464:
1119:
211:
1149:
1160:
259:
1144:
1126:
1154:
1138:
443:. The tail is long and thin and they move by sweeping movements of the large pectoral fins. The erectile spine in front of the dorsal fin is sometimes venomous. There is no stomach (that is, the gut is simplified and the 'stomach' is merged with the intestine), and the mouth is a small aperture surrounded by lips, giving the head a parrot-like appearance. The only surviving members of the group are the rabbit fish (
230:
25:
1132:
502:
period. The record is extensive, but most fossils are of teeth, and the body forms of numerous species are not known, or at best poorly understood. Some experts further group the orders
463:
518:", and treat it as a sister group to Chimaeriformes. However, as almost all members of Paraselachimorpha are poorly understood, most experts suspect this taxon to be either
2212:
2120:
1701:
973:
415:
invertebrates. They lack a stomach, their food moving directly into the intestine. Extinct holocephalans were much more diverse in lifestyles, including
2081:
1784:
1118:
2107:
544:(the branch of chondrichthyans containing true sharks and rays) about 421 million years ago. Analysis of the 280 million-year-old holocephalian
942:
923:
1706:
864:
108:
1148:
2179:
966:
2217:
46:
2112:
816:"Revealing less derived nature of cartilaginous fish genomes with their evolutionary time scale inferred with nuclear genes"
1711:
89:
1777:
1718:
61:
42:
537:
such as sharks and rays. Under this classification scheme, "Holocephali" would have a much more restricted definition.
2042:
2019:
258:
1159:
998:
959:
403:
Chimaeriformes, commonly known as chimaeras, rat fish, or ghost sharks, include three living families and a little
68:
1153:
35:
2125:
1686:
1696:
315:
210:
1143:
1125:
75:
1770:
1744:
1137:
1959:
636:
393:
period. Little is known about these primitive forms, and the only surviving group in the subclass is the
2156:
2047:
57:
893:
515:
2207:
2135:
2068:
2006:
1739:
1691:
1378:
624:
588:
2184:
600:
576:
914:
1427:
1275:
796:
648:
374:
253:
219:
2161:
2143:
2055:
1882:
1601:
1549:
1506:
1500:
1463:
1451:
1421:
1076:
982:
938:
919:
847:
788:
713:
617:
594:
582:
533:
of holocephalians, i.e. all fish more closely related to living holocephalians than to living
523:
507:
503:
404:
343:
337:
1997:
435:
times, though in other respects they are aberrant. They live close to the bottom and feed on
2171:
2148:
1734:
1645:
1628:
1342:
1257:
837:
827:
780:
700:
Kriwet, Jurgen; Engelbrecht, Andrea; Mors, Thomas; Reguero, Marcelo; Pfaff, Cathrin (2016).
630:
557:
1131:
1920:
1614:
1577:
1457:
1408:
1229:
1031:
702:"Ultimate Eocene (Priabonian) Chondrichthyans (Holocephali, Elasmobranchii) Of Antarctica"
394:
382:
407:
of surviving holocephalans. These fishes move by using sweeping movements of their large
82:
1872:
1852:
1835:
1794:
1589:
1567:
1512:
1494:
1445:
1437:
1414:
1372:
1348:
1093:
842:
815:
666:
654:
606:
541:
534:
511:
485:
397:
378:
355:
349:
331:
300:
246:
2201:
1825:
1538:
1385:
1360:
1309:
1222:
1104:
1084:
768:
642:
546:
491:
451:
440:
408:
164:
2011:
800:
2024:
1905:
1877:
1857:
1762:
1650:
1529:
1469:
1354:
1287:
1182:
769:"Relationships of the Chimaeriformes and the basal radiation of the Chondrichthyes"
411:. They are deep sea fish with slender tails, living close to the seabed to feed on
229:
832:
2094:
2060:
1991:
1887:
1867:
1637:
1623:
1559:
1293:
1248:
1195:
1040:
530:
420:
241:
139:
24:
1915:
1819:
1655:
1554:
1399:
1336:
1281:
1263:
1062:
1053:
784:
529:
184:
149:
1982:
865:
280 million-year-old fossil reveals evolutionary origins of shark-like fishes
792:
717:
1862:
1609:
1366:
1330:
1321:
1235:
519:
469:
432:
270:
189:
133:
851:
431:
Members of this taxon preserve today some features of elasmobranch life in
540:
Based on genetic research, it is estimated the
Holocephali split from the
2034:
1976:
1942:
1897:
1813:
1807:
1572:
1517:
1474:
1269:
1068:
1046:
1008:
499:
479:
445:
436:
390:
290:
200:
179:
174:
159:
154:
144:
701:
1910:
1660:
1200:
1190:
1172:
412:
194:
169:
2099:
1023:
386:
280:
1953:
951:
2086:
550:
demonstrates that early members of the group were more shark-like.
