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{{Short description|Taxonomy of lobe-finned fishes}}
{{Short description|Taxonomy of lobe-finned fishes}}
{{context|date=May 2016}}
{{context|date=May 2016}}
[[Sarcopterygii]] or the lobe-finned fishes ([[coelacanth]]s and [[lungfish]]es) were usually classified as either a class or a subclass of [[Osteichthyes]] based on the traditional [[Linnaean classification]]. Identification of the group is based on several characteristics, such as the presence of fleshy, lobed, paired fins, which are joined to the body by a single bone.<ref>{{cite book | vauthors = Clack JA | date = 2012 | title = Gaining Ground: The origin and evolution of tetrapods | publisher = Indiana University Press | isbn = 978-0-253-35675-8 }}</ref>
'''Cladistic classification of Sarcopterygii''' is the classication of [[Sarcopterygii]] as a clade containing not only the lobe-finned fishes ([[Coelacanth|coelacanths]] and [[lungfish]]) but also the tetrapods, which are closely related to lungfish. The taxon Sarcopterygii was traditionally classified as a [[Paraphyly|paraphyletic]] group considered either a class or a subclass of [[Osteichthyes]] (bony fish). Identification of the group is based on several characteristics, such as the presence of fleshy, lobed, paired fins, which are joined to the body by a single bone.<ref>{{cite book | vauthors = Clack JA | date = 2012 | title = Gaining Ground: The origin and evolution of tetrapods | publisher = Indiana University Press | isbn = 978-0-253-35675-8 }}</ref>


==Taxonomic and fossil history==
==Taxonomic and fossil history==
The properties defining the sarcopterygians are in contrast to the other group of bony fish, the [[Actinopterygii]], which have ray-fins made of bony rods, called ''[[lepidotrichia]]''.
The properties defining the sarcopterygians are in contrast to the other group of bony fish, the [[Actinopterygii]], which have ray-fins made of bony rods, called ''[[lepidotrichia]]''.
These two bony fish groups were classified together as Osteichthyes at one time, the whole combined group was seen as the sister group to the [[tetrapod]]s ([[mammal]]s, [[bird]]s, [[reptile]]s, and [[amphibian]]s).
These two bony fish groups were classified together as Osteichthyes at one time, the whole combined group was seen as parallel to the [[tetrapod]]s ([[mammal]]s, [[bird]]s, [[reptile]]s, and [[amphibian]]s).


The extensive fossil record and numerous morphological and molecular studies have shown, however, that lungfish and some fossil lobe-finned fish are more closely related to tetrapods than they are to coelacanths; as a result tetrapods are nested within Sarcopterygii.<ref>{{cite book | vauthors = Tudge C | date = 2000 | title = The variety of life | location = Oxford | publisher = Oxford University Press | isbn = 978-0-19-860426-6 }}</ref><ref>{{cite book | vauthors = Heiser JB, Janis CM, Pough FH | date = 2005 | title = Vertebrate life | publisher = Pearson/Prentice Hall | isbn = 978-0-321-77336-4 }}</ref> This abides to [[cladistic]]s in that in order for a [[clade]] to be monophyletic, it must have an ancestral species and all descendants of that common ancestor based on shared characteristics. As such mammals, birds and reptiles, and amphibians are highly derived lobe-finned fish despite looking nothing like the standard sarcopterygian anatomically speaking.
The extensive fossil record and numerous morphological and molecular studies have shown, however, that lungfish and some fossil lobe-finned fish ("rhipidistians") are more closely related to tetrapods than they are to coelacanths; as a result tetrapods are nested within Sarcopterygii.<ref>{{cite book | vauthors = Tudge C | date = 2000 | title = The variety of life | location = Oxford | publisher = Oxford University Press | isbn = 978-0-19-860426-6 }}</ref><ref>{{cite book | vauthors = Heiser JB, Janis CM, Pough FH | date = 2005 | title = Vertebrate life | publisher = Pearson/Prentice Hall | isbn = 978-0-321-77336-4 }}</ref> This abides to [[cladistic]]s in that in order for a group to be valid, it must [[Monophyly|have an ancestral species and all descendants of that common ancestor]] based on shared characteristics. As such mammals, sauropsids (birds and "reptiles"), and amphibians are highly derived sarcopterygians despite superficially looking nothing like the standard lobe-finned fish anatomically speaking. However, similarities can be noticed in their limb bones and [[tooth enamel]].<ref>{{Cite book |last=Benton |first=Michael J. |url=https://www.worldcat.org/oclc/867852756 |title=Vertebrate Palaeontology |publisher=Wiley-Blackwell |year=2014 |isbn=978-1-118-40764-6 |edition=4th |location=Chichester, West Sussex |pages=74 |oclc=867852756}}</ref> Additionally, lungfish and tetrapods share a divided [[Atrium (heart)|atrium]].<ref>{{Cite book |last=Pough |first=F. Harvey |url=https://www.worldcat.org/oclc/1022979490 |title=Vertebrate Life |publisher=Oxford University Press |others=Christine M. Janis, Sergi López-Torres |year=2018 |isbn=978-1-60535-607-5 |edition=10th |location=New York |pages=123 |oclc=1022979490}}</ref>


