Riodinidae | |
---|---|
Riodinidae, from Reise der Österreichischen Fregatte Novara um die Erde (1861–1876) | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Arthropoda |
Class: | Insecta |
Order: | Lepidoptera |
Superfamily: | Papilionoidea |
Family: | Riodinidae Grote, 1895 |
Subfamilies | |
Riodinidae is the family of metalmark butterflies. 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. [1] Although mostly Neotropical in distribution, the family is also represented both in the Nearctic, Palearctic, Australasian ( Dicallaneura ), Afrotropic ( Afriodinia , Saribia ), and Indomalayan realms. [2]
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, some are bright yellow reminiscent of Coliadinae and others (examples Barbicornis , Rhetus arcius , Helicopis , Chorinea ) have tails as do Papilionidae. The colouration ranges from muted colours in the temperate zone species to iridescent blue and green wings and transparent wings in tropical species. [3] 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 poisonous moths of several families and there are often extensive mimicry rings of similar-looking species, grouped around a model. [4] Mimicry causes often closely related species to have completely different wing patterns, for example the genus Thisbe . [5] Many species mimic the stain and stripe pattern of toxic Nymphalidae. Batesian mimicry seems to be more common than in any other insect family of similar size. [6] Reasons for this are unknown. Another example is Ithomeis 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 by morphological autapomorphy is difficult. [7] 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), 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.
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. [8] 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 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 have different chromosome numbers and are reproductively isolated.
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. [9]
Riodinidae is currently treated as a distinct family within the superfamily Papilionoidea, but in the past they were held to be the subfamily Riodininae of the Lycaenidae. Earlier, they were considered to be part of the now defunct family Erycinidae, whose species are divided between this family and the subfamily Libytheinae.
Today, most systematists prefer to accept an independent family even if there are counter arguments. [10] Based on morphological studies Ackery et al. [11] in the manual of zoology (Kristensen 1998, cf. literature) placed Riodininae within the Lycaenidae. Kristensen et al. [12] accepted the updating of the manual in 2007 raising the classification to family rank at least on a provisional basis.
Molecular phylogenetics (based on homologous DNA sequences) establishes a sister group relationship between the Riodinidae and the Lycaenidae accepted almost unanimously. [13] [14] [15]
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 being subsumed entirely within the Old World Nemeobiinae. [16]
Two subfamily model (Stichel, 1928)
Three subfamily model (Callaghan and Lamas, 2004)
The fossil genus Lithopsyche is sometimes placed here but sometimes within the Lycaenidae.
Species occur in a variety of habitats, but have a unique distribution focus in the tropical rain forests of South America. [17] 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: 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. [18]
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 Euselasiinae. [19]
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). Predatory feeding has also been shown in Alesa amesis. [20] A number of species associate with and are protected by ants during one or more stages of their life cycle. [21]
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. [22]
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 caterpillars 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 species. Pupae are hairy and attached with silk to either the host plant or to ground debris or leaf litter. No cocoon 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 allomones 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. [23]
The larvae feed on plants of the families Araceae, Asteraceae, Bromeliaceae, Bombacaceae, Cecropiaceae, Clusiaceae, Dilleniaceae, Euphorbiaceae, Fabaceae, Lecythidaceae, Loranthaceae, Malpighiaceae, Marantaceae, Melastomataceae, Myrtaceae, Orchidaceae, Rubiaceae, Sapindaceae, Zingiberaceae as well as bryophytes and lichens. [24] [25]
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.
Lepidoptera or lepidopterans is an order of winged insects which includes butterflies and moths. About 180,000 species of the Lepidoptera have been described, representing 10% of the total described species of living organisms, making it the second largest insect order with 126 families and 46 superfamilies, and one of the most widespread and widely recognizable insect orders in the world.
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 have a reduced pair of forelegs and many hold their colourful wings flat when resting. They are also called brush-footed butterflies or four-footed butterflies, because they are known to stand on only four legs while the other two are curled up; in some species, these forelegs have a brush-like set of hairs, which gives this family its other common name. Many species are brightly coloured and include popular species such as the emperors, monarch butterfly, admirals, tortoiseshells, and fritillaries. However, the under wings are, in contrast, often dull and in some species look remarkably like dead leaves, or are much paler, producing a cryptic effect that helps the butterflies blend into their surroundings.
The large blue is a species of butterfly in the family Lycaenidae. The species was first defined in 1758 and first recorded in Britain in 1795. In 1979 the species became mostly extinct in Britain but has been successfully reintroduced with new conservation methods. The species is classified as "near threatened" on the IUCN Red List of Threatened Species. Today P. arion can be found in Europe, the Caucasus, Armenia, western Siberia, Altai, north-western Kazakhstan and Sichuan.
The superfamily Papilionoidea contains all the butterflies except for the moth-like Hedyloidea.
Lycaenidae is the second-largest family of butterflies, with over 6,000 species worldwide, whose members are also called gossamer-winged butterflies. They constitute about 30% of the known butterfly species.
