Lepidoptera fossil record

s spans 130–95 mya from their earliest forms to domination of vegetation. Lepidoptera and Trichoptera (caddisflies) are more closely related to one another than to any other taxa, sharing many similarities that are lacking in other insect orders; for example the females of both orders are heterogametic, meaning they have two different sex chromosomes, whereas in most species the males are heterogametic and the females have two identical sex chromosomes. The adults in both orders display a particular wing venation pattern on their forewings. The larvae of both orders have mouth structures and a gland with which they make and manipulate silk. Willi Hennig grouped the two orders into the Amphiesmenoptera superorder; they are sisters, and together are sister to the extinct order Tarachoptera. Micropterigidae, Agathiphagidae and Heterobathmiidae are the oldest and most basal lineages of Lepidoptera. The adults of these families do not have the curled tongue or proboscis, that are found in most members order, but instead have chewing mandibles adapted for a special diet. Micropterigidae larvae feed on leaves, fungi, or liverworts (much like the Trichoptera). Adult Micropterigidae chew the pollen or spores of ferns. In the Agathiphagidae, larvae live inside kauri pines and feed on seeds. In Heterobathmiidae the larvae feed on the leaves of Nothofagus, the southern beech tree. These families also have mandibles in the pupal stage, which help the pupa emerge from the seed or cocoon after metamorphosis. The Eriocraniidae have a short coiled proboscis in the adult stage, and though they retain their pupal mandibles with which they escaped the cocoon, their mandibles are non-functional thereafter. Most of these non-ditrysian families, are primarily leaf miners 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. 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. The Yponomeutoidea is the first group to have significant numbers of species whose larvae feed on herbaceous plants, as opposed to woody plants. 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. Many also have tympanal organs, that allow them to hear. These organs evolved eight times, at least, because they occur on different body parts and have structural differences. 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. The Rhopalocera, comprising the Papilionoidea (butterflies), Hesperioidea (skippers), and the Hedyloidea (moth-butterflies), are the most recently evolved. There is quite a good fossil record for this group, with the oldest skipper dating from . ==Fossil Lepidoptera taxa==

This is a list of all described fossil Lepidoptera species. Taxa marked with are extinct. Superfamily Bombycoidea Family Saturniidae • Rothschildia • Rothschildia fossilis (originally in Attacus) Family Sphingidae • Mioclanis • Mioclanis shanwangiana • Sphingidites • Sphingidites weidneri Superfamily Copromorphoidea Family Copromorphidae • Copromorpha • †Copromorpha fossilis Superfamily Cossoidea Family Cossidae • †Adelopsyche • †Adelopsyche frustrans (Colorado, Florissant) • †Gurnetia • †Gurnetia durranti (Isle of Wight) Superfamily Eolepidopterigoidea Family Eolepidopterigidae • †Daiopterix • †Daiopterix rasnitsyni • †Daiopterix olgae • †Eolepidopterix • Eolepidopterix jurassica • Gracilepteryx • †Gracilepteryx pulchra • †Netoxena • †Netoxena nana • †Psamateia • †Psamateia calipsa • †Undopterix (sometimes in Undopterigidae Kozlov, 1988) • †Undopterix sukatshevae • †Undopterix cariensis Superfamily Eriocranioidea Family Eriocraniidae • †Eriocranites • †Eriocranites hercynicus Superfamily Gelechioidea