Research Article |
Corresponding author: Shen-Horn Yen ( shenhornyen@hotmail.com ) Academic editor: Thomas Simonsen
© 2017 Chia-Hsuan Wei, David J. Lohman, Djunijanti Peggie, Shen-Horn Yen.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Wei C-H, Lohman DJ, Peggie D, Yen S-H (2017) An illustrated checklist of the genus Elymnias Hübner, 1818 (Nymphalidae, Satyrinae). ZooKeys 676: 47-152. https://doi.org/10.3897/zookeys.676.12579
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We review the genus Elymnias Hübner, 1818, a morphologically diverse satyrine butterfly clade involved in multifarious Batesian mimicry relationships throughout Asia and Africa. A variety of different model species are mimicked, and many Elymnias species are sexually dimorphic mimics, with males and females resembling different model species. We revise species and subspecies delimitations in light of an integrative taxonomic investigation using external morphology, male and female genital morphology, and a multi-locus molecular phylogeny. There is little interspecific genitalic variation among species in this group, and previous taxonomists therefore relied almost entirely on wing patterns. Our molecular phylogenetic analysis reveals several examples of polymorphism or wing pattern divergence within a single species currently classified as two or more different species. We also found examples of wing pattern convergence among disparate lineages that mimic the same widespread model species. Frequently, two or more phenotypically similar species were classified as a single species. This comprehensive checklist reviews all names associated with Elymnias to align its taxonomy with the evolutionary history of the group. All available information on nomenclature, type localities, repositories of type specimens, and geographical distributions is summarized, and images of adult specimens and genitalia are provided along with distribution maps of all species and selected subspecies. We identify 2 species incertae sedis, establish 15 monophyletic species groups (including 1 species unplaced in any species group), and make 49 taxonomic changes, including 35 new synonyms, 7 new combinations (2 of which have new status), 1 resurrected combination, 1 resurrected subspecies, and 7 status changes.
Batesian mimicry, convergent evolution, Elymniini , polymorphism, sexual dimorphism, Southeast Asia
Elymnias Hübner, 1818 (Nymphalidae: Satyrinae) is a species-rich and widespread butterfly genus distributed throughout the Old World tropics (
We recognize a single Afrotropical species with two subspecies and 52 Australasian species with 181 subspecies distributed from Nepal to Sri Lanka in the west, throughout tropical and subtropical Asia, and extending east to Taiwan and south to Australia’s Cape York and the Bismark Archipelago of Papua New Guinea. A few species are widespread across several countries or landmasses, but many are restricted to single islands. Several new species have been discovered recently (
Species in the genus differ markedly in wing color, pattern, shape, and size, making Elymnias one of the most morphologically heterogeneous butterfly genera (
Larvae of all species with known life histories feed exclusively on palms (Arecaceae) (
After the description of Linnaeus’s Papilio hypermenstra (now Elymnias hypermnestra hypermnestra) in 1763, the accumulation of new species/subspecies publications was slow and confined to few localities, for instance, E. nesaea (Linnaeus, 1764) from Java, E. panthera (Fabricius, 1787) from Borneo, and E. vitellia (Stoll, [1781]) from Ambon. Most species and subspecies were described during between the middle of the 19th century and early 20th century in the following works:
The first checklist of Elymnias was compiled by
This checklist enumerates and verifies all current combinations and synonyms, and provides original literature citations, type localities, repositories of type specimens, photographs of specimens and genitalia, and maps of each subspecies’ geographical range and type locality. Integrative taxonomic practice employing multi-locus molecular phylogenetics in concert with data from wing and genitalic characters has informed taxonomic decisions to retain or revise contentious classification and nomenclature. This checklist is meant to clarify taxonomic problems in the genus and aid biologists interested in studying butterfly biodiversity, but will also serve as a framework for future studies on the phylogeny, biogeography, wing pattern evolution, and speciation of this fascinating radiation of Batesian mimetic butterfly species.
The taxonomic changes we propose are based on examinations of hundreds of specimens in dozens of museums, quantification of wing and genitalic characters including over 100 dissections of males and females (
Verification of type specimens was based on information provided in the original literature as well as critical review of the collection of specific authors, especially Fruhstorfer. All taxonomic treatments proposed in the present study, including the availability of infrasubspecific taxa, follow regulations and suggestions of the latest version of ICZN (1999).
All publications with original descriptions of new taxa or describing new taxonomic acts were consulted to verify the status and collection localities of type material. Geographical information was obtained directly from specimen labels and from literature to provide accurate locality data and minimize misinterpretation of geographical localities caused by misidentified or mislabeled specimens.
All images of specimens photographed in various museum collections are used here with permission from each museum. Except for the photographs provided by KUTH (Department of Entomology, Kasetsart University), David J. Lohman, and the Museum of Comparative Zoology, Harvard University, all the other photographs were taken by Chia-Hsuan Wei and Shen-Horn Yen.
The following abbreviations are used to specify the repository of type material. Specimens, including type specimens, were borrowed and/or photographed from many of these institutions and private collections.
DNPFIC Forest Insect Collection, Department of National Parks, Wildlife and Plant Conservation, Thailand
ECMP Entomology Collection, Bureau of Science, Manila, Philippines
HPC Hiroto Hanafusa Private Collection, Japan
HSPC Hiroyuki Soeda Personal Collection, Japan
IPC Yutaka Inayoshi Private Collection, Chiang Mai, Thailand
JPC Julian Jumalon Private Collection, Cebu City, Philippines
KMSPC Kazu-Michi Suzuki Private Collection, Japan
KUTH Department of Entomology, Kasetsart University, Thailand
LSL Linnaean Society of London, London, London, UK
MEPR Museo Entomologico Pietro Rossi, Duino, Italy
MUS Malaysia Universiti Sarawak, Kota Samarahan, Malaysia
NBC National Biodiversity Center, Leiden, Netherlands
NHMT The National History Museum at Tring, Tring, UK
NSYSU National Sun Yat-Sen University, Kaohsiung, Taiwan
NWSUAF Northwest A & F University, Shaanxi, China
OMPC Kikumaro Okano Private Collection, Japan
OPC Kiyoshi Okubo Private Collection, Japan
SMFD Naturmuseum Senckenberg, Frankfurt am Main, Germany
SMTD Staatliches Museum für Tierkunde, Dresden, Germany
SPC Kotaro Saito Private Collection, Japan
TME Toyosato Museum of Entomology, Tsukuba, Japan
TPC Tsukada Private Collection, Japan
UPC Yoshinobu Uémura Private Collection, Japan
We ascribe to the Biological Species Concept, which defines species as reproductively isolated groups of populations (
Given the within-species morphological variability known from this and other mimetic butterfly taxa (
For convenience, we have divided the genus into 15 monophyletic species groups (
A variety of sources were used to infer the distribution maps that we provide, including museum data, taxonomic and other publications (
This annotated checklist is formatted in the following way:
valid species name author, year
Specimens: Figs X, Y, Z; Male genitalia: Figs X, Y, Z; Distribution: Fig: X
ssp. recognized valid subspecies name , author, year
Original combination of subspecies, Author, Year. TL: Type locality provided in original description (Current name of type locality in a standardized format- Country: Province, locality). TS: Depository of type specimen. Original citation.
Junior synonym original combination, Author, Year. TL: Type locality provided in original description (Current name of type locality); TS: Depository of type specimen. Original citation.
ELYMNIAS Hübner, 1818 (Type species: Elymnias jynx Hübner, 1818, = Papilio hypermnestra Linnaeus, 1763) Zuträge Samml. exot. Schmett. 1:12.
Didonis Hübner, [1819] (Type species: Papilio vitellia Stoll, 1781)
Verz. bek. Schmett. 2: 17.
Dyctis Boisduval, 1832 (Type species: Dyctis agondas Boisduval, 1832)
Voy. Astrolabe. 1: 138.
Agrusia Moore, 1894 (Type species: Melanitis esaca Westwood, 1851)
Lepidoptera Indica 2 (18): 169.
Bruasa Moore, 1894 (Type species: Melanitis penanga Westwood, 1851)
Lepidoptera Indica 2 (18): 164–165.
Melynias Moore, 1894 (Type species: Papilio lais Cramer, [1777])
Lepidoptera Indica 2 (18): 156–163.
Mimadelias Moore, 1894 (Type species: Elymnias vasudeva Moore, 1858)
Lepidoptera Indica 2 (18): 165–168.
Elymniopsis Fruhstorfer, 1907 (Type species: Papilio phegea Fabricius, 1793)
Dt. ent. Z. Iris 20 (3): 173–174.
bammakoo (Westwood, 1851)
Specimens: Fig.
ssp. bammakoo (Westwood, 1851)
Papilio phegea Fabricius, 1793. TL: Uganda. TS:
Melanitis bammakoo Westwood, 1851. TL: Ashanti (Ghana: Ashanti Region). TS:
Elymnias phegea var. intermedia Aurivillius, 1898. TL: not indicated. TS:
Elymnias phegea ab. angustata Bartel, 1905. TL: Kamerun, Barombi Station (Cameroon: Southwest Region, Barombi Mbo). TS: unknown. Novit. Zool. 12: 129.
Elymiopsis bammakoo var. hybrida Niepelt, 1915. TL: Kassai River (Democratic Republic of Congo: Kasai River). TS:
Elymniopsis lise Hemming, 1960. TL: Uganda. TS:
ssp. rattrayi Sharpe, 1902
Elymnias rattrayi Sharpe, 1902. TL: Entebbe (Uganda: Central Uganda, Entebbe). TS:
Elymnias ugandae Grünberg, 1908. TL: Uganda. TS: unknown. Sitzungsber. Ges. Naturf. Freunde. Berlin. 1908: 51.
Elymniopsis ugandae f. rattrayi Lewis, 1974. TL: Uganda. TS: unknown. Butterflies of the World, p. 266, pl. 115, fig. 15.
Elymniopsis bammakoo rattrayi (Sharpe, 1902). TL: Uganda. TS: unknown. Butterflies of West Africa, p. 283.
paradoxa Staudinger, 1894
Specimens: Fig.
Elymnias paradoxa Staudinger, 1894. TL: Kubary (Papua New Guinea: Madang Province, Mt. Kubari). TS:
Elymnias erastus Grose-Smith, 1894. TL: Sattelberg (Papua New Guinea: Morobe Province, Huon Peninsula, Sattelberg). TS:
papua Wallace, 1869
Specimens: Fig.
ssp. papua Wallace, 1869
Elymnias papua Wallace, 1869. TL: New Guinea
Elymnias viridescens Grose-Smith, 1894. TL: Humboldt Bay (Indonesia: Papua, Jayapura, Yos Sudarso Bay). TS:
Dyctis viridescens var. kakarona Hagen, 1897. TL: Sattelberg (Papua New Guinea: Morobe Province, Huon Peninsula, Sattelberg). TS: NHMT. Jarhb. Nass. Ver. Nat. 50: 78.
