A significant evolutionary milestone during the Silurian Period (about 443 to 419 million years ago) was the appearance of jawed and bony fish around 420 B.C.T.
During the Silurian Period, the first bony fish, the Osteichthyes, appeared, represented by the Acanthodians covered with bony scales. The Acanthodians reached considerable diversity and developed movable jaws, adapted from the supports of the front two or three gill arches.
A class of spiny sharks, appeared by the late Silurian, about 420 million years ago, and became extinct before the end of the Permian, about 250 million years ago. However, scales and teeth attributed to this group, as well as more derived gnathostomes such as Chondrichthyes and Osteichthyes, date from the Ordovician (~460 million years ago). Acanthodians were generally small shark-like fishes varying from toothless filter-feeders to toothed predators. They were once often classified as an order of the class Placodermi, another group of primitive fishes, but recent authorities tend to place the acanthodians nearer to or within the living gnathostomes. They are distinguished in two respects: they were the earliest known jawed vertebrates, and they had stout spines supporting their fins, fixed in place and non-movable (like a shark's dorsal fin).
Gnathostomata is the group of vertebrates with jaws. The term derives from Greek gnathos ("jaw") + stoma ("mouth"). Gnathostome diversity comprises roughly 60,000 species, which accounts for 99% of all living vertebrates. In addition to opposing jaws, living gnathostomes also have teeth, paired appendages, and a horizontal semi-circular canal of the inner ear, along with physiological and cellular anatomical characters such as the myelin sheathes of neurons. Another is an adaptive immune system that uses V(D)J recombination to create antigen recognition sites, rather than using genetic recombination in the variable lymphocyte receptor gene.
The group is traditionally a superclass, broken into three top-level groupings: Chondrichthyes, or the cartilaginous fish; Placodermi, an extinct clade of armored fish; and Teleostomi, which includes the familiar classes of bony fish, birds, mammals, reptiles, and amphibians. Some classification systems have used the term Amphirhina. It is a sister group of the jawless craniates Agnatha.
New fossil finds suggests thelodonts as the closest relatives of the Gnathostomata.
It is believed that the jaws evolved from anterior gill support arches that had acquired a new role, being modified to pump water over the gills by opening and closing the mouth more effectively — the buccal pump mechanism. The mouth could then grow bigger and wider, making it possible to capture larger prey. This close and open mechanism would with time become stronger and tougher, being transformed into real jaws.
Placoderms used sharp bony plates as teeth instead, and newer research indicates the jaws in placoderms evolved independently of those in the other Gnathostomata.
The Gnathostomata first appeared in the Ordovician period and became common in the Devonian period.
Osteichthyes, also called bony fish, are a taxonomic group of fish that have bone, as opposed to cartilaginous, skeletons. The vast majority of fish are osteichthyes, which is an extremely diverse and abundant group consisting of over 30,000 species. It is the largest class of vertebrates in existence today. Osteichthyes is divided into the ray-finned fish (Actinopterygii) and lobe-finned fish (Sarcopterygii). The oldest known fossils of bony fish are about 420 million years ago, which are also transitional fossils, showing a tooth pattern that is in between the tooth rows of sharks and bony fishes.
In most classification systems the Osteichthyes are paraphyletic with land vertebrates. That means that the nearest common ancestor of all Osteichthyes includes tetrapods amongst its descendants. Actinopterygii (ray-finned fish) are monophyletic, but the inclusion of Sarcopterygii in Osteichthyes causes Osteichthyes to be paraphyletic. Paradoxically, Sarcopterygii is considered monophyletic, as it includes tetrapods.
Most bony fish belong to the ray-finned fish (Actinopterygii); there are only eight living species of non-tetrapod lobe-finned fish (Sarcopterygii), including the lungfish and coelacanths.
Traditionally, the bony fish had been treated as a class within the vertebrates, with Actinopterygii and Sarcopterygii as subclasses. However, some recent works have elevated Osteichthyes to superclass, with Actinopterygii and Sarcopterygii as classes.
Chondrichthyes, from the Greek chondr- ("cartilage") + ichthys ("fish"), or cartilaginous fishes are jawed fish with paired fins, paired nares, scales, a heart with its chambers in series, and skeletons made of cartilage rather than bone. The class is divided into two subclasses: Elasmobranchii (sharks, rays and skates) and Holocephali (chimaeras, sometimes called ghost sharks, which are sometimes separated into their own class).
Within the infraphylum Gnathostomata, cartilaginous fishes are distinct from all other jawed vertebrates, the extant members of which all fall into Teleostomi.
Teleostomi is a clade -- a group consisting of an ancestor and all its descendants, a single "branch" on the "tree of life" -- of jawed vertebrates that 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 placodermi.
The clade Teleostomi should not be confused with the similar-sounding fish clade Teleostei.
The origins of the teleostomes are obscure, but their first known fossils are Acanthodians ("spiny sharks") from the Late Ordovician Period. Living teleostomes constitute the clade Euteleostomi, which includes all osteichthyans and tetrapods. Even after the acanthodians perished at the end of the Permian Period (252 million years ago), their euteleostome relatives flourished such that today they comprise 99% of living vertebrate species.
Teleostomes have two major adaptations that relate to aquatic respiration. First, the early teleostomes probably had some type of operculum, however, it was not the one-piece affair of living fish. The development of a single respiratory opening seems to have been an important step. The second adaptation, the teleostomes also developed a primitive lung with the ability to use some atmospheric oxygen. This developed, in later species, into the lung and (later) the swim bladder, used to keep the fish at neutral buoyancy.
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