Lepidoptera fossil record explained

The Lepidoptera fossil record encompasses all butterflies and moths that lived before recorded history. The fossil record for Lepidoptera is lacking in comparison to other winged species, and tending 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 mines are also seen in fossil leaves, although the interpretation of them is tricky.^[1] Putative fossil stem group representatives of Amphiesmenoptera (the clade comprising Trichoptera and Lepidoptera) are known from the Triassic.^[2]

Previously, the earliest known lepidopteran fossils were three wings of Archaeolepis mane, a primitive moth-like species from the Jurassic, about, found in Dorset, UK, which show scales with parallel grooves under a scanning electron microscope and a characteristic wing venation pattern shared with Trichoptera (caddisflies).^{[3] [4]} In 2018, the discovery of exquisite fossilised scales from the Triassic-Jurassic boundary were reported in the journal Science Advances. They were found as rare palynological elements in the sediments of the Triassic-Jurassic boundary from the cored Schandelah-1 well, drilled near Braunschweig in northern Germany. This pushes back the fossil record and origin of glossatan lepidopterans by about 70 million years, supporting molecular estimates of a Norian (c. 212 million years) divergence of glossatan and non-glossatan lepidopterans. The authors of the study proposed that lepidopterans evolved a proboscis as an adaptation to drinkfrom droplets and thin films of water for maintaining fluid balance in the hot and arid climate of the Triassic.^[5]

Only two more sets of Jurassic lepidopteran fossils have been found, as well as 13 sets from the Cretaceous, which all belong to primitive moth-like families.^[1] Many more fossils are found from the Cenozoic, and particularly the Eocene Baltic amber. The oldest genuine butterflies of the superfamily Papilionoidea have been found in the Early Eocene (Ypresian) MoClay or Fur Formation of Denmark. The best preserved fossil lepidopteran is considered to be the Eocene Prodryas persephone from the Florissant Fossil Beds.^{[6] [7]}

Phylogeny

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.^[8]

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).^[9] 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.^[9]

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.^[9] 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.^[1]

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.^[1]

The Yponomeutoidea is the first group to have significant numbers of species whose larvae feed on herbaceous plants, as opposed to woody plants.^[1] 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.^[1] 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.^[1] 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.^[1] The Rhopalocera, comprising the Papilionoidea (butterflies), Hesperioidea (skippers), and the Hedyloidea (moth-butterflies), are the most recently evolved.^[9] There is quite a good fossil record for this group, with the oldest skipper dating from .^[1]

Fossil Lepidoptera taxa

This is a list of all described fossil Lepidoptera species.^{[10] [11] [12] [13] [14] [15]} 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

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

• †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.^[16]

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 ^[17]
  • †Choristopsyche tenuinervis ^[17]
  • †Choristopsyche perfecta ^[17]
  • †Choristopsyche asticta ^[17]
• †Paristopsyche ^[17]
  • †Paristopsyche angelineae ^[17]

See also

• List of extinct butterflies
• Prehistoric insects

External links

• International Palaeoentomological Society
• Palaeoentomology in Russia in both English and Russian
• Tales from the Butterfly Garden: Lepidoptera Curious
• Monophyly of Noctuoidea