Elymnias papua bivittata van Eecke, 1915. syn. n. TL: Bivakeiland, Koofbivak, New Guinea (Indonesia: Papua, Asmat Regency, Bivak Island). TS: NBC. Nova Guinea 13 (1): 65, pl. 3, fig. 5 & 5a.
ssp. lactentia Fruhstorfer, 1907
Elymnias papua lactentia Fruhstorfer, 1907. TL: Waigiu Island (Indonesia: West Papua, Raja Ampat Regency, Waigeo). TS:
ssp. cinereomargo Joicey & Noakes, 1915
Elymnias viridescens cinereomargo Joicey & Noakes, 1915. TL: Biak Island (Indonesia: Papua, Biak). TS:
ssp. climena Talbot, 1932
Elymnias climena Talbot, 1932. TL: Mysol Island (Indonesia: West Papua, Raja Ampat Regency, Misool). TS:
ssp. euploeoides Talbot, 1932
Elymnias euploeoides Talbot, 1932. TL: Batchian (Indonesia: North Maluku, Bacan). TS:
esaca (
Specimens: Figs
ssp. esaca (Westwood, 1851)
Melanitis esaca Westwood, 1851. TL: East Indies. TS:
Elymnias godferyi Distant, 1883. TL: Sungei Ujong (Peninsular Malaysia: Negeri Sembilan, Sungei Ujong). TS:
Dyctis esacoides de Nicéville, [1893]. TL: Perak (Peninsular Malaysia: Perak), Battak Mountains
ssp. egialina (C. & R. Felder, 1863)
Melanitis egialina C. & R. Felder, 1863. TL: Luzon (Philippines: Luzon). TS:
Wien. ent. Monats. 7 (4): 121.
Melanitis ligya C. & R. Felder, 1863. nom. nud. TL: Luzon (Philippines: Luzon). TS:
Melanitis pallas C. & R. Felder, 1863. nom. nud. TL: Luzon (Philippines: Luzon). TS:
ssp. borneensis Wallace, 1869
Elymnias borneensis Wallace, 1869. TL: Sarawak (East Malaysia: Sarawak). TS:
Elymnias (Mimadelias) esaca taeniola Fruhstorfer, 1907. syn. n. TL: southeast Borneo. TS:
ssp. andersonii (Moore, 1886)
Dyctis andersonii Moore, 1886. TL: Mergui (Myanmar: Thanintharyi, Mergui Archipelago). TS:
Elymnias (Mimadelias) oberthuri Fruhstorfer, 1902. syn. n. TL: Renong, Siam (Thailand: Ranong). TS:
ssp. maheswara Fruhstorfer, 1894
Elymnias (Dyctis) maheswara Fruhstorfer, 1894. TL: Gede Vulcan (Indonesia: West Java, Mt. Gede). TS:
ssp. leontina Fruhstorfer, 1898
Elymnias esaca leontina Fruhstorfer, 1898. TL: Nias (Indonesia: North Sumatra, Nias). TS:
ssp. pseudodelias Fruhstorfer, 1907
Elymnias (Mimadelias) esaca pseudodelias Fruhstorfer, 1907. TL: Sumatra (Indonesia: Sumatra). TS:
ssp. georgi Fruhstorfer, 1907
Elymnias (Mimadelias) esaca georgi Fruhstorfer, 1907. TL: Mindanao (Philippines: Mindanao). TS:
ssp. saifuli Hanafusa, 1993
Elymnias esaca saifuli Hanafusa, 1993. TL: Siberut Island (Indonesia: West Sumatra, Mentawai Islands, Siberut). TS: HPC. Futao (11): 3.
ssp. popularis Hanafusa, 1994
Elymnias esaca popularis Hanafusa, 1994. TL: Tanahmasa Island (Indonesia: North Sumatra, South Nias Regency, Batu Islands, Tanahmasa). TS: HPC. Futao (17): 19.
ssp. splendida Tateishi, 2001
Elymnias esaca splendida Tateishi, 2001. TL: Singkep Island (Indonesia: Riau Islands, Lingga Archipelago, Singkep Island). TS:
ssp. lingga Tateishi, 2001
Elymnias esaca lingga Tateishi, 2001. TL: Lingga Island (Indonesia: Riau Islands, Lingga Archipelago, Lingga Island). TS:
ssp. nigricans Tateishi, 2001
Elymnias esaca nigricans Tateishi, 2001. TL: Enggano Island (Indonesia: Bengkulu, Enggano Island). TS:
ssp. andrewi Schröder & Treadaway, 2003
Elymnias esaca andrewi Schröder & Treadaway, 2003. TL: Philippines: Oriental Mindoro, Mt. Halcon. TS: SMFD. Nachr. ent. Ver. Apollo 23 (4): 193, pl. 1, figs 3–4.
ssp. leytensis Schröder & Treadaway, 2003
Elymnias esaca leytensis Schröder & Treadaway, 2003. TL: Philippines: Southern Leyte, Saint Bernard, Hinabian. TS: SMFD. Nachr. ent. Ver. Apollo 23 (4): 194, pl. 1, figs 7–8.
ssp. tateishii Lamas, 2010
Elymnias esaca tateishii Lamas, 2010. SHILAP 38 (150): 198. (replacement name of Elymnias esaca lautensis Teteishi, 2001).
Elymnias esaca lautensis Tateishi, 2001. TL: Laut Island (Indonesia: South Kalimantan, Kota Baru, Laut Island). TS:
vasudeva Moore, 1857
Specimens: Fig.
ssp . vasudeva Moore, 1857
Elymnias (Mimadelias) vasudeva vasudeva Moore, 1857. TL: Darjeeling (India: West Bengal, Darjeeling). TS:
Elymnias thycana Wallace, 1869. syn. n. TL: India. TS:
Mimadelias deva Moore, 1894. syn. n. TL: Khasia Hills, Assam (India: Meghalaya, Khasi Hills). TS:
Mimadelias burmensis Moore, 1893. syn. n. TL: Tenasserim (Myanmar: Tanintharyi, Tenasserim). TS:
Elymnias vacudera [sic] sinensis Chou, Zhang & Xie, 2000. syn. n. TL: Yunnan (China: Yunnan). TS: NWSUAF. Entomotaxonomia 22 (3): 224, figs 7–8.
dara Distant & Pryer, 1887
Specimens: Fig.
ssp. dara Distant & Pryer, 1887
Elymnias dara Dinstant & Pryer, 1887. TL: north Borneo. TS:
ssp. albofasciata Staudinger, 1889
Elymnias albofasciata Staudinger, 1889. TL: Philippines: Palawan. TS:
ssp. deminuta Staudinger, 1889
Elymnias albofasciata var. deminuta Staudinger, 1889. TL: Lawang (Indonesia: East Java, Malang, Lawang). TS:
ssp. bengena Fruhstorfer, 1907
Elymnias dara bengena Fruhstorfer, 1907. TL: Palabuan (Indonesia: West Java, Sukabumi, Pelabuhan Ratu). TS:
ssp. darina Fruhstorfer, 1907
Elymnias dara darina Fruhstorfer, 1907. TL: Battak Mountains (Indonesia: North Sumatra). TS:
ssp. daedalion (de Nicéville, 1890)
Dyctis daedalion de Nicéville, 1890. TL: Myittha (Myanmar: Mandalay, Kyaukse, Myittha). TS:
patna (Westwood, 1851)
Specimens: Fig.
ssp. patna (Westwood, 1851)
Melanitis patna Westwood, 1851. TL: East India. TS:
Elymnias patna bercovitzi Joicey & Talbot, 1921. TL: Five Finger Mountains (China: Hainan, Wuzhi Mountain). TS:
Melanyias patnoides Moore, 1893. syn. n. TL: Burma, Karen Hills, East Pegu (Myanmar: Bago). TS:
Elymnias patna stictica Fruhstorfer, 1902. syn. n. TL: Than-Moi, Nordtonkin (Vietnam: Lang Son, Than Moi). TS:
ssp. hanitschi Martin, 1909
Elymnias patna hanitschi Martin, 1909. TL: Malayische Halbinsel (Thai-Malay Peninsula). TS: NHMT. Dt. ent. Z. Iris 22 (1): 52.
peali Wood-Mason, 1883
Specimens: Fig.
Elymnias peali Wood-Mason, 1883. TL: Aideo, Sibsagar district, Assam (India: Assam, Sivasagar). TS:
ceryx (Boisduval, 1836)
Specimens: Fig.
Melanitis ceryx Boisduval, 1836. TL: West Java (Indonesia: West Java). TS:
Elymnias hestinia Fruhstorfer, 1911. TL: Java (Indonesia: Java). TS:
kuenstleri Honrath, [1885]
Specimens: Fig.
ssp. kuenstleri Honrath, [1885]
Elymnias künstleri (=kuenstleri) Honrath, [1885]. TL: Perak and Malacca (Peninsular Malaysia: Perak and Malacca). TS:
ssp. gauroides Fruhstorfer, 1894
Elymnias gauroides Fruhstorfer, 1894. TL: Tjisewu, West Java (Indonesia: West Java, Cisewu). TS:
ssp. rileyi Corbet, 1933
Elymnias kuenstleri rileyi Corbet, 1933. TL: Borneo. TS:
Elymnias borneensis Riley, 1923. TL: Borneo. TS: unknown. Entomologist 56 (717): 36.
ssp. dohrnii de Nicéville, 1895
Elymnias (Melynias) dohrnii de Nicéville, 1895. TL: Bohorok, East Sumatra (Indonesia: North Sumatra, Langkat Regency, Bohorok). TS:
Elymnias kuenstleri mariae Toxopeus, 1936. syn. n. TL: Bekoelen (Indonesia: South Sumatra, Bengkulu). TS: NBC. Ent. Med. Ned. Ind. 2: 46, fig. 1.
ceryxoides de Nicéville, 1895. stat. rev.
Specimens: Fig.
Elymnias (Melynias) ceryxoides de Nicéville, 1895. TL: Battak Mountains (Indonesia: North Sumatra). TS:
Elymnias ceryx ceryxoides f. nigritia Fruhstorfer, 1907. TL: Vulkan Singalang (Indonesia: West Sumatra, Agam Regency, Mt. Singgalang). TS:
pellucida Fruhstorfer, 1895
Specimens: Fig.
Elymnias pellucida Fruhstorfer, 1895. TL: Kinabalu (East Malaysia: Sabah, Mt. Kinabalu). TS:
Elymnias annea Pryer & Cator, 1894. TL: Borneo. TS:
Elymnias aroa Shelford, 1902. TL: Mount Penrissen, Sarawak (East Malaysia: Sarawak, Mt. Penrissen). TS:
penanga (Westwood, 1851)
Specimens: Fig.
ssp. penanga (Westwood, 1851)
Melanitis penanga Westwood, 1851. TL: Penang (Peninsular Malaysia: Penang). TS:
Melaninis mehida Hewitson, 1863. TL: Singapore. TS:
Elymnias abrisa Distant, 1886. TL: Province Wellesley (Peninsular Malaysia: Penang, Seberang Perai). TS:
Elymnias penanga penanga f. hislopi (♀) Eliot, 1967. TL: Langkawi (Peninsular Malaysia: Kedah, Langkawi). TS:
Elymnias penanga f. immaculata Martin, 1909. TL: Indonesia: Sumatra. TS: NHMT. Dt. ent. Z. Iris 22 (1): 55.
