''Entelognathus primordialis'' (“primordial complete jaw”) is a maxillate placoderm from the late

Silurian The Silurian ( ) is a geologic period and system spanning 24.6 million years from the end of the Ordovician Period, at million years ago ( Mya), to the beginning of the Devonian Period, Mya. The Silurian is the shortest period of the Paleoz ...

(

Ludlow epoch In the geological timescale, the Ludlow Epoch (from 427.4 ± 0.5 million years ago to 423.0 ± 2.3 million years ago) occurred during the Silurian Period, after the end of the Homerian Age. It is named for the town of Ludlow in Shropshire, Englan ...

) of

Qujing Qujing () is a prefecture-level city in the east of Yunnan province, China, bordering Guizhou province to the east and the Guangxi Zhuang Autonomous Region to the southeast; thus, it was called "Key between Yunnan and Guizhou" () and "Throat of Y ...

Yunnan Yunnan , () is a landlocked province in the southwest of the People's Republic of China. The province spans approximately and has a population of 48.3 million (as of 2018). The capital of the province is Kunming. The province borders the ...

, 419 million years ago. A team led by Min Zhu of the Academy of Sciences'

Institute of Vertebrate Paleontology and Paleoanthropology The Institute of Vertebrate Paleontology and Paleoanthropology (IVPP; ) of China is a research institution and collections repository for fossils, including many dinosaur and pterosaur specimens (many from the Yixian Formation). As its name sugg ...

in Beijing discovered the intact, articulated fossil in rock formations at

Xiaoxiang Xiaoxiang (), also transliterated ''XiaoXiang'', ''Hsiao Hsiang'', and ''Chiu Chiang'', in some older sources, refers to the "lakes and rivers" region in south-central China south of the middle-reaches of the Yangtze River and corresponding, more ...

reservoir.

Specimen and taxonomy

The holotype of ''E. primordialis'' is the uncrushed and mostly intact anterior half of an individual with the articulating head and trunk armor preserved in three dimensions. The holotype is about long, and the live animal is estimated to have been over long. In overall form, the animal resembles primitive

arthrodire Arthrodira (Greek for "jointed neck") is an order of extinct armored, jawed fishes of the class Placodermi that flourished in the Devonian period before their sudden extinction, surviving for about 50 million years and penetrating most marine eco ...

s, but the anatomy of the jaws strongly suggests the anatomies of

bony fish Osteichthyes (), popularly referred to as the bony fish, is a diverse superclass of fish that have skeletons primarily composed of bone tissue. They can be contrasted with the Chondrichthyes, which have skeletons primarily composed of cartil ...

and

tetrapod Tetrapods (; ) are four-limbed vertebrate animals constituting the superclass Tetrapoda (). It includes extant and extinct amphibians, sauropsids ( reptiles, including dinosaurs and therefore birds) and synapsids ( pelycosaurs, extinct t ...

s. Specifically, this is the first stem gnathostome with dermal marginal jaw bones. These bones are the

premaxilla The premaxilla (or praemaxilla) is one of a pair of small cranial bones at the very tip of the upper jaw of many animals, usually, but not always, bearing teeth. In humans, they are fused with the maxilla. The "premaxilla" of therian mammal has ...

maxilla The maxilla (plural: ''maxillae'' ) in vertebrates is the upper fixed (not fixed in Neopterygii) bone of the jaw formed from the fusion of two maxillary bones. In humans, the upper jaw includes the hard palate in the front of the mouth. T ...

, and

dentary In anatomy, the mandible, lower jaw or jawbone is the largest, strongest and lowest bone in the human facial skeleton. It forms the lower jaw and holds the lower teeth in place. The mandible sits beneath the maxilla. It is the only movable bone ...

. Most known placoderms had simple beak-like jaws made of bone plates. The researchers' cladistic diagram suggests that ''E. primordialis'' forms a

polytomy An internal node of a phylogenetic tree is described as a polytomy or multifurcation if (i) it is in a rooted tree and is linked to three or more child subtrees or (ii) it is in an unrooted tree and is attached to four or more branches. A tr ...

with arthrodires,

ptyctodont The ptyctodontids ("folded-teeth") are placoderms of the order Ptyctodontida, containing the family Ptyctodontidae. With their big heads, big eyes, reduced armor and long bodies, the ptyctodontids bore a superficial resemblance to modern day chi ...

s, and all advanced

gnathostome Gnathostomata (; from Greek: (') "jaw" + (') "mouth") are the jawed vertebrates. Gnathostome diversity comprises roughly 60,000 species, which accounts for 99% of all living vertebrates, including humans. In addition to opposing jaws, living ...

s (namely bony fish, tetrapods,

acanthodians Acanthodii or acanthodians is an extinct class of gnathostomes (jawed fishes), typically considered a paraphyletic group. They are currently considered to represent a grade of various fish lineages leading up to the extant Chondrichthyes, which ...

, and

chondrichthyes Chondrichthyes (; ) is a class that contains the cartilaginous fishes that have skeletons primarily composed of cartilage. They can be contrasted with the Osteichthyes or ''bony fishes'', which have skeletons primarily composed of bone tissue. ...

Etymology

The generic name translates as "complete jaw", referring to how the animal had a complete set of dermal marginal jaw bones. The specific name translates as "primordial".

Evolutionary significance

Prior to the discovery of ''Entelognathus'', scientists assumed that the last common ancestor of jawed vertebrates was a shark-like animal, with no distinct jawbones, and that modern jaws evolved in early bony fishes. This discovery shows that modern jaws evolved earlier. It is possible that Chondrichthyes started with distinct jaws and then dispensed with them. This has been called the earliest known animal with what looks like a face.

References

External links

National Geographic - Fish fossil has oldest known face, may influence evolution

Reuters - Fossil fish find in China fills in evolutionary picture

Discover Magazine - First jawed fish had one ugly face

Scientist hails 'jaw-dropping' fish fossil discovery
(BBC, 2013-09-27) Placoderm genera Placoderms of Asia Silurian fish of Asia Placodermi enigmatic taxa Prehistoric animals of China Transitional fossils Fossil taxa described in 2013 2013 in China {{Placoderm-stub

Specimen and taxonomy

Etymology

Evolutionary significance

See also

References

External links