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*** Order [[Chimaeriformes]]
* ''[[Incertae sedis]]''
**†''[[Bandringa]]''<ref>{{Cite web |title=Mazon Monday #19: Species Spotlight: Bandringa rayi #MazonCreek #fossils #MazonMonday #shark |url=https://www.esconi.org/esconi_earth_science_club/2020/08/mazon-monday-19-species-spotlight-bandringa-rayi-mazoncreek-fossils-mazonmonday-shark.html |access-date=2020-10-04 |website=Earth Science Club of Northern Illinois - ESCONI}}</ref>
**†''[[Delphyodontos]]''<ref>{{Cite web|url=http://people.sju.edu/~egrogan/BearGulch/pages_fish_species/Delphyodontos_dacriformes.html|title=Bear Gulch - Delphyodontos dacriformes|work=Fossil Fishes of Bear Gulch|access-date=2019-05-15|archive-date=25 February 2015|archive-url=https://web.archive.org/web/20150225041927/http://people.sju.edu/~egrogan/BearGulch/pages_fish_species/Delphyodontos_dacriformes.html|url-status=dead}}</ref>
**†
** †[[Mcmurdodontidae]]<ref>{{Citation |last1=Long |first1=John |title=Fossil chondrichthyan remains from the Middle Devonian Kevington Creek Formation, South Blue Range, Victoria |date=2021-10-28 |url=https://researchnow-admin.flinders.edu.au/ws/portalfiles/portal/48435020/Long_et_al_2021_MId_dev_sharks_Victoria_JOHN_MAISEY_VOLUME.pdf|work=Ancient Fishes and their Living Relatives |pages=239–245 |editor-last=Pradel |editor-first=Alan |access-date=2023-11-30 |place=Munich, Germany |publisher=Verlag, Dr Friedrich Pfeil |isbn=978-3-89937-269-4 |last2=Thomson |first2=Victoria |last3=Burrow |first3=Carole |last4=Turner |first4=Susan |editor2-last=Denton |editor2-first=John S.S. |editor3-last=Janvier |editor3-first=Philippe}}</ref>
** †''[[Nanocetorhinus]]''<ref name=UnderwoodetSchlogl>{{cite journal |author=Charlie J. Underwood and Jan Schlogl |year=2012 |title=Deep water chondrichthyans from the Early Miocene of the Vienna Basin (Central Paratethys, Slovakia) |journal=Acta Palaeontologica Polonica |volume=58 |issue=3 |pages=
** †''[[Plesioselachus]]''<ref name=":0">{{cite journal |last1=Anderson |first1=M. Eric |last2=Long |first2=John A. |last3=Gess |first3=Robert W. |last4=Hiller |first4=Norton |year=1999 |title=An unusual new fossil shark (Pisces: Chondrichthyes) from the Late Devonian of South Africa |url=http://museum.wa.gov.au/research/records-supplements/records/unusual-new-fossil-shark-pisces-chondrichthyes-late-devonian-so |journal=Records of the Western Australian Museum |volume=57 |pages=151–156}}</ref>
** †[[Psammodontiformes]]
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}}
'''Chondrichthyes''' ({{IPAc-en|k|ɒ|n|ˈ|d|r|ɪ|k|θ|i
The class is divided into two subclasses: [[Elasmobranchii]] ([[shark]]s, [[Batoidea|ray]]s, [[
==Anatomy==
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===Skeleton===
The skeleton is cartilaginous. The [[notochord]] is gradually replaced by a vertebral column during development, except in [[Holocephali]], where the notochord stays intact. In some deepwater sharks, the column is reduced.<ref>
As they do not have [[bone marrow]], [[red blood cell]]s are produced in the [[spleen]] and the epigonal organ (special tissue around the [[gonad]]s, which is also thought to play a role in the immune system). They are also produced in the [[Leydig's organ]], which is only found in certain cartilaginous fishes. The subclass [[Holocephali]], which is a very specialized group, lacks both the Leydig's and epigonal organs.
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===Nervous system===
[[File:Skate Brain Regions.png|thumb|Regions of a Chondrichthyes brain colored and labeled on dissected skate. The [[Anatomical terms of location#Cranial and caudal|rostral]] end of the skate is to the right.]]
