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''Megamastax'' (meaning "big mouth") is a
genus Genus (; : genera ) is a taxonomic rank above species and below family (taxonomy), family as used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In bino ...
of
lobe-finned fish Sarcopterygii (; )—sometimes considered synonymous with Crossopterygii ()—is a clade (traditionally a class or subclass) of vertebrate animals which includes a group of bony fish commonly referred to as lobe-finned fish. These vertebrates ar ...
which lived during the late
Silurian The Silurian ( ) is a geologic period and system spanning 23.5 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 third and shortest period of t ...
period Period may refer to: Common uses * Period (punctuation) * Era, a length or span of time *Menstruation, commonly referred to as a "period" Arts, entertainment, and media * Period (music), a concept in musical composition * Periodic sentence (o ...
, about 423
million years ago Million years ago, abbreviated as Mya, Myr (megayear) or Ma (megaannum), is a unit of time equal to (i.e. years), or approximately 31.6 teraseconds. Usage Myr is in common use in fields such as Earth science and cosmology. Myr is also used w ...
, in
China China, officially the People's Republic of China (PRC), is a country in East Asia. With population of China, a population exceeding 1.4 billion, it is the list of countries by population (United Nations), second-most populous country after ...
. Before the discovery of ''Megamastax'', it was thought that jawed vertebrates (
gnathostomes Gnathostomata (; from Ancient Greek: (') 'jaw' + (') '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 a ...
) were limited in size and variation before the
Devonian The Devonian ( ) is a period (geology), geologic period and system (stratigraphy), system of the Paleozoic era (geology), era during the Phanerozoic eon (geology), eon, spanning 60.3 million years from the end of the preceding Silurian per ...
period. ''Megamastax'' is known only from jaw bones and it is estimated that it reached about long.


Discovery and naming

Fossils of ''Megamastax'' were found in a layer of the Kuanti Formation in
Qujing Qujing ( zh, c= , p=Qūjìng) is a prefecture-level city in the east of Yunnan province, China, bordering Guizhou province to the north and east and the Guangxi Zhuang Autonomous Region to the south; thus, it was called "Key between Yunnan and Gui ...
,
Yunnan Yunnan; is an inland Provinces of China, province in Southwestern China. The province spans approximately and has a population of 47.2 million (as of 2020). The capital of the province is Kunming. The province borders the Chinese provinces ...
,
China China, officially the People's Republic of China (PRC), is a country in East Asia. With population of China, a population exceeding 1.4 billion, it is the list of countries by population (United Nations), second-most populous country after ...
. The
holotype A holotype (Latin: ''holotypus'') is a single physical example (or illustration) of an organism used when the species (or lower-ranked taxon) was formally described. It is either the single such physical example (or illustration) or one of s ...
of ''Megamastax amblyodus'' is IVPP V18499.1, a complete left
mandible In jawed vertebrates, the mandible (from the Latin ''mandibula'', 'for chewing'), lower jaw, or jawbone is a bone that makes up the lowerand typically more mobilecomponent of the mouth (the upper jaw being known as the maxilla). The jawbone i ...
. Two additional specimens, IVPP V18499.2 (a partial left mandible) and IVPP V18499.3 (a right
maxilla In vertebrates, the maxilla (: maxillae ) 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. The two maxil ...
) have been referred to the species. The generic name of ''Megamastax'' is Greek for "big mouth", derived from ''megalos'' (big) and ''mastax'' (mouth). The species name, ''amblyodus'', translates to "blunt tooth".


