Metriorhynchidae is an

extinct Extinction is the termination of a kind of organism or of a group of kinds (taxon), usually a species. The moment of extinction is generally considered to be the death of the last individual of the species, although the capacity to breed and ...

family Family (from la, familia) is a Social group, group of people related either by consanguinity (by recognized birth) or Affinity (law), affinity (by marriage or other relationship). The purpose of the family is to maintain the well-being of its ...

of specialized, aquatic

metriorhynchoid Metriorhynchoidea is an extinct superfamily of thalattosuchian crocodyliforms from the Early Jurassic to the Early Cretaceous (Toarcian - Valanginian, possibly as late as early Aptian) of Europe, North America and South America. Metriorhynch ...

crocodyliform Crocodyliformes is a clade of crurotarsan archosaurs, the group often traditionally referred to as "crocodilians". They are the first members of Crocodylomorpha to possess many of the features that define later relatives. They are the only pseudo ...

s from the

Middle Jurassic The Middle Jurassic is the second epoch of the Jurassic Period. It lasted from about 174.1 to 163.5 million years ago. Fossils of land-dwelling animals, such as dinosaurs, from the Middle Jurassic are relatively rare, but geological formations co ...

to the

Early Cretaceous The Early Cretaceous ( geochronological name) or the Lower Cretaceous (chronostratigraphic name), is the earlier or lower of the two major divisions of the Cretaceous. It is usually considered to stretch from 145 Ma to 100.5 Ma. Geology Pro ...

period (

Bajocian In the geologic timescale, the Bajocian is an age and stage in the Middle Jurassic. It lasted from approximately 170.3 Ma to around 168.3 Ma (million years ago). The Bajocian Age succeeds the Aalenian Age and precedes the Bathonian Age. Stratig ...

to early

Aptian The Aptian is an age in the geologic timescale or a stage in the stratigraphic column. It is a subdivision of the Early or Lower Cretaceous Epoch or Series and encompasses the time from 121.4 ± 1.0 Ma to 113.0 ± 1.0 Ma (million years ago), a ...

) of

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 ...

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 Car ...

and

South America South America is a continent entirely in the Western Hemisphere and mostly in the Southern Hemisphere, with a relatively small portion in the Northern Hemisphere at the northern tip of the continent. It can also be described as the southe ...

. The name Metriorhynchidae was coined by the

Austrian Austrian may refer to: * Austrians, someone from Austria or of Austrian descent ** Someone who is considered an Austrian citizen, see Austrian nationality law * Austrian German dialect * Something associated with the country Austria, for example: ...

zoologist Zoology ()The pronunciation of zoology as is usually regarded as nonstandard, though it is not uncommon. is the branch of biology that studies the Animal, animal kingdom, including the anatomy, structure, embryology, evolution, Biological clas ...

Leopold Fitzinger Leopold Joseph Franz Johann Fitzinger (13 April 1802 – 20 September 1884) was an Austrian zoologist. Fitzinger was born in Vienna and studied botany at the University of Vienna under Nikolaus Joseph von Jacquin. He worked at the Vienna Naturhis ...

in 1843.Fitzinger LJFJ. 1843. ''Systema Reptilium''. Wien: Braumüller et Seidel, 106 pp. The group contains two subfamilies, the

Metriorhynchinae Metriorhynchinae is a subfamily of metriorhynchid crocodyliforms from the late Middle Jurassic to the Early Cretaceous (Callovian - Valanginian) of Europe, North America and South America. Named by Fitzinger, in 1843, it contains the metrior ...

and the

Geosaurinae Geosaurinae is a subfamily of metriorhynchid crocodyliforms from the Middle Jurassic to the Early Cretaceous (Bathonian - Aptian) of Europe, North America and South America. Named by Richard Lydekker, in 1889, it contains the metriorhynchids '' S ...

. They represent the most marine adapted of all archosaurs.

Description

Metriorhynchids are fully aquatic

s. Their forelimbs were small and paddle-like, and unlike living

crocodylian Crocodilia (or Crocodylia, both ) is an order of mostly large, predatory, semiaquatic reptiles, known as crocodilians. They first appeared 95 million years ago in the Late Cretaceous period (Cenomanian stage) and are the closest living re ...

s, they lost their

osteoderms Osteoderms are bony deposits forming scales, plates, or other structures based in the dermis. Osteoderms are found in many groups of extant and extinct reptiles and amphibians, including lizards, crocodilians, frogs, temnospondyls (extinct ...

