Editing Coelacanth (section)

== Evolution and taxonomy ==
[[File:Libys sp.jpg|thumb|Specimen of ''[[Libys]]'' (Latimeriidae) from the Upper Jurassic of Germany]]
[[File:Mawsonia scaling.png|thumb|320x320px|Estimated size of the largest known individual of the Jurassic-Cretaceous freshwater coelacanth ''[[Mawsonia (fish)|Mawsonia]]'' compared to a human]]
[[File:Allenypterus montanus (Restoration).jpg|thumb|Life restoration of the basal coelacanth ''[[Allenypterus]]'' from the Carboniferous of North America]]
Coelacanths are members of the class Actinistia, with many researchers considering the term "coelacanth" to cover all members of Actinistia.<ref name=":0">{{Cite journal |last1=Toriño |first1=Pablo |last2=Soto |first2=Matías |last3=Perea |first3=Daniel |date=2021-02-25 |title=A comprehensive phylogenetic analysis of coelacanth fishes (Sarcopterygii, Actinistia) with comments on the composition of the Mawsoniidae and Latimeriidae: evaluating old and new methodological challenges and constraints |url=https://www.researchgate.net/publication/349631865 |journal=Historical Biology |volume=33 |issue=12 |pages=3423–3443 |bibcode=2021HBio...33.3423T |doi=10.1080/08912963.2020.1867982 |issn=0891-2963 |s2cid=233942585}}</ref><ref name=":1" /> The order Coelacanthiformes has been used for a subgroup of actinistians, containing the modern coelacanths.<ref>{{Cite journal |last=Arratia |first=Gloria |last2=Schultze |first2=Hans-Peter |date=2015-09-03 |title=A new fossil actinistian from the Early Jurassic of Chile and its bearing on the phylogeny of Actinistia |url=http://www.tandfonline.com/doi/full/10.1080/02724634.2015.983524 |journal=Journal of Vertebrate Paleontology |language=en |volume=35 |issue=5 |pages=e983524 |doi=10.1080/02724634.2015.983524 |issn=0272-4634}}</ref> According to the fossil record, the divergence of coelacanths, [[lungfish]], and [[tetrapods]] is thought to have occurred during the [[Silurian]].<ref>{{Cite journal|last1=Lu|first1=Jing|last2=Giles|first2=Sam|last3=Friedman|first3=Matt|last4=Zhu|first4=Min|date=2017-12-05|title=A new stem sarcopterygian illuminates patterns of character evolution in early bony fishes|journal=Nature Communications|language=en|volume=8|issue=1|pages=1932|doi=10.1038/s41467-017-01801-z|pmid=29203766|issn=2041-1723|pmc=5715141|bibcode=2017NatCo...8.1932L}}</ref> Over 100 fossil species of coelacanth have been described.<ref name=":0" /> The oldest identified coelacanth fossils are around 420–410 million years old, dating to the [[Pragian]] stage of the early [[Devonian]]. These include ''[[Eoactinistia]]'' from Australia, known only from a fragmentary jaw, as well as ''[[Euporosteus|Euporosteus yunnanensis]]'' from China, known from a partial skull that indicates it to be the earliest anatomically modern coelacanth.<ref name="ref4" /><ref name=":1">{{Cite journal |last=Zhu |first=Min |last2=Yu |first2=Xiaobo |last3=Lu |first3=Jing |last4=Qiao |first4=Tuo |last5=Zhao |first5=Wenjin |last6=Jia |first6=Liantao |date=2012-04-10 |title=Earliest known coelacanth skull extends the range of anatomically modern coelacanths to the Early Devonian |url=https://www.nature.com/articles/ncomms1764 |journal=Nature Communications |language=en |volume=3 |issue=1 |pages=772 |doi=10.1038/ncomms1764 |issn=2041-1723}}</ref> Some authors have also suggested that the slightly older [[Onychodontiformes|onychodont]] ''[[Styloichthys]]'' may also be an early coelacanth.<ref>{{cite journal|last=Friedman|first=Matt|date=10 August 2007|title=Styloichthys as the oldest coelacanth: implications for early osteichthyan interrelationships |url=https://www.tandfonline.com/doi/abs/10.1017/S1477201907002052 |journal=Journal of Systematic Palaeontology|volume=5|issue=3|pages=289–343|doi=10.1017/S1477201907002052|bibcode=2007JSPal...5..289F |s2cid=83712134|access-date=2007-12-28}}</ref> 