1844:
1479:
462:
416:
993:
1957:
1766:
955:
933:
Pough, F. Harvey; Janis, Christine M.; Heiser, John B. (2013).
18:
2073:
498:
1642:
1597:
1564:
1546:
1396:
1318:
1245:
1212:
1100:
1089:
1080:
1073:
1058:
1036:
1019:
237:
1966:
1933:
1896:
1843:
1834:
1727:
1679:
1588:
1537:
1528:
1436:
1395:
1317:
1308:
1244:
1211:
1181:
1171:
1007:
49:. Unsourced material may be challenged and removed.
754:
730:
767:Lund, Richard; Grogan, Eileen D. (1997-03-01).
757:, pp. 103, 105, Paleozoic Holocephalians.
1778:
967:
369:("complete heads"), sometimes given the name
8:
894:"Chimaeras — The Neglected Chondrichthyans"
467:Various fossil Holocephali. Top to bottom:
1954:
1840:
1785:
1771:
1763:
1534:
1314:
1178:
974:
960:
952:
552:
228:
209:
122:
1921:Myliobatiformes (stingrays and relatives)
841:
831:
109:Learn how and when to remove this message
2213:Extant Early Devonian first appearances
1853:Hexanchiformes (frilled and cow sharks)
692:
875:
814:Renz, AJ; Meyer, A; Kuraku, S (2013).
900:. ReefQuest Centre for Shark Research
773:Reviews in Fish Biology and Fisheries
7:
2136:8ebfe1ff-47e1-474d-a782-062f249ec796
1916:Rajiformes (skates and guitarfishes)
47:adding citations to reliable sources
1873:Heterodontiformes (bullhead sharks)
742:
706:Journal of Vertebrate Paleontology
14:
1883:Carcharhiniformes (ground sharks)
1878:Orectolobiformes (carpet sharks)
1158:
1152:
1147:
1142:
1136:
1130:
1124:
1117:
419:-like predatory forms and slow,
257:
23:
1906:Torpediniformes (electric rays)
389:are of teeth and come from the
34:needs additional citations for
1863:Pristiophoriformes (sawsharks)
937:(9th,international ed.).
892:Martin, R. Aidan; et al.
755:Pough, Janis & Heiser 2013
733:, pp. 99, 101, Table 5-1.
731:Pough, Janis & Heiser 2013
572:†Superorder Paraselachimorpha
16:Subclass of cartilagenous fish
1:
1888:Lamniformes (mackerel sharks)
1868:Squatiniformes (angel sharks)
1858:Squaliformes (dogfish sharks)
613:Superorder Holocephalimorpha
385:Chondrichthyes. The earliest
1719:List of transitional fossils
833:10.1371/journal.pone.0066400
449:), and the elephant fishes (
555:
2236:
1943:Chimaeriformes (chimaeras)
999:Timeline of fish evolution
912:Nelson, Joseph S. (2006).
