, figs. 9 ♂ D, 10 ♀ V. Refs.: Barnes & McDunnough, 1913, Contr. Nat. Hist. Lepid. N. Am. 2(3): 96-97, pl. 2, figs. 1 ♂ D, 2 ♀ D, 3 ♂ V, 4 ♀ V; “C. Guppy and Kondla (2002), J. Lepid. Soc. 56(3): 181, suggested that this name be emended to “ranieri” since “Mt. Ranier” was misspelled in the text in the original description. The name as given in the description heading was derived from the correct spelling of the name of the mountain, and needs no emendation (Code Article 32.5.1)“.

• dia (Linnaeus, 1767)

  • ssp. alpina (Elwes, 1899); original combination: Argynnis dia var. alpina

  • ssp. calida (Jachontov, 1911); original combination: Argynnis dia var. calida

  • ssp. dia (Linnaeus, 1767); original combination: Papilio dia

  • ssp. disconota (Krulikovsky, 1909); original combination: Argynnis dia var. disconota

  • ssp. nigrescens (Schavrov, 1886); original combination: Argynnis dia nigrescens

  • ssp. semota (Tuzov, 1999); original combination: Clossiana dia semota

  • ssp. setania (Fruhstorfer, 1909); original combination: Argynnis dia setania

• epithore (Edwards, 1864)

  • ssp. chermocki Perkins & Perkins, 1966; original combination: Boloria epithore chermocki. Type loc.: “2.9 miles E of Dolph, Yamhill Co., Oregon”. Types: LACM. Refs.: Perkins & Perkins, 1966, J. Lepid. Soc. 20(2): 109-112, pl. 1, figs. 9 ♂ D, 10 ♂ V (holotype); pl. 2 figs. 11 ♀ D, 12 ♀ V.

    • syn. uslui Koçak, 1984, Proposed to replace Boloria epithore borealis Perkins, 1973, with the same data (Code Article 60.3) Refs.: Koçak, 1984, Priamus 3(3): 96.

    • syn. borealis Perkins, 1973, preocc. by Argynnis thore borealis Staudinger, 1861, Stettiner Entomol. Ztg. 22(10-12): 351. Type loc.: “Shingle Creek Road, Keremeos, British Columbia”. Types: CNC. Refs.: Perkins, 1973, In: E. Perkins and Meyer, Bull. Allyn Mus. (11): 6-8, pl. 1, figs. 13 ♂ D (holotype); 14 ♂ V, pl. 2, figs. 15 ♀ D, 16 ♀ V.

    • syn. obscuripennis (Gunder, 1926). Type loc.: “Chilcolin [sic], British Columbia, Canada” [Chilcotin]. Types: “Holotype” in AMNH. Refs.: Gunder, 1926, Entomol. News 37(1): 7, no. 9, pl. 1, fig. 9 ♀ D (holotype).

  • ssp. epithore (Edwards, 1864); original combination: Argynnis Epithore Boisduval in lit. Type loc.: “California”; neotype from “Saratoga, Santa Clara County, California”. Types: Neotype in CMNH, designated by Klots (1965), in: F. Brown, Trans. Am. Entomol. Soc. 91(3): 335, fig. 34a ♂ D&V. Refs.: Edwards, 1864, Proc. Entomol. Soc. Philad. 2(4): 501, 503-504, no. 5.

    • syn. epithore (Boisduval, 1869), preocc. Type loc.: “les hautes montagnes de l’est”; defined as “vic. of Santa Cruz Mtns., Santa Cruz County, California” by E. Perkins and Meyer (1973), Bull. Allyn Mus. (11): 4. Types: Lectotype in CMNH, designated by E. Perkins and Meyer (1973), ibid.; figured by F. Brown (1965), Trans. Am. Entomol. Soc. 91(3): 335, fig. 34b ♂ D&V. Refs.: Boisduval, 1869, Annls. Soc. Entomol. Belg. 12: 58, no. 50.

  • ssp. sierra Perkins, 1973; original combination: Boloria epithore sierra. Type loc.: “Sentinel Dome, Yosemite National Park, Mariposa County, California”. Types: FLMNH. Refs.: Perkins, 1973, In: E. Perkins and Meyer, Bull. Allyn Mus. (11): 10-11, pl. 1, figs. 1 ♂ D (holotype), 2 ♂ V, pl. 2, figs. 3 ♀ D, 4 ♀ V.

    • syn. eldorado (Strand, 1915)

    • syn. wawonae (Gunder, 1924)

  • ssp. sigridae (Shepard, 2001); original combination: Clossiana epithore sigridae. Type loc.: “BC, St. Elias Mts., Tats Lake, 770 m” [British Columbia]. Types: RBCM. Refs.: Shepard, 2001, Butterflies B. C.: 289-290, figs. ♂ D&V (holotype), ♀ D&V.

• erda (Christoph, 1892)

  • ssp. dulkeiti (Kurenzov, 1970); original combination: Brenthis dulkeiti

  • ssp. erda (Christoph, 1892); original combination: Argynnis erda

  • ssp. kitoica (Belik, 1996); original combination: Clossiana erda kitoica

  • ssp. puella (Churkin, 1999); original combination: Clossiana erda puella

• erubescens (Staudinger, 1901)

  • ssp. ancilla (Churkin, 2002); original combination: Clossiana hegemone ancilla

  • ssp. chotana (Bang-Haas, 1915); original combination: Argynnis hegemone var. chotana

    • syn. franciscana (Verhulst, 1999)

    • syn. ali (Churkin, 2002)

  • ssp. erubescens (Staudinger, 1901); original combination: Argynnis hegemone var. erubescens

  • ssp. eximia (Churkin, 2002); original combination: Clossiana hegemone eximia

  • ssp. haberhaueri Hemming, 1933; original combination: Boloria haberhaueri

    • syn. hegemone (Staudinger, 1881)

    • syn. hegemone (Alphéraky, 1881)

  • ssp. houri Wyatt, 1961; original combination: Boloria hegemone houri

  • ssp. panterai Korb, 2005; original combination: Boloria erubescens panterai

  • ssp. tienschanica (Wagner, 1913); original combination: Argynnis hegemone tienschanica

  • ssp. tuzovi (Churkin, 2002); original combination: Clossiana hegemone tuzovi

• euphrosyne (Linnaeus, 1758)

  • ssp. anka (Fruhstorfer, 1907); original combination: Argynnis euphrosyne anka. Type loc.: Caucasus.

  • ssp. dagestanica (Sovinsky, 1905); original combination: Argynnis euphrosyne var. dagestanica

  • ssp. densoi (Fruhstorfer, 1909); original combination: Argynnis euphrosyne densoi

  • ssp. euphrosyne (Linnaeus, 1758); original combination: Papilio euphrosyne

  • ssp. fingal (Herbst, 1800); original combination: Papilio euphrosyne fingal

    • syn. lapponica (Esper, 1798)

  • ssp. kamtschadalus (Seitz, 1908); original combination: Argynnis euphrosyne kamtschadalis

  • ssp. nephele (Herrich-Schaeffer [1847]); original combination: Argynnis euphrosyne nephele

  • ssp. orphanoides (Huang & Murayama, 1992); original combination: Clossiana euphrosyne orphanoides

  • ssp. oprhanus (Fruhstorfer, 1907); original combination: Argynnis euphrosyne orphanus

  • ssp. rusalka (Fruhstorfer, 1909); original combination: Argynnis euphrosyne rusalka

  • ssp. umbra (Seitz, 1909); original combination: Argynnis euphrosyne f. umbra

• freija (Thunberg, 1791)

  • ssp. asahidakeana (Matsumura, 1926); original combination: Argynnis asahidakeana

  • ssp. browni (Higgins, 1953); original combination: Clossiana freija browni. Type loc.: “Independance Pass, Pitkin Co” [Colorado]. Types: AMNH. Refs.: Higgins, 1953, Entomol. 86(9): 210, no. 9.

  • ssp. freija (Thunberg, 1791); original combination: Papilio Freija P[apilio]. n[ymphalis]. ph[aleratus].. Type loc.: “Vestrobothnia juxta flumina locis locios graminosis.in Lapponia”; lectotype labeled “Lapp. Liljeblad” [western Gulf of Bothnia and Lappland, Sweden]. Types: Lectotype in UZIU, designated by Karsholt and Nielsen (1986), Entomol. Scand. 16(4): 446. Refs.: Thunberg, 1791 (1789), Entomol. Sist. Ins. Svec. (2): 34-36, pl. 2, figs. 14, [14a] ♂ D&V.

    • syn. lapponica (Esper, 1789). Type loc.: “Der lapplandisch P. Dia” [Lappland]. Types: Type(s) location not known, probably lost. Refs.: Esper, 1789, Die Schmett. 1(suppl.)(1)(3/4): 21-22, pl. 97 (Cont. 52), fig. 3 ♂ D&V.

    • syn. freya (Godart, 1819), Proposed to replace Papilio freija Thunberg, 1791, erroneously thought to be preoccupied, or possibly an unjustified emendation Refs.: Godart, 1819, In: Enc. méth. 9(Ins.)(1): (254), 273, no. 37; cited many references to freija.

    • syn. jakutensis (Wnukowsky, 1927). Type loc.: “Siberia septentrionalis-orientalis (Jakutsk)” [Yakutsk, Sakha Autonomous Republic, Russia]. Types: Syntypes in UTR. Refs.: Wnukowsky, 1927, Mitt. Münchn. Entomol. Ges. 17(7-12): 70, no. 4.

    • syn. zamolodchikova (Churkin, 2001). Type loc.: “Russia, East Chukotka, 13 km SE Ioni L., Gilmimleviem R” [Magadan Oblast, Chukotka Autonomous Okrug, Russia]. Types: Holotype in DSM. Refs.: Churkin, 2001, Helios 2: 99, pl. 8, figs. 10 ♂ D (holotype), 11 ♂ V, 12 ♀ D.

    • syn. gunderi (Harper, 1933). Type loc.: “McCreary, Manitoba, Canada”. Types: “Holotype” in CAS. Refs.: Harper, 1933, Pan.-Pac. Entomol. 9(3): 99.

  • ssp. pallida (Elwes, 1899); original combination: Argynnis freija var. pallida

  • ssp. tarquinius (Curtis, 1835); original combination: [Melitaea] Tarquinius. Type loc.: “Arctic Regions”; determined from itineraries in vol. 1 and a letter from John Ross in vol. 2, Addenda: 732-735, to be the area bounded by 69.30 o-70.9° N lat. and 91.52°-93.0°W long, in the vicinity of the isthmus of Boothia Peninsula, named by the expedition. [Kitikmeot-Keewatin regions, Nunavut]. Types: Syntypes possibly in NHMUK. Refs.: Curtis, 1835, In: Ross, Voy. Search Northwest Pass., Appendix: lxviii-lxix, no. 14.

• frigga (Thunberg, 1791)

  • ssp. alpestris (Elwes, 1899); original combination: Argynnis frigga var. alpestris

  • ssp. annae (Suschkin, 1906); original combination: Argynnis frigga annae. Type loc.: Tarbagatai.

  • ssp. frigga (Thunberg, 1791); original combination: Papilio Frigga P[apilio]. n[ymphalis]. ph[aleratus]. Type loc.: “Lapponiae sylvis”; lectotype from “Sweden: Lappland”. Types: Lectotype in UUZM, designated by Karsholt and Nielsen (1986), Entomol. Scand. 16(4): 446. Refs.: Thunberg, 1791, Entomol. Sist. Ins. Svec. (2): 33-34.

    • syn. maritima (Kardakoff, 1928). Type loc.: “Primorye: Ternei Bay” [Russia]. Types: Location of type(s) not known. Refs.: Kardakoff, 1928, Entomol. Mitt. 17(4): 270. pl. 7, fig. 4.

    • syn. helvenacius Sedykh, 1977. Type loc.: “Polar Urals” [Polar Ural, Komi Republic, Russia]. Types: Syntypes possibly in ZIN. Refs.: Sedykh, 1977, Geogr. Aspect. Conserv. Flora Faun. NE Europ. USSR: 102.

  • ssp. gibsoni (Barnes & Benjamin, 1926); original combination: Brenthis frigga race gibsoni. Type loc.: “Barter Island, Northern Alaska”. Types: Syntypes in CNC; one figured by Gibson (1920), Rept. Can. Arctic Exped. 3(i): 57i, pl. 5, fig. 1 ♂ D. Refs.: Barnes & Benjamin, 1926, Bull. South. Calif. Acad. Sci. 25(3): 92, no. 197d; Layberry, et al. (1998), Butterflies Can.: 178, suggested that this may be a species-level taxon.

    • syn. alaskensis (Lehmann, 1913), Secondarily preoccupied by Brenthis pales alaskensis W. Holland, 1900, Entomol. News 11(3): 383, no. 6. Type loc.: “nördöstlichen Alaska (69° 40’ nördl. Breite, 141° westl. Länge)”. Types: One probable syntype in ZMUC, others may be in SMF. Refs.: Lehmann, 1913, In: Seitz, Grossschmett. Erde 5(173): 424; (178): pl. 87e, fig. [3] ♀ D.

    • syn. lehmanni (Holland, 1928), Proposed to replace Argynnis frigga fa. alaskensis Lehmann, 1913, with the same data (Code Article 60.3). According to Code Article 59.3, this permanently invalidates alaskensis since the name gibsoni has traditionally been used for this taxon. Refs.: Holland (1928): 44.

  • ssp. maritima (Kardakoff, 1928); original combination: Argynnis frigga maritima

  • ssp. saga (Staudinger, 1861); original combination: [Argynnis Frigga] [var.] Saga. Type loc.: “Labrador” [Newfoundland and Labrador]. Types: NHMUK. Refs.: Staudinger, 1861, Stettiner Entomol. Ztg. 22(10/12): 350, under no. 16.

  • ssp. sagata (Barnes & Benjamin, 1923); original combination: Brenthis frigga race sagata. Type loc.: “Hall Valley, Colorado” [Park County]. Types: NMNH. Refs.: Barnes & Benjamin, 1923, Can. Entomol. 55(6): 146.

• gong (Oberthür, 1844)

  • ssp. gong (Oberthür, 1844); original combination: Argynnis gong

    • syn. xizangensis (Huang, 2000)

  • ssp. splendens (Murayama, 1983); original combination: Clossiana gong splendens

• improba (Butler, 1877)

  • ssp. acrocnema Gall & Sperling, 1980; original combination: Boloria acrocnema. Type loc.: “Mt. Uncomphagre, 13.0 km NW of Lake City, Hinsdale Co., Colorado, elev. 4080-4140m”. Types: PMNH. Refs.: Gall & Sperling, 1980, J. Lepid. Soc. 34(2): 231-236, figs. 1&7 ♂ D&V, 13 ♂ D, 16 ♂ V (holotype), 2&8 ♂ D&V, 3&9 ♂ D&V, 4&10 ♀ D&V, 5&11 ♀ D&V, 6&12 ♀ D&V, 19-21 ♂ genitalia.

  • ssp. harryi (Ferris, 1984); original combination: Clossiana improba harryi. Type loc.: “near the summit of Bears Ears Trail, 0.5 mi. (0.8 km) NW of Mt. Chauvenet, 11, 750’ (3583m), Shoshone National Forest, Fremont Co., Wyoming” [Wind River Mountains]. Types: FLMNH. Refs.: Ferris, 1984, Bull. Allyn Mus. (89): 2-5, figs. 2 ♂ D, 3 ♂ V (holotype); 4 ♀D, 5 ♀ V.

  • ssp. improba (Butler, 1877); original combination: Argynnis improba. Type loc.: “between 67 1/2 and 68° [.] Winter Cove and Cambridge Bay” [Nunavut]. Types: Syntype in NHMUK. Refs.: Butler, 1877, Entomol. Mon. Mag. 13(153): 206.

  • ssp. improbula (Bryk, 1920); original combination: Argynnis improba improbula

    • syn. elegans (Bruun & Schantz, 1950)

    • syn. nigrofasciata (Bruun & Schantz, 1950)

    • syn. obscura (Bruun & Schantz, 1950)

    • syn. obsoleta (Bruun & Schantz, 1950)

    • syn. transversata (Bruun & Schantz, 1950)

  • ssp. nunatak Scott, 1998; original combination: Boloria improba nunatak. Type loc.: “Torrens River, Alberta”, presumably from the mountains nearby. Types: AMNH. Refs.: Scott, 1998, Papilio (n.s.) 11: 3-4.

  • ssp. youngi (Holland, 1900); original combination: B[renthis]. youngi. Type loc.: “mountains between Forty-Mile and Mission Creeks, N. E. Alaska”. Types: CMNH. Refs.: Holland, 1900, Entomol. News 11(3): 383-384, no. 9; figured by W. Holland (1931), Butterfly Book (rev. ed.): pl. 55, fig. 28 ♂ V (holotype); Ferris (1984), Bull. Allyn Mus. (89): 1-2, suggested that this taxon represents an individual form, and therefore a synonym of B. i. improba.

• iphigenia (Graeser, 1888)

  • ssp. insularia (Korshunov, 1996); original combination: Clossiana iphigenia insularia

  • ssp. iphigenia (Graeser, 1888); original combination: Argynnis iphigenia

  • ssp. sachalinensis (Matsumura, 1908); original combination: Argynnis sachalinensis Refs.: Matsumura (1908): 158.

• jerdoni (Lang, 1868); original combination: Argynnis jerdoni

  • syn. cashmirensis (Moore, 1874); original combination: Argynnis cashmirensis.

  • syn. chitralensis Moore, 1900; original combination: Boloria chitralensis.

• kriemhild (Strecker, 1879); original combination: Argynnis Kriemhild. Type loc.: “Utah.Arizona.Rio Florida, Colorado”; lectotype from “Utah”. Types: Lectotype in FMNH, designated by Skinner (1914), Entomol. News 25(7): 324; figured by W. Holland (1931), Butterfly Book (rev. ed.): pl. 74, figs. 1 ♂ D, 2 ♂ V. Refs.: Strecker, 1879, In: C. A. H. McCauley, Rept. San Juan Recon., Ind. Exec. Doc. House Rep. 3rd Sess. 45th Congr. 5(1)pt. 2(3), appendix SS: 1854, pl. 1, figs. 5 ♀ D, 6 ♀ V.

  • syn. - (Skinner, 1913); original combination: Argynnis laurenti. Type loc.: “Silver Lake (Brightons), Utah” [Brighton, Salt Lake County]. Types: Syntypes (#7024) in CMNH; one figured by W. Holland (1931), Butterfly Book (rev. ed.): pl. 56, fig. 17 ♂ V. The identity of this taxon was established by W. Barnes and McDunnough (1914), Entomol. News 25(7): 323-324. Refs.: Skinner, 1913, Entomol. News 24(10): 450.

  • syn. - (Reuss, 1923), emend.; original combination: Clossiana lücki. Type loc.: “Utah, Nord-Amerika”. Types: Syntypes in Reuss collection. Refs.: Reuss, 1923, Soc. Entomol. 38(4): 14.

• natazhati (Gibson, 1920)

  • ssp. bankslandia Shepard, 1998; original combination: Boloria natazhati bankslandia. Type loc.: “N.W.T., Victoria Is. 71o17’N, 114oW” [Northwest Territories]. Types: CNC. Refs.: Shepard, 1998, Syst. W. N. Am. Butterflies (60): 732, figs. 5 ♂ D, 6 ♂ V.

  • ssp. nabokovi Stallings & Turner, 1947; original combination: Boloria freija nabokovi. Type loc.: “Alaska Military Highway.Mile 102, North of Summit 2, Ravine, Elevation 6000 ft.British Columbia”; defined as “the Sentinel Range of the Rocky Mts., B. C., at 1830 m elevation” by Troubridge and Wood (1990), J. Lepid. Soc. 44(3): 186; suggested to be “a ravine north of Summit Lake, mile 392 [now km. 621.7] Alaska Highway, British Columbia, Canada” by J. Shepard and Kondla (1993), Pan-Pac. Entomol. 69(3): 273. Types: Holotype (#27642) in MCZ. Refs.: Stallings & Turner, 1947, Can. Entomol. 78(7/8): 134-135.

  • ssp. natazhati (Gibson, 1920); original combination: Brenthis natazhati. Type loc.: “141st meridian, north of mount Natazhat, international boundary survey, elevation 8,600 feet” [southern Yukon Territory]. Types: CNC. Refs.: Gibson, 1920, Rept. Can. Arctic Exped. 3(i): 21i-22i, 52i, 56i, pl. 3, figs. 6 ♂ V (holotype), 12 ♀ D; pl. 5, fig. 6 ♂ D (holotype).

• oscarus (Eversmann, 1844)

  • ssp. australis (Graeser, 1888); original combination: Argynnis oscarus var. australis

    • syn. major (Graeser, 1888)

    • syn. maxima (Fixsen, 1887)

  • ssp. oscaroides (Ménétriés, 1859); original combination: Argynnis oscarus (?) var. oscaroides

  • ssp. oscarus (Eversmann, 1844); original combination: Argynnis oscarus

• perryi (Butler, 1882); original combination: Argynnis perryi Refs.: Butler, 1882; Ann. mag. nat. Hist. (5) 9 (49): 16.

  • syn. dilutior (Fixsen, 1887); original combination: Argynnis selene var. dilutior.

• polaris (Boisduval [1828])

  • ssp. polaris (Boisduval [1828]); original combination: Argynnis Polaris. Type loc.: “Cap Nord”; stated to be “cap Nord, c’est-à-dire l’extrémité de la Laponie norwégienne” by Duponchel [1832], in: Godart, Hist. Nat. Lepid. 1(Diurnes), Suppl. 1(6): 125-126, no. 44. [Finmark, Norway]. Types: Location of type(s) not known, possibly in NHMUK or MNHN. Refs.: Boisduval, [1828], Europ. Lepid. Ind. Meth: 15-16; figured by Boisduval [1833], Icon. Hist. Lépid. Europ. 1(10): pl. 20, figs. 1 ♂ D, 2 ♂ V, and (1836), Spec. gén. Lépid. 1: pl. 11, fig. 1 ♂ D&V.

    • syn. gronlandica ([Skinner], 1892). Type loc.: “Greenland” implied from plate caption; suggested to be “West Coast.McCormick Bay, Herbert Island and Disco.West Coast of Greenland“ by Skinner and Mengel (1892), Proc. Acad. Nat. Sci. Philad. 44(5): 157. Types: Syntype (#7781) in CMNH. Refs.: [Skinner], 1892, Entomol. News 3(3): 49, nos. 9-10, pl. 2, figs. 9 ♀ D, 10 ♀ V (holotype); Masters (1972), Entomol. Rec. & J. Var. 84(7/8): 178, treated this as a subspecies-level taxon.

    • syn. homeyeri (Pagenstecher, 1902). Type loc.: “Gronland”. Types: type location unknown. Refs.: Pagenstecher, 1902, Fauna Arctica 2(2): 236; cited Wocke, 1874, in: Die Zweite Deutsche Nordpolarfahrt 2: 408.

    • syn. americana (Strand, 1905). Type loc.: “Havnen, Rice Strait (Ellesmere Ld.).Havnefjorden (Jones Sd.).Gaasefjorden (Ellesmere Ld.).die Westseite von Gaasefjorden”. Types: Type(s) probably in ZMHU. Refs.: Strand, 1905, Rept. 2nd Norweg. Arctic Exped. “Fram” 1898-1902 1(3): 10-11, no. 3.

    • syn. kurenzovi Wyatt, 1961. Type loc.: “[U.S.S.R]: N.O. Sibirien: [Tschukotka]: Berge von Tschukotka: Geröllzone” [Magadan Oblast, Chukotka Autonomous Okrug, Russia]. Types: LNKD. Refs.: Wyatt, 1961, Z. wien. entomol. Ges. 4672: 98-99, pl. 5, figs. 1 ♀ D, 2 ♀ V.

    • syn. digna (Churkin, 2001). Type loc.: “Russia, Wrangel Is., Neizvestnaya R. (middle stream)” [Chukotka Autonomous Okrug, Russia]. Types: DSM. Refs.: Churkin, 2001, Helios 2: 97-98, pl. 8, figs. 4 ♂ D (holotype), 5 ♂ V, 6 ♀ D.

