Gnathostomata

Jawed vertebrates Temporal range: Early Silurian–Holocene, 439–0 Ma[1] PreꞒ Ꞓ O S D C P T J K Pg N (Possible Late Ordovician record, 444 Ma)[1]
Example of jawed vertebrates: Dunkleosteus (Placodermi) , a lemon shark (Chondrichthyes), a red-bellied piranha (Actinopterygii) and a Nile crocodile (Tetrapoda).
Scientific classification
Domain:	Eukaryota
Kingdom:	Animalia
Phylum:	Chordata
Clade:	Olfactores
Subphylum:	Vertebrata
Infraphylum:	Gnathostomata Gegenbauer, 1874
Subgroups
†"Placodermi" Eugnathostomata Total group Chondrichthyes †"Acanthodii" Chondrichthyes (cartilaginous fishes) Osteichthyes (bony fish, including tetrapods)

Gnathostomata (/ˌnæθoʊˈstɒmətə/; from Ancient Greek: γνάθος (gnathos) 'jaw' + στόμα (stoma) 'mouth') are jawed vertebrates. Gnathostome diversity comprises roughly 60,000 species, which accounts for 99% of all extant vertebrates, including all living bony fishes (both ray-finned and lobe-finned, including their terrestrial tetrapod relatives) and cartilaginous fishes, as well as extinct prehistoric fish such as placoderms and acanthodians. Most gnathostomes have retained ancestral traits like true teeth, a stomach,^[2] and paired appendages (pectoral and pelvic fins, limbs, wings, etc.).^[3] Other traits are elastin,^[4] horizontal semicircular canal of the inner ear, myelinated neurons, and an adaptive immune system which has discrete secondary lymphoid organs (spleen and thymus)^[5] and uses V(D)J recombination to create antigen recognition sites, rather than using genetic recombination in the variable lymphocyte receptor gene.^[6]

It is now assumed that Gnathostomata evolved from ancestors that already possessed two pairs of paired fins.^[7] Until recently these ancestors, known as antiarchs, were thought to have lacked pectoral or pelvic fins.^[7] In addition to this, some placoderms were shown to have a third pair of paired appendages, that had been modified to claspers in males and pelvic basal plates in females — a pattern not seen in any other vertebrate group.^[8] The jawless Osteostraci are generally considered the closest sister taxon of Gnathostomata.^[3][9][10]

Jaw development in vertebrates is likely a product of bending the first pair of gill arches. This development would help suck water into the mouth by the movement of the jaw, so that it would then pass over the gills via buccal pumping for gas exchange. The repetitive use of the newly formed jaw bones would eventually lead to the ability to bite in some gnathostomes.^[11]

Newer research suggests that a branch of placoderms was most likely the ancestor of present-day gnathostomes. A 419-million-year-old fossil of a placoderm named Entelognathus had a bony oral skeleton and anatomical details associated with cartilaginous and bony fish, demonstrating that the absence of a bony skeleton in cartilaginous fish is a derived trait.^[12] The fossil findings of primitive bony fishes such as Guiyu oneiros and Psarolepis, which lived contemporaneously with Entelognathus and had pelvic girdles more in common with placoderms than with other bony fish, show that it was a relative rather than a direct ancestor of the extant gnathostomes.^[13] It also indicates that spiny sharks and Chondrichthyes represent a single sister group to the bony fishes.^[12] Fossil findings of juvenile placoderms, which had true teeth that grew on the surface of the jawbone and had no roots, making them impossible to replace or regrow as they broke or wore down as they grew older, proves the common ancestor of all gnathostomes had teeth and place the origin of teeth along with, or soon after, the evolution of jaws.^[14][15]

Late Ordovician-aged microfossils of what have been identified as scales of either acanthodians^[16] or "spiny sharks",^[17] may mark Gnathostomata's first appearance in the fossil record. Undeniably unambiguous gnathostome fossils, mostly of primitive acanthodians, begin appearing by the early Silurian, and become abundant by the start of the Devonian.