==Classification==
==Current taxonomy==
Multiple Linnean classifications have been proposed with the explicit intent to incorporate Sarcopterygii as a monophyletic taxon instead of maintaining its traditional paraphyletic definition.<ref>{{Cite journal | vauthors = Ahlberg PE |date=1991 |title=A re-examination of sarcopterygian interrelationships, with special reference to the Porolepiformes |url=https://academic.oup.com/zoolinnean/article-lookup/doi/10.1111/j.1096-3642.1991.tb00905.x |journal=Zoological Journal of the Linnean Society |language=en |volume=103 |issue=3 |pages=241–287 |doi=10.1111/j.1096-3642.1991.tb00905.x}}</ref><ref>{{cite book |title=Fishes of the World. |vauthors=Nelson JS, Grande TC, Wilson MV |date=April 2016 |publisher=John Wiley & Sons |isbn=978-1-118-34233-6}}</ref><ref>{{cite journal | vauthors = Betancur-R R, Wiley EO, Arratia G, Acero A, Bailly N, Miya M, Lecointre G, Ortí G | display-authors = 6 | title = Phylogenetic classification of bony fishes | journal = BMC Evolutionary Biology | volume = 17 | issue = 1 | pages = 162 | date = July 2017 | pmid = 28683774 | pmc = 5501477 | doi = 10.1186/s12862-017-0958-3 | doi-access = free | bibcode = 2017BMCEE..17..162B }}</ref><ref>{{Cite journal | vauthors = Tedersoo L |date=2017 |title=Proposal for practical multi-kingdom classification of eukaryotes based on monophyly and comparable divergence time criteria |url=http://biorxiv.org/lookup/doi/10.1101/240929 |journal=bioRxiv |language=en |doi=10.1101/240929|s2cid=90691603 }}</ref>
The list below shows the taxonomy of the extant members of class Sarcopterygii at the [[order (biology)|ordinal level]]. While this does reflect the evolutionary relationships within the group, it also retains the rankings seen in the Linnaean classification as suggested by some scientists.<ref>{{cite book | vauthors = Nelson JS, Grande TC, Wilson MV | title = Fishes of the World. | publisher = John Wiley & Sons | date = April 2016 | isbn = 978-1-118-34233-6 }}</ref> The evolutionary sequences are based from current phylogenetic work on the various subclades.<ref name="pmid22593086">{{cite journal | vauthors = Crawford NG, Faircloth BC, McCormack JE, Brumfield RT, Winker K, Glenn TC | title = More than 1000 ultraconserved elements provide evidence that turtles are the sister group of archosaurs | journal = Biology Letters | volume = 8 | issue = 5 | pages = 783–6 | date = October 2012 | pmid = 22593086 | pmc = 3440978 | doi = 10.1098/rsbl.2012.0331 }}</ref><ref name="pmid23624526">{{cite journal | vauthors = Wang Z, Pascual-Anaya J, Zadissa A, Li W, Niimura Y, Huang Z, Li C, White S, Xiong Z, Fang D, Wang B, Ming Y, Chen Y, Zheng Y, Kuraku S, Pignatelli M, Herrero J, Beal K, Nozawa M, Li Q, Wang J, Zhang H, Yu L, Shigenobu S, Wang J, Liu J, Flicek P, Searle S, Wang J, Kuratani S, Yin Y, Aken B, Zhang G, Irie N | display-authors = 6 | title = The draft genomes of soft-shell turtle and green sea turtle yield insights into the development and evolution of the turtle-specific body plan | journal = Nature Genetics | volume = 45 | issue = 6 | pages = 701–706 | date = June 2013 | pmid = 23624526 | pmc = 4000948 | doi = 10.1038/ng.2615 }}</ref><ref name="pmid23382987">{{cite journal | vauthors = McCormack JE, Harvey MG, Faircloth BC, Crawford NG, Glenn TC, Brumfield RT | title = A phylogeny of birds based on over 1,500 loci collected by target enrichment and high-throughput sequencing | journal = PLOS ONE | volume = 8 | issue = 1 | pages = e54848 | date = 2013 | pmid = 23382987 | pmc = 3558522 | doi = 10.1371/journal.pone.0054848 | arxiv = 1210.1604 | bibcode = 2013PLoSO...854848M | doi-access = free }}</ref><ref>{{cite journal | vauthors = Kuhl H, Frankl-Vilches C, Bakker A, Mayr G, Nikolaus G, Boerno ST, Klages S, Timmermann B, Gahr M | display-authors = 6 | title = An Unbiased Molecular Approach Using 3'-UTRs Resolves the Avian Family-Level Tree of Life | journal = Molecular Biology and Evolution | volume = 38 | issue = 1 | pages = 108–127 | date = January 2021 | pmid = 32781465 | pmc = 7783168 | doi = 10.1093/molbev/msaa191 }}</ref><ref name="pmid25755933">{{cite journal | vauthors = May-Collado LJ, Kilpatrick CW, Agnarsson I | title = Mammals from 'down under': a multi-gene species-level phylogeny of marsupial mammals (Mammalia, Metatheria) | journal = PeerJ | volume = 3 | issue = | pages = e805 | date = 2015 | pmid = 25755933 | pmc = 4349131 | doi = 10.7717/peerj.805 }}</ref><ref name="pmid26733575">{{cite journal | vauthors = Tarver JE, Dos Reis M, Mirarab S, Moran RJ, Parker S, O'Reilly JE, King BL, O'Connell MJ, Asher RJ, Warnow T, Peterson KJ, Donoghue PC, Pisani D | display-authors = 6 | title = The Interrelationships of Placental Mammals and the Limits of Phylogenetic Inference | journal = Genome Biology and Evolution | volume = 8 | issue = 2 | pages = 330–44 | date = January 2016 | pmid = 26733575 | pmc = 4779606 | doi = 10.1093/gbe/evv261 }}</ref>