Elymnias penanga penanga f. johnsoni Talbot, 1929. TL: Penang (Peninsular Malaysia: Penang). TS:
ssp. sumatrana Wallace, 1869
Elymnias sumatrana Wallace, 1869. TL: Sumatra (Indonesia: Sumatra). TS:
ssp. konga Grose-Smith, 1889
Elymnias konga Grose-Smith, 1889. TL: Kina Balu Mountain, (East Malaysia: Sabah, Mt. Kinabalu). TS:
Elymnias borneensis Grose-Smith, 1892. TL: Northeast Borneo. TS:
Elymnias penanga trepsichroides Shelford, 1904. TL: North Borneo. TS:
Elymnias penanga konga f. mehidina, Fruhstorfer, 1907. TL: Borneo. TS:
Elymnias penanga konga f. ptychandrina, Fruhstorfer, 1907. TL: North Borneo. TS:
ssp. chelensis de Nicéville, 1890
Elymnias chelensis de Nicéville, 1890. TL: Khasi Hills (India: Meghalaya, Khasi Hills). TS:
hypermnestra (Linnaeus, 1763)
Specimens: Figs
ssp. hypermnestra (Linnaeus, 1763)
Papilio hypermnestra Linnaeus, 1763. TL: Java (Indonesia: Java). TS: LSL. Amoenitates Acad. 6: 407.
Papilio protogenia Cramer, 1779. TL: Java (Indonesia: Java). TS: NBC. Uitl. Kapellen. 2 (16): 141, pl. 189, fig. F–G.
Hamadryas jynx Hübner, 1808. TL: not indicated. TS: unknown. Erste Zutr. Samml. exot. Schmett. p. 4.
Elymnias jynx Hübner, 1818. TL: East Indies. TS: unknown. Zuträge Samml. exot. Schmett. 1: 12.
Elymnias hypermnestra hypermnestra f. perpusilla Fruhstorfer, 1907. TL: Java (Indonesia: Java). TS:
Elymnias hypermnestra f. atrata Roepke, 1942. TL: Java (Indonesia: Java). TS: NBC. Rhop. Javan. (4): 422.
ssp. undularis (Drury, 1773)
Papilio undularis Drury, 1773. TL: East Indies. TS:
Biblis undularis Westwood, 1837. TL: East Indies, Java (Indonesia: Java). TS:
Melanitis undularis Westwood, 1851. TL: East India, Java (Indonesia: Java). TS:
ssp. fraterna Butler, 1871
Elymnias fraterna Butler, 1871. TL: Ceylon (Sri Lanka). TS:
ssp. nigrescens Butler, 1871
Elymnias nigrescens Butler, 1871. TL: Sarawak (East Malaysia: Sarawak). TS:
Elymnias hecate Butler, 1871. TL: Labuan, Borneo (East Malaysia: Labuan). TS:
Elymnias nigrescens nigrescens f. pseudagina Fruhstorfer, 1907. TL: Sarawak, Borneo (East Malaysia: Sarawak). TS:
Elymnias nigrescens nigrescens f. edela Fruhstorfer, 1907. TL: Pontianak (Indonesia: West Kalimantan, Pontianak). TS:
Elymnias nigrescens nigrescens f. virilis Fruhstorfer, 1907. TL: Lawas (East Malaysia: Sarawak, Lawas). TS:
Elymnias nigrescens nigrescens f. hecate Fruhstorfer, 1907. TL: Labuan (East Malaysia: Labuan). TS:
ssp. cottonis (Hewitson, 1874). comb. n.
Melanitis cottonis Hewitson, 1874. TL: Andaman Islands (India: Andaman Islands). TS:
Elymnias cottonis cottonis Fruhstorfer, 1907. TL: Andaman Islands (India: Andaman Islands). TS:
ssp. tinctoria Moore, [1879]
Elymnias tinctoria Moore, [1879]. TL: Meetan, Moolai (Myanmar: Tanintharyi) TS:
Elymnias hypermnestra tinctoria f. paraleuca Fruhstorfer, 1907. TL: Mergui-Archiel, Tenasserim (Myanmar: Thanintharyi, Mergui Archipelago). TS:
ssp. hainana Moore, 1878
Elymnias hainana Moore, 1878. TL: Hainan (China: Hainan). TS:
Elymnias nigrescens formosana Fruhstorfer, 1903. TL: Takau (Taiwan: Kaohsiung). TS:
Elymnias nigrescens tonkiniana Fruhstorfer, 1902. syn. n. TL: Tonkin, Haiphong (Vietnam: Haiphong). TS:
Elymnias hypermnestra nigrescens f. depicta Fruhstorfer, 1907. syn. n. TL: Tonkin (northern Vietnam). TS:
Elymnias hypermnestra septentrionalis Chou & Huang, 1994. syn. n. TL: Nanning (China: Guangxi, Nanning). TS: NWSUAF. Monographia Rhopalocerum Sinensium 1: 375, fig. 27.
ssp. discrepans Distant, 1882
Elymnias discrepans Distant, 1882. TL: Penang, Province Wellesley (Peninsular Malaysia: Penang, Seberang Perai). TS:
ssp. orientalis Röber, 1891
Elymnias orientalis Röber, 1891. TL: Flores (Indonesia: East Nusa Tenggara, Flores). TS: unknown. Tijdschr. Ent. 34: 311.
Elymnias nigrescens dohertyi Fruhstorfer, 1902. TL: Ende Island (Indonesia: East Nusa Tenggara, Flores, Ende Island). TS:
ssp. baliensis Fruhstorfer, 1896
Elymnias protegenia baliensis Fruhstorfer, 1896. TL: Bali (Indonesia: Bali). TS:
Elymnias nigrescens bulelenga Rothschild, 1915. TL: Buleleng (Indonesia: Bali, Buleleng Regency). TS:
ssp. violetta Fruhstorfer, 1902
Elymnias undularis violetta Fruhstorfer, 1902. TL: Muok-Lek (Thailand: Saraburi, Muak Lek). TS:
Elymnias hypermnestra violetta f. epixantha Fruhstorfer, 1907. TL: Bangkok (Thailand: Bangkok). TS:
Elymnias hypermnestra violetta f. obfuscata Riley, 1932. TL: Siam (Thailand). TS:
ssp. meridionalis Fruhstorfer, 1902
Elymnias undularis meridionalis Fruhstorfer, 1902. TL: south Annam (southern Vietnam). TS:
Elymnias meridionalis f. orphnia, Fruhstorfer, 1907. TL: south Annam (southern Vietnam). TS:
ssp. beatrice Fruhstorfer, 1902. comb. n.
Elymnias nigrescens, Distant, 1882. Rhopalocera Malayana: 61.
Elymnias nigrescens beatrice Fruhstorfer, 1902. nomen n. for Distant’s nigrescens. TL: Singapore, Perak (Peninsular Malaysia: Perak), Lingga (Indonesia: Riau Islands, Lingga Archipelago, Lingga Island), Deli, (Indonesia: North Sumatra Province, Deli Serdang Regency), Sumatra (Indonesia: Sumatra), Wellesley Province (Peninsular Malaysia: Penang, Seberang Perai), Billiton (Indonesia: Bangka-Belitung Province, Belitung). TS:
Elymnias nigrescens ab. agina Fruhstorfer, 1902. unavailable name. TL: Singapore, Sumatra (Indonesia: Sumatra), Perak (Peninsular Malaysia: Perak). TS:
Elymnias nigrescens beatrice f. ornamenta Fruhsorfer, 1907. unavailable name. TL: Malay (Peninsular Malaysia). TS:
Elymnias hypermnestra agina, Corbet, 1943. Proc. Roy. Ent. Soc. Lond. (B) 12: 117–119.
ssp. sumbana Fruhstorfer, 1902
Elymnias nigrescens sumbana Fruhstorfer, 1902. TL: Sumba (Indonesia: East Nusa Tenggara, Sumba). TS:
ssp. decolorata Fruhstorfer, 1907
Elymnias nigrescens beatrice forma decolorata Fruhstorfer, 1907. unavailable name. TL: Sumatra (Indonesia: Sumatra). TS:
Elymnias hypermnestra decolorata, Aoki, Yamaguchi & Uémura, 1982. Butterflies of the Southeast Asian Islands 3: 175–176.
ssp. sumbawana Fruhstorfer, 1907
Elymnias nigrescens sumbawana Fruhstorfer, 1907. TL: Tambora, Sumbawa (Indonesia: West Nusa Tenggara, Sumbawa, Mt. Tambora). TS:
ssp. timorensis Fruhstorfer, 1907
Elymnias nigrescens timorensis Fruhstorfer, 1907. TL: Timor. TS:
ssp. alorensis Talbot, 1932
Elymnias nigrescens alorensis Talbot, 1932. TL: Alor (Indonesia: East Nusa Tenggara, Alor). TS:
ssp. nimota Corbet, 1937
Elymnias hypermnestra nimota Corbet, 1937. TL: Tioman (Peninsular Malaysia: Pahang, Rompin, Tioman Island). TS:
ssp. kangeana Aoki & Uémura, 1982
Elymnias hypermnestra kangeana Aoki & Uémura, 1982. TL: Kangean (Indonesia: East Java, Sumenap Regency, Kangean). TS: TPC. Mem. Tsukada Coll. 4: 2.
ssp. robinsona Monastyrskii & Devyatkin, 2003
Elymnias hypermnestra robinsona Monastyrskii & Devyatkin, 2003. TL: Con Dao, Con Son Island (Vietnam: Ba Ria–Vung Tau Province, Con Dao Archipelago, Con Son Island). TS:
ssp. jennifferae Suzuki, 2006. comb. n.
Elymnias cottonis jennifferae Suzuki, 2006. TL: Little Andaman (India: Andaman Islands, Little Andaman Island). TS: KMSPC. Futao (52): 13.
ssp. uemurai Lamas, 2010 (replaced Elymnias nigrescens meliophila Fruhstorfer, 1896a). Shilap 38 (150): 198.
Elymnias nigrescens meliophila Fruhstorfer, 1896a. TL: Lombok (Indonesia: West Nusa Tenggara, Lombok). TS:
caudata Butler, 1871
Specimens: Fig.
Elymnias caudata Butler, 1871. TL: Canara (India: Karnataka, Kanara). TS:
merula Swinhoe, 1915. incertae sedis
Specimen: Fig.
Elymnias merula Swinhoe, 1915. TL: Kandy, Ceylon (Sri Lanka: Central Province, Kandy). TS:
leucocyma Godart, 1819. incertae sedis
Distribution: Fig.
