Bony fish

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Bony fish

Obsolete systematic group

The taxon dealt with here is not part of the systematics presented in the German-language Wikipedia. More information can be found in the article text.

Dark giant grouper (Epinephelus lanceolatus) in the Georgia Aquarium in Atlanta, USA

Dark giant grouper ( Epinephelus lanceolatus )
in the Georgia Aquarium in Atlanta , USA

Systematics
without rank: Bilateria
Over trunk : Neumünder (Deuterostomia)
Trunk : Chordates (chordata)
Sub-stem : Vertebrates (vertebrata)
Superclass : Jaw mouths (Gnathostomata)
Paraphyletic taxon :
Row : Bony fish
Scientific name
Osteichthyes
Huxley , 1880

Bony fish (Osteichthyes, from ancient Greek ὀστέον ostéon "bones" and ἰχθύς ichthýs "fish") or bony fish in the broader sense are, according to traditional understanding, those fish whose skeleton, in contrast to that of cartilage fish (Chondrichthyes), is completely or partially ossified . The bony fish in the narrower sense, the real bony fish (Teleostei), are to be distinguished from the Osteichthyes as a subordinate taxon .

The bony fish are divided into two large groups: the ray fins (Actinopterygii) and the meat fins (Sarcopterygii). From the point of view of modern systematics ( cladistics ), however, the terrestrial vertebrates (Tetrapoda) also belong to the meat fins and thus to the bony fish. In the traditional system, however, the land vertebrates are not counted among the bony fish. The bony fish in the classical sense are therefore not a natural family group (not a monophyletic taxon). The classic taxon "Osteichthyes" is therefore used less and less in the zoological and palaeontological system, and the taxon name only serves as an informal collective term for fish with a bony skeleton or with a cartilaginous but originally bony skeleton (e.g. sturgeon and sunfish ).

General

With the exception of the lampreys , all fish species living in inland waters of Central Europe belong to the bony fish and - with the exception of the sturgeon-like - to the real bony fish.

The bony fish are the most species-rich group of vertebrates (vertebrata). Even today, new species are continually being discovered both in the sea and in freshwater. The meat-fin fish , in whose fossil relationship the common ancestor of all terrestrial vertebrates can be found, are only represented with eight recent (today living) species. The ray fins make up with at least 29,000 recent species, more than 96% of the total recent fish fauna.

Ray fins and flesh fins have appeared in the fossil record since the Upper Silurian . In the subsequent Devonian, they rapidly split into numerous types ( adaptive radiation ).

features

External characteristics of a lantern fish :
1   gill cover
2   lateral line
3   dorsal fin (dorsalis)
4   adipose fin
5   tail stalk
6   caudal fin (caudalis)
7   anal fin (analis)
8   luminous organs
9   ventral fins (ventral)
10   pectoral fins (pectorales)

Most bony fish have a spindle-shaped body that offers little resistance to movement in the water. Paired and unpaired fins are used for propulsion and stabilization.

The fins anatomy can be used to distinguish between ray fins and flesh fins: The fins of the ray fins are formed by a series of radii, narrow rays of bone substance or cartilage . On the other hand, the fins of the meat fins are supported by a single, muscle-moving basal bone. The extremities of the terrestrial vertebrates emerged from the paired pectoral and ventral fins of this group .

There can be one, two or three dorsal fins depending on the fish family. Some, like the Old and New World knifefish , have also reduced them. Between the dorsal fin and the caudal fin, some taxa, like many Ostariophysi , still have a small, radiant adipose fin . The fins and the number and type of fin rays are important features in taxonomy .

The length of adult bony fish varies depending on the species between just under a centimeter ( Paedocypris progenetica ) and 8 meters ( Regalecus glesne ). The heaviest bony fish in the world is the sunfish ( Mola mola ) , which are up to three meters long, four meters high and weigh up to 2.3 tons .

Variety of body shapes

Great sea fish ( Phycodurus eques )

Some bony fish differ greatly in shape from the usual fish blueprint. The best-known example are the flatfish (Pleuronectiformes), which in the course of their ontogenesis lie on one side of the body, which from there forms the underside. The eye on this side moves to the other side of the body, which is from there on top; the pectoral fin on the blind side can atrophy. The flatfish lose their symmetry.

Other fish families, such as the frogfish (Antennariidae), the stonefish (Synanceiidae) and the shredded fish (Solegnathinae), dissolve their contours with a multitude of body outgrowths in order to be camouflaged either as predators or as potential prey.

The pelagic sunfish are taller than they are long, have no ventral or caudal fin and are also known as the "swimming head".

skin

The skin of the fish consists of two layers: the connective tissue-containing dermis, in which there are scales and color cells, and the epidermis with mucous glands. The mucus has a protective effect on the skin and the scales inside, as it has an antibacterial effect. The scales reduce the flow resistance because they increase the speed limit for the development of turbulence ( ideally the ctenoid scales ).

