Chimaeras are cartilaginous fish in the order Chimaeriformes //, known informally as ghost sharks, rat fish, spookfish, or rabbit fish; the last three names are not to be confused with rattails, Opisthoproctidae, or Siganidae, respectively.
At one time a "diverse and abundant" group (based on the fossil record), their closest living relatives are sharks and rays, though their last common ancestor with them lived nearly 400 million years ago. Today, they are largely confined to deep water.
Description and habits
Chimaeras live in temperate ocean floors down to 2,600 m (8,500 ft) deep, with few occurring at depths shallower than 200 m (660 ft). Exceptions include the members of the genus Callorhinchus, the rabbit fish and the spotted ratfish, which locally or periodically can be found at shallower depths. Consequently, these are also among the few species from the chimaera order kept in public aquaria. They live in all the oceans except for the Arctic and Antarctic oceans.
They have elongated, soft bodies, with a bulky head and a single gill-opening. They grow up to 150 cm (4.9 ft) in length, although this includes the lengthy tail found in some species. In many species, the snout is modified into an elongated sensory organ.
Like other members of the class Chondrichthyes, chimaera skeletons are composed of cartilage. Their skin is smooth and naked, lacking placoid scales (except in the claspers), and their color can range from black to brownish gray. For defense, most chimaeras have a venomous spine in front of the dorsal fin. They use these fins to "fly" through water.
Chimaeras resemble sharks in some ways: they employ claspers for internal fertilization of females and they lay eggs with leathery cases. They also use electroreception to find their prey. However, unlike sharks, male chimaeras also have retractable sexual appendages on the forehead (a type of tentaculum) and in front of the pelvic fins. The females lay eggs in spindle-shaped, leathery egg cases.
They also differ from sharks in that their upper jaws are fused with their skulls and they have separate anal and urogenital openings. They also have gill covers or opercula like bony fishes. Instead of sharks' many sharp and replaceable teeth, they have just three pairs of large permanent grinding tooth plates. These are unique among vertebrates due to their mineralization. They are made of osteodentin and is covered by an enamel called pleromin formed by mesenchyme-derived cells and made by the mineral whitlockite. Most vertebrates have an enamel of apatite.
Tracing the evolution of these species has been problematic given the paucity of good fossils. DNA sequencing has become the preferred approach to understanding speciation.
The group containing Chimeras and their close relatives (Holocephali) is thought to have originated about 420 million years ago during the Silurian. The oldest known Chimaeriform is Protochimaera from the Early Carboniferous (338-322 million years ago) of Russia, which is more closely related to modern chimeras (Chimaeroidei) than any other known extinct groups of Chimaeriformes. The earliest known remains attributable to modern chimaeras are known from the Early Jurassic (Pleinsbachian) of Europe, but egg cases from the Late Triassic of Yakutia, Russia and New Zealand that resemble those of rhinochimaerids and callorhinchids respectively indicates that they had a global distribution prior to the end of the Triassic. Unlike modern chimaeras, Mesozoic representatives are often found in shallow water settings. The extant species fall into three families—the Callorhinchidae, Rhinochimaeridae and Chimaeridae with the callorhinchids being the most basal clade.
As other fish, chimaeras have a number of parasites. Chimaericola leptogaster (Chimaericolidae) is a monogenean parasite of the gills of Chimaera monstrosa; the species can attain 50 mm (2.0 in) in length.
In some classifications, the chimaeras are included (as subclass Holocephali) in the class Chondrichthyes of cartilaginous fishes; in other systems, this distinction may be raised to the level of class. Chimaeras also have some characteristics of bony fishes.
