Carangiformes

Carangiformes; Temporal range: Late Paleocene–present PreꞒ Ꞓ O S D C P T J K Pg N
	Bluefin trevally (Caranx melampygus)
	Remora remora
Scientific classification
Kingdom:	Animalia
Phylum:	Chordata
Class:	Actinopterygii
Clade:	Percomorpha
Order:	Carangiformes; Jordan, 1923
Type species
	Caranx praeustus ; Anonymous [ Bennett ], 1830;

Carangiformes is a large and diverse order of ray-finned fishes within the clade Percomorpha. It is part of a sister clade to the Ovalentaria, alongside its sister group, the Anabantaria (including Anabantiformes and Synbranchiformes). The order includes ecologically diverse groups such as the jacks and trevallies, flatfishes, barracudas, billfishes, and archerfishes.

The Carangiformes have been long regarded as a monotypic order with only the family Carangidae within it by some authorities, and the other current families within the order have been previously classified as part of the wider order Perciformes. The 5th edition of Fishes of the World classify six families within Carangiformes,^[2] with more recent authorities expanding the order to include up to 30 families, based on phylogenetic evidence.^[3]

Characteristics

While the expanded order Carangiformes is primarily defined by molecular data, the core group (suborder Carangoidei) shares specific synapomorphies:

One or two tubular ossifications (prenasals) extending from the nasal bone, a trait shared with the archerfishes.^[4]
Small, adherent cycloid scales.^[4]

Most carangiforms are medium to large-sized carnivorous predators, ranging from 22 cm (8.7 in) to 2.5 m (8.2 ft) in length. Body shapes vary from slender and fusiform (e.g., barracudas, cobia) to deep-bodied and laterally compressed (e.g., trevallies, moonfish, flatfishes). All species are marine, inhabiting primarily tropical and subtropical waters.

Taxonomy and phylogeny

The order Carangiformes has historically been either subsumed within Perciformes or used exclusively for the families in the suborder Carangoidei (Carangidae, Coryphaenidae, Rachycentridae, Echeneidae, and Nematistiidae). However, recent genetic studies have redefined the group to resolve the paraphyly of Perciformes, incorporating many more groups such as the highly specialized flatfishes.

The earliest known carangiforms are species of the moonfish genus Mene from the Late Paleocene of Peru and Tunisia.^[5]

Internal relationships of Carangoidei

Within the suborder Carangoidei, the family Carangidae (jacks) is paraphyletic. Two of its subfamilies (Naucratinae and Caranginae) are more closely related to the "Echeneoidea" clade (cobias, remoras, and dolphinfishes) than they are to the other carangid subfamilies (Scomberoidinae and Trachinotinae). This relationship is illustrated in the cladogram below based on Girard et al. (2020):^[6]

Carangoidei

Carangidae (part)

	Naucratinae (amberjacks)

	Caranginae (jacks, trevallies)

Trachinotidae

	Scomberoidinae (queenfish, leatherjackets)

	Trachinotinae (pompanos)

Echeneoidea

Coryphaenidae (dolphinfishes)

	Rachycentridae (cobia)

	Echeneidae (remoras)

Classification

The following classification follows Eschmeyer's Catalog of Fishes (2025):^[7]