  • ssp. stellata Masters, 1972; original combination: Boloria Polaris Stellata. Type loc.: “Churchill, Manitoba”. Types: AMNH. Refs.: Masters, 1972, Entomol. Rec. & J. Var. 84(7/8): 177-178, pl. 11, fig. A ♂ D, B ♂ V (holotype), G ♀ D, H ♀ V.

• selene ([Denis & Schiffermüller], 1775)

  • ssp. albequina (Holland, 1928); original combination: B[renthis]. albequina. Type loc.: “White Horse Pass, Yukon Territory”. Types: Syntypes in CMNH. Refs.: Holland, 1928, Ann. Carnegie Mus. 19(1): 40-43, no. 6; figured by W. Holland (1931), Butterfly Book. (rev. ed.): pl. 55, figs. 19 ♂ D, 20 ♀ V, 21 ♂ V; McDunnough (1932), Can. Entomol. 64(12): 269, and Ferris (1971), Sci. Monog. Agric. Exp. Sta., U. Wyo.: 46 (as a synonym of B. s. atrocostalis).

    • syn. baxteri (Holland, 1928). Type loc.: “White Horse Pass, Yukon Territory”. Types: “Holotype” in CMNH.

  • ssp. atrocostalis (Huard, 1927); original combination: Argynnis atrocostalis. Type loc.: Chicoutini, Quebec. Refs.: Huard, 1927; Nat. Can. 54 (6): 131.

    • syn. serratimarginata (Gunder, 1926)

    • syn. jenningsae (Holland, 1928)

  • ssp. chibiana (Matsumura, 1927); original combination: Argynnis selene chibiana. Type loc.: “Chicoutimi” [Quebec]. Types: Location of type(s) not known; McDunnough (1928), Can. Entomol. 60(11): 274, claimed to have examined the “type” so it is presumed to be in CNC. Refs.: Huard, 1927, Nat. Can. 54(6)[(2)34(60]: 131, no. 8.

    • syn. serratimarginata (Gunder, 1926). Type loc.: “Vernon, British Columbia, Canada”. Types: AMNH. Refs.: Gunder, 1926, Entomol. News 37(1): 7, no. 10, pl. 1, fig. 10 ♂ D (“holotype”).

    • syn. jenningsae (Holland, 1928). Type loc.: “Jellicoe, on Thunder Bay, Ontario”. Types: “Holotype” (#5921) in CMNH. Refs.: Holland, 1928, Ann. Carnegie Mus. 19(1): 36, no. 1a; figured by W. Holland (1931), Butterfly Book (rev. ed.): pl. 56, fig. 9 ♂ D.

  • ssp. dilutior (Fixsen, 1887); original combination: Argynnis selene var. dilutior

  • ssp. myrina (Cramer [1777]); original combination: Pap[ilio]. Nymphj[alis]. Phal[erata]. Myrina. Type loc.: “Noord-Amerika by Nieuwjork” “l’Amerique Septentrionale, à la Nouvelle York”; suggested to be “from southeastern New York” by Kohler (1977), J. Lepid. Soc. 31(4): 240. Types: Type(s) probably lost. Refs.: Cramer, [1777], Uitl. Kapellen 2(16): 141, pl. 189, figs. B ♂ D, C ♂ V; (16): 150 (index).

    • syn. myrissa (Godart, 1824), Proposed to replace Papilio myrina Cramer, 1777, erroneously thought preoccupied; a junior objective synonym of myrina Refs.: Godart, [1824], In: Ency. méth. 9(Ins.)(2): (253), 268, 806.

    • syn. marilandica (Clark, 1941). Type loc.: “the boggy pasture on the south side of the road from the Beltsville, M., railway station to the Department of Agriculture experiment farm” [Prince George’s County, Maryland]. Types: Holotype (#55470) in NMNH; figured by A. Clark (1932), Bull. U. S. Nat. Mus. (157): pl. 3. figs. 5 ♂ D, 6 ♂ V, and A. Clark and L. Clark (1951), Smithsonian Misc. Coll. 116(7): frtspc., figs. 1 ♀ V, 2 ♂ D (holotype); pl. 9, figs. B ♂ D, C ♂ V. Refs.: Clark, 1941, J. Wash. Acad. Sci. 31(9): 384.

    • syn. nubes (Scudder, 1889). Type loc.: “Sanborton, N. H” [Belknap County, New Hampshire]. Types: “Holotype” (#29298) in MCZ (The text clearly indicates that this taxon was described as an individual variant.). Refs.: Scudder, 1889, Butterflies E. U. S. & Can. 1(5): 595-596.

    • syn. nivea (Gunder, 1928). Type loc.: “Staten Island, N. Y” [New York]. Types: “Holotype” in AMNH. Refs.: Gunder, 1928, Can. Entomol. 60(7): 163, pl. A, fig. 4 ♂ D (holotype).

    • syn. gunderana (Forbes, 1960), nom. nud. Forbes (1960), attributed this name to Dyar, who never described the taxon. Refs.: Forbes, 1960, Mem. Cornell Univ. Agric. exp. Sta. 371 (part 4): 150, under no. 1.

  • ssp. nebraskensis (Holland, 1928); original combination: B[renthis]. myrina nebraskensis. Type loc.: “Dodge County, Nebraska”. Types: Lectotype in CMNH, designated by W. Holland (1931), Butterfly Book (rev. ed.): 105, pl. 55 fig. 8 ♂ D.

  • ssp. sabulicollis Kohler, 1977; original combination: Boloria selene sabulocollis. Type loc.: “Smith Lake, Sheridan Co., Nebraska”. Types: AMNH. Refs.: Kohler, 1977, J. Lepid. Soc. 31(4): 252-253, figs. 3(a) ♂ D (holotype), 3(b) ♂ V (holotype), 3(c) ♂ D, 3(d) ♂ V, 3(e) ♂ D, 3(f) ♂ V, 6(g) ♀ D, 6(h) ♀ V, 7(a) ♀ D, 7(b) ♀ V, 7(c) ♀ D, 7(d) ♀ V.

  • ssp. selene ([Denis & Schiffermüller], 1775); original combination: P[apilio] Selene. Type loc.: “Wienergegend” from the title of the work. [Vienna, Austria]. Types: Type(s) probably lost. Refs.: [Denis & Schiffermüller], 1775, Ankündung eines syst. Werkes Schmett. Wiener Gegend: 321, no. 11.

    • syn. marphisa (Herbst, 1800). Type loc.: not stated. Types: Type(s) probably lost. Refs.: Herbst, 1800, In: Jablonsky, Natursyst. Ins., Schmett. 10: 105-107, no. 42, pl. 270, figs. 8 ♂ D, 9 ♂ V.

    • syn. rinaldus (Herbst, 1800). Type loc.: “Botnia occidentali” [western Gulf of Bothnia, Sweden]. Types: Type(s) probably lost. Refs.: Herbst, 1800, In: Jablonsky, Natursyst. Ins., Schmett. 10: 108-110, pl. 271, figs. 1 ♂ D, 2 ♂ V.

    • syn. selenia (Freyer, 1847). Type loc.: “Lechstluß”. Types: Holotype probably lost. Refs.: Freyer, 1847, Neuer Beitr. Schmett. Abbild. Natur. 6(83): 21-22, pl. 493, fig. 2 ♂ D&V.

    • syn. hela (Staudinger, 1861). Type loc.: “schwedischen Lapland” [Swedish Lappland]. Types: Type(s) probably in ZMHU. Refs.: Staudinger, 1861, In: Wocke & Staudinger, Stettiner Entomol. Ztg. 22(10-12): 347, under no. 11.

    • syn. obscurum Sedykh, 1977. Type loc.: “Polar Ural Station, on the crest of the Polar Urals; in a meadow amidst mountain dry and marshy tundra” [Komi Republic, Russia]. Types: Syntypes possibly in ZIN. Refs.: Sedykh, 1977, Geogr. Aspect. Conserv. Flora Faun. NE Europ. USSR: 101.

  • ssp. sugitanii (Seok, 1938); original combination: Argynnis selene sugitanii. Type loc.: Korea. Refs.: Seok, 1938; Zephyrus 7: 241.

  • ssp. terraenovae (Holland, 1928), emend.; original combination: B[renthis]. myrina terrae-novae. Type loc.: “Newfoundland”; suggested to be “the eastern part of the island” by Kohler (1977), J. Lepid. Soc. 31(4): 262. [Newfoundland and Labrador]. Types: CMNH. Refs.: Holland, 1928, Ann. Carnegie Mus. 19(1): 36, no. 1b; figured by W. Holland (1931), Butterfly Book (rev. ed.): pl. 55, fig. 13 ♂ D (holotype).

  • ssp. thalia (Hübner, 1790); original combination: Papilio thalia

  • ssp. tollandensis (Barnes & Benjamin, 1925); original combination: Brenthis myrina tollandensis. Type loc.: “Tolland, Moffat [sic!] County, Colorado, altitude approximately 10,000 ft” [Gilpin County]. Types: NMNH. Refs.: Barnes & Benjamin, 1925, Entomol. News 36(2): 44.

• selenis (Eversmann, 1837)

  • ssp. onorensis (Matsumura, 1925); original combination: Argynnis selenis onorensis

  • ssp. selenis (Eversmann, 1837); original combination: Argynnis selenis

  • ssp. sibirica (Erschoff, 1870); original combination: Argynnis selenis var. sibirica

    • syn. festiva (Krulikovsky, 1893)

    • syn. chosensis (Matsumura, 1927)

    • syn. meinhardi (Sheljuzhko, 1929)

    • syn. kononovi (Kurentzov, 1970)

• thore (Hübner [1803-1804])

  • ssp. borealis (Staudinger, 1861); original combination: Argynnis thore var. borealis

  • ssp. excellens (Krulikovsky, 1893); original combination: Argynnis thore var. excellens

    • syn. splendida (Krulikovsky, 1895)

  • ssp. karafutonis (Matsumura, 1919); original combination: Argynnis thore var. karafutonis

  • ssp. herculea (Fruhstorfer, 1907); original combination: Clossiana thore herculea

  • ssp. hypercala (Fruhstorfer, 1907); original combination: Argynnis thore hypercala

    • syn. lacrimosa (Korb, 1999)

  • ssp. hyperlampra (Fruhstorfer, 1907); original combination: Argynnis thore hyperlampra

  • ssp. hyperusia (Fruhstorfer, 1907); original combination: Argynnis thore hyperusia

    • syn. meridionalis (Kardakoff, 1928)

    • syn. brenda (Hemming, 1933)

  • ssp. thore (Hübner [1803-1804]); original combination: Papilio thore

  • ssp. transuralensis (Sheljuzhko, 1931); original combination: Brenthis thore transuralensis

• tritonia (Böber, 1812)

  • ssp. titania (Esper, 1794); original combination: Papilio titania

  • ssp. amphilochus (Ménétriés, 1859); original combination: Melitaea amphilochus

  • ssp. barkalovi (Dubatolov, 2010); original combination: Clossiana tritonia barkalovi

  • ssp. dubatolovi (Korshunov, 1987); original combination: Clossiana distincta dubatolovi

  • ssp. machati (Korshunov, 1987); original combination: Clossiana distincta machati

  • ssp. matveevi (Gorbunov & Korshunov, 1995); original combination: Clossiana matveevi

  • ssp. tritonia (Böber, 1812); original combination: Papilio tritonia. Type loc.: [Lake Baikal, Irkutsk, Russia]. Refs.: Böber, 1812; Mém. Soc. Imp. Nat. Moscou 3: 20.

    • syn. elatus (Staudinger, 1892)

Subgenus Proclossiana

Subgenus: Proclossiana Reuss, 1926

Type species: Proclossiana eunomia (Esper, 1800); original combination: Papilio aphirape Hübner, [1799-1800]

Synonyms: None

References: Reuss (1926 b): 69

1 species:

• eunomia (Esper, 1800)

  • ssp. acidalia (Böber, 1809); original combination: Papilio acidalia Refs.: Böber, 1809, Mém. Soc. Imp. Nat. Moscou 2: 309.

    • syn. asiatica (Staudinger, 1901) Refs.: Staudinger, 1901, Cat. Lep. palaearct. Faunengeb. 1: 34.

    • syn. stromi (Korshunov, 1996)

  • ssp. caelestis (Hemming, 1933), repl. name; original combination: Argynnis aphirape caelestis Refs.: Hemming, 1933, Entomol. 66(847): 275, no. 5.

    • syn. alticola (Barnes & McDunnough, 1913), preocc. by Argynnis alticola Sushkin & Tsetverikov, 1907. Type loc.: “Hall Valley, Colo” [Park County, Colorado]. Types: syntypes in NMNH. Refs.: Barnes & McDunnough, 1913, Contr. Nat. Hist. Lepid. N. Am. 2(3): 97-98.

  • ssp. dawsoni (Barnes & McDunnough, 1916); original combination: Brenthis aphirape dawsoni. Type loc.: “Hymers, Ont” [Ontario]. Types: syntypes in NMNH and LACM. Refs.: Barnes & McDunnough, 1916, Can. Entomol. 48(7): 222-223.

    • syn. harperi (Gunder, 1934). Type loc.: “Churchill, Man., Canada” [Manitoba]. Types: CAS. Refs.: Gunder, 1934, Can. Entomol. 66(6): 127.

  • ssp. denali (Klots, 1940); original combination: Brenthis (Clossiana) aphirape denali. Type loc.: “McKinley National Park” [Denali National Park, Alaska]. Types: AMNH. Refs.: Klots, 1940, J. N. Y. Entomol. Soc. 48(4): 413-414.

  • ssp. eunomia (Esper, 1800); original combination: Pap[ilio]. Nimph[alis]. phal[erata]. Eunomia. Type loc.: “Königsberg in Preussen” [Kaliningrad, Kaliningrad Oblast, Russia]. Types: type(s) probably lost. Refs.: Esper, 1800, Die Schmett. 1(suppl.)(1)(8/9): 94, pl. 110 (Cont. 65), fig. 5 ♂ D&V.

    • syn. aphirape (Hübner [1800]). Type loc.: Schwaben’ [Bavaria, Germany]. Types: type(s) probably lost. Refs.: Hübner, [1800], Samml. Europ. Schmett. 1: pl. 5, figs. 23 ♂ D, 24 ♂ V, 25 ♀ D [1800]; 8, no. 1 [1806].

    • syn. tomyris (Herbst, 1800). Type loc.: “Preussen” [Northern Germany and Poland]. Types: type(s) probably lost. Refs.: Herbst, 1800, In: Jablonsky, Natursyst. Ins. Schmett. 10: 102-104, no. 41, pl. 270, figs. 6 ♀ D, 7 ♀ V.

  • ssp. exspectata Morgun, 2011; original combination: Boloria eunomia exspectata. Type loc.: Caucasus, Teberda vic., Karachaevo-Tcherkesia. Refs.: Morgun, 2011; Caucasian Ent. Bull. 7 (1): 86.

  • ssp. itelmena Gorbunov, 2007; original combination: Boloria eunomia itelmena. Type loc.: Central Kamchatka.

  • ssp. laddi (Klots, 1940); original combination: Clossiana aphirape laddi. Type loc.:  “Snowy Range, Albany Co., Wyoming”. Types: AMNH. Refs.: Klots (1940): 4.

  • ssp. nichollae (Barnes & Benjamin, 1926); original combination: Brenthis aphirape race nichollae. Type loc.: “Rocky Mts”; Nicholl (1906), 36th Ann. Rept. Entomol. Soc. Ont. (1905): 72 defined the region as “Mt. Assiniboine.11,800 feet.just west of the Divide.the southernmost outlier of the glacier fields of the northern Rockies”; suggested to be “near the Columbia Ice Fields” by Pike (1981), J. Res. Lepid. 18(3): 157-158. [British Columbia]. Types: NMNH. Refs.: Barnes & Benjamin, 1926, Bull. South. Calif. Acad. Sci. 25(3): 92, no. 188d; Ferris and Groothuis (1971), J. Res. Lepid. 9(4): 243 (taxonomy).

  • ssp. ossianus (Herbst, 1800); original combination: Papilio ossianus Refs.: Herbst, 1800, Naturs. Schmett. 10: pl. 270, f. 4-5.

  • ssp. tenera Morgun, 2011; original combination: Boloria eunomia tenera. Type loc.: Transcaucasia (Tsakhdadzor, Armenia). Refs.: Morgun, 2011; Caucasian Ent. Bull. 7 (1): 86.

  • ssp. triclaris (Hübner [1821]); original combination: Argynnis Triclaris. Type loc.: not stated; ; listed as “Labrador” by W. F. Kirby (1908), in: P. Wytsman, Fac. ed. Hübner Samml. exot. Schmett. 3(addit. notes): 17; suggested to be “Moravian missionaries in Labrador” by W. Holland (1928), Ann. Carnegie Mus. 19(1): 38-39, no. 5, stated to be “Labrador” by Hemming (1937), Hübner 2: 123 (manuscript #36); subsequently defined as “Nain, Labrador, Canada” by Pike (1981), J. Res. Lepid. 18(3): 160. [Newfoundland and Labrador]. Types: type(s) probably lost. Refs.: Hübner, [1821], Samml. exot. Schmett. 2: pl. [19], figs. 1 ♂ D, 2 ♂ V, 3 ♀ D, 4 ♀ V; [1] (index).

    • syn. lais (Scudder, 1875). Type loc.: “Caribou Island, Straits of Belle Isle, Labrador.also.at Mackenzie River.Colorado”. This name was listed in the synonymy of Argynnis triclaris Hübner, [1821], but not described; it was a nomen nudum until the name helena was proposed to replace Argynnis lais W. H. Edwards, 1884, by dos Passos and Grey (1957), J. N. Y. Entomol. Soc. 63: 95-96. This action made Argynnis lais Scudder, 1875, available (Code Article 11.6.1). Refs.: Scudder, 1875, Proc. Boston Soc. Nat. Hist. 17(3): [294]-296.

  • ssp. ursadentis Ferris & Groothuis, 1971; original combination: Boloria eunomia ursadentis. Type loc.: “Beartooth Pass, 1/2 mile S. of Montana-Wyoming border, Park Co., Wyoming”. Types: FLMNH. Refs.: Ferris & Groothuis, 1971, J. Res. Lepid. 9(4): 245-247, figs. 4 ♂ D, 5 ♀ D, 6 ♂ V, 7 ♀ V, 8 ♀ V (aberrant), 9 ♀ D.

Genus Euptoieta

Genus: Euptoieta Doubleday [1848]

Type species: Euptoieta claudia (Cramer, [1775]); original combination: Papilio claudia Cramer, [1775]

Synonyms: None

References: Doubleday ([1848]): 168

8 species:

• bogotana Staudinger, 1885; original combination: Euptoieta bogotana

  • syn. colombia Strecker, 1899; original combination: Euptoieta colombia.

• claudia (Cramer [1775])

  • ssp. claudia (Cramer [1775]); original combination: Pap[ilio]. Nymph[alis]. Phalerat[a]. Claudia. Type loc.: “Jamaika” “Jamaïque”. Types: type(s) probably lost. Refs.: Cramer [1775] Uitl. Kapellen 1(6): 109, pl. 69, figs. E ♂ D, F ♂ V; (8): 152 (index).

    • syn. clausius (Herbst, 1798), repl. name of Papilio claudia Cramer, 1775, erroneously thought to be preoccupied

    • syn. thais (Doubleday, 1848), nom. nud.

  • ssp. daunius (Herbst, 1798); original combination: Papilio Daunius. Type loc.: “Ostindien” [East Indies]. Types: type(s) probably lost. Refs.: Jablonsky, Natursyst. Ins., Schmett. 9: 184-185, no. 11, pl. 256, figs. 1, 2.

    • syn. mariamne (Scudder, 1889), nom. nud.

    • syn. claudia tr.f. dodgei Gunder, 1927. Type loc.: “near Scribner, Dodge County, Nebraska”. Types: “holotype” probably in CAS. Refs.: Gunder (1927) Entomol. News 38(5): 135-136, no. 6, pl. 2, fig. 6 ♀ D (holotype).

    • syn. claudia tr.f. fumosa Field, 1936. Type loc.: “Hatfield, Ark” [Polk County, Arkansas]. Types: “holotype” probably in NMNH. Refs.: Field (1936) J. Entomol. & Zool. 28(2): 22-23.

    • syn. claudia tr.f. albaclaudia Field, 1936. Type loc.: “Miami, Fl” [Miami-Dade County, Florida]. Types: “holotype” in NMNH.

• hegesia (Cramer [1779])

  • ssp. hegesia (Cramer [1779]); original combination: Pap[ilio]. Nymph[alis]. Phal[erata]. Hegesia. Type loc.: “Nord-Amerika, en Nieuwjork, en op Eiland Jamaïka” “l’Amerique septentrionale, à la Nouvelle-York, & dans l’Isle de la Jamaïque”; neotype from “Montego Bay, Jamaica”.

    • syn. hegesia karibica Stichel, 1938

    • syn. hegesia domingensis Cucurullo, 1959

  • ssp. meridiania Stichel, 1938; original combination: [Euptoieta] h[egesia] meridiania. Type loc.: “Süd-Brasilien (typ.: Espirito Santo)” [Espirito Santo, Brazil]. Types: type(s) probably in NHMUK or lost. Refs.: Stichel (1938) Lepid. Cat. 86: 111; cited Brenthis claudia (sensu Geyer (1837), Zutr. Samml. exot. Schmett. 5: 11, pl. [143], figs. 831, 832).

    • syn. hegesia hoffmanni Comstock, 1944. Type loc.: “Mexico.Escuiñapa, Sinaloa”. Types: AMNH. Refs.: Comstock (1944) Sci. Surv. Puerto Rico & Virgin Is. 12(4): 446, note 1, fig. 6 FW marking.

  • ssp. watsoni Comstock, 1944; original combination: Euptoieta hegesia watsoni

• hortensia (Blanchard, 1852); original combination: Argynnis hortensia

  • syn. valdiviana (Philippi, 1859); original combination: Argynnis valdiviana.

  • syn. ramirezi Giacomelli, 1926; original combination: Euptoieta ramirezi.

  • syn. claudia ab. infuscata Hayward, 1940; original combination: Euptoieta claudia ab. infuscata.

• perdistincta Hall, 1930; original combination: Euptoieta perdistincta

  • syn. hana (Clark, 1936); original combination: Brenthis hana.

• poasina Schaus, 1913; original combination: Eupetoieta[sic] poasina

• sunides (Hewitson, 1877); original combination: Argynnis sunides

  • syn. hegemone Dognin, 1891, nom. nud.; original combination: Euptoieta hegemone.

• thekla Hall, 1919; original combination: Euptoieta claudia var. thekla

  • syn. thekla Weeks, 1905, nom. nud.; original combination: Euptoieta thekla.

Genus Pardopsis

Genus: Pardopsis Trimen, 1887

Type species: Pardopsis punctatissima (Boisduval, 1833); original combination: Acraea punctatissima Boisduval, 1833

Synonyms: None

References: Trimen (1887): 182

1 species:

• punctatissima (Boisduval, 1833); original combination: Acraea punctatissima

  • syn. NA (Boisduval, 1847); original combination: Acraea stictica.

Genus Yramea

Genus: Yramea Reuss, 1920

Type species: Yramea cytheris (Drury, [1773]); original combination: Papilio cytheris Drury, [1773]

Synonyms:

• Chilargynnis Bryk, 1944. Type species: Yramea cytheris (Drury, [1773]); original combination: Papilio cytheris Drury, [1773]. Refs.: Bryk (1945): 8

References: Reuss (1920): 192

6 species:

• cytheris (Drury [1773])

  • ssp. cytheris (Drury [1773]); original combination: Papilio cytheris

    • syn. cytheris falklandica (Watkins, 1924)

  • ssp. siga (Hübner, 1832); original combination: Argynnis siga

    • syn. anna (Blanchard, 1852)

    • syn. chilensis (Reed, 1877), nom. nud.

