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Until finds of other early tetrapods and closely related dating in the late 20th century, Ichthyostega stood alone as the transitional fossil fish fish and tetrapods, combining a fishlike tail and gills fish an amphibian skull and limbs.
The fins evolved into the legs of the first dating land vertebrates, amphibians. They are distinguished in two respects: The fish would be bred female and sterile, though a very small percentage might still be able to breed. They included the structure of its middle ear , and its fins show the precursors of the forearm bones, the radius and ulna. The first animals to venture onto dry land were arthropods.
The evolution of fish began about million years ago during the Cambrian dailycoupons.pro was during this time that the early chordates developed the skull and the vertebral column, leading to the first craniates and vertebrates. The United States Fish and Wildlife Service is the principal Federal agency dedicated to fish and wildlife conservation. The Service's history spans nearly years, dating from the establishment of its predecessor agency, the Bureau of Fisheries, in GM fish set for supermarket shelves after U.S.
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This has led to the Devonian being known as the age of fishes. It was from the lobe-finned fish that the tetrapods evolved, the four-limbed vertebrates, represented today by amphibians , mammals , reptiles and birds. Transitional tetrapods first appeared during the early Devonian, and by the late Devonian the first tetrapods appeared.
The diversity of jawed vertebrates may indicate the evolutionary advantage of a jawed mouth ; but it is unclear if the advantage of a hinged jaw is greater biting force, improved respiration, or a combination of factors. Fish do not represent a monophyletic group, but a paraphyletic one, as they exclude the tetrapods. Fish, like many other organisms, have been greatly affected by extinction events throughout natural history.
The Ordovician—Silurian extinction events led to the loss of many species. The late Devonian extinction led to the extinction of the ostracoderms and placoderms by the end of the Devonian, as well as other fish. The spiny sharks became extinct at the Permian—Triassic extinction event ; the conodonts became extinct at the Triassic—Jurassic extinction event. The Cretaceous—Paleogene extinction event , and the present day Holocene extinction , have also affected fish variety and fish stocks.
Conventional classification has living vertebrates as a subphylum grouped into eight classes based on traditional interpretations of gross anatomical and physiological traits. In turn, these classes are grouped into the vertebrates that have four limbs the tetrapods and those that do not: The extant vertebrate classes are: Fish may have evolved from an animal similar to a coral-like sea squirt a tunicate , whose larvae resemble early fish in important ways.
The first ancestors of fish may have kept the larval form into adulthood as some sea squirts do today , although this path cannot be proven. Vertebrates , among them the first fishes , originated about million years ago during the Cambrian explosion , which saw the rise in organism diversity. The first ancestors of fish, or animals that were probably closely related to fish, were Pikaia , Haikouichthys and Myllokunmingia.
Pikaia had a primitive notochord , a structure that could have developed into a vertebral column later. Unlike the other fauna that dominated the Cambrian, these groups had the basic vertebrate body plan: These were followed by indisputable fossil vertebrates in the form of heavily armoured fishes discovered in rocks from the Ordovician Period — Ma.
The first jawed vertebrates appeared in the late Ordovician and became common in the Devonian , often known as the "Age of Fishes". The Devonian also saw the rise of the first labyrinthodonts , which was a transitional between fishes and amphibians.
The colonisation of new niches resulted in diversification of body plans and sometimes an increase in size. The Devonian Period to Ma brought in such giants as the placoderm Dunkleosteus , which could grow up to seven meters long, and early air-breathing fish that could remain on land for extended periods. Among this latter group were ancestral amphibians. The reptiles appeared from labyrinthodonts in the subsequent Carboniferous period. The anapsid and synapsid reptiles were common during the late Paleozoic , while the diapsids became dominant during the Mesozoic.
In the sea, the bony fishes became dominant. The later radiations, such as those of fish in the Silurian and Devonian periods, involved fewer taxa, mainly with very similar body plans. The first animals to venture onto dry land were arthropods.