A renewed effort to explore deep water and to undertake taxonomic analysis of specimens in museum collections led to a boom during the first decade of the 21st century in the number of new species identified. A preliminary study found 8% of species to be threatened. There are 50 extant species in six genera and four families are described; an additional three genera and two families are only known from fossils):
- †Suborder Myriacanthoidei Patterson 1965 (Late Triassic-Late Jurassic)
- †Family Chimaeropsidae
- †Chimaeropsis Zittel 1887 Belgium, Early Jurassic (Sinemurian)
- †Family Myriacanthidae Woodward 1889
- †Acanthorhina Fraas 1910 Posidonia Shale Formation, Germany, Early Jurassic (Toarcian)
- †Agkistracanthus Duffin and Furrer 1981 Austria, England and Switzerland, Late Triassic-Early Jurassic (Rhaetian-Sinemurian)
- †Alethodontus Duffin 1983 Germany, Early Jurassic (Sinemurian)
- †Halonodon Duffin 1984 Belgium and Luxembourg, Early Jurassic (Sinemurian)
- †Metopacanthus Zittel 1887 Posidonia Shale Formation, Germany, Early Jurassic (Toarcian)
- †Oblidens Duffin and Milàn 2017 Hasle Formation, Denmark, Early Jurassic (Pliensbachian)
- †Myriacanthus Agassiz 1837 United Kingdom, Late Triassic-Early Jurassic (Rhaetian-Sinemurian)
- †Recurvacanthus Duffin 1981 United Kingdom, Early Jurassic (Sinemurian)
- †Family Chimaeropsidae
- †Suborder Protochimaeroidei Lebedev & Popov, 2021
- Suborder Chimaeroidei Patterson 1965
- †Eomanodon Ward and Duffin 1989 United Kingdom, Early Jurassic (Pleinsbachian)
- Family Callorhinchidae Garman, 1901
- Genus Callorhinchus Lacépède, 1798
- †Brachymylus A. S. Woodward 1894 Germany, Early Jurassic (Pleinsbachian)
- †Bathytheristes Duffin 1995 Posidonia Shale Formation, Germany, Early Jurassic (Toarcian)
- †Ottangodus Popov, Delsate & Felten, 2019 France, Middle Jurassic (Bajocian)
- Family Chimaeridae Bonaparte, 1831
- Genus Chimaera Linnaeus, 1758
- Chimaera argiloba Last, W. T. White & Pogonoski, 2008 (whitefin chimaera)
- Chimaera bahamaensis Kemper, Ebert, Didier & Compagno, 2010 (Bahamas ghost shark)
- Chimaera compacta Iglésias, Kemper & Naylor, 2022
- Chimaera cubana Howell-Rivero, 1936
- Chimaera fulva Didier, Last & W. T. White, 2008 (southern chimaera)
- Chimaera jordani S. Tanaka (I), 1905 (Jordan's chimaera)
- Chimaera lignaria Didier, 2002 (carpenter's chimaera)
- Chimaera macrospina Didier, Last & W. T. White, 2008 (longspine chimaera)
- Chimaera monstrosa Linnaeus, 1758 (rabbit fish)
- Chimaera notafricana Kemper, Ebert, Compagno & Didier, 2010 Cape chimaera
- Chimaera obscura Didier, Last & W. T. White, 2008 (shortspine chimaera)
- Chimaera opalescens Luchetti, Iglésias & Sellos, 2011
- Chimaera owstoni S. Tanaka (I), 1905 (Owston's chimaera)
- Chimaera panthera Didier, 1998 (leopard chimaera)
- Chimaera phantasma Jordan & Snyder, 1900 (silver chimaera)
- Genus Hydrolagus Gill, 1863
- Hydrolagus affinis Brito Capello, 1868 (smalleyed rabbitfish)
- Hydrolagus africanus Gilchrist, 1922 (African chimaera)
- Hydrolagus alberti Bigelow & Schroeder, 1951
- Hydrolagus alphus Quaranta, Didier, Long & Ebert, 2006 (whitespot ghost shark)
- Hydrolagus barbouri Garman, 1908
