Bright forest blue Pseudodipsas cephenes, copyright Bob Miller and Ian Hill.

Belongs within: Obtectomera.
Contains: Lycaena, Eumaeini, Liphyrinae, Candalidini, Polyommatini, Deudorix, Theclini, Hypochrysops, Philiris.

The Lycaenidae are a cosmopolitan group of butterflies that are commonly associated with ants as larvae. Their classification into subfamilies has historically been subject to disagreement and many authors have included the metalmarks of the Riodinidae as a subgroup of the Lycaenidae.

Characters (from Nielsen & Common 1991): Small to large; eyes often emarginate at base of antenna, or contiguous with antennal socket, often hairy; maxillary palps absent; labial palps ascending; epiphysis absent, fore leg in male somewhat reduced in most groups, normal in female, spurs 0-2-2, 0-1-1 or 0-0-0; fore wing often lacking one or two branches of R, 1A+2A sometimes forked; hind wing with Sc+R1 diverging from Rs near base, usually with two anal veins, margin often produced into one or more tails. Egg with diameter usually greater than height, or nearly spherical, often densely pitted or with projections. Larva usually onisciform, usually with retractile head, sometimes with dense short hairs; abdomen often with medial dorsal gland on segment 7 and pair of dorsolateral, eversible organs on 8; prolegs with median fleshy lobe, more or less interrupting the uni- to multiordinal row of crochets; frequently associated with ants. Pupa usually attached at anal end by central, silken girdle; sometimes lying loose, cremaster absent.

    |--Aphnaeini [Aphnaeinae] KP19
    |    |--Aloeides AV04
    |    |--Aphnaeus AV04
    |    |--Chloroselas AV04
    |    |--Chrysoritis AV04
    |    |--Axiocerses AV04
    |    |    |--A. amanga [incl. A. mendeche] B01a
    |    |    `--A. harpax [incl. A. h. var. croesus, A. h. var. perion, A. h. var. punicea, A. h. var. tjoane] B01a
    |    `--Cigaritis AV04
    |         |--C. massinissa Lucas 1849 E12
    |         |--C. siphax Lucas 1846 E12
    |         `--C. takanonis KP19
    `--+--Polyommatinae KP19
       |    |  i. s.: Hemiargus FS54
       |    |           |--H. gyas FS54
       |    |           |    |--H. g. gyas FS54
       |    |           |    `--H. g. zachaeina Butler 1870 FS54
       |    |           `--H. hanno (Stoll 1790) FS54
       |    |--Candalidini B00b
       |    `--Polyommatini HH02
       `--Theclinae KP19
            |  i. s.: Deudorix B00b
            |         Hypolycaena Felder & Felder 1862 B00b
            |           |--H. danis (Felder & Felder 1865) B00b
            |           |    |--H. d. danis B00b
            |           |    `--H. d. turneri (Waterhouse 1903) B00b
            |           |--H. pachalica B01a
            |           |--H. philippus B01a
            |           `--H. phorbas (Fabricius 1793) B00b
            |                |--H. p. phorbas B00b
            |                `--H. p. ingura Tindale 1923 B00b
            |         Rapala Moore 1881 B00b
            |           `--R. varuna (Horsfield 1829) B00b
            |                |--R. v. varuna B00b
            |                `--R. v. simsoni (Miskin 1874) B00b
            |         Bindahara Moore 1881 B00b
            |           `--B. phocides (Fabricius 1793) B00b
            |                |--B. p. phocides B00b
            |                `--B. p. yurgama Couchman 1965 B00b
            |         Calycopis KP19
            |           |--C. cecrops KP19
            |           `--C. isobeon Im92
            |--Theclini AV04
            `--Luciini B00b
                 |--Hypochrysops B00b
                 |--Philiris B00b
                 |--Parachrysops Bethune-Baker 1904 B00b
                 |--Titea Eliot 1973 B00b
                 |--Lucia Swainson 1833 B00b
                 |    `--L. limbaria (Swainson 1833) B00b
                 |--Paralucia Waterhouse & Turner 1905 B00b
                 |    |--P. aurifer (Blanchard 1848) B00b
                 |    |--P. pyrodiscus (Doubleday 1847) [incl. P. pyrodiscus lucida Crosby 1951] B00b
                 |    `--P. spinifera Edwards & Common 1978 B00a
                 `--Pseudodipsas Felder & Felder 1860 B00b
                      |--P. cephenes Hewitson 1874 B00b
                      `--P. eone (Felder & Felder 1860) B00b
                           |--P. e. eone B00b
                           `--P. e. iole Waterhouse & Lyell 1914 B00b

