Riodinidae is the

family Family (from ) is a Social group, group of people related either by consanguinity (by recognized birth) or Affinity (law), affinity (by marriage or other relationship). It forms the basis for social order. Ideally, families offer predictabili ...

of metalmark

butterflies Butterflies are winged insects from the lepidopteran superfamily Papilionoidea, characterized by large, often brightly coloured wings that often fold together when at rest, and a conspicuous, fluttering flight. The oldest butterfly fossi ...

. The common name "metalmarks" refers to the small, metallic-looking spots commonly found on their wings. The 1,532 species are placed in 146 genera. Although mostly

Neotropical The Neotropical realm is one of the eight biogeographic realms constituting Earth's land surface. Physically, it includes the tropical terrestrial ecoregions of the Americas and the entire South American temperate zone. Definition In biogeogra ...

in distribution, the family is also represented both in the

Nearctic The Nearctic realm is one of the eight biogeographic realms constituting the Earth's land surface. The Nearctic realm covers most of North America, including Greenland, Central Florida, and the highlands of Mexico. The parts of North America ...

Palearctic The Palearctic or Palaearctic is a biogeographic realm of the Earth, the largest of eight. Confined almost entirely to the Eastern Hemisphere, it stretches across Europe and Asia, north of the foothills of the Himalayas, and North Africa. Th ...

Australasian Australasia is a subregion of Oceania, comprising Australia, New Zealand (overlapping with Polynesia), and sometimes including New Guinea and surrounding islands (overlapping with Melanesia). The term is used in a number of different contexts, ...

('' Dicallaneura''),

Afrotropic The Afrotropical realm is one of the Earth's eight biogeographic realms. It includes Sub-Saharan Africa, the southern Arabian Peninsula, the island of Madagascar, and the islands of the western Indian Ocean. It was formerly known as the Ethiopi ...

('' Afriodinia'', ''

Saribia ''Saribia'' is a genus of butterflies in the family Riodinidae. Species *'' Saribia decaryi'' (Le Cerf, 1922) *'' Saribia ochracea'' Riley, 1932 *'' Saribia perroti'' Riley, 1932 *'' Saribia tepahi'' (Boisduval, 1833) References Extern ...

''), and

Indomalayan realm The Indomalayan realm is one of the eight biogeographic realms. It extends across most of South and Southeast Asia and into the southern parts of East Asia. Also called the Oriental realm by biogeographers, Indomalaya spreads all over the Ind ...

Description

The family includes small to medium-sized species, from 12 to 60 mm wingspan, often with vibrant structural colouring. The wing shape is very different within the family. They may resemble butterflies in other groups, some are similar to

Satyrinae The Satyrinae, the satyrines or satyrids, commonly known as the browns, are a subfamily of the Nymphalidae (brush-footed butterflies). They were formerly considered a distinct family, Satyridae. This group contains nearly half of the known divers ...

, some are bright yellow reminiscent of

Coliadinae Coliadinae, the sulphurs or yellows, are a subfamily of butterflies with about 300 described species. There are 36 species in North America, where they range from Mexico to northern Canada. In most species, males are easily distinguished from ...

and others (examples ''

Barbicornis ''Barbicornis'' is a monotypic butterfly genus of the family Riodinidae with its single species ''Barbicornis basilis'' present in Paraguay, Brazil and Argentina. The species is easily recognizable by the very small hindwings provided with a ...

'', ''

Rhetus arcius ''Rhetus arcius'', the long-tailed metalmark, s a species of Neotropical butterfly, first described in Carl Linnaeus' 1763 ''Centuria Insectorum file:Centuria Insectorum.png, The first page of ''Centuria Insectorum'', as included in ''Amoenita ...

