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*** Order [[Chimaeriformes]]
* ''[[Incertae sedis]]''
** †''[[Acanthorhachis]]''<ref>{{Cite web|title=Fossilworks: Acanthorhachis|url=http://www.fossilworks.org/cgi-bin/bridge.pl?a=taxonInfo&taxon_no=289828|access-date=17 December 2021|website=fossilworks.org}}</ref>
**†''[[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>
**†''[[ListracanthusListracanthidae]]''<ref name="mutter2006">{{cite journal | first1 = R.J. | last1 = Mutter | first2 = A.G. | last2 = Neuman | title = An enigmatic chondrichthyan with Paleozoic affinities from the Lower Triassic of western Canada | journal = Acta Palaeontologica Polonica | volume = 51 | issue = 2 | pages = 271–282 | url = https://www.app.pan.pl/article/item/app51-271.html}}</ref><ref>{{Cite web|title=Fossilworks: Acanthorhachis|url=https://paleobiodb.org/classic/checkTaxonInfo?taxon_no=289828|access-date=17 December 2021|website=fossilworks.org}}</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=487-509487–509 |url=http://app.pan.pl/article/item/app20110101.html |doi=10.4202/app.2011.0101|doi-access=free }}</ref>
** †''[[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|.|iː|z}}; {{etymology|grc|''{{wiktWikt-lang|grc|χόνδρος}}'' (khóndros{{grc-transl|χόνδρος}})|cartilage||''{{wiktWikt-lang|grc|ἰχθύς}}'' (ikhthús{{grc-transl|ἰχθύς}})|fish}}) is a [[class (biology)|class]] of [[jawed fish]] that contains the '''cartilaginous fishesfish''' thator '''chondrichthyans''', which all have skeletons[[skeleton]]s primarily composed of [[cartilage]]. They can be contrasted with the [[Osteichthyes]] or ''bony fishesfish'', which have skeletons primarily composed of [[bone tissue]]. Chondrichthyes are jawed[[aquatic animal|aquatic]] [[vertebrate]]s with paired [[Fishpaired fin|fins]], paired [[nare]]s, scales[[placoid scale]]s, and[[conus a heart with its chambersarteriosus]] in series.the Extant[[heart]], chondrichthyesand rangea inlack sizeof from[[operculum the(fish)|opercula]] 10and cm[[swim (3bladder]]s.9 in)Within [[finlessthe sleeperinfraphylum ray[[Gnathostomata]], tocartilaginous thefishes 10are mdistinct (32from ft)all [[whaleother shark]].jawed vertebrates.
 
The class is divided into two subclasses: [[Elasmobranchii]] ([[shark]]s, [[Batoidea|ray]]s, [[Skateskate (fish)|skate]]s, and [[sawfish]]) and [[Holocephali]] ([[chimaera]]s, sometimes called ghost sharks, which are sometimes separated into their own class). Extant chondrichthyans range in size from the {{cvt|10|cm}} [[finless sleeper ray]] to the over {{cvt|10|m}} [[whale shark]].
 
Within the infraphylum [[Gnathostomata]], cartilaginous fishes are distinct from all other jawed vertebrates.
 