    • syn. cytheris subtusviola Bryk, 1944

• inca (Staudinger, 1894)

  • ssp. inca (Staudinger, 1894); original combination: Argynnis inca

  • ssp. cora (Lehmann, 1913); original combination: Argynnis (Brenthis) cora

• lathonioides (Blanchard, 1852); original combination: Argynnis lathonioides

  • syn. dexamene (Boisduval, 1859); original combination: Argynnis dexamene.

  • syn. dioides (Boisduval, 1859), nom. nud.; original combination: Argynnis dioides.

  • syn. montana (Reed, 1877); original combination: Argynnis montana.

  • syn. euterpe (Reed, 1877), nom. nud.; original combination: Argynnis euterpe.

  • syn. darwini (Staudinger, 1899); original combination: Argynnis darwini.

• lynx Lamas & Grados, 2004; original combination: Yramea lynx. Type loc.: Peru: Ancash, Laguna Conococha, 3900m. Types: MHNUSM. Distr.: open grasslands in high Andes of central Peru (above 3400m). Refs.: Lamas and Grados (2004): 47. Ety.: wildcat (wing coloration resembling the pelage of a spotted cat, e.g. the Andean cat Oreailurus jacobita)

• modesta (Blanchard, 1852)

  • ssp. araucania Benyamini, 2014; original combination: Yramea modesta araucania. Type loc.: Chile: Vn. Laima, 1200-1400m. Types: MNHNC.

  • ssp. modesta (Blanchard, 1852); original combination: Argynnis modesta

• sobrina (Weymer, 1890); original combination: Argynnis sobrina

  • syn. tessellata (Rosenberg & Talbot, 1914); original combination: Boloria tessellata.

  • syn. blancasi Hayward, 1960; original combination: Yramea blancasi.

Tribe Heliconiini

Type genus: Heliconius Kluk, 1780

Classification: The tribe Heliconiini is divided into eight genera based on morphological characters (Michener 1942 a): Agraulis (8 species), Dione (3 species), Dryadula (1 species), Dryas (4 species), Eueides (12 species), Heliconius (47 species), Philaethria (10 species) and Podotricha (2 species).

Characteristics: Larvae are spiny and feed on Passiflora (Passifloraceae).

Phylogenetic relationships: Extensive historical and on-going gene flow in this group makes phylogenetic inference challenging. Molecular phylogenies support the eight genera as monophyletic clades, but the relationships between many genera require further investigation.

Distribution: Neotropics; see occurrence maps by Neil Rosser

References: Michener (1942 a) (keys); Emsley (1963); Emsley (1965); Brown Jr. (1976); Brown (1981) (review); Brown and Fernández Yepez (1984) (Venezuela); Brown Jr. et al. (1992) (chromosomes); Penz and Peggie (2003) (phylogeny); Kozak et al. (2015) (molecular phylogeny); Kozak et al. (2021) (genome phylogeny); Cicconardi et al. (2023) (genome phylogeny)

Genus Agraulis

Genus: Agraulis Boisduval & Le Conte [1835]

Type species: Agraulis vanillae (Linnaeus, 1758); original combination: Papilio vanillae Linnaeus, 1758

Synonyms: None

References: Boisduval (1833 b): 142; Núñez et al. (2022) (phylogeny, taxonomy); Farfán et al. (2022) (as subgenus of Dione); Penz (2022) (as genus)

8 species:

• dodona (Lamas & Farfán, 2022); original combination: Dione (Agraulis) dodona. Type loc.: Pacaychacra, Arequipa, Peru (1500 m). Refs.: Farfán et al. (2022): 203.

• forbesi Michener, 1942; original combination: Agraulis vanillae forbesi. Type loc.: Lima, Peru. Types: Cornell University; paratypes: AMNH. Distr.: Coastal region of Peru. Refs.: Michener (1942 b): 3; Núñez et al. (2022): 167 (to species). Ety.: T. M. Forbes

• galapagensis Holland, 1890; original combination: Agraulis vanillae var. galapagensis. Type loc.: Chatham Island, Galapagos. Distr.: Galapagos Islands. Refs.: Howard (1890): 194; Núñez et al. (2022): 167 (to species). Ety.: Galapagos

• incarnata (Riley, 1926)

  • ssp. incarnata (Riley, 1926); original combination: D[ione] vanillae incarnata. Type loc.: “near Durango City, Mexico” [Durango]. Types: Lectotype (#Rh10118) in NHMUK. Distr.: Central America, southern USA. Refs.: Riley (1926): 243; Michener (1942 b) (lectotype); Zhang et al. (2020): 24 (to species; as Dione); Núñez et al. (2022) (to species).

    • syn. vanillae ab. comstocki (Gunder, 1925). Type loc.: “Monrovia, Los Angeles County, California”. Types: “Holotype” in AMNH. Refs.: Gunder, 1925, Entomol. News 36(1): 5-6, no. 9, pl. 1, fig. T ♂ D.

    • syn. vanillae insularis tr.f. fumosus (Gunder, 1927). Type loc.: “Los Angeles, Los Angeles County, California”. Types: “Holotype” in AMNH. Refs.: Gunder, 1927, Entomol. News 38(5): 137, no. 9, pl. 2, fig. 9 ♀ D (holotype).

    • syn. vanillae f. incarnata tr.f. margineapertus (Gunder, 1928). Type loc.: “Los Angeles” [Los Angeles County, California]. Types: “Holotype” in AMNH. Refs.: Gunder, 1928, Can. Entomol. 60(7): 163, pl. A, fig. 3.

    • syn. vanillae f. incarnata tr.f. hewlettae (Gunder, 1930). Type loc.: “Ontario, Riverside Co., Calif” [California]. Types: “Holotype” in AMNH. Refs.: Gunder, 1930, Bull. Brooklyn Entomol. Soc. 24(5): 327-328, pl. 31, fig. 7 ♀ D&V (holotype).

  • ssp. nigrior Michener, 1942; original combination: Agraulis vanillae nigrior. Type loc.: Upper Matecumbe Key, Florida. Types: AMNH. Refs.: Michener (1942 b): 7.

• insularis Maynard, 1889; original combination: Agraulis insularis. Type loc.: Bahamas. Distr.: Bahamas and the Greater Antilles. Refs.: Maynard (1889): 89; Núñez et al. (2022): 167 (as species).

• lucina Felder & Felder, 1862; original combination: Agraulis lucina Distr.: Eastern slope of the Andes in Ecuador, Peru, Brazil. Refs.: Núñez et al. (2022): 167 (as species).

  • syn. vanillae catella (Stichel [1908]), hyb.

• maculosa (Stichel [1908]); original combination: Dione vanillae maculosa Refs.: Núñez et al. (2022): 167 (to species).

  • syn. argentea (Larrañaga, 1923).

  • syn. vanillae f. superargentata (Giacomelli, 1925).

• vanillae (Linnaeus, 1758); original combination: Papilio vanillae. Type loc.: probably Suriname (Zhang et al. 2020). Refs.: Linnaeus (1758) 482.

  • syn. passiflorae (Fabricius, 1793), repl. name.

Genus Dione

Genus: Dione Hübner [1819]

Type species: Dione juno (Cramer, 1779); original combination: Papilio juno Cramer, 1779

Synonyms: None

References: Hübner ([1819]): 31

3 species:

• glycera (Felder & Felder, 1861)

  • syn. glycera (Ménétriés, 1857), nom. nud.

  • syn. glycera gnophota Stichel, 1907. Distr.: Venezuela, Colombia, Ecuador, Peru, Bolivia (Riley 1926).

• juno (Cramer, 1779)

  • ssp. andicola (Bates, 1864)

  • ssp. huascuma (Reakirt, 1866); original combination: Agraulis huascuma. Type loc.: “Mexico”. Types: Syntypes in FMNH. Refs.: Reakirt, 1866, Proc. Acad. Nat. Sci. Philad. 18(3): 243, no. 14; Recorded from Texas by Bordelon (2013), News Lepid. Soc. 55(suppl. S1): 65.

    • syn. huascama auctorum, missp.

  • ssp. juno (Cramer, 1779); original combination: [Papilio] Helicon[ius]. Juno. Type loc.: Not stated; given as “Surinam”by Lamas et al. (2004), Atl. Neotrop. Lepid. Chcklst. Part 4A: 264. Types: Location of type(s) not known, possibly lost. Refs.: Cramer, 1779, Uitl. Kapellen 3(18): 38, pl. 215, figs. B ♂ D, C ♂ V; (24): 175 (index).

  • ssp. miraculosa Hering, 1926

  • ssp. suffumata Brown & Mielke, 1972, ICZN, Art. 45.6.2

    • syn. juno ab. suffumata Hayward, 1931

• moneta Hübner [1825]

  • ssp. moneta Hübner [1825]; original combination: Dione Moneta. Type loc.: Not stated; suggested to be “South Brazil” by A. Hall (1921), Entomol. 54(703): 276. Types: Location of type(s) not known, probably lost. Refs.: Hübner, [1825], Samml. exot. Schmett. 2: 20, pl. [20], figs. 1 ♂ D, 2 ♂ V.

  • ssp. butleri Stichel [1908]; original combination: Dione moneta butleri. Type loc.: “Colombia, Rio Magdal. sup.Ecuador, Balzapamba”. Types: Syntypes (#549, 620, 623) in NHMUK; one of these figured by Ackery and Smiles (1976), Bull. Brit. Mus. (Nat. Hist.) Entomol. 32(5): pl. 1, fig. 9 ♂ D&V. Refs.: Stichel, [1908], In: Wytsman, Genera Ins. 63: 19, no. 3b (nota 1); figured by Seitz (1912), Grossschmett. Erde 5(152): pl. 84e, figs. [1] ♂ V, [2] ♂ D.

  • ssp. poeyii Butler, 1873; original combination: D[ione]. Poeyii. Type loc.: Not stated. N. Riley (1926), Entomol. 59(760): 244, determined that the description of this taxon was based on figures and text in E. Doubleday 1847, Gen. diurn. Lepid. 1(13): pl. 22, fig. 1 ♂ D; (18): 154; lectotype from “Oaxaca, Mexico”. Types: Lectotype in NHMUK, designated by Lamas (1998), Revta. peruana Entomol. 40: 113. Distr.: Central America, southern USA. Refs.: Butler, 1873, Ann. Mag. Nat. Hist. (4)12: 227, no. 28.

Genus Dryadula

Genus: Dryadula Michener, 1942

Type species: Dryadula phaetusa (Linnaeus, 1758); original combination: Papilo phaetusa Linnaeus, 1758

Synonyms: None

References: Michener (1942 a): 4

1 species:

• phaetusa (Linnaeus, 1758); original combination: P[apilio]. N[ymphalis]. [Phalerata] Phaetusa. Type loc.: “Indiis”. Types: Lectotype in UUZM (#1946), designated by Honey and Scoble (2001), Zool. J. Linn. Soc. 132(3): 361. Refs.: Linnaeus, 1758, Syst. Nat. (ed. 10) 1: 478, no. 123; cited Rösel von Rosenhof (1761), Der monat. Insect.-Belustig. 4: pl. 2, fig. 1.

  • syn. phaetusa f. stupenda (Stichel [1908]); original combination: C[olaenis]. phaetusa forma stupenda. Type loc.: Panama (Bugaba)”. Types: “Holotype” (#533) in NHMUK; figured by Ackery and Smiles (1976), Bull. Brit. Mus. (Nat. Hist.) Entomol. 32(5): pl. 1, fig. 3 ♂ D&V. Refs.: Stichel, [1908], In: Wytsman, Genera Ins. 63: 13, no. 2α (nota 1).

  • syn. phaetusa f. deleta (Stichel [1908]); original combination: C[olaenis]. phaetusa forma deleta. Type loc.: “Paraguay”; holotype from “Paraguay, Nuev. Germania”. Types: “Holotype” (#532) in NHMUK; figured by Ackery and Smiles (1976), Bull. Brit. Mus. (Nat. Hist.) Entomol. 32(5): pl. 1, fig. 4 ♂ D&V. Refs.: Stichel, [1908], In: Wytsman, Genera Ins. 63: 13, no. 2β (nota 2).

  • syn. phaetusa f. lutulenta (Stichel [1908]); original combination: C[olaenis]. phaetusa forma ♀ lutulenta. Type loc.: “Paraguay”; holotype from “Paraguay, Nuev. Germania”. Types: “Holotype” (#532) in NHMUK; figured by Ackery and Smiles (1976), Bull. Brit. Mus. (Nat. Hist.) Entomol. 32(5): pl. 1, fig. 5 ♀ D&V. Refs.: Stichel, [1908], In: Wytsman, Genera Ins. 63: 13, no. 2γ.

Genus Dryas

Genus: Dryas Hübner [1807]

Type species: Dryas iulia (Fabricius, 1775); original combination: Papilio iulia Fabricius, 1775

Synonyms:

• Alcionea Rafinesque, 1815. Type species: Dryas iulia alcionea (Cramer, 1779); original combination: Papilio alcionea Cramer, 1779. Refs.: Rafinesque (1815): 127
• Colaenis Hübner [1819]. Type species: Dryas iulia (Fabricius, 1775); original combination: Papilio iulia Fabricius, 1775. Refs.: Hübner ([1819]): 32

References: Hübner (1807): pl. [43]; Núñez et al. (2022) (phylogeny, taxonomy)

4 species:

• alcionea (Cramer, 1779)

  • ssp. alcionea (Cramer, 1779); original combination: Pap[ilio]. Hel[iconius]. Alcionea. Type loc.: “Surinamen” “Suriname”. Types: Type(s) probably lost. Refs.: Cramer (1782): 38; Núñez et al. (2022): 171 (to species).

    • syn. iulia titio (Stichel [1908]). Type loc.: “Bolivia”. Types: Syntypes in NHMUK. Refs.: Stichel, [1908], In: Wytsman, Genera Ins. 63: 12, in no. 1b (nota 1).

  • ssp. moderata (Riley, 1926), ICZN, Art. 45.5.1 (Riley adopted Colaenis iulia delila form moderata Stichel, [1908], to represent a subspecies-level taxon, with the same data); original combination: C[olaenis]. julia moderata Refs.: Riley (1926): 241; Núñez et al. (2022): 171 (as subspecies of alcionea).

    • syn. iulia delila f. moderata (Stichel [1908]). Type loc.: “Honduras”. Types: “Holotype” (#525) in ZMHU.

  • ssp. framptoni (Riley, 1926); original combination: Colaenis iulia framptoni. Type loc.: St. Vincent. Types: NHMUK. Refs.: Riley (1926): 241; Riley (1926): 241; Núñez et al. (2022): 171 (as subspecies of alcionea).

• dominicana (Hall, 1917)

  • ssp. dominicana (Hall, 1917); original combination: Colaenis iulia var. dominicana Refs.: Núñez et al. (2022): 171 (to species).

  • ssp. warneri (Hall, 1936); original combination: Colaenis iulia warneri Refs.: Núñez et al. (2022): 171 (as subspecies of dominicana).

• iulia (Fabricius, 1775)

  • ssp. carteri (Riley, 1926); original combination: Colaenis iulia carteri. Type loc.: Bahamas: Nassau. Types: NHMUK. Distr.: Bahamas. Refs.: Riley (1926): 240; Núñez et al. (2022). Ety.: Sir Gilbert Carter

  • ssp. delila (Fabricius, 1775)

    • syn. cillene (Cramer, 1779)

  • ssp. fucatus (Boddaert, 1783); original combination: Papilio fucatus

    • syn. julia hispaniola (Hall, 1917)

  • ssp. iulia (Fabricius, 1775); original combination: P[apilio]. N[ymphalis]. P[halerata]. Iulia. Type loc.: “America”. Types: Lectotype in NHMUK. Refs.: Fabricius (1775): 509; Wetherbee (1991), Two Cent. Expl. Hispaniola Butterflies: 18 (lectotype).

    • syn. iulia juncta Comstock, 1944. Type loc.: “Adjuntas, Porto Rico” [Puerto Rico]. Types: AMNH. Refs.: Comstock, 1944, Sci. Surv. Puerto Rico & Virgin Is. 12(4): 441-443, fig. 3 venation, pl. 6, fig. 1 ♂ D.

  • ssp. largo Clench, 1975; original combination: Dryas iulia largo. Type loc.: “Key Largo, Monroe Co., Florida”. Types: CMNH. Refs.: Clench, 1975, J. Lepid. Soc. 29(4): 230.

  • ssp. nudeola (Bates, 1934), ICZN, Art. 45.5.1; original combination: Colaenis iulia nudeola

    • syn. iulia cillene f. nudeola (Stichel [1908])

  • ssp. zoe Miller & Steinhauser, 1992; original combination: Dryas iulia zoe

• lucia (Riley, 1926)

  • ssp. lucia (Riley, 1926); original combination: Colaenis iulia lucia Refs.: Riley (1926): 241; Núñez et al. (2022): 171 (to species).

  • ssp. martinica Enrico & Pinchon, 1969; original combination: Dryas iulia martinica Refs.: Núñez et al. (2022): 171 (as subspecies of lucia).

Genus Eueides

Genus: Eueides Hübner, 1816

Type species: Eueides isabella (Stoll, 1781); original combination: Nereis dianasa Hübner [1806]

Synonyms:

• Mechanitis [Illiger], 1807. Type species: Eueides isabella (Stoll, 1781); original combination: Nereis dianasa Hübner [1806]. Refs.: [Illiger] (1807): 1180; ICZN (1954)
• Semelia Doubleday [1845]. Type species: Eueides vibilia (Godart, 1819); original combination: Cethosia vibilia Godart, 1819. Refs.: Doubleday ([1845]): 64
• Evides Agassiz, 1846. Type species: Eueides isabella (Stoll, 1781); original combination: Nereis dianasa Hübner [1806]. Refs.: Agassiz (1846): 147, 153
• Semelia Erichson [1848]. Type species: Eueides lybia (Fabricius, 1775); original combination: Papilio lybia Fabricius, 1775. Refs.: Erichson (1848): 596
• Semelia Boisduval, 1870. Type species: Eueides vibilia (Godart, 1819); original combination: Cethosia vibilia Godart, 1819. Refs.: Boisduval (1870): 35

References: Hübner (1816): 11

13 species:

• aliphera (Godart, 1819)

  • ssp. aliphera (Godart, 1819)

  • ssp. cyllenella Seitz, 1912

    • syn. aliphera ab. cillenula Seitz, 1913, missp.

    • syn. cyllenula Seitz, 1924, emend.

  • ssp. gracilis Stichel, 1903; original combination: Eueides aliphera gracilis. Type loc.: Costa Rica, Honduras. Distr.: Central America: Costa Rica, Nicaragua, Guatemala, Honduras, Mexico. Refs.: Stichel (1903): 23.

• emsleyi Brown, 1976

  • ssp. emsleyi Brown, 1976

  • ssp. esmeraldensis Cast, 2013. Type loc.: Ecuador, Esmeraldas, route Lita-San Lorenzo. Types: MNHN. Distr.: Ecuador. Refs.: Cast (2013).

• heliconioides Felder & Felder, 1861

  • ssp. eanes Hewitson, 1861

  • ssp. heliconioides Felder & Felder, 1861

    • syn. eanes ab. eanides Stichel, 1903. Type loc.: Bolivia: La Paz; Peru: Marcapato. Refs.: Stichel (1903): 30.

    • syn. eanes ab. farragosa Stichel, 1903, hyb. Type loc.: Peru: Marcapato. Refs.: Stichel (1903): 30.

    • syn. eanes ab. aides Stichel, 1903. Type loc.: Peru: Hillapani, Huanaco; Bolivia: La Paz (Cuzco). Refs.: Stichel (1903): 30.

    • syn. eanes ab. riffarthi Stichel, 1903, hyb. Type loc.: southern Peru; Bolivia: La Paz (Cuzco). Refs.: Stichel (1903): 31.

    • syn. eanes ab. felderi Stichel, 1903 Refs.: Stichel (1903): 31. Ety.: Cajetan, Freiherr von Felder

    • syn. eanes ab. pluto Stichel, 1903. Type loc.: Peru: Hillapani. Refs.: Stichel (1903): 32.

  • ssp. koenigi Holzinger & Holzinger, 1993

    • syn. eanes koeningi Holzinger & Holzinger, 1993, missp.

• isabella (Stoll, 1781)

  • ssp. arquata Stichel, 1903. Type loc.: Colombia: Cauca; Chiriqui. Refs.: Stichel (1903): 9.

    • syn. isabella hübneri ab. spoliata Stichel, 1903 Refs.: Stichel (1903): 9.

    • syn. isabella isabella f. perimacula Boullet & Le Cerf, 1910

  • ssp. cleobaea Geyer, 1832

    • syn. anaxa Ménétriés, 1857

  • ssp. dianasa (Hübner [1806]); original combination: Nerëis fulva dianasa Refs.: Hübner (1806): pl. [8]; Hübner (1816): 11.

    • syn. dianasa ab. decolorata Stichel, 1903. Type loc.: Bahia, Brazil. Refs.: Stichel (1903): 3.

  • ssp. dissoluta Stichel, 1903. Type loc.: Peru & Bolivia. Refs.: Stichel (1903): 6.

    • syn. isabella hippolinus ab. margaritifera Stichel, 1903. Type loc.: Peru. Refs.: Stichel (1903): 5.

    • syn. isabella pellucida ab. vegetissima Stichel, 1903. Type loc.: Ecuador. Refs.: Stichel (1903): 7.

    • syn. isabella hübneri var. olga Neustetter, 1916

  • ssp. dynastes Felder & Felder, 1861

  • ssp. ecuadorensis Strand, 1912

  • ssp. eva (Fabricius, 1793); original combination: P[apilio] H[eliconius]. Eva. Type loc.: “Surinami” probably in error; suggested to be “Panama” by Lamas (1973), Revta. bras. Entomol. 17(19): 128. Types: Type(s) probably lost. Refs.: Fabricius, 1793, Entomol. Syst. 3(1): 162, no. 501; cited Jones’ Icones 2: pl. 20, fig. 1; Lamas (1973), Revta. bras. Entomol. 17(19): 127-130, detailed the use of this name over zorcaon, based on his examination of copies of William Jones’ unpublished drawings in the NHMUK.

    • syn. anaxa Ménétriés, 1855, nom. nud.

    • syn. zorcaon Reakirt, 1866. Type loc.: “Mexico (near Vera Cruz).Honduras, Guatemala”. Types: Syntypes in FMNH. Refs.: Reakirt, 1866, Proc. Acad. Nat. Sci. Philad. 18(3): 243, no. 12.

    • syn. cleobaea zorcaon ab. adusta Stichel, 1903. Type loc.: “Chiriqui, Honduras”; syntypes from “Panama, Bugaba; Honduras; Mexico, Vera Cruz”. Types: “Syntypes” in NHMUK; a possible syntype figured by Seitz (1912), in: Seitz, Grossschmett. Erde 5(151): pl. 80g, [2] ♂ D; a syntype was figured by Ackery and Smiles (1976), Bull. Brit. Mus. (Nat. Hist.) Entomol. 32(5): pl. 5, fig. 48 ♀ D&V. Refs.: Stichel (1903): 11, no. 3bα.

  • ssp. hippolinus Butler, 1873

    • syn. isabella hippolinus ab. personata Stichel, 1903. Type loc.: Peru. Refs.: Stichel (1903): 5.

    • syn. isabella hippolinus ab. brunnea Stichel, 1903. Type loc.: Peru. Refs.: Stichel (1903): 6.

    • syn. imitans Seitz, 1912

  • ssp. huebneri Ménétriés, 1857

    • syn. pellucida Srnka, 1885

    • syn. isabella seitzi Stichel, 1903. Type loc.: Ecuador & Colombia. Refs.: Stichel (1903): 6.

  • ssp. isabella (Stoll, 1781)

  • ssp. melphis (Godart, 1819)

    • syn. cleobaea var. monochroma Boullet & Le Cerf, 1910

  • ssp. nigricornis Maza, 1982

• lampeto Bates, 1862

  • ssp. acacetes Hewitson, 1869

    • syn. lampeto ab. fuliginosa Stichel, 1903. Type loc.: Ecuador. Refs.: Stichel (1903): 12.