Order Carangiformes
- Suborder Centropomoidei
  - Family Latidae Jordan 1888 (giant perches)
  - Family Centropomidae Poey 1967 (snooks)
  - Family Lactariidae Boulenger 1904 (false trevallies)
  - Family Sphyraenidae Rafinesque 1815 (barracudas)
- Suborder Pleuronectoidei
  - Family Polynemidae Rafinesque 1815 (threadfins or tassel-fishes)
  - Family Psettodidae Regan 1910 (spiny turbots)
  - Family Citharidae de Buen 1935 (largescale flounders)
  - Family Scophthalmidae Chabanaud 1933 (turbots)
  - Family Cyclopsettidae Campbell et al. 2019 (sand whiffs or large-tooth flounders)
  - Family Bothidae Smitt 1892 (lefteye flounders)
  - Family Paralichthyidae Regan 1910 (sand flounders)
  - Family Pleuronectidae Rafinesque 1815 (righteye flounders)
    - Subfamily Atheresthinae Vinnikov, Thomson & Munroe 2018
    - Subfamily Pleuronichthyinae Vinnikov, Thomson & Munroe 2018
    - Subfamily Microstominae Cooper & Chapleau 1998 (smallmouth flounders)
    - Subfamily Hippoglossinae Gill 1864 (halibuts)
    - Subfamily Pleuronectinae Rafinesque 1815 (true flounders)
  - Family Paralichthodidae Regan 1920 (peppered flounders)
  - Family Oncopteridae Jordan & Goss 1889 (remo flounders)
  - Family Rhombosoleidae Regan 1910 (South Pacific flounders)
  - Family Achiropsettidae Heemstra 1990 (southern flounders or armless flounders)
  - Family Achiridae Rafinesque 1815 (American soles)
  - Family Samaridae Jordan & Goss 1889 (crested flounders)
  - Family Poecilopsettidae Norman 1934 (bigeye flounders)
  - Family Soleidae Bonaparte 1833 (soles)
  - Family Cynoglossidae Jordan 1888 (tonguefishes)
    - Subfamily Symphurinae Ochiai 1963 (straightsnout tongue soles)
    - Subfamily Cynoglossinae Jordan 1888 (hookedsnout tongue soles)
- Suborder Toxotoidei
  - Family Leptobramidae Ogilby 1913 (beachsalmons)
  - Family Toxotidae Bleeker 1859 (archerfishes)
- Suborder Nematistioidei
  - Family Nematistiidae Gill 1862 (roosterfishes)
- Suborder Menoidei
  - Family Menidae Fitzinger 1873 (moonfishes)
  - Family Xiphiidae Rafinesque 1815 (swordfishes)
  - Family Istiophoridae Rafinesque 1815 (billfishes and marlins)
- Suborder Carangoidei
  - Family Carangidae Rafinesque 1815 (jacks or jack mackerels)
    - Subfamily Naucratinae Bleeker 1859 (amberjacks)
    - Subfamily Caranginae Rafinesque 1815 (trevallies and kingfishes)
    - Subfamily Scomberoidinae Jordan & Gilbert 1883 (leatherjackets and queenfishes)
    - Subfamily Trachinotinae Gill 1861 (pompanos)
  - Family Echeneidae Rafinesque 1810 (remoras and sharksuckers)
  - Family Rachycentridae Gill 1896 (cobias)
  - Family Coryphaenidae Rafinesque 1815 (dolphinfishes)

The Coryphaenidae, Rachycentridae, and Echeneidae have been suggested to comprise a monophyletic grouping, which has been recovered as a sister clade to the Carangidae.^[8] A basal member of this clade is thought to be from the early Eocene.

The following fossil families are also known:^[9]

Order Carangiformes
- ?Family †Pygaeidae Jordan, 1905
- Suborder Pleuronectoidei
  - Family †Amphistiidae Boulenger, 1902
  - Family †Joleaudichthyidae Chabanaud, 1937
- Suborder Menoidei
  - Family †Palaeorhynchidae Günther, 1880
  - Family †Hemingwayidae Sytchevskaya & Prokofiev, 2002
  - Family †Blochiidae Bleeker, 1859
  - Family †Xiphiorhynchidae Regan, 1909^[10]
- Suborder Carangoidei
  - Family †Ductoridae Blot, 1969^[11]
  - Family †Opisthomyzonidae Jordan, 1923

References

^ "Taxon: Order Carangiformes Jordan, 1923 (fish)". Taxonomicon. Retrieved 15 November 2019.
^ J. S. Nelson; T. C. Grande; M. V. H. Wilson (2016). Fishes of the World (5th ed.). Wiley. pp. 380–383. ISBN 978-1-118-34233-6. Archived from the original on 2019-04-08. Retrieved 2019-11-15.
^ Girard, Matthew G.; Davis, Matthew P.; Smith, W. Leo (2020-05-08). "The Phylogeny of Carangiform Fishes: Morphological and Genomic Investigations of a New Fish Clade". Copeia. 108 (2): 265. doi:10.1643/CI-19-320. ISSN 0045-8511.
^ ^a ^b E. O. Wiley & G. David Johnson: A teleost classification based on monophyletic groups. Joseph S. Nelson, Hans-Peter Schultze & Mark V. H. Wilson: Origin and Phylogenetic Interrelationships of Teleosts. Verlag Dr. Friedrich Pfeil, München 2010, ISBN 978-3-89937-107-9
^ Friedman, Matt; V. Andrews, James; Saad, Hadeel; El-Sayed, Sanaa (2023-06-16). "The Cretaceous–Paleogene transition in spiny-rayed fishes: surveying "Patterson's Gap" in the acanthomorph skeletal record André Dumont medalist lecture 2018". Geologica Belgica. doi:10.20341/gb.2023.002. ISSN 1374-8505.
^ Girard, Matthew G.; Davis, Matthew P.; Smith, W. Leo (2020-05-08). "The Phylogeny of Carangiform Fishes: Morphological and Genomic Investigations of a New Fish Clade". Copeia. 108 (2): 265. doi:10.1643/CI-19-320. ISSN 0045-8511.
^ "Eschmeyer's Catalog of Fishes Classification - California Academy of Sciences". www.calacademy.org. Retrieved 2024-12-27.
^ J. S. Nelson; T. C. Grande; M. V. H. Wilson (2016). Fishes of the World (5th ed.). Wiley. pp. 380–383. ISBN 978-1-118-34233-6. Archived from the original on 2019-04-08. Retrieved 2019-11-15.
^ Friedman, Matt; V. Andrews, James; Saad, Hadeel; El-Sayed, Sanaa (2023-06-16). "The Cretaceous–Paleogene transition in spiny-rayed fishes: surveying "Patterson's Gap" in the acanthomorph skeletal record André Dumont medalist lecture 2018". Geologica Belgica. doi:10.20341/gb.2023.002. ISSN 1374-8505.
^ Rust, Seabourne; Wium, Morne; Otero, Rodrigo A.; Terezow, Marianna (2026-02-01). "Fossil billfish (Xiphioidei) from the Eocene of Hampden, North Otago, New Zealand". Gondwana Research. 150: 301–311. doi:10.1016/j.gr.2025.09.021. ISSN 1342-937X.
^ Friedman, Matt; Johanson, Zerina; Harrington, Richard C.; Near, Thomas J.; Graham, Mark R. (2013-09-07). "An early fossil remora (Echeneoidea) reveals the evolutionary assembly of the adhesion disc". Proceedings of the Royal Society B: Biological Sciences. 280 (1766) 20131200. doi:10.1098/rspb.2013.1200. PMC 3730593. PMID 23864599.

