Actinopterygii

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Ray-finned fish
Temporal range:
Late SilurianPresent, 425–0  Ma [1]
Actinopterygii.jpgElectric eelSpotfin lionfishJapanese pineconefishTwo-banded seabream
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
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 fish or actinopterygians, is a class of bony fish [2] that comprise over 50% of living vertebrate species. [3] They 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 lobed fins of the sister class Sarcopterygii (lobe-finned fish). Resembling folding fans, the actinopterygian fins can easily change shape 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).

Contents

The vast majority of actinopterygians are teleosts. By species count, they dominate the subphylum Vertebrata, and constitute nearly 99% of the over 30,000 extant species of fish. [4] They are the most abundant nektonic aquatic animals and are ubiquitous throughout freshwater and marine environments from the deep sea to subterranean waters 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 to the giant oarfish, at 11 m (36 ft). The largest ever known ray-finned fish, the extinct Leedsichthys from the Jurassic, has been estimated to have grown to 16.5 m (54 ft).

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 typical ray-finned fish are shown in the adjacent diagram.

The swim bladder is a more derived structure and used for buoyancy. [5] 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. [6] [5] In early forms the swim bladder could still be used for breathing, a trait still present in Holostei (bowfins and gars). [7] In some fish like the arapaima, the swim bladder has been modified for breathing air again, [8] and in other lineages it have been completely lost. [9]

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. [10]

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). [11] [12] [13] [14] [15]

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 (Gasterosteus aculeatus) 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 (Gasterosteus aculeatus) 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. [16] 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. [17]

Most families use external rather than internal fertilization. [18] Of the oviparous teleosts, most (79%) do not provide parental care. [19] 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. [19] The oldest case of viviparity in ray-finned fish is found in Middle Triassic species of Saurichthys . [20] Viviparity is relatively rare and is found in about 6% of living teleost species; male care is far more common than female care. [19] [21] Male territoriality "preadapts" a species for evolving male parental care. [22] [23]

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. [24]

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. [25]

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). [26] [27] 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. [28] Approximate divergence dates for the different actinopterygian clades (in millions of years, mya) are from Near et al., 2012. [26]

Vertebrates

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. [26]

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. [29] Crown group actinopterygians most likely originated near the Devonian-Carboniferous boundary. [30] The earliest fossil relatives of modern teleosts are from the Triassic period ( Prohalecites , Pholidophorus ), [31] [32] although it is suspected that teleosts originated already during the Paleozoic Era. [26]

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. [33] 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. [33]
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. [34]
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 Eocene carangiform +Mene oblonga Mene oblonga 23.JPG
Fossil of the Eocene carangiform Mene oblonga
Fossil of the Eocene pleuronectiform +Amphistium paradoxum Amphistium.JPG
Fossil of the Eocene 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 [27] [35] with notes when this differs from Nelson, [3] ITIS [36] and FishBase [37] and extinct groups from Van der Laan 2016 [38] and Xu 2021. [39]

Related Research Articles

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

Osteichthyes, commonly referred to as the bony fish but in the 21st century also treated as a clade that includes the tetrapods, is a diverse superclass of vertebrate animals 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 extant fish are members of Osteichthyes, an extremely diverse and abundant group consisting of 45 orders, over 435 families and 28,000 species. It is the largest class of vertebrates in existence today.

<span class="mw-page-title-main">Bowfin</span> Bony fish related to gars in the infraclass Holosteii

The bowfin is a bony fish, native to North America. Common names include mudfish, mud pike, dogfish, grindle, grinnel, swamp trout, and choupique. It is regarded as a relict, being one of only two surviving species of the Halecomorphi, a group of fish that first appeared during the Early Triassic, around 250 million years ago. The bowfin is often considered a "primitive fish" because they have retained some morphological characteristics of their early ancestors. It is one of two species in the genus Amia, along with Amia ocellicauda, the eyespot bowfin. The closest living relatives of bowfins are gars, with the two groups being united in the clade Holostei.

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

The Amiiformes order of fish has only two extant species, the bowfins: Amia calva and Amia ocellicauda, the latter recognized as a separate species in 2022. 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. The order first appeared in the Triassic, and the extinct members include both marine and freshwater species, many of which are morphologically disparate from bowfins, such as the caturids.

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

Semionotiformes is an order of ray-finned fish known from the Middle Triassic (Anisian) to the Late Cretaceous (Maastrichtian). Their closest living relatives are gars (Lepisosteidae), with both groups belonging to the clade Ginglymodi within the Holostei. The group includes both freshwater (Semionotidae) and marine adapted forms. Many members of the family Macrosemiidae, had elongated dorsal fins, often associated with an adjacent area of skin which was free of scales. These fins were likely undulated for use in precision swimming. The body morphology of macrosemiids suggests that they were slow swimmers that were capable of maneuvering around complex topography, such as reef environments.

<span class="mw-page-title-main">Euteleostomi</span> Clade including most vertebrates

Euteleostomi is a successful clade that includes more than 90% of the living species of vertebrates. Both its major subgroups are successful today: Actinopterygii includes most extant bony fish species, and Sarcopterygii includes the tetrapods.

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

Actinopteri is the sister group of Cladistia (bichirs) 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 the single living genus, Amia with two species, the bowfins, 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, which are putative "semionotiforms" such as Acentrophorus and Archaeolepidotus, are known from the Middle to Late Permian and are among the earliest known neopterygians.

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

The Palaeonisciformes, commonly known as "palaeoniscoids" are an extinct grouping of primitive ray-finned fish (Actinopterygii), spanning from the Silurian/Devonian to the Cretaceous. They are generally considered paraphyletic, but their exact relationships to living ray-finned fish are uncertain. While some and perhaps most palaeoniscoids likely belong to the stem-group of Actinopteryii, it has been suggested that some may belong to the crown group, with some of these possibly related to Cladistia and/or Chondrostei. Many palaeoniscoids share a conservative body shape and a similar arrangement of skull bones.

<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. 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. Pachycormiformes are morphologically diverse, containing both tuna and swordfish-like carnivorous forms, as well as edentulous suspension-feeding forms.

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

Halecostomi is the name of a group of neopterygian fish uniting the halecomorphs and the teleosts, the largest group of extant ray-finned fish.

<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 of tropical Africa. 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">Halecomorphi</span> Clade of fishes

Halecomorphi is a taxon of ray-finned bony fish in the clade Neopterygii. The only extant Halecomorph species are the bowfin and eyespot 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 with more than 17 000 known species 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 are two living species in Amia, Amia calva and Amia ocellicauda, 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> Tribe of fishes

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.

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  40. In Nelson, Polypteriformes is placed in its own subclass Cladistia.
  41. In Nelson and ITIS, Syngnathiformes is placed as the suborder Syngnathoidei of the order Gasterosteiformes.