Actinopterygii · Lablypedia

Ray-finned fish are so called because of their lightly built fins made of webbings of skin supported by radially extended thin bony spines called lepidotrichia, as opposed to the bulkier, fleshy fins of the sister clade Sarcopterygii (lobe-finned fish). Resembling folding fans, the actinopterygian fins can easily change shape, orientation and wetted area, providing superior thrust-to-weight ratios per movement compared to sarcopterygian and chondrichthyian fins. The fin rays attach directly to the proximal or basal skeletal elements, the radials, which represent the articulation between these fins and the internal skeleton (e.g., pelvic and pectoral girdles).

Characteristics

Ray-finned fishes occur in many variant forms. The main features of typical ray-finned fish are shown in the adjacent diagram. The swim bladder is a more derived structure and used for buoyancy. Except from the bichirs, which just like the lungs of lobe-finned fish have retained the ancestral condition of ventral budding from the foregut, the swim bladder in ray-finned fishes derives from a dorsal bud above the foregut. In early forms the swim bladder could still be used for breathing, a trait still present in Holostei (bowfins and gars). In some fish like the arapaima, the swim bladder has been modified for breathing air again, and in other lineages it has been completely lost. The teleosts have urinary and reproductive tracts that are fully separated, while the Chondrostei have common urogenital ducts, and partially connected ducts are found in Cladistia and Holostei. 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. Teleosts and chondrosteans (sturgeons and paddlefish) also differ from the bichirs and holosteans (bowfin and gars) in having gone through a whole-genome duplication (paleopolyploidy). The WGD is estimated to have happened about 320 million years ago in the teleosts, which on average has retained about 17% of the gene duplicates, and around 180 (124–225) million years ago in the chondrosteans. It has since happened again in some teleost lineages, like Salmonidae (80–100 million years ago) and several times independently within the Cyprinidae (in goldfish and common carp as recently as 14 million years ago).

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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

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. 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. Most families use external rather than internal fertilization. Of the oviparous teleosts, most (79%) do not provide parental care. 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. The oldest case of viviparity in ray-finned fish is found in Middle Triassic species of †Saurichthys. Viviparity is relatively rare and is found in about 6% of living teleost species; male care is far more common than female care. Male territoriality "preadapts" a species for evolving male parental care. 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.

Classification and fossil record

Actinopterygii is divided into the subclasses Cladistia, 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. The classification of ray-finned fishes can be summarized as follows:

Cladistia, which include bichirs and reedfish
Actinopteri, which include:
- Chondrostei, which include Acipenseriformes (paddlefishes and sturgeons)
- Neopterygii, which include:
  - Teleostei (most living fishes)
  - Holostei, which include:
    - Lepisosteiformes (gars)
    - Amiiformes (bowfin)

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). 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. Approximate divergence dates for the different actinopterygian clades (in millions of years, mya) are from Near et al., 2012.

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. 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. Crown group actinopterygians most likely originated near the Devonian-Carboniferous boundary. The earliest fossil relatives of modern teleosts are from the Triassic period (Prohalecites, Pholidophorus), although it is suspected that teleosts originated already during the Paleozoic Era.

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 Eschmeyer's Catalog of Fishes and Phylogenetic Classification of Bony Fishes with notes when this differs from Nelson, ITIS and FishBase and extinct groups from Van der Laan 2016 and Xu 2021.