'', ''

Helicopis ''Helicopis'' is a Neotropical genus of butterflies of the family Riodinidae. List of species * ''Helicopis gnidus'' (Fabricius, 1787 * ''Helicopis cupido'' (Linnaeus, 1758) * ''Helicopis endymiaena'' (Hübner,

819 __NOTOC__ Year 819 ( DCCCXIX) was a common year starting on Saturday of the Julian calendar. Events By place Europe * Spring – Emperor Louis I marries Judith of Bavaria in Aachen.Rogers, Barbara, Bernhard W. Scholz, and Nithard ...

References Funet
Ri ...

'', ''

Chorinea ''Chorinea'' is a Neotropical metalmark butterfly genus. Species Listed alphabetically:Papilionidae Swallowtail butterflies are large, colorful Butterfly, butterflies in the family Papilionidae, and include over 550 species. Though the majority are tropical, members of the family inhabit every continent except Antarctica. The family includ ...

. The colouration ranges from muted colours in the temperate zone species to iridescent blue and green wings and transparent wings in tropical species. The golden or silvery metallic spots on the wings in many species of the Americas gave them the English common name "metalmarks". A number of species

mimic In evolutionary biology, mimicry is an evolved resemblance between an organism and another object, often an organism of another species. Mimicry may evolve between different species, or between individuals of the same species. In the simples ...

poisonous moths of several families and there are often extensive mimicry rings of similar-looking species, grouped around a model. Mimicry causes often closely related species to have completely different wing patterns, for example the genus ''

Thisbe In Greek mythology, Pyramus and Thisbe () are a pair of ill-fated lovers from Babylon, whose story is best known from Ovid's narrative poem ''Metamorphoses''. The tragic myth has been retold by many authors. Pyramus and Thisbe's parents, driv ...

''. Many species mimic the stain and stripe pattern of toxic

Nymphalidae The Nymphalidae are the largest family of butterflies, with more than 6,000 species distributed throughout most of the world. Belonging to the superfamily Papilionoidea, they are usually medium-sized to large butterflies. Most species ha ...

. Batesian mimicry seems to be more common than in any other insect family of similar size. Reasons for this are unknown. Another example is ''

Ithomeis ''Ithomeis'' is a genus in the butterfly family Riodinidae present only in the Neotropical realm. Species *''Ithomeis aurantiaca'' Bates, 1862 present in Guyana, Brazil, Colombia, Venezuela, Ecuador and Peru *''Ithomeis eulema'' Hewitson, 1870 ...

'' where different subspecies resemble the species they mimic in different parts of the geographic range more than they resemble each other. The delimitation from the closely related

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

by morphological

autapomorphy In phylogenetics, an autapomorphy is a distinctive feature, known as a Synapomorphy, derived trait, that is unique to a given taxon. That is, it is found only in one taxon, but not found in any others or Outgroup (cladistics), outgroup taxa, not ...

is difficult. The first pair of legs of the males, which arises on the prothorax, is less than half as long as the legs of the pterothorax and they are not used for walking. The individual segments of the tarsus are sometimes fused together and fused with the tibia, and the pretarsi have no claws. This feature is also found in some Lycaenidae (and also the

Monotrysia The Monotrysia are a group of moths in the lepidopteran order, not currently considered to be a natural group or clade. The group is so named because the female has a single genital opening for mating and laying eggs, in contrast to the rest of ...

), but in these the legs are always much longer. The sensory hairs on the tarsi of the female forelimbs are arranged in a group. These groups which are arranged in pairs can be found in the other taxa of the Papilionoidea. The third problematic apomorphy is the absence of the rear projections (apophyses) of the female genitalia. This feature (absence) is found as well in some species of the subfamily of

Poritiinae Poritiinae is a subfamily of butterflies, the larvae of which are unusual for feeding on algae and foliate lichen. Systematics * Tribe Poritiini - Oriental ** '' Cyaniriodes'' de Nicéville, 1890 (sometimes placed in Lycaeninae) ** '' Poris ...

