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Mastodonsauridae is a family of capitosauroid
temnospondyl Temnospondyli (from Greek τέμνειν, ''temnein'' 'to cut' and σπόνδυλος, ''spondylos'' 'vertebra') is a diverse order of small to giant tetrapods—often considered Labyrinthodontia, primitive amphibians—that flourished worldwid ...
s.
Fossil A fossil (from Classical Latin , ) is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserved ...
s belonging to this family have been found in
North America North America is a continent in the Northern Hemisphere and almost entirely within the Western Hemisphere. It is bordered to the north by the Arctic Ocean, to the east by the Atlantic Ocean, to the southeast by South America and the C ...
,
Greenland Greenland ( kl, Kalaallit Nunaat, ; da, Grønland, ) is an island country in North America that is part of the Kingdom of Denmark. It is located between the Arctic and Atlantic oceans, east of the Canadian Arctic Archipelago. Greenland is ...
,
Europe Europe is a large peninsula conventionally considered a continent in its own right because of its great physical size and the weight of its history and traditions. Europe is also considered a Continent#Subcontinents, subcontinent of Eurasia ...
,
Asia Asia (, ) is one of the world's most notable geographical regions, which is either considered a continent in its own right or a subcontinent of Eurasia, which shares the continental landmass of Afro-Eurasia with Africa. Asia covers an area ...
, and
Australia Australia, officially the Commonwealth of Australia, is a sovereign country comprising the mainland of the Australian continent, the island of Tasmania, and numerous smaller islands. With an area of , Australia is the largest country b ...
. The family Capitosauridae is
synonymous A synonym is a word, morpheme, or phrase that means exactly or nearly the same as another word, morpheme, or phrase in a given language. For example, in the English language, the words ''begin'', ''start'', ''commence'', and ''initiate'' are all ...
with Mastodonsauridae.


Description


Size

Mastodonsaurids were generally large amphibians, with some length estimates ranging from 3 to 6 meters. Some genera, like ''
Mastodonsaurus ''Mastodonsaurus'' (meaning "teat tooth lizard") is an extinct genus of temnospondyl amphibian from the Middle Triassic of Europe. It belongs to a Triassic group of temnospondyls called Capitosauria, characterized by their large body size and p ...
'' and ''
Cyclotosaurus ''Cyclotosaurus'' is an extinct genus of temnospondyl within the family Mastodonsauridae. It was of great size for an amphibian, had an elongated skull up to . Etymology The name means "round eared lizard" in Ancient Greek, derived from round ...
'' were specially large, reaching at least 4 meters or more. Other genera like '' Parotosuchus'' and '' Paracyclotosaurus'' only reached 2 meters or more in length, but it's still pretty large compared to modern-day amphibians.


Distinguishing features

Some mastodonsaurids including ''Parotosuchus'' were covered in a scaly skin, unlike the smooth skin of modern-day
amphibian Amphibians are four-limbed and ectothermic vertebrates of the class Amphibia. All living amphibians belong to the group Lissamphibia. They inhabit a wide variety of habitats, with most species living within terrestrial, fossorial, arbor ...
s, and probably moved with an eel-like motion in the water. Another peculiar mastodonsaurid is ''Sclerothorax'', with unusual features including its elongated neural spines. The neural spines are tallest at the front of the spine. ''Sclerothorax'' also has a very large
pectoral girdle The shoulder girdle or pectoral girdle is the set of bones in the appendicular skeleton which connects to the arm on each side. In humans it consists of the clavicle and scapula; in those species with three bones in the shoulder, it consists ...
; the
interclavicle An interclavicle is a bone which, in most tetrapods, is located between the clavicles. Therian mammals (marsupials and placentals) are the only tetrapods which never have an interclavicle, although some members of other groups also lack one. In t ...
bone is longer than the skull, meaning that ''Sclerothorax'' has the proportionally largest interclavicle of any temnospondyl. The body of ''Sclerothorax'' is covered by small scale-like bony ossicles that would have been embedded in the
dermis The dermis or corium is a layer of skin between the epidermis (with which it makes up the cutis) and subcutaneous tissues, that primarily consists of dense irregular connective tissue and cushions the body from stress and strain. It is divided ...
in life. The skull of ''Sclerothorax'' is short, wide, and roughly rectangular in shape. The snout is wider than the cheek region. The bones along the midline of the
skull roof The skull roof, or the roofing bones of the skull, are a set of bones covering the brain, eyes and nostrils in bony fishes and all land-living vertebrates. The bones are derived from dermal bone and are part of the dermatocranium. In compara ...
are also very wide relative to those of other temnospondyls.


