Actinopterygii

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Ray-finned fish
Temporal range:
Late SilurianPresent, 425–0  Ma [1]
Actinopterygii-0001.jpg
Scientific classification Red Pencil Icon.png
Kingdom: Animalia
Phylum: Chordata
Superclass: Osteichthyes
Class: Actinopterygii
Klein, 1885
Subclasses

Actinopterygii ( /ˌæktɪnɒptəˈrɪi/ ; from actino-  'having rays',and Ancient Greek πτέρυξ (ptérux) 'wing, fins'), members of which are known as ray-finned fishes, is a class of bony fish. [2] They comprise over 50% of living vertebrate species. [3]

Contents

The ray-finned fishes are so called because their fins are webs of skin supported by bony or horny spines (rays), as opposed to the fleshy, lobed fins that characterize the class Sarcopterygii (lobe-finned fish). These actinopterygian fin rays attach directly to the proximal or basal skeletal elements, the radials, which represent the link or connection between these fins and the internal skeleton (e.g., pelvic and pectoral girdles).

By species count, actinopterygians dominate the vertebrates, and they constitute nearly 99% of the over 30,000 species of fish. [4] They are ubiquitous throughout freshwater and marine environments from the deep sea to the highest mountain streams. Extant species can range in size from Paedocypris , at 8 mm (0.3 in), to the massive ocean sunfish, at 2,300 kg (5,070 lb), and the long-bodied oarfish, at 11 m (36 ft). The vast majority of Actinopterygii (~99%) are teleosts.

Characteristics

Anatomy of a typical ray-finned fish (cichlid)
A: dorsal fin, B: fin rays, C: lateral line, D: kidney, E: swim bladder, F: Weberian apparatus, G: inner ear, H: brain, I: nostrils, L: eye, M: gills, N: heart, O: stomach, P: gall bladder, Q: spleen, R: internal sex organs (ovaries or testes), S: ventral fins, T: spine, U: anal fin, V: tail (caudal fin). Possible other parts not shown: barbels, adipose fin, external genitalia (gonopodium) Anatomia dei pesci.jpg
Anatomy of a typical ray-finned fish (cichlid)
A: dorsal fin, B: fin rays, C: lateral line, D: kidney, E: swim bladder, F: Weberian apparatus, G: inner ear, H: brain, I: nostrils, L: eye, M: gills, N: heart, O: stomach, P: gall bladder, Q: spleen, R: internal sex organs (ovaries or testes), S: ventral fins, T: spine, U: anal fin, V: tail (caudal fin). Possible other parts not shown: barbels, adipose fin, external genitalia (gonopodium)

Ray-finned fishes occur in many variant forms. The main features of a typical ray-finned fish are shown in the adjacent diagram. The swim bladder is the more derived structure. [5]

Ray-finned fishes have many different types of scales; but all teleosts have leptoid scales. The outer part of these scales fan out with bony ridges, while the inner part is crossed with fibrous connective tissue. Leptoid scales are thinner and more transparent than other types of scales, and lack the hardened enamel or dentine-like layers found in the scales of many other fish. Unlike ganoid scales, which are found in non-teleost actinopterygians, new scales are added in concentric layers as the fish grows. [6]

Ray-finned and lobe-finned fishes sometimes possesses lungs used for aerial respiration. Only bichirs retain ventrally budding lungs. [5]

Body shapes and fin arrangements

Ray-finned fish vary in size and shape, in their feeding specializations, and in the number and arrangement of their ray-fins.

Reproduction

Three-spined stickleback males (red belly) build nests and compete to attract females to lay eggs in them. Males then defend and fan the eggs. Painting by Alexander Francis Lydon, 1879 Gasterosteus aculeatus 1879.jpg
Three-spined stickleback males (red belly) build nests and compete to attract females to lay eggs in them. Males then defend and fan the eggs. Painting by Alexander Francis Lydon, 1879

In nearly all ray-finned fish, the sexes are separate, and in most species the females spawn eggs that are fertilized externally, typically with the male inseminating the eggs after they are laid. Development then proceeds with a free-swimming larval stage. [7] However other patterns of ontogeny exist, with one of the commonest being sequential hermaphroditism. In most cases this involves protogyny, fish starting life as females and converting to males at some stage, triggered by some internal or external factor. Protandry, where a fish converts from male to female, is much less common than protogyny. [8]

