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Holocephali Temporal range: Late Devonian–Recent PreꞒ Ꞓ O S D C P T J K Pg N
Chimaera monstrosa, a rat fish
Life r esotoration of Romerodus, a eugeneodontid
Scientific classification
Domain:	Eukaryota
Kingdom:	Animalia
Phylum:	Chordata
Class:	Chondrichthyes
Subclass:	Holocephali Bonaparte, 1832
Orders
Holocephali sensu lato (Euchondrocephali) †Petalodontiformes †Iniopterygiformes †Eugeneodontida †Symmoriiformes? Holocephali sensu stricto †Cochliodontiformes †Menaspiformes †Chondrenchelyiformes Chimaeriformes

Holocephali ("complete heads"), sometimes given the name Euchondrocephali, is a subclass of cartilaginous fish in the class Chondrichthyes. The earliest fossils are of teeth and come from the Devonian period. The only living holocephalans are the chimaeras (Chimaeriformes), though during the late Paleozoic (Carboniferous and Permian) Holocephali was much more diverse, including an array of forms including those considerably different from modern Chimaeriformes, including shark-like predatory forms and slow, durophagous fish.

Chimaeras, also known as rat fish, or ghost sharks, include three living families and a little over 50 species of surviving holocephalans. These fishes move by using sweeping movements of their large pectoral fins. They are deep sea fish with slender tails, living close to the seabed to feed on benthic invertebrates. They lack a stomach, their food moving directly into the intestine.

Characteristics

Members of this taxon preserve today some features of elasmobranch life in Paleozoic times, though in other respects they are aberrant. They live close to the bottom and feed on molluscs and other invertebrates. 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 (Chimaera), and the elephant fishes (Callorhinchus).

Evolution

The fossil record of the Holocephali starts during the Devonian 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 Petalodontiformes, Iniopterygiformes, and Eugeneodontida into the taxon "Paraselachimorpha", 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 paraphyletic or a wastebasket taxon.

Lund & Grogan (1997) coined the subclass Euchondrocephali to refer to the total group of holocephalians, i.e. all fish more closely related to living holocephalians than to living elasmobranchs such as sharks and rays. Under this classification scheme, "Holocephali" would have a much more restricted definition. Other authors have used Holocephali in a broad sense covering all fish more closely related to chimaeras than to sharks and rays.

Based on genetic research, it is estimated the Holocephali split from the Elasmobranchii (the branch of chondrichthyans containing true sharks and rays) about 421 million years ago.

While historically considered closely related to elasmobranchs, recent studies have found members of the shark-like "symmoriiformes", which may be paraphyletic, to be early diverging relatives of Holocephali, (with some of these studies choosing to exclude them from Holocephali proper).

References

1. Kriwet, Jurgen; Engelbrecht, Andrea; Mors, Thomas; Reguero, Marcelo; Pfaff, Cathrin (2016). "Ultimate Eocene (Priabonian) Chondrichthyans (Holocephali, Elasmobranchii) Of Antarctica". Journal of Vertebrate Paleontology. 36 (4): e1160911. doi:10.1080/02724634.2016.1160911. hdl:11336/54351. ISSN 0272-4634. JSTOR 24740249.
2. Pough, Janis & Heiser 2013, pp. 99, 101, Table 5-1.
3. Martin, lead section.
4. Pough, Janis & Heiser 2013, pp. 103, 105, Paleozoic Holocephalians.
5. Lund, Richard; Grogan, Eileen D. (1997-03-01). "Relationships of the Chimaeriformes and the basal radiation of the Chondrichthyes". Reviews in Fish Biology and Fisheries. 7 (1): 65–123. doi:10.1023/A:1018471324332. ISSN 1573-5184. S2CID 40689320.
6. Nelson, Joseph S.; Grande, Terry C.; Wilson, Mark V. H. (2016-02-22). Fishes of the World (1 ed.). Wiley. pp. 47–49. doi:10.1002/9781119174844. ISBN 978-1-118-34233-6.
7. Renz, AJ; Meyer, A; Kuraku, S (2013). "Revealing less derived nature of cartilaginous fish genomes with their evolutionary time scale inferred with nuclear genes". PLOS ONE. 8 (6): e66400. doi:10.1371/journal.pone.0066400. PMC 3692497. PMID 23825540.
8. Klug, Christian; Coates, Michael; Frey, Linda; Greif, Merle; Jobbins, Melina; Pohle, Alexander; Lagnaoui, Abdelouahed; Haouz, Wahiba Bel; Ginter, Michal (December 2023). "Broad snouted cladoselachian with sensory specialization at the base of modern chondrichthyans". Swiss Journal of Palaeontology. 142 (1). doi:10.1186/s13358-023-00266-6. ISSN 1664-2376. PMC 10050047. PMID 37009301.
9. Nelson 2006.

Bibliography

• Martin, R. Aidan; et al. "Chimaeras — The Neglected Chondrichthyans". Biology of Sharks and Rays. ReefQuest Centre for Shark Research. Retrieved 2016-02-22.
• Nelson, Joseph S. (2006). Fishes of the World. Wiley. ISBN 978-0-471-25031-9.
• Pough, F. Harvey; Janis, Christine M.; Heiser, John B. (2013). Vertebrate Life (9th,international ed.). Pearson. ISBN 978-0-321-78235-9.

v t e Evolution of fish
Fish Timeline of fish evolution
Forerunners	Basal member †Pikaia Cephalochordate †Cathaymyrus Lancelet Olfactores †Haikouella Tunicate †Myllokunmingiidae? (†Haikouichthys, †Myllokunmingia) †Zhongxiniscus?
Jawless fish	Cyclostomata Hagfish Hyperoartia Lamprey †Conodonts †Protoconodonta? †Paraconodontida †Conodonta †Ostracoderms †Pteraspidomorphi †Thelodonti †Anaspida †Cephalaspidomorphi †Galeaspida †Pituriaspida †Osteostraci
Jawed fish	†Placoderms †Antiarchi †Arthrodira †Brindabellaspida †Petalichthyida †Phyllolepida †Ptyctodontida †Rhenanida †Acanthothoraci †Pseudopetalichthyida? †Stensioellida? †Acanthodii †Acanthodiformes †"Climatiiformes" †Diplacanthiformes †Ischnacanthiformes Cartilaginous Elasmobranchii †Ctenacanthiformes †Hybodontiformes †Phoebodontiformes †Xenacanthida Ray Shark Holocephali †Eugeneodontida †Iniopterygiformes †Petalodontiformes †Symmoriiformes Chimaera Bony Lobe-finned †Onychodontiformes Coelacanth Rhipidistia †Porolepiformes Lungfish Tetrapodomorpha Ray-finned †"Palaeonisciformes" Cladistia Polypteriformes Chondrostei Acipenseriformes Neopterygii †Pycnodontiformes Halecomorphi Ginglymodi Teleost
Lists	Lists of prehistoric fish spiny sharks placoderms cartilaginous bony lobe-finned List of transitional fossils
Related	Prehistoric life Transitional fossils Vertebrate paleontology
† extinct

• Holocephali
• Extant Early Devonian first appearances
• Vertebrate subclasses