Lycaenidae

Lycaenidae sp.

Superregnum: Eukaryota
Cladus: Unikonta
Cladus: Opisthokonta
Cladus: Holozoa
Regnum: Animalia
Subregnum: Eumetazoa
Cladus: Bilateria
Cladus: Nephrozoa
Cladus: Protostomia
Cladus: Ecdysozoa
Cladus: Panarthropoda
Phylum: Arthropoda
Subphylum: Hexapoda
Classis: Insecta
Cladus: Dicondylia
Subclassis: Pterygota
Cladus: Metapterygota
Infraclassis: Neoptera
Cladus: Eumetabola
Cladus: Endopterygota
Superordo: Panorpida
Cladus: Amphiesmenoptera
Ordo: Lepidoptera
Subordo: Glossata
Cladus: Coelolepida
Cladus: Myoglossata
Cladus: Neolepidoptera
Infraordo: Heteroneura
Cladus: Eulepidoptera
Cladus: Ditrysia
Cladus: Apoditrysia
Cladus: Obtectomera
Superfamilia: Papilionoidea

Familia: Lycaenidae
Subfamiliae (7): Aphnaeinae – Curetinae – Lycaeninae – Miletinae – Polyommatinae – Poritiinae – Theclinae

Overview of genera (423)

Actizera – Acupicta – Acytolepis – Aethiopana – Agriades – Ahlbergia – Ahmetia – Alaena – Allosmaitia – Allotinus – Aloeides – Amblopala – Amblypodia – Ancema – Anthene – Antigius – Aphnaeus – Apporasa – Aquisextana – Araotes – Araragi – Arawacus – Arcas – Archaeogerydus – Argyraspodes – Argyrocheila – Argyrocupha – Arhopala – Aricia – Arletta – Arrhenothrix – Artipe – Artopoetes – Aslauga – Athamanthia – Athysanota – Atlides – Aurea – Austrozephyrus – Axiocerses – Azanus – Baliochila – Baspa – Batelusia – Bidaspa – Biduanda – Bindahara – Boldenaria – Bothrinia – Bowkeria – Brephidium – Britomartis – Bullis – Cacyreus – Caerulea – Caleta – Callicista – Callictita – Callophrys – Calycopis – Calystryma – Candalides – Castalius – Catapaecilma – Catochrysops – Catopyrops – Celastrina – Cesa – Chaetoprocta – Chalybs – Charana – Cheritra – Cheritrella – Chilades – Chliaria – Chloroselas – Chlorostrymon – Chrysoritis – Chrysozephyrus – Cigaritis – Citrinophila – Cnodontes – Contrafacia – Cooksonia – Cordelia – Coreana – Creon – Crudaria – Cupidesthes – Cupido – Cupidopsis – Curetis – Cyaniriodes – Cyaniris – Cyanophrys – Cyclargus – Cyprotides – Dacalana – Danis – Dapidodigma – Darasana – Deloneura – Deramas – Deudorix – Diopetes – Discolampa – Drina – Drupadia – Durbania – Durbaniella – Durbaniopsis – Echinargus – Eicochrysops – Electrostrymon – Elkalyce – Eooxylides – Epidemia – Epimastidia – Epitola – Epitolina – Eresina – Eresinopsides – Erikssonia – Erina – Erora – Erysichton – Esakiozephyrus – Etesiolaus – Euaspa – Euchrysops – Euliphyra – Euliphyrodes – Eumaeus – Eumedonia – Euphilotes – Eupsychellus – Euthecta – Evenus – Falcuna – Famegana – Favonius – Feniseca – Flos – Freyeria – Glaucopsyche – Gonerilia – Habrodais – Harpendyreus – Heliophorus – Helleia – Hemiargus – Hemiolaus – Heodes – Hermelycaena – Hewitsonia – Horaga – Howarthia – Hyllolycaena – Hypaurotis – Hypochlorosis – Hypochrysops – Hypojamides – Hypokopelates – Hypolycaena – Hypomyrina – Hypophytala – Hypostrymon – Hypothecla – Hyrcanana – Hysudra – Iaspis – Icaricia – Incisalia – Iolana – Iolaus – Ionolyce – Iophanus – Ipidecla – Iraota – Iratsume – Iridana – Itylos – Jacoona – Jaera – Jalmenus – Jamides – Japonica – Keraunogramma – Kopelates – Kretania – Lachnocnema – Laeosopis – Lampides – Lamprospilus – Larinopoda – Lepidochrysops – Leptomyrina – Leptotes – Lestranicus – Leucantigius – Liphyra – Liptena – Liptenara – Logania – Loweia – Loxura – Lucia – Luthrodes – Lycaena – Lycaenesthes – Lycaenopsis – Lysandra – Macusia – Madeleinea – Magnastigma – Mahathala – Maneca – Manto – Mantoides – Megalopalpus – Megisba – Melanolycaena – Micandra – Michaelus – Micropentila – Microscena – Miletus – Mimacraea – Mimeresia – Ministrymon – Mithras – Mitoura – Monodontides – Mota – Myrina – Nabokovia – Nacaduba – Nadisepa – Narathura – Neaveia – Neocheritra – Neoepitola – Neolucia – Neolycaena – Neomyrina – Neopithecops – Neozephyrus – Nesiostrymon – Nesolycaena – Neurellipes – Neurypexina – Nilasera – Niphanda – Noreena – Notarthrinus – Oboronia – Ocaria – Oenomaus – Ogyris – Olynthus – Oraidium – Oreolyce – Ornipholidotos – Orthomiella – Oxylides – Paiwarria – Palaeochrysophanus – Palaeophilotes – Pamela – Panchala – Panthiades – Parachrysops – Paraduba – Paragerydus – Paralucia – Paralycaeides – Parasiomera – Paraslauga – Parelodina – Parrhasius – Penaincisalia – Pentila – Pepliphorus – Petrelaea – Phaedrotes – Phaeostrymon – Phasis – Phengaris – Philiolaus – Philiris – Philotes – Philotiella – Phlyaria – Phytala – Pilodeudorix – Pistoria – Pithecops – Plebejus – Plebulina – Podanotum – Poecilmitis – Polyommatus – Poriskina – Poritia – Powellana – Praephilotes – Pratapa – Prosotas – Protantigius – Pseudaletis – Pseudalmenus – Pseuderesia – Pseudiolaus – Pseudochliaria – Pseudochrysops – Pseudodipsas – Pseudogyris – Pseudolucia – Pseudolycaena – Pseudonacaduba – Pseudoneaveia – Pseudophilotes – Pseudozizeeria – Psychonotis – Ptelina – Ptox – Purlisa – Quercusia – Rachana – Rapala – Rathinda – Ravenna – Rekoa – Remelana – Rhinelephas – Ritra – Rysops – Sancterila – Sandia – Satadra – Satyrium – Scolitantides – Semanga – Sheffieldia – Shijimia – Shijimiaeoides – Shirozua – Siderus – Sidima – Simiskina – Sinia – Sinthusa – Sithon – Sospita – Spalgis – Stalachtis – Stempfferia – Strymon – Stugeta – Suasa – Sukidion – Surendra – Symbiopsis – Syrmoptera – Tajuria – Talicada – Taraka – Tarucus – Telipna – Teratoneura – Teratozephyrus – Teriomima – Tetrarhanis – Thaduka – Thamala – Tharsalea – Thaumaina – Thecla – Theclinesthes – Theclopsis – Theorema – Thereus – Theritas – Thermoniphas – Thersamonia – Thestius – Thestor – Thrix – Ticherra – Titea – Tmolus – Tomares – Tongeia – Toxochitona – Trichonis – Triclema – Trimenia – Tumerepedes – Turanana – Tylopaedia – Udara – Una – Upolampes – Uranobothria – Uranothauma – Ussuriana – Vacciniina – Vadebra – Vaga – Virachola – Wagimo – Xamia – Yasoda – Zarona – Zeltus – Zeritis – Zesius – Zetona – Zinaspa – Zintha – Zizeeria – Zizina – Zizula

[source: Catalogue of Life: 2012 Annual Checklist (must be considered tentative)]
Name

Lycaenidae Leach, 1815.
References
Primary references

Eliot, J.N. 1973. The higher classification of the Lycaenidae (Lepidoptera): a tentative arrangement. Bulletin of the British Museum (Natural History), entomology 28(6): 371-505. Reference page.
Leach, W.E. 1815. Entomology [pp. 57–172]. In: Brewster, D. (Ed). Brewster’s Edinburgh Encyclopedia. Volume IX [part I]. W. Blackwood, J. Waugh, etc., Edinburgh, 764 pp. BHL Reference page.