. In almost all Riodinidae, the coxae of the front legs are extended on males jutting out over the trochanter (only hinted at in ''Styx infernalis'' and ''Corrachia leucoplaga''). If there are similar projections in Lycaenidae (in genera ''Curetis'', ''Feniseca'' and ''Poritia''), they are built differently in detail and may be, for example, dorsally convex. In addition, almost all Riodinidae in contrast to the Lycaenidae have a humeral vein in the hindwings and the costa is thickened (exceptions in the subfamily Hamearinae). The head in relation to the eyes is wider than in Lycaenidae, making the antennal bases further away from the eye. The relatively long antennae often reach half of the front wing length. Riodinidae have an unusual variety in

chromosome A chromosome is a package of DNA containing part or all of the genetic material of an organism. In most chromosomes, the very long thin DNA fibers are coated with nucleosome-forming packaging proteins; in eukaryotic cells, the most import ...

numbers, only some very basal groups have the number typical for butterflies (between 29 and 31) or the number characteristic of Lycaenidae (23 to 24). Numbers between 9 and 110 occur. In some cases, representatives of a morphologically indistinguishable

cryptospecies In biology, a species complex is a group of closely related organisms that are so similar in appearance and other features that the boundaries between them are often unclear. The taxa in the complex may be able to hybridize readily with each oth ...

have different chromosome numbers and are reproductively isolated.

Distinguishing features

Like the lycaenids, the males of this family have reduced forelegs while the females have full-sized, fully functional forelegs. The foreleg of males is often reduced and has a uniquely shaped first segment (the coxa) which extends beyond its joint with the second segment, rather than meeting it flush. They have a unique venation on the hindwing: the costa of the hindwing is thickened out to the humeral angle and the humeral vein is short.

Taxonomy and systematics

Riodinidae is currently treated as a distinct family within the superfamily

Papilionoidea Butterflies are winged insects from the lepidopteran superfamily Papilionoidea, characterized by large, often brightly coloured wings that often fold together when at rest, and a conspicuous, fluttering flight. The oldest butterfly fossi ...

, but in the past they were held to be the subfamily Riodininae of the

. Earlier, they were considered to be part of the now defunct family Erycinidae, whose species are divided between this family and the subfamily

Libytheinae The Libytheinae are a nymphalid subfamily known as snout butterflies, containing two valid genera and about ten species: six in ''Libythea'' and four in '' Libytheana''. The common name refers to the thick labial palps (pedipalps) that look like ...

. Today, most systematists prefer to accept an independent family even if there are counter arguments. Based on morphological studies Ackery et al. in the manual of zoology (Kristensen 1998, cf. literature) placed Riodininae within the Lycaenidae. Kristensen et al. accepted the updating of the manual in 2007 raising the classification to family rank at least on a provisional basis.

Molecular phylogenetics Molecular phylogenetics () is the branch of phylogeny that analyzes genetic, hereditary molecular differences, predominantly in DNA sequences, to gain information on an organism's evolutionary relationships. From these analyses, it is possible to ...

(based on homologous DNA sequences) establishes a

sister group In phylogenetics, a sister group or sister taxon, also called an adelphotaxon, comprises the closest relative(s) of another given unit in an evolutionary tree. Definition The expression is most easily illustrated by a cladogram: Taxon A and ...

relationship between the Riodinidae and the Lycaenidae accepted almost unanimously.

Subfamilies

The family Riodinidae has been historically been classified using a two subfamily (Stichel, 1928) or three subfamily (Callaghan and Lamas, 2004) system. Genetic data from Seraphim ''et al.'' (2018) supports the two subfamily interpretation, with the subfamily

Euselasiinae Euselasiinae is a subfamily of Riodinidae. The species are confined to the Neotropical realm. Genera From Funet *'' Corrachia'' Schaus, 1913 *'' Euselasia'' (Hübner, 1819) a populous genus with many species. *''Hades'' (Westwood, 1851) *'' Me ...

being subsumed entirely within the Old World

Nemeobiinae Nemeobiinae is a subfamily of Riodinidae, the metalmark family. The subfamily's members consist entirely of Old World members of the Riodinid family. Recent revisions to the subfamily have begun to include members located within the New World as ...