    • syn. lampeto ab. amoena Stichel, 1903. Type loc.: Ecuador: Santa Inez, 1250 m; Bolivia. Refs.: Stichel (1903): 13.

    • syn. ambatensis Oberthür, 1916

  • ssp. apicalis Röber, 1927

    • syn. lampeto fuliginosus f. pallida Riffarth, 1907

  • ssp. brownsbergensis Gernaat & Beckles, 2010. Type loc.: Brownsberg, Suriname. Types: NCBN. Refs.: Gernaat and Beckles (2010): 172. Ety.: Brownsberg Nature Park in Suriname

  • ssp. carbo Seitz, 1913, ICZN, Art. 45.6.2

    • syn. lampeto ab. carbo Stichel, 1903. Type loc.: Ecuador: Santa Inez. Refs.: Stichel (1903): 13.

  • ssp. concisa Lamas, 1985

  • ssp. lampeto Bates, 1862

    • syn. lampeto romani Bryk, 1953

  • ssp. nigrofulva Kaye, 1906

    • syn. lampeto copiosus Stichel, 1906

• libitina Staudinger, 1885

  • ssp. libitina Staudinger, 1885

  • ssp. malleti Lamas, 1998

• lineata Salvin & Godman, 1868

• lybia (Fabricius, 1775)

  • ssp. lybia (Fabricius, 1775)

    • syn. hypsipyle (Cramer [1777])

  • ssp. orinocensis Brown & Fernández, 1985

  • ssp. otelloi Brown & Fernández, 1985

• olympia (Fabricius, 1793)

  • ssp. lybioides Staudinger, 1876

  • ssp. olympia (Fabricius, 1793)

    • syn. leucomma Bates, 1866

  • ssp. salcedoi Brown & Fernández, 1985

• pavana Ménétriés, 1857

  • syn. thyana Felder & Felder, 1860.

  • syn. pavana f. ferruginea Raymundo, 1931.

• procula Doubleday [1847]

  • ssp. asidia Schaus, 1920

  • ssp. browni Holzinger & Holzinger, 1974

  • ssp. edias Hewitson, 1861

    • syn. edias luminosus Stichel, 1903. Type loc.: Venezuela. Refs.: Stichel (1903): 16.

  • ssp. eurysaces Hewitson, 1864

  • ssp. kuenowii Dewitz, 1877

    • syn. edias umbratilis Röber, 1927

  • ssp. procula Doubleday [1847]

  • ssp. vulgiformis Butler & Druce, 1872

• tales (Cramer [1775])

  • ssp. barcellinus Zikán, 1937

    • syn. eanes f. lucretius Zikán, 1937

  • ssp. calathus Stichel, 1909

  • ssp. cognata Weymer, 1890

  • ssp. franciscus Brown & Holzinger, 1973

  • ssp. guyanensis Lacomme, 2011. Type loc.: French Guiana. Refs.: Lacomme (2011).

  • ssp. michaeli Zikán, 1937

  • ssp. pseudeanes Boullet & Le Cerf, 1910

  • ssp. pythagoras Kirby [1900]

    • syn. thales Hübner [1810], not Cramer, 1775

    • syn. thales heraldicus Stichel, 1903. Type loc.: Para, Brazil. Refs.: Stichel (1903): 27.

    • syn. tales f. zernyi Neustetter, 1928

  • ssp. surdus Stichel, 1903. Type loc.: Brazil (Amazonas: Obidos, Cameta); Guayana. Refs.: Stichel (1903): 27.

    • syn. thales surdus ab. aquilifer Stichel, 1903. Type loc.: Brazil (Amazonas: Obidos). Refs.: Stichel (1903): 28.

    • syn. thales f. reducta Neustetter, 1931

  • ssp. tabernula Lamas, 1985

  • ssp. tales (Cramer [1775])

    • syn. thalestris (Godart, 1819), repl. name

  • ssp. xenophanes Felder & Felder, 1865

    • syn. crystalina Hall, 1921

• vibilia (Godart, 1819)

  • ssp. louisi Brévignon, 2006 Distr.: French Guiana. Refs.: Brevignon (2006).

    • syn. vibilia romeroi Holzinger & Holzinger, 1994, unavail. (ICZN, Art. 15.1)

  • ssp. unifasciatus Butler, 1873

    • syn. vibilia var. arcita Stichel, 1903, nom. nud. Refs.: Stichel (1903): 20.

  • ssp. vialis Stichel, 1903. Type loc.: Colombia, Costa Rica, Guatemala. Refs.: Stichel (1903): 21.

  • ssp. vibilia (Godart, 1819)

    • syn. mereaui (Hübner, 1819), nom. nud.

    • syn. mereaui (Hübner, 1823)

    • syn. ab. pallens Stichel, 1903. Type loc.: Esp. Santo, Brazil. Refs.: Stichel (1903): 19.

  • ssp. vicinalis Stichel, 1903. Type loc.: Palmar, Ecuador (100 m). Refs.: Stichel (1903): 21.

Genus Neruda

Genus: Neruda Turner, 1976

Type species: Heliconius aoede (Hübner, 1816); original combination: Migonitis aoede Hübner, 1816

Synonyms: None

References: Turner (1976): 301 (as subgenus)

4 species:

• aoede (Hübner, 1816)

  • ssp. aliciae Neukirchen, 2000

  • ssp. aoede (Hübner, 1816); original combination: Migonitis aoede Refs.: Hübner (1816): 12.

    • syn. aoede (Hübner, 1808), nom. nud.

    • syn. aoede (Hübner, 1813), nom. nud.

    • syn. aoede (Hübner, 1818), nom. nud. Refs.: Hübner (1818): 23, pl.[23], figs 129-130.

  • ssp. astydamia (Erichson [1849])

  • ssp. auca Neukirchen, 1997

  • ssp. ayacuchensis (Neukirchen, 1992); original combination: Heliconius (Neruda)

  • ssp. bartletti (Druce, 1876)

    • syn. vedius (Staudinger, 1885)

    • syn. aoede f. metella (Neustetter, 1932)

  • ssp. centurius (Neukirchen, 1994)

  • ssp. cupidineus (Stichel, 1906)

    • syn. bartletti var. reducta (Oberthür, 1920)

  • ssp. emmelina (Oberthür, 1902)

  • ssp. eurycleia (Brown, 1973)

  • ssp. faleria (Fruhstorfer, 1910)

  • ssp. lucretius (Weymer, 1891)

    • syn. aoede aoede f. postalbimacula (Bryk, 1953)

  • ssp. manu (Lamas, 1976)

  • ssp. philipi (Brown, 1976)

• godmani (Staudinger, 1882)

• metharme (Erichson [1849])

  • ssp. makiritare Brown & Fernández, 1985

  • ssp. metharme (Erichson [1849])

    • syn. thetis (Boisduval, 1870)

    • syn. anaclia (Riffarth, 1901), nom. nud.

  • ssp. perseis (Stichel, 1923)

• metis Moreira & Mielke, 2010; original combination: Neruda metis. Type loc.: Serra do Penitente, Balsas, Maranhão, Brazil. Types: Departamento de Zoologia, Universidade Federal do Paraná, Brazil. Distr.: only known from type locality. Refs.: Moreira and Mielke (2010): 86 (♀︎). Ety.: Metis, a moon of Jupiter (“In Greek mythology, Metis was the first wife of Zeus (the Roman Jupiter) and the mother of Athena”)

Genus Heliconius

Genus: Heliconius Kluk, 1780

Type species: Heliconius charithonia (Linnaeus, 1767); original combination: Papilio charithonia Linnaeus, 1767

Synonyms:

• Heliconius Latreille [1804]. Type species: Heliconius charithonia (Linnaeus, 1767); original combination: Papilio charithonia Linnaeus, 1767. Refs.: Latreille (1804): 185
• Migonitis Hübner, 1816. Type species: Heliconius erato (Linnaeus, 1758); original combination: Papilio erato (Linnaeus, 1764). Refs.: Hübner (1816): 12
• Sunias Hübner, 1816. Type species: Heliconius melpomene (Linnaeus, 1758); original combination: Papilio melpomene Linnaeus, 1758. Refs.: Hübner (1816): 12
• Ajantis Hübner, 1816. Type species: Heliconius sapho (Linnaeus, 1782); original combination: Papilio sapho Linnaeus, 1782. Refs.: Hübner (1816): 13
• Apostraphia Hübner, 1816. Type species: Heliconius charithonia (Linnaeus, 1767); original combination: Papilio charithonia Linnaeus, 1767. Refs.: Hübner (1816): 13
• Sicyonia Hübner, 1816. Type species: Heliconius sara (Fabricius, 1793); original combination: Papilio rhea Cramer, 1775. Refs.: Hübner (1816): 13
• Heliconia Latreille, 1818. Type species: Heliconius charithonia (Linnaeus, 1767); original combination: Papilio charithonia Linnaeus, 1767. Refs.: NA
• Laparus Billberg, 1820. Type species: Heliconius doris (Linnaeus, 1771); original combination: Papilio doris (Linnaeus, 1771). Refs.: Billberg (1820): 77
• Crenis Hübner, 1821. Type species: Heliconius doris doris (Linnaeus, 1771); original combination: Crenis brylle Hübner (1821). Refs.: NA
• Phlogris Hübner [1825]. Type species: Heliconius melpomene (Linnaeus, 1758); original combination: Papilio melpomene Linnaeus, 1758. Refs.: NA
• Sicyonia Hübner [1826] Refs.: An unjustified emendation of Sicyonia Hübner, 1816
• Podalirius Gistel, 1848. Type species: Heliconius charithonia (Linnaeus, 1767); original combination: Papilio charithonia Linnaeus, 1767. Refs.: Gistel (1848): ix
• Blanchardia Buchecker [1880]. Type species: Heliconius hecuba (Hewitson, [1858]); original combination: Blanchardia dismorphia Buchecker [1880]. Refs.: NA

Phylogenetic relationships: Molecular phylogenetic studies using a small number of loci (Brower 1994; Brower and Egan 1997; Beltrán et al. 2002; Beltrán et al. 2007; Kozak et al. 2015) supported at least six clades within Heliconius: Aoede, Doris, Wallacei, Melpomene (including the silvaniforms), Erato and Sara. The relationships among these groups and among species within each group remain uncertain.

Distribution: Widespread across the Neotropics

References: Kluk (1802): 82; Emsley (1965); ICZN (2014): 204

43 species:

• antiochus (Linnaeus, 1767)

  • ssp. antiochus (Linnaeus, 1767). Type loc.: “Indiis”.

    • syn. zobeide Butler, 1869

    • syn. antiochus ab. divisus Staudinger, 1897

    • syn. antiochus ab. alba Riffarth, 1900

    • syn. antiochus alba ab. trimaculata Krüger, 1933

  • ssp. aranea (Fabricius, 1793). Type loc.: not stated.

  • ssp. araneides Staudinger, 1897. Type loc.: Venezuela: Puerto Cabello.

  • ssp. conchitae Cast & Neild, 2024. Type loc.: Venezuela: La Fría. Distr.: south-western slopes of the Cordillera de Mérida, in the north-west of Venezuela. Refs.: Cast and Neild (2024): 6. Ety.: María de la Concepción (“Conchita”) Romero Montesino

  • ssp. salvinii Dewitz, 1877. Type loc.: Venezuela: Orinoco.

• charithonia (Linnaeus, 1767)

  • ssp. antiquus Lamas, 1988, repl. name

    • syn. charithonia punctata Hall, 1936, preocc. (not Neustetter, 1907)

  • ssp. bassleri Comstock & Brown, 1950

  • ssp. charithonia (Linnaeus, 1767); original combination: Papillio charithonia Refs.: Linnaeus (1767): 757.

    • syn. charitonia (Linnaeus, 1767), missp.

  • ssp. churchi Comstock & Brown, 1950

  • ssp. ramsdeni Comstock & Brown, 1950

  • ssp. simulator Röber, 1921

  • ssp. tuckeri Comstock & Brown, 1950; original combination: Heliconius charitonius tuckeri. Type loc.: “Winter Park, Florida” [Orange County]. Types: AMNH. Refs.: Comstock and Brown (1950): 13, no. F, 15-16, fig. 3, no. F ♂ D.

    • syn. charitonia tuckerorum Lamas, 1988, emend.

  • ssp. vazquezae Comstock & Brown, 1950; original combination: Heliconius charitonius vazquezae. Type loc.: “Campeche, Mexico”. Types: AMNH. Refs.: Comstock and Brown (1950): 13, no. 6, 16, fig. 3, no. G ♂ D.

• congener Weymer, 1890

  • ssp. aquilionaris Brown, 1976

  • ssp. congener Weymer, 1890

    • syn. apseudes var. paranapurae Staudinger, 1897

  • ssp. ocannensis Stichel, 1905, repl. name

    • syn. ocannus Staudinger, 1885, preocc. (not Buchecker, [1880])

• demeter Staudinger, 1897

  • ssp. beebei Turner, 1966

  • ssp. bouqueti Nöldner, 1901

    • syn. xanthoceras (Oberthür, 1902)

    • syn. eueidina (Oberthür, 1916)

    • syn. egeriformis (Joicey & Kaye, 1917)

    • syn. automatia (Oberthür, 1925)

    • syn. eratoformis Neustetter, 1931

  • ssp. demeter Staudinger, 1897

    • syn. f. similis Neustetter, 1931

    • syn. demeter angeli Neukirchen, 1997

  • ssp. joroni Lamas & Rosser, 2018. Type loc.: Peru, San Martín, Tarapoto, San Roque. Types: Natural History Museum at the San Marcos National University, Lima, Peru (MUSM). Refs.: Rosser et al. (2019): 238. Ety.: Mathieu Joron

  • ssp. karinae Neukirchen, 1990

  • ssp. neildi Neukirchen, 1997

  • ssp. terrasanta Brown & Benson, 1975

  • ssp. titan Neukirchen, 1995

  • ssp. turneri Brown & Benson, 1975

  • ssp. zikani Brown & Benson, 1975

• eratosignis (Joicey & Talbot, 1925)

  • ssp. eratosignis (Joicey & Talbot, 1925) Refs.: Rosser et al. (2019) (to species).

  • ssp. tambopata Lamas, 1985

  • ssp. ucayalensis Holzinger & Holzinger, 1975 Host plants: Passiflora skiantha (Rosser et al. 2019).

  • ssp. ulysses Brown & Benson, 1975

• eleuchia (Hewitson [1854])

  • ssp. eleuchia (Hewitson [1854])

  • ssp. eleusinus Staudinger, 1885

    • syn. sapho primularis ab. deflava Joicey & Kaye, 1917, hyb.

    • syn. ceres (Oberthür, 1920), hyb.

    • syn. primularis ceres Stichel, 1923, preocc. (not Oberthür, 1920), hyb.

  • ssp. primularis Butler, 1869

• leucadia Bates, 1862

  • ssp. andromeda Neukirchen, 1996

  • ssp. birgitae Neukirchen, 1996

  • ssp. leucadia Bates, 1862

  • ssp. pseudorhea Staudinger, 1897

• hewitsoni (Hewitson, 1875)

  • syn. hewitsoni Staudinger, 1876, preocc. (not Hewitson, 1875).

• peruvianus (Felder & Felder, 1859)

  • syn. aganice Riffarth, 1901, nom. nud.

• ricini (Linnaeus, 1758)

  • ssp. insulanus (Stichel, 1909)

  • ssp. ricini (Linnaeus, 1758)

    • syn. myrti (Fabricius, 1775)

    • syn. polyhymnia (Shaw, 1806)

• sapho (Drury, 1782)

  • ssp. candidus Brown, 1976

  • ssp. chocoensis Brown & Benson, 1975

  • ssp. leuce (Doubleday, 1847)

    • syn. sappho (Hübner [1831]), not Drury, 1782

    • syn. leuce Bates, 1864, preocc. (not Doubleday, 1847)

  • ssp. sapho (Drury, 1782)

• sara (Fabricius, 1793)

  • ssp. apseudes (Hübner [1813])

    • syn. apseudes (Hübner [1808]), nom. nud.

  • ssp. brevimaculata Riffarth, 1901, ICZN, Art. 45.5.1

    • syn. apseudes var. magdalena ab.? (var.?) brevimaculata Staudinger, 1897

  • ssp. elektra Neukirchen, 1998

  • ssp. fulgidus Stichel, 1906

  • ssp. magdalena Bates, 1864

    • syn. apseudes var. magdalena ab. albimaculata Staudinger, 1897

    • syn. sara ab. albula Riffarth, 1900

    • syn. sarae lilianae Emsley, 1965, nom. nud.

  • ssp. sara (Fabricius, 1793)

    • syn. rhea (Cramer [1775]), preocc. (not Poda, 1761)

    • syn. thamar (Hübner [1806]) Refs.: Hübner (1806): pl. [15].

    • syn. rhea ab. albinea Riffarth, 1899

    • syn. sara praesignis Stichel, 1919

    • syn. sara thamar f. nana Stammeshaus, 1982

  • ssp. sprucei Bates, 1864

  • ssp. theudela (Hewitson, 1874)

  • ssp. veraepacis Bates, 1864

    • syn. f. albifasciatus Hoffmann, 1940

  • ssp. williami Neukirchen, 1994

• doris (Linnaeus, 1771)

  • ssp. dives (Oberthür, 1920)

    • syn. doris alberato Avinoff, 1926

    • syn. doris azurea Avinoff, 1926

    • syn. doris viridis f. viridana Stichel, 1906

    • syn. doris eratonia f. suavior Kaye, 1914

    • syn. doris metharmina f. fascinator Kaye, 1914

  • ssp. doris (Linnaeus, 1771)

    • syn. quirina (Cramer [1775])

    • syn. amathusia (Cramer [1777])

    • syn. delila (Hübner [1813])

    • syn. crenis (Hübner, 1816), repl. name Refs.: Hübner (1816): 12.

    • syn. brylle (Hübner, 1821), repl. name

    • syn. dorimena (Doubleday, 1847), nom. nud.

    • syn. mars Staudinger, 1885

    • syn. erato ab. metharmina Staudinger, 1897

    • syn. erato ab. tecta Riffarth, 1900

    • syn. doris-caerulea Riffarth, 1901, nom. nud.

    • syn. doris rubra Stichel, 1906, nom. nud.

    • syn. doris nigra Stichel, 1906, nom. nud.

    • syn. doris caeruleatus Stichel, 1906

    • syn. doris delila f. albina Boullet & Le Cerf, 1909

    • syn. var. le moulti Boullet & Le Cerf, 1910

    • syn. ab. gibbsi Kaye, 1919

    • syn. doris dialis Stichel, 1923

    • syn. f. albescens Neustetter, 1926

  • ssp. morralli Cast, 2020. Type loc.: Trinidad. Types: Michel Cast Collection (A). Distr.: Trinidad. Refs.: Cast (2020): 86. Ety.: John Morrall

  • ssp. obscurus Weymer, 1891

    • syn. erato aristomache Riffarth, 1901

  • ssp. viridis Staudinger, 1885

    • syn. erato var. eratonius Staudinger, 1897

    • syn. erato var. transiens Staudinger, 1897

    • syn. erato luminosus Riffarth, 1901

• hecuba (Hewitson [1858])

  • ssp. bonplandi Neukirchen, 1991

  • ssp. cassandra Felder & Felder, 1862

  • ssp. choarina (Hewitson, 1872)

    • syn. dismorphia (Buchecker [1880]), nom. nud.

    • syn. f. cacica Neustetter, 1928

  • ssp. creusa Holzinger & Holzinger, 1989

    • syn. hecuba crispus var. crespinus Krüger, 1925, hyb.

  • ssp. crispus Staudinger, 1885

  • ssp. flava Brown, 1979, ICZN, Art. 45.5.1

    • syn. hecuba choarinus f. flava Neustetter, 1928

  • ssp. hecuba (Hewitson [1858])

    • syn. hecuba intermedius Riffarth, 1907, hyb.

  • ssp. lamasi Neukirchen, 1991

  • ssp. salazari Neukirchen, 1993

  • ssp. tolima Fassl, 1912

    • syn. hecuba tolima Seitz, 1913, preocc. (Fassl, 1912)

  • ssp. walteri Salazar, 1998

• hierax (Hewitson, 1869)

  • ssp. hierax (Hewitson, 1869)

    • syn. f. semibrunea Niepelt, 1923

  • ssp. leveri Attal, 1999

• xanthocles Bates, 1862

  • ssp. buechei Neukirchen, 1992

  • ssp. cleoxanthe Holzinger & Holzinger, 1972

  • ssp. donatia Fruhstorfer, 1910

    • syn. xanthocles melete f. meridionalis Neustetter, 1925

  • ssp. explicata Brown, 1976

    • syn. xanthocles melete f. explicata Stichel, 1923

    • syn. xanthocles melete f. paranympha Stichel, 1923

  • ssp. hippocrene Holzinger & Brown, 1982

    • syn. hippocrene Stichel, 1906

  • ssp. melete Felder & Felder, 1865

    • syn. melittus f. fassli Neustetter, 1912

    • syn. cethosia Seitz, 1912

    • syn. xanthocles flavosia Kaye, 1920

    • syn. cethosia completa (Oberthür, 1920)

  • ssp. melior Staudinger, 1897

  • ssp. melittus Staudinger, 1897

  • ssp. napoensis Holzinger & Brown, 1982

    • syn. xanthocles melete f. latior Neustetter, 1932

  • ssp. paraplesius Bates, 1867

    • syn. olede Riffarth, 1901

  • ssp. quindecim Lamas, 1976

  • ssp. rindgei Holzinger & Brown, 1982

  • ssp. similatus Zikán, 1937

  • ssp. vala Staudinger, 1885

    • syn. caternaulti (Oberthür, 1902)

  • ssp. xanthocles Bates, 1862

  • ssp. zamora Holzinger & Brown, 1982

• astraea Staudinger, 1897

  • ssp. astraea Staudinger, 1897

  • ssp. rondonia Brown, 1973

• burneyi (Hübner [1831])

  • ssp. ada Neustetter, 1925

  • ssp. anjae Neukirchen, 1995

  • ssp. boliviensis Neukirchen, 1995

    • syn. burneyi jeanneae f. reducta Boullet & Le Cerf, 1909

  • ssp. burneyi (Hübner [1831])

    • syn. burneyi var. jeanneae Boullet & Le Cerf, 1909

    • syn. burneyi f. oberthüri Boullet & Le Cerf, 1909, preocc. (not Riffarth, 1903)

    • syn. burneyi f. roseni Krüger, 1928

  • ssp. catharinae Staudinger, 1885

    • syn. burneyi serpensis Kaye, 1919, hyb.

    • syn. burneyi f. catharinae ab. vitiosa Krüger, 1933

    • syn. burneyi f. miraculosa Kotzsch, 1936

  • ssp. huebneri Staudinger, 1897

  • ssp. jamesi Neukirchen, 1995

  • ssp. koenigi Neukirchen, 1995

  • ssp. lindigii Felder & Felder, 1865

  • ssp. mirtarosa Orellana, 2006. Type loc.: Finca Vuelta Larga, Sucre, Venezuela. Types: MIZA. Distr.: Sucre, Venezuela. Refs.: Orellana (2006): 71. Ety.: Mirta Rosa Borges de Orellana

  • ssp. skinneri Brown & Fernández, 1985

• egeria (Cramer [1775])

  • ssp. christiani Neukirchen, 1997

  • ssp. egeria (Cramer [1775])

    • syn. isaea (Hübner, 1816), repl. name Refs.: Hübner (1816): 12.

    • syn. ergatis (Godart, 1819), repl. name

    • syn. ab. clearista (Oberthür, 1923)

  • ssp. egerides Staudinger, 1897

  • ssp. homogena Bryk, 1953

    • syn. egeria-hyas f. asterope Zikán, 1937

  • ssp. hyas Weymer, 1883

  • ssp. keithbrowni Neukirchen, 1992

  • ssp. mariasibyllae Neukirchen, 1991

• wallacei Reakirt, 1866

  • ssp. araguaia Brown, 1976

  • ssp. colon Weymer, 1891

    • syn. clytia (Cramer [1776]), preocc. (not Linnaeus, 1758)

    • syn. clytia var. elsa Riffarth, 1899

    • syn. wallacei brevimaculata ab. halli Kaye, 1919, hyb.