Most families use external rather than internal fertilization. [9] Of the oviparous teleosts, most (79%) do not provide parental care. [10] Viviparity, ovoviviparity, or some form of parental care for eggs, whether by the male, the female, or both parents is seen in a significant fraction (21%) of the 422 teleost families; no care is likely the ancestral condition. [10] The oldest case of viviparity in ray-finned fish is found in Middle Triassic species of Saurichthys . [11] Viviparity is relatively rare and is found in about 6% of living teleost species; male care is far more common than female care. [10] [12] Male territoriality "preadapts" a species for evolving male parental care. [13] [14]

There are a few examples of fish that self-fertilise. The mangrove rivulus is an amphibious, simultaneous hermaphrodite, producing both eggs and spawn and having internal fertilisation. This mode of reproduction may be related to the fish's habit of spending long periods out of water in the mangrove forests it inhabits. Males are occasionally produced at temperatures below 19 °C (66 °F) and can fertilise eggs that are then spawned by the female. This maintains genetic variability in a species that is otherwise highly inbred. [15]

Classification and fossil record

Evolution of ray-finned fish.png

Actinopterygii is divided into the classes Cladistia and Actinopteri. The latter comprises the subclasses Chondrostei and Neopterygii. The Neopterygii, in turn, is divided into the infraclasses Holostei and Teleostei. During the Mesozoic (Triassic, Jurassic, Cretaceous) and Cenozoic the teleosts in particular diversified widely. As a result, 96% of living fish species are teleosts (40% of all fish species belong to the teleost subgroup Acanthomorpha), while all other groups of actinopterygians represent depauperate lineages. [16]

The classification of ray-finned fishes can be summarized as follows:

The cladogram below shows the main clades of living actinopterygians and their evolutionary relationships to other extant groups of fishes and the four-limbed vertebrates (tetrapods). [17] [18] The latter include mostly terrestrial species but also groups that became secondarily aquatic (e.g. Whales and Dolphins). Tetrapods evolved from a group of bony fish during the Devonian period. [19] Approximate divergence dates for the different actinopterygian clades (in millions of years, mya) are from Near et al., 2012. [17]

Vertebrates
Jawed vertebrates
Euteleostomi
Sarcopterygii
Rhipidistia
Tetrapods
Amniota

Sauropsids (reptiles, birds) British reptiles, amphibians, and fresh-water fishes (1920) (Lacerta agilis).jpg

Mammals Phylogenetic tree of marsupials derived from retroposon data (Paucituberculata).png

Amphibians Salamandra salamandra (white background).jpg

Lungfish Barramunda coloured.jpg

Actinistia

Coelacanths Coelacanth flipped.png

(lobefins)
Actinopterygii
Cladistia

Polypteriformes (bichirs, reedfishes) Cuvier-105-Polyptere.jpg

Actinopteri
Chondrostei

Acipenseriformes (sturgeons, paddlefishes) Atlantic sturgeon flipped.jpg

Neopterygii
Holostei

Lepisosteiformes (gars) Alligator gar fish (white background).jpg

Amiiformes (bowfins) Amia calva (white background).jpg

275 mya

Teleostei Common carp (white background).jpg

310 mya
360 mya
400 mya
('bony fish')

Cartilaginous fishes (sharks, rays, ratfish) White shark (Duane Raver).png

Jaw-less fishes (hagfish, lampreys) Nejonoga, Iduns kokbok.jpg

The polypterids (bichirs and reedfish) are the sister lineage of all other actinopterygians, the Acipenseriformes (sturgeons and paddlefishes) are the sister lineage of Neopterygii, and Holostei (bowfin and gars) are the sister lineage of teleosts. The Elopomorpha (eels and tarpons) appear to be the most basal teleosts. [17]

The earliest known fossil actinopterygian is Andreolepis hedei , dating back 420 million years (Late Silurian), remains of which have been found in Russia, Sweden, and Estonia. [20] Crown group actinopterygians most likely originated near the Devonian-Carboniferous boundary. [21] The earliest fossil relatives of modern teleosts are from the Triassic period ( Prohalecites , Pholidophorus ), [22] [23] although it is suspected that teleosts originated already during the Paleozoic Era. [17]

