Physics:Flatfish

From HandWiki

Flatfish are a group of ray-finned fish belonging to the suborder Pleuronectoidei of the order Carangiformes. Their collective common name is due to their habit of lying on one side of their laterally-compressed body (flattened side-to-side) upon the seafloor; in this position, both eyes lie on the side of the head facing upwards, while the other side of the head and body (the "blind side") lies on the substrate. This loss of symmetry, a unique adaptation in vertebrates, stems from one eye "migrating" towards the other during the juvenile's metamorphosis. Due to interspecific variation, some species tend to face their left side upward, some their right side, and others face either side upward; members of Pleuronectidae lie on their left side, with eyes on the right, Paralichthyidae lie on their right side, with eyes on the left,[1] while the "primitive" genus Psettodes may develop into "right-facing" or "left-facing" individuals.

They are one of the most diverse groups of demersal fish. Their cryptic coloration and habits, a form of camouflage, conceals them from potential predators. Many species are of interest for fisheries.

Common names

Illustration of several common European flatfish species

There are a multitude of common names for flatfish, as they are a widespread group of fish and important food fish across the world. The following are common flatfish names in English:

As these are merely common names, they do not conform with the "natural" relationships that are recovered through scientific studies of morphology or genetics. As examples, the three species consistently called "halibut" are themselves part of the right-eye flounder family, while the spiny turbots are not at all closely related to "true" turbot, but are consistently recovered in a "primitive" or basal position at the base of flatfish phylogenetic trees.

Distribution

Flatfishes are found in oceans worldwide, ranging from the Arctic, through the tropics, to Antarctica. Species diversity is centered in the Indo-West Pacific and declines following both latitudinal and longitudinal gradients away from this centre of diversity.[2] Most species are found in depths between 0 and 500 m (1,600 ft), but a few have been recorded from depths in excess of 1,500 m (4,900 ft). None have been confirmed from the abyssal or hadal zones of the deep sea; a reported observation of a flatfish from the Bathyscaphe Trieste's dive into the Mariana Trench (at a depth of almost 11 km (36,000 ft)) has been questioned by ichthyologists, and recent authorities do not recognize it as valid.[3] Among the deepwater species is Symphurus thermophilus, a tonguefish which congregates around "ponds" of sulphur at hydrothermal vents on the seafloor; no other flatfish is known from hydrothermal vent ecosystems.[4]

Conversely, many species will enter brackish or fresh water, and a smaller number of soles (families Achiridae and Soleidae) and tonguefish (Cynoglossidae) are entirely restricted to fresh water.[5][6][7]

Description

Winter flounder; Pleuronectidae

The most obvious characteristic of the flatfish is their asymmetry, with both eyes lying on the same side of the head in the adult fish. In some families, the eyes are usually on the right side of the body (dextral or right-eyed flatfish), and in others, they are usually on the left (sinistral or left-eyed flatfish). The primitive spiny turbots include equal numbers of right- and left-sided individuals, and are generally less asymmetrical than the other families.[8] Other distinguishing features of the order are the presence of protrusible eyes, another adaptation to living on the seabed (benthos), and the extension of the dorsal fin onto the head.

Zebrias zebra; Soleidae
Four frames of the same peacock flounder, a sand colored flatfish with a pattern of blue rings, taken a few minutes apart which shows its ability to change colors to match its surroundings. The last photo shows it buried under sand with only its eyes visible
This sequence of photos shows an individual peacock flounder (Bothidae) changing its coloration over different substrates.