  • ssp. flavescens Weymer, 1891

    • syn. sara (Hübner [1809]), not Fabricius, 1793

    • syn. clytia var. colon Weymer, 1891, hyb.

    • syn. clytia var. sulphurea Staudinger, 1897, preocc. (not Weymer, 1894)

    • syn. hagar Kirby [1900]

    • syn. f. quadrimaculata Neustetter, 1925

    • syn. clytia f. flavescens ab. graphitica Krüger, 1933

    • syn. f. inez Stammeshaus, 1982

  • ssp. kayei Neustetter, 1929, repl. name

    • syn. wallacei latus Kaye, 1925, preocc. (not Riffarth, 1900)

  • ssp. mimulinus Butler, 1873

  • ssp. wallacei Reakirt, 1866

    • syn. clytia ab. parvimaculata Riffarth, 1900, hyb.

    • syn. clytia f. flavescens ab. wucherpfennigi Krüger, 1933, hyb.

    • syn. ab. erichi Krüger, 1933, hyb.

• clysonymus Latreille [1817]

  • ssp. clysonymus Latreille [1817]

    • syn. clysonimus fischeri Fassl, 1912, hyb.

    • syn. clysonimus flavopunctatus Fassl, 1912, hyb.

    • syn. micrus Seitz, 1912

    • syn. clysonymus apicalis Joicey & Kaye, 1917, nom. nud.

    • syn. clysonymus apicalis ab. semirubra Joicey & Kaye, 1917, hyb.

    • syn. clysonymus perbellus Stichel, 1923

    • syn. hygianus fischeri f. leonis Holzinger & Holzinger, 1970, hyb.

    • syn. hygianus fischeri f. leoncitonis Holzinger & Holzinger, 1970, hyb.

  • ssp. hygiana (Hewitson, 1867)

    • syn. hygiana ab. albescens Kaye, 1916

  • ssp. montanus Salvin, 1871

  • ssp. tabaconas Brown, 1976

• erato (Linnaeus, 1758)

  • ssp. adana Turner, 1967, ICZN, Art. 45.6.2

    • syn. hydara ab. adana Seitz, 1913

    • syn. hydarus hydarus f. vitellina Stichel, 1919

  • ssp. amalfreda Riffarth, 1901

    • syn. elimaea (Erichson [1849]), hyb.

    • syn. cybele ab. cybelina Staudinger, 1897

    • syn. vesta ab. leda Staudinger, 1897, hyb.

    • syn. phyllis ab. amalfreda Riffarth, 1900

    • syn. erato cybelinus f. helena Riffarth, 1907, hyb.

    • syn. erato erato ab. cybelellus Joicey & Kaye, 1917, hyb.

    • syn. melpomene f. pyritosa Zikán, 1937

    • syn. melpomene-pyritosa var. fumigata Zikán, 1937

    • syn. erato andremona f. juanita Neustetter, 1938, hyb.

    • syn. f. heydei Stammeshaus, 1982

  • ssp. amazona Staudinger, 1897

    • syn. vesta (Hübner [1807]), not Cramer, 1777

    • syn. philadelphus Kirby [1900]

    • syn. androdaixa Seitz, 1912, hyb.

  • ssp. amphitrite Riffarth, 1901

    • syn. erato estrella f. simplex Riffarth, 1906, hyb.

    • syn. erato phyllis f. sperata Riffarth, 1907, hyb.

    • syn. melpomene hyperplea Dyar, 1913

    • syn. ab. unipuncta Joicey & Kaye, 1917

  • ssp. chestertonii (Hewitson, 1872)

    • syn. damysus Hopffer, 1874

    • syn. hydara ab. nocturna Riffarth, 1900, hyb.

  • ssp. colombina Staudinger, 1897

    • syn. hydara var. antigona Riffarth, 1900

  • ssp. cruentus Lamas, 1998

    • syn. erato punctata Beutelspacher, 1992, preocc. (not Neustetter, 1907)

  • ssp. cyrbia (Godart, 1819)

    • syn. cyrbia ab. diformata Riffarth, 1900, hyb.

    • syn. cyrbia cyrbia f. bella Riffarth, 1907

  • ssp. demophoon Ménétriés, 1855

    • syn. chiriquensis Riffarth, 1900, nom. nud.

  • ssp. dignus Stichel, 1923

    • syn. dignus f. discerpta Stichel, 1923, hyb.

    • syn. erato estrella f. problemata Neustetter, 1928, hyb.

    • syn. estrella f. meliorina Neustetter, 1928, hyb.

    • syn. estrella f. glaucina Neustetter, 1928, hyb.

    • syn. dignus f. elvira Neustetter, 1928, hyb.

  • ssp. emma Riffarth, 1901

    • syn. augusta Riffarth, 1901, nom. nud.

    • syn. erato estrella f. palmata Stichel, 1906, hyb.

    • syn. erato estrella f. agnata Stichel, 1906, hyb.

  • ssp. erato (Linnaeus, 1758)

    • syn. vesta (Cramer [1777]), hyb.

    • syn. erythrea (Cramer [1777]), hyb.

    • syn. andremona (Cramer [1780]), hyb.

    • syn. udalrica (Cramer [1780]), hyb.

    • syn. ulrica (Hübner, 1816), missp. Refs.: Hübner (1816): 12.

    • syn. cynisca (Godart, 1819)

    • syn. vesta tellus (Oberthür, 1902), hyb.

    • syn. erato oberthürii Riffarth, 1903, hyb.

    • syn. f. fuliginosa Riffarth, 1907

    • syn. ab. hemicycla Joicey & Kaye, 1917, hyb.

    • syn. ab. protea Joicey & Kaye, 1917, hyb.

    • syn. ab. constricta Joicey & Kaye, 1917, hyb.

    • syn. f. albida Joicey & Kaye, 1919

    • syn. ab. telloides Joicey & Kaye, 1925, hyb.

    • syn. f. roseoflava Neustetter, 1926, hyb.

    • syn. f. fumata Neustetter, 1926, hyb.

    • syn. f. latiflava Neustetter, 1931, hyb.

    • syn. f. nigrobasalis Neustetter, 1931, hyb.

  • ssp. estrella Bates, 1862

  • ssp. etylus Salvin, 1871

    • syn. ab. insignis Kaye, 1916, hyb.

  • ssp. favorinus Hopffer, 1874

    • syn. amaryllis var.? pseudamaryllis Staudinger, 1897

    • syn. erato eratophylla Joicey & Kaye, 1917, hyb.

    • syn. favorinus pseudoanacreon Neustetter, 1932, hyb.

  • ssp. guarica Reakirt, 1868

    • syn. guayana Herrich-Schäffer, 1865, nom. nud.

    • syn. euryas (Boisduval, 1870)

  • ssp. hydara (Hewitson, 1867)

    • syn. callycopis (Cramer [1777]), hyb.

    • syn. carolina (Herbst, 1790), artefact

    • syn. coralii Butler, 1877, hyb.

    • syn. erato f. palantia Möschler, 1883, hyb.

    • syn. cyrbia var. juno Riffarth, 1900, hyb.

    • syn. petiverana ab. tristis Riffarth, 1900, hyb.

    • syn. phyllis var. viculata Riffarth, 1900, hyb.

    • syn. phyllis ab. dryope Riffarth, 1900, hyb.

    • syn. phyllis ab. callista Riffarth, 1900, hyb.

    • syn. palantes Riffarth, 1901, nom. nud.

    • syn. erato callycopis f. belticopis Joicey & Kaye, 1917, hyb.

    • syn. erato rubrizona Joicey & Kaye, 1917, hyb.

  • ssp. lativitta Butler, 1877

    • syn. vesta var. vestalis Staudinger, 1885

    • syn. erato estrella f. ochracea Riffarth, 1907, hyb.

    • syn. erato estrella f. feyeri Niepelt, 1908, hyb.

    • syn. erato estrella f. anactorina Stichel, 1923, hyb.

    • syn. erato estrella f. sanguinella Stichel, 1923, hyb.

    • syn. erato estrella f. perplexa Stichel, 1923, hyb.

    • syn. microclea beata f. zoraida Neustetter, 1925, hyb.

    • syn. erato estrella f. aurivillii Bryk, 1953, hyb.

  • ssp. lichyi Brown & Fernández, 1985

  • ssp. luscombei Lamas, 1976

  • ssp. magnifica Riffarth, 1900

  • ssp. microclea Kaye, 1907

    • syn. f. microfluens Descimon & Mast, 1971, hyb.

  • ssp. notabilis Salvin & Godman, 1868

    • syn. erato estrella f. beata Riffarth, 1907, hyb.

    • syn. erato simplex f. rosacea Riffarth, 1907, hyb.

    • syn. erato estrella f. ilia Niepelt, 1908, hyb.

    • syn. erato estrella f. rothschildi Niepelt, 1909, hyb.

    • syn. microclea notabilis radiata (Oberthür, 1916), hyb.

    • syn. notabilis rubrescens Röber, 1921, hyb.

    • syn. microclea notabilis f. flavopunctatus Neustetter, 1925, hyb.

  • ssp. petiverana (Doubleday, 1847). Type loc.: “Mexico, Honduras”; syntype from “Mexico”. Types: Syntype in NHMUK. Refs.: Doubleday, 1847, In: Gen. diurn. Lepid. 1(10): 103, no. 18; cited Petiver (1702), Gazophylacium: pl. 4, fig. 2.

    • syn. mexicana (Boisduval, 1870). Type loc.: “Mexique et à Honduras”. Types: “Petiver’s figure might be considered to represent a syntype; if extant, the specimen(s) upon which the figure was based might be in NHMUK or MNHP” (Pelham). Refs.: Boisduval, 1870. Consid. Lépid. Guat.: 28; cited Heliconia demophoon Ménétriés, 1855, Enum. corp. anim. (1): 86, no. 331-332, pl. 2, fig. 4, and Petiver (1702), Gazophylacium: pl. 4, fig. 2.

    • syn. petiverea Riffarth, 1901, nom. nud.

  • ssp. phyllis (Fabricius, 1775)

    • syn. roxane (Cramer [1775])

    • syn. phyllidis Grose-Smith & Kirby, 1892

    • syn. anacreon Grose-Smith & Kirby, 1892, hyb.

    • syn. amatus Staudinger, 1897, hyb.

    • syn. phyllis ab. artifex Stichel, 1899

    • syn. f. diffluens Riffarth, 1907

    • syn. erato anacreon f. anaitis Riffarth, 1907, hyb.

    • syn. f. athene Neustetter, 1909

    • syn. phyllis ab. miletus d’Almeida, 1928

    • syn. phyllis ab. cohaerens Hayward, 1931

    • syn. phyllis ab. alicia Schweizer & Kay, 1941

  • ssp. reductimacula Bryk, 1953

    • syn. f. venusta Bryk, 1953

  • ssp. tobagoensis Barcant, 1982

  • ssp. venus Staudinger, 1882

    • syn. molina Grose-Smith, 1898, hyb.

    • syn. erato f. extrema Kaye, 1919, nom. nud.

    • syn. erato extrema Kaye, 1919, hyb.

  • ssp. venustus Salvin, 1871

    • syn. anactorie (Doubleday, 1847), hyb.

    • syn. anactorie ab. sanguineus Staudinger, 1894, hyb.

    • syn. phyllis ab. ottonis Riffarth, 1900, hyb.

    • syn. locris Riffarth, 1901, nom. nud.

    • syn. erato diva Stichel, 1906, hyb.

    • syn. phyllis ab. confluens Seitz, 1913, hyb.

    • syn. phyllis f. anacreonides Neustetter, 1925, hyb.

    • syn. phyllis f. krügeri Neustetter, 1925, hyb.

    • syn. phyllis f. flavomixta Neustetter, 1925, hyb.

    • syn. phyllis f. leonora Krüger, 1927, hyb.

    • syn. phyllis anacreon f. clelia Neustetter, 1927, hyb.

    • syn. phyllis anactorie f. henrici Krüger, 1929, hyb.

    • syn. phyllis venustus f. pseudoleda Neustetter, 1932, hyb.

    • syn. xanthocles f. denieri Hayward, 1939

• hecalesia (Hewitson [1854])

  • ssp. eximius Stichel, 1923

  • ssp. formosus Bates, 1866

  • ssp. gynaesia (Hewitson, 1875)

  • ssp. hecalesia (Hewitson [1854])

  • ssp. longarena (Hewitson, 1875)

    • syn. hecalesia ernestus Brown & Benson, 1975, hyb.

  • ssp. octavia Bates, 1866

  • ssp. romeroi Brown & Fernández, 1985

• hermathena (Hewitson [1854]) Refs.: Brown Jr. and Benson (1977).

  • ssp. curua Freitas & Ramos, 2019. Type loc.: Salto do Curuá, Altamira, PA, Brazil. Types: Museu de Zoologia da Universidade Estadual de Campinas (ZUEC), Campinas, São Paulo, Brazil. Host plants: Passiflora hexagonocarpa (Freitas et al. 2019). Refs.: Freitas et al. (2019): 469. Ety.: Salto do Curuá (Curua Waterfalls)

  • ssp. duckei Brown & Benson, 1977. Type loc.: Bazil, Para: scrub along northeastern inlet at bend of Rio Nhamund’a, 20 km due north of Faro. Types: Museu Nacional, Rio de Janeiro; paratypes: NHMUK. Refs.: Brown Jr. and Benson (1977): 101. Ety.: Adolph Ducke (botanist)

    • syn. hermathena f. rubropunctata d’Abrera, 1984, nom. nud.

  • ssp. hermathena (Hewitson [1854]) Host plants: Passiflora hexagonocarpa (Brown Jr. and Benson 1977).

  • ssp. renatae Brown & Fernández, 1985

  • ssp. sabinae Neukirchen, 1992

  • ssp. sheppardi Brown & Benson, 1977. Type loc.: Bazil: Amazonas, Reserva Campina of the Instituto Nacional de Pesquisas da Amazonia, on the Manaus-Caracarai highway 45 km north of the Pollcia Rodoviari. Types: Museu Nacional, Rio de Janeiro; paratypes: NHMUK, ZUEC. Host plants: Passiflora faroana (Brown Jr. and Benson 1977). Refs.: Brown Jr. and Benson (1977): 99. Ety.: Dr. and Mrs. Philip M. Sheppard

    • syn. hermathena sheppardorum Lamas, 1988, emend.

  • ssp. vereatta Stichel, 1912

    • syn. hermathena f. hydarina Stichel, 1912

• himera (Hewitson, 1867)

• hortense (Guérin-Méneville [1844])

• telesiphe (Doubleday, 1847)

  • ssp. cretacea Neustetter, 1916

    • syn. telesiphe ab. nivea Kaye, 1916

  • ssp. sotericus O. Salvin, 1871

  • ssp. telesiphe (Doubleday, 1847)

• atthis (Doubleday, 1847)

  • syn. bourcieri (Becker, 1851), nom. nud.

  • syn. atthis f. gerstneriana Descimon & Mast, 1971.

• besckei (Ménétriés, 1857)

  • syn. besckii (Ménétriés, 1857), missp.

  • syn. epiphyllis Riffarth, 1901, nom. nud.

  • syn. besckei principalis Röber, 1921.

  • syn. besckei f. intermixta Neustetter, 1931.

• cydno (Doubleday, 1847)

  • ssp. alithea (Hewitson, 1869)

    • syn. cydno ab. haenschi Riffarth, 1900

    • syn. cydno broncus Stichel, 1906, hyb.

    • syn. f. egregia Riffarth, 1907

    • syn. alithea (var.?) ab. minor Neustetter, 1907. Type loc.: Llanos, Ecuador. Refs.: Neustetter (1907): (184).

    • syn. alithea f. neustetteri Riffarth, 1908

  • ssp. barinasensis Masters, 1973

  • ssp. chioneus Bates, 1864

  • ssp. cordula Neustetter, 1913

    • syn. cydno ab. larseni Niepelt, 1916

    • syn. cydno flaveola Joicey & Kaye, 1917, hyb.

    • syn. cydno perijaensis Masters, 1973

  • ssp. cydnides Staudinger, 1885

    • syn. cydno var. epicydnides Staudinger, 1897

    • syn. cydno var. subcydnides Staudinger, 1897

    • syn. interrupta Riffarth, 1901, nom. nud.

    • syn. f. werneri Neustetter, 1928

    • syn. f. azteka Neustetter, 1928

    • syn. f. semicydnides Holzinger & Holzinger, 1968

    • syn. f. tenebrosa Holzinger & Holzinger, 1968

  • ssp. cydno (Doubleday, 1847)

    • syn. cydno ab. mediocydno Neustetter, 1907. Type loc.: Colombia. Refs.: Neustetter (1907): (183).

    • syn. cydno f. punctata Neustetter, 1907. Type loc.: Colombia. Refs.: Neustetter (1907): (184).

  • ssp. gadouae Brown & Fernández, 1985

    • syn. hahneli Staudinger, 1885, hyb.

  • ssp. galanthus ates, 1864

    • syn. diotrephes (Hewitson, 1869)

    • syn. cydno ab. stübeli Riffarth, 1900

    • syn. piera Riffarth, 1901, nom. nud.

    • syn. galanthus f. exornata Riffarth, 1907, hyb.

    • syn. galanthus ab. subrufescens Schaus, 1913, hyb.

  • ssp. hermogenes (Hewitson [1858])

    • syn. temerinda (Hewitson, 1873)

    • syn. hermogenes ab. lutescens Kaye, 1916

    • syn. f. anthelea Neustetter, 1925

    • syn. cydno dolores Apolinar, 1926

  • ssp. lisethae Neukirchen, 1995. Type loc.: Colombia: Huila. Types: Zoologischen Museum der Humboidt-Universitat, Berlin. Refs.: Neukirchen (1955): 190.

  • ssp. wanningeri Neukirchen, 1991

  • ssp. weymeri Staudinger, 1897

    • syn. weymeri ab. gustavi Staudinger, 1897

    • syn. weymeri submarginatus Fassl, 1912

    • syn. submarginalis Seitz, 1912, missp.

    • syn. weymeri ab. sulphureomaculata Fassl, 1914

    • syn. cydno gustavi ab. flavomaculata Seitz, 1916

    • syn. hermogenes ab. leucosticta Apolinar, 1926

    • syn. hermogenes ab. xanthosticta Apolinar, 1926

    • syn. cydno gerstneri Holzinger & Holzinger, 1968, hyb.

    • syn. cydno gerstneri f. pseudoweymeri Holzinger & Holzinger, 1968, hyb.

    • syn. cydno gerstneri f. denhezi Holzinger & Holzinger, 1968, hyb.

    • syn. cydno gerstneri f. flavissima Holzinger & Holzinger, 1968, hyb.

  • ssp. zelinde Butler, 1869

    • syn. f. inca Neustetter, 1928

    • syn. cydno cydnides f. flavidior Neustetter, 1928, hyb.

    • syn. cydno cydnides f. confluens Neustetter, 1928, hyb.

    • syn. cydno cydnides f. albidior Neustetter, 1928, hyb.

• elevatus Nöldner, 1901

  • ssp. bari (Oberthür, 1902)

  • ssp. elevatus Nöldner, 1901

    • syn. f. griseoviridis Neustetter, 1938

    • syn. elevatus schmidt-mummi Takahashi, 1977

    • syn. elevatus willmotti Neukirchen, 1997

  • ssp. jigginsi Costa & Neild, 2023. Type loc.: Venezuela: Auyán Tepui. Types: MIZA. Distr.: Southern slopes of Auyán Tepui, Venezuela. Refs.: Costa et al. (2023). Ety.: Chris Jiggins

  • ssp. lapis Lamas, 1976

  • ssp. perchlora Joicey & Kaye, 1917

  • ssp. pseudocupidineus Neustetter, 1931

    • syn. elevatus f. nigromacula Neustetter, 1932

    • syn. elevatus f. nöldneri Neustetter, 1938

  • ssp. roraima Turner, 1966

  • ssp. schmassmanni Joicey & Talbot, 1925

    • syn. melpomene thelxiope f. aquilina Neustetter, 1925

  • ssp. sonjae Neukirchen, 1997

  • ssp. taracuanus Bryk, 1953

  • ssp. tumatumari Kaye, 1906

  • ssp. willmotti Neukirchen, 1997

  • ssp. zoelleri Neukirchen, 1990

• ethilla (Godart, 1819)

  • ssp. adela Neustetter, 1912

  • ssp. aerotome Felder & Felder, 1862

  • ssp. cephallenia Felder & Felder, 1865

  • ssp. chapadensis Brown, 1973

  • ssp. claudia Godman & Salvin, 1881

    • syn. assimilis Röber, 1921

  • ssp. ethilla (Godart, 1819)

    • syn. metalilis var. flavidus Weymer, 1894

    • syn. ethilla metalilis f. depuncta Boullet & Le Cerf, 1909

  • ssp. eucoma (Hübner [1831])

    • syn. eucoma (Hübner, 1816), nom. nud. Refs.: Hübner (1816): 11.

  • ssp. flavofasciatus Weymer, 1894

  • ssp. flavomaculatus Weymer, 1894

  • ssp. hyalina Neustetter, 1928

  • ssp. jaruensis Brown, 1976

  • ssp. latona Neukirchen, 1998

  • ssp. mentor Weymer, 1883

    • syn. orchamus Weymer, 1912

  • ssp. metalilis Butler, 1873

  • ssp. michaelianius Lamas, 1988, repl. name

    • syn. aurotome f. clarus Michael, 1926, preocc. (not Fabricius, 1793)

  • ssp. narcaea (Godart, 1819)

    • syn. eucrate (Hübner [1823])

    • syn. satis Weymer, 1875

    • syn. eucrate var. infuscata Staudinger, 1885

    • syn. narcaea ab. brunnescens Neustetter, 1907. Type loc.: Rio de Janeiro, Brazil. Types: Wiener Hofmuseum. Refs.: Neustetter (1907): (180).

    • syn. connexa Seitz, 1912

  • ssp. nebulosa Kaye, 1916

  • ssp. neukircheni Lamas, 1998

  • ssp. numismaticus Weymer, 1894

  • ssp. penthesilea Neukirchen, 1994

  • ssp. polychrous Felder & Felder, 1865

    • syn. physcoa Seitz, 1912

  • ssp. semiflavidus Weymer, 1894

    • syn. eucoma var. daguanus Staudinger, 1897

    • syn. eucoma var. juntana Riffarth, 1900

  • ssp. thielei Riffarth, 1900

    • syn. numatus isabellinus ab. fusca Boullet & Le Cerf, 1909

  • ssp. tyndarus Weymer, 1897

  • ssp. yuruani Brown & Fernández, 1985

• hecale (Fabricius, 1776)

  • ssp. anderida (Hewitson [1853])

    • syn. zagora Riffarth, 1901, nom. nud.

    • syn. rebeli Neustetter, 1907. Type loc.: Bogota, Colombia. Types: Wiener Hofmuseum. Distr.: Colombia. Refs.: Neustetter (1907): (182).

    • syn. anderida estebana Kaye, 1914

  • ssp. annetta Riffarth, 1900

  • ssp. australis Brown, 1976

  • ssp. barcanti Brown, 1976

  • ssp. clearei Hall, 1930

  • ssp. discomaculatus Weymer, 1891

  • ssp. ennius Weymer, 1891

  • ssp. felix Weymer, 1894

    • syn. felix var. concors Weymer, 1894

    • syn. versicolor Weymer, 1894

    • syn. quitalena felix f. umbrina Neustetter, 1931, hyb.

  • ssp. fornarina (Hewitson, 1854)

    • syn. anderida fornarina f. bouvieri Boullet & Le Cerf, 1909

    • syn. anderida f. styx Niepelt, 1921

  • ssp. fraternitas Brévignon, 2006. Type loc.: French Guiana. Refs.: Brevignon (2006).