In chondrichthyans, the nervous system is composed of a small brain,
===Immune system===
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==Evolution==
{{See also|Evolution of fish}}
{{Further|List of transitional fossils#Chondrichthyes|List of prehistoric cartilaginous fish}}
Cartilaginous fish are considered to have evolved from [[acanthodians]]. The discovery of ''[[Entelognathus]]'' and several examinations of acanthodian characteristics indicate that bony fish evolved directly from placoderm like ancestors, while acanthodians represent a paraphyletic assemblage leading to Chondrichthyes. Some characteristics previously thought to be exclusive to acanthodians are also present in basal cartilaginous fish.<ref>{{cite journal |author1=Min Zhu |author2=Xiaobo Yu |author3=Per Erik Ahlberg |author4=Brian Choo |author5=Jing Lu |author6=Tuo Qiao |author7=Qingming Qu |author8=Wenjin Zhao |author9=Liantao Jia |author10=Henning Blom |author11=You'an Zhu |year=2013 |title=A Silurian placoderm with osteichthyan-like marginal jaw bones |journal=[[Nature (journal)|Nature]] |volume=502 |issue=7470 |pages=188–193 |doi=10.1038/nature12617 |pmid=24067611|bibcode=2013Natur.502..188Z |s2cid=4462506 }}</ref> In particular, new phylogenetic studies find cartilaginous fish to be well nested among acanthodians, with ''[[Doliodus]]'' and ''[[Tamiobatis]]'' being the closest relatives to Chondrichthyes.<ref>{{cite journal|doi=10.26879/601|title=The diplacanthid fishes (Acanthodii, Diplacanthiformes, Diplacanthidae) from the Middle Devonian of Scotland|journal=Palaeontologia Electronica|year=2016|last1=Burrow|first1=CJ|last2=Den Blaauwen|first2=J.|last3=Newman|first3=MJ|last4=Davidson|first4=RG|doi-access=free}}</ref> Recent studies vindicate this, as ''[[Doliodus]]'' had a mosaic of chondrichthyan and acanthodian traits.<ref>{{cite journal|doi=10.1206/3875.1|title=Pectoral Morphology in ''Doliodus'': Bridging the 'Acanthodian'-Chondrichthyan Divide|journal=American Museum Novitates|issue=3875|pages=1–15|year=2017|last1=Maisey|first1=John G.|last2=Miller|first2=Randall|last3=Pradel|first3=Alan|last4=Denton|first4=John S.S.|last5=Bronson|first5=Allison|last6=Janvier|first6=Philippe|s2cid=44127090|url=https://www.archive.org/download/pectoralmorphol00mais/pectoralmorphol00mais.pdf |archive-url=https://ghostarchive.org/archive/20221009/https://www.archive.org/download/pectoralmorphol00mais/pectoralmorphol00mais.pdf |archive-date=2022-10-09 |url-status=live}}</ref> Dating back to the Middle and Late [[Ordovician]] Period, many isolated scales, made of [[dentine]] and bone, have a structure and growth form that is chondrichthyan-like. They may be the remains of [[stem group|stem]]-chondrichthyans, but their classification remains uncertain.<ref name="Andreev2015">{{cite journal |last1 = Andreev |first1 = Plamen S.|last2=Coates |first2 =Michael I. |last3=Shelton |first3=Richard M. |last4=Cooper |first4=Paul R. |last5=Smith |first5=M. Paul |last6=Sansom |first6=Ivan J. |year=2015 |title=Ordovician chondrichthyan-like scales from North America |journal=[[Palaeontology (journal)|Palaeontology]] |volume=58 |issue=4 |pages=691–704 |doi=10.1111/pala.12167|s2cid = 140675923|doi-access=free }}</ref><ref name="Sansom2012">{{cite journal |last1=Sansom |first1=Ivan J. |last2=Davies |first2= Neil S. |last3=Coates |first3=Michael I. |last4=Nicoll |first4=Robert S. |last5=Ritchie |first5=Alex |year=2012 |title=Chondrichthyan-like scales from the Middle Ordovician of Australia |journal=[[Palaeontology (journal)|Palaeontology]] |volume=55 |issue=2 |pages=243–247 |doi=10.1111/j.1475-4983.2012.01127.x|bibcode=2012Palgy..55..243S |doi-access=free }}</ref><ref name="Andreev2016">{{cite journal |last1=Andreev |first1=Plamen |last2=Coates |first2=Michael I. |last3=Karatajūtė-Talimaa |first3=Valentina |last4=Shelton |first4=Richard M. |last5=Cooper |first5=Paul R. |last6=Wang |first6=Nian-Zhong |last7=Sansom |first7=Ivan J. |year=2016 |title=The systematics of the Mongolepidida (Chondrichthyes) and the Ordovician origins of the clade |journal=PeerJ |volume=4 |page=e1850 |doi=10.7717/peerj.1850|pmid=27350896 |pmc=4918221 |doi-access=free }}</ref>