Anatomy and physiology

skeleton

The skeleton consists of bones , in primitive (but also derived) species partly made of cartilage . At the skull , the gill skeleton is made up of seven gill arches, one of which is transformed to the lower jaw forward. In the trunk region, the vertebrae carry ribs , the entire length of the spinous processes on the back, and in the tail area also on the abdomen. Bones of ossified connective tissue are often found in the muscle sheaths of bony fish . For the control and locomotion serving fins , which up to the dorsal fin (with salmon-like , Characins , catfish-like ) reinforced with bony fin rays (radii, resulting from scale rows). The pectoral and pectoral fins are paired and articulate on the shoulder girdle and the pelvis, respectively. The unpaired dorsal, anal and caudal fin (s) are connected to the spine via fin supports (radialia). The aforementioned adipose fin has no skeleton, but can be reinforced like a horn.

Skeleton of a perch

The illustration opposite shows the skeleton of a perch ( Perca fluviatilis ). The names of the most important bones in the largely internationally valid nomenclature are:

1  premaxillary (toothed), 2  maxillary , 3  lower jaw, consisting of dental (toothed), articular and angular, 4  eye socket (orbita), 5  (six) suborbitalia (the anterior, largest is called lacrimal), 6  preopercular, 7  interopercular, 8  Suboperculare, 9  operculare, 10  shoulder girdle with scapulare and coracoideum, 11  pectoralis (13 radii on four radials), 12  pelvic bones (pelvis), 13  pelvicalis (six radii; chest), 14  (40) vertebrae (vertebrae), 15  neurapophyses, 16  Haemapophyses, 17  ribs (costae), 18  bones (epipleuralia), 19, 20 fin carriers (pterygiophores), 21, 22 two dorsales, 23  analis, 24  hypuralia , 25  caudalis (with 17 radii).

Muscles

The muscles of the bony fish are divided into trunk and fin muscles . The trunk muscles consist of individual segments, the number of which usually corresponds to that of the vertebrae . The sunfish only has 17 vertebrae, the snipe eel ( Nemichthys scolopaceus ) over 600. Naturally, a large number of muscles are attached to the skull for food acquisition and breathing. They are segmented into myomers . The partitions between the myomers are called myosepta. The myomers are W-shaped when viewed from the side. In some species (especially herbivores) a muscular chewing stomach develops.

Circulation and breathing

Schematic representation of the blood circulation:
  • Oxygenated blood
  • Oxygenated blood
  • Bony fish have closed blood circulation with a simple heart close to the gills that consists of an atrium and a ventricle. It pumps venous blood directly into the gills, which consist of numerous leaflets with capillaries , fine skin protuberances. The gills absorb dissolved oxygen from the water over their large surface area and pass it on to the blood. The very high effectiveness of oxygen uptake via the gills (up to 70% of the available oxygen) can be attributed to the principle of countercurrent exchange in the gills and the very high affinity of some bony fish hemoglobins for oxygen.

    The gills are protected in the gill cavity by the bony gill cover (operculum). By lowering the floor of the mouth and lifting the gill cover at the same time, breathing water flows in through the mouth, while this is prevented by a soft membrane on the gill cover. When squeezing out, two flaps (valves) close in the front mouth area and the water is squeezed backwards through the gill cover gap; it should be noted that these two processes take place almost synchronously, so that water can constantly flow through the gill slits.

    In some bony fish (e.g. eels ) the gill cavity is only opened to the outside through a small gap, which means that the gills remain moist for a certain period of time, even when dry. Some more or less amphibious fish species have additional respiratory organs : Mudskippers can absorb oxygen from the skin by breathing . More common is the intestinal breathing , for example, when Loach , while air is swallowed, the gas exchange takes place on the walls instead of the highly perfused foregut. Some fish ( labyrinth fish ) breathe air through cranial chambers by using protuberances in the gill area to absorb oxygen. Fish which swim bladder is still connected to the intestine, such as the gars , also serving swim bladder for air breathing. In Flösselhechten and lungfish these intestinal evadinations have already evolved into lungs.

    The blood also serves the transportation of the endocrine glands formed hormones . Both the glands and the hormones are basically similar to those of humans.