* Order †?Asarotiformes Schaeffer 1968

Order †?Discordichthyiformes Minikh 1998
Order †?Paphosisciformes Grogan & Lund 2015
Order †Cheirolepidiformes Kazantseva-Selezneva 1977
Order †Paramblypteriformes Heyler 1969
Order †Rhadinichthyiformes
Order †Palaeonisciformes Hay 1902
Order †Tarrasiiformes sensu Lund & Poplin 2002
Order †Ptycholepiformes Andrews et al. 1967
Order †Haplolepidiformes Westoll 1944
Order †Aeduelliformes Heyler 1969
Order †Platysomiformes Aldinger 1937
Order †Dorypteriformes Cope 1871
Order †Eurynotiformes Sallan & Coates 2013
Subclass Cladistia Pander 1860
- Order Polypteriformes Bleeker 1859 (bichirs and reedfishes)
- Order †?Scanilepiformes Selezneya 1985
Subclass Actinopteri Cope 1972 s.s.
- Order †Elonichthyiformes Kazantseva-Selezneva 1977
- Order †Phanerorhynchiformes
- Order †Bobasatraniiformes Berg 1940
- Order †Guildayichthyiformes Lund 2000
- Order †Saurichthyiformes Aldinger 1937
Subclass Chondrostei Müller, 1844
- Order †Birgeriiformes Heyler 1969
- Order †Chondrosteiformes Aldinger, 1937
- Order Acipenseriformes Berg 1940 (includes sturgeons and paddlefishes)
Subclass Neopterygii Regan 1923 sensu Xu & Wu 2012
- Order †Pholidopleuriformes Berg 1937
- Order †Redfieldiiformes Berg 1940
- Order †Platysiagiformes Brough 1939
- Order †Polzbergiiformes Griffith 1977
- Order †Perleidiformes Berg 1937
- Order †Louwoichthyiformes Xu 2021
- Order †Peltopleuriformes Lehman 1966
- Order †Colobodontiformes Xu et al 2024
- Order †Luganoiiformes Lehman 1958
- Order †Pycnodontomorpha Nursall, 2010
- Infraclass Holostei Müller 1844
  - Division Halecomorphi Cope 1872 sensu Grande & Bemis 1998
    - Order †Parasemionotiformes Lehman 1966
    - Order †Ionoscopiformes Grande & Bemis 1998
    - Order Amiiformes Huxley 1861 sensu Grande & Bemis 1998 (bowfins)
  - Division Ginglymodi Cope 1871
    - Order †Dapediiformes Thies & Waschkewitz 2015
    - Order †Semionotiformes Arambourg & Bertin 1958
    - Order Lepisosteiformes Hay 1929 (gars)
- Teleosteomorpha Arratia 2000 sensu Arratia 2013
  - Order †Prohaleciteiformes Arratia 2017
  - Division Aspidorhynchei Nelson, Grand & Wilson 2016
    - Order †Aspidorhynchiformes Bleeker 1859
    - Order †Pachycormiformes Berg 1937
  - Infraclass Teleostei Müller 1844 sensu Arratia 2013
    - Order †?Araripichthyiformes
    - Order †?Ligulelliiformes Taverne 2011
    - Order †?Tselfatiiformes Nelson 1994
    - Order †Pholidophoriformes Berg 1940
    - Order †Dorsetichthyiformes Nelson, Grand & Wilson 2016
    - Order †Leptolepidiformes
    - Order †Crossognathiformes Taverne 1989
    - Order †Ichthyodectiformes Bardeck & Sprinkle 1969
    - Teleocephala de Pinna 1996 s.s.
      - Megacohort Elopocephalai Patterson 1977 sensu Arratia 1999 (Elopomorpha Greenwood et al. 1966)
        Order Elopiformes Gosline 1960 (ladyfishes and tarpon)
        Order Albuliformes Greenwood et al. 1966 sensu Forey et al. 1996 (bonefishes)
        Order Notacanthiformes Goodrich 1909 (halosaurs and spiny eels)
        Order Anguilliformes Jarocki 1822 sensu Goodrich 1909 (true eels)
      - Megacohort Osteoglossocephalai sensu Arratia 1999
        Supercohort Osteoglossocephala sensu Arratia 1999 (Osteoglossomorpha Greenwood et al. 1966)
        Order †Lycopteriformes Chang & Chou 1977
        Order Hiodontiformes McAllister 1968 sensu Taverne 1979 (mooneye and goldeye)
        Order Osteoglossiformes Regan 1909 sensu Zhang 2004 (bony-tongued fishes)
        Supercohort Clupeocephala Patterson & Rosen 1977 sensu Arratia 2010
        Cohort Otomorpha Wiley & Johnson 2010 (Otocephala; Ostarioclupeomorpha)
        Subcohort Clupei Wiley & Johnson 2010 (Clupeomorpha Greenwood et al. 1966)
        Order †Ellimmichthyiformes Grande 1982
        Order Clupeiformes Bleeker 1859 (herrings and anchovies)
        Subcohort Alepocephali
        Order Alepocephaliformes Marshall 1962
        Subcohort Ostariophysi Sagemehl 1885
        Section Anotophysa (Rosen & Greenwood 1970) Sagemehl 1885
        Order †Sorbininardiformes Taverne 1999
        Order Gonorynchiformes Regan 1909 (milkfishes)
        Section Otophysa Garstang 1931
        Order Cypriniformes Bleeker 1859 sensu Goodrich 1909 (barbs, carp, danios, goldfishes, loaches, minnows, rasboras)
        Order Characiformes Goodrich 1909 (characins, pencilfishes, hatchetfishes, piranhas, tetras, dourado / golden (genus Salminus) and pacu)
        Order Gymnotiformes Berg 1940 (electric eels and knifefishes)
        Order Siluriformes Cuvier 1817 sensu Hay 1929 (catfishes)
