Osteichthyes | |
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Example of Osteichthyes: Queensland lungfish and West Indian Ocean coelacanth (two Sarcopterygii), iridescent shark and American black sturgeon (two Actinopterygii) | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Subphylum: | Vertebrata |
Infraphylum: | Gnathostomata |
Clade: | Eugnathostomata |
Clade: | Teleostomi |
Superclass: | Osteichthyes Huxley, 1880 |
Classes | |
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Osteichthyes ( /ˌɒstiːˈɪkθi.iːz/ ), also known as osteichthyans or commonly referred to as the bony fish, is a diverse superclass of vertebrate animals that have endoskeletons primarily composed of bone tissue. They can be contrasted with the Chondrichthyes (cartilaginous fish) and the extinct placoderms and acanthodians, which have endoskeletons primarily composed of cartilage. The vast majority of extant fish are members of Osteichthyes, being an extremely diverse and abundant group consisting of 45 orders, over 435 families and 28,000 species. [2] It is the largest class of vertebrates in existence today, encompassing most aquatic vertebrates, as well as all semi-aquatic and terrestrial vertebrates.
The group is divided into two main clades, the ray-finned fish (Actinopterygii, which makes up the vast majority of extant fish) and the lobe-finned fish (Sarcopterygii, which gave rise to all land vertebrates, i.e. tetrapods). The oldest known fossils of bony fish are about 425 million years old from the late Silurian, [1] which are also transitional fossils showing a tooth pattern that is in between the tooth rows of sharks and true bony fishes. [3] Despite the name, these early basal bony fish had not yet evolved ossification and their skeletons were still mostly cartilaginous, and the main distinguishing feature that set them apart from other fish clades were the development of foregut pouchs that eventually evolved into the swim bladders and lungs, respectively.
Osteichthyes can be compared to Euteleostomi. In paleontology the terms are synonymous. In ichthyology the difference is that Euteleostomi presents a cladistic view which includes the terrestrial tetrapods that evolved from lobe-finned fish. Until recently, the view of most ichthyologists has been that Osteichthyes were paraphyletic and include only fishes. [4] However, since 2013 widely cited ichthyology papers have been published with phylogenetic trees that treat the Osteichthyes as a clade including tetrapods. [5] [6] [7] [4]
Bony fish are characterized by a relatively stable pattern of cranial bones, rooted, medial insertion of mandibular muscle in the lower jaw. The head and pectoral girdles are covered with large dermal bones. The eyeball is supported by a sclerotic ring of four small bones, but this characteristic has been lost or modified in many modern species. The labyrinth in the inner ear contains large otoliths. The braincase, or neurocranium, is frequently divided into anterior and posterior sections divided by a fissure.
Early bony fish had simple respiratory diverticula (an outpouching on either side of the esophagus) which helped them breathe air in low-oxygen water as a form of supplementary enteral respiration. In ray-finned fish these have evolved into swim bladders, the changing sizes of which help to alter the body's specific density and buoyancy. In elpistostegalians, a crown group of lobe-finned fish that gave rise to the land-dwelling tetrapods, these respiratory diverticula became further specialized for obligated air breathing and evolved into the modern amphibian, reptilian, avian and mammalian lungs. [8] [9] [10] Early bony fish did not have fin spines like most modern fish, but instead had the fleshy paddle-like fins similar other non-bony clades of fish, although the lobe-finned fish evolved articulated appendicular skeletons within their paired fins, which gave rise to tetrapods' limbs. They also evolved a pair of opercula (gill covers), which can actively draw water across the gills so they can breathe without having to swim.
Bony fish do not have placoid scales like cartilaginous fish, instead they consist of three types of scales that do not penetrate the epidermis in the process. The three categories of scales for Osteichthyes which are cosmoid scales, ganoid scales, teleost scales. The teleost scales are also then divided into two subgroups which are the cycloid scales, and the ctenoid scales. All these scales have a base of bone that they all originate from, the only difference is that the teleost scales only have one layer of bone. Ganoid scales have lamellar bone, and vascular bone that lies on top of the lamellar bone, then enamel lies on top of both layers of bone. Cosmoid scales have the same two layers of bone that ganoid scales have expect they gave dentin in-between the enamel and vascular bone and lamellar (vascular and lamellar two subcategories for bone found in scales). All these scales are found underneath the epidermis and do not break the epidermis of the fish. Unlike the placoid scales that poke through the epidermis of the fish.
...it is increasingly widely accepted that tetrapods, including ourselves, are simply modified bony fishes, and so we are comfortable with using the taxon Osteichthyes as a clade, which now includes all tetrapods...