Chondrostei Atlantic sturgeon flipped.jpg Chondrostei (cartilage bone) is a subclass of primarily cartilaginous fish showing some ossification. Earlier definitions of Chondrostei are now known to be paraphyletic, meaning that this subclass does not contain all the descendants of their common ancestor. There used to be 52 species divided among two orders, the Acipenseriformes (sturgeons and paddlefishes) and the Polypteriformes (reedfishes and bichirs). Reedfish and birchirs are now separated from the Chondrostei into their own sister lineage, the Cladistia. It is thought that the chondrosteans evolved from bony fish but lost the bony hardening of their cartilaginous skeletons, resulting in a lightening of the frame. Elderly chondrosteans show beginnings of ossification of the skeleton, suggesting that this process is delayed rather than lost in these fish. [24] This group had once been classified with the sharks: the similarities are obvious, as not only do the chondrosteans mostly lack bone, but the structure of the jaw is more akin to that of sharks than other bony fish, and both lack scales (excluding the Polypteriforms). Additional shared features include spiracles and, in sturgeons, a heterocercal tail (the vertebrae extend into the larger lobe of the caudal fin). However the fossil record suggests that these fish have more in common with the Teleostei than their external appearance might suggest. [24]
Neopterygii Salmo salar flipped.jpg Neopterygii (new fins) is a subclass of ray-finned fish that appeared somewhere in the Late Permian. There were only few changes during its evolution from the earlier actinopterygians. Neopterygians are a very successful group of fishes because they can move more rapidly than their ancestors. Their scales and skeletons began to lighten during their evolution, and their jaws became more powerful and efficient. While electroreception and the ampullae of Lorenzini is present in all other groups of fish, with the exception of hagfish, neopterygians have lost this sense, though it later re-evolved within Gymnotiformes and catfishes, who possess nonhomologous teleost ampullae. [25]
Fossil of the Devonian +cheirolepidiform +Cheirolepis canadensis Cheirolepis canadensis.jpg
Fossil of the Devonian cheirolepidiform Cheirolepis canadensis
Fossil of the Carboniferous +elonichthyiform +Elonichthys peltigerus Elonichthys peltigerus.jpg
Fossil of the Carboniferous elonichthyiform Elonichthys peltigerus
Fossil of the Permian +aeduelliform +Aeduella blainvillei Aeduella sp.JPG
Fossil of the Permian aeduelliform Aeduella blainvillei
Fossil of the Permian +palaeonisciform +Palaeoniscum freieslebeni PalaeoniscusFreieslebenensis-NaturalHistoryMuseum-August23-08.jpg
Fossil of the Permian palaeonisciform Palaeoniscum freieslebeni
Fossil of the Triassic +bobasatraniiform +Bobasatrania canadensis Bobasatrania canadensis 1.jpg
Fossil of the Triassic bobasatraniiform Bobasatrania canadensis
Fossil of the Triassic +perleidiform +Thoracopterus magnificus Thoracopterus magnificus.JPG
Fossil of the Triassic perleidiform Thoracopterus magnificus
Fossils of the Triassic +prohaleciteiform +Prohalecites sp., the earliest teleosteomorph Prohalecites sp Rasa 1.JPG
Fossils of the Triassic prohaleciteiform Prohalecites sp., the earliest teleosteomorph
Fossil of the Jurassic +aspidorhynchiform +Aspidorhynchus sp. Aspidorhynchus sp.jpg
Fossil of the Jurassic aspidorhynchiform Aspidorhynchus sp.
Fossil of the Jurassic +pachycormiform +Pachycormus curtus Pachycormus curtus SMNS 55300.jpg
Fossil of the Jurassic pachycormiform Pachycormus curtus
Fossil of the Cretaceous acipenseriform +Yanosteus longidorsalis Yanosteus longidorsalis MHNT.jpg
Fossil of the Cretaceous acipenseriform Yanosteus longidorsalis
Fossil of the Cretaceous aulopiform +Nematonotus longispinus Nematonotus longispinus.jpg
Fossil of the Cretaceous aulopiform Nematonotus longispinus
Fossil of the Cretaceous +ichthyodectiform +Thrissops formosus Thrissops formosus 3.JPG
Fossil of the Cretaceous ichthyodectiform Thrissops formosus
Fossil of the Cretaceous carangiform +Mene oblonga Mene oblonga 23.JPG
Fossil of the Cretaceous carangiform Mene oblonga
Fossil of the Cretaceous pleuronectiform +Amphistium paradoxum Amphistium.JPG
Fossil of the Cretaceous pleuronectiform Amphistium paradoxum
Fossil of a ray-finned perch (+Priscacara serrata) from the Lower Eocene about 50 million years ago Priscacara serrata FMNH PF13014 img1.jpg
Fossil of a ray-finned perch ( Priscacara serrata ) from the Lower Eocene about 50 million years ago
Fossil of the Miocene syngnathiform +Nerophis zapfei Nerophis (7992564775).jpg
Fossil of the Miocene syngnathiform Nerophis zapfei
Skeleton of the angler fish, Lophius piscatorius. The first spine of the dorsal fin of the anglerfish is modified so it functions like a fishing rod with a lure Lophius piscatorius MHNT.jpg
Skeleton of the angler fish, Lophius piscatorius . The first spine of the dorsal fin of the anglerfish is modified so it functions like a fishing rod with a lure
Skeleton of another ray-finned fish, the lingcod Lingcodskeleton1600ppx.JPG
Skeleton of another ray-finned fish, the lingcod
Blue catfish skeleton Blue catfish skeleton.jpg
Blue catfish skeleton

