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Hexanchiformes

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Hexanchiformes
Temporal range: Early Jurassic–Recent[1] Possible Permian record
Broadnose sevengill shark (Notorynchus cepedianus)
Frilled shark (Chlamydoselachus anguineus)
Scientific classification Edit this classification
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Class: Chondrichthyes
Subclass: Elasmobranchii
Division: Selachii
Superorder: Squalomorphi
Clade: Hexanchida
Order: Hexanchiformes
F. de Buen, 1926
Families

see text

The Hexanchiformes /hɛkˈsæŋkɪfɔːrmiːz/ are a primitive order of sharks,[a] numbering just seven extant species in two families. Fossil sharks that were apparently very similar to modern sevengill species are known from Jurassic specimens.[2]

Taxonomy

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Due to their primitive anatomy, hexanchiforms were previously considered the most basal group of sharks. However, more recent phylogenetic studies indicate that while primitive, they in fact belong to the superorder Squalomorphii, which also contains dogfishes, angelsharks, and sawsharks, although they are thought to be the most basal member of the group.[3][4]

Description

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Hexanchiform sharks have one spineless dorsal fin located over or behind the pelvic fins and one anal fin. The vertebral column extends into the long dorsal lobe of the caudal fin, while the ventral lobe is either small or absent. They have either six or seven gill slits, located in front of the pectoral fins. They have a large mouth, with eyes on either side of the head. The spiracles are small and located well above and behind the eyes.[5] The eyes have no nictitating membrane.

The frilled sharks of the genus Chlamydoselachus are very different from the cow sharks, and have been proposed to be moved to a distinct order, Chlamydoselachiformes. However, genetic studies have found them to be each others' closest relatives, and they share certain derived features supporting them both being in the same order.[3][4]

Shark teeth similar to modern hexanchids and echinorhinids are known from Devonian deposits in Antarctica and Australia, as well as Permian deposits in Japan. If these are in fact hexanchids, this may be the only extant order of elasmobranchs to have survived the Permian extinction (and by extension, the oldest extant order of elasmobranchs). However, the Australian/Antarctic shark teeth, from the family Mcmurdodontidae, have also been found to lack a multilayer enameloid layer covering the tooth crown, something found in all modern sharks and most Devonian sharks, indicating that they are neoselachians of uncertain affinity or even indeterminate chondrichthyans. The occurrence of derived sharks in the Devonian is also irreconcilable with the results of all phylogenetic estimates in the group.[6][7][8]

It is debated whether the extinct families Orthacodontidae and Paraorthacodontidae belong to the Hexanchiformes or the extinct Synechodontiformes. However, the Shark-References database currently lists them as members of the Hexanchiformes.[9][10][11]

Distribution

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Species are widespread and found across most of the world. They are most common in cold deep water in the tropics, but are also found closer to the shore in more temperate regions.[5]