Fishes of the World (5th ed) [4]
Traditionally, Osteichthyes was considered a class, recognised on the presence of a swim bladder, only three pairs of gill arches hidden behind a bony operculum, and a predominantly bony skeleton. [11] Under this classification system, Osteichthyes was considered paraphyletic with regard to land vertebrates, as the common ancestor of all osteichthyans includes tetrapods amongst its descendants. While the largest subclass, Actinopterygii (ray-finned fish), is monophyletic, with the inclusion of the smaller sub-class Sarcopterygii, Osteichthyes was regarded as paraphyletic.
This has led to the current cladistic classification which splits the Osteichthyes into two full classes. Under this scheme Osteichthyes is monophyletic, as it includes the tetrapods making it a synonym of the clade Euteleostomi. Most bony fish belong to the ray-finned fish (Actinopterygii).
Actinopterygii | ray-finned fish | Actinopterygii, members of which are known as ray-finned fishes, is a class or subclass of the bony fishes. The ray-finned fishes are so called because they possess lepidotrichia or "fin rays", their fins being webs of skin supported by bony or horny spines ("rays"), as opposed to the fleshy, lobed fins that characterize the class Sarcopterygii which also possess lepidotrichia. 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). In terms of numbers, actinopterygians are the dominant class of vertebrates, comprising nearly 99% of the over 30,000 species of fish (Davis, Brian 2010). 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, to at least 11 m (36 ft). |
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Sarcopterygii | lobe-finned fish | Sarcopterygii (fleshy fin), members of which are known as lobe-finned fish, is a group of the bony fishes. Traditionally, it is a class or subclass that excludes Tetrapoda, a group of typically terrestrial vertebrates that descends from lobe-finned fish. However, under modern cladistic classification schemes, Sarcopterygii is a clade that includes the tetrapods. The living sarcopterygians are the coelacanths, lungfish, and the tetrapods. Early lobe-finned fishes had fleshy, lobed, paired fins, joined to the body by a single bone. [12] Their fins differ from those of all other fish in that each is borne on a fleshy, lobelike, scaly stalk extending from the body. Pectoral and pelvic fins have articulations resembling those of tetrapod limbs. These fins evolved into legs of the first tetrapod land vertebrates, amphibians. They also possess two dorsal fins with separate bases, as opposed to the single dorsal fin of actinopterygians (ray-finned fish). The braincase of sarcoptergygians primitively has a hinge line, but this is lost in tetrapods and lungfish. Many early lobe-finned fishes have a symmetrical tail. All lobe-finned fishes possess teeth covered with true enamel. |
A phylogeny of living Osteichthyes, including the tetrapods, is shown in the cladogram below. [5] [13] [14] [15] Whole-genome duplication took place in the ancestral Osteichthyes. [16]
Osteichthyes/ | |
Euteleostomi |
All bony fish possess gills. For the majority this is their sole or main means of respiration. Lungfish and other osteichthyan species are capable of respiration through lungs or vascularized swim bladders. Other species can respire through their skin, intestines, and/or stomach. [17]
Osteichthyes are primitively ectothermic (cold blooded), meaning that their body temperature is dependent on that of the water. But some of the larger marine osteichthyids, such as the opah, [18] [19] swordfish [20] [21] and tuna [22] have independently evolved various levels of endothermy. Bony fish can be any type of heterotroph: numerous species of omnivore, carnivore, herbivore, filter-feeder or detritivore are documented.
Some bony fish are hermaphrodites, and a number of species exhibit parthenogenesis. Fertilization is usually external, but can be internal. Development is usually oviparous (egg-laying) but can be ovoviviparous, or viviparous. Although there is usually no parental care after birth, before birth parents may scatter, hide, guard or brood eggs, with sea horses being notable in that the males undergo a form of "pregnancy", brooding eggs deposited in a ventral pouch by a female.