    Intestines

    Internal anatomy of a bony fish:
    1 liver, 2 stomach, 3 intestines, 4 heart, 5 swim bladder, 6 kidney, 7 testes, 8 ureters, 9 vas deferens, 10 urinary bladder, 11 gills

    The intestine can be divided into: oral cavity (teeth extremely varied, rarely missing, e.g. in cypriniformes ), gill intestine (pharynx with crevices to the gills, usually with trap formation; at the end the pharyngeal bones with teeth that are very rarely missing), Esophagus, stomach (varied, absent e.g. in the Cypriniformes), midgut (very different in length, depending on diet; at the beginning porter blind tubes can extend from it, 0 to 1000 in number, function unclear; liver and Pancreas often not yet divorced), rectum (in herbivores sometimes with appendix; a spiral fold in it as in cartilaginous fish is still found in all non-Teleostei among the Osteichthyes, but is more and more regressed).

    Swim bladder

    Swimming bladder of a rudd

    In most species of bony fish, the function of the swim bladder is to regulate the specific weight of the fish (the density ) so that they can float in the water without exertion or only sink very slowly to the bottom. It emerged from a protuberance on the upper side of the foregut; it can, as with the carp , still be connected to it (physostom) or, as with the perch , form chambers separated from the intestine (physoclist).

    To regulate the specific weight, gases ( oxygen , carbon dioxide , nitrogen ) are released from the blood into the swim bladder via the so-called gas gland or the connecting duct to the intestine, the ductus pneumaticus . To reduce the volume of the swim bladder, the gas either reaches the foregut (and is spat out) via the ductus pneumaticus or back into the blood via a heavily perfused area in the swim bladder wall, the oval.

    Cartilaginous fish, but also bottom-dwelling or particularly well-swimming bony fish, lack the swim bladder - they sink to the bottom if they do not move. Through bone reduction, fat storage, etc. but the sunfish ( Mola ), for example, can also float on the sea surface without a swim bladder.

    Nervous system and sensory organs

    Fish brain
    Schematic longitudinal section through the eye of a bony fish

    The nervous system is simple, the brain is small, and there is no pronounced cerebral cortex. The fish's sense of smell is usually very pronounced, and the nasal and oral cavities are separated from each other. The organs of balance and hearing consist of closed, fluid-filled bladders. In some species (carp, tetras, catfish) they are connected to the swim bladder by moving bones and are known as Weber's apparatus . In structure they are similar to the inner ear of mammals, but a cochlea is not formed. Most fish species can perceive signals below 1  kHz , some have improved their hearing ability through special structures and extended the hearing range to 5 kHz or more.

    eye

    The structure and function of the bony fish's eye is largely a typical vertebrate eye and is therefore basically similar to the human eye . A significant difference to the eye of the land vertebrates is that the opening in the iris (Iris), the pupil is usually rigid, so that the amount of incident light into the eye can not be regulated. The biggest difference to the eye of terrestrial vertebrates, however, is the shape of the lens . This is spherical and the only partial organ of the eye that is responsible for the refraction of light rays, whereas in terrestrial vertebrates most of the light refraction (approx. 85%) occurs when passing through the cornea due to the clear difference in density between air and cornea. A consequence of the spherical shape, however, would be a strong spherical aberration and thus the creation of a blurred image on the retina . To reduce this error, the lens body has inhomogeneous optical properties: the refractive index decreases from the center to the edge of the lens. Such a lens is called a sliding index lens or a Matthiessen lens . In addition, the fisheye lens is not flexible and therefore has a fixed focal length . The accommodation , that is the conversion of near to far vision, is performed by contraction of the so-called Linsenretraktormuskels ( Musculus retractor lentis ), whereby the lens along the optical axis is moved closer to the retina.  Paragraph up to this point, unless otherwise quoted, after   and  The eye of the bony fish is therefore preset for close-up vision.

    For the eye of the cartilaginous fish it used to be that the accommodation runs exactly the other way around: the eye is preset to distant vision and the contraction of a protractor muscle ensures the conversion to near vision. However, this could not be confirmed by various studies on several shark species, so that it is now unclear how and whether the eyes of the cartilaginous fish accommodate at all. Many fish are capable of color and also perceive ultraviolet light.

    sense of touch

    The sense of touch is particularly well developed in bottom-dwelling fish. Especially on the lips and barbels there are sensory cells in the epidermis that transmit touch and, of course, taste stimuli. As a “sixth sense”, fish have organs for the perception of water currents, the lateral line organs , which extend laterally approximately in the middle over a large part of the body length. They consist of a series of skin protuberances in which there are sensory cells with sensory hairs that are excited by changes in flow. If this “side line” is missing (e.g. in herring ), there are also such organs on the head in (almost) all bony fish.

    Locomotion

    The main purpose of locomotion is the caudal fin - when the body moves sideways. Wrasse and surgeon fish , however, mainly move by means of the pectoral fins and use the tail fin only as a rudder (labriform). In the puffer fish relatives, the dorsal and anal fins, which are exactly opposite each other, are used for locomotion. The wave-like movement of the anal fin only serves to advance the New and Old World knife fish . You can also swim backwards skillfully by reversing the wave motion. In addition to the sense of balance, the light back reflex also serves to adjust the body orientation .