Some fish had lungs and strong, bony fins and could crawl onto the land also. Jawless fishes belong to the superclass Agnatha in the phylum Chordata , subphylum Vertebrata. Agnatha comes from the Greek , and means "no jaws". Although a minor element of modern marine fauna , jawless fish were prominent among the early fish in the early Paleozoic. Two types of Early Cambrian animal apparently having fins, vertebrate musculature, and gills are known from the early Cambrian Maotianshan shales of China: They have been tentatively assigned to Agnatha by Janvier.
A third possible agnathid from the same region is Haikouella. A possible agnathid that has not been formally described was reported by Simonetti from the Middle Cambrian Burgess Shale of British Columbia.
Many Ordovician, Silurian, and Devonian agnathians were armoured with heavy bony-spiky plates. The first armoured agnathans—the Ostracoderms , precursors to the bony fish and hence to the tetrapods including humans —are known from the middle Ordovician , and by the Late Silurian the agnathans had reached the high point of their evolution.
Most of the ostracoderms, such as thelodonts , osteostracans , and galeaspids , were more closely related to the gnathostomes than to the surviving agnathans, known as cyclostomes. Cyclostomes apparently split from other agnathans before the evolution of dentine and bone, which are present in many fossil agnathans, including conodonts. The agnathans as a whole are paraphyletic ,  because most extinct agnathans belong to the stem group of gnathostomes.
Phylogenetic groups are given definitions based on their relationship to one another, rather than purely on physical traits such as the presence of a backbone. This nesting pattern is often combined with traditional taxonomy, in a practice known as evolutionary taxonomy. The cladogram below for jawless fish is based on studies compiled by Philippe Janvier and others for the Tree of Life Web Project.
Conodonts resembled primitive jawless eels. They appeared Ma and were wiped out Ma. These "teeth" have been variously interpreted as filter-feeding apparatuses or as a "grasping and crushing array". The preserved musculature hints that some conodonts Promissum at least were efficient cruisers but incapable of bursts of speed. Ostracoderms shell-skinned are armoured jawless fishes of the Paleozoic.
The term does not often appear in classifications today because it is paraphyletic or polyphyletic , and has no phylogenetic meaning. The eyes were particularly shielded.
Earlier chordates used their gills for both respiration and feeding, whereas ostracoderms used their gills for respiration only. They had up to eight separate pharyngeal gill pouches along the side of the head, which were permanently open with no protective operculum.
Unlike invertebrates that use ciliated motion to move food, ostracoderms used their muscular pharynx to create a suction that pulled small and slow moving prey into their mouths.
The first fossil fishes that were discovered were ostracoderms. The Swiss anatomist Louis Agassiz received some fossils of bony armored fish from Scotland in the s. He had a hard time classifying them as they did not resemble any living creature. He compared them at first with extant armored fish such as catfish and sturgeons but later realizing that they had no movable jaws, classified them in into a new group "ostracoderms".
Ostracoderms existed in two major groups, the more primitive heterostracans and the cephalaspids. Later, about million years ago, the jawed fish evolved from one of the ostracoderms. After the appearance of jawed fish, most ostracoderm species underwent a decline, and the last ostracoderms became extinct at the end of the Devonian period. The vertebrate jaw probably originally evolved in the Silurian period and appeared in the Placoderm fish , which further diversified in the Devonian.
The two most anterior pharyngeal arches are thought to have become the jaw itself and the hyoid arch, respectively. The hyoid system suspends the jaw from the braincase of the skull, permitting great mobility of the jaws.
Already long assumed to be a paraphyletic assemblage leading to more derived gnathostomes, the discovery of Entelognathus suggests that placoderms are directly ancestral to modern bony fish. As in most vertebrates , fish jaws are bony or cartilaginous and oppose vertically, comprising an upper jaw and a lower jaw. The jaw is derived from the most anterior two pharyngeal arches supporting the gills, and usually bears numerous teeth.
The skull of the last common ancestor of today's jawed vertebrates is assumed to have resembled sharks. It is thought that the original selective advantage garnered by the jaw was not related to feeding, but to increased respiration efficiency. The jaws were used in the buccal pump observable in modern fish and amphibians that pumps water across the gills of fish or air into the lungs in the case of amphibians.
Over evolutionary time the more familiar use of jaws to humans , in feeding, was selected for and became a very important function in vertebrates. Many teleost fish have substantially modified their jaws for suction feeding and jaw protrusion , resulting in highly complex jaws with dozens of bones involved.