Selected references

Bálint, Z. & Johnson, K. 1997. Reformation of the Polyommatus section with a taxonomic and biogeographic overview (Lepidoptera, Lycaenidae, Polyommatini). Neue Entomologische Nachrichten 40: 1–68. PDF. Reference page.
Bridges, C.A. 1994. Catalogue of the Family-Group, Genus-Group and Species-Group Names of the Riodinidae & Lycaenidae (Lepidoptera) of the World. Bridges, Urbana, Illinois. BHL Reference page.
Espeland, M., Breinholt, J.W., Willmott, K.R., Warren, A.D., Vila, R., Toussaint, E.F.A., Maunsell, S.C., Aduse-Poku, K., Talavera, G., Eastwood, R., Jarzyna, M.A., Guralnick, R., Lohman, D.J., Pierce, N.E. & Kawahara, A.Y. 2018. A Comprehensive and Dated Phylogenomic Analysis of Butterflies. Current Biology 28(5): 770–778. DOI: 10.1016/j.cub.2018.01.061 Reference page.
Hemming, A. F., 1967. The generic names of the butterflies and their type species (Lepidoptera: Rhopalocera). Bulletin of the British Museum (Natural History) 1967, Suppl. 9: 1-509.Reference page.
Sáfián, S. & Collins, S.C. 2015. Establishment of a new genus for Eresiomera paradoxa (Schultze, 1917) and related taxa (Lepidoptera: Lycaenidae) with description of two new species. Zootaxa 4018(1): 124–136. DOI: 10.11646/zootaxa.4018.1.7. Preview (PDF) Reference page.
Stempffer, H. 1967. The genera of the African Lycaenidae (Lepidoptera: Rhopalocera). Bulletin of the British Museum (Natural History), entomology. Supplement 10: 322pp, 1pl. BHL Reference page.