Skull structures

There is a lot of known information about the skulls of mastodonsaurids, for example, '' Stanocephalosaurus'', one of the most well-known mastodonsaurids, has a lot of data about its skull. The skull of ''Stanocephalosaurus'' has key features characteristic to other temnospondyls, with the most apparent being the flattened shape of its
skull roof The skull roof, or the roofing bones of the skull, are a set of bones covering the brain, eyes and nostrils in bony fishes and all land-living vertebrates. The bones are derived from dermal bone and are part of the dermatocranium. In compara ...
. The dorsal as well as ventral surfaces of the skull of ''Stanocephalosaurus'' have well ornamented honeycombed patterns throughout. Palatal views of skulls in this genus show narrow interpalatal vacuities, as well as paired palatal
tusk Tusks are elongated, continuously growing front teeth that protrude well beyond the mouth of certain mammal species. They are most commonly canine teeth, as with pigs and walruses, or, in the case of elephants, elongated incisors. Tusks share c ...
s alternately functioning on the
vomer The vomer (; lat, vomer, lit=ploughshare) is one of the unpaired facial bones of the skull. It is located in the midsagittal line, and articulates with the sphenoid, the ethmoid, the left and right palatine bones, and the left and right max ...
and
palatine A palatine or palatinus (in Latin; plural ''palatini''; cf. derivative spellings below) is a high-level official attached to imperial or royal courts in Europe since Roman times.
, with the vomerine tusks directed backwards. This animal shows an increased expansion of the pterygoids and palatines, as well as narrow posterior
nostril A nostril (or naris , plural ''nares'' ) is either of the two orifices of the nose. They enable the entry and exit of air and other gasses through the nasal cavities. In birds and mammals, they contain branched bones or cartilages called tur ...
s. The vomer also bears two vomerine rows: a transversal denticle row and a longitudinal denticle row that runs along the inner margin of the choana. The palatines and ectopterygoids are broader than in other temnospondyls, which is somewhat common within mastodonsaurids. ''Stanocephalosaurus'' also has a long, triangular skull which is much longer than broad, and has an elongated preorbital region continuously narrowing anteriorly to an obtuse rounded snout. This type of skull is very similar to that of ''Mastodonsaurus'', which is also triangular in shape. The ''Stanocephalosaurus'' has at least 100 straight and conical teeth on 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 b ...
and
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. The ...
combined, gradually increasing in size towards the front. While there is a lot of known information about ''Stanocephalosaurus'', some of the skull regions which were previously inaccessible or too poorly preserved on ''Stanocephalosaurus'' have been observed with X-ray
micro-CT X-ray microtomography, like tomography and X-ray computed tomography, uses X-rays to create cross-sections of a physical object that can be used to recreate a virtual model ( 3D model) without destroying the original object. The prefix ''micro- ...
scans, including the otic capsule, delta groove of the exoccipital, as well as parts of the arterial and nervous system. Air pockets around the
stapes The ''stapes'' or stirrup is a bone in the middle ear of humans and other animals which is involved in the conduction of sound vibrations to the inner ear. This bone is connected to the oval window by its annular ligament, which allows the foo ...
of ''Stanocephalosaurus'' have been hypothesized to act as resonance chambers, meaning that the spoon-shaped inner ear bone could be related to underwater hearing. This trait is possibly associated with early
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 ther ...
evolution, which probably also appears on other members of this family and can act as a link to
anuran A frog is any member of a diverse and largely carnivorous group of short-bodied, tailless amphibians composing the order Anura (ανοὐρά, literally ''without tail'' in Ancient Greek). The oldest fossil "proto-frog" ''Triadobatrachus'' is ...
tympanum evolution. The narrow head and elongated snout of ''Stanocephalosaurus'' suggests that stress levels during biting are slightly higher than temnospondyls with a wider and shorter skull. Its skull also has an elongated preorbital region compared to other mastodonsaurids. The
vertebra The spinal column, a defining synapomorphy shared by nearly all vertebrates,Hagfish are believed to have secondarily lost their spinal column is a moderately flexible series of vertebrae (singular vertebra), each constituting a characteristic ...
e of ''Stanocephalosaurus'' are rhachitomous, with a neural arch and a bipartite centrum that is divided into a large, unpaired wedge-shaped intercentrum and smaller paired pleurocentra. In anterior and posterior views, the intercentrum is a dorsally half-ring, surrounding the persistent
notochord In anatomy, the notochord is a flexible rod which is similar in structure to the stiffer cartilage. If a species has a notochord at any stage of its life cycle (along with 4 other features), it is, by definition, a chordate. The notochord consi ...
from ventral and lateral sides. Lateral and ventral surfaces of the intercentrum are smooth, suggesting a continuation of
cartilage Cartilage is a resilient and smooth type of connective tissue. In tetrapods, it covers and protects the ends of long bones at the joints as articular cartilage, and is a structural component of many body parts including the rib cage, the neck a ...
due to the unfinished medial surface. The posterodorsal margin of the intercentrum also shows a parapophysis for articulation with the capitulum of the ribs. Another well-known mastodonsaurid in terms of fossil knowledge is ''Mastodonsaurus'', notably from numerous skull specimens found in Germany. The large, oval eye sockets are midway along the skull. The jaws are lined with conical teeth, two large
tusk Tusks are elongated, continuously growing front teeth that protrude well beyond the mouth of certain mammal species. They are most commonly canine teeth, as with pigs and walruses, or, in the case of elephants, elongated incisors. Tusks share c ...
s project up from the end of the lower jaw, fitting through openings on the
palate The palate () is the roof of the mouth in humans and other mammals. It separates the oral cavity from the nasal cavity. A similar structure is found in crocodilians, but in most other tetrapods, the oral and nasal cavities are not truly sep ...