("armour scutes"). Their body shape maximised hydrodynamy (swimming efficiency), as they did have a shark-like tail fluke. Like ichthyosaurs and plesiosaurs, metriorhynchids developed smooth, scaleless skin. Metriorhynchids were the only group of

archosaur 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 d ...

s to become fully adapted to the marine realm, becoming

pelagic The pelagic zone consists of the water column of the open ocean, and can be further divided into regions by depth (as illustrated on the right). The word ''pelagic'' is derived . The pelagic zone can be thought of as an imaginary cylinder or wa ...

in lifestyle.Steel R. 1973. ''Crocodylia. Handbuch der Paläoherpetologie, Teil 16''. Stuttgart: Gustav Fischer Verlag,116 pp. With tail flukes, reduced limb musculature, and long bones

histologically Histology, also known as microscopic anatomy or microanatomy, is the branch of biology Biology is the scientific study of life. It is a natural science with a broad scope but has several unifying themes that tie it together as a single ...

comparable to other obligately aquatic animals, they were almost certainly incapable of terrestrial locomotion; combined with an unusually tall hip opening, as also seen in other obligately aquatic reptiles including the

viviparous Among animals, viviparity is development of the embryo inside the body of the parent. This is opposed to oviparity which is a reproductive mode in which females lay developing eggs that complete their development and hatch externally from the m ...

Keichousaurus ''Keichousaurus'' (key-cho-saurus) is a genus of marine reptile in the pachypleurosaur family which went extinct at the close of the Triassic in the Triassic-Jurassic extinction event. The name derives from Kweichow (now Guizhou Province) in Ch ...

'', these characters suggest that metriorhynchids gave live birth. A fossil of a pregnant ''Dakosaurus'' female recovered from the Late Jurassic

Plattenkalk Plattenkalk is a very finely grained limestone chemically precipitated in a stratified water column under conditions where bioturbation does not occur. The reasons for the quiet depositional environment and the processes of sediment accumulation v ...

, Bavaria, preserves the complete skeleton of a neonate with small, paddle-like forelimbs unsuited for walking on land, similar to those of adults, further supporting live birth in metriorhynchids. Recent research posits that despite their successful adaptation to a pelagic lifestyle, basal metriorhynchids were uniquely disadvantaged among aquatic tetrapods in evolving into sustained swimmers due to little to no posterodorsal retraction of the external nares (unlike other reptilian groups such as

mesosaur Mesosaurs ("middle lizards") were a group of small aquatic reptiles that lived during the early Permian period, roughly 299 to 270 million years ago. Mesosaurs were the first known aquatic reptiles, having apparently returned to an aquatic life ...

phytosaur Phytosaurs (Φυτόσαυροι in greek) are an extinct group of large, mostly semiaquatic Late Triassic archosauriform reptiles. Phytosaurs belong to the order Phytosauria. Phytosauria and Phytosauridae are often considered to be equivalent ...

thalattosaur Thalattosauria (Greek for "sea lizards") is an extinct order of prehistoric marine reptiles that lived in the middle to late Triassic period. Thalattosaurs were diverse in size and shape, and are divided into two superfamilies: Askeptosauroidea a ...

ians,

saurosphargid Saurosphargidae is an extinct family of marine reptiles known from the early Middle Triassic (Anisian stage) of Europe and China. The type species of the family is '' Saurosphargis volzi'', named by Friedrich von Huene in 1936 based on a single ...

ichthyosaur Ichthyosaurs (Ancient Greek for "fish lizard" – and ) are large extinct marine reptiles. Ichthyosaurs belong to the order known as Ichthyosauria or Ichthyopterygia ('fish flippers' – a designation introduced by Sir Richard Owen in 1842, altho ...

iforms,

sauropterygia Sauropterygia ("lizard flippers") is an extinct taxon of diverse, aquatic reptiles that developed from terrestrial ancestors soon after the end-Permian extinction and flourished during the Triassic before all except for the Plesiosauria became ...

ns, pleurosaurids or

mosasaur Mosasaurs (from Latin ''Mosa'' meaning the 'Meuse', and Greek ' meaning 'lizard') comprise a group of extinct, large marine reptiles from the Late Cretaceous. Their first fossil remains were discovered in a limestone quarry at Maastricht on th ...

oids, as well as mammalian

cetacea Cetacea (; , ) is an infraorder of aquatic mammals that includes whales, dolphins, and porpoises. Key characteristics are their fully aquatic lifestyle, streamlined body shape, often large size and exclusively carnivorous diet. They propel them ...

ns or

sirenia The Sirenia (), commonly referred to as sea-cows or sirenians, are an order of fully aquatic, herbivorous mammals that inhabit swamps, rivers, estuaries, marine wetlands, and coastal marine waters. The Sirenia currently comprise two distinct f ...

ns). The family has a wide geographic distribution, with material found in

Argentina Argentina (), officially the Argentine Republic ( es, link=no, República Argentina), is a country in the southern half of South America. Argentina covers an area of , making it the second-largest country in South America after Brazil, th ...