Taxonomy

The listing below is a summary of all extinct (indicated by a dagger, †) and living groups of Actinopterygii with their respective taxonomic rank. The taxonomy follows Phylogenetic Classification of Bony Fishes [18] [26] with notes when this differs from Nelson, [3] ITIS [27] and FishBase [28] and extinct groups from Van der Laan 2016 [29] and Xu 2021. [30]

Related Research Articles

<span class="mw-page-title-main">Osteichthyes</span> Diverse group of fish with skeletons of bone rather than cartilage

Osteichthyes, popularly referred to as the bony fish, is a diverse superclass of fish that have skeletons primarily composed of bone tissue. They can be contrasted with the Chondrichthyes, which have skeletons primarily composed of cartilage. The vast majority of fish are members of Osteichthyes, which is an extremely diverse and abundant group consisting of 45 orders, and over 435 families and 28,000 species. It is the largest class of vertebrates in existence today. The group Osteichthyes is divided into the ray-finned fish (Actinopterygii) and lobe-finned fish (Sarcopterygii). The oldest known fossils of bony fish are about 425 million years old, which are also transitional fossils, showing a tooth pattern that is in between the tooth rows of sharks and bony fishes.

<span class="mw-page-title-main">Amiiformes</span> Order of fishes

The Amiiformes order of fish has only one extant species, the bowfin. These Amiiformes are found in the freshwater systems of North America, in the United States and parts of southern Canada. They live in freshwater streams, rivers, and swamps.

<span class="mw-page-title-main">Semionotiformes</span> Extinct order of fishes

Semionotiformes is an order of primitive, ray-finned, primarily freshwater fish from the Middle Triassic (Anisian) to the Late Cretaceous (Maastrichtian). The best-known genus is Semionotus of Europe and North America. Their closest living relatives are gars (Lepisosteidae), with both groups belonging to the clade Ginglymodi within the Holostei.

<span class="mw-page-title-main">Actinopteri</span> Group of fishes

Actinopteri is the sister group of Cladistia in the class Actinopterygii.

<span class="mw-page-title-main">Neopterygii</span> Subclass of fishes

Neopterygii is a subclass of ray-finned fish (Actinopterygii). Neopterygii includes the Holostei and the Teleostei, of which the latter comprise the vast majority of extant fishes, and over half of all living vertebrate species. While living holosteans include only freshwater taxa, teleosts are diverse in both freshwater and marine environments. Many new species of teleosts are scientifically described each year.

<span class="mw-page-title-main">Elopomorpha</span> Superorder of fishes

The superorder Elopomorpha contains a variety of types of fishes that range from typical silvery-colored species, such as the tarpons and ladyfishes of the Elopiformes and the bonefishes of the Albuliformes, to the long and slender, smooth-bodied eels of the Anguilliformes. The one characteristic uniting this group of fishes is they all have leptocephalus larvae, which are unique to the Elopomorpha. No other fishes have this type of larvae.