Family Autostichidae • †Symmocites • †Symmocites rohdendorfi Family Elachistidae • †Elachistites • †Elachistites inclusus (Baltic region, Eocene) • †Elachistites sukatshevae (Baltic region, Eocene) Family Ethmiidae • Ethmia • †Ethmia mortuella (Colorado, Florissant) Family Oecophoridae • †Borkhausenites • †Borkhausenites angustipenella • †Borkhausenites bachofeni • †Borkhausenites crassella • †Borkhausenites implicatella • †Borkhausenites incolumella • †Borkhausenites ingentella • †Borkhausenites vulneratella • †Depressarites • †Depressarites blastuliferella • †Depressarites levipalpella • †Epiborkhausenites (Bartonian, Baltic amber, Lithuania) • †Epiborkhausenites obscurotrimaculatus • †Glesseumeyrickia • †Glesseumeyrickia henrikseni • †Hexerites (originally in Thyrididae) • †Hexerites primalis • †Microsymmocites • †Microsymmocites • †Neoborkhausenites • †Neoborkhausenites incertella (originally in Borkhausenites) • †Palaeodepressaria • †Palaeodepressaria hannemanni • †Paraborkhausenites • †Paraborkhausenites innominatus • †Paraborkhausenites vicinella (originally in Borkhausenites) Family Symmocidae • †Oegoconiites • †Oegoconiites borisjaki (Baltic region, Oligocene amber) Superfamily Geometroidea Family Geometridae '' holotype forewing • †Geometridites • †Geometridites jordani (Willershausen, Pliocene) • †Geometridites larentiiformis • †Geometridites repens • Hydriomena • †Hydriomena? protrita (Priabonian, Florissant Formation, Colorado) Superfamily Gracillarioidea Family Bucculatricidae • Bucculatrix • †Bucculatrix platani (Kazakhstan, Late Cretaceous) Family Gracillariidae • †Gracillariites • †Gracillariites lithuanicus • †Gracillariites mixtus • Two undescribed Phyllocnistis species • One undescribed Lithocolletis species Superfamily Hepialoidea Family Hepialidae • †Oiophassus • †Oiophassus nycterus • †Prohepialus • †Prohepialus incertus (Menat, France, Cenozoic) • †Protohepialus • †Protohepialus comstocki Superfamily Adeloidea Family Adelidae • Adela • †Adela kuznetzovi • †Adela similis • †Adelites • †Adelites electrella • †Adelites purpurascens • †Adelites serraticornella • An undescribed †Adelites species Family Incurvariidae • †Incurvarites • †Incurvarites alienella • †Prophalonia (originally placed in Tortricidae) • †Prophalonia gigas • †Prophalonia scutitarsella Superfamily Micropterigoidea Family Micropterigidae • †Auliepterix • †Auliepterix minima • †Auliepterix mirabilis • †Baltimartyria • †Baltimartyria rasnitsyni • †Baltimartyria proavitella • Micropterix • †Micropterix anglica • †Micropterix gertraudae • †Micropterix immensipalpa (sometimes placed in Eriocraniidae as Electrocrania immensipalpa) • †Moleropterix • †Moleropterix kalbei (sometimes placed in Eolepidopterigidae) • †Palaeolepidopterix • †Palaeolepidopterix aurea • †Palaeosabatinca • †Palaeosabatinca zherichini • †Parasabatinca • †Parasabatinca aftimacrai • †Parasabatinca caldasae • Sabatinca • †Sabatinca perveta Superfamily Nepticuloidea Family Nepticulidae • †Foliofossor • †Foliofossor cranei (Paleocene; England; mines in Platanus sp. leaves) (originally placed in Agromyzidae) • †Stigmellites • †Stigmellites araliae (Czech Republic; mine in Araliaceae sp. leaf) • †Stigmellites baltica (Eocene; Baltic amber; mine) • †Stigmellites caruini-orientalis (Pliocene; Hessen, Germany; mine in Carpinus orientalis fossilis leaf) • †Stigmellites heringi (Pliocene; Hessen, Germany; mine in Berberis sp. leaf) • †Stigmellites kzyldzharica (Kazakhstan; mine in Platanus sp. leaf) • †Stigmellites messelensis (Eocene; Messel, Germany; mine) • †Stigmellites pliotityrella (Pliocene; Hessen, Germany; mine in