=== Ahlberg (1991) ===
'''Class [[Sarcopterygii]]''' Romer, 1955
Class [[Osteichthyes]]
*'''Subclass [[Actinistia]]''' Cope, 1871

**Order [[Coelacanthiformes]] Berg, 1937
*'''Subclass [[Rhipidistia]]''' Cope, 1871
* Subclass [[Actinopterygii]]
* Subclass '''Sarcopterygii'''
**'''Infraclass [[Dipnoi]]''' Müller, 1844
** Plesion [[Onychodontiformes|Onychodontia]]
***Order [[Ceratodontiformes]] Berg, 1940
** Infraclass [[Coelacanth|Actinistia]]
***Order [[Lepidosireniformes]] Müller, 1844
**'''Infraclass [[Tetrapoda]]''' Goodrich, 1930
** Infraclass [[Rhipidistia]]
*** Superdivision [[Tetrapodomorpha]]
***'''Parvclass [[Lissamphibia]]''' Haeckel, 1866
****Order [[Caecilian|Apoda]] Oppel, 1811
**** Plesion [[Rhizodontida]]
**** Plesion [[Osteolepiformes]]
****Order [[Urodela]] Duméril, 1806
**** Plesion [[Panderichthyidae]]
****Order [[Frog|Anura]] Duméril, 1806
**** Division [[Tetrapod|Tetrapoda]]
***'''Parvclass [[Sauria]]''' Macartney, 1802
*** Superdivision [[Dipnomorph|Dipnomorpha]]
****'''Cohort [[Lepidosauria]]''' Haeckel, 1866
**** Plesion [[Porolepiformes]]
*****Order [[Rhynchocephalia]] Günther, 1867
**** Plesion ''[[Powichthys]]''
*****Order [[Squamata]] Oppel, 1811
****'''Cohort [[Testudinata]]''' Klein, 1760
**** Plesion ''[[Youngolepis]]''
**** Plesion ''[[Diabolepis]]''
*****Order [[Testudines]] Batsch, 1788
**** Division [[Lungfish|Dipnoi]]
****'''Cohort [[Archosauria]]''' Cope, 1869

