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{{Taxobox
| color = pink
| name = [[Butterfly|Butterflies]], [[moth]]s and [[Skipper (butterfly)|skippers]]
| image = Parthenos_sylvia_philippensis.jpg
| image_width = 250px
| image_caption = The '''Clipper''' ''Parthenos sylvia''
| regnum = [[Animal]]ia
| phylum = [[Arthropod]]a
| classis = [[Insect]]a
| subclassis = [[Pterygota]]
| infraclassis = [[Neoptera]]
| superordo = [[Endopterygota]]
| ordo = '''Lepidoptera'''
| ordo_authority = [[Carolus Linnaeus|Linnaeus]], [[Systema Naturae|1758]]
}}
The order '''Lepidoptera''' is the second most speciose [[order (biology)|order]] in the [[class (biology)|class]] [[Insect]]a and includes the [[butterfly|butterflies]], [[moth]]s and [[skipper (butterfly)|skipper]]s. Members of the order are referred to as '''lepidopterans'''. A person who collects or studies this order is referred to as a [[Lepidopterist]]. This order has more than 180,000 [[species]]<ref>
{{cite web| url = http://www.ucl.ac.uk/taxome/ | title = The Lepidoptera Taxome Project Draft Proposals and Information
| accessdate = 2007-03-05
| publisher = Centre for Ecology and Evolution, University College London
}}
</ref> in 128 [[Family (biology)|families]] and 47 [[Superfamily|superfamilies]]. It is second only to the Coleoptera (the [[beetle]]s) in number of described species. The name is derived from [[Ancient Greek]] [[wikt:λεπίδος|λεπίδος]] (scale) and [[wikt:πτερόν|πτερόν]] (wing).
==Characteristics of Lepidoptera==
Lepidopterans undergo [[metamorphosis (biology)|complete metamorphosis]] going through a four-stage [[Biological life cycle|life cycle]] of egg - [[larva]] / [[caterpillar]] - [[pupa]]/[[Pupa#Chrysalis|chrysalis]] -[[imago]]/adult.
The larvae have a toughened (sclerotized) head capsule, chewing mouthparts, and a soft body, that may have hair-like or other projections, 3 pairs of true legs, and additional prolegs (up to 5 pairs). They can be confused with the larvae of [[Symphyta|sawflies]]. Lepidopteran larvae can be differentiated by the presence of crochets on the prolegs which are absent in the [[Symphyta]] (sawflies). Most caterpillars are herbivores, but a few are carnivores (some eat ants or other caterpillars) and detritivores.<ref name="dugdale">Dugdale, JS, 1996. Natural history and identification of litter-feeding Lepidoptera larvae (Insecta) in beech forests, Orongorongo Valley, New Zealand, with especial reference to the diet of mice (''Mus musculus''). Journal of The Royal Society of New Zealand, Volume 26, Number 2, pp 251-274</ref>
Adults have two pairs of membranous wings covered, usually completely, by minute [[Scale (zoology)|scales]]. In some species, wings are reduced or absent (often in the female but not the male). Antennae are prominent. In moths, males frequently have more feathery antennae than females, for detecting the female [[pheromone]]s at a distance.
Adult mouth parts, called a proboscis, are adapted for sucking nectar. Some species have reduced mouthparts (some species do not feed as adults), and others have them modified to pierce and suck blood or fruit juices (some Noctuids).<ref name="scoble">Scoble, MJ 1995. The Lepidoptera: form, function and diversity. Oxford, UK: The Oxford University Press; 404 p.</ref>
See also: [[difference between a butterfly and a moth]].
== Families of Lepidoptera ==
[[Image:Lepidoptere(s).jpg|thumb|left|140px|[[Forester Moth]] (Zygaenidae)]]
There are about 130 families in this order with variations depending on the taxonomic treatment (see the family template box at the bottom of this section).
The Lepidoptera are divided into several suborders, the largest being [[Glossata]], the vast majority of which are [[Ditrysia]].
Several other classifications of lepidopteran families are used in older literature. These include the [[Rhopalocera]] (club-horned) consisting of what are commonly called butterflies and the [[Heterocera]] (varied-horned) consisting of the moths. However, Rhophalocera is a natural ([[monophyletic]]) group, while Heterocera is a [[paraphyletic]] assemblage.
Another non-standard classification separates the Lepidoptera into [[Microlepidoptera]] for the smaller species (mostly moths) and [[Macrolepidoptera]] for the larger species.