The ocean sunfish is the heaviest bony fish in the world, [23] in late 2021, Portuguese fishermen found a dead sunfish near the coast of Faial Island, Azores, with a weight of 2,744 kilograms (6,049 lb) and 3.6 metres (12 ft) tall and 3.5 metres (11 ft) long established the biggest ocean sunfish ever captured. [24]
The longest is the king of herrings, a type of oarfish. Other very large bony fish include the Atlantic blue marlin, some specimens of which have been recorded as in excess of 820 kilograms (1,810 lb), the black marlin, some sturgeon species, and the giant and goliath grouper, which both can exceed 300 kilograms (660 lb) in weight. In contrast, Paedocypris progenetica and the stout infantfish can measure less than 8 millimetres (0.31 in). [25] [26] The beluga sturgeon is the largest species of freshwater bony fish extant today, and Arapaima gigas is among the largest of the freshwater fish. The largest bony fish ever was Leedsichthys , which dwarfed the beluga sturgeon as well as the ocean sunfish, giant grouper and all the other giant bony fishes alive today. [27]
Comparison of cartilaginous and bony fishes [28] | ||
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Characteristic | Sharks (cartilaginous) | Bony fishes |
Habitat | Mainly marine | Marine and freshwater |
Shape | Usually dorso-ventrally flattened | Usually bilaterally flattened |
Exoskeleton | Separate dermal placoid scales | Overlapping dermal cosmoid, ganoid, cycloid or ctenoid scales |
Endoskeleton | Cartilaginous | Mostly bony |
Caudal fin | Heterocercal | Heterocercal or diphycercal |
Pelvic fins | Usually posterior. | Mostly anterior, occasionally posterior. |
Intromittent organ | Males use pelvic fins as claspers for transferring sperm to a female | Do not use claspers, though some species use their anal fins as gonopodium for the same purpose |
Mouth | Large, crescent shaped on the ventral side of the head | Variable shape and size at the tip or terminal part of the head |
Jaw suspension | Hyostylic | Hyostylic and autostylic |
Gill openings | Usually five pairs of gill slits which are not protected by an operculum. | Five pairs of gill slits protected by an operculum (a lateral flap of skin). |
Type of gills | Larnellibranch with long interbranchial septum | Filiform with reduced interbranchial septum |
Spiracles | The first gill slit usually becomes spiracles opening behind the eyes. | No spiracles |
Afferent branchial vessels | Five pairs from ventral aorta to gills | Only four pairs |
Efferent branchial vessels | Nine pairs | Four pairs |
Conus arteriosus | Present in heart | Absent |
Cloaca | A true cloaca is present only in cartilaginous fishes and lobe-finned fishes. | In most bony fishes, the cloaca is absent, and the anus, urinary and genital apertures open separately [29] |
Stomach | Typically J-shaped | Shape variable. Absent in some. |
Intestine | Short with spiral valve in lumen | Long with no spiral valve |
Rectal gland | Present | Absent |
Liver | Usually has two lobes | Usually has three lobes |
Swim bladder | Absent | Usually present |
Brain | Has large olfactory lobes and cerebrum with small optic lobes and cerebellum | Has small olfactory lobes and cerebrum and large optic lobes and cerebellum |
Restiform bodies | Present in brain | Absent |
Ductus endolymphaticus | Opens on top of head | Does not open to exterior |
Retina | Lacks cones | Most fish have double cones, a pair of cone cells joined to each other. |
Accommodation of eye | Accommodate for near vision by moving the lens closer to the retina | Accommodate for distance vision by moving the lens further from the retina [30] |
Ampullae of Lorenzini | Present | Absent |
Male genital duct | Connects to the anterior part of the genital kidney | No connection to kidney |
Oviducts | Not connected to ovaries | Connected to ovaries |
Urinary and genital apertures | United and urinogenital apertures lead into common cloaca | Separate and open independently to exterior |
Eggs | A small number of large eggs with plenty of yolk | A large number of small eggs with little yolk |
Fertilisation | Internal | Usually external |
Development | Ovoviviparous types develop internally. Oviparous types develop externally using egg cases | Normally develop externally without an egg case |
Actinopterygii, members of which are known as ray-finned fish or actinopterygians, 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. 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.
Chondrichthyes is a class of jawed fish that contains the cartilaginous fish or chondrichthyians, which all have skeletons primarily composed of cartilage. They can be contrasted with the Osteichthyes or bony fish, which have skeletons primarily composed of bone tissue. Chondrichthyes are aquatic vertebrates with paired fins, paired nares, placoid scales, conus arteriosus in the heart, and a lack of opecula and swim bladders. Within the infraphylum Gnathostomata, cartilaginous fishes are distinct from all other jawed vertebrates.
Sarcopterygii — sometimes considered synonymous with Crossopterygii — is a clade of bony fish commonly referred to as lobe-finned fish. They are characterised by prominent muscular limb buds (lobes) within their fins, which are supported by articulated appendicular skeletons. This is in contrast to the other clade of bony fish, the Actinopterygii, which have only skin-covered bony spines supporting the fins.
