The Lycaenidae as construed here includes the blues, coppers and hairstreaks, but not the metalmarks, which are viewed as a sister-family Riodinidae, based on the combined phylogenetic analysis of morphological and molecular data by Wahlberg et al. (2005).
The largest and most biologically diverse of all butterfly families, containing more than 6000 species (however this figure includes the riodinids, which have also been considered a family in their own right). The apomorphies included here are those used to justify a Lycaenidae that includes the riodinids and the curetines (also considered a family by some).
The Lycaenidae as construed here includes the blues, coppers and hairstreaks, but not the metalmarks, which are viewed as a sister-family Riodinidae, based on the combined phylogenetic analysis of morphological and molecular data by Wahlberg et al. (2005). The vast majority of lycaenids are in the clade that includes the blues (Polyommatinae), coppers (Lycaeninae) and hairstreaks (Theclinae). Eliot (who viewed these taxa as tribes of a more inclusive Lycaeninae in Corbet et al. 1992) implied that Aphnaeinae is the sister group to the other three subfamilies based on characters of the larval head capsule.
These are small butterflies with iridescent colors. The three most important subfamilies are the Hairstreaks (Theclinae), Coppers (Lycaeninae), and Blues (Polyommotinae). Butterflies in this family often visit composite flowers (Asteraceae) and small wildflowers from other families. Theclinae (Hairstreaks): These butterflies are silvery grey with rows of red or blue dots on the wing undersides, while the uppersides (exposed when the wings are outstretched) are a drab gray, brown, or black. Sometimes there is a small tail on each hindwing. The caterpillars feed on various trees and shrubs, including willows, wild cherries, hawthorns, oaks, hickories, and sumac. The species Strymon melinus (Gray Hairstreak), also feeds on some herbaceous plants, such as Mallows and Smartweeds. Lycaeninae (Coppers): These butterflies are orange and silvery gray with scattered black dots on the wing undersides. The uppersides have vivid orange and black patterns, although they are sometimes greyish or purplish in overall appearance. The larvae feed primarily on Rumex spp. (Dock) and Polygonum spp. (Smartweed and Knotweed). Polyommotinae (Blues): The Blues are silvery grey, or blue on the wing undersides, with rows of black dots and a patch of orange on the hindwings. The wing uppersides are silvery blue with white or black edges. Some species have small tails on the hind wings, and females are sometimes gray on the uppersides. The caterpillars of Blues usually feed on legumes, although the caterpillars of Celastrina argiolus (Spring/Summer Azure) feed on various small trees and shrubs, including Dogwood, Wild Cherry, Sumac, New Jersey Tea, and Viburnum. The caterpillars of Blues often secrete a honey dew that attracts ants.
There are nearly 5,000 species in this family around the world, but most only live in the tropics. We only have about 145 in the United States, and 32 species in Michigan.
Biogeographic Regions: nearctic (Native ); palearctic (Native ); oriental (Native ); ethiopian (Native ); neotropical (Native ); australian (Native ); antarctica (Native ); oceanic islands (Native )
Nearctic, Palearctic, Oriental, Ethiopian, Neotropical, Australian, Oceanic Island
Geographic Range description:
Cosmopolitan distribution, including species endemic to New Zealand, Hawaii and many other Pacific islands). Greatest diversity in the tropics. From Scoble, 1992: The Lipeninae are Afrotropical. Poritiinae are Oriental. Liphyrinae are mainly African with a few species in the Oriental and Australasian regions. Miletinae occur mostly in Africa or the Orient, some are present in the Holarctic region. Curetinae are Oriental with a few Palaearctic representatives. Theclinae are represented in all the main zoogeographical areas. Lycaeninae occur mainly in the Holarctic region. Polyommatinae are represented in all major biogeographical areas.
This group of butterflies get their common names from their wings. Some groups have shiny blue on their wings, others are the color of copper. Some have very thin little "tails" on their wings that are called hairstreaks. Many are not so colorful, and have gray or brown wings with spots of black, white, or orange. They are usually small butterflies, with wingspans of 25 mm or less. Some species have males with reduced front legs like the Brushfoot family, but females always have all 6 legs for walking and standing.