==Evolution==
===History of study===
Linnaeus in ''Systema Naturae'' (1758) recognized three divisions of the Lepidoptera: ''Papilio, Sphinx,'' and ''[[Phalaena]]'' with seven subgroups in ''Phalaena''.<ref name="scoble"/> These persist today as 9 of the superfamilies of Lepidoptera. Other works on classification followed including those by Denis & [[Ignaz Schiffermüller]] (1775), Fabricius (1775) and [[Pierre André Latreille]] (1796). [[Jacob Hübner]] described many genera, and the Lepidopteran genera were catalogued by [[Ferdinand Ochsenheimer|Ochsenheimer]] and [[Georg Friedrich Treitschke|Treitschke]] in a series of volumes on the Lepidopteran fauna of Europe published between 1807 and 1835.<ref name="scoble"/> G.A.W. Herrich-Schaffer (several volumes, 1843-1856), and [[Edward Meyrick]] (1895) based their classifications primarily on wing venation. Sir [[George Francis Hampson]] worked on the 'microlepidoptera' during this period.
Among the first entomologists to study fossil insects and their evolution was Samuel H Scudder (1837-1911), who worked on butterflies.<ref name="grimaldi">Grimaldi, D, and M S Engel, 2005. Evolution of the Insects. Cambridge University Press.</ref> He published a study of the Florissant deposits of Colorado. Andreas V. Martynov (1879-1938) recognized the close relationship between Lepidoptera and Trichoptera in his studies on phylogeny.<ref name="grimaldi"/>
Major contributions in the 20th century included the creation of the monotrysia and ditrysia (based on female genital structure) by Borner in 1925 and 1939.<ref name="scoble"/> [[Willi Hennig]] (1913-1976) developed the [[cladistic]] methodology and applied it to insect phylogeny. Niels P. Kristensen, E. S. Nielsen and D. R. Davis studied the relationships among [[monotrysia]]n families and Kristensen worked more generally on insect [[phylogeny]] and higher Lepidoptera too.<ref name="scoble"/><ref name="grimaldi"/>. While it is often found that DNA-based phylogenies differ from those based on [[Morphology (biology)|morphology]], this has not been the case for the Lepidoptera; DNA phylogenies correspond to a large extent to morphology-based phylogenies.<ref name="grimaldi"/>
Many attempts have been made to group the superfamilies of the Lepidoptera into natural groups, most of which fail because one of the two groups is not [[monophyly|monophyletic]]: Microlepidotera and Macrolepidoptera, Heterocera and Rhopalocera, Jugatae and Frenatae, Monotrysia and Ditrysia.<ref name="scoble"/>
===Fossils===
Lepidoptera tend not to be as common as some other insects in the habitats that are most conducive to fossilization, such as lakes and ponds, and their juvenile stage has only the head capsule as a hard part that might be preserved. Yet there are fossils, some preserved in amber and some in very fine sediments. [[Leaf miner|Leaf mines]] are also seen in fossil leaves, although the interpretation of them is tricky.<ref name="grimaldi"/>
The earliest fossil is ''Archaeolepis mane'' from the Jurassic, about 190 million years ago in Dorset, UK.<ref name="grimaldi"/> It consists of wings and shows scales with parallel grooves under a scanning electron microscope and the characteristic wing venation pattern shared with [[Trichoptera]].<ref name="grimaldi"/> Only 2 more sets of Jurassic Lepidopteran fossils have been found, and 13 sets in the Cretaceous.<ref name="grimaldi"/> From there, many more fossils are found from the Tertiary, and particularly the Eocene Baltic amber.