The surface of the fish facing away from the sea floor is pigmented, often serving to camouflage the fish, but at times displaying striking patterns. Some flatfishes are also able to change their pigmentation to match the background using their chromatophores, in a manner similar to some cephalopods. The side of the body without the eyes, facing the seabed, is usually colourless or very pale.[8]

In general, flatfishes rely on their camouflage for avoiding predators, but some have aposematic traits such as conspicuous eyespots (e.g., Microchirus ocellatus) and several small tropical species (at least Aseraggodes, Pardachirus and Zebrias) are poisonous.[9][10][11] Juveniles of Soleichthys maculosus mimic toxic flatworms of the genus Pseudobiceros in both colours and swimming pattern.[12][13] Conversely, a few octopus species have been reported to mimic flatfishes in colours, shape and swimming mode.[14]

Flatfishes range in size from the sand flounder Tarphops oligolepis, measuring about 6.5 cm (2.6 in) in length,[15] and weighing 2 g (0.071 oz),[8] to the Hippoglossus halibuts, with the Atlantic halibut measuring up to 4.7 m (15 ft) long,[16] and the Pacific halibut weighing up to 363 kg (800 lb).[17][8]

Many species such as flounders and spiny turbots eat smaller fish, and have well-developed teeth. These species sometimes hunt in the midwater, away from the bottom, and show fewer "extreme" adaptations than other families. The soles, by contrast, are almost exclusively bottom-dwellers (more strictly demersal), and feed on benthic invertebrates. They show a more extreme asymmetry, and may lack teeth on one side of the jaw.[8]

Development

European flounder, like other flatfish, experience an eye migration during their lifetime.

Flatfishes lay eggs that hatch into larvae resembling typical, symmetrical, fish. These are initially elongated, but quickly develop into a more rounded form. The larvae typically have protective spines on the head, over the gills, and in the pelvic and pectoral fins. They also possess a swim bladder, and do not dwell on the bottom, instead dispersing from their hatching grounds as ichthyoplankton.[8] Bilaterally symmetric fish such as goldfish maintain balance using a system within their inner ears which involves the otolith, but larval and metamorphizing flatfish require visible light (such as sunlight) to properly orient themselves.[18]

The length of the planktonic stage varies between different types of flatfishes, but through the influence of thyroid hormones,[19] they eventually begin to metamorphose into the adult form. One of the eyes migrates across the top of the head and onto the other side of the body, leaving the fish blind on one side. The larva also loses its swim bladder and spines, and sinks to the bottom, laying its blind side on the underlying surface.[20][18]

Hybrids

Hybrids are well known in flatfishes. The Pleuronectidae have the largest number of reported hybrids of marine fishes.[21] Two of the most famous intergeneric hybrids are between the European plaice (Pleuronectes platessa) and European flounder (Platichthys flesus) in the Baltic Sea,[22] and between the English sole (Parophrys vetulus) and starry flounder (Platichthys stellatus) in Puget Sound. The offspring of the latter species pair is popularly known as the hybrid sole and was initially believed to be a valid species in its own right.[21]

Evolution

Flatfishes have been cited as dramatic examples of evolutionary adaptation. In The Blind Watchmaker, Richard Dawkins explains the flatfishes' evolutionary history as such:

...bony fish as a rule have a marked tendency to be flattened in a vertical direction.... It was natural, therefore, that when the ancestors of [flatfish] took to the sea bottom, they should have lain on one side.... But this raised the problem that one eye was always looking down into the sand and was effectively useless. In evolution this problem was solved by the lower eye 'moving' round to the upper side.[23]

Scientists have been proposing since the 1910s that flatfishes evolved from more "typical" percoid ancestors.[24] The fossil record indicated that flatfishes might have been present before the Eocene, based on fossil otoliths resembling those of modern pleuronectiforms dating back to the Thanetian and Ypresian stages (57-53 million years ago).[25] Despite this, the origin of the unusual morphology of flatfishes was enigmatic up to the 2000s, with earlier researchers having suggested that it came about as a result of saltation rather than gradual evolution through natural selection, because a partially migrated eye was considered to have been maladaptive.

Specimen of Amphistium.