Biblis leucocyma Godart, 1819. TL: Java (Indonesia: Java). TS: unknown. Encyc. Méth. 9: 326.
nepheronides Fruhstorfer, 1907
Specimens: Fig.
ssp. nepheronides Fruhstoerfer, 1907
Elymnias nepheronides Fruhstorfer, 1907. TL: Flores Island (Indonesia: East Nusa Tenggara, Flores). TS:
Elymnias detanii Aoki & Uémura, 1982. TL: Flores (Indonesia: East Nusa Tenggara, Flores). TS:
ssp. tamborana Okubo, 2010
Elymnias tamborana Okubo, 2010. TL: Mt. Ngegep, Sumbawa (Indonesia: West Nusa Tenggara, Sumbawa, Mt. Sengenges). TS: OPC. Trans. Lep. Soc. Jpn. 60 (4): 255–257.
harterti Honrath, 1889
Specimens: Fig.
ssp. harterti Honrath, 1889
Elymnias harterti Honrath, 1889. TL: Perak (Peninsular Malaysia: Perak). TS:
ssp. brookei Shelford, 1904
Elymnias brookei Shelford, 1904. TL: Sarawak (East Malaysia: Sarawak). TS:
Elymnias smithi Moulton, 1915. syn. n. TL: Mt. Molu (East Malaysia: Sarawak, Mt. Molu). TS:
ssp. lautensis Medicielo & Hanafusa, 1994
Elymnias harterti lautensis Medicielo & Hanafusa, 1994. TL: Laut Island (Indonesia: South Kalimantan, Kota Baru Regency, Laut Island). TS: HPC. Futao (15): 17, pl. 4, figs 17–18.
ssp. arbaimuni Hanfusa, 2005
Elymnias haterti [sic] arbaimuni Hanfusa, 2005. TL: Indonesia: Jambi Province, Kuala Tungkal, Suban. TS: HPC. Futao (49): 11, pl. 1, figs 11–12.
parce Staudinger, 1889
Specimens: Fig.
ssp. parce Staudinger, 1889
Elymnias panthera parce Saudinger, 1889. TL: Palawan (Philippines: Palawan). TS:
ssp. justini Schröder & Treadaway, 2003
Elymnias parce justini Schröder & Treadaway, 2003. TL: Philippines: Palawan, Busuanga Island. TS: SMFD. Nachr. ent. Ver. Apollo 23 (4): 194, pl. 1, fig. 21.
panthera (Fabricius, 1787)
Specimens: Figs
ssp. panthera (Fabricius, 1787)
Papilio panthera Fabricius, 1787. TL: Tranquebariae (India: Tamil Nadu, Tharangambadi). TS:
ssp. dusara (Horsfield, [1829])
Melanitis dusara Horsfield, [1829]. TL: West Java (Indonesia: West Java). TS:
ssp. mimus Wood-Mason & de Nicéville, 1881
Elymnias mimus Wood-Mason & de Nicéville, 1881. TL: Nicobar Islands (India: Nicobar Islands). TS: uknown. J. Asiat. Soc. Bengal 50: 230.
ssp. dolorosa Butler, 1883.
Elymnias dolorosa Butler, 1883. TL: Nias Island (Indonesia: North Sumatra, Indonesia, Nias). TS:
ssp. lutescens Butler, 1867. comb. n., stat. n.
Elymnias lutescens Butler, 1867. TL: Malacca, Singapore and Penang (Singapore & Peninsular Malaysia: Penang and Malacca). TS:
Elymnias panthera var. labuana Staudinger, 1889. syn. n. TL: Labuan Island (East Malaysia: Labuan). TS:
Elymnias panthera lacrima Fruhstorfer, 1904. syn. n. TL: [North Borneo], [Banka] (Indonesia: Banka-Belitung Province, Banka Island). TS:
Elymnias defasciata Fruhstorfer, 1911, syn. n. TL: Borneo. TS: TS:
Elymnias panthera alfredi Fruhstorfer, 1907. syn. n. TL: Southeast Borneo. TS:
Elymnias panthera alfredi f. pantherina Fruhstorfer, 1907. unavailable name. TL: Southeast Borneo. TS:
Elymnias panthera alfredi f. alfredi Fruhstorfer, 1907. unavailable name. TL: Southeast Borneo. TS:
ssp. enganica Doherty, 1891
Elymnias enganica Doherty, 1891. TL: Engano (Indonesia: Bengkulu, Enggano Island). TS:
ssp. lacrimosa Fruhstorfer, 1898
Elymnias panthera lacrimosa Fruhstorfer, 1898. TL: Bawean Island (Indonesia: East Java, Gresik Regency, Bawean). TS:
ssp. suluana Fruhstorfer, 1899
Elymnais panthera suluana Fruhstorfer, 1899. TL: Sulu Island (Philippines: Sulu Province, Sulu Island). TS:
ssp. bangueyana Fruhstorfer, 1899
Elymnias panthera bangueyana Fruhstorfer, 1899. TL: Banguey Island (Malaysia: Sabah, Banggi Island). TS:
ssp. dulcibella Fruhstorfer, 1907
Elymnias panthera f. dulcibella Fruhstorfer, 1907. TL: East Java (Indonesia: East Java). TS:
ssp. tautra Fruhstorfer, 1907
Elymnias panthera tautra Fruhstorfer, 1907. TL: Northeast Sumatra (Indonesia: North Sumatra). TS:
ssp. arikata Fruhstorfer, 1907
Elymnias panthera arikata Fruhstorfer, 1907. TL: Natuna Island (Indonesia: Riau Province, Natuna Island). TS:
ssp. balina Martin, 1909
Elymnias panthera balina Martin, 1909. TL: Bali Island (Indonesia: Bali). TS: NHMT. Dt. ent. Z. Iris 22 (1): 58.
ssp. exsulata van Eecke, 1918
Elymnias panthera exsulata van Eecke, 1918. TL: Pulu [sic] Lasia (Indonesia: North Sumatra, Lasia Island). TS: NBC. Zoologische Mededeelingen 4 (2): 82.
ssp. winkleri Kalis, 1933
Elymnias panthera winkleri Kalis, 1933. TL: Sabang, Weh Island (Indonesia: Aceh, Sabang, Weh Island). TS: MEPR. Tijdschrift voor Entomologie 76 (1–2): 80.
ssp. mira Corbet, 1942
Elymnias panthera mira Corbet, 1942. TL: Sipora Island (Indonesia: West Sumatra, Mentawai Regency, Sipora). TS:
ssp. tiomanica Eliot, 1978
Elymnias panthera tiomanica Eliot, 1978. TL: Tioman (Peninsular Malaysia: Pahang, Rompin, Tioman Island). TS:
ssp. belitungensis Okano, 1986
Elymnias panthera belitungensis Okano, 1986. TL: Belitung Island (Indonesia: Bangka-Belitung Province, Belitung). TS: OMPC. Tokurana 11 (1): 1, figs 1–6.
ssp. ruricolaris Hanafusa, 1989
Elymnias panthera ruricolaris Hanafusa, 1989. TL: Karimata Island (Indonesia: West Kalimatan Province, Karimata Island). TS: HPC. Futao (3): 10, pl. 3, figs 1–4.
ssp. banyakensis Hanafusa, 1993
Elymnias panthera banyakensis Hanafusa, 1993. TL: Kepulauan Banyak (Indonesia: Aceh, Banyak Islands). TS: HPC. Futao (13): 8.
ssp. attenuata Hanafusa, 1994
Elymnias panthera attenuata Hanafusa, 1994. TL: Tanahmasa Island (Indonesia: North Sumatra Province, Tanahmasa Island). TS: HPC. Futao (4): 13.
ssp. redangensis Hanafusa, 2001
Elymnias panthera redangensis Hanafusa, 2001. TL: Redang Island (Peninsular Malaysia: Terengganu, Redang Island). TS: HPC. Futao (37): 14, pl.1, figs 5–8.
ssp. zeta Abang, Treadaway & Schröder, 2004
Elymnias panthera zeta Abang, Treadaway & Schröder, 2004. TL: Balambangan Island (East Malaysia: Sabah, Balambangan Island). TS: MUS. Futao (47): 10, pl. 3, figs 33–36.
obnubila Marshall & de Nicéville, 1883
Specimens: Fig.
Elymnias obnubila Marshall & de Nicéville, 1883. TL: Mergui (Myanmar: Thanintharyi, Mergui Archipelago). TS:
congruens Semper, 1887
Specimens: Fig.
ssp. congruens Semper, 1887
Elymnias congruens Semper, 1887. TL: N. Mindanao (Philippines: northern Mindanao). TS: SMFD. Reisen. Philipp. 2: 61, pl. 11, fig. 8–10.
Elymnias congruens photinus Fruhstorfer, 1907. syn. n. TL: N. Mindanao (Philippines: northern Mindanao). TS:
Elymnias congruens phaios Fruhstorfer, 1907. syn. n. TL: S. Mindanao (Philippines: southern Mindanao). TS:
Elymnias congruens rafaela Fruhstorfer, 1907. syn. n. TL: Bazilan (Philippines: Sulu Archipelago, Basilan). TS:
ssp. subcongruens Semper, 1892
Elymnias subcongruens Semper, 1892. TL: Mindoro (Philippines: Mindoro). TS: SMFD. Reisen. Philipp. 7: 329.
ssp. endida Fruhstorfer, 1911
Elymnias congruens endida Fruhstorfer, 1911. TL: Bohol (Philippines: Bohol). TS:
ssp. salipi Schroeder & Treadaway, 1989
Elymnias salipi Schroeder & Treadaway, 1989. TL: Philippines: Tawi-Tawi Archipelago, Sanga Sanga Island, Boloboc. TS: SMFD. Ent. Z. 99 (22): 327, fig. 6.
ssp. jekei Schroeder & Treadaway, 1989
Elymnias jekei Schroeder & Treadaway, 1989. TL: Philippines: Luzon, Nueva Ecija, near Carranglan. TS: SMFD. Ent. Z. 99 (22): 328, fig. 6.
ssp. neergaardorum Schroeder & Treadaway, 2003
Elymnias neergaardorum Schroeder & Treadaway, 2003. TL: Phillipines: Masbate. TS: SMFD. Nachr. ent. Ver. Apollo 23 (4): 194, pl. 1, figs 14–15.
miyagawai Saito & Kishi, 2012
Specimens: Fig.
Elymnias miyagawai Saito & Kishi, 2012. TL: Vietnam: Lam Dong. TS: SPC. Butterflies (62): 4, figs 1–2, 10.
nesaea (Linnaeus, 1764)
Specimens: Fig.
ssp. nesaea (Linnaeus, 1764)
Papilio (Nymphalis) nesaea Linnaeus, 1764. TL: [Java] (Indonesia: Java). TS: LSL. Mus. Lud. Ulr. Reg.: 302.
Papilio lais Cramer, 1777. TL: Java (Indonesia: Java). TS: unknown. Uitl. Kapellen 2 (10): 21, pl. 110, f. A–B.
Elymnias nesaea hermia Fruhstorfer, 1907. syn. n. TL: near Lawang, (Indonesia: East Java, Lawang). TS:
ssp. timandra Wallace, 1869
Elymnias timandra Wallace, 1869. TL: Sylhet (Bangladesh: Sylhet Division), Moulmein (Myanmar: Mon State, Mawlamyine). TS:
Elymnias nesaea cortona Fruhstorfer, 1911. syn. n. TL: Burma (Myanmar). TS:
ssp. laisidis de Nicéville, 1896
Elymnias (Melynias) laisidis de Nicéville, 1896. TL: Sumatra (Indonesia: Sumatra). TS:
ssp. baweana Hagen, 1896
Elymnias baweana Hagen, 1896. TL: Bawean Island (Indonesia: East Java, Gresik, Bawean). TS: NHMT. Jahrb. Nass. Nat. 49: 184, pl. 4, fig. 6.
ssp. neolais de Nicéville, 1898
Elymnias (Melynias) neolais de Nicéville, 1898. TL: Nias Island (Indonesia: North Sumatra, Nias). TS:
ssp. apelles Fruhstorfer, 1902
Elymnias lais apelles Fruhstorfer, 1902. TL: Bangkok (Thailand: Bangkok). TS:
ssp. vordemani Snellen van Vollenhoven, 1902
Elymnias vordemani Snellen van Vollenhoven, 1902. TL: Kangean Island (Indonesia: East Java, Sumenap, Kangean). TS: NBC. Tijdschr. Ent. 45: 77, pl. 8, fig. 1.
ssp. hypereides Fruhstorfer, 1903 36
Elymnias lais hypereides Fruhstorfer, 1903. TL: North Borneo. TS:
Elymnias nesaea coelifrons Fruhstorfer, 1907. syn. n. TL: Southeast Borneo (Indonesia: South or East Kalimantan). TS:
ssp. kamarina Fruhstorfer, 1906
Elymnias lais kamarina Fruhstorfer, 1906. TL: Batu Island (Indonesia: North Sumatra, South Nias Regency, Batu Islands). TS:
ssp. lioneli Fruhstorfer, 1907
Elymnias nesaea lioneli Fruhstorfer, 1907. TL: Malaysia. TS:
ssp. tawicola Schröder & Treadaway, 1989
Elymnias nesaea tawicola Schröder & Treadaway, 1989. TL: Philippines: Tawi-Tawi Archipelago, Sibutu Island, Cavan Cavan. TS: SMFD. Ent. Z. 99 (22): 326, fig. 4.
casiphone Geyer, [1827]
Specimens: Fig.
ssp. casiphone Geyer, [1827]
Elymnias casiphone Geyer, [1827]. TL: not indicated. TS: unknown. Samml. exot. Schmett. 3: pl. [9], f. 1–2.