Additional references

Clench, H. K., 1971. Two new hairstreaks from Mexico (Lepidoptera: Lycaenidae). Bull. Allyn Museum 3: 1–6. Full article: [1] Reference page.
Heikkilä, M., Kaila, L., Mutanen, M., Peña, C. & Wahlberg, N. 2011. Cretaceous origin and repeated tertiary diversification of the redefined butterflies. Proceedings of the Royal Society B 279(1731): 1093–1099. DOI: 10.1098/rspb.2011.1430 Open access. Reference page.
Hirowatari, T. 1992. A generic classification of the tribe Polyommatini of the Oriental and Australian regions (Lepidoptera, Lycaenidae, Polyommatinae). Bulletin of the University of Osaka Prefecture (B) 44 (Suppl.) PDF. Reference page.
Kerr, J.F.R., 1967: New records of Lycaenidae (Lepidoptera) in Australia and a description of a new subspecies. Australian Journal of Entomology 6 (1): 49–51. DOI: 10.1111/j.1440-6055.1967.tb02138.x. Full article: [2].
Korb, S.K. 2018. On the blue butterfly taxa established by L.G. Courvoisier with the type locality “Narün” (Lepidoptera, Lycaenidae). Zootaxa 4429(3): 589–594. DOI: 10.11646/zootaxa.4429.3.11 Paywall Reference page.
Macqueen, J., 1965: Notes on Australian Lycaenidae (Lepidoptera). Australian Journal of Entomology 4 (1): 56–57. DOI: 10.1111/j.1440-6055.1965.tb00648.x. Full article: [3].
Matsumura, S., 1926. Some new and unrecorded Lycaenids-Species from Japan, Girca and Formosa. Ins. matsumur. 1(1): 23-31, pl 2. web. Reference page.
Munguira, M.L., Martin, J., García-Barros, E., Shahbazian, G. & Cancela, J.P. 2015: Morphology and morphometry of Lycaenid eggs (Lepidoptera: Lycaenidae). Zootaxa 3937(2): 201–247. DOI: 10.11646/zootaxa.3937.2.1. Reference page.
Murayama, S. & Okamura, H., 1973. Butterflies of Luzon island in the Philippines, with descriptions of new species and sub-species. Tyô to Ga 24(1): 10-25., 48 figs. Reference page.
Schröder, H.G. & Treadaway, C. G. 1989. Zur Kenntnis philippinischer Lycaenidae, 5 (Lepidoptera). Entomologische Zeitschrift 99(19): 273–288, 7 figs. Reference page.
Schröder, H.G. & Treadaway, C. G. 1990. Zur Kenntnis philippinischer Lycaenidae, 6. (Lepidoptera). Entomologische Zeitschrift 100(9): 168–171. ill. Reference page.
Schröder, H.G. & Treadaway, C. G. 1990. Zur Kenntnis philippinischer Lycaenidae, 7. (Lepidoptera). Entomologische Zeitschrift 100(21): 404–407, ill. Reference page.
Schroeder, H.G. & C.G. Treadaway, 1999: Zur Kenntnis philippinischer Lycaenidae, 12. (Lepidoptera). Entomologische Zeitschrift 109 (5): 206–212.
Schroeder, H.G. & Treadaway, C.G., 2000: Zur Kenntnis philippinischer Lycaenidae, 13 (Lepidoptera). — NF 20 (3/4): 271–280.
Schröder, H.G. & Treadaway, C. G. 2000. Zur kenntnis philippinischer Lycaenidae, 14 (Lepidoptera). Nachrichten des Entomologischen Vereins Apollo N.F. 21(2): 61–64. Reference page.
Schroeder, H.G. & Treadaway, C.G., 2002: Zur Kenntnis philippinischer Lycaenidae, 15 (Lepidoptera). — NF 22 (4): 239–242.
Schroeder, H.G. & Treadaway, C.G., 2002: Zur Kenntnis philippinischer Lycaenidae, 16 (Lepidoptera). — NF 23 (3): 135–139.
Schroeder, H.G. ; C.G. Treadaway & J. Nuyda, 1999: Zur kenntnis philippinischer Lycaenidae, 11. (Lepidoptera). Entomologische Zeitschrift 109 (4): 145–154.
Torres-Núñez, R. ; J.P.W. Hall ; K.R. Willmott & K. Johnson, 1996: A new genus of Elfin butterflies from the northern high Andes (Lepidoptera: Lycaenidae). Tropical Lepidoptera 7 (1): 81–86. Full article: [4].

Links

Markku Savela: Funet: Lycaenidae.

Lycaenidae – Taxon details on Atlas of Living Australia (ALA).
Australian Faunal Directory (AFD)
Brower, Andrew V. Z. 2008. Lycaenidae [Leach] 1815. Gossamer-winged butterflies. Version 25 April 2008 (under construction). http://tolweb.org/Lycaenidae/12175/2008.04.25 in The Tree of Life Web Project, http://tolweb.org/
Butterflies of America

BHL bibliography
Hassler, M. {{{1}}}. Lycaenidae. World Plants: Synonymic Checklists of the Vascular Plants of the World In: Roskovh, Y., Abucay, L., Orrell, T., Nicolson, D., Bailly, N., Kirk, P., Bourgoin, T., DeWalt, R.E., Decock, W., De Wever, A., Nieukerken, E. van, Zarucchi, J. & Penev, L., eds. {{{1}}}. Species 2000 & ITIS Catalogue of Life. Published online. Accessed: {{{1}}} {{{2}}} {{{3}}}. Reference page.
Lycaenidae – Taxon details on Encyclopedia of Life (EOL).
Global Biodiversity Information Facility. 2019. GBIF Backbone Taxonomy. Checklist dataset. Taxon: Lycaenidae.
ION
Pitkin, B. and Jenkins, P., 2005. Butterflies & Moths of the World: Generic Names & their Type-species[5]

