Editing Lepidoptera (section)

=== Phylogeny ===

[[File:Phylogenetic chart of Lepidoptera.svg|right|thumb|upright=1.2|Phylogenetic hypothesis of major lepidopteran lineages superimposed on the geologic time scale. Radiation of [[angiosperm]]s spans {{Ma|130|95}} from their earliest forms to domination of vegetation.]]

Lepidoptera and [[Trichoptera]] (caddisflies) are [[sister group]]s, sharing many similarities that are lacking in others; for example the females of both orders are [[ZW sex-determination system|heterogametic]], meaning they have two different [[gamete|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 in the two orders have mouth structures and glands with which they make and manipulate [[silk]]. [[Willi Hennig]] grouped the two orders into the superorder [[Amphiesmenoptera]]; together they are sister to the extinct order [[Tarachoptera]].<ref name=Tarachoptera>{{Cite journal |author1=Wolfram Mey |author2=Wilfried Wichard |author3=Patrick Müller |author4=Bo Wang |year=2017 |title=The blueprint of the Amphiesmenoptera – Tarachoptera, a new order of insects from Burmese amber (Insecta, Amphiesmenoptera) |journal=Fossil Record |volume=20 |issue=2 |pages=129–145 |doi=10.5194/fr-20-129-2017 |doi-access=free |bibcode=2017FossR..20..129M }}</ref> Lepidoptera descend from a diurnal moth-like common ancestor that either fed on dead or living plants.<ref>{{cite journal|last=Kaila|first=Lauri|author2=Marko Mutanen |author3=Tommi Nyman |title=Phylogeny of the mega-diverse Gelechioidea (Lepidoptera): Adaptations and determinants of success|journal=Molecular Phylogenetics and Evolution|date=27 August 2011|volume=61|pages=801–809|pmid=21903172|doi=10.1016/j.ympev.2011.08.016|issue=3|bibcode=2011MolPE..61..801K }}</ref>

The [[cladogram]], based on molecular analysis, shows the order as a [[clade]], sister to the Trichoptera, and more distantly related to the [[Diptera]] (true flies) and [[Mecoptera]] (scorpionflies).<ref name=Kjer>{{Cite journal |last1=Kjer |first1=Karl M. |last2=Simon |first2=Chris|author2-link=Chris Simon (biologist) |last3=Yavorskaya |first3=Margarita |last4=Beutel |first4=Rolf G. |date=2016 |title=Progress, pitfalls and parallel universes: a history of insect phylogenetics |journal=Journal of the Royal Society Interface |volume=13 |issue=121 |page=121 |doi=10.1098/rsif.2016.0363 |pmid=27558853 |name-list-style=amp |pmc=5014063 }}</ref>

{{clade
|label1=[[Holometabola]]
|1={{clade
   |label1=[[Hymenopterida]]
   |1=[[Hymenoptera]] (sawflies, wasps) [[File:European wasp white bg.jpg|70px]]
   |label2=[[Aparaglossata]]
   |2={{clade
      |label1=[[Neuropteroidea]]      
      |1={{clade
         |label1=[[Coleopterida]]
         |1={{clade
            |1=[[Coleoptera]] (beetles) [[File:Pseudacrossus przewalskyi (Reitter, 1887).jpg|50px]]
            |2=[[Strepsiptera]] (twisted-wing parasites) [[File:Elenchus koebelei.jpg|70px]]
          }}
         |label2=[[Neuropterida]]
         |2={{clade
            |1=[[Raphidioptera]] (snakeflies) [[File:Raphidia icon.png|70px]]
            |label2=
            |2={{clade
               |1=[[Megaloptera]] (alderflies and allies) [[File:Corydalus cornutus illustration (rotated).png|70px]]
               |2=[[Neuroptera]] (Lacewings and allies) [[File:Osmylus_(white_background).jpg|70px]]
             }}
          }}
       }}
      |label2=[[Panorpida]]
      |2={{clade
         |label1=[[Amphiesmenoptera]]
         |1={{clade
            |1='''Lepidoptera''' (butterflies, moths) [[File:Arctia villica SLU.JPG|70px]]
            |2=[[Trichoptera]] (caddisflies) [[File:RHYACOPHILA DORSALIS Male Pont Forge de Sailly Watigny 02 MHNT.jpg|70px]] 
          }}
         |label2=Antliophora
         |2={{clade 
            |1=[[Diptera]] [[File:Common house fly, Musca domestica.jpg|70px]] 
            |2={{clade
               |1=[[Mecoptera]] (scorpionflies)  [[File:Scorpionfly (white background).jpg|70px]]
               |2=[[Siphonaptera]] (fleas) [[File:Pulex irritans female ZSM (white background).jpg|40px]]
             }}
          }}
       }}
    }}
 }}
}}