*****Order [[Crocodylia]] Owen, 1842
=== Nelson ''et al''. (2016) ===
*****'''Subcohort [[Aves]]''' Linnaeus, 1758
Class [[Osteichthyes]]
******'''Infracohort [[Palaeognathae]]''' Pycraft, 1900

*******Order [[Struthioniformes]] Latham, 1790
* Subclass [[Actinopterygii]]
*******Order [[Rheiformes]] Forbes, 1884
* Subclass '''Sarcopterygii'''
*******Order [[Tinamiformes]] Huxley, 1872
** Infraclass [[Coelacanth|Actinistia]]
*******†Order [[Dinornithiformes]] Bonaparte, 1853
** †Infraclass [[Onychodontiformes|Onychodontida]]
*******†Order [[Aepyornithiformes]] Newton, 1884
** Infraclass [[Lungfish|Dipnomorpha]]
*******Order [[Apterygiformes]] Haeckel, 1866
** †Infraclass [[Rhizodontida]]
*******Order [[Casuariiformes]] Sclater 1880
** †Infraclass [[Osteolepiformes|Osteolepidida]]
******'''Infracohort [[Neognathae]]''' Pycraft, 1900
** †Infraclass Elpistostegalia
*******'''Section [[Galloanserae]]''' Sclater, 1880
** Infraclass [[Tetrapod|Tetrapoda]]
********Order [[Anseriformes]] Wagler, 1831

********Order [[Galliformes]] Temminck, 1820
=== Betancur-Rodrigues ''et al''. (2017) ===
*******'''Section [[Neoaves]]''' Sibley et al., 1988
Superclass '''Sarcopterygii'''
********'''Subsection [[Mirandornithes]]''' Sangster, 2005

*********Order [[Phoenicopteriformes]] Fürbringer, 1888
* Class [[Coelacanth|Coelacanthimorpha]]
*********Order [[Podicipediformes]] Fürbringer, 1888
* Class [[Dipnotetrapodomorpha]]
********'''Subdivision [[Gruimorphae]]''' Bonaparte, 1854
** Subclass [[Lungfish|Dipnomorpha]]
*********Order [[Gruiformes]] Bonaparte, 1854
** Subclass [[Tetrapodomorpha]]
*********Order [[Charadriiformes]] Huxley, 1867

********'''Subsection [[Columbaves]]''' Prum et al., 2015
Other classifications do not use Sarcopterygii as a ranked taxon but still nonetheless still reject traditional paraphyletic assemblages. In the scheme below, sarcopterygian groups are marked in bold letters.
*********Order [[Musophagiformes]] Seebohm, 1890

*********Order [[Cuculiformes]] Wagler, 1830
=== Tedersoo (2017) ===
*********Order [[Otidiformes]] Wagler, 1830
Phylum [[Vertebrate|Craniata]]
*********Order [[Columbiformes]] Latham, 1790