Notes and References

1. Book: Grimaldi, D. and Engel, M. S.. Evolution of the Insects . 2005 . . 978-0-521-82149-0.
2. Book: Powell . Jerry A. . Cardé. Ring T.. Vincent H. . Resh . Encyclopedia of Insects . https://books.google.com/books?id=wrMcPwAACAAJ . 14 November 2010 . 2 (illustrated) . 557–587. 2009 . Academic Press . 978-0-12-374144-8 . Lepidoptera.
3. Book: Grimaldi. David A.. Michael S. Engel. Evolution of the insects. 2005. Cambridge University Press. 978-0-521-82149-0. 561.
4. Book: Davies. Hazel. Butler. Carol A.. Do butterflies bite?: fascinating answers to questions about butterflies and moths. 15 July 2011. June 2008. Rutgers University Press. 978-0-8135-4268-3. 48.
5. 10.1126/sciadv.1701568 . 29349295 . 5770165 . A Triassic-Jurassic window into the evolution of Lepidoptera . 10 January 2018 . Timo J. B.. van Eldijk. Torsten. Wappler. Paul K.. Strother. Carolien M. H.. van der Weijst. Hossein. Rajaei. Henk. Visscher. Bas. van de Schootbrugge. Science Advances . 4 . e1701568 . 1. 2018SciA....4.1568V.
6. Book: Meyer. Herbert William. Smith. Dena M.. Paleontology of the Upper Eocene florissant formation, Colorado. 15 July 2011. 2008. Geological Society of America. 978-0-8137-2435-5. 6.
7. Web site: Lepidoptera – latest classification . Unacknowledged . Discoveries in Natural History & Exploration . University of California . 15 July 2011.
8. Wolfram Mey . Wilfried Wichard . Patrick Müller . Bo Wang . 2017 . The blueprint of the Amphiesmenoptera – Tarachoptera, a new order of insects from Burmese amber (Insecta, Amphiesmenoptera) . Fossil Record . 20 . 2 . 129–145 . 10.5194/fr-20-129-2017. free.
9. Book: Scoble, Malcolm J. . The Lepidoptera: Form, Function and Diversity . Oxford University Press. Oxford University . September 1995. 1 . 4–5. 2 . 978-0-19-854952-9 . https://books.google.com/books?id=gnpd_5iNTiwC.
10. Las mariposas fósiles. Razones de su escasez y su influencia sobre el conocimiento de la filogenia y distribución de Zygaenini (Lepidoptera: Zygaenidae). Boletín de la S.E.A. . 26 . 1999. 521–532. Fossil butterflies. Causes of their rarity and how they influence our knowledge of phylogeny and distribution of Zygaenini (Lepidoptera: Zygaenidae). Fidel Fernández-Rubio.
11. Book: Niels P. Kristensen. Handbuch der Zoologie: eine Naturgeschichte der Stämme des Tierreiches. 25 July 2011. Walter de Gruyter. 978-3-11-015704-8. 19–. 1998-12-03.
12. https://web.archive.org/web/20071214214433/http://testing.leptree.net/fossil_information leptree
13. Die Psychidae des baltischen Bernsteins. Nota Lepidopterologica. 32. 1. 13–22. Thomas Sobczyk. Max J. Kobbert. amp. 2009. dead. https://web.archive.org/web/20120320021039/http://www.soceurlep.eu/tl_files/nota/bd32_1/02_Sobczyk.pdf. 20 March 2012. dmy-all.
14. http://www.nhm.ac.uk/jdsml/research-curation/research/projects/butmoth/index.dsml?AUTHORqtype=starts+with&noRecMsg=Sorry+-+no+records+found.++Please+try+again.&sort=GENUS&SUBTRIBEqtype=starts+with&YEARqtype=equals&searchPageURL=index.dsml&GENUSqtype=starts+with&TRIBEqtype=starts+with&GENUS=Pierides&SUBFAMILYqtype=starts+with&FAMILYqtype=starts+with Lepidoptera Genera
15. Web site: Beccaloni . George . etal . February 2005 . Scientific name search . The Global Lepidoptera Names Index . Natural History Museum, London.
16. Sohn . Jae-Cheon . Labandeira . Conrad . Davis . Donald . Mitter . Charles . An annotated catalog of fossil and subfossil Lepidoptera (Insecta: Holometabola) of the world . Zootaxa . 30 April 2012 . 3286 . 1 . 59 . 10.11646/zootaxa.3286.1.1 . free .
17. Qiao X, Shih CK, Petrulevičius JF, Ren Dong R . 2013. Fossils from the Middle Jurassic of China shed light on morphology of Choristopsychidae (Insecta, Mecoptera). ZooKeys. 318. 91–111. 10.3897/zookeys.318.5226. 29 July 2013. 23950679. 3744206. free .