22-461: The Esociformes ( / ˈ iː s ɒ s ɪ f ɔːr m iː z / ) is a small order of freshwater ray-finned fish , with two families, Umbridae and Esocidae . The pikes of genus Esox give the order its name. This order is closely related to the Salmoniformes , the two comprising the superorder Protacanthopterygii , and are often included in their order. The esociform fishes first appeared in
44-508: A holarctic distribution. Two additional genera have been described from fossils dating to the Cretaceous of North America. Umbra remains the only extant genus in this family, and can be found in eastern North America and Europe. Three additional genera have been described from fossils dating from the Paleocene of Europe; however, genetic studies on the extant species of Umbra have recovered
66-459: 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
88-717: A split between the North American and European species dating to the Late Cretaceous and earliest half of the Paleogene . While the family Esocidae traditionally only contained the genus Esox, recent genetic and paleontological research have recovered Novumbra and Dallia as members of the family Esocidae, being closer related to Esox than Umbra . Umbra is the only remaining extant species in Umbridae. Various fossils have been described as members of Esociformes and are placed on
110-726: 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 have 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
132-587: Is a class of bony fish that comprise over 50% of living vertebrate species. 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
154-540: 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: The cladogram below shows
176-432: 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
198-500: The Cyprinidae (in goldfish and common carp as recently as 14 million years ago). Ray-finned fish vary in size and shape, in their feeding specializations, and in the number and arrangement of their ray-fins. 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
220-587: 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). Ray-finned fishes occur in many variant forms. The main features of typical ray-finned fish are shown in
242-481: 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
SECTION 10
#1732854875378264-651: 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
286-930: 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 . 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 with notes when this differs from Nelson, ITIS and FishBase and extinct groups from Van der Laan 2016 and Xu 2021. [REDACTED] [REDACTED] [REDACTED] [REDACTED] [REDACTED] [REDACTED] [REDACTED] Wetted area In fluid dynamics ,
308-402: 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,
330-463: 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
352-706: The different actinopterygian clades (in millions of years , mya) are from Near et al., 2012. Jaw-less fishes ( hagfish , lampreys ) [REDACTED] Cartilaginous fishes ( sharks , rays , ratfish ) [REDACTED] Coelacanths [REDACTED] Lungfish [REDACTED] Amphibians [REDACTED] Mammals [REDACTED] Sauropsids ( reptiles , birds ) [REDACTED] Polypteriformes ( bichirs , reedfishes ) [REDACTED] Acipenseriformes ( sturgeons , paddlefishes ) [REDACTED] Teleostei [REDACTED] Amiiformes ( bowfins ) [REDACTED] Lepisosteiformes ( gars ) [REDACTED] The polypterids (bichirs and reedfish) are
374-432: 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. Actinopterygii is divided into the subclasses Cladistia , Chondrostei and Neopterygii . The Neopterygii , in turn,
396-549: The following tree accordingly. Esox [REDACTED] Novumbra [REDACTED] Dallia [REDACTED] † Oldmanesox † Estesesox † Boltyshia † Palaeoesox [REDACTED] † Proumbra Umbra [REDACTED] Actinopterygii Actinopterygii ( / ˌ æ k t ɪ n ɒ p t ə ˈ r ɪ dʒ i aɪ / ; from actino- 'having rays' and Ancient Greek πτέρυξ (ptérux) 'wing, fins'), members of which are known as ray-finned fish or actinopterygians ,
418-438: 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
440-461: 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
462-566: The mid- Cretaceous — early products of the Euteleostei radiation of that time. They diverged from their sister group Salmoniformes about 110 million years ago, with the extant species having evolved from a common ancestor that lived about 90 million years ago. Today, they are found in weed-choked freshwater habitats in North America and northern Eurasia . The three extant esocid genera ( Esox , Novumbra , and Dallia ) together comprise
SECTION 20
#1732854875378484-494: 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). 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 . They are the most abundant nektonic aquatic animals and are ubiquitous throughout freshwater and marine environments from
#377622