This started to change in 2008 with a study on the two fossil fish genera; Amphistium and Heteronectes, which dated to about 50 million years ago. These genera retain primitive features not seen in modern types of flatfishes, such as their heads being less asymmetric than modern flatfishes, retaining one eye on each side of their heads, although the eye on one side is closer to the top of the head than on the other.[26][27] The more recently described fossil genera Quasinectes and Anorevus have been proposed to show similar morphologies and have also been classified as "stem-pleuronectiforms".[28][29] Such findings lead palaeontologist Matt Friedman to conclude that the evolution of flatfish morphology "happened gradually, in a way consistent with evolution via natural selection—not suddenly [saltationally] as researchers once had little choice but to believe."[27]

To explain the survival advantage of a partially migrated eye, it has been proposed that primitive flatfishes like Amphistium rested with the head propped up above the seafloor (a behaviour sometimes observed in modern flatfishes), enabling them to use their partially migrated eye to see things closer to the seafloor.[30] While known basal genera like Amphistium and Heteronectes support a gradual acquisition of the flatfish morphology, they were probably not direct ancestors to living pleuronectiforms, as fossil evidence (for example Eobothus) indicate that most flatfish lineages living today were present in the Eocene and contemporaneous with them.[26] It has been suggested that the more primitive forms were eventually outcompeted.[27]

Taxonomy

Due to their highly distinctive morphology, flatfishes were previously treated as belonging to their own order, Pleuronectiformes. However, more recent taxonomic studies have found them to group within a diverse group of nektonic marine fishes known as the Carangiformes, which also includes jacks and billfish. Specifically, flatfish have been recovered to be closely related to various groups, such as the threadfins (often recovered as a sister group to flatfish), archerfish, and beachsalmons. Due to this, they are now treated as a suborder of the Carangiformes,[31][32] as represented in Eschmeyer's Catalog of Fishes.[33]

Classification

The following classification is based on Eschmeyer's Catalog of Fishes (2025):[34]

  • 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, Chanet, Chen, Lee & Chen, 2019 (sand whiffs or large-tooth flounders)
    • Family Grammatobothidae Tongboonkua, Chanet & Chen, 2025 (small lefteye flounders)[35]
    • Family Monolenidae Tongboonkua, Chanet & Chen, 2025 (deepwater lefteye flounders)[35]
    • Family Taeniopsettidae Amaoka, 1969[35]
    • Family Bothidae Smitt, 1892 (lefteye flounders)
    • Family Paralichthyidae Regan, 1910 (sand flounders)
    • Family Pleuronectidae Rafinesque, 1815 (righteye 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)

Fossil taxa

The following basal fossil flatfish from the Paleogene are also known:[36]

  • Genus ?†Anorevus Bannikov & Zorzin, 2020 (Early Eocene of Italy)[28]
  • Genus †Eobothus Eastman, 1914 (Early Eocene of Italy)
  • Genus †Heteronectes Friedman, 2008 (Middle Eocene of France)
  • Genus †Imhoffius Chabanaud, 1940 (Middle Eocene of France)[37]
  • Genus †Keasichthys Murray & Champagne, 2025 (Early Oligocene of Oregon, US)[38]
  • Genus †Numidiopleura Gaudant & Gaudant, 1969 (Eocene of Tunisia)[37][39]
  • Genus ?†Quasinectes Bannikov & Zorzin, 2019 (Early Eocene of Italy)[29]
  • Family †Amphistiidae Boulenger, 1902
  • Family †Joleaudichthyidae Chabanaud, 1937

Phylogeny

Threadfins such as Polynemus have been recovered closer to the primitive spiny turbots than those are to other flatfish, or as a sister group to a monophyletic flatfish group

There has been some disagreement whether flatfish as a whole are a monophyletic group. Some palaeontologists think that some percomorph groups unrelated to flatfishes were also "experimenting" with head asymmetry during the Eocene,[28][29] and certain molecular studies conclude that the primitive family of Psettodidae evolved their flat bodies and asymmetrical head independently of other flatfish groups.[40][41] The following phylogeny is from Lü et al. 2021; a whole-genome analysis using concatenated sequences of coding sequence (CDS) (codon1 + 2 + 3, GTRGAMMA model; codon1 + 2, GTRGAMMA model) and 4dTV (fourfold degenerate synonymous site, GTRGAMMA model) derived from 1,693 single-copy genes. Notably, Pleuronectiformes is found to be polyphyletic as seen here:[42]

 
Polynemidae

Polydactylus sextarius

 
 
