View source for Odonata

== Evolution ==

=== Fossil history ===
[[File:Meganeura.png|thumb|The giant [[Upper Carboniferous]] dragonfly ancestor, ''[[Meganeura]] monyi'', had a wingspan of about {{convert|680|mm|in|abbr=on}},<ref>{{cite book |title=The Biology of Dragonflies |url=https://books.google.com/books?id=J584AAAAIAAJ&pg=PA324 |publisher=Cambridge University Press Archive |date=2018 |page=324 |id=GGKEY:0Z7A1R071DD |quote=No Dragonfly at present existing can compare with the immense ''[[Meganeura monyi]]'' of the [[Upper Carboniferous]], whose expanse of wing was somewhere about twenty-seven inches.}}</ref>]]
Members of the [[crown group]] Odonata first appeared during the [[Late Triassic]],<ref name=palaeo-electronica>{{cite journal |last1=Kohli |first1=Manpreet Kaur |last2=Ware |first2=Jessica L. |last3=Bechly |first3=Günter |year=2016 |title=How to date a dragonfly: Fossil calibrations for odonates |journal=Palaeontologia Electronica |volume=19 |issue=1 |page=576 |doi=10.26879/576 |doi-access=free |bibcode=2016PalEl..19..576K |url=http://palaeo-electronica.org/content/fc-8 }}</ref> though members of their [[Crown group|total group]], [[Odonatoptera]], first appeared in the [[Late Carboniferous]], making them one of the earliest groups of winged insects.<ref name=palaeo-electronica/> The fossils of odonates and their cousins, including [[Paleozoic]] "giant dragonflies" like ''[[Meganeuropsis]] permiana'' from the [[Permian]] of North America, reached wing spans of up to {{convert|71|cm|in|abbr=on}} and a body length of {{convert|43|cm|in|abbr=on}}, making it the largest insect of all time.<ref>{{cite book |last1=Mitchell |first1=F.L. |last2=Lasswell |first2=J. |year=2005 |title=A Dazzle of Dragonflies |publisher=Texas A&M University Press |page=47}}</ref> This insect belonged to the extinct order [[Meganisoptera]], the griffenflies, related to odonates but not part of the modern order Odonata in the restricted sense. They have one of the most complete fossil records going back 319 million years.<ref name=Bybee-CórdbAglr-etal-2016>{{Cite journal |last1=Bybee |first1=Seth |last2=Córdoba-Aguilar |first2=Alex |last3=Duryea |first3=M. Catherine |last4=Futahashi |first4=Ryo |last5=Hansson |first5=Bengt |last6=Lorenzo-Carballa |first6=M. Olalla |last7=Schilder |first7=Ruud |last8=Stoks |first8=Robby |last9=Suvorov |first9=Anton |display-authors=3 |date=December 2016 |title=Odonata (dragonflies and damselflies) as a bridge between ecology and evolutionary genomics |journal=Frontiers in Zoology |volume=13 |issue=1 |page=46 |doi=10.1186/s12983-016-0176-7 |doi-access=free |issn=1742-9994 |pmc=5057408 |pmid=27766110 }}</ref>

The Odonata is closely related to [[mayflies]] and several extinct orders in a group called the [[Palaeoptera]], but this grouping might be [[paraphyletic]]. What they do share with mayflies is the nature of [[insect flight|how the wings are articulated and held in rest]].<ref name="k975">{{cite journal | last1=Ninomiya | first1=Tomoya | last2=Yoshizawa | first2=Kazunori | title=A revised interpretation of the wing base structure in Odonata | journal=Systematic Entomology | volume=34 | issue=2 | date=2009 | issn=0307-6970 | doi=10.1111/j.1365-3113.2008.00455.x | pages=334–345| bibcode=2009SysEn..34..334N | hdl=2115/42988 | hdl-access=free }}</ref>

[[Tarsophlebiidae]] is a prehistoric family of Odonatoptera that can be considered either a basal lineage of Odonata or their immediate [[sister taxon]].<ref name="x206">{{cite journal| last1=Fleck | first1=G. | last2=Bechly | first2=G. | last3=Martinez-Delclòs | first3=X. | last4=Jarzembowski | first4=E.A. | last5=Nel | first5=A. | title= A revision of the Mesozoic dragonfly family Tarsophlebiidae, with a discussion on the phylogenetic positions of the Tarsophlebiidae and Sieblosiidae (Odonatoptera: Panodonata). | journal=Geodiversitas | date=2004-06-01 | url=https://www.researchgate.net/publication/278817520 | access-date=2024-06-04}}</ref>