*********Order [[Pterocliformes]] Huxley, 1868
* Class [[Cyclostomi]]
*********Order [[Mesitornithiformes]] Wetmore, 1960
* Class [[Chondrichthyes]]
********Order [[Opisthocomiformes]] L'Herminier, 1837
* Class [[Cladistia]]
********'''Subsection [[Strisores]]''' Baird, 1858
* Class [[Actinopteri]]
*********Order [[Steatornithidae|Steatornithiformes]] Sharpe, 1891
* '''Class [[Coelacanth|unspecified]]'''
*********Order [[Nyctibiidae|Nyctibiiformes]] Sharpe, 1891
* '''Class [[Lungfish|Ceratodontimorpha]]'''
*********Order [[Caprimulgidae|Caprimulgiformes]] Ridgway, 1881
* '''Class [[Amphibian|Amphibia]]'''
*********Order [[Podargidae|Podargiformes]] Sharpe, 1891
* '''Class [[Mammal|Mammalia]]'''
*********Order [[Aegothelidae|Aegotheliformes]] Simonetta, 1967
* '''Class [[Rhynchocephalia|Sphenodontea]]'''
*********Order [[Apodiformes]] Peters, 1940
* '''Class [[Squamata|Squamatea]]'''
********'''Subsection [[Ardeae]]''' Wagler, 1830
* '''Class [[Turtle|Testudinea]]'''
*********Order [[Phaethontidae|Phaethontiformes]] Sharpe, 1891
* '''Class [[Crocodilia|Crocodylea]]'''
*********Order [[Eurypygiformes]] Fürbringer, 1888
* '''Class [[Bird|Aves]]'''
*********Order [[Gaviiformes]] Wetmore & Miller, 1926
*********Order [[Sphenisciformes]] Sharpe, 1891
*********Order [[Procellariiformes]] Fürbringer, 1888
*********Order [[Ciconiiformes]] Bonaparte, 1854
*********Order [[Suliformes]] Sharpe, 1891
*********Order [[Pelecaniformes]] Sharpe, 1891
********'''Subsection [[Telluraves]]''' Yuri et al., 2013
*********'''Division [[Afroaves]]''' Ericson, 2012
**********Order [[Cathartiformes]] Coues, 1884
**********Order [[Accipitriformes]] Vieillot, 1816
**********Order [[Strigiformes]] Wagler, 1830
**********Order [[Coliiformes]] Murie, 1872
**********Order [[Leptosomiformes]] Sharpe, 1991
**********Order [[Trogoniformes]] AOU, 1886
**********Order [[Bucerotiformes]] Fürbringer, 1888
**********Order [[Coraciiformes]] Forbes, 1884
**********Order [[Piciformes]] Meyer & Wolf, 1810
*********'''Division [[Australaves]]''' Ericson, 2012
**********Order [[Cariamiformes]] Fürbringer, 1888
**********Order [[Falconiformes]] Sharpe, 1874
**********Order [[Psittaciformes]] Wagler, 1830
**********Order [[Passeriformes]] Linnaeus, 1758
***'''Parvclass [[Mammalia]]''' Linnaeus, 1758
****Order [[Monotremata]] Bonaparte, 1837
****'''Supercohort [[Theria]]''' Parker & Haswell, 1897
*****'''Cohort [[Marsupialia]]''' Illiger, 1811
******Order [[Didelphimorphia]] Gill, 1872
******Order [[Paucituberculata]] Trouessart, 1898
******'''Subcohort [[Australidelphia]]''' Szalay, 1982
*******Order [[Microbiotheria]] Ameghino, 1889
*******Order [[Diprotodontia]] Owen, 1866
*******Order [[Notoryctemorphia]] Kirsch, 1977
*******Order [[Dasyuromorphia]] Gill, 1872
*******Order [[Peramelemorphia]] Ameghino, 1889
*****'''Cohort [[Placentalia]]''' Owen, 1837
******'''Subcohort [[Atlantogenata]]''' Waddell, Okada & Hasegawa, 1999
*******'''Infracohort [[Xenarthra]]''' Cope, 1889
********Order [[Cingulata]] Illiger, 1811
********Order [[Pilosa]] Flower, 1883
*******'''Infracohort [[Afrotheria]]''' Stanhope et al., 1998
********Order [[Afrosoricida]] Stanhope et al., 1998
********Order [[Macroscelidea]] Butler, 1956
********Order [[Tubulidentata]] Huxley, 1872
********Order [[Hyracoidea]] Huxley, 1869
********Order [[Sirenia]] Illiger, 1811
********Order [[Proboscidea]] Illiger, 1810
******'''Subcohort [[Boreoeutheria]]''' Springer & de Jong, 2001
*******'''Infracohort [[Laurasiatheria]]''' Waddell, Okada & Hasegawa, 1999
********Order [[Eulipotyphla]] Douady et al., 2002
********Order [[Chiroptera]] Blumenbach, 1779
********Order [[Pholidota]] Weber, 1904
********Order [[Carnivora]] Bowdich, 1821
********Order [[Perissodactyla]] Owen, 1848
********Order [[Artiodactyla]] Owen, 1848
*******'''Infracohort [[Euarchontoglires]]''' Murphy et al., 2001
********Order [[Lagomorpha]] Brandt, 1855
********Order [[Rodent]]ia Bowdich, 1821
********Order [[Scandentia]] Wagner, 1855
********Order [[Dermoptera]] Illiger, 1811
********Order [[Primates]] Linnaeus, 1758