===Phylogeny===
{{userboxtop|toptext= }}
{{Clade|style=font-size:75%;line-height:75%
|1={{clade
|label1=Apoditrysia
|1={{clade
|Label1=Obtectomera
|1={{clade
|label1=Macrolepidoptera
|1= {{clade
|label1=Rhopalocera
|1= {{clade
|1=[[Papilionoidea]] (true butterflies)
|2=[[Hesperiidae]] (skippers)
|3=[[Hedylidae]] (American moth-butterflies)
}}
|2= {{clade
|1=[[Geometroidea]] (geometer moths)
|2=[[Drepanoidea]] (hooktip moths)
}}
|3=[[Axiidae (Lepidoptera)|Axiidae]] (gold moths)
|4=[[Callidulidae]] (old world butterfly moths)
|5=[[Noctuoidea]] (Owlet, tiger moths)
|6= {{clade
|1=[[Bombycoidea]] (Silk moths, hawk moths)
|2=[[Lasiocampoidea]] (lappet moths)
|3=[[Mimallonidae]] (sackbearer moths)
}}
}}
|2=[[Thyrididae]] (picture winged leaf moths)
|3=[[Hyblaeidae]] (teak moths)
|4=[[Copromorphoidea]] (fruitworm moths)
|5=[[Pyraloidea]] (snout moths)
|6=[[Immidae]]
|7=[[Whalleyana]]
}}
|2=[[Tortricoidea]] (leafrollers)
|3=[[Zygaenoidea]] (burnet moths)
|4=[[Pterophoridae]] (plume moths)
|5=[[Alucitoidea]] (many-plumed moths)
|6=[[Epermeniidae]] (fringe-tufted moths)
|7=[[Schreckensteinia]] (bristle legged moths)
|8=[[Choreutidae]] (metalmark moths)
|9=[[Urodidae]] (false-burnet moths)
|10={{clade
|1=[[Sesioidea]] (clearwing moths)
|2=[[Cossoidea]] (carpenter moths)
}}
}}
|2=[[Gelechioidea]] (twirler moths)
|3=[[Yponomeutoidea]] (ermine moths)
|4=[[Gracillarioidea]] (leafminers)
|5=[[Tineoidea]] (bagworm moths)
}}
}}
<center><small>A proposed phylogeny of the principal lepidopteran groups.<ref>[http://tolweb.org/Ditrysia/11868 Tree of Life] Accessed January 2007</ref></small></center>
{{userboxbottom}}
It has long been noted that the Lepidoptera and the [[Trichoptera]] (caddisflies) share many similarities that are lacking in other insect orders. Among these are:
* females, rather than males, are [[ZW sex-determination system|heterogametic]] (i.e. their sex chromosomes differ)
* dense [[setae]] on the wings (modified into scales in Lepidoptera)
* a particular wing venation pattern on the forewings
* larvae with mouth structures and glands to make and manipulate silk.<ref name="grimaldi"/>
Thus the two sister orders are grouped into the [[Amphiesmenoptera]]. The group probably evolved in the [[Jurassic]], diverging from the extinct Necrotaulidae.<ref name="grimaldi"/> Lepidoptera differ from the [[Trichoptera]] in several features, including wing venation, form of the scales on the wings, loss of the [[cerci]], loss of an [[ocellus]], and changes to the legs.<ref name="grimaldi"/>
The oldest, most ''basal'' lineages of Lepidoptera have as adults, not the curled tongue or [[proboscis]] characteristic of most members of the order, but chewing [[mandibles]] ([[Micropterigidae]], [[Agathiphagidae]] and [[Heterobathmiidae]]). Micropterigidae larvae feed on decaying leaves (much like the [[Trichoptera]]), [[fungi]], [[liverwort]]s or live leaves.<ref name="scoble"/> The adults chew pollen or spores of ferns. In the Agathiphagidae, larvae feed inside seeds of [[kauri pine]]s, and in Heterobathmiidae the larvae mine leaves of ''[[Nothofagus]]'', the southern beech. These families also have mandibles in the pupal stage, which help the pupa emerge from the seed or cocoon just before adult emergence.<ref name="scoble"/>
The [[Eriocraniidae]] have a short coiled proboscis in the adult stage, and retain mandibles for the purpose of escaping the cocoon, but they are non-functional thereafter.<ref name="scoble"/> They, and most of the other non-ditrysian families, are primarily [[leaf miner]]s in the larval stage. In addition to the proboscis, there is a change in the scales among these basal lineages, with later lineages showing more complex perforated scales.<ref name="grimaldi"/>
With the evolution of the [[Ditrysia]] in the mid-[[Cretaceous]], there was a major reproductive change. The Ditrysia, which comprise 98% of the Lepidoptera, have two separate openings for reproduction in the females (as well as a third opening for excretion), one for mating, and one for laying eggs. The two are linked internally by a seminal duct. (In more basal lineages there is one [[cloaca]], or later, two openings and an external sperm canal.) Of the early lineages of Ditrysia, [[Gracillarioidea]] and [[Gelechioidea]] are mostly leaf miners, but more recent lineages feed externally. In the [[Tineoidea]], most species feed on plant and animal detritus and fungi, and build shelters in the larval stage.<ref name="grimaldi"/>
The [[Yponomeutoidea]] is the first group to have significant numbers of species whose larvae feed on herbaceous plants, as opposed to woody plants.<ref name="grimaldi"/> They evolved about the time that flowering plants underwent an expansive [[adaptive radiation]] in the mid-[[Cretaceous]], and the Gelechioidea that evolved at this time also have great diversity. Whether the processes involved [[co-evolution]] or sequential evolution, the diversity of the Lepidoptera and the angiosperms increased together.