    Reproductive biology

    Maulbrütender threadfin cardinalfish ( Apogon leptacanthus )

    The kidneys are a paired (or unpaired fused) elongated organ below the spine. In all other "fish", the ureters (or tubes separated from them) also serve to drain the germ cells - but the real bony fish have developed their own egg and seminal ducts for this purpose. Your reproductive organs are to the side and above the intestine.

    In most species, fertilization takes place without copulation ; instead, the female roe (the eggs) and the male milk (the seeds) are released into the water at about the same time. (The reproductive female fish is called "Rogner", the reproductive male fish is called "Milchner".) The number of eggs fluctuates extremely: Sturgeons lay several million, the brood-caring sticklebacks no more than a hundred. In brood-caring species, it is often the males who take care of them.

    There are different forms of brood-caring species: open , cave and mouth brooders .

    • In open brooders, the eggs are laid by the female on plants or other solid materials and then inseminated by the male.
    • In cave breeders, the eggs are laid on the ceiling of caves. After hatching, the young are placed on leaves or in pits for a while (until the yolk sac is used up) and guarded. Then the larvae are led by the parents, with the parents either taking turns or one parent, usually the male, guarding the territorial boundaries. If a large part of the brood is lost, it can happen that the parents eat the few surviving larvae and quickly start a new brood again.
    • In the case of mouthbrooders, the females take the eggs into their mouths after they have been deposited, the male swims over the eggs and releases his semen in the process. The young of the mouthbrooders are already well developed when they leave the mouth, but after hatching they return to the mouth for a while if there is danger.

    The larvae of the bony fish develop into adults through metamorphosis . During metamorphosis, the larval organs are resorbed or rejected and the existing systems of the adult organs are developed to function.

    Examples of peculiarities in reproduction:

    • Cuckoo-whiskered catfish have specialized in laying their eggs next to the eggs of the host fish, which are then taken up by the female in the mouth and "hatched" there. The faster developing catfish larvae then eat the eggs or larvae of the host fish.
    • After hatching, discus fish secrete a skin secretion that is grazed by the larvae and serves as first food.
    • In labyrinth fish , the males often form a foam nest on the surface of the water between floating plants and then drive the female under the foam nest to lay eggs. The male wraps his body around the female and turns it on its back so that the eggs swim upwards. After insemination, the female is driven away and the brood is guarded by the male alone until the larvae hatch, after which the male does not take care of the brood either.
    • There are also (rarely) bony fish that have copulatory organs, such as the livebearers toothcarps (Poeciliinae) or the false thorn catfish (Auchenipteridae).
    • Some bony fish, especially marine perch relatives, do not have a genetically determined sex, it is only expressed through contact with partners or environmental conditions and can be changed in the course of life.

    Habitats

    Oceans

    The oceans make up about 70% of the earth's surface and are therefore the largest living space. The bony fish, which have chosen the upper layer of the open ocean down to a depth of 200 meters as a habitat, make up only a little more than one percent of all species, for the most part members of the mackerel and tuna (Scombridae) and the flying fish ( Exocoetidae). Another five percent of bony fish live pelagically below 200 meters. Most of them are herring-like (Clupeiformes) and lantern fish (Myctophidae).

    Most of the marine bony fish live near the coasts. The cold coasts are mainly inhabited by cod (Gadidae), eel-nut (Zoarcidae), slime fish (Blennioidei), armored cheeks (Scorpaeniformes, not related to a family) and Antarctic fish (Notothenioidei). A little more than five percent of the bony fish species prefer this habitat, which is also an important fishing area because the cold seas near the coast are very rich in nutrients and the fish occur in large schools. The warm coasts of the continents and tropical islands are the most biodiverse marine habitat because of the diverse ecological niches in coral reefs and mangroves, which make up more than 40% of the total bony fish fauna. Most of the species that occur here belong to the perch relatives (Percomorphaceae). B. to the perch-like (Perciformes), the slime fish-like (Blennioidei) and the goby-like (Gobiiformes). There are also a lot of eels (Anguilliformes) on warm coasts, e.g. B. the moray eels .

    The continental slopes and the near-ground zone of the deep sea , on and above which slightly more than six percent of all bony fish live, form special habitats . Grenadiers (Macrouridae), intestinal fish-like (Ophidiiformes), eel-nuts (Zoarcidae), many eels (Anguilliformes) and armored cheeks (Scorpaeniformes) have their habitat here.