. Two subfamily model (Stichel, 1928) *

Riodininae __NOTOC__ Riodininae is the largest of the three subfamilies within the metalmark butterfly family Family (from ) is a Social group, group of people related either by consanguinity (by recognized birth) or Affinity (law), affinity (by ma ...

Three subfamily model (Callaghan and Lamas, 2004) *

The

fossil A fossil (from Classical Latin , ) is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserve ...

genus '' Lithopsyche'' is sometimes placed here but sometimes within the

Biology

Species occur in a variety of habitats, but have a unique distribution focus in the

tropical rain forests Tropical rainforests are dense and warm rainforests with high rainfall typically found between 10° north and south of the Equator. They are a subset of the tropical forest biome that occurs roughly within the 28° latitudes (in the torrid zo ...

of South America. Many species are rarely found and have a relatively small distribution area. Species of the genus ''Charis'' were therefore used to reconstruct the history of the forest of the

Amazon basin The Amazon basin is the part of South America drained by the Amazon River and its tributary, tributaries. The Amazon drainage basin covers an area of about , or about 35.5 percent of the South American continent. It is located in the countries ...

: each of the 19 species has a vicariant distribution area, three originally separate forests (upper, lower Amazonas, Guyana) can be derived from the relationship between the species. The food plants for the caterpillars total more than 40 plant families. Mostly young leaves or flowers are used, and rarely fallen, dead leaves or lichen are eaten. The larvae feed mostly individually not gregariously. However, gregarious caterpillars are found within the Euselasiinae (''Euselasia''), Riodinini (''Melanis'') and Emesini (''Emesis''), with some species demonstrating processionary behaviours. Available evidence from ''Euselasia'' and ''Hades'' suggests the gregarious trait may be widespread among members of the subfamily

. The larva of ''Setabis lagus'' (Riodininae: Nymphidiini), is predatory. There are records of predation on larvae of ''Horiola'' species (family Membracidae) as well as scale insects (

Coccidae Coccidae, from Ancient Greek (''kókkos''), meaning "kernel" or "Kermes (insect), Kermes", are a family of scale insects belonging to the superfamily Scale insect, Coccoidea. They are commonly known as soft scales, wax scales or tortoise scales ...

). Predatory feeding has also been shown in ''Alesa amesis''. A number of species associate with and are protected by ants during one or more stages of their life cycle. A study in Ecuador based on adult male feeding records for 124 species in 41 genera of Riodinidae (out of a total of 441 species in 85 genera collected in the study) demonstrated that rotting fish and other carrion was the most frequently used food source in terms of numbers of individuals and taxa, attracting 89 species from 32 genera. Other food substrates visited in this study included flowers, damp sand or

mud-puddling Puddling is a behaviour in which an organism seeks out nutrients in certain moist substances such as rotting plant matter, mud, and carrion, and sucks up the fluid. Where the conditions are suitable, conspicuous insects such as butterfly, butterf ...

Life cycle

The eggs vary in shape, but often appear round and flattened, some have the shape of a dome or turban. They are similar to the eggs of the Lycaenidae. The

caterpillar Caterpillars ( ) are the larval stage of members of the order Lepidoptera (the insect order comprising butterflies and moths). As with most common names, the application of the word is arbitrary, since the larvae of sawflies (suborder ...

s are usually hairy and plump, and are the common overwintering stage. The caterpillars are usually longer than those of the Lycaenidae except in the

myrmecophilous thumb , Myrmecophilous aphids being tended by ants Myrmecophily ( , ) consists of positive, mutualistic, interspecies associations between ants and a variety of other organisms, such as plants, other arthropods, and fungi. It may also incl ...

species.