Hamearis lucina, the Duke of Burgundy, the only member of the genus Hamearis, is a European butterfly in the family Riodinidae. For many years, it was known as the "Duke of Burgundy fritillary", because the adult's chequered pattern is strongly reminiscent of "true" fritillaries of the family Nymphalidae.
Choreutidae, or metalmark moths, are a family of insects in the lepidopteran order whose relationships have been long disputed. It was placed previously in the superfamily Yponomeutoidea in family Glyphipterigidae and in superfamily Sesioidea. It is now considered to represent its own superfamily. The relationship of the family to the other lineages in the group "Apoditrysia" need a new assessment, especially with new molecular data.
Hedylidae, the "American moth-butterflies", is a family of insects in the order Lepidoptera, representing the superfamily Hedyloidea. They have traditionally been viewed as an extant sister group of the butterfly superfamily Papilionoidea, but a 2014 phylogenetic analysis has suggested Hedylidae is a subgroup of Papilionoidea, and not a sister group, and are more accurately referred to as butterflies rather than moths. They are represented by a single Neotropical genus Macrosoma with 35 currently recognized species.
Miletinae is a subfamily of the family Lycaenidae of butterflies, commonly called harvesters and woolly legs, and virtually unique among butterflies in having predatory larvae. Miletinae are entirely aphytophagous. The ecology of the Miletinae is little understood, but adults and larvae live in association with ants, and most known species feed on Hemiptera, though some, like Liphyra, feed on the ants themselves. The butterflies, ants, and hemipterans, in some cases, seem to have complex symbiotic relationships benefiting all.
Riodininae is the largest of the three subfamilies within the metalmark butterfly family, Riodinidae.
Butterfly evolution is the origin and diversification of butterflies through geologic time and over a large portion of the Earth's surface. The earliest known butterfly fossils are from the mid Eocene epoch, between 40-50 million years ago. Their development is closely linked to the evolution of flowering plants, since both adult butterflies and caterpillars feed on flowering plants. Of the 220,000 species of Lepidoptera, about 45,000 species are butterflies, which probably evolved from moths. Butterflies are found throughout the world, except in Antarctica, and are especially numerous in the tropics; they fall into eight different families.
Philip James DeVries is a tropical biologist whose research focuses on insect ecology and evolution, especially butterflies. His best-known work includes symbioses between caterpillars, ants and plants, and community level biodiversity of rainforest butterflies. DeVries is Professor Emeritus at the University of New Orleans, Research Associate at the American Museum of Natural History (NY), Florida Museum of Natural History (FL), and The Milwaukee Public Museum (WI).
Carla Maria Penz is a butterfly comparative morphologist and systematist, and the former Doris Zemurray Stone Chair in Biodiversity at the University of New Orleans. Her research also focuses on natural history and behavior, mostly of neotropical butterflies. She is currently Professor Emeritus at the University of New Orleans, Research Associate at the American Museum of Natural History (NY), Florida Museum of Natural History (FL), and The Milwaukee Public Museum (WI).
Singing caterpillars is a term coined by Philip James DeVries, referring to the fact that the larvae of ant-associated butterfly species of the families Riodinidae and Lycaenidae produce substrate borne sounds that attract ants. The study of these symbiotic associations was pioneered by Phil DeVries in Central America, and Naomi Pierce in Australia. Recently, Lucas Kaminski and collaborators are expanding the studies of riodinid-ant symbioses in Brazil.
Charis is a genus of the Riodinini tribe of metalmark butterflies. Nineteen species have been identified within the Charis cleonus complex and eight species within the Charis gynaea group (clade). Charis butterflies are common in the Neotropics and often live in primary and secondary growth. The Charis cleonus group exhibits contemporary parapatric distributions throughout Amazonia and are thought to have speciated allopatrically; residing in "areas of endemism". Some evidence suggests that Charis are reproductively isolated by mating preferences for different topographic areas and different times—of which, may have promoted speciation between the various groups.
Eurybia elvina, commonly known as the blind eurybia, is a Neotropical metalmark butterfly. Like many other riodinids, the caterpillars are myrmecophilous and have tentacle nectary organs that exude a fluid similar to that produced by the host plant Calathea ovandensis. This mutualistic relationship allows ants to harvest the exudate, and in return provide protection in the form of soil shelters for larvae. The larvae communicate with the ants by vibrations produced by the movement of its head. The species was described and given its binomial name by the German lepidopterist Hans Stichel in 1910.
Adelotypa annulifera is a species of riodinid butterfly found in South America. It was first described by Frederick DuCane Godman in 1903.
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 well, however, the subfamily continues to encompass the entirety of the Old World Riodinids.
Styx is a monotypic genus of butterflies in the metalmark family Riodinidae. It consists of one species, Styx infernalis, described by Otto Staudinger in 1875. It is endemic to Peru, where it inhabits tropical montane cloud forests between the elevations of 1000-1600 meters.