Coelacanths were never a diverse group in comparison to other groups of fish, and reached a peak diversity during the [[Early Triassic]] (252–247 million years ago),<ref name="Cavin&al.2020" /> coinciding with a burst of diversification between the Late Permian and Middle Triassic.<ref name=":0" /> Most [[Mesozoic]] coelacanths belong to the suborder Latimerioidei, which contains two major subdivisions, the marine [[Latimeriidae]], which contains modern coelacanths, as well as the extinct [[Mawsoniidae]], which were native to [[Brackish water|brackish]], freshwater as well as marine environments.<ref>{{Cite journal|last1=Cavin|first1=Lionel|last2=Cupello|first2=Camila|last3=Yabumoto|first3=Yoshitaka|last4=Léo|first4=Fragoso|last5=Deersi|first5=Uthumporn|last6=Brito|first6=Paul M.|date=2019|title=Phylogeny and evolutionary history of mawsoniid coelacanths|url=http://www.kmnh.jp/wp-content/uploads/2019/05/A17-3-Cavin.pdf|journal=Bulletin of the Kitakyushu Museum of Natural History and Human History, Series A|volume=17|pages=3–13}}</ref>

Paleozoic coelacanths are generally small (~{{Cvt|30–40|cm|disp=or}} in length), while Mesozoic forms were larger.<ref name=":0" /> Several specimens belonging to the Jurassic and Cretaceous mawsoniid coelacanth genera ''[[Trachymetopon]]'' and ''[[Mawsonia (fish)|Mawsonia]]'' likely reached or exceeded {{Convert|5|m|ft|abbr=off}} in length, making them amongst the largest known fishes of the Mesozoic, and amongst the largest bony fishes of all time.<ref>{{Cite journal|last1=Cavin|first1=Lionel|last2=Piuz|first2=André|last3=Ferrante|first3=Christophe|last4=Guinot|first4=Guillaume|date=2021-06-03|title=Giant Mesozoic coelacanths (Osteichthyes, Actinistia) reveal high body size disparity decoupled from taxic diversity|journal=Scientific Reports|language=en|volume=11|issue=1|pages=11812|doi=10.1038/s41598-021-90962-5|pmid=34083600|issn=2045-2322|pmc=8175595|bibcode=2021NatSR..1111812C}}</ref>