Caterpillars in this family have small heads and legs, and sometimes look like slugs covered with tiny hairs. Most are green or brown, but a few are yellowish or reddish. Some can pull in their heads completely, like a slug.
Other Physical Features: ectothermic ; bilateral symmetry
Sexual Dimorphism: female larger; male more colorful
Description of egg morphology:
From Scoble, 1992: "The diameter of the egg usually exceeds that of its height, or the egg may be almost spherical. Often, lycaenid eggs are pitted or bear projections. In eggs of most species parts of the highly porous chorion is perforated by numerous pores enable it to act as a plastron when eggs are submerged in rain water (Downey and Allyn, 1981)"
From Scoble 1992: "The larva (Downey in Stehr, 1987) is usually onisciform, resembling the shape of a woodlouse." "The abdomen often bears a dorsal gland on segment A7, a pair of eversible organs on segment A8, and minute, round "pore cupolas' associated particularly with the dorsal gland and the eversible organs."
Crochet arrangement description:
From Scoble 1992: Crochets are usually "tri- to multiordinal, and arranged in an interrupted mesoseries."
From Scoble, 1992: "The pupal is often girdled. Where not girdled, it may be positioned at an angle to the substrate or be suspended with the head downwards."
Adult Thorax Morphology
Foreleg in males usually reduced with the tarsal segments typically fused and the pretarsus reduced but in females normally developed. The forelegs are used for walking in most species. Tibial spurs may be entirely absent 0-0-2 or 0-1-1
The humeral vein is present or absent in the hindwing (Scoble 1992; Eliot 1973; Munroe, 1982). Vein CuP is usually absent from both wings.
Wing coupling description:
From Scoble 1992: "There is no frenular-retinacular system in true butterflies, wing coupling is said to be effected by an amplexiform system made possible by the expanded humeral area. The forewing-metathoracic locking-device is absent.
From Scoble 1992: "At least some stalking of the radial veins of the forewing occurs."
Abdomen tympanum description:
Although there are no tympanal structures, in the pupae rows of tubercles on the posterior edge of A5 are rubbed against tubercles on the anterior edge of A6, to produce sound.
Thorax tympanum description:
Palp tympanum description:
Adult Head Morphology
Number of labial palp segments:
Labial palpus modification:
Usually well developed and ascending but occasionally reduced (Scoble, 1992)
Number of chaetosomata:
Adult head description:
From Watson and Dallwitz 2007: "Eyes conspicuously white-rimmed, notched or emarginate at the bases of the antennae and contiguous with the bases of the antennal sockets; hairy, or glabrous."
Antennal base adjacent to margin of compound eye, and usually indenting it. Reduction of the metathoracic wing case of the pupa. Spatulate lobe on larval abdominal prolegs which interrupts the mesoseries of crochets. Absence of larval eversible prothoracic gland
Adults in this family are usually found close to the food plants of the caterpillars. This usually means around forest edges, open fields, along streams, and other open but vegetated areas.
Caterpillars in this family are found on their food plants, or in the company of ants.
Habitat Regions: temperate ; tropical ; terrestrial
Terrestrial Biomes: tundra ; taiga ; desert or dune ; chaparral ; forest ; rainforest ; scrub forest ; mountains
Wetlands: marsh ; swamp ; bog
Caterpillars in this family eat a wider variety of foods than other butterfly families. Some species eat leaves, but others specialize in flowers or fruit. One species is a predator! This is very rare in butterflies and moths. It feeds on Aphididae, and females lay their eggs near their insect prey.
Some adults sip nectar, but many prefer tree sap or puddles.
Adults help pollinate flowers. Caterpillars usually aren't a major factor for plant populations, but sometimes are when they occur in large populations. Many species in this group have caterpillars that have mutualistic relationships with ants. The caterpillars have glands that produce liquids that the ants like to drink, and in exchange the ants protect the caterpillars from predators, and in a few species even take the caterpillars into their nests.