Teleostei, members of which are known as teleosts, is, by far, the largest infraclass in the class Actinopterygii, the ray-finned fishes, and contains 96% of all extant species of fish. Teleosts are arranged into about 40 orders and 448 families. Over 26,000 species have been described. Teleosts range from giant oarfish measuring 7.6 m (25 ft) or more, and ocean sunfish weighing over 2 t, to the minute male anglerfish Photocorynus spiniceps, just 6.2 mm (0.24 in) long. Including not only torpedo-shaped fish built for speed, teleosts can be flattened vertically or horizontally, be elongated cylinders or take specialised shapes as in anglerfish and seahorses.
Fish anatomy is the study of the form or morphology of fish. It can be contrasted with fish physiology, which is the study of how the component parts of fish function together in the living fish. In practice, fish anatomy and fish physiology complement each other, the former dealing with the structure of a fish, its organs or component parts and how they are put together, such as might be observed on the dissecting table or under the microscope, and the latter dealing with how those components function together in living fish.
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.
Actinopteri is the sister group of Cladistia (bichirs) in the class Actinopterygii.
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.
Acanthodii or acanthodians is an extinct class of gnathostomes. They are currently considered to represent a paraphyletic grade of various fish lineages basal to extant Chondrichthyes, which includes living sharks, rays, and chimaeras. Acanthodians possess a mosaic of features shared with both osteichthyans and chondrichthyans. In general body shape, they were similar to modern sharks, but their epidermis was covered with tiny rhomboid platelets like the scales of holosteians.
Teleostomi is an obsolete taxon of jawed vertebrates that supposedly includes the tetrapods, bony fish, and the wholly extinct acanthodian fish. Key characters of this group include an operculum and a single pair of respiratory openings, features which were lost or modified in some later representatives. The teleostomes include all jawed vertebrates except the chondrichthyans and the extinct class Placodermi.
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.
Andreolepis is an extinct genus of prehistoric fish, which lived around 420 million years ago. It was described by Walter Gross in 1968 based on scales found in the Hemse Formation in Gotland, Sweden. It is placed in the monogeneric family Andreolepididae and is generally regarded as a primitive member of the class Actinopterygii based on its ganoid scale structure; however some new research regards it as a stem group of osteichthyans.
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.
A fish scale is a small rigid plate that grows out of the skin of a fish. The skin of most jawed fishes is covered with these protective scales, which can also provide effective camouflage through the use of reflection and colouration, as well as possible hydrodynamic advantages. The term scale derives from the Old French escale, meaning a shell pod or husk.
Fins are moving appendages protruding from the body of fish that interact with water to generate thrust and help the fish swim. Apart from the tail or caudal fin, fish fins have no direct connection with the spine and are supported only by muscles.
The evolution of fish began about 530 million years ago during the Cambrian explosion. It was during this time that the early chordates developed the skull and the vertebral column, leading to the first craniates and vertebrates. The first fish lineages belong to the Agnatha, or jawless fish. Early examples include Haikouichthys. During the late Cambrian, eel-like jawless fish called the conodonts, and small mostly armoured fish known as ostracoderms, first appeared. Most jawless fish are now extinct; but the extant lampreys may approximate ancient pre-jawed fish. Lampreys belong to the Cyclostomata, which includes the extant hagfish, and this group may have split early on from other agnathans.
The evolution of tetrapods began about 400 million years ago in the Devonian Period with the earliest tetrapods evolved from lobe-finned fishes. Tetrapods are categorized as animals in the biological superclass Tetrapoda, which includes all living and extinct amphibians, reptiles, birds, and mammals. While most species today are terrestrial, little evidence supports the idea that any of the earliest tetrapods could move about on land, as their limbs could not have held their midsections off the ground and the known trackways do not indicate they dragged their bellies around. Presumably, the tracks were made by animals walking along the bottoms of shallow bodies of water. The specific aquatic ancestors of the tetrapods, and the process by which land colonization occurred, remain unclear. They are areas of active research and debate among palaeontologists at present.
Cladistic classification of Sarcopterygii is the classication of Sarcopterygii as a clade containing not only the lobe-finned fishes but also the tetrapods, which are closely related to lungfish. The taxon Sarcopterygii was traditionally classified as a paraphyletic group considered either a class or a subclass of Osteichthyes. Identification of the group is based on several characteristics, such as the presence of fleshy, lobed, paired fins, which are joined to the body by a single bone.
Fishes are a paraphyletic group and for this reason, the class Pisces seen in older reference works is no longer used in formal taxonomy. Traditional classification divides fish into three extant classes, and with extinct forms sometimes classified within those groups, sometimes as their own classes:
The evolution of fishes took place over a timeline which spans the Cambrian to the Cenozoic, including during that time in particular the Devonian, which has been dubbed the "age of fishes" for the many changes during that period.
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