<span class="mw-page-title-main">Holostei</span> Group of bony fish

Holostei is a group of ray-finned bony fish. It is divided into two major clades, the Halecomorphi, represented by a single living species, the bowfin, as well as the Ginglymodi, the sole living representatives being the gars (Lepisosteidae), represented by seven living species in two genera. The earliest members of the clade appeared during the Early Triassic, over 250 million years ago.

<span class="mw-page-title-main">Palaeonisciformes</span> Extinct order of fishes

The Palaeonisciformes (Palaeoniscida) are an extinct order of early ray-finned fishes (Actinopterygii). Palaeonisciformes sensu lato first appeared in the fossil record in the Late Silurian and last appeared in the Late Cretaceous. The name is derived from the Ancient Greek words παλαιός and ὀνίσκος, probably pertaining to the organization of the fishes' scales, similar to the exoskeletal plating of woodlice.

Acanthomorpha is an extraordinarily diverse taxon of teleost fishes with spiny-rays. The clade contains about one third of the world's modern species of vertebrates: over 14,000 species.

<span class="mw-page-title-main">Aspidorhynchiformes</span> Extinct order of ray-finned fishes

Aspidorhynchiformes is an extinct order of ray-finned fish. It contains only a single family, the Aspidorhynchidae. Members of the group are noted for their elongated, conical rostrums, of varying length, formed from fused premaxillae. They are generally interpreted as stem-group teleosts. The range of the group extends from the Middle Jurassic to the late Paleocene.

<span class="mw-page-title-main">Pachycormiformes</span> Extinct order of ray-finned fishes

Pachycormiformes is an extinct order of marine ray-finned fish known from the Early Jurassic to the end of the Cretaceous. It only includes a single family, Pachycormidae. They were characterized by having serrated pectoral fins, reduced pelvic fins and a bony rostrum. Their exact relations with other fish are unclear, but they are generally considered to be teleosteomorphs, more closely related to teleosts than to Holostei. Pachycormiformes are morphologically diverse, containing both tuna and swordfish-like carnivorous forms, as well as edentulous suspension-feeding forms, with the latter including the largest ray finned fish known to have existed, Leedsichthys, with an estimated maximum length of 16 metres.

<i>Prohalecites</i> Extinct genus of ray-finned fishes

Prohalecites is an extinct genus of ray-finned fish from the Ladinian and possibly Carnian (Triassic) of Italy. It is the oldest known teleosteomorph, a group that includes extant teleosts and their close fossil relatives.

<span class="mw-page-title-main">Cladistia</span> Clade of ray-finned fishes

Cladistia is a clade of bony fishes whose only living members are the bichirs. Their major synapomorphies are a heterocercal tail in which the dorsal fin has independent rays, and a posteriorly elongated parasphenoid.

<span class="mw-page-title-main">Crossognathiformes</span> Extinct order of ray-finned fishes

Crossognathiformes is an extinct order of ray-finned fish that lived from the Late Jurassic to the Eocene. Its phylogenetic placement is disputed; some authors have recovered it as part of the teleost stem group, while others place it in a basal position within crown group Teleostei.

<span class="mw-page-title-main">Halecomorphi</span> Clade of fishes

Halecomorphi is a taxon of ray-finned bony fish in the clade Neopterygii. The sole living Halecomorph is the bowfin, but the group contains many extinct species in several families in the order Amiiformes, as well as the extinct orders Ionoscopiformes, Panxianichthyiformes, and Parasemionotiformes. The fossil record of halecomorphs goes back at least to the Early Triassic epoch.

<span class="mw-page-title-main">Percomorpha</span> Clade of ray-finned fishes

Percomorpha is a large clade of ray-finned fish that includes the tuna, seahorses, gobies, cichlids, flatfish, wrasse, perches, anglerfish, and pufferfish.

<span class="mw-page-title-main">Otocephala</span> Clade of ray-finned fishes

Otocephala is a clade of ray-finned fishes within the infraclass Teleostei that evolved some 230 million years ago. It is named for the presence of a hearing (otophysic) link from the swimbladder to the inner ear. Other names proposed for the group include Ostarioclupeomorpha and Otomorpha.