=== Phylogeny ===

The phylogenetic tree of the orders and suborders of odonates according to Bybee et al. 2021:<ref name="Bybee 2021">{{cite journal | last1=Bybee | first1=Seth M. | last2=Kalkman | first2=Vincent J. | last3=Erickson | first3=Robert J. | last4=Frandsen | first4=Paul B. | last5=Breinholt | first5=Jesse W. | last6=Suvorov | first6=Anton | last7=Dijkstra | first7=Klaas-Douwe B. | last8=Cordero-Rivera | first8=Adolfo | last9=Skevington | first9=Jeffrey H. | last10=Abbott | first10=John C. | last11=Sanchez Herrera | first11=Melissa | last12=Lemmon | first12=Alan R. | last13=Moriarty Lemmon | first13=Emily | last14=Ware | first14=Jessica L. |display-authors=3 | title=Phylogeny and classification of Odonata using targeted genomics | journal=Molecular Phylogenetics and Evolution | publisher=Elsevier BV | volume=160 | year=2021 | issn=1055-7903 | doi=10.1016/j.ympev.2021.107115 | article-number=107115| pmid=33609713 | doi-access=free | bibcode=2021MolPE.16007115B | hdl=11093/2768 | hdl-access=free }}</ref>

{{clade
|style=font-size:100%;line-height:110%
|label1 =Odonata
|1={{clade
   |label1 =
   |1={{clade
   |label1 =
      |1 =[[Zygoptera]] (damselflies) [[File:Megaloprepus coerulatus Drury.png|100px]]
      |2={{clade
         |label1 =[[Epiprocta]]
         |1={{clade
            |label1 =
            |1={{clade
               |label1 =
               |1 =[[Anisozygoptera]]  [[File:Epiophlebia_superstes_f.png|70px]]
               |2 =[[Anisoptera]] (dragonflies) [[File:Illustrations of Exotic Entomology II 47 ('LIBELLULA AXILENA').jpg|100px]]
               }}
            }}
         }}
      }}
   }}
}}

=== Taxonomy ===
In some treatments,{{sfnp|Trueman|Rowe|2008}} the Odonata are understood in an expanded sense, essentially synonymous with the [[superorder]] [[Odonatoptera]], but not including the prehistoric [[Meganisoptera]]. In this approach, instead of Odonatoptera, the term Odonatoidea is used. The [[systematics]] of the "Palaeoptera" are by no means resolved; what can be said however is that regardless of whether they are called "Odonatoidea" or "Odonatoptera", the Odonata and their extinct relatives do form a [[clade]].{{sfnp|Trueman|2008}}

The [[Anisoptera]] was long treated as a suborder, with a third suborder, the Anisozygoptera (ancient dragonflies). However, the combined suborder [[Epiprocta]] (in which Anisoptera is an infraorder) was proposed when it was thought that the "Anisozygoptera" was paraphyletic, composed of mostly extinct offshoots of dragonfly evolution. The four living [[species]] placed in that group are (in this treatment) in the infraorder [[Epiophlebioptera]], whereas the fossil [[taxa]] that were formerly there are now dispersed about the Odonatoptera (or Odonata ''[[sensu lato]]'').{{sfnp|Lohmann|1996}}{{sfnp|Rehn|2003}} World Odonata List considers [[Anisoptera]] as a suborder along with [[Zygoptera]] and [[Anisozygoptera]] as well-understood and widely preferred terms.<ref>{{World Odonata List}}</ref><ref name=Bybee-Kalkman-etal-2021>{{cite journal |last1=Bybee |first1=Seth M. |last2=Kalkman |first2=Vincent J. |last3=Erickson |first3=Robert J. |last4=Frandsen |first4=Paul B. |last5=Breinholt |first5=Jesse W. |last6=Suvorov |first6=Anton |last7=Dijkstra |first7=Klaas-Douwe B. |last8=Cordero-Rivera | first8=Adolfo |last9=Skevington |first9=Jeffrey H. |last10=Abbott |first10=John C. |last11=Sanchez Herrera | first11=Melissa |last12=Lemmon |first12=Alan R. |last13=Moriarty Lemmon | first13=Emily |last14=Ware |first14=Jessica L. |display-authors=6 |year=2021a |title=Phylogeny and classification of Odonata using targeted genomics |journal=Molecular Phylogenetics and Evolution |volume=160 |article-number=107115 |issn=1055-7903 |doi=10.1016/j.ympev.2021.107115 |doi-access=free |pmid=33609713 |bibcode=2021MolPE.16007115B |hdl=11093/2768 | hdl-access=free }}</ref>

Cladogram of Epiprocta after Rehn et al. 2003{{sfnp|Rehn|2003|pages=181–240}}:{{clade|{{clade
|1=[[Zygoptera]] (damselflies)
|label2='''Epiprocta'''
|2={{clade
|1=[[Epiophlebioptera]]
|2={{clade
|1={{extinct}}[[Isophlebioptera]]
|2={{clade
|1={{extinct}}[[Heterophlebioptera]]
|2={{clade
|1={{extinct}}[[Stenophlebioptera]]
|2={{clade
|1={{extinct}}[[Aeschnidioptera]]
|2=[[Anisoptera]] (dragonflies)
}}}}}}}}}}}}|label1=Odonata}}Cladogram of [[Odonatoptera]] including Odonata by Deregnaucourt et al. 2023.<ref>{{Cite journal |last1=Deregnaucourt |first1=Isabelle |last2=Bardin |first2=Jérémie |last3=Villier |first3=Loïc |last4=Julliard |first4=Romain |last5=Béthoux |first5=Olivier |date=August 2023 |title=Disparification and extinction trade-offs shaped the evolution of Permian to Jurassic Odonata |journal=iScience |language=en |volume=26 |issue=8 |article-number=107420 |doi=10.1016/j.isci.2023.107420 |pmc=10424082 |pmid=37583549|bibcode=2023iSci...26j7420D }}</ref>