Pupa A pupa (; : pupae) is the life stage of some insects undergoing transformation between immature and mature stages. Insects that go through a pupal stage are holometabolous: they go through four distinct stages in their life cycle, the stages th ...

e are hairy and attached with silk to either the host plant or to ground debris or leaf litter. No

cocoon Cocoon may refer to: * Cocoon (silk), a pupal casing made by moth caterpillars and other insect larvae Music * Cocoon Recordings, a German record label * Cocoon (band), a French band * Cocoon (club), a techno club in Frankfurt am Main, Germany * ' ...

is seen. Several genera of Riodinidae have evolved intimate associations with ants, and their larvae are tended and defended by ant associates. This also is the case with several lineages of Lycaenidae and contributed to arguments for the uniting the two families. It is now recognized that myrmecophily arose several times among Riodinidae and Lycaenidae clades. Like their sister family Lycaenidae, numerous species of Riodinidae are myrmecophiles (involving about 280 ant species). The larvae of many species have special organs, which have a soothing or tempting effect on ants. Many Riodinidae larvae have so-called "tentacle nectary organs" on the eighth segment of the abdomen that secrete a fluid which is eaten by ants. Other tentacle organs on the third thoracic segment have been shown to emit

allomone An allomone (from Ancient Greek ' "other" and pheromone) is a type of semiochemical produced and released by an individual of one species that affects the behaviour of a member of another species to the benefit of the originator but not the rec ...

s which influence ants. Studies suggest caterpillar acoustic signals are used to enhance their symbioses with ants (see singing caterpillars). The location of riodinid organs that function in caterpillar-ant symbioses differs from those found in the Lycaenidae, suggesting that the organs in these two families of butterflies are not homologous in origin.

Food plants

The larvae feed on plants of the families

Araceae The Araceae are a family of monocotyledonous flowering plants in which flowers are borne on a type of inflorescence called a spadix. The spadix is usually accompanied by, and sometimes partially enclosed in, a spathe (or leaf-like bract). Also ...

Asteraceae Asteraceae () is a large family (biology), family of flowering plants that consists of over 32,000 known species in over 1,900 genera within the Order (biology), order Asterales. The number of species in Asteraceae is rivaled only by the Orchi ...

Bromeliaceae The Bromeliaceae (the bromeliads) are a family of monocot flowering plants of about 80 genera and 3700 known species, native mainly to the tropical Americas, with several species found in the American subtropics and one in tropical west Africa, ...

Bombacaceae Bombacaceae were long recognised as a family of flowering plants or Angiospermae. The family name was based on the type genus '' Bombax''. As is true for many botanical names, circumscription and status of the taxon has varied with taxonomic poi ...

Cecropiaceae The Urticaceae are a family, the nettle family, of flowering plants. The family name comes from the genus ''Urtica''. The Urticaceae include a number of well-known and useful plants, including nettles in the genus ''Urtica'', ramie (''Boehmeria ...

Clusiaceae The Clusiaceae or Guttiferae Juss. (1789) (''nom. alt. et cons.'' = alternative and valid name) are a family (biology), family of plants including 13 genera and ca 750 species. Several former members of Clusiacae are now placed in Calophyllaceae ...

Dilleniaceae Dilleniaceae is a family of flowering plants with 11 genera and about 430 known species. It is known to gardeners for the genus '' Hibbertia'', which contains many commercially valuable garden species. Description and distribution The family is ...

Euphorbiaceae Euphorbiaceae (), the spurge family, is a large family of flowering plants. In English, they are also commonly called euphorbias, which is also the name of Euphorbia, the type genus of the family. Most spurges, such as ''Euphorbia paralias'', ar ...

Fabaceae Fabaceae () or Leguminosae,International Code of Nomen ...

Lecythidaceae The Lecythidaceae ( ) comprise a family of about 20 genera and 250–300 species of woody plants native to tropical South America, Africa (including Madagascar), Asia and Australia. Well known members of the family include the cannonball tree ...