Lycaenidae incertae sedis:
  Antipodalycaena PPE03
  Lachnocnemini AV04
    |--Lachnocnema bibulus B01a
    `--Thestor AV04
  Chrysophanus T27
    |--C. abbotii B01d
    |--C. boldenarum T27
    |--C. feredayi L27
    |--C. salustius L27
    `--C. thersamon [incl. C. omphale] B01a
  Thecloxurina fassii GE05
  Arcas ducalis GE05
  Lithopsyche antiqua GE05
  Aquisextana irenaei GE05
  Thereus GE05
  Feniseca tarquinius GE05
  Cupido minimus (Fuessly 1775) JP05
  Magnastigma Nicolay 1977 BF08
    `--*M. tegula (Hewitson 1868) [=Thecla tegula] BF08
  Agrodiaetus F05
    |--A. damocles AV04
    |    |--A. d. damocles AV04
    |    `--A. d. rossicus AV04
    `--A. fabressei F05
         |--A. f. fabressei F05
         `--A. f. subbaeticus Gil-T. & Gil-Uceda 2005 F05
  Palaeochrysophanus Im92
    |--P. eurybia (Ochsenheimer 1808) [=Papilio eurybia] B08
    `--P. hippothoe Im92
  Nothodanis schaeffera (Eschscholtz 1821) B00b
  Heoda Johnson, Miller & Herrera 1992 B00b
  Tingra amenaida [incl. T. amenaida var. mombasae] B01a
  Argiolaus lalos B01a
  Stugeta bowkeri [incl. S. bowkeri f. mombasae] B01a
  Spindasis B01a
    |--S. fusca B01c
    |--S. lilacina B01c
    |--S. minima Butler 1901 B01c
    |--S. natalensis B01a
    |--S. nyassae B01a
    |--S. takanonis (Matsumura 1906) Iw92
    `--S. victoriae B01a
  Cacyreus B01a
    |--C. lingeus B01a
    `--C. palemon B01d
  Castalius melaena B01a
  Tarucus telicanus B01a
  Azanus B01b
    |--A. jesous B01a
    `--A. ubaldus B01b
  Chilades B01c
    |--C. alberta Butler 1901 B01c
    `--C. trochilus B01a
  Cupidopsis jobates B01a
  Zizera B01a
    |--Z. gaika B01d
    `--Z. knysna B01a
  Uranothauma B01b
    |--U. cordatus B01b
    `--U. nubifer B01b
  Epitola B01c
    |--E. divisa Butler 1901 B01c
    `--E. honorius [incl. E. teres] B01c
  Iraota B01c
    |--I. nicevillei Butler 1901 B01c
    `--I. timoleon [incl. I. maecenas] B01c
  Fenesica torquinus [incl. Polyommatus porsenna] R13
  Miletinae HH02
  Lipteninae NC91
  Poritiinae NC91
  Curetis GE05 [Curetinae NC91]
    `--C. regula GE05
  Brephidiinae C82
    |--Brephidium C82
    `--Oraidium C82
  Lycaeninae J03
    |  i. s.: Iophanus J03
    |         Lycaena WM66
    `--Eumaeini J03
  Liphyrinae HH02
  Lysandra RD77
    |--L. corydon RD77
    `--L. nivescens W70
  Gerydus chinensis RD77
  Spalgis epius RD77
  Lycaenites P92

*Type species of generic name indicated


[AV04] Als, T. D., R. Vila, N. P. Kandul, D. R. Nash, S.-H. Yen, Y.-F. Hsu, A. A. Mignault, J. J. Boomsma & N. E. Pierce. 2004. The evolution of alternative parasitic life histories in large blue butterflies. Nature 432: 386–390.

[B08] Bálint, Z. 2008. Lepidoptera collections of historical importance in the Hungarian Natural History Museum. Annales Historico-Naturales Musei Nationalis Hungarici 100: 17–35.

[BF08] Bálint, Z., & C. Faynel. 2008. Review of the genus Brangas Hübner, 1819 (Lepidoptera: Lycaenidae) with description of a new genus. Annales Historico-Naturales Musei Nationalis Hungarici 100: 271–306.

[B00a] Braby, M. F. 2000a. Butterflies of Australia: their identification, biology and distribution vol. 1. CSIRO Publishing: Collingwood (Victoria).

[B00b] Braby, M. F. 2000b. Butterflies of Australia: their identification, biology and distribution vol. 2. CSIRO Publishing: Collingwood (Victoria).

[B01a] Butler, A. G. 1901a. An account of a collection of butterflies made by the Rev. K. St. Aubyn Rogers between Mombasa and the forests of Taveta. Annals and Magazine of Natural History, series 7, 7: 22–35.