Elymnias kamara Moore, [1858]. syn. n. TL: Java (Indonesia: Java). TS:
Elymnias kamara pareuploea Fruhstorfer, 1911. TL: [Java] (Indonesia: Java). TS:
Elymnias kamara pseudalumna Fruhstorfer, 1911. TL: Java (Indonesia: Java). TS:
ssp. erinyes de Nicéville, 1895. comb. rev.
Elymnias (Melynias) erinyes de Nicéville, 1895. TL: Battak Mountains (Indonesia: North Sumatra). TS:
ssp. praetextata Fruhstorfer, 1896
Elymnias casiphone praetextata Fruhstorfer, 1896. TL: Lombok (Indonesia: West Nusa Tenggara, Lombok). TS:
Elymnias kamara lombokiana Fruhstorfer, 1911. syn. n. TL: Lombok Island (Indonesia: West Nusa Tenggara, Lombok). TS:
ssp. exclusa de Nicéville, 1898. comb. n.
Elymnias (Melynias) exclusa de Nicéville, 1898. TL: Bali (Indonesia: Bali). TS:
Elymnias casiphone djilantik Martin, 1909, syn. n. TL: Bali (Indonesia: Bali). TS: NHMT. Dt. ent. Z. Iris 22 (1): 49.
ssp. alumna Fruhstorfer, 1907
Elymnias casiphone alumna Fruhstorfer, 1907. TL: East Java (Indonesia: East Java). TS:
malelas (Hewitson, 1863)
Specimens: Fig.
ssp. malelas (Hewitson, 1863)
Melanitis malelas Hewitson, 1863. TL: East India. TS:
Elymnias malelas malelas ab. subdecorata Fruhstorfer, 1911. unavailable name. TL: Assam (India: Meghalaya). TS:
Elymnias malelas ivena Fruhstorfer, 1911. syn. n. TL: Thailand, N. Vietnam. TS:
Elymnias malelas nilamba Fruhstorfer, 1911. syn. n. TL: India. TS:
saueri Distant, 1882
Specimens: Fig.
ssp. saueri Distant, 1882
Elymnias saueri Distant, 1882. TL: Malaysia, Province Wellesley (Peninsular Malaysia: Penang, Seberang Perai). TS:
kochi Semper, 1887
Specimens: Fig.
ssp. kochi Semper, 1887
Elymnias kochi Semper, 1887. TL: Philippines: Central Luzon. TS: SMFD. Reisen Philipp. (2) 55: 63, pl. 12, fig. 4.
casiphonides Semper, 1892
Specimens: Fig.
ssp. casiphonides Semper, 1892
Elymnias casiphonides Semper, 1892. TL: Philippines: Mindanao. TS: SMFD. Reisen Philipp. (7): 330.
ssp. sanrafaela Schröder & Treadaway, 1980
Elymnias casiphonides sanrafaela Schröder & Treadaway, 1980. TL: Philippines: Samar, San Rafael TS: SMFD. Ent. Z. 90 (21): 238, fig. 3.
nelsoni Corbet, 1942
Specimens: Fig.
ssp. nelsoni Corbet, 1942
Elymnias nelsoni Corbet, 1942. TL: Mentawei Islands (Indonesia: West Sumatra, Mentawai Islands). TS:
amoena Tsukada & Nishiyama, 1979
Specimens: Fig.
ssp. amoena Tsukada & Nishiyama, 1979
Elymnias amoena Tsukada & Nishiyama, 1979. TL: Sumba (Indonesia: East Nusa Tenggara, Sumba). TS: TPC. Mem. Tsukada Coll. 1: 15, figs 19–20.
kanekoi Tsukada & Nishiyama, 1980
Specimens: Fig.
ssp. kanekoi Tsukada & Nishiyama, 1980
Elymnias kanekoi Tsukada & Nishiyama, 1980. TL: north Negros (Philippines: Negros Occidental). TS: TPC. Mem. Tsukada Coll. 2: 14, f. 8–9, 14
saola Monastyrskii, 2004
Specimens: Fig.
ssp. saola Monastyrskii, 2004
Elymnias saola Monastyrskii, 2004. TL: Vietnam: Nghe An Province, Pu Mat Nature Reserve. TS:
melias (C. & R. Felder, 1863)
Specimens: Fig.
ssp. melias (C. & R. Felder, 1863)
Melanitis melias C. & R. Felder, 1863. TL: Lugban (Philippines: Luzon, Quezon, Lucban) and Burias Island (Philippines: Masbate, Burias Island). TS:
ssp. malis Semper, 1887
Elymnias melias malis Semper, 1887. TL: Casiguran (Philippines: Central Luzon, Aurora, Casiguran). TS: SMFD. Reisen Philipp. (2): 62, pl. 12, figs 2–3.
Elymnias palmifolia Schultze, 1908. TL: Cagayang (Philippines: Northern Luzon, Cagayan). TS: ECMP. Philipp. J. Sci 3 (1): 27, pl. 1, fig. 1.
beza (Hewitson, 1877)
Specimens: Fig.
ssp. beza (Hewitson, 1877)
Melanitis beza Hewitson, 1877. TL: Philippines: Mindanao. TS:
Elymnias kochi plateni Fruhstorfer, 1907. syn. n. TL: Philippines: Mindanao. TS:
ssp. samarana Schröder & Treadaway, 1980
Elymnias beza samarana Schröder & Treadaway, 1980. TL: Philippines: Samar, San Rafael. TS: SMFD. Ent. Z. 90 (21): 236, fig. 2.
sansoni Jumalon, 1975
Specimens: Fig.
ssp. sansoni Jumalon, 1975
Elymnias sansoni Jumalon, 1975. TL: Philippines: Negros. TS: JPC. Trans. Lep. Soc. Jpn. 26 (2): 47.
ssp. aklanensis Uémura & Kitamura, 2001
Elymnias sansoni aklanensis Uémura & Kitamura, 2001. TL: Philippines: Panay, Aklan Province, Makato, Castillo. TS: TME. Butterflies 29: 5.
luteofasciata Okubo, 1980
Specimens: Fig.
Elymnias luteofasciata Okubo, 1980. TL: Philippines: Mindanao, Davao, Penangudloton, Upian River, Calinan. TS: OPC. Tyô to Ga 31 (1,2): 60.
vitellia (Stoll, [1781])
Specimens: Fig.
ssp. vitellia (Stoll, [1781])
Papilio vitellia Stoll, [1781]. TL: Ambon (Indonesia: Maluku, Ambon). TS: unknown. Uitl. Kapellen. 4 (30): 116, pl. 349, fig. E–F.
Melanitis stellaris Snellen van Vollenhoven, 1861. TL: [New Guinea]. TS: NBC. Tijdschr. Ent. 4 (5/6): 159, pl. 8, fig. 3.
Elymnias vitellia f. basium Fruhstorfer, 1907. unavailable name. TL: Saparua (Indonesia: Maluku, Saparau). TS:
Elymnias vitellia ceramensis Martin, 1909. TL: Ceram (Indonesia: Maluku, Seram). TS: NHMT. Dt. ent. Z. Iris 22 (1): 65.
ssp. viminalis Wallace, 1869
Elymnias viminalis Wallace, 1869. TL: Buru Island (Indonesia: Maluku, Buru). TS:
agondas (Boisduval, 1832)
Specimens: Figs
ssp. agondas (Boisduval, 1832)
Dyctis agondas Boisduval, 1832. TL: Vanikoro (Solomon Islands: Temotu Province, Vanikoro). TS: unknown. Voy. Astrolabe. 1: 138.
Dyctis bioculatus Westwood, 1851. syn. n. TL: Arfak Mountains (Indonesia: West Papua). TS:
Elymnias agondas muscosa Fruhstorfer, 1907. TL: Kapaur (Indonesia: West Papua, Fakfak). TS:
Elymnias agondas tampyra Fruhstorfer, 1907. TL: Kumusi River (Papua New Guinea: Northern Province, Kumusi River). TS:
Elymnias agondas hagias Fruhstorfer, 1914. TL: Eilandenfluß (Indonesia: Papua, Pulau River). TS:
ssp. melane (Hewitson, 1858)
Melanitis melane Hewitson, 1858. TL: [Key Island] (Indonesia: Maluku, Kei Island). TS:
Elymnias (Dyctis) mela de Nicéville, 1902. TL: Key Island (Indonesia: Maluku, Kei Island). TS:
Elymnias (Dyctis) meletus de Nicéville, 1902. TL: Key Island (Indonesia: Maluku, Kei Island). TS:
Elymnias (Dyctis) melitia de Nicéville, 1902. TL: Key Island (Indonesia: Maluku, Kei Island). TS:
ssp. melantho Wallace, 1869
Elymnias melantho Wallace, 1869. TL: Gagie Island (Indonesia: West Papua, Raja Ampat Regency, Gag Island). TS:
Elymnias agondas moranda Fruhstorfer, 1904. TL: Waigeu (Indonesia: West Papua, Raja Ampat Regency, Waigeo). TS:
ssp. glaucopis Staudinger, 1894
Elymnias glaucopis Staudinger, 1894. TL: Sattelberg (Papua New Guinea: Morobe Province, Huon Peninsula, Sattelberg). TS:
Elymnias agondas glaucopis Fruhsforfer, 1907. TL: New Guinea. TS:
ssp. melanippe Grose-Smith, 1894
Elymnias melanippe Grose-Smith, 1894. TL: Sattelberg (Papua New Guinea: Morobe Province, Huon Peninsula, Sattelberg). TS:
Elymnias vertenteni Hulstaert, 1925. TL: Irian Jaya (Indonesia: Papua or West Papua). TS: NBC. Ann. Mag. nat. Hist. (9) 15 (88): 447.