Jawed vertebrates and jawed fish evolved from earlier jawless fish, and the cladogram below for jawed vertebrates is a continuation of the cladogram in the section above. Acanthodians and Chondrichthyes cartilaginous fishes. Placoderms , class Placodermi plate skinned , are extinct armoured prehistoric fish, which appeared about Ma in the Early to Middle Silurian.
They were mostly wiped out during the Late Devonian Extinction event, Ma, though some survived and made a slight recovery in diversity during the Famennian epoch before dying out entirely at the close of the Devonian, mya; they are ultimately ancestal to modern vertebrates.
Their head and thorax were covered with massive and often ornamented armoured plates. The rest of the body was scaled or naked, depending on the species. The armour shield was articulated, with the head armour hinged to the thoratic armour.
This allowed placoderms to lift their heads, unlike ostracoderms. Placoderms were the first jawed fish; their jaws likely evolved from the first of their gill arches. The chart on the right shows the rise and demise of the separate placoderm lineages: Spiny sharks , class Acanthodii, are extinct fishes that share features with both bony and cartilaginous fishes, though ultimately more closely related to and ancestral to the latter. Despite being called "spiny sharks", acanthodians predate sharks, though they gave rise to them.
They evolved in the sea at the beginning of the Silurian Period, some 50 million years before the first sharks appeared. Eventually competition from bony fishes proved too much, and the spiny sharks died out in Permian times about Ma.
In form they resembled sharks, but their epidermis was covered with tiny rhomboid platelets like the scales of holosteans gars , bowfins. Cartilaginous fishes, class Chondrichthyes , consisting of sharks , rays and chimaeras , appeared by about million years ago, in the middle Devonian , evolving from acanthodians. The class contains the sub classes Holocephali chimaera and Elasmobranchii sharks and rays. The radiation of elasmobranches in the chart on the right is divided into the taxa: Bony fishes , class Osteichthyes, are characterised by bony skeleton rather than cartilage.
They appeared in the late Silurian , about million years ago. The recent discovery of Entelognathus strongly suggests that bony fishes and possibly cartilaginous fishes, via acanthodians evolved from early placoderms. The bony and cartilaginous fish groups that emerged after the Devonian, were characterised by steady improvements in foraging and locomotion. Lobe-finned fishes , fish belonging to the class Sarcopterygii, are mostly extinct bony fishes, basally characterised by robust and stubby lobe fins containing a robust internal skeleton, cosmoid scales and internal nostrils.
Their fins are fleshy, lobed , paired fins, joined to the body by a single bone. The pectoral and pelvic fins are articulated in ways resembling the tetrapod limbs they were the precursors to. The fins evolved into the 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 ray-finned fish. The braincase of lobe-finned fishes 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. Lobe-finned fishes, such as coelacanths and lungfish , were the most diverse group of bony fishes in the Devonian. Taxonomists who subscribe to the cladistic approach include the grouping Tetrapoda within the Sarcopterygii, and the tetrapods in turn include all species of four-limbed vertebrates. The lobe-finned fish apparently followed two different lines of development and are accordingly separated into two subclasses, the Rhipidistia including the lungfish, and the Tetrapodomorpha , which include the Tetrapoda and the Actinistia coelacanths.
The first lobe-finned fishes, found in the uppermost Silurian ca Ma , closely resembled spiny sharks , which became extinct at the end of the Paleozoic. In the early—middle Devonian - Ma , while the predatory placoderms dominated the seas, some lobe-finned fishes came into freshwater habitats. In the Early Devonian Ma , the lobe-finned fishes split into two main lineages — the coelacanths and the rhipidistians.
The former never left the oceans and their heyday was the Late Devonian and Carboniferous , from to Ma, as they were more common during those periods than in any other period in the Phanerozoic; coelacanths still live today in the oceans genus Latimeria. The Rhipidistians, whose ancestors probably lived in estuaries , migrated into freshwater habitats. They in turn split into two major groups: The lungfish's greatest diversity was in the Triassic period; today there are fewer than a dozen genera left.