  • ssp. hecale (Fabricius, 1776)

    • syn. pasithoë (Cramer [1775]), preocc. (not Linnaeus, 1767)

    • syn. pasithoë var. fulvescens Lathy, 1906

  • ssp. holcophorus Staudinger, 1897

    • syn. metaphorus var. semiphorus Staudinger, 1897

    • syn. eucherius Weymer, 1906

  • ssp. humboldti Neustetter, 1928

    • syn. humboldti f. alexander Neustetter, 1928

  • ssp. ithaca Felder & Felder, 1862

    • syn. vittatus Butler, 1873

    • syn. marius Weymer, 1890

    • syn. ithaka f. cajetani Neustetter, 1909

    • syn. ithaca f. hero Weymer, 1912

    • syn. aristiona indecisa Joicey & Kaye, 1917

    • syn. ithaka ithaka f. sulphureofasciata Neustetter, 1925

    • syn. ithaka vittatus f. nigroapicalis Neustetter, 1925

  • ssp. latus Riffarth, 1900

    • syn. gradatus xinguensis Neustetter, 1925

  • ssp. melicerta Bates, 1866

    • syn. clara (Fabricius, 1793), preocc. (not Cramer, 1775)

    • syn. albucilla Bates, 1866

    • syn. etholea Riffarth, 1901, nom. nud.

    • syn. anderida melicerta f. zygia Riffarth, 1907

    • syn. anderida melicerta f. muzoënsis Neustetter, 1909

  • ssp. metellus Weymer, 1894

    • syn. vetustus boyi Röber, 1923

  • ssp. naxos Neukirchen, 1998

  • ssp. nigrofasciatus Weymer, 1894

  • ssp. novatus Bates, 1867

    • syn. schulzi Riffarth, 1899

    • syn. schultzi Seitz, 1924, emend.

  • ssp. paraensis Riffarth, 1900

  • ssp. paulus Neukirchen, 1998

  • ssp. quitalena (Hewitson [1853])

  • ssp. rosalesi Brown & Fernández, 1976

  • ssp. shanki Lamas & Brown, 1976

  • ssp. sisyphus Salvin, 1871

    • syn. jonas Weymer, 1894

  • ssp. sulphureus Weymer, 1894

    • syn. quitalenus denticulatus Riffarth, 1907

    • syn. sulphureus subsulphureus Röber, 1919, nom. dub.

  • ssp. vetustus Butler, 1873

    • syn. clarissa Riffarth, 1901, nom. nud.

  • ssp. zeus Neukirchen, 1995

    • syn. felix var. concors Weymer, 1894 in part

  • ssp. zuleika (Hewitson, 1854)

    • syn. jucundus Bates, 1864

    • syn. xanthicus Bates, 1864

    • syn. chrysantis Godman & Salvin, 1881

    • syn. zuleika ab. albipunctata Riffarth, 1900

    • syn. zuleika ab. dentata Neustetter, 1907. Type loc.: Panama. Refs.: Neustetter (1907): (183).

• heurippa (Hewitson [1854])

• ismenius Latreille [1817]

  • ssp. boulleti Neustetter, 1928

    • syn. sylvana sylvana f. sticheli Boullet & Le Cerf, 1909

  • ssp. clarescens Butler, 1875

  • ssp. fasciatus Godman & Salvin, 1877

  • ssp. ismenius Latreille [1817]

    • syn. fritscheï (Möschler, 1872)

    • syn. ocanna Buchecker [1880]

    • syn. faunus Staudinger, 1885

    • syn. faunus var. antioquensis Staudinger, 1885

    • syn. var. (ab.?) hermanni Riffarth, 1899

    • syn. distincta Riffarth, 1901, nom. nud.

    • syn. f. immoderata Stichel, 1906

    • syn. ab. albofasciatus Neustetter, 1907. Type loc.: Bogota, Colombia. Refs.: Neustetter (1907): (181).

    • syn. f. defasciatus Neustetter, 1909

    • syn. ismenius abadiae Apolinar, 1926

  • ssp. metaphorus Weymer, 1883

    • syn. catilina Riffarth, 1901, nom. nud.

    • syn. ismenius faunus f. hoppi Neustetter, 1928, hyb.

  • ssp. occidentalis Neustetter, 1928

  • ssp. telchinia (Doubleday, 1847)

  • ssp. tilletti Brown & Fernández, 1976

• luciana Lichy, 1970

  • ssp. luciana Lichy, 1970

  • ssp. watunna Lichy, 1960

• melpomene (Linnaeus, 1758)

  • ssp. aglaope Felder & Felder, 1862

    • syn. melpomene ab. mirabilis Riffarth, 1900, hyb.

    • syn. melpomene riffarthi Stichel, 1906, hyb.

    • syn. melpomene riffarthi f. rubescens Stichel, 1906, hyb.

    • syn. f. cognata Riffarth, 1907

    • syn. aglaope f. flavotenuiata Neustetter, 1931

    • syn. aglaope f. magnimacula Neustetter, 1932, hyb.

    • syn. aglaope f. daira Neustetter, 1932, hyb.

  • ssp. amandus Grose-Smith & Kirby, 1892

    • syn. penelope ab. pelopeia Staudinger, 1894, hyb.

    • syn. penelope ab. penelamanda Staudinger, 1894, hyb.

    • syn. aphrodyte Staudinger, 1897, hyb.

    • syn. penelope ab. penelopeia Staudinger, 1897, emend.

    • syn. phyllis f. flammea Niepelt, 1925, hyb.

    • syn. penelope f. biedermanni Niepelt, 1926, hyb.

    • syn. penelope f. amandoides Neustetter, 1926, hyb.

    • syn. penelope f. carnea Neustetter, 1926, hyb.

  • ssp. amaryllis Felder & Felder, 1862

    • syn. amaryllis ab. pseudo-penelamanda Michael, 1912, hyb.

    • syn. melpomene melpophylla Joicey & Kaye, 1917, hyb.

  • ssp. anduzei Brown & Fernández, 1985

  • ssp. bellula Turner, 1971, ICZN, Art. 45.5.1

    • syn. amaryllis amaryllis f. bellula Stichel, 1923

    • syn. amaryllis amaryllis f. permira Stichel, 1923, hyb.

    • syn. amaryllis amaryllis f. degener Stichel, 1923, hyb.

    • syn. amaryllis amaryllis anacreontica Stichel, 1923, hyb.

    • syn. amaryllis amaryllis f. perrara Stichel, 1923, hyb.

    • syn. melpomene aglaope f. parva Neustetter, 1928, hyb.

    • syn. melpomene mocoa Brower, 1996

  • ssp. burchelli Poulton, 1910

    • syn. f. curvifascia Talbot, 1928, hyb.

  • ssp. cythera (Hewitson, 1869)

    • syn. vulcanus ab. modesta Riffarth, 1900, hyb.

    • syn. hypna Riffarth, 1901, nom. nud.

    • syn. vulcanus sticheli Riffarth, 1907, hyb.

    • syn. vulcanus tenuistriga Kaye, 1920

  • ssp. ecuadorensis Emsley, 1964, ICZN, Art. 45.5.1

    • syn. melpomene aglaope f. ecuadorensis Neustetter, 1909

    • syn. equatorensis Seitz, 1912, missp.

    • syn. melpomene equadoriensis Seitz, 1913, missp.

  • ssp. euryades Riffarth, 1900

    • syn. amaryllis euryades f. rubrica Stichel, 1919, hyb.

  • ssp. flagrans Stichel, 1919

  • ssp. intersectus Neustetter, 1928

  • ssp. madeira Riley, 1919

  • ssp. malleti Lamas, 1988

    • syn. melpomene aglaope f. iris Riffarth, 1907, hyb.

    • syn. melpomene aglaope f. isolda Niepelt, 1908, hyb.

    • syn. melpomene aglaope f. adonides Niepelt, 1908, hyb.

    • syn. melpomene aglaope f. rubripicta Niepelt, 1908, hyb.

    • syn. melpomene aglaope f. gisela Niepelt, 1908, hyb.

    • syn. melpomene aglaope f. anna Neustetter, 1909, hyb.

    • syn. melpomene aglaope f. dione Neustetter, 1909, hyb.

    • syn. melpomene aglaope f. gratiosa Niepelt, 1909, hyb.

    • syn. amaryllis amaryllis f. rufata Stichel, 1923, hyb.

    • syn. amaryllis amaryllis f. aglaspis Stichel, 1923, hyb.

    • syn. melpomene aglaope f. aurofasciata Neustetter, 1928, hyb.

    • syn. melpomene aglaope f. paula Neustetter, 1928, hyb.

    • syn. aglaope f. paulina Niepelt, 1928, hyb.

    • syn. aglaope f. carminata Niepelt, 1928, hyb.

  • ssp. martinae Cast & Le Crom, 2012. Type loc.: Otanche, Boyaca, Colombia. Types: MNHN. Distr.: Colombia. Refs.: Cast and Le Crom (2012).

    • syn. melpomene martinae f. flavomartina Cast & Le Crom, 2012

    • syn. melpomene martinae f. mixta Cast & Le Crom, 2012

  • ssp. melpomene (Linnaeus, 1758)

    • syn. lucia (Stoll, 1781), hyb.

    • syn. var. tyche Bates, 1862, hyb.

    • syn. var. hippolyte Bates, 1862, hyb.

    • syn. rufolimbatus Butler, 1873, hyb.

    • syn. mutabilis Butler, 1877, hyb.

    • syn. funebris Möschler, 1877, hyb.

    • syn. amor Staudinger, 1885, hyb.

    • syn. timareta ab.? erebia Riffarth, 1900, hyb.

    • syn. ab. atrosecta Riffarth, 1900

    • syn. ab. lucinda Riffarth, 1900, hyb.

    • syn. ab. melpomenides Riffarth, 1900, hyb.

    • syn. ab. karschi Riffarth, 1900, hyb.

    • syn. var. melanippe Riffarth, 1900, hyb.

    • syn. ab. diana Riffarth, 1900, hyb.

    • syn. gaea Riffarth, 1901, nom. nud.

    • syn. jussa Riffarth, 1901, nom. nud.

    • syn. justina Riffarth, 1901, nom. nud.

    • syn. ab. collis Joicey & Kaye, 1917, hyb.

    • syn. ab. primus Joicey & Kaye, 1917, hyb.

    • syn. ab. melpina Joicey & Kaye, 1917, hyb.

    • syn. melpomene cybele ab. dianides Joicey & Kaye, 1917, hyb.

    • syn. melpomene cybele ab. elegantula Joicey & Kaye, 1971, hyb.

    • syn. melpomene cybele ab. maris Joicey & Kaye, 1917, hyb.

    • syn. melpomene eltringhami Joicey & Kaye, 1917, hyb.

    • syn. ab. compacta Joicey & Kaye, 1919, hyb.

    • syn. ab. faivrei Joicey & Kaye, 1919, hyb.

    • syn. f. flavorubra Neustetter, 1926

    • syn. f. lydia Neustetter, 1926, hyb.

    • syn. f. luteipicta Neustetter, 1926, hyb.

    • syn. f. aurelia Neustetter, 1926, hyb.

    • syn. f. bang-haasi Neustetter, 1926, hyb.

    • syn. f. rubroflammea Neustetter, 1927, hyb.

    • syn. f. laurentina Neustetter, 1927, hyb.

    • syn. f. rufolinea Neustetter, 1928, hyb.

    • syn. f. trimacula Neustetter, 1931, hyb.

    • syn. melpomene melpomene alabanda Lever, Chazal & Lacomme, 2008, hyb. Refs.: Lever et al. (2008).

  • ssp. meriana Turner, 1967, repl. name

    • syn. cybele (Cramer [1777]), preocc. (not Fabricius, 1775)

    • syn. deinia Möschler, 1877, hyb.

    • syn. faustina Staudinger, 1885, hyb.

    • syn. melpomene ab. eulalia Riffarth, 1900, hyb.

    • syn. melpomene funebris f. obscurata Riffarth, 1907

    • syn. fascinatrix Seitz, 1912, hyb.

    • syn. melpomene cybele ab. faustalia Joicey & Kaye, 1917, hyb.

    • syn. melpomene cybele ab. negroida Joicey & Kaye, 1917, hyb.

    • syn. melpomene cybele ab. cybeleia Joicey & Kaye, 1919, hyb.

    • syn. melpomene f. lavinia Neustetter, 1926, hyb.

    • syn. f. alba Stammeshaus, 1982

    • syn. melpomene meriana novalis Lever, Chazal & Lacomme, 2008, hyb.

    • syn. melpomene meriana penthesilea Lever, Chazal & Lacomme, 2008, hyb.

    • syn. melpomene meriana hyperion Lever, Chazal & Lacomme, 2008, hyb.

  • ssp. michellae Neukirchen, 1997

  • ssp. nanna Stichel, 1899

    • syn. bidentatus Riffarth, 1901, nom. nud.

    • syn. mayi d’Almeida, 1928

  • ssp. penelope Staudinger, 1894

    • syn. penelope ab. pluto Staudinger, 1897, hyb.

    • syn. melpomene ab. margarita Riffarth, 1900, hyb.

    • syn. f. praxedis Neustetter, 1925, hyb.

    • syn. penelope f. noctis Neustetter, 1926, hyb.

    • syn. penelope f. excellens Neustetter, 1926, hyb.

    • syn. penelope f. aida Neustetter, 1926, hyb.

    • syn. penelope f. amneris Neustetter, 1926, hyb.

    • syn. penelope f. rufofascia Neustetter, 1926, hyb.

    • syn. melpomene burchelli f. obscurifascia Talbot, 1928, hyb.

    • syn. penelope f. flavodiscalis Neustetter, 1931, hyb.

  • ssp. plesseni Riffarth, 1907

    • syn. unimaculata (Hewitson, 1869), hyb.

    • syn. radiatus Riffarth, 1901, nom. nud.

    • syn. batesi plesseni f. rubicunda Niepelt, 1907

    • syn. batesi plesseni f. pura Niepelt, 1907

    • syn. melpomene aglaope f. niepelti Riffarth, 1907, hyb.

    • syn. batesi plesseni f. adonis Riffarth, 1907, hyb.

    • syn. batesi plesseni f. diadema Niepelt, 1908, hyb.

    • syn. melpomene aglaope f. fraterna Niepelt, 1909

    • syn. microclea virgo (Oberthür, 1916)

    • syn. xenoclea plesseni f. dido Neustetter, 1925, hyb.

    • syn. xenoclea plesseni f. clytie Neustetter, 1927, hyb.

    • syn. xenoclea plesseni f. mimetica Neustetter, 1928, hyb.

  • ssp. pyrforus Kaye, 1907

    • syn. vulcanus f. immarginata Neustetter, 1938

  • ssp. rosina (Boisduval, 1870)

  • ssp. schunkei Lamas, 1976

    • syn. melpomene aglaope ab. rubra Stichel, 1906

    • syn. aglaope var. incarnata Stichel, 1906, nom. nud.

  • ssp. tessa Barcant, 1982

  • ssp. thelxiope (Hübner [1806]) Refs.: Hübner (1806): pl. [12].

  • ssp. thelxiopeia Staudinger, 1897

    • syn. thelxiope var. aglaopeia Staudinger, 1897

    • syn. melpomene var. augusta Riffarth, 1900, hyb.

    • syn. judith Riffarth, 1901, nom. nud.

    • syn. milesia Riffarth, 1901, nom. nud.

    • syn. melpomene thelxiope ab. punctarius Joicey & Kaye, 1917, hyb.

    • syn. melpomene thelxiope ab. lucindella Joicey & Kaye, 1917, hyb.

    • syn. melpomene thelxiope ab. majestica Joicey & Kaye, 1917, hyb.

    • syn. melpomene thelxiope ab. stygianus Joicey & Kaye, 1917, hyb.

    • syn. melpomene thelxiope ab. negroidens Joicey & Kaye, 1917, hyb.

    • syn. melpomene f. athalia Neustetter, 1927, hyb.

    • syn. melpomene f. penelopides Neustetter, 1927, hyb.

    • syn. melpomene f. nigrointerrupta Neustetter, 1931, hyb.

    • syn. melpomene thelxiopeia diotima Lever, Chazal & Lacomme, 2008, hyb.

  • ssp. vicina Ménétriés, 1857

  • ssp. vulcanus Butler, 1865

    • syn. amaryllis rosina f. fumosa Neustetter, 1925, hyb.

  • ssp. xenoclea (Hewitson [1853])

    • syn. batesi Riffarth, 1900

    • syn. xenoclea var. superba Lathy, 1906, hyb.

    • syn. xenoclea var. confluens Lathy, 1906, hyb.

    • syn. xenoclea ab. zio Röber, 1919, hyb.

    • syn. f. integra Descimon & Mast, 1971, hyb.

• nattereri Felder & Felder, 1865 Distr.: Brazil: Espírio Santo to Bahia (Atlantic Forest). Refs.: Brown (1970); Brown (1972); Cardoso et al. (2017) (distr.; eastern Brazil).

  • syn. fruhstorferi Riffarth, 1899.

• numata (Cramer [1780])

  • ssp. arcuella Druce, 1874

    • syn. seraphion Weymer, 1894, hyb.

    • syn. numatus praelautus Stichel, 1906

    • syn. aristiona aurora f. deflavata Neustetter, 1932, hyb.

  • ssp. aristiona (Hewitson [1853])

    • syn. aristiona (Doubleday, 1847), nom. nud.

    • syn. aristiona var. peruana Hopffer, 1879

    • syn. aristiona var. splendidus Weymer, 1894, hyb.

  • ssp. aulicus Weymer, 1883

  • ssp. aurora Bates, 1862

    • syn. gordius Weymer, 1894, hyb.

    • syn. aristiona aurora f. michaeli Neustetter, 1931

    • syn. aristiona idalion f. excelsa Neustetter, 1932, hyb.

  • ssp. bicoloratus Butler, 1873

    • syn. bicoloratus var. phalaris Weymer, 1894, hyb.

  • ssp. elegans Weymer, 1894

    • syn. floridus Weymer, 1894, hyb.

  • ssp. ethra (Hübner [1831])

    • syn. dryalus Hopffer, 1869, repl. name

    • syn. ethra ab. brasiliensis Neustetter, 1907. Type loc.: Esp. Santo, Brazil. Refs.: Neustetter (1907): (180).

    • syn. ethra var.? hopfferi Neustetter, 1907. Type loc.: Brazil. Refs.: Neustetter (1907): (181).

  • ssp. euphone Felder & Felder, 1862

    • syn. idalion Weymer, 1894

    • syn. tleson Riffarth, 1901, nom. nud.

    • syn. aganippe Riffarth, 1901, nom. nud.

    • syn. aristiona lepidus Riffarth, 1907

    • syn. idalion f. confluens Neustetter, 1912, preocc. (not Lathy, 1906)

    • syn. euphone ab. nephele Seitz, 1916

    • syn. euphone ab. confluxus Seitz, 1916, nom. nud.

    • syn. aristiona euphorbus Stichel, 1923

    • syn. mixta Apolinar, 1927

  • ssp. euphrasius Weymer, 1890, hyb.

    • syn. gradatus Weymer, 1894, hyb.

  • ssp. geminatus Weymer, 1894

  • ssp. holzingeri Fernández & Brown, 1976

  • ssp. ignotus Joicey & Kaye, 1917

  • ssp. illustris Weymer, 1894

    • syn. silvana mirificus Stichel, 1906

    • syn. novatus subnubilus Stichel, 1906

  • ssp. isabellinus Bates, 1862, hyb.

  • ssp. jiparanaensis Neustetter, 1931

    • syn. f. mediatrix Neustetter, 1931

  • ssp. laura Neustetter, 1932, hyb.

  • ssp. lenaeus Weymer, 1891

    • syn. colepta Riffarth, 1901, nom. nud.

  • ssp. lyrcaeus Weymer, 1891

  • ssp. mavors Weymer, 1894

    • syn. numata superioris ab. translata Joicey & Kaye, 1917

  • ssp. messene Felder & Felder, 1862

    • syn. sikinos Riffarth, 1901, nom. nud.

    • syn. aristiona messene f. juncta Neustetter, 1925

    • syn. aristiona colombiana Apolinar, 1927

    • syn. aristiona messene f. euphrasinus Neustetter, 1928

  • ssp. mirus Weymer, 1894

    • syn. leopardus Weymer, 1894

    • syn. spadicarius Weeks, 1901

    • syn. arethusa Riffarth, 1901, nom. nud.

    • syn. novatus obscurior Stichel, 1906

    • syn. novatus artemis Riffarth, 1907

    • syn. ainsolitus Avinoff, 1926

    • syn. novatus f. confluens Neustetter, 1931, preocc. (not Lathy, 1906)

  • ssp. nubifer Butler, 1875

  • ssp. numata (Cramer [1780])

    • syn. pione (Hübner, 1816), repl. name Refs.: Hübner (1816): 11.

    • syn. var. melanops Weymer, 1894

    • syn. var. guiensis Riffarth, 1900

    • syn. numata isabellinus f. intermedia Boullet & Le Cerf, 1909

    • syn. f. melanopors Joicey & Kaye, 1917

  • ssp. peeblesi Joicey & Talbot, 1925

  • ssp. pratti Joicey & Kaye, 1917

  • ssp. robigus Weymer, 1875

    • syn. numilia (Herrich-Schäffer, 1865), nom. nud.

  • ssp. silvana (Stoll, 1781)

    • syn. diffusus Butler, 1873

    • syn. silvana var. divisus Kaye, 1906

    • syn. numata silvaniformis Joicey & Kaye, 1917

    • syn. silvana atakama Neustetter, 1931

  • ssp. sourensis Brown, 1976

  • ssp. superioris Butler, 1875

    • syn. numata var. maecenas Weymer, 1894

    • syn. numatus sincerus Riffarth, 1907

  • ssp. talboti Joicey & Kaye, 1917

  • ssp. tarapotensis Riffarth, 1901

    • syn. staudingeri Weymer, 1894, hyb.

    • syn. staudingeri var. pretiosus Weymer, 1894, hyb.

    • syn. aristiona lepidus f. gracilis Riffarth, 1907

    • syn. aristiona timaeus f. aristeus Neustetter, 1931, hyb.

    • syn. aristiona staudingeri f. lutea Neustetter, 1931, hyb.

  • ssp. timaeus Weymer, 1894

  • ssp. zobrysi Fruhstorfer, 1910

• pachinus Salvin, 1871; original combination: Heliconius cydno pachinus

• pardalinus Bates, 1862

  • ssp. ariadne Neukirchen, 1995

  • ssp. butleri Brown, 1976

  • ssp. dilatus Weymer, 1894

    • syn. pardalinus radiosus f. colorata Stichel, 1919

  • ssp. julia Neukirchen, 2000

  • ssp. lucescens Weymer, 1894

  • ssp. maeon Weymer, 1891

  • ssp. orteguaza Brown, 1976

  • ssp. pardalinus Bates, 1862

  • ssp. radiosus Butler, 1873

    • syn. fortunatus Weymer, 1883

    • syn. spurius Weymer, 1894

  • ssp. sergestus Weymer, 1894

    • syn. sergestus ab. ninacura Michael, 1926, hyb.

  • ssp. tithoreides Staudinger, 1900

    • syn. tithoreides f. minor Riffarth, 1908, nom. nud.