ssp. melanthes Grose-Smith & Kirby, 1897
Elymnias melanthes Grose-Smith & Kirby, 1897. TL: Woodlark Island (Papua New Guinea: Milne Bay, Woodlark Island). TS:
Elymnias agondas melanthes f. infernalis (♀) Fruhstorfer, 1914. TL: Not indicated. TS:
Elymnias agondas melanthes f. virginalis (♀) Fruhstorfer, 1914. TL: Not indicated. TS:
ssp. melagondas Fruhstorfer, 1900
Elymnias melagondas Fruhstorfer, 1900. TL: New Guinea. TS:
Elymnias agondas melagondas f. taenarides (♀) Fruhstorfer, 1914. TL: Milnebai (Papua New Guinea: Milne Bay). TS:
ssp. australiana Fruhstorfer, 1900
Elymnias australiana Fruhstorfer, 1900. TL: Cape York (Australia: Queensland, Cape York). TS:
ssp. aruana Fruhstorfer, 1900
Elymnias aruana Fruhstorfer, 1900. TL: Aru (Indonesia: Maluku, Indonesia). TS:
ssp. goramensis Fruhstorfer, 1900
Elymnias goramensis Fruhstorfer, 1900. TL: Goram Island (Indonesia: Maluku, East Seram Regency, Gorong Island). TS:
ssp. agondina Fruhstorfer, 1904
Elymnias agondina Fruhstorfer, 1904. TL: Salewatti (Indonesia: West Papua, Raja Ampat Islands, Salawati). TS:
ssp. dampierensis Rothschild, 1915
Elymnias dampierensis Rothschild, 1915. TL: Dampier (Papua New Guinea: Madang, Karkar Island). TS: NHMT. Novit. Zool. 22 (2): 201.
ssp. multocellata van Eecke, 1915
Elymnias multocellata van Eecke, 1915. TL: Kloofbivak (Indonesia: Papua). TS: NBC. Nova Guinea 13 (1): 66, pl. 3, f. 6.
ssp. thryallis Kirsch, 1876. comb. n.
Elymnias thryallis Kirsch, 1876. TL: Mysore, Kordo (Indonesia: Papua, Biak). TS: SMTD. Mitt. zool. Mus. Dresden 1: 119, pl. 6, fig. 4.
Elymnias glauconia Staudinger, 1894. TL: Kubary (Papua New Guinea: Jiwaka, Mt. Kubari). TS:
Elymnias glauconia var. chloera Staudinger, 1894. TL: New Guinea. TS:
Elymnias thryallis f. brunnescens Fruhstorfer, 1911. TL: New Guinea. TS:
Elymnias thryallis f. pseudosalpinx Fruhstorfer, 1911. TL: New Guinea. TS:
Elymnias thryallis f. terentilina Fruhstorfer, 1911. TL: New Guinea. TS:
Elymnias thryallis f. violacea Fruhstorfer, 1911. TL: Waigiu Island (Indonesia: West Papua, Raja Ampat Regency, Waigeo). TS:
cybele (C. & R. Felder, 1860)
Specimens: Fig.
ssp. cybele (C. & R. Felder, 1860)
Melanitis cybele C. & R. Felder, 1860. TL: Batschian Island (Indonesia: North Maluku, Bacan). TS:
Dyctis astrifera Butler, 1874. TL: Batchian (Indonesia: North Maluku, Bacan). TS:
Elymnias cybele opaca Fruhstorfer, 1907. syn. n. TL: Halmaheira (Indonesia: North Maluku, Halmahera). TS:
Elymnias cybele ternatana syn. n. Fruhstorfer, 1907. TL: Ternate (Indonesia: North Maluku, Ternate). TS:
ssp. obiana Fruhstorfer, 1904
Elymnias obiana Fruhstorfer, 1904. TL: Obi Island (Indonesia: North Maluku, Obi). TS:
ssp. adumbrata Fruhstorfer, 1907. subsp. rev.
Elymnias cybele adumbrata Fruhstorfer, 1907. TL: Buru (Indonesia: Maluku, Buru). TS:
cumaea (C. & R. Felder, [1867])
Specimens: Fig.
ssp. cumaea (C. & R. Felder, [1867])
Melanitis cumaea C. & R. Felder, [1867]. TL: Halmahera (Indonesia: North Maluku Halmahera). TS:
ssp. thyone Fruhstorfer, 1904. comb. n., stat. n.
Elymnias thyone Fruhstorfer, 1904. TL: [North Celebes] (Indonesia: North Sulawesi, Indonesia). TS:
ssp. toliana Fruhstorfer, 1899
Elymnias cumaea toliana Fruhstorfer, 1899. TL: Toli Toli (Indonesia: Central Sulawesi, Tolitoli). TS:
Elymnias pseudeuploea Fruhstorfer, 1911. unavailable name. TL: Sulawesi (Indonesia: Sulawesi). TS:
hewitsoni Wallace, 1869
Specimens: Fig.
ssp. hewitsoni Wallace, 1869
Elymnias hewitsoni Wallace, 1869. TL: Macassar (Indonesia: South Sulawesi, Makassar). TS:
Elymnias hewitsoni hewitsoni f. sumptuosa Fruhstorfer, 1907. TL: Tanetta (Indonesia: Central Sulawesi, Poso Regency, Tentena). TS:
ssp. meliophila Fruhstorfer, 1896
Elymnias meliophila Fruhstorfer, 1896. TL: Saleyer (Indonesia: South Sulawesi, Selayar Islands, Selayar). TS:
ssp. atys Fruhstorfer, 1904
Elymnias hewitsoni atys Fruhstorfer, 1904. TL: Bouthain, south Celebes (Indonesia: South Sulawesi, Moncong Lompobatang). TS:
mimalon (Hewitson, 1861)
Specimens: Fig.
ssp. mimalon (Hewitson, 1861)
Melanitis mimalon (Hewitson, 1861). TL: Toli-Toli (Indonesia: Central Sulawesi, Tolitoli). TS:
Elymnias mimalon mimalon f. leucostigmata Fruhstorfer, 1907. TL: Toli-Toli (Indonesia: Central Sulawesi, Tolitoli). TS:
ssp. ino Fruhstorfer, 1904
Elymnias mimalon ino Fruhstorfer, 1904. TL: Tawaya, Celebes (Indonesia: Central Sulawesi, Towaya). TS:
ssp. nysa Fruhstorfer, 1907
Elymnias mimalon nysa Fruhstorfer, 1907. TL: South Celebes (Indonesia: Southeast Sulawesi). TS:
hicetas Wallace, 1869
Specimens: Fig.
ssp. hicetas Wallace, 1869
Elymnias hicetas Wallace, 1869. TL: Macassar, south Celebes (Indonesia: South Sulawesi, Makassar). TS:
Elymnias hicetas bonthainensis Fruhstorfer, 1899. syn. n. TL: Bua Kraeng (Indonesia: South Sulawesi, Mt. Bawakaraeng). TS:
ssp. hicetina Fruhstorfer, 1904
Elymnias hicetas hicetina Fruhstorfer, 1904. TL: Tombugu (Indonesia: Central Sulawesi, Tombuko). TS:
ssp. butona Fruhstorfer, 1904
Elymnais hicetas butona Fruhstorfer, 1904. TL: North Buton (Indonesia: Southeast Sulawesi, Buton). TS:
ssp. rarior Martin, 1929
Elymnias hicetas rarior Martin, 1929. TL: Celebes (Indonesia: Sulawesi). TS: NHMT. Mitt. münchn. ent. Ges. 19: 160.
holofernes (Butler, 1882)
Specimens: Fig.
Dyctis holofernes Butler, 1882. TL: Duke-of-York Island (Papua New Guinea: East New Britain, Duke of York Island). TS:
bornemanni Ribbe, 1889. stat. n.
Specimens: Fig.
Elymnias bornemanni Ribbe, 1889. TL: Bangkai (Indonesia: Central Sulawesi, Banggai). TS: SMTD (?). Dt. ent. Z. Iris 2 (1): 183, pl. 3, f. 1–2.
phrikonis Fruhstorfer, 1899. stat. n.
Specimens: Fig.
Elymnias cumaea phrikonis Fruhstorfer, 1899. TL: Sula Besi and Sula-Mangoli (Indonesia: North Maluku, Sula Islands, Sanana and Mangole). TS:
Elymnias cumaea relicina Fruhstorfer, 1907. syn. n. TL: Sula Besi (Indonesia: North Maluku, Sula Islands, Sanana). TS:
sangira Fruhstorfer, 1899
Specimens: Fig.
Elymnias cumaea sangira Fruhstorfer, 1899. TL: Sangir, Sulawesi (Indonesia: North Sulawesi, Sangihe Islands, Sangir Besar). TS:
umbratilis Joicey & Noakes, 1915. stat. n.
Specimens: Fig.
Elymnias cybele umbratilis Joicey & Noakes, 1915. TL: Biak (Indonesia: Papua, Biak). TS:
resplendens Martin, 1929. stat. n.
Specimens: Fig.
Elymnias cumaea resplendens Martin, 1929. TL: Celebes (Indonesia: Sulawesi). TS: NHMT. Mitt. münchn. ent. Ges. 19: 162.
singhala Moore, [1875]
Specimens: Fig.
Elymnias singhala Moore, [1875]. TL: Colombo, Ceylon (Sri Lanka: Western Province, Colombo). TS:
Wing patterns of Elymnias butterflies appear to be highly evolvable, which facilitates Batesian mimetic resemblance to a variety of phenotypically dissimilar model species. Many Elymnias are found on islands in the Indo-Australian Archipelago, and the isolation provided by islands seems to provide the opportunity for divergence and local adaptation, facilitating resemblance to different model species in different locales. The remarkable capacity for phenotypic evolution of wing patterns has resulted in sexually dimorphic mimicry, convergence of distantly related taxa on similar wing patterns, and marked phenotypic divergence among conspecific popuations. These phenomena have previously confounded attempts to produce an accurate taxonomic framework because few if any morphological characters are taxonomically or phylogenetically informative. Wing veination, male genitalia, and female genitalia are remarkably uniform among species of Elymnias; only slight variation in male genitalia might be useful for discriminating some species. Species delimitation and diagnosis in Elymnias has therefore traditionally relied almost entirely on wing patterns. Our molecular phylogeny, which uses genetic markers presumed to be unrelated to wing phenotypes, has detected multiple instances of similar wing patterns in non-sister Elymnias lineages that mimic the same, widespread model species. This similarity seems to be the result of convergent evolution, and we have therefore split these taxa into two or more monophyletic lineages (e.g., E. cumaea and E. cybele have each been split into four and three different species, respectively). On the other hand, some Elymnias species—like other mimetic butterfly taxa (
Strong dimorphism caused many early workers to describe males and females as separate species, most of which have been synonymized. In this paper we confirmed
Females of several Elymias species, including E. agondas, E. hypermnestra, and E. esaca, are morphologically variable across their range. Rather than recognize every wing pattern variant as a different subspecies, we have synonymized many subspecies into geographically cohesive taxa, for example, within the islands of Borneo or New Guinea.