The most recent fossil latimeriid is ''[[Megalocoelacanthus|Megalocoelacanthus dobiei]]'', whose disarticulated remains are found in late [[Santonian]] to middle [[Campanian]], and possibly earliest [[Maastrichtian]]-aged marine strata of the Eastern and Central United States,<ref name="Schwimmer&al.1994">{{cite journal|last1=Schwimmer|first1=D.R.|author2=Stewart, J.D.|author3=Williams, G.D.|title=Giant fossil coelacanths of the Late Cretaceous in the eastern United States|journal=Geology|date=1994|volume=2|issue=6|pages=503–506|doi=10.1130/0091-7613(1994)022<0503:GFCOTL>2.3.CO;2|bibcode=1994Geo....22..503S}}</ref><ref name="gottfried">{{cite journal|author1=Gottfried, Michael D. |author2=Rogers, Raymond R. |author3=Rogers, K. Curry   |title=First record of Late Cretaceous coelacanths from Madagascar|url=https://www.researchgate.net/publication/265667981|journal= Recent Advances in the Origin and Early Radiation of Vertebrates |year=2004|pages= 687–691}}</ref><ref name="Dutel&al.2012">{{cite journal|last1=Dutel|first1=H.|author2=Maisey, J.P.|author3=Schwimmer, D.R.|author4=Janvier, P.|author5=Herbin, M.|author6=Clément, G.|title= The Giant Cretaceous Coelacanth (Actinistia, Sarcopterygii) Megalocoelacanthus dobiei Schwimmer, Stewart & Williams, 1994, and Its Bearing on Latimerioidei Interrelationships|journal=PLOS ONE|year=2012|volume= 7|issue= 11|pages= e49911|doi=10.1371/journal.pone.0049911|pmid= 23209614|pmc= 3507921|bibcode=2012PLoSO...749911D|doi-access=free}}</ref> the most recent mawsoniids are ''[[Axelrodichthys|Axelrodichthys megadromos]]'' from early Campanian to early Maastrichtian freshwater continental deposits of France,<ref name="Cavin&al.2005">{{cite journal|last1=Cavin|first1=L.|author2=Forey, P.L.|author3=Tong, H.|author4=Buffetaut, E.|title=Latest European coelacanth shows Gondwanan affinities|journal=Biology Letters|date=2005|volume=1|issue=2|pages=176–177|doi=10.1098/rsbl.2004.0287|pmid=17148159|pmc=1626220}}</ref><ref name="Cavin&al.2016">{{cite journal|last1=Cavin|first1=L.|author2=Valentin, X.|author3=Garcia, G.|title=A new mawsoniid coelacanth (Actinistia) from the Upper Cretaceous of Southern France|journal=Cretaceous Research|date=2016|volume=62|pages=65–73|doi=10.1016/j.cretres.2016.02.002|bibcode=2016CrRes..62...65C }}</ref><ref name="Cavin&al.2020">{{cite journal|last1=Cavin|first1=L.|author2=Buffetaut, E.|author3=Dutour, Y.|author4=Garcia, G.|author5=Le Loeuff, J.|author6=Méchin, A.|author7=Méchin, P.|author8=Tong, H.|author9=Tortosa, T.|author10=Turini, E.|author11=Valentin, X.|title=The last known freshwater coelacanths: New Late Cretaceous mawsoniids remains (Osteichthyes: Actinistia) from Southern France|journal=PLOS ONE|date=2020|volume=15|issue=6|pages=e0234183|doi=10.1371/journal.pone.0234183|pmc=7274394|pmid=32502171|bibcode=2020PLoSO..1534183C|doi-access=free}}</ref> as well as an indeterminate marine mawsoniid from Morocco, dating to the late Maastrichtian<ref name="Brito&al.2021">{{cite journal|last1=Brito|first1=P.M.|author2=Martill, D.M.|author3=Eaves, I.|author4=Smith, R.E.|author5=Cooper, S.L.A.|year=2021|title=A marine Late Cretaceous (Maastrichtian) coelacanth from North Africa|journal=Cretaceous Research|volume=122|page=104768|doi=10.1016/j.cretres.2021.104768|bibcode=2021CrRes.12204768B |s2cid=233551515|url=https://researchportal.port.ac.uk/portal/en/publications/a-marine-late-cretaceous-maastrichtian-coelacanth-from-north-africa(c8521ebb-5597-4a15-9320-961113ee06d9).html }}</ref> A small bone fragment from the [[Europe]]an [[Paleocene]] has been considered the only plausible post-Cretaceous record, but this identification is based on comparative bone histology methods of doubtful reliability.<ref name="Schwimmer&al.1994" /><ref>{{cite journal |last1=Ørvig |first1=Tor |title=A vertebrate bone from the Swedish Paleocene |journal=Geologiska Föreningen i Stockholm Förhandlingar |date=1 June 1986 |volume=108 |issue=2 |pages=139–141 |doi=10.1080/11035898609452636 |issn=0016-786X}}</ref>