898:Biology of Sharks and Rays
1802:
1753:
1687:Lists of prehistoric fish
1115:
989:
328:
323:
254:Scientific classification
252:
236:
227:
217:
208:
125:
1911:Pristiformes (sawfishes)
1745:Vertebrate paleontology
785:10.1023/A:1018471324332
556:Taxonomy according to
495:
2218:Vertebrate subclasses
2157:Paleobiology Database
466:
1740:Transitional fossils
1379:Pseudopetalichthyida
678:* position uncertain
637:Chondrenchelyiformes
567:Subclass Holocephali
43:improve this article
915:Fishes of the World
242:Helicoprion davisii
1795:cartilaginous fish
1428:Ischnacanthiformes
1276:Cephalaspidomorphi
649:Cochliodontiformes
496:
379:cartilaginous fish
220:Chimaera monstrosa
2195:
2194:
2144:Open Tree of Life
1960:Taxon identifiers
1951:
1950:
1929:
1928:
1845:Selachii (sharks)
1760:
1759:
1675:
1674:
1671:
1670:
1602:Palaeonisciformes
1550:Onychodontiformes
1507:Petalodontiformes
1501:Iniopterygiformes
1464:Phoebodontiformes
1452:Ctenacanthiformes
1422:Diplacanthiformes
1304:
1303:
1077:Myllokunmingiidae
983:Evolution of fish
944:978-0-321-78235-9
925:978-0-471-25031-9
684:
683:
679:
676:
618:Psammodontiformes
595:Iniopterygiformes
583:Petalodontiformes
524:wastebasket taxon
516:Paraselachimorpha
508:Iniopterygiformes
504:Petalodontiformes
364:
363:
344:Iniopterygiformes
338:Petalodontiformes
319:
119:
118:
111:
93:
2225:
2188:
2187:
2175:
2174:
2165:
2164:
2152:
2151:
2139:
2138:
2129:
2128:
2116:
2115:
2103:
2102:
2090:
2089:
2077:
2076:
2064:
2063:
2051:
2050:
2038:
2037:
2028:
2027:
2015:
2014:
2002:
2001:
2000:
1987:
1986:
1985:
1955:
1841:
1787:
1780:
1773:
1764:
1735:Prehistoric life
1646:Pycnodontiformes
1644:
1629:Acipenseriformes
1599:
1566:
1548:
1535:
1398:
1343:Brindabellaspida
1320:
1315:
1258:Pteraspidomorphi
1247:
1214:
1179:
1163:
1162:
1156:
1151:
1146:
1141:
1140:
1134:
1129:
1128:
1121:
1102:
1091:
1082:
1075:
1060:
1038:
1021:
976:
969:
962:
953:
948:
929:
908:
906:
905:
879:
873:
867:
862:
856:
855:
845:
835:
811:
805:
804:
764:
758:
752:
746:
740:
734:
728:
722:
721:
697:
677:
674:
631:Squalorajiformes
553:
514:into the taxon "
371:Euchondrocephali
314:
262:
261:
239:
232:
213:
199:
136:
129:Temporal range:
123:
114:
107:
103:
100:
94:
92:
51:
27:
19:
2235:
2234:
2228:
2227:
2226:
2224:
2223:
2222:
2198:
2197:
2196:
2191:
2183:
2178:
2170:
2168:
2160:
2155:
2147:
2142:
2134:
2132:
2124:
2119:
2111:
2106:
2098:
2093:
2085:
2080:
2072:
2067:
2059:
2054:
2046:
2041:
2033:
2031:
2023:
2018:
2010:
2005:
1996:
1995:
1990:
1981:
1980:
1975:
1962:
1952:
1947:
1925:
1898:Batoidea (rays)
1892:
1830:
1798:
1791:
1761:
1756:
1749:
1723:
1667:
1615:Polypteriformes
1584:
1578:Tetrapodomorpha
1524:
1458:Hybodontiformes
1432:
1409:Acanthodiformes
1391:
1300:
1240:
1230:Paraconodontida
1207:
1175:
1167:
1166:
1165:
1157:
1135:
1123:
1113:
1032:Cephalochordate
1003:
985:
980:
945:
935:Vertebrate Life
932:
926:
911:
903:
901:
891:
888:
883:
882:
874:
870:
863:
859:
813:
812:
808:
766:
765:
761:
753:
749:
745:, lead section.