Toxotidae

Toxotes chatareus

 
Psettodoidei

Psettodes erumei

Psettodidae
 
 
Pleuronectoidei
 

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However, threadfins (Polynemidae) aren't universally found to be nested within the group of flatfish, as recovered by a study of ultraconserved elements from the threadfin family in Girard et al. 2022,[43] or as represented in the World Register of Marine Species,[44] where Pleuronectiformes is retained as a name for the flatfish group.[45] Numerous scientists continue to argue for a monophyletic group of all flatfish,[46] though the debate continues.[47]

Over 800 described species are placed into 16 families.[48] When they were treated as an order, the flatfishes are divided into two suborders, Psettodoidei and Pleuronectoidei, with > 99% of the species diversity found within the Pleuronectoidei.[49] The largest families are Soleidae, Bothidae and Cynoglossidae with more than 150 species each. There also exist two monotypic families (Paralichthodidae and Oncopteridae). Some families are the results of relatively recent splits. For example, the Achiridae were classified as a subfamily of Soleidae in the past, and the Samaridae were considered a subfamily of the Pleuronectidae.[9][50] The families Paralichthodidae, Poecilopsettidae, and Rhombosoleidae were also traditionally treated as subfamilies of Pleuronectidae, but are now recognised as families in their own right.[50]Cite error: Closing </ref> missing for <ref> tag New species are described with some regularity and undescribed species likely remain.[9]

Timeline of genera

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Relation to humans

Template:Common fish

Fishing and aquaculture

Flatfish are commonly fished using bottom trawls.[51][52] Large species such as the halibuts are specifically targeted by fisheries, resulting in heavy fishing pressures and bycatch.[53][54][55] Some species are aquacultured, such as the tonguefish Cynoglossus semilaevis.[56][57]

As food

Flatfish is considered a whitefish[58] because of the high concentration of oils within its liver. Its lean flesh makes for a unique flavor that differs from species to species. Methods of cooking include grilling, pan-frying, baking and deep-frying.

  • The European plaice is the principal commercial flatfish in Europe.
    The European plaice is the principal commercial flatfish in Europe.
  • American soles are found in both freshwater and marine environments of the Americas.
    American soles are found in both freshwater and marine environments of the Americas.
  • Halibut are the largest of the flatfishes, and provide lucrative fisheries.
    Halibut are the largest of the flatfishes, and provide lucrative fisheries.
  • The turbot is a large, left-eyed flatfish found in sandy shallow coastal waters around Europe.
    The turbot is a large, left-eyed flatfish found in sandy shallow coastal waters around Europe.
  • Flatfish (left‐eyed flounder)
    Flatfish (left‐eyed flounder)