Vernacular names
беларуская: Блакітніцы
čeština: Modráskovití
dansk: Blåfugle
Deutsch: Bläulinge
English: Gossamer-winged butterflies
suomi: Sinisiivet
français: lycénidés
magyar: Boglárkalepkefélék, boglárkalepkék, hangyaboglárka lepkék, lángszinérfélék
日本語: シジミチョウ科
한국어: 부전나빗과
lietuvių: Melsviai
polski: Modraszkowate
русский: Голубянки
svenska: Juvelvingar
Tiếng Việt: Họ Bướm xanh
中文: 灰蝶科

Lycaenidae is the second-largest family of butterflies (behind Nymphalidae, brush-footed butterflies), with over 6,000 species worldwide,[1] whose members are also called gossamer-winged butterflies. They constitute about 30% of the known butterfly species.

The family comprises seven subfamilies, including the blues (Polyommatinae), the coppers (Lycaeninae), the hairstreaks (Theclinae), and the harvesters (Miletinae).

Description, food, and life cycle

Adults are small, under 5 cm usually, and brightly coloured, sometimes with a metallic gloss.

Larvae are often flattened rather than cylindrical, with glands that may produce secretions that attract and subdue ants. Their cuticles tend to be thickened. Some larvae are capable of producing vibrations and low sounds that are transmitted through the substrates they inhabit. They use these sounds to communicate with ants.[2][3]

Adult individuals often have hairy antenna-like tails complete with black and white annulated (ringed) appearance. Many species also have a spot at the base of the tail and some turn around upon landing to confuse potential predators from recognizing the true head orientation. This causes predators to approach from the true head end resulting in early visual detection.[4]
Ant tending a lycaenid larva

Lycaenids are diverse in their food habits and apart from phytophagy, some of them are entomophagous, feeding on aphids, scale insects, and ant larvae. Some lycaenids even exploit their association with ants by inducing ants to feed them by regurgitation, a process called trophallaxis. Not all lycaenid butterflies need ants, but about 75% of species associate with ants,[2] a relationship called myrmecophily. These associations can be mutualistic, parasitic, or predatory depending on the species.

In some species, larvae are attended and protected by ants while feeding on the host plant, and the ants receive sugar-rich honeydew from them, throughout the larval life, and in some species during the pupal stage. In other species, only the first few instars are spent on the plant, and the remainder of the larval lifespan is spent as a predator within the ant nest. It becomes a parasite, feeding on ant regurgitations, or a predator on the ant larvae.[2] The caterpillars pupate inside the ants' nest and the ants continue to look after the pupae. Just before the adults emerge, the wings of the butterfly inside the pupal case detach from it, and the pupa becomes silvery. The adult butterfly emerges from the pupa after three to four weeks, still inside the ant nest. The butterfly must crawl out of the ant nest before it can expand its wings.

Several evolutionary adaptations enable these associations, including small glands on the skin of the caterpillars called "pore cupola organs". Caterpillars of many species have a gland on the seventh abdominal segment that produces honeydew and is called the "dorsal nectary gland" (also called "Newcomer's gland"). An eversible organ called the "tentacular organ" is present on the eighth abdominal segment and this is cylindrical and topped with a ring of spikes and emits chemical signals which are believed to help in communicating with ants.[5]
Subfamilies
Mating Cyaniris semiargus
Caterpillar of the Atala (Eumaeus atala)
Leaf blue (Amblypodia anita) from Theclinae

Many taxonomists only include the Lycaeninae, Theclinae, Polyommatinae, Poritiinae, Miletinae, and Curetinae under the Lycaenidae.[6][7] The Aphnaeinae, which used to be a tribe (Aphnaeini) within the Theclinae, were recently given subfamily rank too.[8]