Living coelacanths have been considered "living fossils" based on their supposedly conservative [[morphology (biology)|morphology]] relative to fossil species;<ref name="ref10">{{cite journal |last=Butler |first=Carolyn |title=Living Fossil Fish |journal=National Geographic |date=March 2011 |pages=86–93}}</ref><ref name="ref1" />{{rp|1}} however, recent studies have expressed the view that coelacanth morphologic conservatism is a belief not based on data.<ref name="Friedman2007" /><ref name="Friedman2006" /><ref name="Rebellatrix" /><ref name="Casane">{{cite journal |doi=10.1002/bies.201200145 |title=Why coelacanths are not 'living fossils' |year=2013 |last1=Casane |first1=Didier |last2=Laurenti |first2=Patrick |journal=BioEssays |volume=35 |issue=4 |pages=332–8 |pmid=23382020|s2cid=2751255 |doi-access=free }}</ref> Fossils suggest that coelacanths were most morphologically diverse during the Devonian and Carboniferous, while Mesozoic species are generally morphologically similar to each other.<ref name=":0" />

[[Cladogram]] showing the relationships of coelacanth genera after Torino, Soto and Perea, 2021.<ref name=":0" />

{{clade |style=font-size:90%; line-height:90%;
|1=''[[Mimipiscis]]'' ([[Actinopterygii]])
|2=''[[Porolepis]]'' ([[Porolepiformes]])
|label3='''Actinistia'''
|3={{clade
|1=''[[Miguashaia]]''
|2=''[[Styloichthys]]''
|3=''[[Gavinia]]''
|4={{clade
|1=''[[Diplocercides]]''
|2={{clade
|1=''[[Serenichthys]]''
|2={{clade
|1=''[[Holopterygius]]''
|2=''[[Allenypterus]]''}} 
|3={{clade
|1=''[[Lochmocercus]]''
|2={{clade
|1={{clade
|1=''[[Polyosteorhynchus]]''
|2={{clade 
|1=''[[Rebellatrix]]''
|2=''[[Hadronector]]''}} }}
|2={{clade
|1=''[[Rhabdoderma]]''
|2={{clade
|1=''[[Caridosuctor]]''
|2={{clade
|1={{clade
|1=''[[Sassenia]]''
|2=''[[Spermatodus]]''}}
|2={{clade
|1={{clade
|1=''[[Piveteauia]]''
|2={{clade
|1=''[[Coccoderma]]''
|2=''[[Laugia]]''}} }}
|2={{clade
|1=''[[Coelacanthus]]''
|2={{clade
|1=''[[Guizhoucoelacanthus]]''
|2={{clade
|1={{clade
|1=''[[Wimania]]''
|2=''[[Axelia]]''}} 
|2={{clade
|1=''[[Whiteia]]''
|2={{clade
|1=''[[Heptanema]]''
|2=''[[Dobrogeria]]''
|label3=[[Latimerioidei]]
|3={LATIMERIOIDEA}
}} }} }} }} }} }} }} }} }} }} }} }} }} }}
|targetA={LATIMERIOIDEA}
|subcladeA={{clade
   |label1=[[Mawsoniidae]]
   |1={{clade
      |1={{clade
         |1=''[[Atacamaia]]''
         |2=''[[Luopingcoelacanthus]]''
         }}
      |2={{clade
         |1=''[[Yunnancoelacanthus]]''
         |2={{clade
            |1=''[[Chinlea]]''
            |2={{clade
               |1=''[[Parnaibaia]]''
               |2={{clade
                  |1=''[[Trachymetopon]]''
                  |2=''[[Lualabaea]]''
                  |3={{clade
                     |1=''[[Axelrodichthys]]''
                     |2=''[[Mawsonia (fish)|Mawsonia]]''
                     }}
                  }} 
               }} 
            }} 
         }} 
      }}
   |label2=[[Latimeriidae]]
   |2={{clade
      |1=''[[Garnbergia]]''
      |2={{clade
         |1=''[[Diplurus]]''
         |2={{clade
            |1={{clade
               |1=''[[Megalocoelacanthus]]''
               |2=''[[Libys]]''
               }} 
            |2={{clade
               |1={{clade
                  |1=''[[Ticinepomis]]''
                  |2=''[[Foreyia]]''
                  }}
               |2={{clade
                  |1=''[[Holophagus]]''
                  |2={{clade
                     |1=''[[Undina (fish)|Undina]]''
                     |2={{clade
                        |1=''[[Macropoma]]''
                        |2={{clade
                           |1=''[[Swenzia]]''
                           |2=''[[Latimeria]]''
                           }} 
                        }} 
                     }} 
                  }} 
               }} 
            }} 
         }} 
      }} 
   }} 
}}