References

  1. ↑ Dawkins, Richard; Wong, Yan (2016). The Ancestor's Tale: A Pilgrimage to the Dawn of Life. Houghton Mifflin Harcourt. ISBN 978-0544859937. 
  2. ↑ Campbell, Matthew A.; Chanet, Bruno; Chen, Jhen-Nien; Lee, Mao-Ying; Chen, Wei-Jen (2019). "Origins and relationships of the Pleuronectoidei: Molecular and morphological analysis of living and fossil taxa" (in en). Zoologica Scripta 48 (5): 640–656. doi:10.1111/zsc.12372. ISSN 1463-6409. https://hal.science/hal-03971070/file/Campbell%20et%20al%202019%20Zool%20Scripta.pdf. 
  3. ↑ Jamieson, A.J., and Yancey, P. H. (2012). On the Validity of the Trieste Flatfish: Dispelling the Myth. The Biological Bulletin 222(3): 171-175
  4. ↑ Munroe, T.A.; and Hashimoto, J. (2008). A new Western Pacific Tonguefish (Pleuronectiformes: Cynoglossidae): The first Pleuronectiform discovered at active Hydrothermal Vents. Zootaxa 1839: 43–59.
  5. ↑ Duplain, R.R.; Chapleau, F; and Munroe, T.A. (2012). A New Species of Trinectes (Pleuronectiformes: Achiridae) from the Upper Río San Juan and Río Condoto, Colombia. Copeia 2012 (3): 541-546.
  6. ↑ Kottelat, M. (1998). Fishes of the Nam Theun and Xe Bangfai basins, Laos, with diagnoses of twenty-two new species (Teleostei: Cyprinidae, Balitoridae, Cobitidae, Coiidae and Odontobutidae). Ichthyol. Explor. Freshwat. 9(1):1-128.
  7. ↑ Monks, N. (2007). Freshwater flatfish, order Pleuronectiformes. Retrieved 18 May 2014
  8. ↑ 8.0 8.1 8.2 8.3 8.4 8.5 Chapleau, Francois; Amaoka, Kunio (1998). Paxton, J.R.. ed. Encyclopedia of Fishes. San Diego: Academic Press. p. xxx. ISBN 0-12-547665-5. 
  9. ↑ 9.0 9.1 9.2 Randall, J. E. (2007). Reef and Shore Fishes of the Hawaiian Islands. ISBN 1-929054-03-3
  10. ↑ Elst, R. van der (1997) A Guide to the Common Sea Fishes of South Africa. ISBN 978-1868253944
  11. ↑ Debelius, H. (1997). Mediterranean and Atlantic Fish Guide. ISBN 978-3925919541
  12. ↑ Practical Fishkeeping (22 May 2012) Video: Tiny sole mimics a flatworm. Retrieved 17 May 2014.
  13. ↑ Australian Museum (5 November 2010). This week in Fish: Flatworm mimic and shark teeth. Retrieved 17 May 2014.
  14. ↑ Hanlon, R.T.; Warson, A.C.; and Barbosa, A. (2010). A "Mimic Octopus" in the Atlantic: Flatfish Mimicry and Camouflage by Macrotritopus defilippi. The Biological Bulletin 218(1): 15-24
  15. ↑ Froese, Rainer and Pauly, Daniel, eds. (2006). "Tarphops oligolepis" in FishBase. April 2006 version.
  16. ↑ Froese, Rainer and Pauly, Daniel, eds. (2006). "Hippoglossus hippoglossus" in FishBase. April 2006 version.
  17. ↑ Froese, Rainer and Pauly, Daniel, eds. (2006). "Hippoglossus stenolepis" in FishBase. April 2006 version.
  18. ↑ 18.0 18.1 Jabr, Feris (7 May 2014). "The Improbable—but True—Evolutionary Tale of Flatfishes". PBS NOVA. https://www.pbs.org/wgbh/nova/article/flatfish-evolution/. 
  19. ↑ Schreiber, Alexander M. (2013). "Flatfish: an asymmetric perspective on metamorphosis". Curr Top Dev Biol 103: 167–94. doi:10.1016/B978-0-12-385979-2.00006-X. PMID 23347519. 
  20. ↑ Bao, Baolong (2022). Flatfish Metamorphosis (1 ed.). Singapore: Springer. doi:10.1007/978-981-19-7859-3. ISBN 978-981-19-7858-6. 
  21. ↑ 21.0 21.1 Garrett, D.L.; Pietsch, T.W.; Utter, F.M.; and Hauser, L. (2007). The Hybrid Sole Inopsetta ischyra (Teleostei: Pleuronectiformes: Pleuronectidae): Hybrid or Biological Species? American Fisheries Society 136: 460–468