    Inland waters

    Although the share of inland waters (fresh waters and salt lakes ) in the world's water resources is only 2.6 to 3%, they are home to around 40% of all bony fish species. Geographical isolation and differences in many conditions, such as subsoil, temperature, flow rate, seasons, oxygen levels, hardness, and pH , resulted in a wide variety of adaptations and species. Most of the bony fish living in fresh water with about 6000 species belong to the carp-like (Cypriniformes), the catfish-like (Siluriformes) and the tetra-like (Characiformes), which together with the species poorer orders of the New World knife fish (Gymnotiformes) and the sandfish-like (Gonorynchiformes; marine!) form the taxon of the Ostariophysi . From the range of these orders one can conclude that the Ostariophysi originated when all continents except Australia , where they do not occur, were still connected. The Ostariophysi split off from a common ancestor with the herring-like (Clupeiformes) and specialized in life in freshwater. They are primary freshwater fish. The most important freshwater fish after the Ostariophysi are the cichlids (Cichlidae), which dominate the freshwater fauna of the East African lakes and Central America, but also occur in West Africa, South America, Madagascar and South India.

    Phylogeny, systematics and fossil record

    External classification: origin of the bony fish

    Drawing of an acanthodier

    When exactly the first bony fish lived is unknown. One has to assume a history of this group from the early Silurian that was not documented by fossils . The oldest known relatively complete fossil of a bony fish comes from deposits of the Upper Silurian ( Ludlowium ) in today's China. This approximately 26 centimeter long, slim, streamlined representative, which bears the scientific name Guiyu oneiros , is assigned to the muscle fins (Sarcopterygii), but still has characteristics that are also to be found in other very original vertebrate groups ( cartilaginous fish , acanthodians , placoderms ) were lost in the further course of the evolution of the bony fish. Andreolepis hedei from northern Europe comes from slightly older strata than Guiyu and is therefore the oldest known bony fish. He seems to be even more original than Guiyu and apparently represents a stage of development before the separation of the muscle fins and ray fins lines. However, his remains are far more fragmentary and less complete than those of Guiyu . Both Guiyu and Andreolepis, as well as some other fragmentarily preserved Upper Silurian fossil finds of bony fish, come from marine deposits; In other words, according to current knowledge, the bony fish originated in the sea.

    Since the bony fish show similarities to the extinct acanthodes, it is assumed that these are the sister group of the bony fish. With them they have the gill structure of four pairs of gills with cleft gill and gill cover in common. Acanthodes and bony fish, in turn, have common ancestors with the cartilaginous fish. The oldest and most pristine jaw mouths are the placoderms, which share a common ancestor with the taxon from cartilaginous fish, acanthodes and bony fish. It is considered certain that the bony fish are not directly related to the cartilaginous fish. According to previous knowledge, the most likely family tree of the bony fish (Osteichthyes) looks like this:

     Jaw mouths 
     (Gnathostomata)  
     Eugnathostomata  
     Teleostomi  
     Osteichthyes 1  

    Meat fin (Sarcopterygii), including terrestrial vertebrates (Tetrapoda)


       

    Ray fins (Actinopterygii)



       

    Acanthodii



       

    Cartilaginous fish (Chondrichthyes)



       

    Placodermi



    1) The taxon that takes up this branch of the family tree is also called Euteleostomi or Osteognathostomata in order to avoid confusion with the paraphyletic taxon Osteichthyes of the classical system (see bony fish and terrestrial vertebrates ).

    Internal systematics: evolutionary history of the bony fish

    Eusthenopteron from Upper Devonian, a typical representative of the muscle fins line that leads to the tetrapods
    Palaeoniscus freilebeni , the " copper slate herring", a representative of the Palaeonisciformes and thus the first flower of the ray fins
    Bake fish ( Lepisosteus sp.), Survivors of the second wave of evolution of the ray fins

    Bony fish fossils can be found in freshwater as early as the early Devonian. Thus, the oldest as derived coelacanth identified (Coelacanthimorpha) fossils from freshwater sediments of Pragiums in China ( Euporosteus yunnanensis ) and Australia ( eoactinistia foreyi ). One of the earliest representatives of the line of development that leads to the modern lungfish and tetrapods, Youngolepis , was again found in China, in freshwater sediments of the Lochkovian . The oldest unequivocal representative of the ray fins Cheirolepis trailli comes from Central Devonian ( Emsium ) layers of the Old Red sandstone in Scotland. At that time, the representatives of the two main lines of bony fish, muscle fins and ray fins, already clearly differ in their body characteristics and have also lost many of the original characteristics that went back to their common ancestry with other basal vertebrate groups. Both the early ray fins and the early muscle fins are active predators, and the muscle fins are the dominant freshwater fish during the Mid-Devonian. The first land vertebrates (Tetrapoda) emerged from them in the course of the Upper Devonian , as evidenced by transitional forms such as Eusthenopteron , Panderichthys and Tiktaalik . In the course of the extinction event at the end of the Permian , the number of muscle fins species is greatly reduced. While the tetrapods flourished again in the Mesozoic Era, their fish-like musculoskeletal relatives remained comparatively poor in species and disappeared completely from the fossil record at the end of the Cretaceous . It was not until the 19th century that the six species of lungfish living today and in 1938 the Comoros coelacanth ( Latimeria chalumnae ) were discovered as " living fossils ".

