Nemeobiinae (but see text)
Zygia metalmark (Lemonias zygia)
in the Pantanal, Brazil
Riodinidae is the family of metalmark butterflies. The common name
"metalmarks" refers to the small metallic-looking spots commonly found
on their wings. There are 1532 species and 146 genera of metalmark
butterflies in the world. Although mostly neotropical in
distribution, the family is represented both in the Nearctic and the
2 Distinguishing features
3 Taxonomy and systematics
3.2 Genera of uncertain position
5 Life cycle
5.1 Food plants
6 Economic significance
9 Further reading
10 External links
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. 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.
Mimicry causes often closely related species to have completely
different wing patterns, for example the genus Thisbe. 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. 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
The delimitation from the closely related
Lycaenidae by morphological
autapomorphy 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
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
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 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.
Taxonomy and systematics
Riodinidae is currently treated as a distinct family within the
superfamily Papilionoidea, but in the past they were held to be the
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. Based on morphological studies
Ackery et al. in the manual of zoology (Kristensen 1998, cf.
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 (based on homologous DNA sequences)
establishes a sister group relationship between the
Riodinidae and the
Lycaenidae accepted almost unanimously.
Duke of Burgundy
Riodinidae consists of three subfamilies. They are:
Euselasiinae – a handful of genera
New World (Americas)
Nemeobiinae – sometimes treated as a tribe, Nemeobiini, but
which of the remaining two subfamilies it would belongs is uncertain.
Riodinidae incertae sedis.
Riodininae – some dozens of genera
New World (Americas)
Genera of uncertain position
Several genera from the
Old World are of more uncertain affiliations;
some of them are monotypic. Such
Riodinidae incertae sedis are:
Hamearis – Duke of Burgundy (tribe Zemerini or distinct subfamily
Taxila – orange harlequin
Stiboges – columbine
Dodona – Punches
The fossil genus
Lithopsyche is sometimes placed here but sometimes in
Amazonas tropical rainforest is the habitat for most species of
Species occur in a variety of habitats, but have a unique distribution
focus in the tropical rain forests 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: each of the 19 species has
a vicariant distribution area, three originally separate forests
(upper, lower Amazonas, Guyana) can be derived from the relationship
of between the species.
The food plants for the caterpillars include 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
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). Predatory feeding
has also been shown in Alesa amesis. A number of species associate
and are protected by ants during one or more stages of their life
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
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
Lycaenidae. The caterpillars are usually hairy, 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. There is no cocoon.
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 linages 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. But there are counter arguments.
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.
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,
Zingiberaceae as well as bryophytes and lichens.
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
Bromeliceae or Orchidaceae.
^ Erik J. van Nieukerken, Lauri Kaila, Ian J. Kitching, Niels P.
Kristensen, David C. Lees, Joël Minet, Charles Mitter, Marko Mutanen,
Jerome C. Regier, Thomas J. Simonsen, Niklas Wahlberg, Shen-Horn Yen,
Reza Zahiri, David Adamski, Joaquin Baixeras, Daniel Bartsch, Bengt
Å. Bengtsson, John W. Brown, Sibyl Rae Bucheli, Donald R. Davis,
Jurate De Prins, Willy De Prins, Marc E. Epstein, Patricia
Gentili-Poole, Cees Gielis, Peter Hättenschwiler, Axel Hausmann,
Jeremy D. Holloway, Axel Kallies, Ole Karsholt, Akito Y. Kawahara,
Sjaak (J.C.) Koster, Mikhail V. Kozlov, J. Donald Lafontaine, Gerardo
Lamas, Jean-François Landry, Sangmi Lee, Matthias Nuss, Kyu-Tek Park,
Carla Penz, Jadranka Rota, Alexander Schintlmeister, B. Christian
Schmidt, Jae-Cheon Sohn, M. Alma Solis, Gerhard M. Tarmann, Andrew D.
Warren, Susan Weller, Roman V. Yakovlev, Vadim V. Zolotuhin, Andreas
Zwick (2011): Order
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classification and survey of taxonomic richness. Zootaxa 3148:
^ Thomas C. Emmel, Edward S. Ross (Hrsg.): Wunderbare und
geheimnisvolle Welt der Schmetterlinge. 1. Auflage. Bertelsmann,
Gütersloh und Berlin 1976 (übersetzt von Irmgard Jung),
^ Mathieu Joron (2008): Batesian Mimicry: Can a Leopard Change Its
Spots — and Get Them Back? Current Biology Volume 18, Issue 11:
^ Carla M. Penz & Philip J. DeVries (2001): A phylogenetic
Thisbe and Uraneis butterflies (Riodinidae,
Nymphidiini). Contributions in Science 485: 1-27.
^ K.S. Brown Jr., B. von Schoultz, A.O. Saura, A. Saura (2012):
Chromosomal evolution in the South American
Papilionoidea). Hereditas 149: 128–138.
^ Rienk de Jong, Philip R. Ackery, Richard I. Vane-Wright (1996):The
higher classification of butterflies (Lepidoptera): problems and
Insect Systematics & Evolution, Volume 27, Issue 1: 65
– 101. doi:10.1163/187631296X00205.