  22. ↑ Food and Agriculture Organization of the United Nations: Platichthys flesus (Linnaeus, 1758).. Retrieved 18 May 2014
  23. ↑ Dawkins, Richard (1991). The Blind Watchmaker. London: Penguin Books. p. 92. ISBN 0-14-014481-1. 
  24. ↑ Regan C.T. (1910). "The origin and evolution of the Teleostean fishes of the order Heterosomata". Annals and Magazine of Natural History 6(35): p. 484-496. doi.org/10.1080/00222931008692879
  25. ↑ Schwarzhans W. (1999). "A comparative morphological treatise of recent and fossil otoliths of the order Pleuronectiformes". Piscium Catalogus. Otolithi Piscium 2. doi:10.13140/2.1.1725.5043
  26. ↑ 26.0 26.1 Friedman M. (2008). "The evolutionary origin of flatfish asymmetry". Nature 454(7201): p. 209–212. doi:10.1038/nature07108
  27. ↑ 27.0 27.1 27.2 "Odd Fish Find Contradicts Intelligent-Design Argument". National Geographic. July 9, 2008. http://news.nationalgeographic.com/news/2008/07/080709-evolution-fish.html. 
  28. ↑ 28.0 28.1 28.2 Bannikov A.F. & Zorzin R (2019). "A new genus and species of incertae sedis percomorph fish (Perciformes) from the Eocene of Bolca in northern Italy, and a new genus for Psettopsis latellai Bannikov, 2005". Studi e ricerche sui giacimenti terziari di Bolca: p. 5-15.
  29. ↑ 29.0 29.1 29.2 Bannikov A.F. & Zorzin R. (2020). "A new genus and species of percomorph fish ("stem pleuronectiform") from the Eocene of Bolca in northern Italy". Miscellanea Paleontologica 17: p. 5–14
  30. ↑ Janvier P. (2008). "Squint of the fossil flatfish". Nature 454(7201): p. 169–170
  31. ↑ 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): 265–298. doi:10.1643/CI-19-320. ISSN 0045-8511. https://meridian.allenpress.com/copeia/article-abstract/108/2/265/436665/The-Phylogeny-of-Carangiform-Fishes-Morphological. 
  32. ↑ Shi, Wei; Chen, Shixi; Kong, Xiaoyu; Si, Lizhen; Gong, Li; Zhang, Yanchun; Yu, Hui (2018-05-25). "Flatfish monophyly refereed by the relationship of Psettodes in Carangimorphariae" (in en). BMC Genomics 19 (1): 400. doi:10.1186/s12864-018-4788-5. ISSN 1471-2164. PMID 29801430. 
  33. ↑ Fricke, Ron; Fong, Jon David. "GENERA/SPECIES BY FAMILY/SUBFAMILY IN Eschmeyer's Catalog of Fishes". California Academy of Sciences. https://researcharchive.calacademy.org/research/ichthyology/catalog/SpeciesByFamily.asp. 
  34. ↑ Fricke, R.; Eschmeyer, W. N.; Van der Laan, R. (2025). "ESCHMEYER'S CATALOG OF FISHES: CLASSIFICATION" (in en). https://www.calacademy.org/eschmeyers-catalog-of-fishes-classification. 
  35. ↑ 35.0 35.1 35.2 Tongboonkua, Pakorn; Chanet, Bruno; Chen, Wei-Jen. "Integrated Molecular and Morphological Analyses Resolve Long-Standing Classification Challenges in the Sinistral Flatfish Family Bothidae (Teleostei: Carangiformes)" (in en). Zoologica Scripta n/a (n/a). doi:10.1111/zsc.70020. ISSN 1463-6409. https://onlinelibrary.wiley.com/doi/abs/10.1111/zsc.70020. 
  36. ↑ Laan, Richard van der (2018-10-11). "Family-group names of fossil fishes" (in en). European Journal of Taxonomy (466). doi:10.5852/ejt.2018.466. ISSN 2118-9773. https://europeanjournaloftaxonomy.eu/index.php/ejt/article/view/597. 