    The Flösselhechte (Polypteriformes) are the most original taxon of the ray fins living today. The oldest fossils come from the early Upper Cretaceous, but the history of this group must go back at least as far as the Carboniferous , where the representatives of their sister group, the Guildayichthyids , which are known only from fossils, lived. Both the Guildayichthyiden and the Flösselike, which are collectively referred to as Cladistia , were apparently never particularly rich in species.

    In contrast, the cartilaginous organoids (Chondrostei) underwent rapid radiation in the carbon. A group of this first great wave of evolution known only through fossils are the Palaeonisciformes . Among them u. a. the most common fish in the copper shale of the Zechstein series , a Permian rock sequence in Central Europe, Palaeoniscus freilebeni . Representatives of the Redfieldiidae family are typical of the Triassic-Jurassic freshwater deposits of the Newark supergroup , a series of rocks on the eastern edge of North America that is associated with the formation of the Atlantic. The sturgeon-like (Acipenseriformes) are still living representatives of the cartilaginous organoids. They are known to be fossilized only from the Lower Jurassic, but recently the discovery of an early Permian sturgeon from China ( Eochondrosteus sinensis ) was reported.

    Already at the end of the Triassic the cartilaginous organoids experience a decline, possibly u. a. triggered by increasing competition from representatives of the basal neopteries (Neopterygii), which evolved rapidly during the Triassic. The North and Central American bonefish (Lepisosteidae) and the North American bald pike or mudfish ( Amia calva ), which together with their extinct relatives are referred to as bone organoids (Holostei), are survivors of this second evolutionary wave of the ray fins, which only ended in the late Cretaceous was. Two representatives from the Eocene , Cyclurus kehreri and Atractosteus strausi , which are closely related to the Holosteer species living today , are among the most common vertebrate fossils in the famous Messel Pit .

    The following cladogram shows the relationship of the basal taxa of the recent bony fish. The number of species living today is given in brackets.

    Dapedium from the Lias vom Holzmaden , a fossil representative of the second evolutionary wave of the ray fins
     Bony fish 
     (Osteichthyes) 
     Muscle Floss 
     (Sarcopterygii) 

    Quastenflosser 
     (Coelacanthimorpha) (2 types)


     Choanata 

    Land vertebrates (Tetrapoda, 
      approx. 30,000 species)


       

    Lungfish 
     (Dipnoi) (6 species)




     Ray fins 
     (Actinopterygii)  
     Actinopteri 
     Neuflosser  (Neopterygii) 

    Real bony fish (Teleostei) 
     (approx. 29,000 species)


       

    Bone organoids 
     (Holostei) (8 species)



       

    Sturgeon (Acipenseriformes) (30 species)



       

    Flössler (Polypteriformes) (18 species)




    Template: Klade / Maintenance / Style
    Leptolepides sprattiformis , a basal Teleoste from the Upper Jura, is one of the most common fish in Solnhofen limestone
    Halec microlepsis , a modern Teleosteer from the Upper Cretaceous Lebanon

    As early as the late Triassic around 220 million years ago, representatives of the real bony fish (Teleostei) appear in the fossil record for the first time . The Teleosteer stand out u. a. characterized by the fact that they no longer have any strongly mineralized scales, which is shown in the fossil record by the fact that only the inner skeleton of these representatives has been handed down as fossils. The basal teleosts also include Leedsichthys problematicus from the Jura , the largest bony fish in the history of the earth, which may even have reached a length of up to 25 meters. However, these basal lines all died out before the end of the Chalk . This could in turn be related to the fact that the Teleosteer in the Upper Jura and especially during the Cretaceous period experienced a further evolutionary surge, in the course of which almost all modern large groups emerged. These more modern representatives are likely to have increasingly competed with the early Teleosts. In the Upper Cretaceous and, as a result of the mass extinction at the Cretaceous-Tertiary border , especially in the Lower Tertiary , the last major phase of rapid evolutionary splitting took place among the Teleosts, which are essentially the perch relatives (Percomorphaceae) and the Ostariophysi (carp, Catfish and relatives) concerned. These two groups are the most species-rich today (perch relatives: approx. 14,000 species, Ostariophysi: approx. 6000 species, including two thirds of all freshwater fish species).