Much of the mismatch between Elymnias’ previous taxonomic framework and its evolutionary history is due to rapid evolutionary change. This resulted in morphologically-delimited nominal species that were polyphyletic. In these cases, our molecular phylogenetic results make delimiting species relatively straightforward. However, there are several cases that are not as clear-cut. For example, we elected to retain E. esaca and E. vasudeva as distinct species despite their paraphyletic relationship because of marked, species-specific morphological differences in these two parapatrically distributed taxa. Population genetic theory predicts incomplete lineage sorting of genetic loci to persist for some time after speciation, resulting in paraphyletic species; the probability of reciprocal monophyly increases with time since divergence (
Our molecular phylogeny identifies several examples of allopatrically or parapatrically distributed populations that form distinct, monophyletic sister groups: E. sansoni sansoni on Negros and E. sansoni aklanensis on Panay; E. patna from India and E. patna from peninsular Malaysia; E. vitellia vitellia from Seram and E. vitellia viminalis from Buru; and E. hypermnestra from Java and the Lesser Sundas and E. hypermnestra from everywhere else. These monophyletic sister lineages would likely be considered different species under a strict phylogenetic species concept, and, in most cases, preliminary Bayesian species delimitation analyses with the program Bayesian Phylogenetics and Phylogeography (BPP;
Although there is one African and several mainland Asian species, most of Elymnias’ diversity is found on the islands of the Indo-Australian Archipelago. Islands are considered laboratories for the study of evolution because they promote isolation and divergence while simplifying the task of delimitating populations and other taxa. Evolutionary study of this taxon provides an excellent opportunity to study the role of archipelagoes in diversification, and the evolutionary genetics of evolutionary novelty and speciation.
New synonyms
Elymnias papua bivittata van Eecke, 1915, of Elymnias papua papua Wallace, 1869
Elymnias (Mimadelias) esaca taeniola Fruhstorfer, 1907, of Elymnias esaca borneensis, Wallace, 1869
Elymnias (Mimadelias) oberthuri Fruhstorfer, 1902, of Elymnias esaca andersonii (Moore, 1886)
Elymnias thycana Wallace, 1869, of Elymnias vasudeva vasudeva Moore, 1857
Mimadelias deva Moore, 1894, of Elymnias vasudeva vasudeva Moore, 1857
Mimadelias burmensis Moore, 1893, of Elymnias vasudeva vasudeva Moore, 1857
Elymnias vacudera [sic] sinensis Chou, Zhang & Xie, 2000, of Elymnias vasudeva vasudeva Moore, 1857
Melanyias patnoides Moore, 1893, of Elymnias patna patna (Westwood, 1851)
Elymnias patna stictica Fruhstorfer, 1902, of Elymnias patna patna (Westwood, 1851)
Elymnias kuenstleri mariae Toxopeus, 1936, of Elymnias kuenstleri Honrath, [1885]
Elymnias nigrescens tonkiniana Fruhstorfer, 1902, of Elymnias hypermnestra hainana Moore, 1878
Elymnias hypermnestra nigrescens f. depicta Fruhstorfer, 1907, of Elymnias hypermnestra hainana Moore, 1878
Elymnias hypermnestra septentrionalis Chou & Huang, 1994, of Elymnias hypermnestra hainana Moore, 1878
Elymnias smithi Moulton, 1915, of Elymnias harterti brookei Shelford, 1904
Elymnias panthera var. labuana Staudinger, 1889, of Elymnias panthera lutescens Butler, 1867
Elymnias panthera lacrima Fruhstorfer, 1904, of Elymnias panthera lutescens Butler, 1867
Elymnias defasciata Fruhstorfer, 1911, of Elymnias panthera lutescens Butler, 1867
Elymnias panthera alfredi Fruhstorfer, 1907, of Elymnias panthera lutescens Butler, 1867
Elymnias congruens photinus Fruhstorfer, 1907, of Elymnias congruens congruens Semper, 1887
Elymnias congruens phaios Fruhstorfer, 1907, of Elymnias congruens congruens Semper, 1887
Elymnias congruens rafaela Fruhstorfer, 1907, of Elymnias congruens congruens Semper, 1887
Elymnias nesaea hermia Fruhstorfer, 1907, of Elymnias nesaea nesaea (Linnaeus, 1764)
Elymnias nesaea cortona Fruhstorfer, 1911, of Elymnias nesaea timandra Wallace, 1869
Elymnias nesaea coelifrons Fruhstorfer, 1907, of Elymnias nesaea hypereides Fruhstorfer, 1903
Elymnias kamara Moore, [1858], of Elymnias casiphone casiphone Geyer, [1827]
Elymnias kamara lombokiana Fruhstorfer, 1911, of Elymnias casiphone praetextata Fruhstorfer, 1896
Elymnias casiphone djilantik Martin, 1909, of Elymnias casiphone exclusa de Nicéville, 1898
Elymnias malelas ivena Fruhstorfer, 1911, of Elymnias malelas malelas (Hewitson, 1863)
Elymnias malelas nilamba Fruhstorfer, 1911, of Elymnias malelas malelas (Hewitson, 1863)
Elymnias kochi plateni Fruhstorfer, 1907, of Elymnias beza beza (Hewitson, 1877)
Dyctis bioculatus Westwood, 1850, of Elymnias agondas agondas (Boisduval, 1832)
Elymnias cybele opaca Fruhstorfer, 1907, of Elymnias cybele cybele (C. & R. Felder, 1860)
Elymnias cybele ternatana Fruhstorfer, 1907, of Elymnias cybele cybele (C. & R. Felder, 1860)
Elymnias hicetas bonthainensis Fruhstorfer, 1899, of Elymnias hicetas hicetas Wallace, 1869
Elymnias cumaea relicina Fruhstorfer, 1907, of Elymnias phrikonis Fruhstorfer, 1899
New combinations
Elymnias hypermnestra cottonis (Hewitson, 1874) (Melanitis cottonis)
Elymnias hypermnestra beatrice Fruhstorfer, 1902 (Elymnias nigrescens beatrice)
Elymnias hypermnestra jennifferae Suzuki, 2006 (Elymnias cottonis jennifferae)
Elymnias panthera lutescens Butler, 1867 (Elymnias lutescens)
Elymnias casiphone exclusa de Nicéville, 1898 (Elymnias (Melynias) exclusa)
Elymnias agondas thryallis Kirsch, 1876 (Elymnias thryallis)
Elymnias cumaea thyone Fruhstorfer, 1904 (Elymnias thyone)
Resurrected combination
Elymnias casiphone erinyes de Nicéville, 1895
Resurrected subspecies
Elymnias cybele adumbrata Fruhstorfer, 1907
Status changes
Elymnias ceryxoides de Nicéville, 1895 stat. rev.
Elymnias panthera lutescens Butler, 1867 stat. n.
Elymnias cumaea thyone Fruhstorfer, 1904 stat. n.
Elymnias bornemanni Ribbe, 1889 stat. n.
Elymnias phrikonis Fruhstorfer, 1899 stat. n.
Elymnias umbratilis Joicey & Noakes, 1915 stat. n.
Elymnias resplendens Martin, 1929 stat. n.
Incertae sedis
Elymnias merula Swinhoe, 1915
Elymnias leucocyma Godart, 1819
Species not placed in any group
Elymnias singhala Moore, [1875]
Format of each legend for specimen figures (1–22):
valid species or subspecies name_♂♀_dorsal/ventral_specimen repository_current name of collection locality (Country: State/Province, Locality). D = dorsal; V = Ventral; ♂ = male ♀ = female.
Format of each legend for male genitalia figures (22–26):
valid species or subspecies_specimen repository_current locality name.
See pages 4–5 for abbreviations of specimen repositories.
Each distribution map (Figs
A bammakoobammakoo ♂ D
A esacaleontina ♂ D+V
A darabengena ♂ D
A kuenstlerikuenstleri ♂ D+V
A hypermnestrahypermnestra ♂ D+V
A hypermnestratinctoria ♂ D
A hypermnestrabaliensis ♂ D
A hypermnestrasumbawana ♂ D
A hartertiharterti ♂ D+V OPC Malaysia: Perak, Batang Padang, Tapah B hartertiharterti ♀ D+V OPC Malaysia: Perak, Batang Padang, Tapah C hartertibrookei ♂ D
A pantherapanthera ♂ D
A obnubila ♂ D+V
A nesaeanesaea ♂ D
A casiphonecasiphone ♂ D+V
A malelas ♂ D
A meliasmelias ♂ D
A agondasagondas ♂ D+V
A agondasmelanthes ♂ D+V
A agondasmelagondas ♂ D+V
A agondasdampierensis ♂ D
A cybelecybele ♂ D
A holofernes ♂ D
A bammakoobammakoo
A hypermnestrahypermnestra
A pantherabalina
A nelsoni
A cybelecybele
We dedicate this paper to Colin G. “Trig” Treadaway for his many contributions to the study of butterflies in the Philippines, and for the samples, photographs, and helpful advice he has provided to us. We are grateful to Shuen-Chen Chang, Rod Eastwood, Yutaka Inayoshi, Krushnamegh Kunte, Alexander Monastyrskii, Kiyoshi Okubo, Kotaro Saito, Hiroyuki Soeda, Yoshinobu Uémura, and Wei-Ting Wu for providing specimens or images. We also thank the various museums and institutional insect collections acknowledged in the manuscript for allowing us to study, photograph, and occasionally dissect specimens in their care, and are grateful to Perry A.C. Buenavente, David A. Grimaldi, Frank Yu-Feng Hsu, Jia-Yuan Liang, Naomi E. Pierce, Robert K. Robbins, Watana Sakchoowong, Andrei Sourakov, Akekawat Vitheepradit, and others for allowing us access to these collections. This work was funded by grants 102-2621-B-110-003-MY3 from Ministry of Science and Technology (Taiwan), DEB-1120380 and DEB-1541557 from the US National Science Foundation, Grant 9285-13 from the Committee for Exploration and Research of the National Geographic Society, and start-up funds from the City College of New York.
The type species of
Didonis Hübner was erroneously thought to be
Papilio biblis Fabricius, 1807 in earlier literature.
Elymniopsis has often been regarded as a distinct genus since its establishment in 1907 by Fruhstorfer, and most references of Afrotropical butterflies list it as a genus of its own (e.g.,
We recognize 5 subspecies of papua in the present study and synonymize bivitatta with papua because no significant morphological differences were found. In addition, no apparent barriers to dispersal seem to exist between the geographic ranges of these two taxa.
Since Wallace only traveled to the Bird’s Head Peninsula on New Guinea, it is likely that the type specimen was collected in what is now the Indonesian province of West Papua.
The distinctiveness of
esaca and
vasudeva has never been doubted and they have been treated as distinct species in all prior studies. The former is distributed throughout most of the Greater Sunda Islands, the Philippines, and the Thai-Malay peninsula, and the range of the latter encompasses northeast India, Myanmar, northern Laos, northern Vietnam and southwest China. The wings of male
esaca are shorter and more attenuate than
vasudeva, and have black ground coloration with a metallic submarginal band in some specimens. The male of
vasudeva is not dramatically different from the female in wing shape or color pattern. However, our molecular phylogenetic analysis (
“Battak Mountains” is the type locality for many butterfly and other animal taxa, but the name appears on no recent map of Sumatra. It seems to refer to the mountainous region historically inhabited by the Batak ethnic groups in northeast Sumatra. From the description in
The subspecies esaca taeniola is synonymized with esaca borneensis because there are no consistent morphological differences between them and no obvious biogeographical barriers within the island Borneo that would restrict gene flow and maintain subspecific differences.
oberthuri was originally described as a species by
Having examined all the type specimens and a long series of other material ranging from northeast India to southwest China, we failed to detect any consistent differences among the subspecies described by (
Toxopeus’s
mariae was described from Sumatra, from which de Nicéville’s
dohrnii was also collected. Since there are no remarkable or consistent morphological differences between the subspecies and because no biogeographic or climatic barriers to dispersal seem to exist on Sumatra, we consider
mariae to be a junior synonym of
dohrnii. de Nicéville’s
dohrnii was once placed as a subspecies either of
pellucida (
The taxonomic status of De Nicéville’s
ceryxoides has been inconsistent. It was originally proposed as a subspecies of
ceryx, and the treatment was followed by
Elymnias penanga is one of the few Elymnias species with polymorphic female color patterns. However, the female forms are not diagnostically different among subspecies. The diagnostic characters that distinguish subspecies are the size, forewing shape, and metallic sheen of the male.