        Cohort Euteleosteomorpha (Greenwood et al. 1966) (Euteleostei Greenwood 1967 sensu Johnson & Patterson 1996)
        Subcohort Lepidogalaxii
        Order Lepidogalaxiiformes Betancur-Rodriguez et al. 2013 (salamanderfish)
        Subcohort Protacanthopterygii Greenwood et al. 1966 sensu Johnson & Patterson 1996
        Order Argentiniformes (barreleyes and slickheads) (formerly in Osmeriformes)
        Order Galaxiiformes
        Order Salmoniformes Bleeker 1859 sensu Nelson 1994 (salmon, trout and pike)
        Subcohort Stomiati
        Order Osmeriformes (smelts)
        Order Stomiiformes Regan 1909 (bristlemouths and marine hatchetfishes)
        Subcohort Neoteleostei Nelson 1969
        Infracohort Ateleopodia
        Order Ateleopodiformes (jellynose fish)
        Infracohort Eurypterygia Rosen 1973
        Section Aulopa [Cyclosquamata Rosen 1973]
        Order Aulopiformes Rosen 1973 (Bombay duck and lancetfishes)
        Section Ctenosquamata Rosen 1973
        Subsection Myctophata [Scopelomorpha]
        Order Myctophiformes Regan 1911 (lanternfishes)
        Subsection Acanthomorpha Betancur-Rodriguez et al. 2013
        Division Lampridacea Betancur-Rodriguez et al. 2013 [Lampridomorpha; Lampripterygii]
        Order Lampriformes Regan 1909 (oarfish, opah and ribbonfishes)
        Division Paracanthomorphacea sensu Grande et al. 2013 (Paracanthopterygii Greenwood 1937)
        Order Percopsiformes Berg 1937 (cavefishes and trout-perches)
        Order †Sphenocephaliformes Rosen & Patterson 1969
        Order Zeiformes Regan 1909 (dories)
        Order Gadiformes Goodrich 1909 (cods)
        Division Polymixiacea Betancur-Rodriguez et al. 2013 (Polymyxiomorpha; Polymixiipterygii)
        Order †Pattersonichthyiformes Gaudant 1976
        Order †Ctenothrissiformes Berg 1937
        Order Polymixiiformes Lowe 1838 (beardfishes)
        Division Euacanthomorphacea Betancur-Rodriguez et al. 2013 (Euacanthomorpha sensu Johnson & Patterson 1993; Acanthopterygii Gouan 1770 sensu])
        Order Trachichthyiformes (fangtooths and pineconefishes)
        Subdivision Berycimorphaceae Betancur-Rodriguez et al. 2013
        Order Beryciformes (alfonsinos and holocentrids) (incl. Holocentriformes, Stephanoberyciformes; Cetomimiformes)
        Subdivision Percomorphaceae Betancur-Rodriguez et al. 2013 (Percomorpha sensu Miya et al. 2003; Acanthopteri)
        Series Ophidiimopharia Betancur-Rodriguez et al. 2013
        Order Ophidiiformes (pearlfishes)
        Series Batrachoidimopharia Betancur-Rodriguez et al. 2013
        Order Batrachoidiformes (toadfishes)
        Series Gobiomopharia Betancur-Rodriguez et al. 2013
        Order Gobiiformes (cardinalfishes, sleepers and gobies)
        Series Syngnathomorpharia Betancur-Rodriguez et al. 2013
        Order Syngnathiformes (seahorses, pipefishes, sea moths, cornetfishes and flying gurnards)
        Order Scombriformes (Tunas and (mackerels)
        Series Carangimopharia Betancur-Rodriguez et al. 2013
        Subseries Anabantaria Betancur-Rodriguez et al. 2014
        Order Synbranchiformes (swamp eels)
        Order Anabantiformes (Labyrinthici) (gouramies, snakeheads, )
        Subseries Carangaria Betancur-Rodriguez et al. 2014
        Order Carangiformes (Jack mackerels, pompanos, flatfishes, billfishes)
        Subseries Ovalentaria Smith & Near 2012 (Stiassnyiformes sensu Li et al. 2009)
        Order Atheriniformes Rosen 1964 (silversides and rainbowfishes)
        Order Cyprinodontiformes Berg 1940 (livebearers, killifishes)
        Order Beloniformes Berg 1940 (flyingfishes and ricefishes)
        Order Cichliformes Betancur-Rodriguez et al. 2013 (Cichlids, Convict blenny, leaf fishes)
        Order Mugiliformes Berg 1940 (mullets)
        Order Blenniiformes Springer 1993 (Blennies, damselfish, Clingfishes)
        Series Eupercaria Betancur-Rodriguez et al. 2014 (Percomorpharia Betancur-Rodriguez et al. 2013)
        Order Perciformes Bleeker 1859
        Order Centrarchiformes Bleeker 1859 (Sunfishes and mandarin fishes)
        Order Labriformes (Wrasses and Parrotfishes)
        Order Acropomatiformes
        Order Acanthuriformes
        Order Lophiiformes Garman 1899 (Anglerfishes)
        Order Tetraodontiformes Regan 1929 (Filefishes and pufferfish)

Characteristics

Body shapes and fin arrangements

Reproduction

Classification and fossil record

Taxonomy

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