    Only 57% of all bony fish families living today are represented in the fossil record. Almost 70 families have died out.

    The success of the true bony fish is based in large part on adapting to life in the open water. In the course of the evolution of bony fish there is a tendency to reduce the number of vertebrae, which makes the fish body stiffer and more aerodynamic. Another cause may be the improved skull kinetics compared to more basal bony fish , associated with the development of a highly developed jaw apparatus . The latter allows predators to hunt more effectively using suckers , but also forms the basis for the development of very special forms of nutrition, such as B. grazing the growth of rocks and coral reefs . Today the Teleosteer represent the largest part of the fish species with over 40 orders and over 400 families .

    Some taxa of the real bony fish, such as the muzzle fish (Stomiiformes), the lizard fish relatives (Aulopiformes) and the lantern fish relatives (Myctophiformes) adapted to a life in the deep sea .

    Cladogram for the internal systematics of real bony fish (Teleostei) according to Betancur-R. et al. (2016):

     Real bony fish 

    Elopomorpha ( tarpon-like , bone-fish , thorn -back eel , eel-like )


       

    Bone-lick-like ( bone-lick-like , moon-eye )


     Clupeocephala  
     Otomorpha  

    Herring-like


       

    Alepocephaliformes


       

    Ostariophysi ( sandfish ,
    carp , New World knife fish , catfish , tetra )




     Euteleosteomorpha 

    Salamander fish


       

    Protacanthopterygii ( golden salmon-like , galaxies , salmon fish , pike-like )


       

    Stomiati ( smelt-like , Maulstachler )


       

    Neoteleostei  (see below)








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     Neoteleostei 

    Deep-sea tadpoles


     Eurypterygia 

    Lizardfish relatives


     Ctenosquamata 

    Lantern fishy


     Acanthomorphata 


    Paracanthopterygii ( Perch-like , Petersfisch-like , Stylephorus chordatus , Cod-like )


       

    Gloss fish-like



       

    Bearded fish


     Spinefish 

    Mucous head-like and Trachichthyiformes


       

    Soldier and hussar fish


       

    Perch relatives









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    The evolution and the phylogenetic relationship among the species, genera and families of the perch relatives (Percomorphaceae) are still largely unexplained. The abundance of almost 15,000 species makes detailed research difficult.

    An overview of all orders and families can be found under systematics of bony fish .

    Bony fish and terrestrial vertebrates

    African lungfish ( Protopterus annectens )

    From the point of view of the cladistic system , the terrestrial vertebrates (Tetrapoda) also belong to the bony fish i. w. S. Without it, the bony fish are a so-called paraphyletic taxon, which means that they do not include all descendants of the common ancestor of the bony fish - precisely because the terrestrial vertebrates belong to these descendants. For example, the lung fish or the coelacanth are more closely related to the terrestrial vertebrates than to other groups of fish. That is why the cladistic system rejects a taxon bony fish without terrestrial vertebrates. To avoid confusion with the paraphyletic Osteichthyes, some cladists introduced the new names Osteognathostomata (bone jaw mouths ), Neognathostomata (new jaw mouths ) and Euteleostomi for the taxon that also includes the terrestrial vertebrates.

    Remarks

    1. The use of the term “chondrostei” is not uniform in the specialist literature. On the one hand, as in this article, it stands for a larger group of original ray fins, on the other hand, it is only used for the sturgeon-like and their closest fossil relatives.
    2. The Holosteer have recently been viewed as paraphyletic. In more recent relationship analyzes, however, they again form a monophylum.

    literature

    • Robert L. Carroll: Paleontology and Evolution of the Vertebrates. Georg Thieme Verlag, Stuttgart 1993, ISBN 3-13-774401-6 .
    • Rudie H. Kuiter , Helmut Debelius : Atlas of the marine fish. Kosmos-Verlag, 2006, ISBN 3-440-09562-2 .
    • Kurt Fiedler: Textbook of Special Zoology, Volume II, Part 2: Fish. Gustav Fischer Verlag, Jena 1991, ISBN 3-334-00338-8 .
    • Karl Albert Frickhinger: Fossils Atlas Fishes. Mergus-Verlag, Melle 1999, ISBN 3-88244-018-X .
    • Hans-Eckard Gruner, Horst Füller, Kurt Günther: Urania animal kingdom, fish, amphibians, reptiles. (7 volumes), Urania-Verlag, 1991, ISBN 3-332-00376-3 .
    • Juraj Holcik: The Freshwater Fishes of Europe. Volume 1 / II, AULA-Verlag, Wiesbaden 1989, ISBN 3-89104-431-3 .
    • Oskar Kuhn: The prehistoric fish-like and fish. A. Ziemsen Verlag, Wittenberg 1967.
    • Joseph S. Nelson : Fishes of the World. John Wiley & Sons, 2006, ISBN 0-471-25031-7 .
    • Günther Sterba : The world's freshwater fish. 2nd Edition. Urania, Leipzig / Jena / Berlin 1990, ISBN 3-332-00109-4 .
    • Volker Storch, Ulrich Welsch: Kükenthal, zoological internship , Kapithel Osteichthyes; Spectrum Academic Publishing House; 26th edition; ISBN 978-3-8274-1998-9 .