741:
737:
729:
725:
699:
698:
694:
689:
662:Incertae sedis
625:Copodontiformes
589:Helodontiformes
461:
429:
427:Characteristics
405:over 50 species
313:
256:
204:
198:
197:
192:
187:
182:
177:
172:
167:
162:
157:
152:
147:
142:
131:
130:
127:
115:
104:
98:
95:
52:
50:
40:
28:
17:
12:
11:
5:
2233:
2232:
2229:
2221:
2220:
2215:
2210:
2200:
2199:
2193:
2192:
2190:
2189:
2176:
2166:
2153:
2140:
2130:
2117:
2104:
2091:
2078:
2065:
2052:
2039:
2029:
2016:
2003:
1988:
1972:
1970:
1964:
1963:
1958:
1949:
1948:
1946:
1945:
1939:
1937:
1931:
1930:
1927:
1926:
1924:
1923:
1918:
1913:
1908:
1902:
1900:
1894:
1893:
1891:
1890:
1885:
1880:
1875:
1870:
1865:
1860:
1855:
1849:
1847:
1838:
1836:Elasmobranchii
1832:
1831:
1829:
1828:
1822:
1816:
1810:
1803:
1800:
1799:
1792:
1790:
1789:
1782:
1775:
1767:
1758:
1757:
1754:
1751:
1750:
1748:
1747:
1742:
1737:
1731:
1729:
1725:
1724:
1722:
1721:
1716:
1715:
1714:
1709:
1704:
1699:
1694:
1683:
1681:
1677:
1676:
1673:
1672:
1669:
1668:
1666:
1665:
1664:
1663:
1658:
1653:
1648:
1634:
1633:
1632:
1631:
1620:
1619:
1618:
1617:
1606:
1605:
1594:
1592:
1586:
1585:
1583:
1582:
1581:
1580:
1575:
1570:
1568:Porolepiformes
1557:
1552:
1543:
1541:
1532:
1526:
1525:
1523:
1522:
1521:
1520:
1515:
1513:Symmoriiformes
1509:
1503:
1497:
1495:Eugeneodontida
1485:
1484:
1483:
1482:
1477:
1472:
1466:
1460:
1454:
1446:Elasmobranchii
1442:
1440:
1434:
1433:
1431:
1430:
1424:
1418:
1415:Climatiiformes
1411:
1404:
1402:
1393:
1392:
1390:
1389:
1382:
1375:
1373:Acanthothoraci
1369:
1363:
1357:
1351:
1349:Petalichthyida
1345:
1339:
1333:
1326:
1324:
1312:
1306:
1305:
1302:
1301:
1299:
1298:
1297:
1296:
1290:
1284:
1272:
1266:
1260:
1253:
1251:
1242:
1241:
1239:
1238:
1232:
1226:
1223:Protoconodonta
1218:
1216:
1209:
1208:
1206:
1205:
1204:
1203:
1193:
1187:
1185:
1176:
1169:
1168:
1116:
1114:
1112:
1111:
1110:
1109:
1098:
1094:Myllokunmingia
1071:
1066:
1051:
1050:
1049:
1044:
1029:
1028:
1027:
1013:
1011:
1005:
1004:
1002:
1001:
996:
990:
987:
986:
981:
979:
978:
971:
964:
956:
950:
949:
943:
930:
924:
909:
887:
884:
881:
880:
868:
857:
806:
759:
747:
735:
723:
691:
690:
688:
685:
682:
681:
672:
671:
670:
669:
667:Symmoriiformes
660:
659:
658:
655:Chimaeriformes
651:
645:
639:
633:
627:
621:
611:
610:
609:
607:Eugeneodontida
603:
601:Debeeriiformes
597:
591:
585:
579:
577:Orodontiformes
569:
568:
562:
561:
542:Elasmobranchii
512:Eugeneodontida
486:Harpagofututor
460:
457:
428:
425:
398:Chimaeriformes
362:
361:
360:
359:
356:Symmoriiformes
352:
350:Eugeneodontida
346:
340:
334:
332:Chimaeriformes
326:
325:
321:
320:
308:
304:
303:
301:Chondrichthyes
298:
294:
293:
288:
284:
283:
278:
274:
273:
268:
264:
263:
250:
249:
234:
233:
225:
224:
215:
214:
206:
205:
193:
188:
183:
178:
173:
168:
163:
158:
153:
148:
143:
138:
137:
128:
117:
116:
99:September 2014
31:
29:
22:
15:
13:
10:
9:
6:
4:
3:
2:
2231:
2230:
2219:
2216:
2214:
2211:
2209:
2206:
2205:
2203:
2186:
2181:
2177:
2173:
2167:
2163:
2158:
2154:
2150:
2145:
2141:
2137:
2131:
2127:
2122:
2118:
2114:
2109:
2105:
2101:
2096:
2092:
2088:
2083:
2079:
2075:
2070:
2066:
2062:
2057:
2053:
2049:
2044:
2040:
2036:
2030:
2026:
2021:
2017:
2013:
2008:
2004:
1999:
1993:
1989:
1984:
1978:
1974:
1973:
1971:
1969:
1965:
1961:
1956:
1944:
1941:
1940:
1938:
1936:
1932:
1922:
1919:
1917:
1914:
1912:
1909:
1907:
1904:
1903:
1901:
1899:
1895:
1889:
1886:
1884:
1881:
1879:
1876:
1874:
1871:
1869:
1866:
1864:
1861:
1859:
1856:
1854:
1851:
1850:
1848:
1846:
1842:
1839:
1837:
1833:
1827:
1826:Gnathostomata
1823:
1821:
1817:
1815:
1811:
1809:
1805:
1804:
1801:
1796:
1788:
1783:
1781:
1776:
1774:
1769:
1768:
1765:
1752:
1746:
1743:
1741:
1738:
1736:
1733:
1732:
1730:
1726:
1720:
1717:
1713:
1710:
1708:
1705:
1703:
1702:cartilaginous
1700:
1698:
1695:
1693:
1690:
1689:
1688:
1685:
1684:
1682:
1678:
1662:
1659:
1657:
1654:
1652:
1649:
1647:
1641:
1640:
1639:
1636:
1635:
1630:
1627:
1626:
1625:
1622:
1621:
1616:
1613:
1612:
1611:
1608:
1607:
1603:
1596:
1595:
1593:
1591:
1587:
1579:
1576:
1574:
1571:
1569:
1563:
1562:
1561:
1558:
1556:
1553:
1551:
1545:
1544:
1542:
1540:
1536:
1533:
1531:
1527:
1519:
1516:
1514:
1510:
1508:
1504:
1502:
1498:
1496:
1492:
1491:
1490:
1487:
1486:
1481:
1478:
1476:
1473:
1471:
1467:
1465:
1461:
1459:
1455:
1453:
1449:
1448:
1447:
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1438:Cartilaginous
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1105:Zhongxiniscus
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1085:Haikouichthys
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779:(1): 65–123.
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558:Joseph Nelson
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547:Dwykaselachus
543:
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535:elasmobranchs
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527:
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494:
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492:Delphyodontos
488:
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475:Heteropetalus
472:
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452:Callorhinchus
448:
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441:invertebrates
438:
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409:pectoral fins
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247:eugeneodontid
244:
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223:, a rat fish
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63:
60: –
59:
58:"Holocephali"
55:
54:Find sources:
48:
44:
38:
37:
32:This article
30:
26:
21:
20:
1967:
1934:
1824:Infraphylum
1692:spiny sharks
1651:Halecomorphi
1488:
1470:Xenacanthida
1355:Phyllolepida
1288:Pituriaspida
1249:Ostracoderms
1183:Cyclostomata
1173:Jawless fish
1122:
1103:
1092:
1083:
1061:
1039:
1022:
934:
913:
902:. Retrieved
897:
886:Bibliography
871:
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776:
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520:paraphyletic
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365:
310:
240:
218:
120:
105:
96:
86:
79:
72:
65:
53:
41:Please help
36:verification
33:
2208:Holocephali
2095:iNaturalist
2025:Holocephali
2012:Holocephali
1998:Holocephali
1992:Wikispecies
1968:Holocephali
1935:Holocephali
1712:lobe-finned
1638:Neopterygii
1624:Chondrostei
1560:Rhipidistia
1539:Lobe-finned
1489:Holocephali
1294:Osteostraci
1196:Hyperoartia
1041:Cathaymyrus
1009:Forerunners
876:Nelson 2006
712:(4): 1–19.