and emerging out from the top of the skull when the jaw is closed. Based on recent research, the body of ''Mastodonsaurus'' was more crocodile-like in shape than in earlier reconstructions, with a longer trunk section and a longer tail. The greatly reduced limb bones have joints that are poorly developed. ''Paracyclotosaurus'' is yet again, another well-known mastodonsaurid, and just like most genera of the family, it had various species. This mastodonsaurid has at least three species, including its
type species In zoological nomenclature, a type species (''species typica'') is the species name with which the name of a genus or subgenus is considered to be permanently taxonomically associated, i.e., the species that contains the biological type specimen( ...
, which is ''P. davidi'', along with two other: ''P. crookshanki'' and ''P. morganorum''. The completeness of the skeleton of this animal led to an accurate description. The small distortion of the head can be overlooked, and the complete preservation of the skull and jaws makes its shape certain at a length of 60 cm. The right corner of the head is essentially undistorted and shows that the skull at its point of greatest depth is almost exactly as high as the lower jaw. The lower jaw also has its symphysis the same height as the skull above it. Therefore, at its point of greatest height, the head was essentially of oval section, 22 cm high and 44 cm wide. The nostril, orbit and tympanic membrane are all directed upward, but whether the eye could be projected upward above the head to give a horizontal view similar to a
frog A frog is any member of a diverse and largely carnivorous group of short-bodied, tailless amphibians composing the order Anura (ανοὐρά, literally ''without tail'' in Ancient Greek). The oldest fossil "proto-frog" ''Triadobatrachus'' is ...
, is difficult to determine. With very similar features to ''Cyclotosaurus'', '' Subcyclotosaurus'' is another good example of mastodonsaurid. Its skull is characterized by the small tabular without any trace of a "horn", but with a round lappet that approaches the squamosal flange lateral to the tympanic membrane, failing to meet it by about its own width. The occiput between the otic notches is proportionately wide, a reflection of the small size of the skull. The skull is otherwise of normal mastodonsaurid structure, being specifically more similar to ''Parotosuchus'' than to any other genera of this family. The skull also has a small internasal vacuity between the dorsal processes of the premaxillae and lateral lines are often shown as continuous grooves with well-defined borders. A deep groove on the
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. The ...
begins immediately behind and lateral to the nostril and passes straight back to the lachrymal, on which bone it turns outward and forward and ends abruptly. Another groove appears to begin on the maxilla, immediately lateral to that described above. It passes back just above the insertion of the teeth for the full length of the bone. The supraorbital groove begins abruptly on the dorsal surface of the premaxilla, immediately passes on to the nasal, and extends back on that bone close to its suture with the lachrymal. It then comes on to the prefrontal, passing on to the frontal where that bone enters the orbital border. Then as a well-defined groove it surrounds the hinder part of the orbit, turns vertically on to the jugal, and then backward to cross the point where jugal, quadratojugal and squamosal meet, continuing over the squamosal to pass back on to the body. '' Xenotosuchus'' is a member of this which is not very well known, and only a few fragments have been discovered from this animal. This genus was originally a species of ''Parotosuchus''. Like many mastodontosaurids, it was a large animal with a large head. Its
amphibian Amphibians are four-limbed and ectothermic vertebrates of the class Amphibia. All living amphibians belong to the group Lissamphibia. They inhabit a wide variety of habitats, with most species living within terrestrial, fossorial, arbor ...
life history meant that the distinct shape of the
skull roof The skull roof, or the roofing bones of the skull, are a set of bones covering the brain, eyes and nostrils in bony fishes and all land-living vertebrates. The bones are derived from dermal bone and are part of the dermatocranium. In compara ...
would change from a generalized
tadpole A tadpole is the larval stage in the biological life cycle of an amphibian. Most tadpoles are fully aquatic, though some species of amphibians have tadpoles that are terrestrial. Tadpoles have some fish-like features that may not be found i ...
-like skull to the distinct adult shape. The head bones are covered in large pits and grooves, indicating extensive dermal armour on the head. Both the upper and lower jaw had
tusk Tusks are elongated, continuously growing front teeth that protrude well beyond the mouth of certain mammal species. They are most commonly canine teeth, as with pigs and walruses, or, in the case of elephants, elongated incisors. Tusks share c ...
s, those of the upper jaw being situated on a second row of teeth on the
vomer The vomer (; lat, vomer, lit=ploughshare) is one of the unpaired facial bones of the skull. It is located in the midsagittal line, and articulates with the sphenoid, the ethmoid, the left and right palatine bones, and the left and right max ...
and
palatine bone In anatomy, the palatine bones () are two irregular bones of the facial skeleton in many animal species, located above the uvula in the throat. Together with the maxillae, they comprise the hard palate. (''Palate'' is derived from the Latin ...
. Contrary to related forms like ''Mastodonsaurus'' and ''Stanocephalosaurus'', the tusks of the lower jaw were of moderate size and did not penetrate 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 b ...
. Like the mentioned ''Xenotosuchus'', the genus ''
Wetlugasaurus ''Wetlugasaurus'' (meaning " Vetluga River lizard") is an extinct genus of temnospondyl amphibian from the Early Triassic (Olenekian) Charkabozh, Kzylsaiskaya, Petropavlovka, Kamennyi Yar and Vetluga Series Formations of northern Russia ...
'' is also one of the smaller and less known mastodonsaurids. It had a
skull The skull is a bone protective cavity for the brain. The skull is composed of four types of bone i.e., cranial bones, facial bones, ear ossicles and hyoid bone. However two parts are more prominent: the cranium and the mandible. In humans, t ...
that measured around , and reached a total length of at least , and although it is pretty large compared to modern-day amphibians, ''Wetlugasaurus'' is still very small compared to most of the other mastodonsaurids.