Chile Chile, officially the Republic of Chile, is a country in the western part of South America. It is the southernmost country in the world, and the closest to Antarctica, occupying a long and narrow strip of land between the Andes to the east a ...

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England England is a country that is part of the United Kingdom. It shares land borders with Wales to its west and Scotland to its north. The Irish Sea lies northwest and the Celtic Sea to the southwest. It is separated from continental Europe b ...

France France (), officially the French Republic ( ), is a country primarily located in Western Europe. It also comprises of Overseas France, overseas regions and territories in the Americas and the Atlantic Ocean, Atlantic, Pacific Ocean, Pac ...

Germany Germany,, officially the Federal Republic of Germany, is a country in Central Europe. It is the second most populous country in Europe after Russia, and the most populous member state of the European Union. Germany is situated betwe ...

Italy Italy ( it, Italia ), officially the Italian Republic, ) or the Republic of Italy, is a country in Southern Europe. It is located in the middle of the Mediterranean Sea, and its territory largely coincides with the homonymous geographical re ...

Mexico Mexico (Spanish: México), officially the United Mexican States, is a country in the southern portion of North America. It is bordered to the north by the United States; to the south and west by the Pacific Ocean; to the southeast by Guatema ...

Poland Poland, officially the Republic of Poland, is a country in Central Europe. It is divided into 16 administrative provinces called voivodeships, covering an area of . Poland has a population of over 38 million and is the fifth-most populous ...

Russia Russia (, , ), or the Russian Federation, is a List of transcontinental countries, transcontinental country spanning Eastern Europe and North Asia, Northern Asia. It is the List of countries and dependencies by area, largest country in the ...

Switzerland ). Swiss law does not designate a ''capital'' as such, but the federal parliament and government are installed in Bern, while other federal institutions, such as the federal courts, are in other cities (Bellinzona, Lausanne, Luzern, Neuchâtel ...

and

Czech Republic The Czech Republic, or simply Czechia, is a landlocked country in Central Europe. Historically known as Bohemia, it is bordered by Austria to the south, Germany to the west, Poland to the northeast, and Slovakia to the southeast. The ...

Classification

Phylogenetic In biology, phylogenetics (; from Greek φυλή/ φῦλον [] "tribe, clan, race", and wikt:γενετικός, γενετικός [] "origin, source, birth") is the study of the evolutionary history and relationships among or within groups o ...

analyses published during the 2000s cast doubt on the idea that many traditional metriorhynchid genera formed natural groups (i.e., include all descendants of a common ancestor). The traditional species of ''

Geosaurus ''Geosaurus'' is an extinct genus of marine crocodyliform within the family Metriorhynchidae, that lived during the Late Jurassic and the Early Cretaceous. ''Geosaurus'' was a carnivore that spent much, if not all, its life out at sea. No ''Geosa ...

'', ''Dakosaurus'' and ''

Cricosaurus ''Cricosaurus'' is an extinct genus of marine crocodyliforms of the Late Jurassic. belonging to the family Metriorhynchidae. The genus was established by Johann Andreas Wagner in 1858 for three skulls from the Tithonian (Late Jurassic) of German ...

'' were found to represent unnatural groups, and the species traditionally classified in these genera were reshuffled in a study published in November 2009 by Mark T. Young and Marco Brandalise de Andrade. The

monophyly In cladistics for a group of organisms, monophyly is the condition of being a clade—that is, a group of taxa composed only of a common ancestor (or more precisely an ancestral population) and all of its lineal descendants. Monophyletic grou ...

of ''

Metriorhynchus ''Metriorhynchus'' is an extinct genus of marine crocodyliform that lived in the oceans during the Late Jurassic. The type species, ''M. brevirostris'' was named in 1829 as a species of ''Steneosaurus'' before being named as a separate genus by ...