  37. ↑ 37.0 37.1 Chanet, Bruno; Schultz, Orwin (1994). "Pleuronectiform fishes from the Upper Badenian (Middle Miocene) of St. Margarethen (Austria)". Ann. Naturhist. Mus. Wien (96): 95–115. https://www.researchgate.net/publication/230844383. 
  38. ↑ Murray, Alison M.; Champagne, Donald E. (January 2025). "A new species of early Oligocene flatfish (Pleuronectiformes) from Oregon, USA" (in en). Geological Magazine 162. doi:10.1017/S0016756825100125. ISSN 0016-7568. Bibcode: 2025GeoM..162E..22M. https://www.cambridge.org/core/journals/geological-magazine/article/new-species-of-early-oligocene-flatfish-pleuronectiformes-from-oregon-usa/29C984BB97C135EF0ADC1CB242CD5143. 
  39. ↑ Campbell, Matthew A.; Chanet, Bruno; Chen, Jhen-Nien; Lee, Mao-Ying; Chen, Wei-Jen (2019). "Origins and relationships of the Pleuronectoidei: Molecular and morphological analysis of living and fossil taxa" (in en). Zoologica Scripta 48 (5): 640–656. doi:10.1111/zsc.12372. ISSN 1463-6409. https://onlinelibrary.wiley.com/doi/abs/10.1111/zsc.12372. 
  40. ↑ Campbell M.A., Chen W-J. & López J.A. (2013). "Are flatfishes (Pleuronectiformes) monophyletic?". Molecular Phylogenetics and Evolution 69(3): p. 664-673. doi.org/10.1016/j.ympev.2013.07.011
  41. ↑ Campbell M.A., López J.A., Satoh T.P., Chen W-J. & Miya M. (2014). "Mitochondrial genomic investigation of flatfish monophyly". Gene 551(2): p. 176-182. doi.org/10.1016/j.gene.2014.08.053
  42. ↑ Zhenming Lü; Li Gong; Yandong Ren; Yongjiu Chen; Zhongkai Wang; Liqin Liu; Haorong Li; Xianqing Chen et al. (19 April 2021). "Large-scale sequencing of flatfish genomes provides insights into the polyphyletic origin of their specialized body plan". Nature Genetics 53 (5): 742–751. doi:10.1038/s41588-021-00836-9. PMID 33875864. 
  43. ↑ Matthew G. Girard; Matthew P. Davis; Carole C. Baldwin; Agnès Dettaï; Rene P. Martin; W. Leo Smith (12 January 2022). "Molecular phylogeny of the threadfin fishes (Polynemidae) using ultraconserved elements". Fish Biology 100 (3): 793–810. doi:10.1111/jfb.14997. PMID 35137410. Bibcode: 2022JFBio.100..793G. https://www.mgirard.fish/files/Girard_et_al_2022.pdf. 
  44. ↑ "Polynemidae Rafinesque, 1815". FishBase. World Register of Marine Species. http://www.marinespecies.org/aphia.php?p=taxdetails&id=125551. 
  45. ↑ "Pleuronectiformes". FishBase. World Register of Marine Species. http://www.marinespecies.org/aphia.php?p=taxdetails&id=10331. 
  46. ↑ Duarte-Ribeiro E, Rosas-Puchuri U, Friedman M, Woodruff G.C., Hughes L.C., Carpenter K.E., White W.T., Pogonoski J.J., Westneat M, Diaz de Astarloa J.M., Williams J.T., Santos M.D., Domínguez-Domínguez O, Ortí G, Arcila D & Betancur-R R. (2024). "Phylogenomic and comparative genomic analyses support a single evolutionary origin of flatfish asymmetry". Nature Genetics 56: p. 1069-1072. doi.org/10.1038/s41588-024-01784-w
  47. ↑ Lü, Zhenming; Li, Haorong; Jiang, Hui; Luo, Hairong; Wang, Wen; Kong, Xiaoyu; Li, Yongxin (2024-05-27). "Reply to: Phylogenomic and comparative genomic analyses support a single evolutionary origin of flatfish asymmetry". Nature Genetics 56 (6): 1073–1074. doi:10.1038/s41588-024-01783-x. PMID 38802565. https://www.x-mol.net/paper/article/1795282921848369152. Retrieved 17 July 2025. 