Classification

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Living species

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Extinct species

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Notidanodon sp. fossil at the Geological Museum, Copenhagen
  • Suborder Chlamydoselachoidi
    • Family Chlamydoselachidae
      • Chlamydoselachus Garman, 1884
        • Chlamydoselachus balli Cappetta, Morrison & Adnet, 2019
        • Chlamydoselachus gracilis Antunes & Cappetta, 2002
        • Chlamydoselachus lawleyi Davis, 1887
        • Chlamydoselachus tobleri Leriche, 1929
      • Dykeius Cappetta, Morrison & Adnet, 2019
        • Dykeius garethi Cappetta, Morrison & Adnet, 2019
      • Rolfodon Cappetta, Morrison & Adnet, 2019
        • Rolfodon bracheri (Pfeil, 1983)
        • Rolfodon fiedleri (Pfeil, 1983)
        • Rolfodon goliath (Antunes & Cappetta, 2002)
        • Rolfodon keyesi (Mannering & Hiller 2008)
        • Rolfodon landinii (Carrillo-Briceño et al. 2014)
        • Rolfodon ludvigseni Cappetta, Morrison & Adnet, 2019
        • Rolfodon tatere (Consoli, 2008)
        • Rolfodon thomsoni (Richter & Ward, 1990)
  • Suborder Hexanchoidei
    • Family Crassodontidanidae
      • Crassodontidanus Kriwet & Klug, 2011
        • Crassodontidanus serratus Fraas, 1855
        • Crassodontidanus wiedenrothi Thies, 1983
      • Notidanoides Maisey, 1986
        • Notidanoides muensteri Agassiz, 1843
      • Notidanus Cuvier, 1816
        • Notidanus amalthei Oppel, 1854
        • Notidanus atrox Ameghino, 1899
        • Notidanus intermedius Wagner, 1862
        • Notidanus nikitini Chabakov & Zonov, 1935
      • Pachyhexanchus Cappetta, 1990
        • Pachyhexanchus pockrandti Ward & Thies, 1987
    • Family Hexanchidae
      • Gladioserratus Underwood, Goswami, Prasad, Verma & Flynn, 2011
        • Gladioserratus aptiensis Pictet, 1864
        • Gladioserratus dentatus Guinot, Cappetta & Adnet, 2014
        • Gladioserratus magnus Underwood, Goswami, Prasad, Verma & Flynn, 2011
      • Heptranchias Rafinesque, 1810
        • Heptranchias ezoensis Applegate & Uyeno, 1968
        • Heptranchias howellii Reed, 1946
        • Heptranchias karagalensis Kozlov in Zhelezko & Kozlov, 1999
        • Heptranchias tenuidens Leriche, 1938
      • Hexanchus Rafinesque, 1810
        • Hexanchus agassizi Cappetta, 1976
        • Hexanchus andersoni Jordan, 1907
        • Hexanchus casieri Kozlov, 1999
        • Hexanchus collinsonae Ward, 1979
        • Hexanchus gracilis Davis, 1887
        • Hexanchus hookeri Ward, 1979
        • Hexanchus microdon Agassiz, 1843
        • Hexanchus tusbairicus Kozlov in Zhelezko & Kozlov, 1999
      • Notidanodon Cappetta, 1975
        • Notidanodon lanceolatus Woodward, 1886
        • Notidanodon pectinatus Agassiz, 1843
      • Notorynchus Ayres, 1855
        • Notorynchus borealus Jordan & Hannibal, 1923
        • Notorynchus kempi Ward, 1979
        • Notorynchus lawleyi Cigala Fulgosi, 1983
        • Notorynchus primigenius Agassiz, 1843
        • Notorynchus serratissimus Agassiz, 1843
        • Notorynchus subrecurvus Oppenheimer, 1907
      • Pachyhexanchus Cappetta, 1990
        • Pachyhexanchus pockrandti Ward & Thies, 1987
      • Paraheptranchias Pfeil, 1981
        • Paraheptranchias repens Probst, 1879
      • Pseudonotidanus Underwood & Ward, 2004
        • Pseudonotidanus semirugosus Underwood & Ward, 2004
      • Welcommia Cappetta, 1990
      • Weltonia Ward, 1979
        • Weltonia ancistrodon Arambourg, 1952
        • Weltonia burnhamensis Ward, 1979
      • Xampylodon Cappetta, Morrison & Adnet, 2019
        • Xampylodon brotzeni (Siverson, 1995)
        • Xampylodon dentatus (Woodward, 1886)
        • Xampylodon loozi (Vincent, 1876)
    • ?Family Orthacodontidae
      • Occitanodus Guinot, Cappetta & Adnet, 2014
        • Occitanodus sudrei Guinot, Cappetta & Adnet, 2014
      • Sphenodus Agassiz, 1843
        • Sphenodus alpinus Gümbel, 1861
        • Sphenodus longidens Agassiz, 1843
        • Sphenodus lundgreni Davis, 1890
        • Sphenodus macer Quenstedt, 1852
        • Sphenodus nitidus Wagner, 1862
        • Sphenodus longidens Agassiz, 1843
        • Sphenodus planus Agassiz, 1843
        • Sphenodus rectidens Emmons, 1858
        • Sphenodus robustidens Seguenza, 1900
        • Sphenodus tithonius Gemmellaro, 1871
        • Sphenodus virgai Gemmellaro, 1871
  • ?Family Komoksodontidae Cappetta, Morrison & Adnet, 2019[14]
    • Komoksodon Cappetta, Morrison & Adnet, 2019
      • Komoksodon kwutchakutch Cappetta, Morrison & Adnet, 2019
  • ?Family Paraorthacodontidae
    • Macrourogaleus Fowler, 1947
      • Macrourogaleus hassei
    • Paraorthacodus Glückman, 1957
      • Paraorthacodus andersoni (Case, 1978)
      • Paraorthacodus antarcticus Klug, Kriwet, Lirio & Nuñez, 2008
      • Paraorthacodus arduennae Delsate, 2001
      • Paraorthacodus clarkii (Eastman, 1901)
      • Paraorthacodus conicus (Davis, 1890)
      • Paraorthacodus eocaenus (Leriche, 1902)
      • Paraorthacodus jurensis (Schweizer, 1964)
      • Paraorthacodus recurvus (Trautschold, 1877)
      • Paraorthacodus rossi Cappetta, Morrison & Adnet, 2019
      • Paraorthacodus turgaicus Glikman, 1964

Species

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Family Image Common name Genera Species Description
Chlamydoselachidae Frilled sharks 1
extant
1
extinct 		2
extant
12
extinct 		Frilled sharks contain only two extant species of deepsea creatures which are typically weakened in areas closer to the surface. The most widely known species still surviving is the frilled shark, known as a living fossil, along with the Southern African frilled shark, found along coastal areas of South Africa. Several extinct species are known.
†Crassodontidanidae †Crassodontidanidae 4 8 Extinct
Hexanchidae Cow sharks 3
extant
5
extinct 		5
extant
31
extinct 		Cow sharks are considered the most primitive of all the sharks, because their skeletons resemble those of ancient extinct forms, with few modern adaptations. Their excretory and digestive systems are also unspecialised, suggesting that they may also resemble those of their primitive shark ancestors. Their most distinctive feature, however, is the presence of a sixth, and, in two genera, a seventh, gill slit, in addition to the five found in all other sharks.[15] They range from 1.4 metres (4.6 ft) to over 5.5 metres (18 ft) in adult body length.
†Komoksodontidae? †Komoksodontidae? 1 1 Extinct
†Orthacodontidae? †Orthacodontidae? 2 12 Extinct
†Paraorthacodontidae? †Paraorthacodontidae? 2 11 Extinct

See also

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Footnotes

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  1. ^ The Chimaeras (Holocephali) are arguably more primitive than the Hexanchiformes, but arguably may not be sharks, depending on whether "sharks" are taken to be superorder Selachimorpha or instead class Chondrichthyes (all cartilaginous fish).