^ Robert K. Robbins (1988): Comparative morphology of the butterfly
foreleg coxa and trochanter (Lepidoptera) and its systematic
implications. Proceedings of the Entomological Society of Washington
90 (2): 133-154.
^ Borror et al. (1989)
^ Zhao F, Huang DY, Sun XY, Shi QH, Hao JS, Zhang LL, Yang Q. (2013):
The first mitochondrial genome for the butterfly family Riodinidae
Abisara fylloides) and its systematic implications. Zoological
Research 34 (E4−5): E109−E119.
^ Philip R. Ackery, Rienk de Jong, Richard I. Vane-Wright: The
Butterflies: Hedyloidea, Hesperioidea, Papilionoidea. In: Niels P.
Kristensen (editor): Lepidoptera, Moths and Butterflies. Volume 1:
Evolution, Systematics, and Biogeography. Walter de Gruvter, Berlin
& New York 1999. vgl. pp. 283-284
^ Niels P. Kristensen, Malcolm J. Scoble, Ole Karsholt (2007):
Lepidoptera phylogeny and systematics: the state of inventorying moth
and butterfly diversity. Zootaxa 1668: 699–747.
^ Dana L. Campbell and Naomi E. Pierce (2003): Phylogenetic
relationships of the Riodinidae: Implications for the evolution of ant
association. In: C. Boggs, P. Ehrlich, W.B. Watt (editors).
Butterflies as Model Systems. Chicago University Press: 395-408.
^ Niklas Wahlberg, Michael F Braby, Andrew V.Z Brower, Rienk de Jong,
Ming-Min Lee, Sören Nylin, Naomi E Pierce, Felix A.H Sperling, Roger
Vila, Andrew D Warren and Evgueni Zakharov (2005): Synergistic effects
of combining morphological and molecular data in resolving the
phylogeny of butterflies and skippers. Proceedings of the Royal
Society Series B 272: 1577-1586. doi:10.1098/rspb.2005.3124
^ aria Heikkilä, Lauri Kaila, Marko Mutanen, Carlos Peña, Niklas
Wahlberg (2012) Cretaceous origin and repeated tertiary
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^ Hall, J.P.W. (2004b)
^ See Savela (2007) for references.
^ J.P.W. Hall (2004): Metalmark Butterflies (Lepidoptera: Riodinidae)
In J.L. Capinera (editor) Encyclopedia of Entomology, Vol. 2 Kluwer
Academic Publishers, 2004. pp. 1383–1386.
^ Jason P.W. Hall & Donald J. Harvey (2002): The phylogeography of
Amazonia revisited: new evidence from Riodinid butterflies. Evolution,
^ P.J. DeVries, I.A. Chacon & D. Murray (1992) Toward a better
understanding of host use and biodiversity in riodinid butterflies
(Lepidoptera). Journal of Research on the Lepidoptera,
^ DeVries, P.J. & C.M. Penz. 2000. Entomophagy, behavior, and
elongated thoracic legs in the myrmecophilous Neotropical butterfly
Alesa amesis (Riodinidae). Biotropica 32: 712-721.
^ DeVries, P. J. 1997. The Butterflies of Costa Rica and their Natural
History. II: Riodinidae. Princeton Univ. Press, New Jersey, p. 288.
^ Jason P.W. Hall & Keith R. Willmott (2000): Patterns of feeding
behaviour in adult male riodinid butterflies and their relationship to
morphology and ecology. Biological Journal of the Linnean Society 69:
^ DeVries, P. J. 1991. Ecological and evolutionary patterns in
riodinid butterflies. IN: Ant-Plant Interactions. C. Huxley & D.
F. Cutler (eds.) Oxford Univ. Press, pp. 143-156.
^ DeVries, P.J. (2001): [Riodinidae]. In Levin, S.A. (ed.):
Encyclopaedia of Biodiversity. Academic Press.
^ Ferrer-Paris, José R.; Sánchez-Mercado, Ada; Viloria, Ángel L.;
Donaldson, John (2013). "Congruence and Diversity of Butterfly-Host
Plant Associations at Higher Taxonomic Levels". PLoS ONE. 8 (5):
e63570. doi:10.1371/journal.pone.0063570. PMC 3662771 .
PMID 23717448. Retrieved 29 June 2016.
Borror, Donald J.; Triplehorn, Charles A. & Johnson, Norman F.
(1989): An introduction to the study of insects (6th ed.).
Philadelphia: Saunders College Pub. ISBN 0-03-025397-7.