  48. ↑ Campbell, Matthew A.; Chanet, Bruno; Chen, Jhen-Nien; Lee, Mao-Ying; Chen, Wei-Jen (2019). "Origins and relationships of the Pleuronectoidei: Molecular and morphological analysis of living and fossil taxa" (in en). Zoologica Scripta 48 (5): 640–656. doi:10.1111/zsc.12372. ISSN 0300-3256. https://hal.science/hal-03971070/file/Campbell%20et%20al%202019%20Zool%20Scripta.pdf. 
  49. ↑ Fishes of the world. John Wiley & Sons. 2016-03-28. ISBN 978-1-118-34233-6. OCLC 958002567. 
  50. ↑ 50.0 50.1 Cooper, J.A.; and Chapleau, F. (1998). Monophyly and intrarelationships of the family Pleuronectidae (Pleuronectiformes), with a revised classification. Fish. Bull. 96 (4): 686–726.
  51. ↑ Segura-Garcia, Iris; Soe, Sabai; Tun, NYO-NYO; Box, Stephen (2021). "Beyond Bycatch: The Species Diversity of Tonguesole (Pleuronectiformes: Cynoglossidae) in Coastal Fisheries of the Tanintharyi Region, Southern Myanmar". Asian Fisheries Science 34: 23–33. doi:10.33997/j.afs.2021.34.1.003. https://www.asianfisheriessociety.org/publication/downloadfile.php?id=1347&file=Y0dSbUx6QXhORGsxTWpNd01ERTJNVGN4TkRVMU1qa3VjR1Jt. Retrieved 17 July 2025. 
  52. ↑ Zhang, Jinyong; Cui, Xiaoyu; Lin, Lin; Liu, Yuan; Ye, Jinqing; Zhang, Weiyue; Li, Hongjun (30 April 2025). "Unraveling Fish Community Diversity and Structure in the Yellow Sea: Evidence from Environmental DNA Metabarcoding and Bottom Trawling". Animals 15 (9): 1283. doi:10.3390/ani15091283. PMID 40362097. 
  53. ↑ Szymkowiak, Marysia; Marrinan, Sarah; Kasperski, Stephen (2020). "The Pacific Halibut, Hippoglossus stenolepis, and Sablefish, Anoplopoma fimbria, Individual Fishing Quota Program: A Twenty-year Retrospective". Marine Fisheries Review 82 (1–2): 1–16. doi:10.7755/MFR.82.1-2.1. https://spo.nmfs.noaa.gov/sites/default/files/pdf-content/mfr821-21.pdf. Retrieved 17 July 2025. 
  54. ↑ "Collaborative Pacific Halibut, Hippoglossus stenolepis, Bycatch Control by Canada and the United States". Marine Fisheries Review 66. https://spo.nmfs.noaa.gov/sites/default/files/pdf-content/MFR/mfr662/mfr6624.pdf. Retrieved 17 July 2025. 
  55. ↑ Figus, Elizabeth; Criddle, Keith R. (March 1, 2019). "Characterizing preferences of fishermen to inform decision-making: A case study of the Pacific halibut (Hippoglossus stenolepis) fishery off Alaska". PLOS 14 (3). doi:10.1371/journal.pone.0212537. PMID 30822324. Bibcode: 2019PLoSO..1412537F. 
  56. ↑ Hu, Yuanri; Li, Yangzhen; Li, Zhongming; Chen, Changshan; Zang, Jiajian; Li, Yuwei; Kong, Xiangqing (December 2020). "Novel insights into the selective breeding for disease resistance to vibriosis by using natural outbreak survival data in Chinese tongue sole (Cynoglossus semilaevis)" (in en). Aquaculture 529. doi:10.1016/j.aquaculture.2020.735670. Bibcode: 2020Aquac.52935670H. https://linkinghub.elsevier.com/retrieve/pii/S0044848620314496. 
  57. ↑ Li, Yangzhen; Hu, Yuanri; Yang, Yingming; Zheng, Weiwei; Chen, Changshan; Li, Zhongming (January 2021). "Selective breeding for juvenile survival in Chinese tongue sole (Cynoglossus semilaevis): Heritability and selection response" (in en). Aquaculture 531. doi:10.1016/j.aquaculture.2020.735901. Bibcode: 2021Aquac.53135901L. https://linkinghub.elsevier.com/retrieve/pii/S0044848620314848. 
  58. ↑ "Flatfish BBC". https://www.bbc.co.uk/food/flatfish. 

Further reading

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Wikidata ☰ Q59577 entry



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