The lungfish evolved the first proto-lungs and proto-limbs; developing the ability to live outside a water environment in the middle Devonian Ma. The first tetrapodomorphs, which included the gigantic rhizodonts, had the same general anatomy as the lungfish, who were their closest kin, but they appear not to have left their water habitat until the late Devonian epoch - Ma , with the appearance of tetrapods four-legged vertebrates.
Tetrapods are the only tetrapodomorphs that survived after the Devonian. Lobe-finned fishes continued until towards the end of Paleozoic era, suffering heavy losses during the Permian-Triassic extinction event Ma. Ray-finned fishes , class Actinopterygii, differ from lobe-finned fishes in that their fins consist of webs of skin supported by spines "rays" made of bone or horn.
There are other differences in respiratory and circulatory structures. Ray-finned fishes normally have skeletons made from true bone, though this is not true of sturgeons and paddlefishes. Ray-finned fishes are the dominant vertebrate group, containing half of all known vertebrate species.
They inhabit abyssal depths in the sea, coastal inlets and freshwater rivers and lakes, and are a major source of food for humans. Pikaia , along with Myllokunmingia and Haikouichthys ercaicunensis immediately below, are all candidates in the fossil record for the titles of "first vertebrate" and "first fish". Pikaia is a genus that appeared about Ma during the Cambrian explosion of multicellular life.
Pikaia gracilens pictured is a transitional fossil between invertebrates and vertebrates,  and may be the earliest known chordate. Pikaia was a sideways-flattened, leaf-shaped animal that swam by throwing its body into a series of S-shaped, zig-zag curves, similar to movement of snakes. Fish inherited the same swimming movement, but they generally have stiffer backbones. It had a pair of large head tentacles and a series of short appendages, which may be linked to gill slits, on either side of its head.
Pikaia shows the essential prerequisites for vertebrates. The flattened body is divided into pairs of segmented muscle blocks , seen as faint vertical lines. The muscles lie on either side of a flexible structure resembling a rod that runs from the tip of the head to the tip of the tail.
By the start of the Early Devonian mya, jawed fishes had divided into four distinct clades: The modern bony fishes, class Osteichthyes , appeared in the late Silurian or early Devonian, about million years ago. Both the cartilaginous and bony fishes may have arisen from either the placoderms or the spiny sharks. A subclass of bony fishes, the ray-finned fishes Actinopterygii , have become the dominant group in the post-Paleozoic and modern world, with some 30, living species.
Sea levels in the Devonian were generally high. Marine faunas were dominated by bryozoa , diverse and abundant brachiopods , the enigmatic hederelloids , microconchids and corals. Lily-like crinoids were abundant, and trilobites were still fairly common.
Among vertebrates, jawless armoured fish ostracoderms declined in diversity, while the jawed fish gnathostomes simultaneously increased in both the sea and fresh water. Armoured placoderms were numerous during the lower stages of the Devonian Period but became extinct in the Late Devonian, perhaps because of competition for food against the other fish species.
Early cartilaginous Chondrichthyes and bony fishes Osteichthyes also become diverse and played a large role within the Devonian seas. The first abundant genus of shark, Cladoselache , appeared in the oceans during the Devonian Period. The great diversity of fish around at the time have led to the Devonian being given the name "The Age of Fish" in popular culture. The first ray-finned and lobe-finned bony fish appeared in the Devonian, while the placoderms began dominating almost every known aquatic environment.
However, another subclass of Osteichthyes, the Sarcopterygii , including lobe-finned fishes including coelacanths and lungfish and tetrapods , was the most diverse group of bony fishes in the Devonian.
Sarcopterygians are basally characterized by internal nostrils, lobe fins containing a robust internal skeleton, and cosmoid scales. During the Middle Devonian — Ma, the armoured jawless ostracoderm fishes were declining in diversity; the jawed fish were thriving and increasing in diversity in both the oceans and freshwater.
The shallow, warm, oxygen-depleted waters of Devonian inland lakes, surrounded by primitive plants, provided the environment necessary for certain early fish to develop essential characteristics such as well developed lungs and the ability to crawl out of the water and onto the land for short periods of time. Cartilaginous fishes, class Chondrichthyes , consisting of sharks , rays and chimaeras , appeared by about million years ago, in the middle Devonian. During the Late Devonian the first forests were taking shape on land.