Elymnias hypermnestra is one of the few satyrine species that is regarded as a minor pest of several species of palms. The name has been used numerous times in the taxonomic, ecological, and agricultural literature (
Roepke’s atrata has rarely been mentioned in previous literature, and, having examined the original description, we regard it as a junior synonym of the nominotypical subspecies, which is also from Java.
Interpretation of the type locality of Drury’s
undularis has been problematic. Drury did not clearly indicate the origin of the specimen he saw but labeled the locality as “East Indies”. During the late 18
th century, the phrase “East Indies” referred to a wide range of possible localities from India to Indonesia. However, authors working after Drury, without clear reason, interpreted
undularis as a taxon distributed in northeast India rather than Indonesia. We examined many specimens from north India to Java at different museums and also compared the original drawings of
hypermnestra (and its junior synonyms) and
undularis in the literature (e.g.,
Hewitson’s
cottonis was described as a full species due to the lack of metallic sheen or any other notable markings on the upper side of both the fore- and hindwings; its conspecificity with
hypermnestra has not previously been suggested. Our unpublished data (
Besides
undularis from northeast India, there are three other subspecies with orange,
Danaus-mimicking females distributed in Myanmar, Thailand, and Vietnam. The differences between females of the subspecies
tinctoria,
violetta, and
meridionalis, are subtle. Besides the female form mimicking
Danaus chrysippus (or
D. genutia), there is another female form with whitish hindwings (the forms
obfuscata and
paraleuca) that possibly mimics either
D. melanippus or
D. affinis in Thailand and Vietnam. Since the current subspecies classification has been adopted by local guidebooks and other publications for so long (
Moore’s hainana was described from specimens from Hainan Island, China, and the name has been applied to the Taiwanese population since the late 19 th century. Since Hainan is between China’s Guangxi Province and northern Vietnam, where septentrionalis and tonkiniana were described, respectively, and because examination of dozens of specimens evince no consistent morphological differences among these subspecies, we synonymize these three names and regard hainana as the valid name.
Penang is a small island in the Andaman Sea lying just off the western coast of peninsular Malaysia. The strait that separates this small island (293 km2) from the peninsula is only 2–8 km wide, yet seems to form a dispersal barrier between the peninsular population (commonly known as agina, but herein changed to beatrice, see discussion below) and insular discrepans. The female type specimen of discrepans seems to be a morphologically intermediate form between orange, Danaus-mimicking tinctoria and dark, Euploea-mimicking phenotypes. This phenotype has not been documented from the mainland. We therefore retain the name discrepans because of the taxon’s distinctive female wing patterns; further studies will ascertain whether this subspecies is genetically distinct from other hypermnestra subspecies.
Elymnias hypermnestra from Malay Peninsula, Singapore, Borneo, and Sumatra have been historically confusing and chaotic. In 1882, Distant reported and described
nigrescens from the Malay Peninsula, Malacca, Billiton (Belitung), and Borneo. Since
Although
caudata was originally proposed as a species of its own, some authors (
Swinhoe’s merula is based on a single male type collected from Sri Lanka. Having examined the type deposited in the Natural History Museum, London, we are convinced that merula should be a synonym of hypermnestra. Lamas (pers. comm.) suggests synonymizing merula with the Sri Lankan fraterna; however, the male of Sri Lankan fraterna is quite different from merula and we cannot at present conclude that synonymizing it with this subspecies is warranted. We presume that the single specimen of merula was accidentally introduced with imported palms or is an aberration; we treat this name as incertae sedis.
The true identity of Godart’s
leucocyma has been problematic since its description in 1819. Godart specified “Java” as the source of the specimen but gave a vague description without any figure.
Our molecular phylogenetic analysis confirms that
detanii and
nepheronides represent opposite sexes of the same species as
Having examined specimens deposited in the NHM, we are confident that harterti and brookei should be regarded as different subspecies of the same species. Moulton’s smithi is identical to Shelford’s brookei so they are synonymized in the present study.
Fabricius stated that the type locality of the nominotypical panthera is Tranquebar (Tharangambadi, Tamil Nadu) in southern India, but the current distribution of this species in India seems to be restricted to the north. It is necessary to confirm the actual distribution of the species in India to verify whether the type locality falls in the actual distribution range or is simply a port from which the specimen was exported during the colonial period.
The subspecies
dusara and
dulcibella were both described from Java, suggesting the names might be synonymous. However, many lepidopteran species have distinctive populations western and eastern parts of the island (
Butler’s
lutescens was proposed as a full species based on specimens from Borneo (collected by Lowe), Malacca, Singapore, and Penang (from Roberts’ collection), and according to
Having examined the type material described by Fruhstorder and Semper, we are convinced that only one subspecies of congruens is distributed on the island Mindanao, and therefore synonymize photinus, phaios, and rafaela with the nominotypical congruens.
Linnaeus did not indicate from which part of Java his specimen of nesaea was collected, and we found no consistent differences among Javan nesaea populations based on examination of specimens at several different museums, so we combine Fruhstorfer’s hermia with the nominotypical nesaea.
Wallace’s timandra from Sylhet, Bangladesh, is not different from Fruhstorfer’s cortona from Myanmar, so we synonymize them.
We examined many nesaea from different regions of Borneo and found no consistent difference among them. Frustorfer’s coelifrons is therefore treated as a junior synonym of hypereides. The relationship between the Malayan lioneli with hypereides can be addressed in future phylogenetic studies.
Lamas (pers. comm.) regards casiphone and kamara as different species, so erinyes, exclusa, and lombokiana are therefore regarded as subspecies of kamara. Our phylogenetic study based on morphology and DNA sequence data, however, show that kamara is conspecific with casiphone; kamara seems to represent a sexually dimorphic, mimetic form in which both males and females differ from the sexually dimorphic mimetic forms of casiphone. We therefore associate all subspecies previously included under kamara with casiphone.
Geyer, when describing
casiphone, did not specify the geographical provenance of his specimen.
de Nicéville’s
erinyes was originally described as a full species, and then downgraded to be a subspecies of
casiphone by
Since we regard kamara as a junior synonym of casiphone, the name lombokiana, originally described as a subspecies of kamara, is treated as a new junior synonym of praetextata in the present study.
Martin’s djilantik and de Nicéville’s exclusa were previously placed under casiphone and kamara, respectively. We regard them as different color forms.
We examined many specimens from northeast India to Vietnam and found no consistent morphological differences to labelport the current subspecies classification. We therefore synonymize ivena and nilamba with malelas.
Distant’s
saueri was originally described as a full species and then placed under
casiphone as a subspecies by
The physiognomy of casiphonides is remarkably similar to female casiphone and female malelas, however, our molecular phylogenetic study places it as the sister group of nesaea. The similarity is almost certainly the result of these different lineages mimicking the same widespread model: female Euploea mulciber.
According to the current taxonomy, two subspecies of
melias are recognized and distributed in Luzon, and it seems unusual for a single island to harbor more than one subspecies. The biotic regions of Luzon, however, are complex. The northern Sierra Madre mountains may serve as a geographical barrier within the island as suggested by
The species agondas has long been considered a highly variable species, the females of which mimic various Taenaris spp. throughout its range in New Guinea and the surrounding islands. Our phylogenetic analysis, however, reveals that thryallis, which was currently placed as a subspecies of cybele, is nested within agondas, and the branch support for this relationship is strong. We therefore conclude that both agondas and cybele are not monophyletic species as currently circumscribed. We sink thryallis into agondas, thus forming a monophyletic species, but further clarification of the subspecific nomenclature is difficult because of the myriad described taxa, vague descriptions of many type localities (frequently “New Guinea”), and our lack of access to material of several rare “subspecies” of agondas, cybele, and other members of the species group.
The true identity of
agondas is mysterious. Boisduval stated that the source of the type specimen was the Solomon Islands (
Our phylogenetic analysis shows that cybele cybele from Bacan is nested within cybele opaca from Halmahera. Morphological examination failed to distinguish these two subspecies; we therefore synonymize them.
The subspecies adumbrata was described from Buru, but it was synonymized with the nominotypical cybele by previous authors. Since the wing pattern of adumbrata is different from that of cybele, and Buru island is presently ~280 km from Halmahera, we revive this subspecies from synonymy with cybele cybele.
According to our phylogenetic analysis,
cumaea is not a monophyletic group and is part of a radiation on Sulawesi that includes
hicetas and
hewitsoni. The subspecies
phrikonis is not allied with other
cumaea subspecies, but comprises the sister group to a clade including
cybele,
vitellia,
holofernes and
agondas. We therefore elevate
phrikonis to the species level.
Nominotypical cumaea is only reported from Halmahera. We have not been able to obtain specimens for inclusion in the molecular phylogeny so we have no indication of how many subspecies previously associated with cumaea should be retained.
Having examined specimens of thyone, we conclude that this taxon should not be regarded as a junior synonym of cumaea cumaea, but we tentatively place thyone as a subspecies of cumaea pending molecular data for inclusion in a phylogenetic study.
In our phylogenetic study, a specimen from North Sulawesi identified as toliana is sister to hicetas, and this pair is sister to bornemanni, which we regard as a full species. Since we have no genetic data from nominotypical cumaea, we are unsure were to place this taxon and tentatively retain toliana as a subspecies of cumaea, even though this arrangement makes cumaea polyphyletic.
Having examined the type specimens of bonthainensis and hicetas, we failed to find any distinguishing morphological characters. These two names are therefore synonymized.
Martin described rarior as a subspecies of hicetas without a clear indication of its type locality in Sulawesi. In our phylogenetic study, two specimens identified as rarior are paraphyletic with regard to hicetas and butona. Since the type locality of rarior is unclear, we retain rarior as a subspecies of hicetas.
The situation of bornemanni is similar to that of phrikonis. It is not closely related to other cumaea subspecies, so we treat it as a full species.
phrikonis has been regarded as a subspecies of cumaea. Our phylogenetic analysis, however, places it as the sister to a clade consisting of cybele, holofernes, umbratilis, vitellia, and agondas. We therefore elevate phrikonis to full species status. Since relicina was described from the same locality as phrikonis we synonymize these two names in the present study.
Elymnias cybele umbratilis was originally described based on five syntypes from Biak and synonymized with thryallis by subsequent authors. Our phylogenetic analysis, however, reveals that umbratilis is a distinct taxon, not closely related to any subspecies of cybele, and sister to holofernes. We therefore revive this taxon and give it full species status.
We have been unable to obtain DNA sequence data from our tissue samples of singhala from old museum specimens. This species is probably not closely related to either the hypermnestra or harterti species group, but its true species group affinity cannot be deduced at this time.