In the so-called "[[macrolepidoptera]]", which constitutes about 60% of Lepidopteran species, there was a general increase in size, better flying ability (via changes in wing shape and linkage of the forewings and hindwings), reduction in the adult mandibles, and a change in the arrangement of the crochets (hooks) on the larval prolegs, perhaps to improve the grip on the host plant.<ref name="grimaldi"/> Many also have [[tympanal organ]]s, that allow them to hear. These organs evolved eight times, at least, because they occur on different body parts and have structural differences.<ref name="grimaldi"/>
The main lineages in the macrolepidoptera are the [[Noctuoidea]], [[Bombycoidea]], [[Lasiocampidae]], [[Mimallonoidea]], [[Geometroidea]] and [[Rhopalocera]]. Bombycoidea plus Lasiocampidae plus Mimallonoidea may be a [[monophyletic]] group.<ref name="grimaldi"/> The Rhopalocera, comprising the [[Papilionoidea]] (Butterflies), [[Hesperioidea]] (skippers), and the [[Hedyloidea]] (moth-butterflies), are the most recently evolved.<ref name="scoble"/> There is quite a good fossil record for this group, with the oldest skipper about 56 million years old.<ref name="grimaldi"/>
<gallery>
Image:Metamorphosis of a Butterfly Merrian 1705.jpg| Painting of butterfly life cycle
Image:Butterfly tongue.jpg| Head of Pyralid moth, showing tongue and compound eye
Image:Lepidoptera wing.jpg| Close up of wing, showing scales
Image:Stathmopoda melanochra (ento-csiro-au).jpg| moth
Image:Monarch Butterfly egg.jpg| Egg of Monarch butterfly ''Danaus plexippus''
Image:Monarch caterpillar and egg.jpg| Larva of Monarch on milkweed
Image:Danaus plexippus chrysalis 2003-10-16.jpg| Chrysalis of Monarch
</gallery>
==See also==
*[[Butterflies]]
*[[Moths]]
*[[Wikipedia:WikiProject Lepidoptera|WikiProject Lepidoptera]]
== External links ==
{{Wikispecies|Lepidoptera}}
{{Wikibookspar|Dichotomous Key|Lepidoptera}}
{{commonscat|Lepidoptera}}
*[http://www.lepidoptera.forumactif.com/ Forum Lepidoptera]
*[http://www.lepidoptera.pl European Butterflies and Moths]
*[http://www.lepidopterology.com/index.htm Lepidopterology.com]
*[http://www.leps.it/ Moths and Butterflies of Europe and North Africa]
*[http://www.ukmoths.force9.co.uk/ British Butterflies and Moths]
*[http://home.hccnet.nl/bernard.fransen/0testsite/0testsite/ Photography of European Butterflies and Moths]
*[http://www.leps.nl/ Butterflies and Moths in the Netherlands]
*[http://www.cirrusimage.com/butterflies_of_North_America.htm Lepidoptera of North America]
*[http://www.goliathus.cz/en/museum-lepidoptera-6.html Lepidoptera] in [http://www.goliathus.cz/en/museum-homepage-0.html online insect museum]
*[http://www.imagenesdeasturias.com/web/panoramicas_graonet/mariposas_de_asturias/index_mariposas.htm Butterflies of Asturias - Spain ]
*[http://www.inra.fr/internet/Produits/PAPILLON/index.htm Lepidoptera of French Antilles]
*[http://pisum.bionet.nsc.ru/kosterin/korgor/index.htm Butterflies of Asian Russia]
*[http://yutaka.it-n.jp/papi.html Butterflies from Indo_China]
*[http://www.mbarnes.force9.co.uk/jamaicamoths/jamaicahome.htm Moths of Jamaica]
*[http://bcrc.bio.umass.edu/kunkel/Moths/ Historic Moth illustrations]
*{{ITIS|ID=117232|taxon=Lepidoptera}}
==Cited references==
<references/>
==Other references==
*Kristensen, NP (Ed.). 1999. Lepidoptera, Moths and Butterflies. Volume 1: Evolution, Systematics, and Biogeography. ''Handbuch der Zoologie. Eine Naturgeschichte der Stämme des Tierreiches / Handbook of Zoology. A Natural History of the phyla of the Animal Kingdom.'' Band / Volume IV Arthropoda: Insecta Teilband / Part 35: 491 pp. Walter de Gruyter, Berlin, New York.
*Nye, IWB & DS Fletcher, 1991. ''Generic Names of Moths of the World.'' Volume 6: xxix + 368 pp. Trustees of the British Museum (Natural History), London.