[B01b] Butler, A. G. 1901b. An account of a collection of butterflies obtained by Lord Delamere, chiefly at Munisu, near Mount Kenya. Annals and Magazine of Natural History, series 7, 7: 197–203.

[B01c] Butler, A. G. 1901c. Descriptions of new species of Lycaenidae in the collection of the British Museum. Annals and Magazine of Natural History, series 7, 7: 288–291.

[B01d] Butler, A. G. 1901d. On a collection of butterflies from the Uganda Protectorate, forwarded by C. Steuart Betton, Esq., in 1900. Annals and Magazine of Natural History, series 7, 7: 562.

[C82] Craw, R. C. 1982. Phylogenetics, areas, geology and the biogeography of Croizat: a radical view. Systematic Zoology 31 (3): 304–316.

[E12] Evenhuis, N. L. 2012. Publication and dating of the Exploration Scientifique de l’Algérie: Histoire Naturelle des Animaux Articulés (1846–1849) by Pierre Hippolyte Lucas. Zootaxa 3448: 1–61.

[F05] Fernández, J. 2005. Noticia de nuevos táxones para la ciencia en el ámbito Íbero-Balear y Macaronésico. Nuevos táxones animales descritos en la península Ibérica y Macaronesia desde 1994 (IX). Graellsia 61 (2): 261–282.

[FS54] Franz, E., & H. Schröder. 1954. Tagfalter (Lep. Rhopalocera) aus El Salvador. Senckenbergiana Biologica 35: 75–87.

[GE05] Grimaldi, D., & M. S. Engel. 2005. Evolution of the Insects. Cambridge University Press: New York.

[HH02] Hall, J. P. W., & D. J. Harvey. 2002. Basal subtribes of the Nymphidiini (Lepidoptera: Riodinidae): phylogeny and myrmecophily. Cladistics 18: 539–569.

[Im92] Imes, R. 1992. The Practical Entomologist. Aurum Press: London.

[Iw92] Iwahashi, J. (ed.) 1992. Reddo Deeta Animaruzu: a pictorial of Japanese fauna facing extinction. JICC: Tokyo.

[J03] Jong, R. de. 2003. Are there butterflies with Gondwanan ancestry in the Australian region? Invertebrate Systematics 17: 143–156.

[JP05] Jungmann, E., & U. Poller. 2005. Neunachweise und Wiederfunde für die Schmetterlingsfauna (Insecta; Lepidoptera) des Altenburger Landes. Mauritiana 19 (2): 317–326.

[KP19] Kawahara, A. Y., D. Plotkin, M. Espeland, K. Meusemann, E. F. A. Toussaint, A. Donath, F. Gimnich, P. B. Frandsen, A. Zwick, M. dos Reis, J. R. Barber, R. S. Peters, S. Liu, X. Zhou, C. Mayer, L. Podsiadlowski, C. Storer, J. E. Yack, B. Misof & J. W. Breinholt. 2019. Phylogenomics reveals the evolutionary timing and pattern of butterflies and moths. Proceedings of the National Academy of Sciences of the USA 116 (45): 22657–22663.

[L27] Lindsay, S. 1927. A list of the Lepidoptera of Deans Bush, Riccarton, Canterbury. Transactions and Proceedings of the New Zealand Institute 57: 693–696.

[NC91] Nielsen, E. S., & I. F. B. Common. 1991. Lepidoptera (moths and butterflies). In: CSIRO. The Insects of Australia: A textbook for students and research workers 2nd ed. vol. 2 pp. 817–915. Melbourne University Press: Carlton (Victoria).

[PPE03] Patrick, B., H. Patrick & E. Edwards. 2003. Tiger moths and pheromones. Weta 25: 14–17.

[P92] Poinar, G. O., Jr. 1992. Life in Amber. Stanford University Press: Stanford.

[R13] Reuter, O. M. 1913. Lebensgewohnheiten und Instinkte der Insekten bis zum Erwachen der sozialen Instinkte. R. Friedländer & Sohn: Berlin.

[RD77] Richards, O. W., & R. G. Davies. 1977. Imms' General Textbook of Entomology 10th ed. vol. 2. Classification and Biology. Chapman and Hall: London.

[T27] Thomson, G. M. 1927. The pollination of New Zealand flowers by birds and insects. Transactions and Proceedings of the New Zealand Institute 57: 106–125.

[WM66] Wallace, A. R., & F. Moore. 1866. List of lepidopterous insects collected at Takow, Formosa, by Mr. R. Swinhoe. Proceedings of the Zoological Society of London 1866: 355–365.

[W70] White, M. J. D. 1970. Cytogenetics. In: CSIRO. The Insects of Australia: A textbook for students and research workers pp. 72–82. Melbourne University Press.

Last updated: 27 February 2021.

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