    • syn. f. garleppi Neustetter, 1928

• timareta (Hewitson, 1867)

  • ssp. florencia Giraldo, Salazar, Jiggins, Bermingham & Linares, 2008. Type loc.: Colombia , Caquetá, Florencia. Types: Natural History Museum ANDES of Universidad de los Andes, Colombia. Distr.: Colombia. Refs.: Giraldo et al. (2008): 8. Ety.: Florencia (Caquetá, Colombia)

  • ssp. linaresi Arias & Lamas, 2017. Type loc.: Colombia, Caquetá, San Vicente del Caguán, Vereda Las Morras, close to the Pato river. Types: Colección de Mariposas del Instituto de Investigación de Recursos Biológicos Alexander von Humboldt, Villa de Leyva, Colombia; paratype: Museo de Historia Natural de la Universidad de los Andes, Bogotá, Colombia. Distr.: San Vicente del Caguán (Caquetá), along the eastern slopes of the Colombian Andes, elevation 1100-1300 m. Refs.: Arias et al. (2017): 814. Ety.: Mauricio Linares

  • ssp. thelxinoe Lamas & Mérot, 2013. Type loc.: Peru, San Martín, entre El Afluente y Nuevo Edén. Types: Museo de Historia Natural, Universidad Nacional Mayor de San Marcos, Lima, Peru (MUSM). Distr.: Peru. Host plants: Passiflora oerstedii Mast, Passiflora riparia Mast and Passiflora (Astrophea) sp. (Mérot et al. 2013). Refs.: Mérot et al. (2013): 845. Ety.: Thelxinoe (Greek goddess)

  • ssp. timareta (Hewitson, 1867)

    • syn. contiguus Weymer, 1890

    • syn. ab. richardi Riffarth, 1900

    • syn. melpomene timareta ab. virgata Stichel, 1902

    • syn. melpomene timareta f. insolita Riffarth, 1907

    • syn. melpomene timareta f. peregrina Stichel, 1909

    • syn. f. strandi Neustetter, 1928

  • ssp. timoratus Lamas, 1998

  • ssp. tristero Brower, 1996; original combination: Heliconius tristero

Genus Philaethria

Genus: Philaethria Billberg, 1820

Type species: Philaethria dido (Linnaeus, 1763); original combination: Papilio dido Linnaeus, 1763

Synonyms:

• Metamandana Stichel [1908]. Type species: Philaethria dido (Linnaeus, 1763); original combination: Papilio dido Linnaeus, 1763. Refs.: Stichel (1908): 6

Classification: Species of Philaethria have distinctive green and black wing patterns, similar to Siproeta stelenes (malachite; Nymphalidae: Nymphalinae: Victorinini) that is also found throughout the Neotropics, but they have distinct wing shapes.

Phylogenetic relationships: Phylogenetic relationships among the species remain to be investigated.

Distribution: Widespread across the Neotropics

References: Billberg (1820): 77; Michener (1942 a) (key to genus); Constantino and Salazar (2010) (key to species)

10 species:

• andrei Brévignon, 2002

  • ssp. andrei Brévignon, 2002. Type loc.: Carapa, Macouria, French Guyana (LCB). Distr.: Lower Amazon basin in northern South America from Venezuela (El Manteco, Bolivar) to French Guiana (Carapa, Macouria) and northern Brazil (Marajó). Altitude 100–200m. Rainforest habitats. Host plants: Passiflora laurifolia. Refs.: Brevignon (2002), Constantino and Salazar (2010) (to subspecies). Ety.: André Brévignon

  • ssp. orinocensis Constantino & Salazar, 2010. Type loc.: Delta Amacuro, Venezuela (MIZA; paratypes: JCSC). Distr.: Open grasslands in the Orinoco river basin in Venezuela up to 100m above sea level and in the state of Bolivar (rio Caura). Refs.: Constantino and Salazar (2010): 16. Ety.: Orinoco River

• browni Constantino & Salazar, 2010. Type loc.: Barinitas, Barinas, Venezuela (MIZA; paratypes: CFC). Distr.: East slope of the Cordillera de Merida in Barinas, Venezuela. 600-1200m above sea level. Premontane forest habitats. Refs.: Constantino and Salazar (2010): 23. Ety.: Keith S. Brown

• constantinoi Salazar, 1991. Type loc.: Tatabro River, 100m, Anchicayá, Valle, Costa Pacifica, Colombia (MHNUC; paratypes: CFC). Distr.: Western slope of the Cordillera Occidental in Colombia (Chocó, Valle, Cauca) and northwestern Ecuador (Esmeraldas). Altitude 100-800m above sea level. Rainforest habitats. Host plants: Passiflora vitifolia, P. edulis, P. ambigua (DeVries 1987: 187). Refs.: Salazar (1991): 274. Ety.: Luis Miguel Constantino

• diatonica (Fruhstorfer, 1912); original combination: Metamandana dido diatonica. Type loc.: San Pedro Sula, Honduras (BMNH). Distr.: Mexico to Panama. Up to 1,110m above sea level. Rainforest habitats. Adults fly above the the forest canopy and forest edges. Refs.: Fruhstorfer (1912 b): 14. Ety.: not stated

• dido (Linnaeus, 1763)

  • ssp. chocoensis Constantino, 1999. Type loc.: Tatabro River, Anchicayá, Valle, Colombia (MHNUC). Distr.: Western slope of Cordillera Occidental in Colombia and western Ecuador; Magdalena valley in Colombia. Host plants: Passiflora vitifolia. Refs.: Constantino (1999): 65. Ety.: Chocó biogeographic region in Colombia

  • ssp. dido (Linnaeus, 1763); original combination: Papillio dido. Type loc.: “America Meridionalis” [South America] (BMNH). Distr.: Amazon Basin. Elevation up to 300m. Lowland rainforest habitats. Host plants: Passiflora auriculata, P. guazumaefolia in Venezuela (Brown & Fernandez, 1985), P. coccinea, P. edulis, P. mucronata, P. speciosa, P. vitifolia in Brazil (Benson et al., 1976, Brown & Mielke, 1972; Brown, 1981), P. vitifolia in Colombia (Constantino, 1998), P. laurifolia in Trinidad (Beebe et al., 1960). Refs.: Linnaeus (1763): 408. Ety.: not stated

  • ssp. panamensis Constantino & Salazar, 2010. Type loc.: Veraguas, Serrania de Tabasará, Panama (MHNUC; allotypes MHNUC; paratypes: FDLC). Distr.: Panama, Costa Rica. Refs.: Constantino and Salazar (2010): 8. Ety.: Panama

• neildi Constantino & Salazar, 2010

  • ssp. neildi Constantino & Salazar, 2010. Type loc.: Barinas, Venezuela (MIZA). Distr.: Eastern slope of Cordillera de Merida, Barinas and Cordillera de La Costa, Aragua in Venezuela. Refs.: Constantino and Salazar (2010): 12. Ety.: Andrew Neild

  • ssp. tachiraensis Constantino & Salazar, 2010. Type loc.: Táchira, Venezuela. (MIZA). Distr.: El Tamá National Park and up to Sierra de Perija in Venezuela. Refs.: Constantino and Salazar (2010): 14. Ety.: State of Táchira in Venezuela

  • ssp. winhardi Constantino & Salazar, 2010. Type loc.: Magdalena, Colombia (MHNUC). Distr.: Sierra Nevada de Santa Marta, Magdalena, Colombia. Refs.: Constantino and Salazar (2010): 14. Ety.: Walter Winhard (Bogotá)

• ostara (Röber, 1906)

  • ssp. araguensis Constantino & Salazar, 2010. Type loc.: Aragua, Venezuela (MIZA). Distr.: Cordillera de La Costa in Aragua, Trujillo and Carabobo in northwestern Venezuela. Refs.: Constantino and Salazar (2010): 17. Ety.: State of Aragua, Venezuela

  • ssp. meridensis Constantino & Salazar, 2010. Type loc.: Barinitas, Barinas, Venezuela (MIZA; paratypes: CFR). Distr.: Eastern slope of Cordillera de Merida, Venezuela. Refs.: Constantino and Salazar (2010): 19. Ety.: Cordillera de Merida, Venezuela

  • ssp. ostara (Röber, 1906); original combination: Metamorpha dido var. ostara. Type loc.: Cauca-Tale, Colombia (AMNH). Distr.: Eastern slope of the Andes in Colombia at high altitudes (450-1600m). Refs.: Röber (1906): 177.

• pygmalion (Fruhstorfer, 1912)

  • ssp. metaensis Constantino & Salazar, 2010. Type loc.: Ariari River, Meta, Colombia (MHNUC). Distr.: Savanna of the Orinoco basin and the eastern slope of the Cordillera Oriental in Colombia. Refs.: Constantino and Salazar (2010): 11. Ety.: Meta, Colombia

  • ssp. pygmalion (Fruhstorfer, 1912); original combination: Metamandana dido pygmalion. Type loc.: Obidos, Brazil (BMNH). Distr.: Rainforest habitats between 100–300m in the Amazon basin from Colombia, Venezuela to central Brazil. Host plants: Passiflora coccinea, P. faroana, P. hexagonocarpa, P. mansoi in Brazil (Benson et al., 1976). P. phaeocaula in Venezuela (Brown & Fernandez, 1985; Beccaloni et al., 2008). Refs.: Fruhstorfer (1912 b): 14; Constantino and Salazar (2010); Barão et al. (2014).

• romeroi Constantino & Salazar, 2010. Type loc.: Cordillera de La Costa, 1100m, Rancho Grande, Aragua, Venezuela (MIZA; paratypes: CFR). Distr.: Rancho Grande, Aragua, Venezuela. Host plants: Unknown. Refs.: Constantino and Salazar (2010): 23. Ety.: Francisco Romero (Maracay, Venezuela)

• wernickei (Röber, 1906); original combination: Metamorpha wernickei. Type loc.: Rio Grande do Sul & Santa Catarina, Brazil. Distr.: Eastern coast of Brazil from Rio Grande do Norte south into Uruguay and Northern Argentina. Host plants: Passiflora caerulea, P. elegans, P. mansoni, P. mucronata, P. rhamnifolia, P. sidaefolia, P. suberosa, P. actinia (Brown & Mielke, 1972; Benson et al., 1976; Brown, 1992; Beccaloni et al., 2008) in Brazil. P. quadrangularis and P. caerulea in Uruguay (Biezanko et al., 1974). Refs.: Röber (1906): 177. Ety.: H. Wernicke (Blasewitz, Germany)

Genus Podotricha

Genus: Podotricha Michener, 1942

Type species: Podotricha judith (Guérin-Méneville, [1844]); original combination: Colaenis euchroia Doubleday [1847]

Synonyms: None

Classification: Forewing: the posterior corner (tornus) is supported by Cu2 rather than A. Hindwing: the base of M2 is transverse.

References: Michener (1942 a): 3 (taxonomy)

2 species:

• judith (Guérin-Méneville [1844])

  • ssp. caucana (Riley, 1926); original combination: Colaenis euchroia caucana. Type loc.: Colombia. Distr.: Cauca Valley, Colombia. Refs.: Riley (1926): 242. Ety.: Cauca Valley, Colombia

  • ssp. judith (Guérin-Méneville [1844]); original combination: Cethosia judith. Type loc.: Colombia. Refs.: Guérin-Méneville and Cuvier (1844): 477. Ety.: not stated

    • syn. euchroia (Doubleday [1847]). Type loc.: Colombia. Refs.: Doubleday ([1847]): 149, pl. 20, fig. 3; Ackery and Smiles (1976) pl. 2.

  • ssp. mellosa (Stichel, 1906); original combination: Colaenis euchoria mellosa. Type loc.: Ecuador. Distr.: Northwestern Ecuador (such as Quito Rio Verde). Refs.: Stichel (1906): 208. Ety.: not stated

  • ssp. straminea (Riley, 1926); original combination: Colaenis euchroia straminea. Type loc.: Ecuador: Balsapamba. Refs.: Riley (1926): 242. Ety.: straw-coloured hindwing submarginal spots

• telesiphe (Hewitson, 1867)

  • ssp. telesiphe (Hewitson, 1867); original combination: Colaenis telesiphe. Type loc.: Ecuador. Distr.: Eastern slope of the Andes at elevation 1500-2600m from Colombia to Bolivia. Refs.: Hewitson (1867): 564. Ety.: Colour resembles Heliconius telesiphe

    • syn. tithraustes var. diaphana (Niepelt, 1915). Type loc.: Peru. Refs.: Niepelt (1915): 58.

  • ssp. tithraustes (Salvin, 1871); original combination: Colaenis tithraustes. Type loc.: Ecuador: Canelos. Distr.: Ecuador. Refs.: Salvin (1871): 415. Ety.: not stated

Tribe Vagrantini

Type genus: ?

Classification: The tribe Vagrantini is divided into ten genera: Algia, Algiachroa, Cirrochroa, Cupha, Lachnoptera, Phalanta, Smerina, Terinos, Vagrans and Vindula

References: Pinratana and Eliot (1996) (phylogeny); Peggie (2003) (phylogeny); Penz and Peggie (2003) (phylogeny); Peggie (2004) (phylogeny)

Genus Algia

Genus: Algia Herrich-Schäffer, 1864

Type species: Algia satyrina (Felder & Felder, [1867]); original combination: Cirrochroa satyrina Felder & Felder, [1867]

Synonyms:

• Paduca Moore, 1886. Type species: Algia fasciata Felder & Felder, 1859; original combination: Atella fasciata Felder & Felder, 1860. Refs.: Moore (1886): 34
• Ducapa Moore, 1900. Type species: Algia fasciata Felder & Felder, 1859; original combination: Atella fasciata Felder & Felder, 1860. Refs.: Moore (1900): 209

References: Herrich-Schäffer (1864): 125

3 species:

• fasciata (Felder & Felder, 1860)

  • ssp. angustata (Fruhstorfer, 1912); original combination: Cirrochroa satyrina angustata

  • ssp. bilbilis (Fruhstorfer, 1912); original combination: Cirrochroa fasciata bilbilis

  • ssp. fasciata (Felder & Felder, 1860); original combination: Atella fasciata

  • ssp. formosana (Matsumura, 1929); original combination: Cirrochroa fasciata formosana

  • ssp. lautus Tsukada, 1985; original combination: Algia fasciata lautus

  • ssp. ortopia (Fruhstorfer, 1912); original combination: Cirrochroa fasciata ortopla

  • ssp. palloris (Fruhstorfer, 1900); original combination: Paduca fasciata palloris

• satyrina (Felder & Felder [1867])

  • ssp. satyrina (Felder & Felder [1867]); original combination: Cirrochroa satyrina

  • ssp. sibylla (Röber, 1887); original combination: Cirrochroa sibylla

  • ssp. similiana (Röber, 1887); original combination: Cirrochroa similiana

• felderi (Kirsch, 1877); original combination: Cirrochroa felderi

Genus Algiachroa

Genus: Algiachroa Parsons, 1989

Type species: Algiachroa woodfordi (Godman & Salvin, 1888); original combination: Messaras woodfordi Godman & Salvin, 1888

Synonyms: None

References: Parsons (1989): 176

1 species:

• woodfordi (Godman & Salvin, 1888); original combination: Messaras woodfordi Refs.: Godman and Salvin (1888): 97.

Genus Cirrochroa

Genus: Cirrochroa Doubleday [1847]

Type species: Cirrochroa aoris Doubleday, [1847]; original combination: Cirrochroa aoris Doubleday, [1847]

Synonyms: None

References: Doubleday ([1847]): pl.21

18 species:

• aoris Doubleday [1847]

  • ssp. aoris Doubleday [1847]; original combination: Cirrochroa aoris

  • ssp. olivacea de Nicéville, 1886; original combination: Cirrhochroa olivacea

• clagia (Godart, 1824)

  • ssp. clagia (Godart, 1824); original combination: Argynnis clagia

  • ssp. clagina Fruhstorfer, 1906; original combination: Cirrochroa clagia clagina

• emalea (Guérin-Méneville, 1843)

  • ssp. bajadeta Moore, 1858; original combination: Cirrochroa bajadeta

  • ssp. emalea (Guérin-Méneville, 1843); original combination: Argynnis emalea

  • ssp. lapoana Kheil, 1884; original combination: Cirrochroa emalea lapaona

  • ssp. martini Fruhstorfer, 1906; original combination: Cirrochroa emalea martini

  • ssp. ravana Moore, 1858; original combination: Cirrochroa ravana

• eremita Tsukada, 1985; original combination: Cirrochroa eremita

• imperatrix Grose-Smith, 1894; original combination: Cirrochroa imperatrix

• malaya Felder & Felder, 1860

  • ssp. calypso Wallace, 1869; original combination: Cirrochroa calypso

    • syn. baluna Fruhstorfer, 1904

  • ssp. malaya Felder & Felder, 1860; original combination: Cirrochroa malaya

    • syn. johannes Butler, 1868

  • ssp. natuna Fruhstorfer, 1904; original combination: Cirrochroa malaya natuna

• menones Semper, 1888; original combination: Cirrochroa menones

• niassica Honrath, 1892; original combination: Cirrochroa niassica

• nicobarica Wood-Mason & de Nicéville, 1881; original combination: Cirrochroa nicobarica

• orissa Felder & Felder, 1860

  • ssp. chione Riley & Godfrey, 1921; original combination: Cirrochroa chione

  • ssp. orissa Felder & Felder, 1860; original combination: Cirrochroa orissa

  • ssp. orissides Fruhstorfer, 1906; original combination: Cirrochroa orissa orissides

• recondita Roos, 1996; original combination: Cirrochroa recondita

• regina Felder & Felder, 1867

  • ssp. ducalis Wallace, 1869; original combination: Cirrochroa regina jobina

  • ssp. jobina Talbot, 1932; original combination: Cirrochroa ducalis

  • ssp. myra Fruhstorfer, 1907; original combination: Cirrochroa regina myra

  • ssp. nasica Fruhstorfer, 1907; original combination: Cirrochroa regina nasica

  • ssp. paulowna Fruhstorfer, 1898; original combination: Cirrochroa ducalis paulowna

  • ssp. princesa Fruhstorfer, 1904; original combination: Cirrochroa regina princesa

  • ssp. regina Felder & Felder, 1867; original combination: Cirrochroa regina

  • ssp. sophene Fruhstorfer, 1907; original combination: Cirrochroa regina sophene

• satellita Butler, 1869

  • ssp. illergeta Fruhstorfer, 1912; original combination: Cirrochroa satellita illergeta

  • ssp. satellita Butler, 1869; original combination: Cirrochroa satellita

• semiramis Felder & Felder, 1867; original combination: Cirrochroa semiramis

• surya Moore, 1878

  • ssp. siamensis Fruhstorfer, 1906; original combination: Cirrochroa surya siamensis

  • ssp. surya Moore, 1878; original combination: Cirrochroa surya

• thais (Fabricius, 1787)

  • ssp. lanka Moore, 1872; original combination: Cirrochroa lanka

  • ssp. thais (Fabricius, 1787); original combination: Papilio thais

• thule Felder & Felder, 1867

  • ssp. massalia Fruhstorfer, 1906; original combination: Cirrochroa thule massalia

  • ssp. thule Felder & Felder, 1867; original combination: Cirrochroa thule

• tyche Fruhstorfer, 1905

  • ssp. anjira Moore, 1877; original combination: Cirrochroa anjira

  • ssp. anthonia Moore, 1877; original combination: Cirrochroa tyche anthonia

  • ssp. aurica Monastyrskii, 2912; original combination: Cirrochroa tyche aurica

  • ssp. laudabilis Eliot, 1978; original combination: Cirrochroa tyche laudabilis

  • ssp. lesseta Fruhstorfer, 1900; original combination: Cirrochroa tyche lesseta

  • ssp. mithila Fruhstorfer, 1912; original combination: Cirrochroa mithila

  • ssp. rotundata Moore, 1872; original combination: Cirrochroa rotundata

  • ssp. thilina Butler, 1879; original combination: Cirrochroa tyche thilina

  • ssp. tyche Fruhstorfer, 1905; original combination: Cirrochroa tyche

Genus Cupha

Genus: Cupha Billberg, 1829

Type species: Cupha erymanthis (Drury, [1773]); original combination: Papilio erymanthis Drury, [1773]

Synonyms: None

References: Billberg (1820): 79

9 species:

• arias (Felder & Felder [1867])

  • ssp. arias (Felder & Felder [1867]); original combination: Messaras arias

  • ssp. cacina Fruhstorfer, 1912; original combination: Cupha arias cacina

  • ssp. celebensis Fruhstorfer, 1900; original combination: Cupha erymanthis celebensis

  • ssp. dapatana (Grose-Smith, 1887); original combination: Messaras dapatana

  • ssp. muna Fruhstorfer, 1898; original combination: Cupha erymanthis muna

  • ssp. sangirica Fruhstorfer, 1912; original combination: Cupha arias sangirica

• aureus Samson, 1980; original combination: Cupha aureus

• crameri (Felder & Felder, 1860)

  • ssp. crameri (Felder & Felder, 1860); original combination: Messaras crameri

  • ssp. kalaoensis Tsukada, 1985; original combination: Cupha crameri kalaoensis

  • ssp. keyana Fruhstorfer, 1912; original combination: Cupha crameri keyana

  • ssp. leonida Fruhstorfer, 1912; original combination: Cupha crameri leonida

  • ssp. palla (Röber, 1891); original combination: Messaras erymanthis var. palla

• erymanthis (Drury [1773])

  • ssp. andamanica Moore, 1900; original combination: Cupha andamanica

  • ssp. disjuncta (Weymer, 1885); original combination: Messaras disjuncta

  • ssp. dohertyi Fruhstorfer, 1904; original combination: Cupha erymanthis dohertyi

  • ssp. erymanthis (Drury [1773]); original combination: Papilio erymanthis

  • ssp. kangeana Fruhstorfer, 1903; original combination: Cupha erymanthis kangeana

  • ssp. lotis (Sulzer, 1776); original combination: Papilio lotis

  • ssp. maja Fruhstorfer, 1898; original combination: Cupha erymanthis maja

  • ssp. melanodotis Monastyrskii, 2012; original combination: Cupha erymanthis melanodotis

  • ssp. nicobarica (Felder & Felder, 1862); original combination: Messaras erymanthis var. nikobarica

  • ssp. nagara Fruhstorfer, 1912; original combination: Cupha erymanthis nagara

  • ssp. palawana Fruhstorfer, 1900; original combination: Cupha erymanthis palawana

  • ssp. peliopteryx (Hagen, 1898); original combination: Messaras peliopteryx

  • ssp. placida Moore, 1881; original combination: Cupha placida

  • ssp. saturatior Fruhstorfer, 1898; original combination: Cupha erymanthis saturatior

  • ssp. synnara Fruhstorfer, 1912; original combination: Cupha erymanthis synnara

  • ssp. tiomana Corbet, 1940; original combination: Cupha erymanthis tiomana

• lampetia (Linnaeus, 1764)

  • ssp. lampetia (Linnaeus, 1764); original combination: Papilio lampetia

  • ssp. mirona Fruhstorfer, 1904; original combination: Cupha lampetia mirona

• maeonides (Hewitson, 1859)

  • ssp. maenada Grose-Smith, 1898; original combination: Cupha maenada

  • ssp. maeonides (Hewitson, 1859); original combination: Messaras maeonides

  • ssp. nigrico Tsukada, 1985; original combination: Cupha maeonides nigrico

  • ssp. rovena Fruhstorfer, 1912; original combination: Cupha maeonides rovena

• melichrysops (Mathew, 1887)

  • ssp. melichrysops (Mathew, 1887); original combination: Messarus melichrysos

  • ssp. pallescens (Godman & Salvin, 1888); original combination: Messaras pallescens

  • ssp. tredecia (Mathew, 1887); original combination: Messarus tredecia

• myronides (Felder & Felder, 1867)

  • ssp. datos Fruhstorfer, 1904; original combination: Cupha lampetia datos

  • ssp. lampetina Fruhstorfer, 1904; original combination: Cupha lampetia lampetina

  • ssp. myronides (Felder & Felder, 1867); original combination: Messaras myronides

• prosope (Fabricius, 1775)

  • ssp. alexis Grose-Smith, 1898; original combination: Cupha alexis

  • ssp. charmides Grose-Smith, 1898; original combination: Cupha charmides

  • ssp. cluentia Fruhstorfer, 1904; original combination: Cupha madestes cluentia

  • ssp. cyclotas Grose-Smith, 1894; original combination: Cupha madestes cluentia

  • ssp. decernia Fruhstorfer, 1912; original combination: Cupha prosope decernia

  • ssp. fumosa Grose-Smith, 1897; original combination: Cupha fumosa

  • ssp. madestes (Hewitson, 1859); original combination: Messaras madestes

  • ssp. oderca Fruhstorfer, 1912; original combination: Cupha madestes oderca

  • ssp. prosope (Fabricius, 1775); original combination: Papilio prosope

  • ssp. turneri (Butler, 1876); original combination: Messaras turneri

  • ssp. wallacei (Felder & Felder [1867]); original combination: Messaras wallacei