Paleobiology

Mastodonsaurids were
piscivorous A piscivore () is a carnivorous animal that eats primarily fish. The name ''piscivore'' is derived . Piscivore is equivalent to the Greek-derived word ichthyophage, both of which mean "fish eater". Fish were the diet of early tetrapod evoluti ...
, mainly preying on small and mid-sized fish, although they also ate land-living animals, such as small
archosaurs Archosauria () is a clade of diapsids, with birds and crocodilians as the only living representatives. Archosaurs are broadly classified as reptiles, in the cladistic sense of the term which includes birds. Extinct archosaurs include non-avian ...
. Some evidences, including the fossils of some smaller temnospondyls bear tooth marks made by mastodonsaurid-like animals. It is very likely that mastodonsaurids caught their food like the living Giant Salamander, waiting until the prey came near, then quickly opening their enormous mouths and swallowing them prey whole. The most probable food would be small fishes, up to some 15 cm. long, which swam in shoals.


Paleoecology

Mastodonsaurids were aquatic animals that rarely left water and may have been completely unable to leave the water, as large quantities of bones have been found that suggests that many species have died en masse when pools dried up during times of
drought A drought is defined as drier than normal conditions.Douville, H., K. Raghavan, J. Renwick, R.P. Allan, P.A. Arias, M. Barlow, R. Cerezo-Mota, A. Cherchi, T.Y. Gan, J. Gergis, D.  Jiang, A.  Khan, W.  Pokam Mba, D.  Rosenfeld, J. Tierney, an ...
. They mostly inhabited swampy pools and fed mainly on fish, whose remains have been found in fossilized
coprolite A coprolite (also known as a coprolith) is fossilized feces. Coprolites are classified as trace fossils as opposed to body fossils, as they give evidence for the animal's behaviour (in this case, diet) rather than morphology. The name is de ...
s.Benes, Josef. Prehistoric Animals and Plants. Prague, Artia, 1979.


See also

*
List of prehistoric amphibians This list of prehistoric amphibians is an attempt to create a comprehensive listing of all genera from the fossil record that have ever been considered to be amphibians, excluding purely vernacular terms. The list includes all commonly accepted ge ...


References


Sources

* * {{Taxonbar, from=Q255408 Capitosaurs Triassic temnospondyls Triassic first appearances Triassic extinctions