657:(chimaeras)
531:total group
421:durophagous
367:Holocephali
311:Holocephali
132:416–0
126:Holocephali
2202:Categories
1820:Vertebrata
1818:Subphylum
1697:placoderms
1656:Ginglymodi
1590:Ray-finned
1555:Coelacanth
1400:Acanthodii
1337:Arthrodira
1322:Placoderms
1310:Jawed fish
1282:Galeaspida
1264:Thelodonti
1063:Haikouella
1054:Olfactores
904:2016-02-22
687:References
439:and other
307:Subclass:
69:newspapers
1755:† extinct
1610:Cladistia
1367:Rhenanida
1331:Antiarchi
1236:Conodonta
1215:Conodonts
793:1573-5184
718:0272-4634
675:† Extinct
470:Cobelodus
459:Evolution
433:Paleozoic
316:Bonaparte
277:Kingdom:
271:Eukaryota
2032:BioLib:
1977:Wikidata
1814:Chordata
1808:Animalia
1806:Kingdom
1573:Lungfish
1518:Chimaera
1270:Anaspida
1069:Tunicate
1047:Lancelet
852:23825540
820:PLOS ONE
801:40689320
500:Devonian
480:Falcatus
446:Chimaera
437:molluscs
391:Devonian
375:subclass
291:Chordata
287:Phylum:
281:Animalia
267:Domain:
201:Devonian
1983:Q149104
1812:Phylum
1793:Extant
1728:Related
1661:Teleost
1201:Lamprey
1191:Hagfish
843:3692497
665:†Order
647:†Order
641:†Order
635:†Order
629:†Order
623:†Order
616:†Order
605:†Order
599:†Order
593:†Order
587:†Order
581:†Order
575:†Order
560:, 2006
413:benthic
387:fossils
381:in the
373:, is a
324:Orders
297:Class:
203:–Recent
83:scholar
2172:113873
2169:uBio:
2162:100271
2149:550652
2133:NZOR:
2113:161008
1797:orders
1024:Pikaia
941:
922:
850:
840:
799:
791:
743:Martin
716:
653:Order
510:, and
489:, and
423:fish.
318:, 1832
85:
78:
71:
64:
56:
2185:10196
2180:WoRMS
2100:60450
2048:34231
2035:15393
1680:Lists
1480:Shark
797:S2CID
522:or a
417:shark
395:order
383:class
90:JSTOR
76:books
2126:7863
2121:NCBI
2108:ITIS
2082:GBIF
2074:8901
2043:BOLD
1707:bony
1530:Bony
994:Fish
939:ISBN
920:ISBN
848:PMID
789:ISSN
714:ISSN
245:, a
140:PreꞒ
62:news
2087:120
2069:EoL
2056:CoL
2020:AFD
2007:ADW
1475:Ray
1079:? (
838:PMC
828:doi
781:doi
455:).
400:.
377:of
45:by
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2061:CK
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1994::
1979::
1413:†"
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918:.
896:.
846:.
836:.
822:.
818:.
795:.
787:.
775:.
771:.
710:36
708:.
704:.
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506:,
483:,
477:,
473:,
190:Pg
134:Ma
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1779:t
1772:v
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1604:"
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1598:†
1565:†
1547:†
1511:†
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907:.
878:.
854:.
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783::
777:7
720:.
620:*
358:?
354:†
348:†
342:†
336:†
238:†
195:N
185:K
180:J
175:T
170:P
165:C
160:D
155:S
150:O
145:Ꞓ
112:)
106:(
101:)
97:(
87:·
80:·
73:·
66:·
39:.
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