Fagus silvatica leaf) • †Stigmellites samsonovi (Kazakhstan; mine in Trochodendroides arctica leaf) • †Stigmellites serpentina (Kazakhstan mine in Trochodendroides arctica leaf) • †Stigmellites sharovi (Kazakhstan mine in Trochodendroides arctica leaf) • †Stigmellites tyshchenkoi (Kazakhstan mine in Platanus latior leaf) • †Stigmellites zelkovae (Pliocene; Hessen, Germany; mine in Zelkova sp. leaf) Superfamily Noctuoidea Family Arctiidae • †Oligamatites • †Oligamatites martynovi (Kazakhstan, Upper Oligocene) • †Stauropolia • †Stauropolia nekrutenkoi (Caucasus, Miocene) Family Lymantriidae • One undescribed Euproctis species Family Noctuidae • †Noctuites • †Noctuites haidingeri (Croatia, Radoboj, Cenozoic) • †Xyleutites • †Xyleutites miocenicus (northern Caucasus, Miocene) (originally in Cossidae) Family Notodontidae • †Cerurites • †Cerurites wagneri (Germany, Willershausen, Cenozoic) Superfamily Papilionoidea Basal or incertae sedis • †Lithodryas – Lycaenidae, Nymphalidae? • †Lithodryas styx • †Lithopsyche – Lycaenidae, Riodinidae? • †Lithopsyche antiqua • †Riodinella • †Riodinella nympha (Colorado, Middle Eocene) – Nymphalidae, Pieridae, Riodinidae? Family Hesperiidae • †Pamphilites • †Pamphilites abdita (Aix-en-Provence, Oligocene) • †Thanatites • †Thanatites vetula (Western Germany, Cenozoic) (originally in Nymphalidae) Family Lycaenidae • †Aquisextana • †Aquisextana irenaei (France, Early Oligocene) Family Nymphalidae • †Apanthesis • †Apanthesis leuce (Colorado, Florissant) • †Barbarothea • †Barbarothea florissanti (Colorado, Florissant) • Doxocopa • †Doxocopa wilmattae (Colorado, Florissant) (originally in Chlorippe) • Hestina • Hestina japonica • †Jupitellia • †Jupitellia charon (originally in Jupiteria) • †Lethites • †Lethites reynesii • Undescribed Limenitis species • †Mylothrites • †Mylothrites pluto (Europe, Oligocene) (originally in Vanessa) • †Neorinella • †Neorinella garciae • †Neorinopis • †Neorinopis sepulta (France, Early Oligocene) • †Nymphalites • †Nymphalites obscurum (Colorado, Florissant) • †Nymphalites scudderi • †Nymphalites zeuneri • †Prodryas • †Prodryas persephone • †Prolibythea • †Prolibythea vagabunda (Colorado, Florissant) • Vanessa • †Vanessa amerindica (Colorado, Florissant) Family Papilionidae '' • †Doritites • †Doritites bosniackii (Italy, Tuscany, Miocene) (sometimes in Luehdorfia) • †Praepapilio • †Praepapilio colorado • †Praepapilio gracilis • †Thaites • †Thaites ruminianus (France, Aix-en-Provence, Oligocene) Family Pieridae • †Coliates • †Coliates proserpina • †Oligodonta • †Oligodonta florissantensis (Colorado, Oligocene) • Pontia • †Pontia freyeri • †Stolopsyche • †Stolopsyche libytheoides (Colorado, Cenozoic) Family Riodinidae • Voltinia • †Voltinia dramba Superfamily Pterophoroidea Family Pterophoridae '' • Merrifieldia • †Merrifieldia oligocenicus () (synonym=†Pterophorus oligocenus) Superfamily Pyraloidea Family Pyralidae • †Gallerites • †Gallerites keleri • †Glendotricha • †Glendotricha olgae • †Pyralites • †Pyralites obscures • †Pyralites preecei Superfamily Sesioidea Family Castniidae • †Dominickus • †Dominickus castinodes (Priabonian, Florissant Formation, Colorado) Superfamily Tineoidea Family Psychidae • Dahlica • Dahlica triquetrella (Baltic amber) • †Palaeopsyche • †Palaeopsyche secundum (Baltic amber) • †Palaeopsyche transversum (Baltic amber) • †Psychites • †Psychites pristinella (Baltic region, Cenozoic, amber) • Siederia • Siederia pineti • Sterrhopterix • †Sterrhopteryx pristinella • Taleporia • Taleporia tubulosa • Bacotia • Bacotia claustrella Family Tineidae • †Architinea • †Architinea balticella • †Architinea