References

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  1. ^ Froese, Rainer; Pauly, Daniel (eds.). "Order Hexanchiformes". FishBase. January 2009 version.
  2. ^ Allen, Thomas B. (1999). The Shark Almanac. New York: The Lyons Press. p. 45. ISBN 1-55821-582-4.
  3. ^ a b Vélez-Zuazo, Ximena; Agnarsson, Ingi (2011-02-01). "Shark tales: A molecular species-level phylogeny of sharks (Selachimorpha, Chondrichthyes)". Molecular Phylogenetics and Evolution. 58 (2): 207–217. doi:10.1016/j.ympev.2010.11.018. ISSN 1055-7903.
  4. ^ a b Tanaka, Keiko; Shiina, Takashi; Tomita, Taketeru; Suzuki, Shingo; Hosomichi, Kazuyoshi; Sano, Kazumi; Doi, Hiroyuki; Kono, Azumi; Komiyama, Tomoyoshi; Inoko, Hidetoshi; Kulski, Jerzy K.; Tanaka, Sho (2013-09-05). "Evolutionary Relations of Hexanchiformes Deep-Sea Sharks Elucidated by Whole Mitochondrial Genome Sequences". BioMed Research International. 2013: e147064. doi:10.1155/2013/147064. ISSN 2314-6133. PMC 3780621.
  5. ^ a b Compagno, Leonard J. V.; Dando, Marc; Fowler, Sarah L.; Compagno, Leonard; Fowler, Sarah (2005). A field guide to the Sharks of the world. Collins field guide. London: Collins. ISBN 978-0-00-713610-0.
  6. ^ Adnet, S.; Guinot, G.; Cappetta, H.; Welcomme, J.-L. (2012). "Oldest evidence of bramble sharks (Elasmobranchii, Echinorhinidae) in the Lower Cretaceous of southeast France and the evolutionary history of orbitostylic sharks". Cretaceous Research. 35: 81–87. doi:10.1016/j.cretres.2011.11.021. ISSN 0195-6671.
  7. ^ Klug, Stefanie; Kriwet, Jürgen (2013). "Node age estimations and the origin of angel sharks, Squatiniformes (Neoselachii, Squalomorphii)". Journal of Systematic Palaeontology. 11 (1): 91–110. doi:10.1080/14772019.2012.674066. ISSN 1477-2019.
  8. ^ Long, John; Thomson, Victoria; Burrow, Carole; Turner, Susan (2021-10-28), Pradel, Alan; Denton, John S.S.; Janvier, Philippe (eds.), "Fossil chondrichthyan remains from the Middle Devonian Kevington Creek Formation, South Blue Range, Victoria" (PDF), Ancient Fishes and their Living Relatives, Munich, Germany: Verlag, Dr Friedrich Pfeil, pp. 239–245, ISBN 978-3-89937-269-4, retrieved 2023-11-30
  9. ^ Cappetta, Henri; Morrison, Kurt; Adnet, Sylvain (2019-12-10). "A shark fauna from the Campanian of Hornby Island, British Columbia, Canada: an insight into the diversity of Cretaceous deep-water assemblages". Historical Biology. 33 (8): 1121–1182. doi:10.1080/08912963.2019.1681421. ISSN 0891-2963.
  10. ^ "Sphenodus hybodoides | Shark-References". shark-references.com. Retrieved 2023-12-01.
  11. ^ "Paraorthacodus andersoni | Shark-References". shark-references.com. Retrieved 2023-12-01.
  12. ^ "New shark species confirmed: Genetic testing finds a different sixgill shark". ScienceDaily. Retrieved 2020-08-21.
  13. ^ "New species of shark discovered through genetic testing". phys.org. Retrieved 2020-08-21.
  14. ^ Cappetta, Henri; Morrison, Kurt; Adnet, Sylvain (2019-12-10). "A shark fauna from the Campanian of Hornby Island, British Columbia, Canada: an insight into the diversity of Cretaceous deep-water assemblages". Historical Biology. 33 (8): 1121–1182. doi:10.1080/08912963.2019.1681421. ISSN 0891-2963.
  15. ^ Matt's, J. & Last P.R. (1998). Paxton, J.R. & Eschmeyer, W.N. (eds.). Encyclopedia of Fishes. San Diego: Academic Press. p. 61. ISBN 0-12-547665-5.
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