The first tetrapods appear in the fossil record over a period, the beginning and end of which are marked with extinction events. This lasted until the end of the Devonian mya. The ancestors of all tetrapods began adapting to walking on land, their strong pectoral and pelvic fins gradually evolved into legs see Tiktaalik.
The first ammonite mollusks appeared. Trilobites , the mollusk-like brachiopods and the great coral reefs , were still common.
The Late Devonian extinction occurred at the beginning of the last phase of the Devonian period, the Famennian faunal stage, the Frasnian-Famennian boundary , about Many fossil agnathan fishes, save for the psammosteid heterostracans , make their last appearance shortly before this event. The Late Devonian extinction crisis primarily affected the marine community, and selectively affected shallow warm-water organisms rather than cool-water organisms.
The most important group affected by this extinction event were the reef-builders of the great Devonian reef-systems. A second extinction pulse, the Hangenberg event closed the Devonian period and had a dramatic impact on vertebrate faunas. Placoderms mostly became extinct during this event, as did most members of other groups including lobe-finned fishes, acanthodians and early tetrapods in both marine and terrestrial habitats, leaving only a handful of survivors. This event has been related to glaciation in the temperate and polar zones as well as euxinia and anoxia in the seas.
Bothriolepis pitted scale was the most successful genus of antiarch placoderms , if not the most successful genus of any placoderm, with over species spread across Middle to Late Devonian strata across every continent.
Dunkleosteus is a genus of arthrodire placoderms that existed from to Ma. Apart from its contemporary Titanichthys below , no other placoderm rivalled it in size. Instead of teeth, Dunkleosteus had two pairs of sharp bony plates, which formed a beak-like structure. Apart from megalodon , it had the most powerful bite of any fish,  generating bite forces in the same league as Tyrannosaurus rex and the modern crocodile. Materpiscis mother fish is a genus of ptyctodontid placoderm from about Ma.
Known from only one specimen, it is unique in having an unborn embryo present inside, and with remarkable preservation of a mineralised placental feeding structure umbilical cord. This makes Materpiscis the first known vertebrate to show viviparity , or giving birth to live young. The first tetrapods are four-legged, air-breathing, terrestrial animals from which the land vertebrates descended, including humans. They evolved from lobe-finned fish of the clade Sarcopterygii , appearing in coastal water in the middle Devonian, and giving rise to the first amphibians.
The group of lobe-finned fishes that were the ancestors of the tetrapod are grouped together as the Rhipidistia ,  and the first tetrapods evolved from these fish over the relatively short timespan — Ma.
The early tetrapod groups themselves are grouped as Labyrinthodontia. They retained aquatic, fry-like tadpoles , a system still seen in modern amphibians. From the s to the early s it was thought that tetrapods evolved from fish that had already acquired the ability to crawl on land, possibly so they could go from a pool that was drying out to one that was deeper. However, in , nearly complete fossils of Acanthostega from about Ma showed that this Late Devonian transitional animal had legs and both lungs and gills, but could never have survived on land: The current hypothesis is that Acanthostega , which was about 1 metre 3.
Its skeleton differed from that of most fish, in ways that enabled it to raise its head to breathe air while its body remained submerged, including: The Devonian proliferation of land plants may help to explain why air-breathing would have been an advantage: There are three major hypotheses as to how tetrapods evolved their stubby fins proto-limbs. The traditional explanation is the "shrinking waterhole hypothesis" or "desert hypothesis" posited by the American paleontologist Alfred Romer.
He believed limbs and lungs may have evolved from the necessity of having to find new bodies of water as old waterholes dried up. They argued that sarcopterygians may have first emerged unto land from intertidal zones rather than inland bodies of water.
The third hypothesis, the "woodland hypothesis", was proposed by the American paleontologist Gregory J. He argues that limbs may have developed in shallow bodies of water in woodlands as a means of navigating in environments filled with roots and vegetation. He based his conclusions on the evidence that transitional tetrapod fossils are consistently found in habitats that were formerly humid and wooded floodplains.
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