'' and ''

Teleidosaurus ''Teleidosaurus'' is an extinct genus of carnivorous metriorhynchoid crocodyliform from Middle Jurassic (late Bajocian to early Bathonian stage) deposits of Normandy, France.Wilkinson LE, Young MT, Benton MJ. 2008. A new metriorhynchid crocodilia ...

'' is also unsupported, and the species of these genera are pending reclassification. The classification presented by Young and Andrade in 2009 was approved in later studies of the Metriorhynchidae. Metriorhynchidae is a

node-based taxon Phylogenetic nomenclature is a method of nomenclature for taxa in biology that uses phylogenetic definitions for taxon names as explained below. This contrasts with the traditional approach, in which taxon names are defined by a '' type'', which ...

defined in 2009 as the least inclusive clade consisting of ''

Metriorhynchus geoffroyii ''Metriorhynchus'' is an extinct genus of marine crocodyliform that lived in the oceans during the Late Jurassic. The type species, ''M. brevirostris'' was named in 1829 as a species of ''Steneosaurus'' before being named as a separate genus by ...

'' and ''

Geosaurus giganteus ''Geosaurus'' is an extinct genus of marine crocodyliform within the family Metriorhynchidae, that lived during the Late Jurassic and the Early Cretaceous. ''Geosaurus'' was a carnivore that spent much, if not all, its life out at sea. No ''Geo ...

''. The

cladogram A cladogram (from Greek ''clados'' "branch" and ''gramma'' "character") is a diagram used in cladistics to show relations among organisms. A cladogram is not, however, an evolutionary tree because it does not show how ancestors are related to d ...

below follows the topology from the 2020 analyses by Young ''et al.'' and reduced to genera only.Young, M.T., Brignon, A., Sachs, S., Hornung J.J., Foffa, D., Kitson, J.J.N., Johnson, M.M., and Steel, L. 2020. Cutting the Gordian knot: a historical and taxonomic revision of the Jurassic crocodylomorph ''Metriorhynchus''. ''Zoological Journal of the Linnean Society'', zlaa092. https://doi.org/10.1093/zoolinnean/zlaa092.

List of genera

The

type genus In biological taxonomy, the type genus is the genus which defines a biological family and the root of the family name. Zoological nomenclature According to the International Code of Zoological Nomenclature, "The name-bearing type of a nominal f ...

of the family Metriorhynchidae is ''

'' from the Middle to Late Jurassic. Other genera included within this family are ''

'', ''

'', and ''

Dakosaurus ''Dakosaurus'' is an extinct genus of crocodylomorph within the family Metriorhynchidae that lived during the Late Jurassic and Early Cretaceous. It was large, with teeth that were serrated and compressed lateromedially (flattened from side to ...

''. Though once considered a metriorhynchid, ''

'' has since been found to be slightly more distantly related to these animals within the superfamily

Metriorhynchoidea Metriorhynchoidea is an extinct superfamily of thalattosuchian crocodyliforms from the Early Jurassic to the Early Cretaceous (Toarcian - Valanginian, possibly as late as early Aptian) of Europe, North America and South America. Metriorhynchids a ...

. Within this family, the genus '' Neustosaurus'' and ''

Enaliosuchus ''Enaliosuchus'' is a dubious genus of extinct marine crocodyliform within the family Metriorhynchidae that lived during the Valanginian stage of the Early Cretaceous. It is known from fossil remains found in France and Germany and it was firs ...

'' are considered ''

nomen dubium In binomial nomenclature, a ''nomen dubium'' (Latin for "doubtful name", plural ''nomina dubia'') is a scientific name that is of unknown or doubtful application. Zoology In case of a ''nomen dubium'' it may be impossible to determine whether a s ...

'' ("doubtful name"). The genus ''

Capelliniosuchus ''Capelliniosuchus'' is an extinct genus of mosasaur. It was discovered in Italy, and described by Simonelli,Simonelli V. 1896. Intoro agli avanzi di coccodrilliano scoperti a San Valentino (provincial di Reggio Emilia) nel 1886. ''Atli della Re ...

'' was once thought to be a metriorhynchid similar to ''Dakosaurus''. However, it was later found to be a

References

{{Taxonbar, from=Q134271 Middle Jurassic crocodylomorphs Early Cretaceous crocodylomorphs Prehistoric reptile families Prehistoric marine crocodylomorphs Middle Jurassic first appearances Early Cretaceous extinctions Late Jurassic crocodylomorphs

Description

Classification

List of genera

See also

References