- Hydrolagus bemisi Didier, 2002 (pale ghost shark)
- Hydrolagus colliei Lay & E. T. Bennett, 1839 (spotted ratfish)
- Hydrolagus deani H. M. Smith & Radcliffe, 1912 (Philippine chimaera)
- Hydrolagus eidolon Jordan & Hubbs, 1925
- Hydrolagus homonycteris Didier, 2008 (black ghostshark)
- Hydrolagus lemures Whitley, 1939 (blackfin ghostshark)
- Hydrolagus lusitanicus Moura, Figueiredo, Bordalo-Machado, Almeida & Gordo, 2005
- Hydrolagus macrophthalmus de Buen, 1959
- Hydrolagus marmoratus Didier, 2008 marbled ghostshark
- Hydrolagus matallanasi Soto & Vooren, 2004 (striped rabbitfish)
- Hydrolagus mccoskeri Barnett, Didier, Long & Ebert, 2006 (Galápagos ghostshark)
- Hydrolagus melanophasma K. C. James, Ebert, Long & Didier, 2009 (Eastern Pacific black ghostshark)
- Hydrolagus mirabilis Collett, 1904 (large-eyed rabbitfish)
- Hydrolagus mitsukurii Jordan & Snyder, 1904 (spookfish)
- Hydrolagus novaezealandiae Fowler, 1911 (dark ghostshark)
- Hydrolagus ogilbyi Waite, 1898
- Hydrolagus pallidus Hardy & Stehmann, 1990
- Hydrolagus purpurescens Gilbert, 1905 (purple chimaera)
- Hydrolagus trolli Didier & Séret, 2002 (pointy-nosed blue chimaera)
- Hydrolagus waitei Fowler, 1907
- Genus Chimaera Linnaeus, 1758
- Family Rhinochimaeridae Garman, 1901
- Genus Harriotta Goode & Bean, 1895
- Genus Neoharriotta Bigelow & Schroeder, 1950
- Genus Rhinochimaera Garman, 1901
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... Perhaps the most intriguing feature of the newly described species, Hydrolagus melanophasma, is a presumed sexual organ that extends from its forehead called a tentaculum. ...
- A unique mineralization mode of hypermineralized pleromin in the tooth plate of Chimaera phantasma contributes to its microhardness
- Inoue, Jun G.; Miya, Masaki; Lam, Kevin; Tay, Boon-Hui; Danks, Janine A.; Bell, Justin; Walker, Terrence I.; Venkatesh, Byrappa (November 2010). "Evolutionary Origin and Phylogeny of the Modern Holocephalans (Chondrichthyes: Chimaeriformes): A Mitogenomic Perspective". Molecular Biology and Evolution. 27 (11): 2576–2586. doi:10.1093/molbev/msq147. PMID 20551041. Retrieved 14 November 2018.
- Lebedev, Oleg A.; Popov, Evgeny V.; Bagirov, Sergey V.; Bolshiyanov, Igor P.; Kadyrov, Rail I.; Statsenko, Evgeny O. (2021-10-21). "The earliest chimaeriform fish from the Carboniferous of Central Russia". Journal of Systematic Palaeontology. 19 (12): 821–846. doi:10.1080/14772019.2021.1977732. ISSN 1477-2019. S2CID 239509836.
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- Popov, Evgeny V.; Delsate, Dominique; Felten, Roland (2019-07-02). "A New Callorhinchid Genus (Holocephali, Chimaeroidei) from the Early Bajocian of Ottange-Rumelange, on the Luxembourg-French Border". Paleontological Research. 23 (3): 220. doi:10.2517/2018PR021. ISSN 1342-8144. S2CID 198423356.
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- Iglésias, S.P., Kemper, J.M. & Naylor, G.J.P. Chimaera compacta, a new species from southern Indian Ocean, and an estimate of phylogenetic relationships within the genus Chimaera (Chondrichthyes: Chimaeridae). Ichthyol Res 69, 31–45 (2022). https://doi.org/10.1007/s10228-021-00810-9
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