{{clade|{{clade
 |label1=†[[Meganisoptera]]
 |1=†[[Meganeuridae]]
 |label2=Nodialata
 |2={{clade
  |label1=†[[Protanisoptera]]
  |1={{clade
   |1=†[[Polytaxineuridae]]
   |2={{clade
    |1=†[[Permaeschnidae]]
    |2={{clade
     |1=†[[Callimokaltaniidae]]
     |2={{clade
      |1=†[[Hemizygopteridae]]
      |2=†[[Ditaxineuridae]] }} }} }} }}
  |label2=Discoidalia
  |2={{clade
   |label1=†[[Triadophlebiomorpha]]
   |1={{clade
    |1=†''[[Iverya]]''
    |2={{clade
     |1=†[[Pirouteliidae]]
     |2={{clade
      |1=†[[Kargalotypidae]]
      |label2=†[[Zygophlebiida]]
      |2={{clade
       |1=†[[Zygophlebiidae]]
       |2=†[[Moltenophlebiidae]]
       |3={{clade
        |1=†[[Permophlebiidae]]
        |2=†[[Xamenophlebiidae]] }} }}
      |label3=†[[Triadophlebiida]]
      |3={{clade
       |1=†[[Triassologus]]
       |2=†[[Sinotriadophlebiidae]]
       |3=†[[Mitophlebiidae]]
       |4={{clade
        |1=†[[Paurophlebiidae]]
        |2=†[[Triadophlebiidae]] }} }} }} }} }}
   |label2=Stigmoptera
   |2={{clade
    |1=†[[Permagrionidae]]
    |2={{clade
     |label1=†[[Archizygoptera]]
     |1={{clade
      |1=†[[Bakteniidae]]
      |2=†[[Kennedyidae]]
      |3=†[[Oboraneuridae]]
      |4=†[[Kaltanoneuridae]]
      |5=†[[Lodeviidae]]
      |6=†[[Luiseiidae]]
      |7=†[[Permepallagidae]]
      |8={{clade
       |1=†[[Voltzialestidae]]
       |label2=†[[Terskejoptera]]
       |2={{clade
        |1=†''[[Terskeja]]''
        |2={{clade
         |1=†[[Protomyrmeleontidae]]
         |2=†[[Moltenagrionidae]] }} }} }} }}
     |label2=Panodonata
     |2={{clade
      |1=†[[Triassolestidae]] (in part)
      |label2=Lobodonata
      |2={{clade
       |1=†[[Mesophlebiidae]]
       |2=†[[Triassolestidae]] (in part)
       |3={{clade
        |1=†[[Tarsophlebiidae]]
|label2='''Odonata''' crown group
        |2={{clade
         |1=[[Zygoptera]] (damselflies)
         |label2=[[Epiproctophora]]
         |2={{clade
          |1={{clade
           |1=†[[Euthemistidae]]
           |2=†[[Selenothemistidae]]
           |3={{clade
            |1=†[[Archithemistidae]]
            |2={{clade
             |1=†[[Campterophlebiidae]]
             |2=†[[Isophlebiidae]] }} }}
           |4={{clade
            |1=†[[Sphenophlebiidae]]
            |2={{clade
             |1=†[[Asiopteridae]]
             |2=†[[Cyclothemistidae]] }} }} }}
          |2={{clade
           |1=†[[Erichschmidtiidae]]
           |2={{clade
            |1=†[[Liassostenophlebiidae]]
            |2={{clade
             |1=†[[Prostenophlebiidae]]
             |2=†[[Stenophlebiidae]] }} }}
           |3={{clade
            |1=†[[Myopophlebiidae]]
            |2={{clade
             |1=†[[Heterophlebiidae]]
             |2={{clade
              |1=†[[Liassophlebiidae]]
              |2=†[[Parastenophlebiidae]] }} }} }}
           |4={{clade
            |1=†[[Juragomphidae]]
            |2=†[[Liassogomphidae]]
            |3={{clade
             |1=†[[Aeschnidiidae]]
             |2=[[Anisoptera]] (modern dragonflies) }} }} }} }} }} }} }} }} }} }} }} }} }}|style=font-size:85%; line-height:85%;|label1=[[Odonatoptera]] (Odonata ''sensu lato'')}}