<i>Amia</i> (fish) Genus of ray-finned fishes

Amia, commonly called bowfin, is a genus of bony fish related to gars in the infraclass Holostei. They are regarded as taxonomic relicts, being the sole surviving species of the order Amiiformes, which dates from the Jurassic to the Eocene, persisting to the present. There is one living species in Amia, Amia calva, and a number of extinct species which have been described from the fossil record.

<span class="mw-page-title-main">Parasemionotiformes</span> Extinct order of ray-finned fishes

Parasemionotiformes is an extinct order of neopterygian ray-finned fish that existed globally during the Triassic period. It comprises the families Parasemionotidae and Promecosominidae. Many of the included genera are monotypic and most species lived during the Early Triassic epoch.

<span class="mw-page-title-main">Scorpaenini</span>

Scorpaenini is a tribe of marine ray-finned fishes, one of two tribes in the subfamily Scorpaeninae. This tribe contains the "typical" or "true" scorpionfishes. The taxonomy of the scorpionfishes is in some flux, the 5th Edition of Fishes of the World treats this taxa as a tribe within the subfamily Scorpaeninae of the family Scorpaenidae within the order Scorpaeniformes, while other authorities treat it as a subfamily within a reduced family Scorpaenidae within the suborder Scorpaenoidei, or the superfamily Scorpaenoidea within the order Perciformes.