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): Markku Savela's
Lepidoptera and some other life
forms: Riodinidae. Version of 7 August 2007. Retrieved 9 September
Charles A. Bridges, 1994. Catalogue of the family-group, genus-group
and species-group names of the
Riodinidae & Lycaenidae
(Lepidoptera) of the world Urbana, Ill.:C.A. Bridges pdf
Campbell, D. L. & Pierce, N. E. 2003: Chapter 18: Phylogenetic
Relationships of the Riodinidae:Implications for the Evolution of Ant
Association. Pp. 395–408. – In: Boggs, C. L.,Watt, B. &
Ehrlich, P. R. (eds): Butterflies. Ecology and Evolution Taking
Flight. The University of Chicago Press, Cambridge University Press,
Chicago and London pdf
Glassberg, Jeffrey Butterflies through Binoculars, The West (2001)
Guppy, Crispin S. and Shepard, Jon H. Butterflies of British Columbia
James, David G. and Nunnallee, David Life Histories of Cascadia
Pelham, Jonathan Catalogue of the Butterflies of the United States and
Pyle, Robert Michael The Butterflies of Cascadia (2002)
Seitz, A., 1916. Family: Erycinidae. In A. Seitz (editor),
Macrolepidoptera of the World, vol. 5: 617–738. Stuttgart: Alfred
Kernen. also available as pdf. Out of date but very useful.
Wikimedia Commons has media related to Riodinidae.
Wikispecies has information related to Riodinidae
Riodinidae in French
Riodinidae at eol
TOL - Implied clade links to species lists.
TOL Images 3 pages.
LEPINDEX Taxonomy project of Natural History Museum, London
Barcode of Life Includes images.
Idaho Museum of Natural History
Calephelis virginiensis, little metalmark on the UF / IFAS Featured
Creatures Web site
Mariposas mexicana Excellent high resolution images of Mexican
Neotropical Butterflies Metalmark Gallery
Butterflies of America Images of type specimens
Butterflies and Moths of North America
Micropterigidae (mandibulate archaic moths)
Agathiphagidae (kauri moths)
Acanthopteroctetidae (archaic sun moths)
Neopseustidae (archaic bell moths)
Hepialidae (swift moths, ghost moths)
Neotheoridae (Amazonian primitive ghost moths)
Palaeosetidae (miniature ghost moths)
Prototheoridae (African primitive ghost moths)
Mnesarchaeidae (New Zealand primitive moths)
Adelidae (fairy longhorn moths)
Prodoxidae (yucca moths)
Andesianidae (Andean endemic moths)
Nepticulidae (pigmy, or midget moths)
Opostegidae (white eyecap moths)
Palaephatidae (Gondwanaland moths)
Tischeriidae (trumpet leaf miner moths)
Acrolophidae (burrowing webworm moths)
Old World spiny-winged moths)
Psychidae (bagworm moths)
Tineidae (fungus moths)
Bucculatricidae (ribbed cocoon makers)
Douglasiidae (Douglas moths)
Acrolepiidae (false diamondback moths)
Glyphipterigidae (sedge moths)
Yponomeutidae (ermine moths)
Coleophoridae (case-bearers, case moths)
Cosmopterigidae (cosmet moths)
Elachistidae (grass-miner moths)
Gelechiidae (twirler moths)
Lecithoceridae (long-horned moths)
Momphidae (mompha moths)
Oecophoridae (concealer moths)
Scythrididae (flower moths)
Xyloryctidae (timber moths)
Zygaenidae (burnet, forester, or smoky moths)
Megalopygidae (flannel moths)
Epipyropidae (planthopper parasite moths)
Dalceridae (slug caterpillars)
Limacodidae (slug, or cup moths)
Cossidae (carpenter millers, or goat moths)
Dudgeoneidae (dudgeon carpenter moths)
Brachodidae (little bear moths)
Castniidae (castniid moths: giant butterfly-moths, sun moths)
Sesiidae (clearwing moths)
Choreutidae (metalmark moths)
Tortricidae (tortrix moths)
Urodidae (false burnet moths)
Schreckensteiniidae (bristle-legged moths)
Epermeniidae (fringe-tufted moths)
Alucitidae (many-plumed moths)
Tineodidae (false plume moths)
Pterophoridae (plume moths)
Copromorphidae (tropical fruitworm moths)
Carposinidae (fruitworm moths)
Hyblaeidae (teak moths)
Pyralidae (snout moths)
Crambidae (grass moth)
Thyrididae (picture-winged leaf moths)
Mimallonidae (sack bearer moths)
Lasiocampidae (eggars, snout moths, or lappet moths)
Anthelidae (Australian lappet moth)
Bombycidae (silk moths)
Brahmaeidae (Brahmin moths)
Carthaeidae (Dryandra moth)
Endromidae (Kentish glory and relatives)
Sphingidae (hawk moths, sphinx moths and hornworms)
Erebidae (underwing, tiger, tussock, litter, snout, owlet moths)
Noctuidae (daggers, sallows, owlet moths, quakers, cutworms, darts)
Nolidae (tuft moths)
Notodontidae (prominents, kittens)
Epicopeiidae (oriental swallowtail moths)
Geometridae (geometer moths)
Cimeliidae (gold moths)
Old World butterfly-moths)
Hedylidae (American moth-butterflies)
Lycaenidae (gossamer-winged butterflies: blues, coppers and relatives)
Nymphalidae (brush-footed, or four-footed butterflies)
Papilionidae (swallowtail butterflies)
Pieridae (whites, yellows, orangetips, sulphurs)
Monotrysia is not a clade.
Taxonomy of the Lepidoptera
Lists by region
Fauna Europaea: 357093