The internal phylogeny of Lepidoptera is still being resolved. While many large clades have been established, interfamilial and superfamilial relationships are poorly understood. A large scale study by Regier et al. attempts to resolve these relationships using three different analysis methods, which is shown in the following cladogram.<ref>{{cite journal |author1=Regier, Jerome |display-authors=etal |title=A Large-Scale, Higher-Level, Molecular Phylogenetic Study of the Insect Order Lepidoptera (Moths and Butterflies) |journal=PLOS ONE |date=12 March 2013|volume=8 |issue=3 |pages=e58568 |doi=10.1371/journal.pone.0058568 |doi-access=free |pmid=23554903 |pmc=3595289 |bibcode=2013PLoSO...858568R }}</ref>

{{clade
|label1='''Lepidoptera'''
|1={{clade
   |1=[[Micropterigoidea]]
   |2={{clade
      |1=[[Agathiphagoidea]]
      |2={{clade
         |1=[[Heterobathmioidea]]
         |label2=[[Glossata]]
         |2={{clade
            |1=[[Eriocranioidea]]
            |label2=[[Coelolepida]]
            |2={{clade
               |1={{clade
                  |1=[[Lophocoronoidea]]
                  |label2=[[Exoporia]]
                  |2={{clade
                     |1=[[Hepialoidea]]
                     |2=[[Mnesarchaeoidea]]
                     }}
                  }}
               |2={{clade
                  |1=[[Neopseustoidea]] <small>(incl. [[Acanthopteroctetidae]])</small>
                  |label2=[[Heteroneura]]
                  |2={{clade
                     |1=[[Nepticuloidea]]
                     |label2=[[Eulepidoptera]]
                     |2={{clade
                        |1={{clade
                           |1=[[Adeloidea]]
                           |2=[[Andesianoidea]]
                           }}
                        |2={{clade
                           |1={{clade
                              |1=[[Tischerioidea]]
                              |2={{clade
                                 |1=''[[Azaleodes]]''
                                 |2=''[[Ptyssoptera]]''
                                 }}
                              }}
                           |2={{clade
                              |1=''[[Palaephatus (moth)|Palaephatus]]''
                              |label2=[[Ditrysia]]
                              |2={{clade
                                 |1=[[Meessiinae]] <small>(traditionally in Tineidae)</small>
                                 |2={{clade
                                    |1=[[Psychidae]]
                                    |2={{clade
                                       |1=[[Eriocottidae]]
                                       |2={{clade
                                          |1=[[Tineidae]]
                                          |2={{clade
                                             |1={{clade
                                                |1=[[Roeslerstammiidae]]
                                                |2={{clade
                                                   |1=[[Gracillariidae]]
                                                   |2=[[Yponomeutoidea]] <small>(incl. [[Bucculatricidae]])</small>
                                                   }}
                                                }}
                                             |2='''[[Apoditrysia]]''' <small>(cont.)</small>
                                             }}
                                          }}
                                       }}
                                    }}
                                 }}
                              }}
                           }}
                        }}
                     }}
                  }}
               }}
            }}
         }}
      }}
   }}
}}