*''Firefly Encyclopedia of Insects and Spiders'', edited by Christopher O'Toole, ISBN 1-55297-612-2, 2002
{{Lepidoptera}}
[[Category:Insects]]
[[Category:Lepidoptera| ]]
| color = pink
| name = [[Butterfly|Butterflies]], [[moth]]s and [[Skipper (butterfly)|skippers]]
| image = Parthenos_sylvia_philippensis.jpg
| image_width = 250px
| image_caption = The '''Clipper''' ''Parthenos sylvia''
| regnum = [[Animal]]ia
| phylum = [[Arthropod]]a
| classis = [[Insect]]a
| subclassis = [[Pterygota]]
| infraclassis = [[Neoptera]]
| superordo = [[Endopterygota]]
| ordo = '''Lepidoptera'''
| ordo_authority = [[Carolus Linnaeus|Linnaeus]], [[Systema Naturae|1758]]
}}
The order '''Lepidoptera''' is the second most speciose [[order (biology)|order]] in the [[class (biology)|class]] [[Insect]]a and includes the [[butterfly|butterflies]], [[moth]]s and [[skipper (butterfly)|skipper]]s. Members of the order are referred to as '''lepidopterans'''. A person who collects or studies this order is referred to as a [[Lepidopterist]]. This order has more than 180,000 [[species]]<ref>
{{cite web| url = http://www.ucl.ac.uk/taxome/ | title = The Lepidoptera Taxome Project Draft Proposals and Information
| accessdate = 2007-03-05
| publisher = Centre for Ecology and Evolution, University College London
}}
</ref> in 128 [[Family (biology)|families]] and 47 [[Superfamily|superfamilies]]. It is second only to the Coleoptera (the [[beetle]]s) in number of described species. The name is derived from [[Ancient Greek]] [[wikt:λεπίδος|λεπίδος]] (scale) and [[wikt:πτερόν|πτερόν]] (wing).
==Characteristics of Lepidoptera==
Lepidopterans undergo [[metamorphosis (biology)|complete metamorphosis]] going through a four-stage [[Biological life cycle|life cycle]] of egg - [[larva]] / [[caterpillar]] - [[pupa]]/[[Pupa#Chrysalis|chrysalis]] -[[imago]]/adult.
The larvae have a toughened (sclerotized) head capsule, chewing mouthparts, and a soft body, that may have hair-like or other projections, 3 pairs of true legs, and additional prolegs (up to 5 pairs). They can be confused with the larvae of [[Symphyta|sawflies]]. Lepidopteran larvae can be differentiated by the presence of crochets on the prolegs which are absent in the [[Symphyta]] (sawflies). Most caterpillars are herbivores, but a few are carnivores (some eat ants or other caterpillars) and detritivores.<ref name="dugdale">Dugdale, JS, 1996. Natural history and identification of litter-feeding Lepidoptera larvae (Insecta) in beech forests, Orongorongo Valley, New Zealand, with especial reference to the diet of mice (''Mus musculus''). Journal of The Royal Society of New Zealand, Volume 26, Number 2, pp 251-274</ref>
Adults have two pairs of membranous wings covered, usually completely, by minute [[Scale (zoology)|scales]]. In some species, wings are reduced or absent (often in the female but not the male). Antennae are prominent. In moths, males frequently have more feathery antennae than females, for detecting the female [[pheromone]]s at a distance.
Adult mouth parts, called a proboscis, are adapted for sucking nectar. Some species have reduced mouthparts (some species do not feed as adults), and others have them modified to pierce and suck blood or fruit juices (some Noctuids).<ref name="scoble">Scoble, MJ 1995. The Lepidoptera: form, function and diversity. Oxford, UK: The Oxford University Press; 404 p.</ref>
See also: [[difference between a butterfly and a moth]].
== Families of Lepidoptera ==
[[Image:Lepidoptere(s).jpg|thumb|left|140px|[[Forester Moth]] (Zygaenidae)]]
There are about 130 families in this order with variations depending on the taxonomic treatment (see the family template box at the bottom of this section).
The Lepidoptera are divided into several suborders, the largest being [[Glossata]], the vast majority of which are [[Ditrysia]].
Several other classifications of lepidopteran families are used in older literature. These include the [[Rhopalocera]] (club-horned) consisting of what are commonly called butterflies and the [[Heterocera]] (varied-horned) consisting of the moths. However, Rhophalocera is a natural ([[monophyletic]]) group, while Heterocera is a [[paraphyletic]] assemblage.
Another non-standard classification separates the Lepidoptera into [[Microlepidoptera]] for the smaller species (mostly moths) and [[Macrolepidoptera]] for the larger species.