== See also ==
== See also ==

Latest revision as of 13:12, 11 March 2024

Cladistic classification of Sarcopterygii is the classication of Sarcopterygii as a clade containing not only the lobe-finned fishes (coelacanths and lungfish) but also the tetrapods, which are closely related to lungfish. The taxon Sarcopterygii was traditionally classified as a paraphyletic group considered either a class or a subclass of Osteichthyes (bony fish). Identification of the group is based on several characteristics, such as the presence of fleshy, lobed, paired fins, which are joined to the body by a single bone.[1]

Taxonomic and fossil history

[edit]

The properties defining the sarcopterygians are in contrast to the other group of bony fish, the Actinopterygii, which have ray-fins made of bony rods, called lepidotrichia. These two bony fish groups were classified together as Osteichthyes at one time, the whole combined group was seen as parallel to the tetrapods (mammals, birds, reptiles, and amphibians).

The extensive fossil record and numerous morphological and molecular studies have shown, however, that lungfish and some fossil lobe-finned fish ("rhipidistians") are more closely related to tetrapods than they are to coelacanths; as a result tetrapods are nested within Sarcopterygii.[2][3] This abides to cladistics in that in order for a group to be valid, it must have an ancestral species and all descendants of that common ancestor based on shared characteristics. As such mammals, sauropsids (birds and "reptiles"), and amphibians are highly derived sarcopterygians despite superficially looking nothing like the standard lobe-finned fish anatomically speaking. However, similarities can be noticed in their limb bones and tooth enamel.[4] Additionally, lungfish and tetrapods share a divided atrium.[5]

Classification

[edit]

Multiple Linnean classifications have been proposed with the explicit intent to incorporate Sarcopterygii as a monophyletic taxon instead of maintaining its traditional paraphyletic definition.[6][7][8][9]

Ahlberg (1991)

[edit]

Class Osteichthyes

Nelson et al. (2016)

[edit]

Class Osteichthyes

Betancur-Rodrigues et al. (2017)

[edit]

Superclass Sarcopterygii

Other classifications do not use Sarcopterygii as a ranked taxon but still nonetheless still reject traditional paraphyletic assemblages. In the scheme below, sarcopterygian groups are marked in bold letters.

Tedersoo (2017)

[edit]

Phylum Craniata

See also

[edit]

References

[edit]
  1. ^ Clack JA (2012). Gaining Ground: The origin and evolution of tetrapods. Indiana University Press. ISBN 978-0-253-35675-8.
  2. ^ Tudge C (2000). The variety of life. Oxford: Oxford University Press. ISBN 978-0-19-860426-6.
  3. ^ Heiser JB, Janis CM, Pough FH (2005). Vertebrate life. Pearson/Prentice Hall. ISBN 978-0-321-77336-4.
  4. ^ Benton, Michael J. (2014). Vertebrate Palaeontology (4th ed.). Chichester, West Sussex: Wiley-Blackwell. p. 74. ISBN 978-1-118-40764-6. OCLC 867852756.
  5. ^ Pough, F. Harvey (2018). Vertebrate Life. Christine M. Janis, Sergi López-Torres (10th ed.). New York: Oxford University Press. p. 123. ISBN 978-1-60535-607-5. OCLC 1022979490.
  6. ^ Ahlberg PE (1991). "A re-examination of sarcopterygian interrelationships, with special reference to the Porolepiformes". Zoological Journal of the Linnean Society. 103 (3): 241–287. doi:10.1111/j.1096-3642.1991.tb00905.x.
  7. ^ Nelson JS, Grande TC, Wilson MV (April 2016). Fishes of the World. John Wiley & Sons. ISBN 978-1-118-34233-6.
  8. ^ Betancur-R R, Wiley EO, Arratia G, Acero A, Bailly N, Miya M, et al. (July 2017). "Phylogenetic classification of bony fishes". BMC Evolutionary Biology. 17 (1): 162. Bibcode:2017BMCEE..17..162B. doi:10.1186/s12862-017-0958-3. PMC 5501477. PMID 28683774.
  9. ^ Tedersoo L (2017). "Proposal for practical multi-kingdom classification of eukaryotes based on monophyly and comparable divergence time criteria". bioRxiv. doi:10.1101/240929. S2CID 90691603.
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