The earliest unequivocal fossils of acanthodian-grade cartilaginous fishes are ''[[Qianodus]]'' and ''[[Fanjingshania]]'' from the early Silurian ([[Aeronian]]) of [[Guizhou]], China around 439 million years ago, which are also the oldest unambiguous remains of any jawed vertebrates.<ref name=":12">{{cite journal |last1=Andreev |first1=Plamen S. |last2=Sansom |first2=Ivan J. |last3=Li |first3=Qiang |last4=Zhao |first4=Wenjin |last5=Wang |first5=Jianhua |last6=Wang |first6=Chun-Chieh |last7=Peng |first7=Lijian |last8=Jia |first8=Liantao |last9=Qiao |first9=Tuo |last10=Zhu |first10=Min |date=September 2022 |title=Spiny chondrichthyan from the lower Silurian of South China |url=https://www.nature.com/articles/s41586-022-05233-8 |journal=Nature |volume=609 |issue=7929 |pages=969–974 |doi=10.1038/s41586-022-05233-8 |pmid=36171377 |bibcode=2022Natur.609..969A |s2cid=252570103}}</ref><ref name=":02">{{Cite journal |last1=Andreev |first1=Plamen S. |last2=Sansom |first2=Ivan J. |last3=Li |first3=Qiang |last4=Zhao |first4=Wenjin |last5=Wang |first5=Jianhua |last6=Wang |first6=Chun-Chieh |last7=Peng |first7=Lijian |last8=Jia |first8=Liantao |last9=Qiao |first9=Tuo |last10=Zhu |first10=Min |date=2022-09-28 |title=The oldest gnathostome teeth |url=http://dx.doi.org/10.1038/s41586-022-05166-2 |journal=Nature |volume=609 |issue=7929 |pages=964–968 |bibcode=2022Natur.609..964A |doi=10.1038/s41586-022-05166-2 |issn=0028-0836 |pmid=36171375 |s2cid=252569771}}</ref> ''Shenacanthus vermiformis'', which lived 436 million years ago, had thoracic armour plates resembling those of placoderms.<ref>
By the start of the Early Devonian, 419 million years ago, [[jawed fish]]es had divided into three distinct groups: the now extinct [[placoderm]]s (a paraphyletic assemblage of ancient armoured fishes), the [[bony fish]]es, and the clade that includes [[spiny sharks]] and early [[cartilaginous fish]]. The modern bony fishes, class [[Osteichthyes]], appeared in the late [[Silurian]] or early Devonian, about 416 million years ago. The first abundant genus of shark, ''[[Cladoselache]]'', appeared in the oceans during the Devonian Period. The first Cartilaginous fishes evolved from ''[[Doliodus]]''-like [[spiny shark]] ancestors.
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| align=center style="background:rgb(180,120,180)" | [[Petalodontiformes|<span style="color:white;">†Petalodontiformes</span>]]
| [[File:Strigilodus tollesonae-novataxa 2023-Hodnett Toomey Olson.jpg|147x147px]]
| align=center | [[Petalodontiformes|Petalodonts]]
| align=center | <small></small>Zangerl, 1981▼
| align=center | 4▼
| align=center |
▲| align=center | <small></small>
| align=center |
| valign=top | Members of the holocephali, some genera resembled parrot fish, but some members of the [[Janassidae]] resembled skates.
▲| align=center |
| align=center | ▼
|-
| align=center style="background:rgb(165,100,165)" | [[Helodontiformes|<span style="color:white;">†Helodontiformes</span>]]
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| align=center |
| align=center |
| valign=top | Members of the holocephali that resembled flying fish, are often characterized by large eyes, large upturned pectoral fins, and club-like tails.