Ecosystem Impact: pollinates
These species are not usually toxic to predators. The caterpillars sometimes make silk nests to hide in, or rely on ants for protection. Some adults rub their hindwings together whenever they land. This may draw predators attention to the eyespots or hairstreaks on their wings, confusing a predator about where their head is.
- Soricidae (eat pupae)
- Sigmodontinae (eat pupae)
- Anura (eat adults)
- Araneae, especially crab spiders and orb-weavers (eat adults)
- Formicidae (eat caterpillars)
- Hymenoptera (eat caterpillars and adults)
- mantids (eat adults)
- Diptera (eat caterpillars)
- Coccinellidae (eat eggs)
- Chrysopidae (eat eggs)
- Acari (eat eggs)
Life History and Behavior
Communication and Perception
These butterflies communicate mainly with their scent and their colors. Males attract mates with scent and display, and females leave a scent mark on plants where they have laid eggs.
Like all Moths and Butterflies, this family has complete metamorphosis. See More Information on Butterflies and Moths for an explanation of this. Some species in this group spin cocoons, others don't but the pupae attach themselves to plant stems with silk. Different species overwinter in different stages, usually larvae or pupae, rarely eggs and never as adults.
Development - Life Cycle: metamorphosis
All Blues, Coppers, and Hairstreaks live for a year or less. Many complete their whole life in a few months.
After mating, females lay eggs one by one on the flowers, twigs or leaves of food plants. This group often has two generations a year, one that spends the winter as caterpillars or pupae and fly as adults in the spring, and a second that complete their life-cycle in the summer and fly as adults in the fall.
Breeding season: May to October in Michigan.
Key Reproductive Features: iteroparous ; seasonal breeding ; sexual ; oviparous
These butterflies don't care for their young.
Parental Investment: no parental involvement
Evolution and Systematics
Systematic and taxonomic history
The relationhip between the lycaenids and riodinids is undetermined. Traditionally the riodinids have been considered a subfamily within the Lycaenidae, but this is not well supported. Harvey (1987) proposed 5 characters (based on early stages) that support a sister relationship between the two groups. However, Robbins (1988a, 1990)
Discussion of Phylogenetic Relationships
The higher classification of Lycaenidae is currently in flux. The subfamilies, tribes and subtribes enumerated within this clade are for the most part the groups listed by Eliot (1973, 1992 [in Corbet et al.]). Note that the higher classifications presented by Eliot in 1973 and 1992 differ from one another in that the 1992 version reduced the three diverse subfamilies Lycaeninae, Polyommatinae and Theclinae to tribes within an inclusive "Lycaeninae," in order to reflect their affinity to one another with respect to Curetinae, Poritiinae and Miletinae. That change has not been generally adopted by the lycaenid research community (R. K. Robbins, pers. comm, 4/08), and has not been incorporated here. Note also that Eliot's classification was intuitive, and the monophyly and implied relationships among these groups should therefore be viewed as tentative, pending revelations from the large amount of molecular evidence that is currently being gathered by Naomi Pierce and colleagues. The relationships shown among the four subfamilies are those inferred by Wahlberg et al. (2005).
Molecular Biology and Genetics
Statistics of barcoding coverage
Specimen Records: 18314
Specimens with Sequences: 16075
Specimens with Barcodes: 15140
Species With Barcodes: 2232
Public Records: 4960
Public Species: 1073
Public BINs: 662
One species in Michigan is considered endangered. It is Lycaeides melissa samuelis, the Karner Blue. Scientists are still not sure if it is just a subspecies of a more common species, or if it is a completely separate species. The Karner Blue only exists in a few small populations scattered from Minnesota to New York. It needs dry pine-oak barrens to live in, and only feeds on one kind of plants (lupines, genus Lupinus). This habitat has mostly been converted into agricultural fields, and is hard to find.
Relevance to Humans and Ecosystems
Economic Importance for Humans: Negative
These butterflies don't have strong effects on people one way or another. A few species sometimes eat crop plants but this is not common and they rarely do enough damage to matter.