    Web links

    Commons : Bonefish  - Collection of images, videos and audio files

    Individual evidence

    1. ^ A b Hans-Peter Schultze: Gnathostomata, Kiefermäuler. In Wilfried Westheide, Reinhard Rieger (Ed.): Special Zoology. Part 2: vertebrates or skulls. 2nd Edition. Spektrum Akademischer Verlag, Heidelberg 2010, ISBN 978-3-8274-2039-8 , pp. 211-215.
    2. a b Michael J. Benton : Paleontology of the vertebrates. Translation of the 3rd English edition (Translator: Hans-Ulrich Pfretzschner). Publishing house Dr. Friedrich Arrow. Munich 2007. 472 pages. ISBN 978-3-89937-072-0 .
    3. WDR.de: Sascha Ott: Chatty like a fish - under water, acoustic communication plays an astonishingly large role. P. 7, (PDF 110kB), Leonardo - Science and more. January 29, 2010.
    4. David J. Randall, Roger Eckert, Warren Burggren, Kathleen French: Tierphysiologie. 4th edition. Thieme, 2002, ISBN 3-13-664004-7 , p. 282.
    5. ^ State Institute for School Quality and Educational Research: Fundamentals of Biophysics . BRIGG Pädagogik Verlag, Munich 2009, ISBN 978-3-87101-640-0 , p. 13-14 .
    6. Jan-Peter Hildebrandt, Horst Beckmann, Uwe Homberg: Penzlin - textbook of animal physiology . 8th edition. Springer Spectrum, Berlin / Heidelberg 2015, ISBN 978-3-642-55369-1 , p. 673 .
    7. ^ Andreas Sebastian Reimann: Anatomical-macroscopic investigations of fish eyes. An interactive guide to the production and photography of ophthalmic preparations as a basis for fish ophthalmology. Dissertation. Veterinary Faculty of the Ludwig Maximilians University in Munich, 2015, urn : nbn: de: bvb: 19-180417 , pp. 3–18.
    8. Quentin Bone, Richard H. Moore: Biology of Fishes. 3. Edition. Taylor & Francis, 2008, ISBN 978-0-415-37562-7 , pp. 313 f.
    9. Volker Storch, Ulrich Welsch: Kükenthal - zoological internship. 26th edition. Spektrum Akademischer Verlag, Heidelberg 2009, ISBN 978-3-8274-1998-9 , p. 353 f.
    10. a b Zhu Min, Zhao Wenjin, Jia Liantao, Lu Jing, Qiao Tuo, Qu Qingming: The oldest articulated osteichthyan reveals mosaic gnathostome characters . In: Nature . tape 458 , 2009, p. 469-474 , doi : 10.1038 / nature07855 .
    11. ^ Hector Botella, Henning Blom, Markus Dorka, Per Erik Ahlberg, Philippe Janvier: Jaws and teeth of the earliest bony fishes . In: Nature . tape 448 , 2007, p. 583-586 , doi : 10.1038 / nature05989 .
    12. ^ Richard Lund: The new Actinopterygian order Guildayichthyiformes from the Lower Carboniferous of Montana (USA) In: Geodiversitas. Vol. 22, No. 2, 2000, pp. 171-206, sciencepress.mnhn.fr .
    13. ^ Lu Liwu, Li Daqing, Yang Liangfeng: Notes on the discovery of Permian Acipenseriformes in China . In: Chinese Science Bulletin . tape 50 , no. 12 , 2005, p. 1279-1280 , doi : 10.1007 / BF03183706 .
    14. Elizabeth C. Sibert, Richard D. Norris. New Age of Fishes initiated by the Cretaceous − Paleogene mass extinction. In: Proceedings of the National Academy of Sciences. Vol. 112, No. 28, 2015, pp. 8537-8542, doi: 10.1073 / pnas.1504985112 .
    15. ^ G. David Johnson, EO Wiley: Percomorpha. Tree of Life Web Project (January 9, 2007 release).
    16. R. Betancur-R., E. Wiley, N. Bailly, A. Acero, M. Miya, G. Lecointre, G. Ortí: Phylogenetic Classification of Bony Fishes - Version 4 (2016)