sepositella • †Dysmasiites • †Dysmasiites carpenteri • †Electromeessia • †Electromeessia zagulijaevi (Baltic region, Eocene amber) • †Glessoscardia • †Glessoscardia gerasimovi • †Martynea • †Martynea rebeli • †Monopibaltia • †Monopibaltia ignitella (Baltic region, Eocene amber) • †Palaeoinfurcitinea • †Palaeoinfurcitinea rohdendorfi (Russia, Eocene amber) • †Palaeoscardiites • †Palaeoscardiites mordvilkoi • †Palaeotinea • †Palaeotinea rasnitsyni • †Paratriaxomasia • †Paratriaxomasia solentensis • †Proscardiites • †Proscardiites martynovi • †Pseudocephitinea • †Pseudocephitinea svetlanae (Russia, Eocene amber) • †Scardiites • †Scardiites meyricki • †Simulotenia • †Simulotenia intermedia • †Tillyardinea • †Tillyardinea eocaenica • Tinea • †Tinea antique • †Tineitella • †Tineitella crystalli (originally in Tineites) • †Tineitella sucinacius (originally in Tineites) • †Tineolamima • †Tineolamima aurella • †Tineosemopsis • †Tineosemopsis decurtatus Superfamily Tortricoidea Family Tortricidae '' '' '' • †Antiquatortia • †Antiquatortia histuroides (Dominican amber) • †Electresia • †Electresia zalesskii (Tanzania, Copal) • †Tortricibaltia • †Tortricibaltia diakonoffi (Baltic amber) • †Tortricidrosis • †Tortricidrosis inclusa (Baltic amber) Superfamily Yponomeutoidea Family Heliodinidae • †Baltonides • †Baltonides roeselliformis (Baltic region, Late Eocene) Family Lyonetiidae • †Prolyonetia (Eocene; Baltic amber, Europe) • †Prolyonetia cockerelli Family Yponomeutidae • †Epinomeuta • †Epinomeuta truncatipennella Superfamily Zygaenoidea Family Zygaenidae '' • Neurosymploca • †Neurosymploca? oligocenica (Lower Stampian, Céreste, Alpes-de-Haute-Provence, France) • Zygaena • †"Zygaena" miocaenica (Germany) • †"Zygaena" turolensis • †Zygaenites • †Zygaenites controversus (Germany, Miocene) Superfamily unassigned Family †Archaeolepidae • †Archaeolepis • †Archaeolepis mane Family †Curvicubitidae • †Curvicubitus • †Curvicubitus triassicus (China, Middle Triassic) Family †Mesokristenseniidae • †Mesokristensenia • †Mesokristensenia angustipenna • †Mesokristensenia latipenna • †Mesokristensenia sinica Superfamily unassigned '' '' '' • †Bombycites • †Bombycites oeningensis (Croatia, Oeningen, described from a pupa) • †Karataunia • †Karataunia lapidaria (Kazakhstan, Upper Jurassic) • †Paleolepidopterites • †Paleolepidopterites destructus (Priabonian, Florissant Formation, Colorado) • †Paleolepidopterites florissantanus (Priabonian, Florissant Formation, Colorado) • †Phylledestes • †Phylledestes vorax (Colorado, Florissant – in the Miocene shales, described from a larva) • †Protolepis • †Protolepis cuprealata (Kazakhstan, Upper Jurassic) • †Spatalistiforma • †Spatalistiforma submerga (Baltic amber) • †Thermojana • †Thermojana sinica (China) (originally placed in Eupterotidae) ==Excluded from Lepidoptera==

Several fossils originally described as lepidopterans have subsequently been assigned to other groups, some as basal Amphiesmenoptera, others into other entirely distinct insect orders. Superorder Amphiesmenoptera Family Eocoronidae • †Eocorona • †Eocorona iani (Queensland, Mid-Triassic) Order Hemiptera Family Palaeontinidae (?) • †Cyllonium • †Cyllonium boidusvalianum • †Cyllonium hewitsonianum Order Mecoptera (?) Family Permochoristidae • †Eoses • †Eoses triassica , disputed to be a synonym of †Mesochorista proavita in the Mecoptera Family Choristopsychidae From the late middle Jurassic (164–165 mya) from the Daohugou fossil beds of Inner Mongolia. • †Choristopsyche tenuinervis • †Choristopsyche perfecta • †Choristopsyche asticta • †Paristopsyche • †Paristopsyche angelineae ==See also==