Genus Lachnoptera

Genus: Lachnoptera Doubleday [1847]

Type species: Lachnoptera anticlia (Hübner, [1819]); original combination: Issoria anticlia Hübner, [1819]

Synonyms: None

References: Doubleday ([1847]): pl.22; Doubleday ([1848]): 161

2 species:

• anticlia (Hübner, 1819); original combination: Issoria anticlia

• ayresii Trimen, 1879; original combination: Lachnoptera ayresii

Genus Phalanta

Genus: Phalanta Horsfield [1829]

Type species: Phalanta phalantha (Drury [1773]); original combination: Papilio phalantha Drury [1773]

Synonyms:

• Atella Doubleday [1847]. Type species: Phalanta phalantha (Drury [1773]); original combination: Papilio phalantha Drury [1773]. Refs.: Doubleday ([1847]): pl.22
• Atella Doubleday [1848]. Type species: Phalanta eurytis (Doubleday, [1847]); original combination: Atella eurytis Doubleday, [1847]. Refs.: Doubleday ([1848]): 165
• Albericia Dufrane, 1945. Type species: Phalanta eurytis (Doubleday, [1847]); original combination: Albericia gomensis Dufrane, 1945. Refs.: Dufrane (1945): 98

References: Horsfield ([1829]): pl.7

5 species:

• alcippe (Stoll [1782])

  • ssp. alcesta Corbet, 1941; original combination: Phalanta alcesta

  • ssp. alcippe (Stoll [1782]); original combination: Papilio alcippe

  • ssp. alcippina (Fruhstorfer, 1904); original combination: Atella alcippe alcippina

  • ssp. alcippoides (Moore, 1900); original combination: Atella alcippoides

  • ssp. andamana (Fruhstorfer, 1904); original combination: Atella alcippe andamana

  • ssp. aridus Tsukada, 1985; original combination: Phalanta alcippe aridus

  • ssp. arruana (Felder & Felder, 1860); original combination: Atella arruana

  • ssp. asinia (Fruhstorfer, 1904); original combination: Atella alcippe asinia

  • ssp. aurica Eliot, 1978; original combination: Phalanta alcippe aurica

  • ssp. bellona Howarth, 1962; original combination: Phalanta alcippe bellona

  • ssp. celebensis (Wallace, 1869); original combination: Atella celebensis

  • ssp. cervina (Butler [1877]); original combination: Atella cervina

  • ssp. cervinides (Fruhstorfer, 1904); original combination: Atella alcippe cervinides

  • ssp. ceylonica (Mander, 1903); original combination: Atella ceylonica

  • ssp. denosa (Fruhstorfer, 1912); original combination: Atella alcippe denosa

  • ssp. drepana (Fruhstorfer, 1906); original combination: Atella alcippe drepana

  • ssp. enganica (Fruhstorfer, 1904); original combination: Atella alcippe enganica

  • ssp. ephyra (Godman & Salvin, 1888); original combination: Atella ephyra

  • ssp. floresiana (Fruhstorfer, 1912); original combination: Atella alcippe floresiana

  • ssp. fraterna (Moore, 1900); original combination: Atella fraterna

  • ssp. handjahi Tsukada, 1985; original combination: Phalanta alcippe handjahi

  • ssp. interposita (Joicey & Talbot, 1916); original combination: Atella alcippe interposita

  • ssp. kinitis (Fruhstorfer, 1904); original combination: Atella alcippe kinitis

  • ssp. omarion (Fruhstorfer, 1912); original combination: Atella alcippe omarion

  • ssp. onyx Tsukada, 1985; original combination: Phalanta alcippe onyx

  • ssp. pallidior (Staudinger, 1899); original combination: Atella alcippe var. pallidior

  • ssp. quinta (Fruhstorfer, 1904); original combination: Atella alcippe quinta

  • ssp. rennellensis Howarth, 1962; original combination: Phalanta alcippe rennellensis

  • ssp. semperi (Moore, 1900); original combination: Atella semperi

  • ssp. tiomana Corbet, 1937; original combination: Phalanta alcippe tiomana

  • ssp. violetta (Fruhstorfer, 1900); original combination: Atella alcippe violetta

• eurytis (Doubleday, 1847)

  • ssp. eurytis (Doubleday, 1847); original combination: Atella eurytis

  • ssp. microps (Rothschild & Jordan, 1903); original combination: Atella columbina microps

• madagascariensis (Mabille [1887]); original combination: Atella madagascariensis

• phalantha (Drury [1773])

  • ssp. araca (Waterhouse & Lyell, 1914); original combination: Atella phalanta araca

  • ssp. aethiopica (Rothschild & Jordan, 1903); original combination: Atella phalantha aethiopica

  • ssp. columbina (Cramer [1779]); original combination: Papilio columbina

  • ssp. granti (Rothschild & Jordan, 1903); original combination: Atella phalantha granti

  • ssp. luzonica (Fruhstorfer, 1906); original combination: Atella phalanta luzonica

  • ssp. phalantha (Drury [1773]); original combination: Papilio phalantha

  • ssp. sabulum Tsukada, 1985; original combination: Phalanta phalantha sabulum

• philiberti (de Joannis, 1893); original combination: Atella philiberti

Genus Smerina

Genus: Smerina Hewitson, 1874

Type species: Smerina vindonissa Hewitson, 1874; original combination: Smerina vindonissa Hewitson, 1875

Synonyms: None

References: Hewitson (1874): 359

1 species:

• manoro (Ward, 1871); original combination: Atella manoro. Type loc.: Madagascar.

  • syn. vindonissa Hewitson, 1874; original combination: Smerina vindonissa.

Genus Terinos

Genus: Terinos Boisduval [1836]

Type species: Terinos clarissa Boisduval [1836]; original combination: Terinos clarissa Boisduval [1836]

Synonyms: None

References: Boisduval (1836): pl.9

7 species:

• alurgis Godman & Salvin, 1880; original combination: Terinos alurgis

• atlita (Fabricius, 1787)

  • ssp. atlita (Fabricius, 1787); original combination: Papilio atlita

  • ssp. fulminans Butler, 1869; original combination: Terinos fulminans

  • ssp. falcata Fruhstorfer, 1898; original combination: Terinos falcata

  • ssp. guangxienis Chou, 1994; original combination: Terinos atlita guangxienis

  • ssp. militum Oberthür, 1877; original combination: Terinos militum

  • ssp. teuthras Hewitson, 1862; original combination: Terinos teuthras

• clarissa Boisduval, 1836

  • ssp. aurensis Eliot, 1978; original combination: Terinos clarissa aurensis

  • ssp. bangueyana Fruhstorfer, 1912; original combination: Terinos clarissa bangueyana

  • ssp. clarissa Boisduval, 1836; original combination: Terinos clarissa

  • ssp. dinnaga Fruhstorfer, 1906; original combination: Terinos clarissa dinnaga

  • ssp. lucia Staudinger, 1889; original combination: Terinos lucilla

  • ssp. luciella Fruhstorfer, 1912; original combination: Terinos clarissa luciella

  • ssp. lucilla Butler, 1870; original combination: Terinos lucilla

  • ssp. ludmilla Staudinger, 1898; original combination: Terinos ludmilla

  • ssp. malayana Fruhstorfer, 1906; original combination: Terinos clarissa malayana

  • ssp. nympha Wallace, 1869; original combination: Terinos nympha

• maddelena Grose-Smith, 1889; original combination: Terinos maddelena

• taxiles Hewitson, 1862

  • ssp. abisares Felder & Felder [1867]; original combination: Terinos abisares

  • ssp. amplior Fruhstorfer, 1906; original combination: Terinos taxiles amplior

  • ssp. angurium Tsukada, 1985; original combination: Terinos taxiles angurium

  • ssp. banggaiensis Detani, 1983; original combination: Terinos taxiles banggaiensis

  • ssp. helleri Fruhstorfer, 1906; original combination: Terinos taxiles helleri

  • ssp. poros Fruhstorfer, 1906; original combination: Terinos taxiles poros

  • ssp. taxiles Hewitson, 1862; original combination: Terinos taxiles

• terpander Hewitson, 1862

  • ssp. bangkanensis Fruhstorfer, 1912; original combination: Terinos terpander bankanensis

  • ssp. blachieri Fruhstorfer, 1914; original combination: Terinos terpander blachieri

  • ssp. intermedia Godfrey, 1916; original combination: Terinos terpander intermedia

  • ssp. natunensis Fruhstorfer, 1901; original combination: Terinos terpander natunensis

  • ssp. niasica Fruhstorfer, 1901; original combination: Terinos robertsia niasica

  • ssp. piepersi Martin, 1909; original combination: Terinos piepersi

  • ssp. robertsia Butler, 1867; original combination: Terinos robertsia

  • ssp. teos de Nicéville, 1893; original combination: Terinos teos

  • ssp. terpander Hewitson, 1862; original combination: Terinos terpander

  • ssp. tiomanensis Eliot, 1978; original combination: Terinos terpander tiomanensis

• tethys Hewitson, 1862

  • ssp. tethys Hewitson, 1862; original combination: Terinos tethys

  • ssp. udaios Fruhstorfer, 1906; original combination: Terinos tethys udaios

  • ssp. wahnesi Heller, 1902; original combination: Terinos wahnesi

Genus Vagrans

Genus: Vagrans Hemming, 1934

Type species: Vagrans egista (Cramer, [1780]); original combination: Papilio egista Cramer, [1780]

Synonyms: None

References: Hemming (1934)

1 species:

• egista (Cramer [1780])

  • ssp. admiralia (Rothschild, 1915); original combination: Issoria sinha admiralia

  • ssp. bowdenia (Butler [1874]); original combination: Atella bowdenia. Type loc.: Friendly Is. Refs.: Butler (1873): 687.

  • ssp. brixia (Fruhstorfer, 1912); original combination: Issoria sinha brixia

  • ssp. buruana (Fruhstorfer, 1904); original combination: Issoria egista buruana

  • ssp. creaghana Pryor & Cator, 1894; original combination: Vagrans egista creaghana

  • ssp. eda (Fruhstorfer, 1904); original combination: Issoria sinha eda

  • ssp. egista (Cramer [1780]); original combination: Papilio egista

  • ssp. elvira (Fruhstorfer, 1904); original combination: Issoria egista elvira

  • ssp. hebridina (Waterhouse, 1920); original combination: Issoria egista hebridina

  • ssp. macromalayana (Fruhstorfer, 1912); original combination: Issoria sinha macromalayana

  • ssp. nupta (Staudinger, 1889); original combination: Atella sinha var. nupta

  • ssp. obiana (Fruhstorfer, 1904); original combination: Issoria egista obiana

  • ssp. offaka (Fruhstorfer, 1904); original combination: Issoria egista offaka

  • ssp. orfeda (Fruhstorfer, 1904); original combination: Issoria egista orfeda

  • ssp. propinqua (Miskin, 1884); original combination: Atella propinqua

  • ssp. scyllaria (Fruhstorfer, 1912); original combination: Issoria sinha scyllaria

  • ssp. shortlandia (Fruhstorfer, 1912); original combination: Issoria sinha shortlandica

  • ssp. sinha (Kollar [1844]); original combination: Terinos sinha

  • ssp. vitiensis (Waterhouse, 1920); original combination: Issoria egista vitiensis

Genus Vindula

Genus: Vindula Hemming, 1934

Type species: Vagrans arsinoe (Cramer, [1777]); original combination: Papillio arsinoe Cramer, [1777]

Synonyms: None

References: Hemming (1934)

4 species:

• arsinoe (Cramer [1777])

  • ssp. ada (Butler [1874]); original combination: Cynthia ada

  • ssp. adina (Fruhstorfer, 1906); original combination: Cynthia arsinoe adina

  • ssp. arsinoe (Cramer [1777]); original combination: Papilio arsinoe

  • ssp. bosnikensis (Joicey & Noakes, 1915); original combination: Cynthia arsinoe bosnikensis

  • ssp. catenes (Godman & Salvin, 1888); original combination: Cynthia catenes

  • ssp. clodia (Godman & Salvin, 1888); original combination: Cynthia clodia

  • ssp. dampierensis (Rothschild, 1915); original combination: Cynthia arsinoe dampierensis

  • ssp. figalea (Fruhstorfer, 1904); original combination: Cynthia arsinoe figalea

  • ssp. insularis (Salvin & Godman, 1877); original combination: Cynthia insularis

  • ssp. lemina (Ribbe, 1898); original combination: Cynthia arsinoe var. lemina

  • ssp. meforica (Fruhstorfer, 1906); original combination: Cynthia arsinoe meforica

  • ssp. melena (Fruhstorfer, 1899); original combination: Cynthia arsinoe melena

  • ssp. moluccana Nieuwenhuis, 1962; original combination: Vindula arsinoe moluccana

  • ssp. pisidike (Fruhstorfer, 1903); original combination: Cynthia arsinoe pisidike

  • ssp. polykaste (Fruhstorfer, 1903); original combination: Cynthia arsinoe polykaste

  • ssp. rebeli (Fruhstorfer, 1906); original combination: Cynthia arsinoe rebeli

• dejone (Erichson, 1834)

  • ssp. ambonensis Nieuwenhuis, 1962; original combination: Vindula dejone ambonensis

  • ssp. austrosundana (Fruhstorfer, 1897); original combination: Cynthia erota austrosundana

  • ssp. bagrada (Fruhstorfer, 1912); original combination: Cynthia erota bagrada

  • ssp. buruana (Fruhstorfer, 1902); original combination: Cynthia arsinoe buruana

  • ssp. bushi Tsukada, 1985; original combination: Vindula dejone bushi

  • ssp. celebensis (Butler, 1874); original combination: Cynthia celebensis

  • ssp. cycnia (de Nicéville, 1897); original combination: Cynthia cycnia

  • ssp. dajakorum (Fruhstorfer, 1906); original combination: Cynthia erota montana f. dajakorum

  • ssp. dejone (Erichson, 1834); original combination: Cynthia dejone

  • ssp. dioneia (Fruhstorfer, 1912); original combination: Cynthia erota dioneia

  • ssp. dorokusana (Fruhstorfer, 1899); original combination: Cynthia arsinoe dorokusuna

  • ssp. erotella (Butler, 1879); original combination: Cynthia erotella

  • ssp. erotoides (de Nicéville [1896]); original combination: Cynthia erotoides

  • ssp. javana (Fruhstorfer, 1899); original combination: Cynthia dejone javana

  • ssp. kabiana (Fruhstorfer, 1912); original combination: Cynthia erota kabiana

  • ssp. natunensis (Fruhstorfer, 1906); original combination: Cynthia erota natunensis

  • ssp. obiensis (Rothsch, 1899); original combination: Cynthia obiensis

  • ssp. rafflesi Pendlebury, 1939; original combination: Vindula rafflesi

  • ssp. satellitica (Fruhstorfer, 1899); original combination: Cynthia arsinoe satellitica

  • ssp. susanoo Tsukada, 1985; original combination: Vindula dejone susanoo

  • ssp. tiomana (Pendlebury, 1933); original combination: Cynthia erota tiomana

• erota (Fabricius, 1793)

  • ssp. asela (Moore, 1872); original combination: Cynthia asela

  • ssp. banta Eliot, 1956; original combination: Vindula erota banta

  • ssp. battaka (Martin, 1894); original combination: Cynthia battaka

  • ssp. boetonensis (Juriaanse & Lindemann, 1923); original combination: Cynthia erota boetonensis

  • ssp. chersonesia Pendlebury, 1939; original combination: Vindula chersonesia

  • ssp. erota (Fabricius, 1793); original combination: Papilio erota

  • ssp. gedeana (Fruhstorfer, 1906); original combination: Cynthia erota javana ab. gedeana

  • ssp. hainana (Holland, 1887); original combination: Cynthia deione var. hainana

  • ssp. kohana (Fruhstorfer, 1906); original combination: Cynthia erota kohana

  • ssp. montana (Fruhstorfer, 1905); original combination: Cynthia erota montana

  • ssp. orahilia (Kheil, 1884); original combination: Cynthia erota orahilia

  • ssp. palawanica (Fruhstorfer, 1899); original combination: Cynthia arsinoe palawanica

  • ssp. pallida (Staudinger, 1885); original combination: Cynthia erota var. pallida

  • ssp. ricussa (Fruhstorfer, 1912); original combination: Cynthia erota ricussa

  • ssp. saloma (de Nicéville, 1886); original combination: Cynthia saloma

  • ssp. sulaensis (Joicey & Talbot, 1924); original combination: Cynthia arsinoe sulaensis

  • ssp. vanroesseli (Jurriaanse & Volbeda, 1923); original combination: Cynthia erota vanroesseli

• sapor (Godman & Salvin, 1888)

  • ssp. albosignata (Talbot, 1932); original combination: Cynthia sapor albosignata

  • ssp. obscura (Ribbe, 1898); original combination: Cynthia sapor var. obscura

  • ssp. sapor (Godman & Salvin, 1888); original combination: Cynthia sapor

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Abbreviations

ab.

aberration

f.

form, forma

hyb.

hybrid

syn.

synonym

ssp.

subspecies

tr.f.

transition form

v.

variety

var.

variety

Type depositories

A

Personal collection of species author

ACF

Clifford Ferris, Laramie, WY, USA

AFEN

Andrew F. E. Neild Collection, London, UK

AMNH

American Museum of Natural History, New York, USA

AO

Andrés Miguel Orellana Collection (San Cristobal, Venezuela)

BCPM

British Columbia Provincial Museum, Victoria, British Columbia, Canada [see current designation, RBCM]

BMNH

British Museum (Natural History), London, UK [see current designation, NHMUK]

CAS

California Academy of Sciences, California, USA

CFC

Constantino family collection, Cali, Colombia

CFR

Romero family collection, Maracay, Venezuela

CMNH

Carnegie Museum of Natural History, Pittsburgh, PA, USA

CNC

Canadian National Collection of Insects, Agriculture Canada, Ottawa, Ontario, Canada

CNF

Norbert Flauger collection, Bejuma, Carabobo, Venezuela

CNIC

Canadian National Collection, Ottawa, Ontario, Canada

CPC

Carlos Prieto collection, Cali, Colombia

CSUC

C. P. Gillette Museum of Arthropod Biodiversity, Ft. Collins, CO, USA

DSM

Darwin State Museum, Moscow, Russia

EMUS

Utah State University Insect Collection, Logan, UT, USA

ESM

Ernesto Schmidt-Mumm collection, IaVH, Villa de Leyva, Colombia

ESUW

University of Wyoming Insect Museum, Laramie, WY, USA

FDLC

Francisco Delgado collection, Departamento de Zoologia, Universidad de Panamá, Veraguas, Panamá

FDLV

Francisco de la Villa collection, Puerto Ayacucho, Venezuela

FLMNH

McGuire Center for Lepidoptera and Biodiversity, Florida Museum of Natural History, University of Florida, Gainesville, Florida, USA

FSCA

Florida State Collection of Arthropods, Gainesville, FL, USA

FSMC

Allyn Museum of Entomology [now located at the McGuire Center for Lepidoptera and Biodiversity], Gainesville, FL, USA; see FLMNH

FMNH

Field Museum of Natural History, Chicago, IL, USA

GR

Gabriel Rodríguez collection, Envigado, Colombia

IAVH

Instituto Alexander Von Humboldt, Villa de Leyva, Colombia

IBUNAM

Instituto de Biología, Universidad Nacional Autónoma de Mexico, Mexico City, Mexico

ICN

Instituto de Ciencias Naturales, Universidad Nacional, Bogotá, Colombia

INBio

Instituto Nacional de Biodiversidad, Sto.Domingo de Heredia, Costa Rica

INHS

Illinois Natural History Survey, Champaign (Urbana), Illinois, USA

IZCAS

Institute of Zoology, Chinese Academy of Sciences, Beijing, China

JAS

Julian A. Salazar collection, Manizales, Colombia

JCSC

Juan Carlos de Sousa collection, Maracay, Venezuela

JFC

Jean F. LeCrom collection, Bogotá, Colombia

KSB

Keith S. Brown collection, Sao Paulo, Brazil

KWJH

Keith R. Willmott & Jason P. Hall collection, Bristol, UK

LACM

Natural History Museum of Los Angeles County, Los Angeles, CA, USA

LCB

Christian Brevignon collection, French Guiana

LNKD

Landessammlung für Naturkunde, Karlsruhe, Germany

MB

Mohamed Benmesbah, Toulouse, France

MBSM

Monte L. Bean Life Science Museum, Brigham Young University, Provo, Utah, USA

MC

Mauro Costa, Caracas, Venezuela

MCPM

Milwaukee Public Museum, Milwalkee, WI, USA

MCZ

Museum of Comparative Zoology, Harvard University, Cambridge, Massachusetts, USA

MECN

Museo Ecuatoriano de Ciencias Naturales, Quito, Ecuador

MEL

Museo Entomológico de León, León, Nicaragua

MEMB

Museo Entomológico Marcial Benavides-Cenicafé, Chinchiná, Colombia

MNHNC

Museo Nacional de Historia Natural de Chile, Santiago, Chile

MHNUC

Museo de Historia Natural, Universidad de Caldas, Manizales, Colombia

MHNUSM

Museo de Historia Natural, Universidad Nacional Mayor de San Marcos, Lima, Peru

MIZA

Museo del Instituto de Zoología Agrícola, Universidad Central de Venezuela, Maracay, Venezuela

MNHN

Muséum national d’Histoire naturelle, Paris, France [= MNHP]

MNKM

Museo Noel Kempff Mercado, Santa Cruz, Bolivia

MUSM

Museo de Historia Natural, Universidad Nacional Mayor de San Marcos, Lima, Peru

MZFC

Museo de Zoologia de la Facultad de Ciencias, UNAM, Mexico City, Mexico [see current designation, IBUNAM]

MZUSP

Museu de Zoologia da Universidade de São Paulo, São Paulo, Brazil

NCBN

National Centre for Biodiversity Naturalis, Leiden, the Netherlands [previously RMNH]

NHMUK

Natural History Museum, London, England, UK

NMNH

National Museum of Natural History, Smithsonian Insitution, Washington DC, USA [formerly United States National Museum; USNM]

NVMC

Nevada State Museum, Carson City, NV, USA

OSU

Ohio State University, Columbus, Ohio, USA

PMNH

Peabody Museum of Natural History, Yale University, New Haven, Connecticut, USA

RBCM

Royal British Columbia Museum, Victoria, British Columbia, Canada [previously BCPM]

RMNH

National Museum of National History (Nationaal Natuurhistorisch Museum), Naturalis, Leiden, the Netherlands; formerly Rijksmuseum van Natuurlijke Historie [see current designation, NCBN]

SDSU

Severin-McDaniel Insect Collection, South Dakota State University, Brookings, South Dakota, USA

SMF

Forschungsinstitut und Naturmuseum Senckenberg, Frankfurt-am-Main, Germany

SZMN

Siberian Zoological Museum, Institute of Animal Systematics and Ecology, Novosibirsk, Russia

UEC

Museo Dept. de Zoologia, Universidade Estadual de Campinas, São Paulo, Brazil [= ZUEC]

UFPC

Departamento de Zoologia, Universidade Federal do Paraná, Curitiba, Paraná, Brazil

USNM

United States National Museum, Washington DC, United States [see current designation, NMNH]

UTR

University of Tomsk, Tomsk, Russia

UUZM

Uppsala Universitet Zoologiska Museum, Uppsala, Sweden [= UZIU, ZIUU]

WWC

Walter Winhard collection, Bogotá, Colombia

ZIN

Zoological Institute, Russian Academy of Sciences, St. Petersburg, Russia

ZMHU

Zoologisches Museum der Humboldt Universität, Berlin, Germany [= ZMHB]

ZMUC

Zoologisk Museum, Universitets Copenhagen, Copenhagen, Denmark

ZUEC

Museu de Zoologia da Universidade Estadual de Campinas, Campinas, São Paulo, Brazil