|-
| align=center style="background:rgb(165,100,165)" | [[Debeeriiformes|<span style="color:white;">†Debeeriiformes</span>]]
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| align=center style="background:rgb(180,120,180)" | [[Symmoriida|<span style="color:white;">†Symmoriida</span>]]
| [[File:
| align=center | [[Symmoriidae|Symmoriids]]
| align=center | <small></small>Zangerl, 1981 (sensu Maisey, 2007)▼
▲| align=center | 4
| align=center |
▲| align=center | <small></small>
| align=center |
| valign=top | Members of the holocephali, they were heavily [[sexually dimorphic]].<ref>{{cite journal | last1 = Coates | first1 = M. | last2 = Gess | first2 = R. | last3 = Finarelli | first3 = J. | last4 = Criswell | first4 = K. | last5 = Tietjen | first5 = K. | year = 2016 | title = A symmoriiform chondrichthyan braincase and the origin of chimaeroid fishes | journal = Nature | volume = 541| issue = 7636| pages = 208–211| doi = 10.1038/nature20806 | pmid = 28052054 | bibcode = 2017Natur.541..208C | s2cid = 4455946 }}</ref>▼
| align=center | ▼
| align=center | ▼
▲| valign=top | <ref>{{cite journal | last1 = Coates | first1 = M. | last2 = Gess | first2 = R. | last3 = Finarelli | first3 = J. | last4 = Criswell | first4 = K. | last5 = Tietjen | first5 = K. | year = 2016 | title = A symmoriiform chondrichthyan braincase and the origin of chimaeroid fishes | journal = Nature | volume = 541| issue = 7636| pages = 208–211| doi = 10.1038/nature20806 | pmid = 28052054 | bibcode = 2017Natur.541..208C | s2cid = 4455946 }}</ref>
|-
| align=center style="background:rgb(165,100,165)" | [[Eugeneodontida|<span style="color:white;">†Eugeneodontida</span>]]
| [[File:Helicoprion_reccon.png|140px]]
| align=center | [[Eugeneodontida|Eugeneodonts]]
| align=center | <small></small>Eugeneodontida
<small></small><small>Zangerl, 1981</small>
| align=center | ▼
▲| align=center | 4
| align=center |
| align=center |
| valign=top | Members of the holocephali, they are characterized by large tooth whorls in their jaws.<ref>{{cite journal |last1=Tapanila |first1=L |last2=Pruitt |first2=J |last3=Pradel |first3=A |last4=Wilga |first4=C |last5=Ramsay |first5=J |last6=Schlader |first6=R |last7=Didier |first7=D |year=2013 |title=Jaws for a spiral-tooth whorl: CT images reveal novel adaptation and phylogeny in fossil Helicoprion |journal=Biology Letters |volume=9 |issue= 2|page=20130057 |doi=10.1098/rsbl.2013.0057 |pmid=23445952 |pmc=3639784 }}</ref>
|-
| align=center style="background:rgb(180,120,180)" | [[Psammodontiformes|<span style="color:white;">†Psammodonti-<br />formes</span>]]
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| [[File:Squatinactis NT small.jpg|140px]]
| align=center |
| align=center | <small></small>Cappetta et al., 1993
| align=center | 1
| align=center | 1
| align=center |
| valign=top |
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| align=center style="background:rgb(180,120,180)" | [[Cladoselachiformes|<span style="color:white;">†Cladoselachi-<br />formes</span>]]
| [[File:
| align=center |
| align=center | <small></small>Dean, 1894
▲| align=center | 1
▲| align=center | 2
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| valign=top | Holocephalans, and potential members of the symmoriida.
| align=center | ▼
| align=center | ▼
|-
| align=center style="background:rgb(165,100,165)" | [[Xenacanthiformes|<span style="color:white;">†Xenacanthiformes</span>]]
| [[File:
| align=center | [[Xenacanthida|Xenacanths]]
| align=center | <small></small>Glikman, 1964▼
▲| align=center | 4
| align=center |
▲| align=center | <small></small>
| align=center |
| valign=top | Eel-like elasmobranchs that were some of the top freshwater predators of the late Paleozoic.
| align=center | ▼
|-
| align=center style="background:rgb(180,120,180)" | [[Ctenacanthiformes|<span style="color:white;">†Ctenacanthi-<br />formes</span>]]
| [[File:
| align=center | [[Ctenacanthiformes|Ctenacanths]]
| align=center | <small></small>Glikman, 1964▼
▲| align=center | 2
| align=center |
▲| align=center | <small></small>
| align=center |
| valign=top | Shark-like elasmobranchs characterized by their robust heads and large dorsal fin spines.
|-
| align=center style="background:rgb(165,100,165)" | [[Hybodontiformes|<span style="color:white;">†Hybodontiformes</span>]]
| [[File:Hybodus
| align=center | [[Hybodontiformes|Hybodonts]]
| align=center | <small></small>Patterson, 1966▼
▲| align=center | 5
| align=center |
▲| align=center | <small></small>
| align=center |
| valign=top | Shark-like elasmobranchs distinguished by their conical tooth shape, and the presence of a spine on each of their two dorsal fins.
|}
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* [[Cartilaginous versus bony fishes]]
* [[Largest organisms#Cartilaginous fish (Chondrichthyes)|Largest cartilaginous fishes]]
* [[Important Shark and Ray Areas]] (ISRA)
* [[Threatened rays]]
* [[Threatened sharks]]
|