References

  1. Zhao, W.; Zhang, X.; Jia, G.; Shen, Y.; Zhu, M. (2021). "The Silurian-Devonian boundary in East Yunnan (South China) and the minimum constraint for the lungfish-tetrapod split". Science China Earth Sciences. 64 (10): 1784–1797. Bibcode:2021ScChD..64.1784Z. doi:10.1007/s11430-020-9794-8. S2CID   236438229.
  2. Kardong, Kenneth (2015). Vertebrates: Comparative Anatomy, Function, Evolution. New York: McGraw-Hill Education. pp. 99–100. ISBN   978-0-07-802302-6.
  3. 1 2 Nelson, Joseph S. (2016). Fishes of the World. John Wiley & Sons. ISBN   978-1-118-34233-6.
  4. (Davis, Brian 2010).
  5. 1 2 Funk, Emily; Breen, Catriona; Sanketi, Bhargav; Kurpios, Natasza; McCune, Amy (2020). "Changing in Nkx2.1, Sox2, Bmp4, and Bmp16 expression underlying the lung-to-gas bladder evolutionary transition in ray-finned fishes". Evolution & Development. 22 (5): 384–402. doi:10.1111/ede.12354. PMC   8013215 . PMID   33463017.
  6. "Actinopterygii Klein, 1885". www.gbif.org. Retrieved 20 September 2021.
  7. Dorit, R.L.; Walker, W.F.; Barnes, R.D. (1991). Zoology . Saunders College Publishing. p.  819. ISBN   978-0-03-030504-7.
  8. Avise, J.C.; Mank, J.E. (2009). "Evolutionary perspectives on hermaphroditism in fishes". Sexual Development. 3 (2–3): 152–163. doi:10.1159/000223079. PMID   19684459. S2CID   22712745.
  9. Pitcher, T (1993). The Behavior of Teleost Fishes. London: Chapman & Hall.
  10. 1 2 3 Reynolds, John; Nicholas B. Goodwin; Robert P. Freckleton (19 March 2002). "Evolutionary Transitions in Parental Care and Live Bearing in Vertebrates". Philosophical Transactions of the Royal Society B: Biological Sciences. 357 (1419): 269–281. doi:10.1098/rstb.2001.0930. PMC   1692951 . PMID   11958696.
  11. Maxwell; et al. (2018). "Re‐evaluation of the ontogeny and reproductive biology of the Triassic fish Saurichthys (Actinopterygii, Saurichthyidae)". Palaeontology. 61: 559–574. doi:10.5061/dryad.vc8h5.
  12. Clutton-Brock, T. H. (1991). The Evolution of Parental Care. Princeton, NJ: Princeton UP.
  13. Werren, John; Mart R. Gross; Richard Shine (1980). "Paternity and the evolution of male parentage". Journal of Theoretical Biology. 82 (4): 619–631. doi:10.1016/0022-5193(80)90182-4. PMID   7382520 . Retrieved 15 September 2013.
  14. Baylis, Jeffrey (1981). "The Evolution of Parental Care in Fishes, with reference to Darwin's rule of male sexual selection". Environmental Biology of Fishes. 6 (2): 223–251. doi:10.1007/BF00002788. S2CID   19242013.
  15. Wootton, Robert J.; Smith, Carl (2014). Reproductive Biology of Teleost Fishes. Wiley. ISBN   978-1-118-89139-1.
  16. Sallan, Lauren C. (February 2014). "Major issues in the origins of ray-finned fish (Actinopterygii) biodiversity". Biological Reviews. 89 (4): 950–971. doi:10.1111/brv.12086. hdl:2027.42/109271. PMID   24612207. S2CID   24876484.
  17. 1 2 3 4 Thomas J. Near; et al. (2012). "Resolution of ray-finned fish phylogeny and timing of diversification". PNAS. 109 (34): 13698–13703. Bibcode:2012PNAS..10913698N. doi: 10.1073/pnas.1206625109 . PMC   3427055 . PMID   22869754.
  18. 1 2 Betancur-R, Ricardo; et al. (2013). "The Tree of Life and a New Classification of Bony Fishes". PLOS Currents Tree of Life. 5 (Edition 1). doi:10.1371/currents.tol.53ba26640df0ccaee75bb165c8c26288. PMC   3644299 . PMID   23653398. Archived from the original on 13 October 2013.
  19. Laurin, M.; Reisz, R.R. (1995). "A reevaluation of early amniote phylogeny". Zoological Journal of the Linnean Society. 113 (2): 165–223. doi:10.1111/j.1096-3642.1995.tb00932.x.
  20. "Fossilworks: Andreolepis". Archived from the original on 12 February 2010. Retrieved 14 May 2008.
  21. Henderson, Struan; Dunne, Emma M.; Fasey, Sophie A.; Giles, Sam (3 October 2022). "The early diversification of ray-finned fishes (Actinopterygii): hypotheses, challenges and future prospects". Biological Reviews. doi:10.1111/brv.12907 . Retrieved 29 October 2022.
  22. Arratia, G. (2015). "Complexities of early teleostei and the evolution of particular morphological structures through time". Copeia. 103 (4): 999–1025. doi:10.1643/CG-14-184. S2CID   85808890.
  23. Romano, Carlo; Koot, Martha B.; Kogan, Ilja; Brayard, Arnaud; Minikh, Alla V.; Brinkmann, Winand; Bucher, Hugo; Kriwet, Jürgen (February 2016). "Permian-Triassic Osteichthyes (bony fishes): diversity dynamics and body size evolution". Biological Reviews. 91 (1): 106–147. doi:10.1111/brv.12161. PMID   25431138. S2CID   5332637.
  24. 1 2 "Chondrosteans: Sturgeon Relatives". paleos.com. Archived from the original on 25 December 2010.
  25. Theodore Holmes Bullock; Carl D. Hopkins; Arthur N. Popper (2005). Electroreception. Springer Science+Business Media, Incorporated. p. 229. ISBN   978-0-387-28275-6.
  26. Betancur-Rodriguez; et al. (2017). "Phylogenetic Classification of Bony Fishes Version 4". BMC Evolutionary Biology. 17 (1): 162. doi:10.1186/s12862-017-0958-3. PMC   5501477 . PMID   28683774.
  27. "Actinopterygii". Integrated Taxonomic Information System . Retrieved 3 April 2006.
  28. R. Froese and D. Pauly, ed. (February 2006). "FishBase". Archived from the original on 5 July 2018. Retrieved 8 January 2020.
  29. Van der Laan, Richard (2016). Family-group names of fossil fishes. doi:10.13140/RG.2.1.2130.1361.
  30. Xu, Guang-Hui (9 January 2021). "A new stem-neopterygian fish from the Middle Triassic (Anisian) of Yunnan, China, with a reassessment of the relationships of early neopterygian clades". Zoological Journal of the Linnean Society. 191 (2): 375–394. doi:10.1093/zoolinnean/zlaa053. ISSN   0024-4082.
  31. In Nelson, Polypteriformes is placed in its own subclass Cladistia.
  32. In Nelson and ITIS, Syngnathiformes is placed as the suborder Syngnathoidei of the order Gasterosteiformes.