Negative Impacts: crop pest
Lycaenidae is the second-largest family of butterflies (behind the brush-footed butterflies), with over 5,000 species worldwide, whose members are also called gossamer-winged butterflies. They constitute about 30% of the known butterfly species.
The family is traditionally divided into the subfamilies of the blues (Polyommatinae), the coppers (Lycaeninae), the hairstreaks (Theclinae) and the harvesters (Miletinae); others include the Lipteninae, Liphyrinae, Curetinae and Poritiinae. A few authorities still include the family Riodinidae within the Lycaenidae. The monotypic former subfamily Styginae represented by Styx infernalis from the Peruvian Andes has been placed within the subfamily Euselasiinae of the family Riodinidae.
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 larva are capable of producing vibrations and low sounds that are transmitted through the substrates they inhabit. They use these sounds to communicate with ants.
Adult individuals often have hairy antenna-like tails complete with black and white annulated 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.
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, 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. The caterpillars pupate inside the ant's nest and the ants continue to look after the pupa. Just before the adult emerges 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 and they include small glands on the skin of the caterpillars called "pore cupola organs". Caterpillars of many species, except those of the Riodininae, have a gland on the seventh abdominal segment that produces honey dew 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 (third segment of thorax in the Riodininae) and this is cylindrical and topped with a ring of spikes and emits chemical signals which are believed to help in communicating with ants.
|Phylogeny of the family.|
- Lipteninae (Afrotropical) may be ranked as a tribe of Poritiinae. (Liptenini) 
- Poritiinae (Oriental and Afrotropical)
- Liphyrinae (mostly African, some Asian) may be ranked as a tribe of Miletinae. (Liphyrini)  Selected species
- Liphyra brassolis – moth butterfly (largest lycaenid)
- Miletinae – harvesters (mostly African, or Oriental, some Holarctic). Probably all feed on aphids or their secretions.
- Curetinae – sunbeams (Oriental or Palaearctic) selected species
- Curetis thetis – Indian sunbeam
- Theclinae – hairstreaks (usually tailed) and elfins (not tailed) (global) may be ranked as a tribe of Lycaeninae (Theclini ) see the clade below right. Selected species
- Lycaeninae – coppers (Holarctic) selected species
- Polyommatinae – blues (global) selected species
- Caleta spp.
- Celastrina ladon – spring azure
- Talicada nyseus – red pierrot
- Cupido comyntas – Eastern tailed-blue
- Cupido minimus – small blue
- Pseudozizeeria maha – pale grass blue
- Euphilotes battoides allyni – El Segundo blue
- Euphilotes pallescens arenamontana – Sand Mountain blue
- Chilades - jewel blues
- Plebejus argus – silver-studded blue
- Icaricia icarioides fenderi – Fender's blue
- Polyommatus icarus – common blue
- Polyommatus semiargus – mazarine blue
- Glaucopsyche lygdamus – silvery blue
- Glaucopsyche lygdamus palosverdesensis – Palos Verdes blue
- Glaucopsyche xerces (extinct) – Xerces blue
- Phengaris xiushani
- Maculinea arion – large blue
- Fiedler, K. 1996. Host-plant relationships of lycaenid butterflies: large-scale patterns, interactions with plant chemistry, and mutualism with ants. Entomologia Experimentalis et Applicata 80(1):259-267 doi:10.1007/BF00194770 
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- Pierce NE, Braby MF, Heath A, Lohman DJ, Mathew J, Rand DB, Travassos MA. 2002. The ecology and evolution of ant association in the Lycaenidae (Lepidoptera.) Annual Review of Entomology 47: 733-771. PDF
- 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
- Australian Museum factsheets Accessed 4 November 2010 on the Wayback Machine.
- Brower, Andrew V. Z. 2008. Lycaenidae. Version 25 April 2008 (temporary).  in The Tree of Life Web Project, 
- Ackery, P. R., R. de Jong, and R. I. Vane-Wright. 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.
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- , Site of Markku Savela
- , Site of Markku Savela
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