=== Timeline of genera ===
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 from: -488.3    till: -443.7    color:ordovician    text:[[Ordovician]]
 from: -443.7    till: -416    color:silurian    text:[[Silurian]]
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 from: -359.2    till: -299   color:carboniferous    text:[[Carboniferous]]
 from: -299    till: -251   color:permian    text:[[Permian]]
 from: -251    till: -199.6  color:triassic    text:[[Triassic]]
 from: -199.6    till: -145.5   color:jurassic    text:[[Jurassic]]
 from: -145.5   till: -66   color:cretaceous    text:[[Cretaceous]]
 from: -66    till: -23.03   color:paleogene    text:[[Paleogene]]
 from: -23.03    till: -2.588   color:neogene    text:[[Neogene|Neog.]]
 from: -2.588    till: 0   color:quaternary    text:[[Quaternary|Q.]]

 bar:eratop
 from:  -542   till:  -251    color:paleozoic    text:[[Paleozoic Era]]
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 from:  -66   till:  0   color:cenozoic    text:[[Cenozoic]]

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 align:left fontsize:M mark:(line,white) width:5 anchor:till align:left

 color:devonian bar:NAM3 from:-412.8    till:-411.2 text:[[Powichthys]]
 color:devonian bar:NAM4 from:-408.4    till:-407 text:[[Youngolepis]]
 color:devonian bar:NAM6 from:-407    till:-359.2 text:[[Onychodus]]
 color:devonian bar:NAM13 from:-397.5    till:-359.2 text:[[Strunius]]
 color:devonian bar:NAM15 from:-391.8    till:-374.5 text:[[Euporosteus]]
 color:devonian bar:NAM20 from:-385.3    till:-381.7 text:[[Miguashaia]]
 color:devonian bar:NAM21 from:-385.3    till:-374.5 text:[[Nesides]]
 color:devonian bar:NAM22 from:-381.7    till:-328.3 text:[[Diplocercides]]
 color:devonian bar:NAM26 from:-364.3    till:-359.2 text:[[Chagrinia]]
 color:carboniferous bar:NAM29 from:-345.3    till:-299 text:[[Rhabdoderma]]
 color:carboniferous bar:NAM30 from:-333.97    till:-328.3 text:[[Coelacanthopsis]]
 color:carboniferous bar:NAM31 from:-328.3    till:-324.9 text:[[Allenypterus]]
 color:carboniferous bar:NAM32 from:-328.3    till:-324.9 text:[[Caridosuctor]]
 color:carboniferous bar:NAM33 from:-328.3    till:-324.9 text:[[Hadronector]]
 color:carboniferous bar:NAM34 from:-328.3    till:-324.9 text:[[Lochmocereus]]
 color:carboniferous bar:NAM35 from:-328.3    till:-324.9 text:[[Polyosteorhynchus]]
 color:carboniferous bar:NAM36 from:-306.5    till:-301.37 text:[[Synaptotylus]]
 color:permian bar:NAM37 from: -299    till:-294.6 text:[[Ectosteorhachis]]
 color:permian bar:NAM38 from:-260    till:-251 text:[[Changxingia]]
 color:permian bar:NAM39 from:-258.0    till:-252 text:[[Coelacanthus]]
 color:permian bar:NAM40 from:-255    till:-251 text:[[Yonngichthys]]
 color:triassic bar:NAM41 from:-251    till:-250.57 text:[[Laugia]]
 color:triassic bar:NAM42 from:-251    till:-248.13 text:[[Sassenia]]
 color:triassic bar:NAM43 from:-251    till:-245 text:[[Sinocoelacanthus]]
 color:triassic bar:NAM44 from:-250.13    till:-249.7 text:[[Piveteauia]]