==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>[{{Cite book|url=https://books.google.com/books?id=cxxSN4YA2i8C&dq=Notochord%3A:+Chimaeroids+%22Some+deepwater+squaloid%2C,+hexanchoid%2C,+and+lamnoid+sharks+%22&pg=PA23 |title=Sharks of the World: An Annotated and Illustrated Catalogue of Shark Species Known to Date]|first1=Leonard J. V.|last1=Compagno|first2=Food and Agriculture Organization of the United|last2=Nations|date=20 November 2001|publisher=Food & Agriculture Org.|isbn=9789251045435 |via=Google Books}}</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, 8-108–10 pairs of cranial nerves, and a spinal cord with spinal nerves.<ref>{{Cite journal|last=Collin|first=Shaun P.|date=2012|title=The Neuroecology of Cartilaginous Fishes: Sensory Strategies for Survival|journal=Brain, Behavior and Evolution|language=en|volume=80|issue=2|pages=80–96|doi=10.1159/000339870|pmid=22986825|s2cid=207717002|issn=1421-9743}}</ref> They have several sensory organs which provide information to be processed. [[Ampullae of Lorenzini]] are a network of small jelly filled pores called [[Electroreception|electroreceptors]] which help the fish sense electric fields in water. This aids in finding prey, navigation, and sensing temperature. The [[Lateral line]] system has modified epithelial cells located externally which sense motion, vibration, and pressure in the water around them. Most species have large well-developed eyes. Also, they have very powerful nostrils and [[Olfactory bulb|olfactory]] organs. Their inner ears consist of 3 large [[semicircular canals]] which aid in balance and orientation. Their sound detecting apparatus has limited range and is typically more powerful at lower frequencies. Some species have [[Electric organ (biology)|electric organs]] which can be used for defense and predation. They have relatively simple brains with the forebrain not greatly enlarged. The structure and formation of myelin in their nervous systems are nearly identical to that of tetrapods, which has led evolutionary biologists to believe that Chondrichthyes were a cornerstone group in the evolutionary timeline of myelin development.<ref>{{Cite journal|last=de Bellard|first=Maria Elena|date=2016-06-15|title=Myelin in cartilaginous fish|journal=Brain Research|volume=1641|issue=Pt A|pages=34–42|doi=10.1016/j.brainres.2016.01.013|issn=0006-8993|pmc=4909530|pmid=26776480}}</ref>
 
===Immune system===
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| align=center | 1
| align=center | 2
| align=center | 5-75–7
| align=center | 5-75–7
| align=center |
| align=center |
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==Evolution==
[[File:Evolution of cartilaginous fishes.png|thumb|400px|right|{{center|1=Radiation of [[cartilaginous fish]]es, based on [[Michael Benton]], 2005.<ref>{{cite book |last=Benton |first=M. J. |year=2005 |title=Vertebrate Palaeontology |publisher=Blackwell |edition=3rd |at=Fig 7.13 on page 185 |isbn=978-0-632-05637-8 |url=https://books.google.com/books?id=VThUUUtM8A4C&pg=PA185 }}</ref>}}]]
{{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>[{{cite journal | url=https://pubmed.ncbi.nlm.nih.gov/36171378/ | pmid=36171378 | year=2022 | last1=Zhu | first1=Y. A. | last2=Li | first2=Q. | last3=Lu | first3=J. | last4=Chen | first4=Y. | last5=Wang | first5=J. | last6=Gai | first6=Z. | last7=Zhao | first7=W. | last8=Wei | first8=G. | last9=Yu | first9=Y. | last10=Ahlberg | first10=P. E. | last11=Zhu | first11=M. | title=The oldest complete jawed vertebrates from the early Silurian of China | journal=Nature | volume=609 | issue=7929 | pages=954–958 | doi=10.1038/s41586-022-05136-8 | bibcode=2022Natur.609..954Z | s2cid=252569910 PubMed]}}</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]]
| [[File:Belantsea montana.JPG|140px]]
| 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 |
| valign=top |
|-
| 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.
| valign=top |
|-
| 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:Stethacanthus BWSymmorium1DB.jpg|140px147x147px]]
| 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:Early SharkCladoselache.jpgpng|140px140x140px]]
| align=center |
| align=center | <small></small>Dean, 1894
| align=center | 1
| align=center | 2
| align=center |
| valign=top | Holocephalans, and potential members of the symmoriida.
| align=center |
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| align=center style="background:rgb(165,100,165)" | [[Xenacanthiformes|<span style="color:white;">†Xenacanthiformes</span>]]
| [[File:Triodus1dbXenacanth.jpgpng|140px147x147px]]
| 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 |
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|-
| align=center style="background:rgb(180,120,180)" | [[Ctenacanthiformes|<span style="color:white;">†Ctenacanthi-<br />formes</span>]]
| [[File:Sphenacanthus1DBDracopristis hoffmanorum.jpgpng|140px147x147px]]
| 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 |
| align=center |
| valign=top |
|-
| align=center style="background:rgb(165,100,165)" | [[Hybodontiformes|<span style="color:white;">†Hybodontiformes</span>]]
| [[File:Hybodus modelhauffianus.jpgpng|140px147x147px]]
| 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.
| align=center |
| align=center |
| valign=top |
|}
 
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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]]