[Taxonomicon-1] "Taxon: Order Carangiformes Jordan, 1923 (fish)". Taxonomicon. Retrieved 15 November 2019.

[Nelson5-2] J. S. Nelson; T. C. Grande; M. V. H. Wilson (2016). Fishes of the World (5th ed.). Wiley. pp. 380–383. ISBN 978-1-118-34233-6. Archived from the original on 2019-04-08. Retrieved 2019-11-15.

[3] Girard, Matthew G.; Davis, Matthew P.; Smith, W. Leo (2020-05-08). "The Phylogeny of Carangiform Fishes: Morphological and Genomic Investigations of a New Fish Clade". Copeia. 108 (2): 265. doi:10.1643/CI-19-320. ISSN 0045-8511.

[Wiley_&_Johnson-4] E. O. Wiley & G. David Johnson: A teleost classification based on monophyletic groups. Joseph S. Nelson, Hans-Peter Schultze & Mark V. H. Wilson: Origin and Phylogenetic Interrelationships of Teleosts. Verlag Dr. Friedrich Pfeil, München 2010, ISBN 978-3-89937-107-9

[5] Friedman, Matt; V. Andrews, James; Saad, Hadeel; El-Sayed, Sanaa (2023-06-16). "The Cretaceous–Paleogene transition in spiny-rayed fishes: surveying "Patterson's Gap" in the acanthomorph skeletal record André Dumont medalist lecture 2018". Geologica Belgica. doi:10.20341/gb.2023.002. ISSN 1374-8505.

[Girard-6] Girard, Matthew G.; Davis, Matthew P.; Smith, W. Leo (2020-05-08). "The Phylogeny of Carangiform Fishes: Morphological and Genomic Investigations of a New Fish Clade". Copeia. 108 (2): 265. doi:10.1643/CI-19-320. ISSN 0045-8511.

[7] "Eschmeyer's Catalog of Fishes Classification - California Academy of Sciences". www.calacademy.org. Retrieved 2024-12-27.

[Nelson52-8] J. S. Nelson; T. C. Grande; M. V. H. Wilson (2016). Fishes of the World (5th ed.). Wiley. pp. 380–383. ISBN 978-1-118-34233-6. Archived from the original on 2019-04-08. Retrieved 2019-11-15.

[9] Friedman, Matt; V. Andrews, James; Saad, Hadeel; El-Sayed, Sanaa (2023-06-16). "The Cretaceous–Paleogene transition in spiny-rayed fishes: surveying "Patterson's Gap" in the acanthomorph skeletal record André Dumont medalist lecture 2018". Geologica Belgica. doi:10.20341/gb.2023.002. ISSN 1374-8505.

[:0-10] Rust, Seabourne; Wium, Morne; Otero, Rodrigo A.; Terezow, Marianna (2026-02-01). "Fossil billfish (Xiphioidei) from the Eocene of Hampden, North Otago, New Zealand". Gondwana Research. 150: 301–311. doi:10.1016/j.gr.2025.09.021. ISSN 1342-937X.

[11] Friedman, Matt; Johanson, Zerina; Harrington, Richard C.; Near, Thomas J.; Graham, Mark R. (2013-09-07). "An early fossil remora (Echeneoidea) reveals the evolutionary assembly of the adhesion disc". Proceedings of the Royal Society B: Biological Sciences. 280 (1766) 20131200. doi:10.1098/rspb.2013.1200. PMC 3730593. PMID 23864599.

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