 color:triassic bar:NAM45 from:-250.13    till:-249.7 text:[[Whiteia]]
 color:triassic bar:NAM46 from:-249.7    till:-248.13 text:[[Axelia]]
 color:triassic bar:NAM47 from:-249.7    till:-248.13 text:[[Mylacanthus]]
 color:triassic bar:NAM48 from:-249.7    till:-248.13 text:[[Scleracanthus]]
 color:triassic bar:NAM49 from:-249.7    till:-248.13 text:[[Wimania]]
 color:triassic bar:NAM50 from:-245    till:-237 text:[[Garnbergia]]
 color:triassic bar:NAM51 from:-242    till:-237 text:[[Heptanema]]
 color:triassic bar:NAM52 from:-237    till:-234 text:[[Alcoveria]]
 color:triassic bar:NAM53 from:-237    till:-234 text:[[Ticinepomis]]
 color:triassic bar:NAM54 from:-237    till:-228 text:[[Hainbergia]]
 color:triassic bar:NAM55 from:-220.3    till:-216.5 text:[[Graphiuricthys]]
 color:jurassic bar:NAM56 from:-199.6    till:-191 text:[[Holophagus]]
 color:jurassic bar:NAM57 from:-199    till:-163.5 text:[[Trachymetopon]]
 color:jurassic bar:NAM57.1 from:-163.5  till:-157.3 text:[[Swenzia]]
 color:jurassic bar:NAM58 from:-161.2    till:-145.5 text:[[Lualabaea]]
 color:jurassic bar:NAM59 from:-152    till:-145 text:[[Undina]]
 color:jurassic bar:NAM60 from:-150.8    till:-149.03 text:[[Coccoderma]]
 color:jurassic bar:NAM60.1 from:-150.8    till:-149.03 text:[[Libys]]
 color:cretaceous bar:NAM60.2 from:-150   till:-95 text:[[Mawsonia (fish)|Mawsonia]]
 color:cretaceous bar:NAM60.3 from:-112    till:-71.5 text:[[Axelrodichthys]]
 color:cretaceous bar:NAM60.6 from:-112    till:-79.2 text:[[Macropoma]]
 color:cretaceous bar:NAM61 from:-99.6    till:-93.5 text:[[Macropomoides]]
 color:cretaceous bar:NAM62 from:-86.3   till:-71.5 text:[[Megalocoelacanthus]]
 color:cretaceous bar:NAM62.1 from:-67   till:-65 text:[[Coelacanth|Mawsoniidae indeterminate]]
 color:cretaceous bar:NAM63 from:-65    till:-64 text: [[Post Cretaceous Coelacanth fossils|Coelacanthiformes indeterminate]]?
 color:quaternary bar:NAM64 from:-1    till:0 text:[[Latimeria]]

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 from: -542    till:  -488.3   color:cambrian  text:[[Cambrian]]
 from: -488.3    till: -443.7    color:ordovician    text:[[Ordovician]]
 from: -443.7    till: -416    color:silurian    text:[[Silurian]]
 from: -416    till: -359.2   color:devonian    text:[[Devonian]]
 from: -359.2    till: -299   color:carboniferous    text:[[Carboniferous]]
 from: -299    till: -251   color:permian    text:[[Permian]]
 from: -251    till: -199.6  color:triassic    text:[[Triassic]]
 from: -199.6    till: -145.5   color:jurassic    text:[[Jurassic]]
 from: -145.5   till: -66   color:cretaceous    text:[[Cretaceous]]
 from: -66    till: -23.03   color:paleogene    text:[[Paleogene]]
 from: -23.03    till: -2.588   color:neogene    text:[[Neogene|Neog.]]
 from: -2.588    till: 0   color:quaternary    text:[[Quaternary|Q.]]

 bar:era
 from:  -542   till:  -251    color:paleozoic    text:[[Paleozoic Era]]
 from:  -251   till:  -66    color:mesozoic    text:[[Mesozoic Era]]
 from:  -66   till:  0   color:cenozoic    text:[[Cenozoic]]
</timeline>