Description

Although ''Megamastax'' can be safely considered a
sarcopterygian Sarcopterygii (; )—sometimes considered synonymous with Crossopterygii ()—is a clade (traditionally a class or subclass) of vertebrate animals which includes a group of bony fish commonly referred to as lobe-finned fish. These vertebrates ar ...
(lobe-finned fish) due to having
cosmine Cosmine is a spongy, bony material that makes up the dentine-like layers in the scales of the lobe-finned fishes of the class Sarcopterygii. Fish scales that include layers of cosmine are known as cosmoid scales. Description As traditionally d ...
on its jaws, coronoid plates, a prearticular bone, and a biconcave
glenoid The glenoid fossa of the scapula or the glenoid cavity is a bone part of the shoulder. The word ''glenoid'' is pronounced or (both are common) and is from , "socket", reflecting the shoulder joint's ball-and-socket form. It is a shallow, pyrif ...
, it is unique among early jawed fish for its jaw and teeth structure. Although most early osteichthyans have only one row of sharp marginal teeth along the edge of the jaw, ''Megamastax'' has two rows of small marginal teeth. In addition, ''Megamastax'' uniquely has a row of large, blunt teeth fused to four coronoid bones on the inside edge of each mandible. Similar genera such as '' Psarolepis'' and '' Guiyu'' have sharp fangs on their five coronoids while poroplepiformes and
tetrapodomorphs Tetrapodomorpha (also known as Choanata) is a clade of vertebrates consisting of tetrapods (four-limbed vertebrates) and their closest sarcopterygian relatives that are more closely related to living tetrapods than to living lungfish. Advanced for ...
have tusk-like teeth and three coronoids. Using the body plans of
Devonian The Devonian ( ) is a period (geology), geologic period and system (stratigraphy), system of the Paleozoic era (geology), era during the Phanerozoic eon (geology), eon, spanning 60.3 million years from the end of the preceding Silurian per ...
osteichthyans as a guide, it can be estimated that the mandibles of ''Megamastax'' occupy between a fifth and a seventh of the total body length. If this is true, then IVPP V18499.1 had a length of and IVPP V18499.2 had a length of . Given these large estimates, ''Megamastax'' was the largest known Silurian vertebrate.


Paleobiology

With rounded coronoid teeth, ''Megamastax'' could have fed on hard-shelled animals. Extant fish with crushing teeth, such as
wolf eel The wolf eel (''Anarrhichthys ocellatus'') is a species of marine ray-finned fish belonging to the family ''Anarhichadidae'', the wolf fishes. It is found in the North Pacific Ocean. Despite its common name and resemblance, it is not a true eel ...
s and horn sharks, generally have such teeth at the front of the jaw, separate from the coronoids. Although ''Megamastax'' differs from these fish by having its teeth and coronoids run the entire length of the jaw, this could simply be a result of having a different method of prey capture. While low body sizes for Silurian vertebrates has traditionally been seen as a result of the Silurian having low levels of atmospheric oxygen (as large vertebrates are susceptible to hypoxia), recent analyses have shown that terrestrial plant life was more well established in the Silurian than once thought, providing a previously unknown source of oxygen. In addition, new Paleozoic atmospheric models indicate that oxygen levels were actually around modern levels by the end of the Silurian period. Although it is not certain that ''Megamastax's'' large size is a result of this atmospheric trend, the presence of such a large fish at this period does weaken the argument that a lack of large Silurian vertebrates can be used as evidence of low oxygen levels.


Paleoecology

Various hard-shelled animals were present in the late Silurian of China, including brachipods,
molluscs Mollusca is a phylum of protostome, protostomic invertebrate animals, whose members are known as molluscs or mollusks (). Around 76,000 extant taxon, extant species of molluscs are recognized, making it the second-largest animal phylum ...
,
trilobite Trilobites (; meaning "three-lobed entities") are extinction, extinct marine arthropods that form the class (biology), class Trilobita. One of the earliest groups of arthropods to appear in the fossil record, trilobites were among the most succ ...
s, and even armored fish like
placoderms Placoderms (from Ancient Greek πλάξ 'plax'', ''plakos plate' and δέρμα 'dermaskin') are vertebrate animals of the class Placodermi, an extinct group of prehistoric fish known from Paleozoic fossils during the Silurian and the Dev ...
and galeaspids. Due to its large size and predatory lifestyle, ''Megamastax'' could be considered the first vertebrate
apex predator An apex predator, also known as a top predator or superpredator, is a predator at the top of a food chain, without natural predators of its own. Apex predators are usually defined in terms of trophic dynamics, meaning that they occupy the hig ...
.


References

{{Taxonbar, from=Q18434336 Prehistoric lobe-finned fish genera Silurian fish of Asia Enigmatic fish taxa