Loranthaceae Loranthaceae, commonly known as the showy mistletoes, is a family of flowering plants. It consists of about 75 genera and 1,000 species of woody plants, many of them hemiparasites. The three terrestrial species are '' Nuytsia floribunda'' (the ...

Malpighiaceae Malpighiaceae is a family of flowering plants in the order Malpighiales. It comprises about 73 genera and 1315 species, all of which are native to the tropics and subtropics. About 80% of the genera and 90% of the species occur in the New World ( ...

Marantaceae The Marantaceae are a family, the arrowroot family, or the prayer plant family, of flowering plants consisting of 31 genera and around 530 species, defining it as one of the most species-rich families in its order. Kennedy, H. (2000). “Diversif ...

Melastomataceae Melastomataceae () is a family of dicotyledonous flowering plants found mostly in the tropics (two-thirds of the genera are from the New World tropics) comprising c. 175 genera and c. 5115 known species. Melastomes are annual or perennial herbs ...

Myrtaceae Myrtaceae (), the myrtle family, is a family of dicotyledonous plants placed within the order Myrtales. Myrtle, pōhutukawa, bay rum tree, clove, guava, acca (feijoa), allspice, and eucalyptus are some notable members of this group. All ...

Orchidaceae Orchids are plants that belong to the family (biology), family Orchidaceae (), a diverse and widespread group of flowering plants with blooms that are often colourful and fragrant. Orchids are cosmopolitan distribution, cosmopolitan plants that ...

Rubiaceae Rubiaceae () is a family (biology), family of flowering plants, commonly known as the coffee, madder, or bedstraw family. It consists of terrestrial trees, shrubs, lianas, or herbs that are recognizable by simple, opposite leaves with Petiole ( ...

Sapindaceae The Sapindaceae are a family (biology), family of flowering plants in the order Sapindales known as the soapberry family. It contains 138 genera and 1,858 accepted species. Examples include Aesculus, horse chestnut, maples, ackee and lychee. The ...

Zingiberaceae Zingiberaceae () or the ginger family is a family of flowering plants made up of about 50 genera with a total of about 1600 known species of aromatic perennial herbs with creeping horizontal or tuberous rhizomes distributed throughout tropical ...

as well as bryophytes and lichens.DeVries, P.J. (2001): iodinidae In Levin, S.A. (ed.): Encyclopaedia of Biodiversity. Academic Press.

Economic significance

The importance of Riodinidae species considered pests is very low. Some species of Euselasiinae feed on Myrtaceae of economic importance such as guava. A few Riodininae are specified as harmful to farmed Bromeliaceae or Orchidaceae.

References

* Borror, Donald J.; Triplehorn, Charles A. & Johnson, Norman F. (1989): ''An introduction to the study of insects'' (6th ed.). Philadelphia: Saunders College Pub. . * DeVries, P.J. (1997): Butterflies of Costa Rica and their natural history. Vol 2 Riodinidae. Princeton University Press. * Hall, J.P.W. (2004b): Metalmark Butterflies (Lepidoptera: Riodinidae), pp. 1383–1386. ''In'' J.L. Capinera (ed.) ''Encyclopedia of Entomology, Vol. 2.''
PDF
* Savela, Markku (2007)

Version of 7 August 2007. Retrieved 9 September 2007.

type specimens In biology, a type is a particular specimen (or in some cases a group of specimens) of an organism to which the scientific name of that organism is formally associated. In other words, a type is an example that serves to anchor or centralizes t ...

Butterflies and Moths of North America
{{Authority control Butterfly families Taxa named by Augustus Radcliffe Grote Papilionoidea Taxa described in 1895

Description

Distinguishing features

Taxonomy and systematics

Subfamilies

Biology

Life cycle

Food plants

Economic significance

References

Further reading

External links