==Evolution==
===History of study===
Linnaeus in ''Systema Naturae'' (1758) recognized three divisions of the Lepidoptera: ''Papilio, Sphinx,'' and ''[[Phalaena]]'' with seven subgroups in ''Phalaena''.<ref name="scoble"/> These persist today as 9 of the superfamilies of Lepidoptera. Other works on classification followed including those by Denis & [[Ignaz Schiffermüller]] (1775), Fabricius (1775) and [[Pierre André Latreille]] (1796). [[Jacob Hübner]] described many genera, and the Lepidopteran genera were catalogued by [[Ferdinand Ochsenheimer|Ochsenheimer]] and [[Georg Friedrich Treitschke|Treitschke]] in a series of volumes on the Lepidopteran fauna of Europe published between 1807 and 1835.<ref name="scoble"/> G.A.W. Herrich-Schaffer (several volumes, 1843-1856), and [[Edward Meyrick]] (1895) based their classifications primarily on wing venation. Sir [[George Francis Hampson]] worked on the 'microlepidoptera' during this period.
Among the first entomologists to study fossil insects and their evolution was Samuel H Scudder (1837-1911), who worked on butterflies.<ref name="grimaldi">Grimaldi, D, and M S Engel, 2005. Evolution of the Insects. Cambridge University Press.</ref> He published a study of the Florissant deposits of Colorado. Andreas V. Martynov (1879-1938) recognized the close relationship between Lepidoptera and Trichoptera in his studies on phylogeny.<ref name="grimaldi"/>
Major contributions in the 20th century included the creation of the monotrysia and ditrysia (based on female genital structure) by Borner in 1925 and 1939.<ref name="scoble"/> [[Willi Hennig]] (1913-1976) developed the [[cladistic]] methodology and applied it to insect phylogeny. Niels P. Kristensen, E. S. Nielsen and D. R. Davis studied the relationships among [[monotrysia]]n families and Kristensen worked more generally on insect [[phylogeny]] and higher Lepidoptera too.<ref name="scoble"/><ref name="grimaldi"/>. While it is often found that DNA-based phylogenies differ from those based on [[Morphology (biology)|morphology]], this has not been the case for the Lepidoptera; DNA phylogenies correspond to a large extent to morphology-based phylogenies.<ref name="grimaldi"/>
Many attempts have been made to group the superfamilies of the Lepidoptera into natural groups, most of which fail because one of the two groups is not [[monophyly|monophyletic]]: Microlepidotera and Macrolepidoptera, Heterocera and Rhopalocera, Jugatae and Frenatae, Monotrysia and Ditrysia.<ref name="scoble"/>
===Fossils===
Lepidoptera tend not to be as common as some other insects in the habitats that are most conducive to fossilization, such as lakes and ponds, and their juvenile stage has only the head capsule as a hard part that might be preserved. Yet there are fossils, some preserved in amber and some in very fine sediments. [[Leaf miner|Leaf mines]] are also seen in fossil leaves, although the interpretation of them is tricky.<ref name="grimaldi"/>
The earliest fossil is ''Archaeolepis mane'' from the Jurassic, about 190 million years ago in Dorset, UK.<ref name="grimaldi"/> It consists of wings and shows scales with parallel grooves under a scanning electron microscope and the characteristic wing venation pattern shared with [[Trichoptera]].<ref name="grimaldi"/> Only 2 more sets of Jurassic Lepidopteran fossils have been found, and 13 sets in the Cretaceous.<ref name="grimaldi"/> From there, many more fossils are found from the Tertiary, and particularly the Eocene Baltic amber.