Curetinae – sunbeams (Oriental or Palaearctic). Selected species:
Curetis thetis – Indian sunbeam
Miletinae – harvesters (mostly African, or Oriental, one Nearctic), probably all feed on aphids or their secretions. Selected species:
Liphyra brassolis – moth butterfly (largest lycaenid)
Poritiinae (Oriental and Afrotropical)
Aphnaeinae (Afrotropical and Oriental)
Theclinae – hairstreaks (usually tailed) and elfins (not tailed) (global). Selected species:
Arhopala – oakblues
Atlides halesus – great purple hairstreak
Eumaeus atala – Atala
Satyrium pruni – black hairstreak
Lycaeninae – coppers (Holarctic). Selected species:
Iophanus pyrrhias – Guatemalan copper
Lycaena boldenarum – boulder copper
Lycaena epixanthe – bog copper
Lycaena rauparaha – Rauparaha's copper
Lycaena dispar – large copper
Lycaena phlaeas – small copper
Lycaena heteronea – blue copper
Polyommatinae – blues (global). Selected species:
Celastrina ladon – spring azure
Chilades – jewel blues
Cupido comyntas – eastern tailed-blue
Cupido minimus – small blue
Cyaniris semiargus – mazarine blue
Euphilotes battoides allyni – El Segundo blue
Euphilotes pallescens arenamontana – Sand Mountain blue
Glaucopsyche lygdamus – silvery blue
Glaucopsyche lygdamus palosverdesensis – Palos Verdes blue
Glaucopsyche xerces (extinct) – Xerces blue
Icaricia icarioides fenderi – Fender's blue
Phengaris arion – large blue
Polyommatus icarus – common blue
Pseudozizeeria maha – pale grass blue
Plebejus argus – silver-studded blue
Talicada nyseus – red Pierrot

Some older classifications used to include other subfamilies such as Liphyrinae (now Liphyrini, a tribe within Miletinae), Lipteninae (now Liptenini, a tribe within Poritiinae), or Riodininae (now a separate family: Riodinidae).

The fossil genus Lithodryas is usually (but not unequivocally) placed here; Lithopsyche is sometimes placed here, but sometimes in the Riodininae.
See also

List of lycaenid genera

References

Pierce, Naomi E.; Braby, Michael F.; Heath, Alan; Lohman, David J.; Mathew, John; Rand, Douglas B. & Travassos, Mark A. (January 2002)."The Ecology and Evolution of Ant Association in the Lycaenidae (Lepidoptera)". Annual Review of Entomology. 47 (1): 259-267. doi:[https://doi.org/10.1146%2Fannurev.ento.47.091201.145257 10.1146/annurev.ento.47.091201.145257.
Pierce, N. E.; Braby, M. F.; Heath, A.; Lohman, D. J.; Mathew, J.; Rand, D. B. & Travassos, M. A. (2002). "The ecology and evolution of ant association in the Lycaenidae (Lepidoptera)". Annual Review of Entomology. 47: 733-771.
DeVries, Philip J. (1992). "Singing Caterpillars, Ants and Symbiosis". Scientific American, 267: 76
Robbins, Robert K. (1981). "The 'False Head' Hypothesis: Predation and Wing Pattern Variation of Lycaenid Butterflies". American Naturalist. 118 (5): 770-775.
"Lycaenid Butterflies and Ants". Australian Museum. Archived 18 November 2007.
Brower, Andrew V. Z. (2008). "Lycaenidae [Leach] 1815". Version 25 April 2008 (under construction). The Tree of Life Web Project.
Ackery, P. R.; de Jong, R. & Vane-Wright, R. I. (1999). "The butterflies: Hedyloidea, Hesperioidea, and Papilionoidea". Pages 264-300 in: Lepidoptera: Moths and Butterflies. 1. Evolution, Systematics, and Biogeography. Handbook of Zoology Vol. IV, Part 35. N. P. Kristensen, ed. De Gruyter, Berlin and New York.

Boyle, J. H.; Kaliszewska, Z. A.; Espeland, M.; Suderman, T. R.; Fleming, J.; Heath, A. & Pierce, N. E. (2015). "Phylogeny of the Aphnaeinae: Myrmecophilous African butterflies with carnivorous and herbivorous life histories". Systematic Entomology. 40 (1): 169–182.