{{clade
|label1='''Apoditrysia'''
|1={{clade
   |1=[[Urodoidea]]
   |2={{clade
      |1={{clade
         |1=[[Choreutoidea]]
         |2={{clade
            |1=[[Douglasiidae]]
            |2=[[Schreckensteinioidea]]
            }}
         }}
      |2={{clade
         |1={{clade
            |1=[[Immoidea]]
            |2=[[Millieriidae]]
            }}
         |2={{clade
            |1={{clade
               |1={{clade
                  |1=[[Tortricoidea]]
                  |2={{clade
                     |1=[[Galacticoidea]]
                     |2=[[Heliocosma|Heliocosmidae]]
                     }}
                  }}
               |2={{clade
                  |1=[[Zygaenoidea]]
                  |2={{clade
                     |1=[[Cossoidea]] ''sensu lato'' <small>(including traditional [[Sesioidea]])</small>
                     |2=[[Cyclotornidae]] <small>(traditionally in Zygaenoidea)</small>
                     }}
                  }}
               }}
            |label2=[[Obtectomera]]
            |2={{clade
               |1=[[Gelechioidea]]
               |2={{clade
                  |1={{clade
                     |1=[[Pyraloidea]]
                     |label2=[[Macroheterocera]]
                     |2={{clade
                        |1=[[Drepanoidea]] <small>(incl. [[Mimallonoidea]])</small>
                        |2={{clade
                           |1=[[Geometroidea]]
                           |2={{clade
                              |1=[[Noctuoidea]]
                              |2={{clade
                                 |1=[[Bombycoidea]]
                                 |2=[[Lasiocampidae]]
                                 }}
                              }}
                           }}
                        }}
                     }}
                  |2={{clade
                     |1={{clade
                        |1=[[Butterfly|Papilionoidea]]
                        |2=[[Pterophoroidea]]
                        }}
                     |2={{clade
                        |1={{clade
                           |1=''[[Pseudurgis]]''
                           |2=[[Thyridoidea]]
                           }}
                        |2={{clade
                           |1={{clade
                              |1=[[Aluctoidea]]
                              |2={{clade
                                 |1=[[Carposinoidea]]
                                 |2=[[Epermenioidea]]
                                 }}
                              }}
                           |2={{clade
                              |1=[[Calliduloidea]]
                              |2={{clade
                                 |1=[[Hyblaeoidea]]
                                 |2=[[Macropiratidae]]
                                 }}
                              }}
                           }}
                        }}
                     }}
                  }}
               }}
            }}
         }}
      }}
   }}
}}

[[Micropterigidae]], [[Agathiphagidae]] and [[Heterobathmiidae]] are the oldest and most [[basal (phylogenetics)|basal]] lineages of Lepidoptera. The adults of these families do not have the curled tongue or [[proboscis]], that are found in most members of the order, but instead have chewing [[mandible (insect mouthpart)|mandible]]s adapted for a special diet. Micropterigidae larvae feed on [[leaves]], [[fungi]], or [[liverworts]] (much like the [[Trichoptera]]).<ref name="scoble">{{Cite book |last=Scoble |first=Malcolm J. |title=The Lepidoptera: Form, Function and Diversity |publisher=Oxford University Press |location=Oxford University |date=September 1995 |edition=1 |pages=4–5 |chapter=2 |isbn=978-0-19-854952-9 |chapter-url=https://books.google.com/books?id=gnpd_5iNTiwC |access-date=11 September 2020 |archive-date=26 June 2020 |archive-url=https://web.archive.org/web/20200626195110/https://books.google.com/books?id=gnpd_5iNTiwC |url-status=live }}</ref> Adult Micropterigidae chew the pollen or spores of ferns. In the Agathiphagidae, larvae live inside [[kauri pine]]s 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]].<ref name="scoble"/>

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.<ref name="scoble"/> Most of these 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 [[coevolution]] 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 dating from {{Ma|56}}.<ref name="grimaldi"/>