===Phylogeny===
{{userboxtop|toptext= }}
{{Clade|style=font-size:75%;line-height:75%
|1={{clade
|label1=Apoditrysia
|1={{clade
|Label1=Obtectomera
|1={{clade
|label1=Macrolepidoptera
|1= {{clade
|label1=Rhopalocera
|1= {{clade
|1=[[Papilionoidea]] (true butterflies)
|2=[[Hesperiidae]] (skippers)
|3=[[Hedylidae]] (American moth-butterflies)
}}
|2= {{clade
|1=[[Geometroidea]] (geometer moths)
|2=[[Drepanoidea]] (hooktip moths)
}}
|3=[[Axiidae (Lepidoptera)|Axiidae]] (gold moths)
|4=[[Callidulidae]] (old world butterfly moths)
|5=[[Noctuoidea]] (Owlet, tiger moths)
|6= {{clade
|1=[[Bombycoidea]] (Silk moths, hawk moths)
|2=[[Lasiocampoidea]] (lappet moths)
|3=[[Mimallonidae]] (sackbearer moths)
}}
}}
|2=[[Thyrididae]] (picture winged leaf moths)
|3=[[Hyblaeidae]] (teak moths)
|4=[[Copromorphoidea]] (fruitworm moths)
|5=[[Pyraloidea]] (snout moths)
|6=[[Immidae]]
|7=[[Whalleyana]]
}}
|2=[[Tortricoidea]] (leafrollers)
|3=[[Zygaenoidea]] (burnet moths)
|4=[[Pterophoridae]] (plume moths)
|5=[[Alucitoidea]] (many-plumed moths)
|6=[[Epermeniidae]] (fringe-tufted moths)
|7=[[Schreckensteinia]] (bristle legged moths)
|8=[[Choreutidae]] (metalmark moths)
|9=[[Urodidae]] (false-burnet moths)
|10={{clade
|1=[[Sesioidea]] (clearwing moths)
|2=[[Cossoidea]] (carpenter moths)
}}
}}
|2=[[Gelechioidea]] (twirler moths)
|3=[[Yponomeutoidea]] (ermine moths)
|4=[[Gracillarioidea]] (leafminers)
|5=[[Tineoidea]] (bagworm moths)
}}
}}
<center><small>A proposed phylogeny of the principal lepidopteran groups.<ref>[http://tolweb.org/Ditrysia/11868 Tree of Life] Accessed January 2007</ref></small></center>
{{userboxbottom}}
It has long been noted that the Lepidoptera and the [[Trichoptera]] (caddisflies) share many similarities that are lacking in other insect orders. Among these are:
* females, rather than males, are [[ZW sex-determination system|heterogametic]] (i.e. their sex chromosomes differ)
* dense [[setae]] on the wings (modified into scales in Lepidoptera)
* a particular wing venation pattern on the forewings
* larvae with mouth structures and glands to make and manipulate silk.<ref name="grimaldi"/>
Thus the two sister orders are grouped into the [[Amphiesmenoptera]]. The group probably evolved in the [[Jurassic]], diverging from the extinct Necrotaulidae.<ref name="grimaldi"/> Lepidoptera differ from the [[Trichoptera]] in several features, including wing venation, form of the scales on the wings, loss of the [[cerci]], loss of an [[ocellus]], and changes to the legs.<ref name="grimaldi"/>
The oldest, most ''basal'' lineages of Lepidoptera have as adults, not the curled tongue or [[proboscis]] characteristic of most members of the order, but chewing [[mandibles]] ([[Micropterigidae]], [[Agathiphagidae]] and [[Heterobathmiidae]]). Micropterigidae larvae feed on decaying leaves (much like the [[Trichoptera]]), [[fungi]], [[liverwort]]s or live leaves.<ref name="scoble"/> The adults chew pollen or spores of ferns. In the Agathiphagidae, larvae feed inside seeds of [[kauri pine]]s, and in Heterobathmiidae the larvae mine leaves of ''[[Nothofagus]]'', the southern beech. These families also have mandibles in the pupal stage, which help the pupa emerge from the seed or cocoon just before adult emergence.<ref name="scoble"/>
The [[Eriocraniidae]] have a short coiled proboscis in the adult stage, and retain mandibles for the purpose of escaping the cocoon, but they are non-functional thereafter.<ref name="scoble"/> They, and most of the other non-ditrysian families, are primarily [[leaf miner]]s in the larval stage. In addition to the proboscis, there is a change in the scales among these basal lineages, with later lineages showing more complex perforated scales.<ref name="grimaldi"/>
With the evolution of the [[Ditrysia]] in the mid-[[Cretaceous]], there was a major reproductive change. The Ditrysia, which comprise 98% of the Lepidoptera, have two separate openings for reproduction in the females (as well as a third opening for excretion), one for mating, and one for laying eggs. The two are linked internally by a seminal duct. (In more basal lineages there is one [[cloaca]], or later, two openings and an external sperm canal.) Of the early lineages of Ditrysia, [[Gracillarioidea]] and [[Gelechioidea]] are mostly leaf miners, but more recent lineages feed externally. In the [[Tineoidea]], most species feed on plant and animal detritus and fungi, and build shelters in the larval stage.<ref name="grimaldi"/>
The [[Yponomeutoidea]] is the first group to have significant numbers of species whose larvae feed on herbaceous plants, as opposed to woody plants.<ref name="grimaldi"/> They evolved about the time that flowering plants underwent an expansive [[adaptive radiation]] in the mid-[[Cretaceous]], and the Gelechioidea that evolved at this time also have great diversity. Whether the processes involved [[co-evolution]] or sequential evolution, the diversity of the Lepidoptera and the angiosperms increased together.
In the so-called "[[macrolepidoptera]]", which constitutes about 60% of Lepidopteran species, there was a general increase in size, better flying ability (via changes in wing shape and linkage of the forewings and hindwings), reduction in the adult mandibles, and a change in the arrangement of the crochets (hooks) on the larval prolegs, perhaps to improve the grip on the host plant.<ref name="grimaldi"/> Many also have [[tympanal organ]]s, that allow them to hear. These organs evolved eight times, at least, because they occur on different body parts and have structural differences.<ref name="grimaldi"/>
The main lineages in the macrolepidoptera are the [[Noctuoidea]], [[Bombycoidea]], [[Lasiocampidae]], [[Mimallonoidea]], [[Geometroidea]] and [[Rhopalocera]]. Bombycoidea plus Lasiocampidae plus Mimallonoidea may be a [[monophyletic]] group.<ref name="grimaldi"/> The Rhopalocera, comprising the [[Papilionoidea]] (Butterflies), [[Hesperioidea]] (skippers), and the [[Hedyloidea]] (moth-butterflies), are the most recently evolved.<ref name="scoble"/> There is quite a good fossil record for this group, with the oldest skipper about 56 million years old.<ref name="grimaldi"/>
<gallery>
Image:Metamorphosis of a Butterfly Merrian 1705.jpg| Painting of butterfly life cycle
Image:Butterfly tongue.jpg| Head of Pyralid moth, showing tongue and compound eye
Image:Lepidoptera wing.jpg| Close up of wing, showing scales
Image:Stathmopoda melanochra (ento-csiro-au).jpg| moth
Image:Monarch Butterfly egg.jpg| Egg of Monarch butterfly ''Danaus plexippus''
Image:Monarch caterpillar and egg.jpg| Larva of Monarch on milkweed
Image:Danaus plexippus chrysalis 2003-10-16.jpg| Chrysalis of Monarch
</gallery>
==See also==
*[[Butterflies]]
*[[Moths]]
*[[Wikipedia:WikiProject Lepidoptera|WikiProject Lepidoptera]]
== External links ==
{{Wikispecies|Lepidoptera}}
{{Wikibookspar|Dichotomous Key|Lepidoptera}}
{{commonscat|Lepidoptera}}
*[http://www.lepidoptera.forumactif.com/ Forum Lepidoptera]
*[http://www.lepidoptera.pl European Butterflies and Moths]
*[http://www.lepidopterology.com/index.htm Lepidopterology.com]
*[http://www.leps.it/ Moths and Butterflies of Europe and North Africa]
*[http://www.ukmoths.force9.co.uk/ British Butterflies and Moths]
*[http://home.hccnet.nl/bernard.fransen/0testsite/0testsite/ Photography of European Butterflies and Moths]
*[http://www.leps.nl/ Butterflies and Moths in the Netherlands]
*[http://www.cirrusimage.com/butterflies_of_North_America.htm Lepidoptera of North America]
*[http://www.goliathus.cz/en/museum-lepidoptera-6.html Lepidoptera] in [http://www.goliathus.cz/en/museum-homepage-0.html online insect museum]
*[http://www.imagenesdeasturias.com/web/panoramicas_graonet/mariposas_de_asturias/index_mariposas.htm Butterflies of Asturias - Spain ]
*[http://www.inra.fr/internet/Produits/PAPILLON/index.htm Lepidoptera of French Antilles]
*[http://pisum.bionet.nsc.ru/kosterin/korgor/index.htm Butterflies of Asian Russia]
*[http://yutaka.it-n.jp/papi.html Butterflies from Indo_China]
*[http://www.mbarnes.force9.co.uk/jamaicamoths/jamaicahome.htm Moths of Jamaica]
*[http://bcrc.bio.umass.edu/kunkel/Moths/ Historic Moth illustrations]
*{{ITIS|ID=117232|taxon=Lepidoptera}}
==Cited references==
<references/>
==Other references==
*Kristensen, NP (Ed.). 1999. Lepidoptera, Moths and Butterflies. Volume 1: Evolution, Systematics, and Biogeography. ''Handbuch der Zoologie. Eine Naturgeschichte der Stämme des Tierreiches / Handbook of Zoology. A Natural History of the phyla of the Animal Kingdom.'' Band / Volume IV Arthropoda: Insecta Teilband / Part 35: 491 pp. Walter de Gruyter, Berlin, New York.
*Nye, IWB & DS Fletcher, 1991. ''Generic Names of Moths of the World.'' Volume 6: xxix + 368 pp. Trustees of the British Museum (Natural History), London.
*''Firefly Encyclopedia of Insects and Spiders'', edited by Christopher O'Toole, ISBN 1-55297-612-2, 2002
{{Lepidoptera}}
[[Category:Insects]]
[[Category:Lepidoptera| ]]
