Metriorhynchidae is an

extinct Extinction is the termination of an organism by the death of its Endling, last member. A taxon may become Functional extinction, functionally extinct before the death of its last member if it loses the capacity to Reproduction, reproduce and ...

family Family (from ) is a Social group, group of people related either by consanguinity (by recognized birth) or Affinity (law), affinity (by marriage or other relationship). It forms the basis for social order. Ideally, families offer predictabili ...

of specialized, aquatic metriorhynchoid

crocodyliform Crocodyliformes is a clade of Crurotarsi, 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 ...

s from the

Middle Jurassic The Middle Jurassic is the second Epoch (geology), epoch of the Jurassic Period (geology), Period. It lasted from about 174.1 to 161.5 million years ago. Fossils of land-dwelling animals, such as dinosaurs, from the Middle Jurassic are relativel ...

to the

Early Cretaceous The Early Cretaceous (geochronology, geochronological name) or the Lower Cretaceous (chronostratigraphy, chronostratigraphic name) is the earlier or lower of the two major divisions of the Cretaceous. It is usually considered to stretch from 143.1 ...

period (

Bajocian In the geologic timescale, the Bajocian is an age and stage in the Middle Jurassic. It lasted from approximately 170.9 ±0.8 Ma to around 168.2 ±1.2 Ma (million years ago). The Bajocian Age succeeds the Aalenian Age and precedes the Bathonia ...

to early

Aptian The Aptian is an age (geology), age in the geologic timescale or a stage (stratigraphy), stage in the stratigraphic column. It is a subdivision of the Early Cretaceous, Early or Lower Cretaceous epoch (geology), Epoch or series (stratigraphy), S ...

) of

Europe Europe is a continent located entirely in the Northern Hemisphere and mostly in the Eastern Hemisphere. It is bordered by the Arctic Ocean to the north, the Atlantic Ocean to the west, the Mediterranean Sea to the south, and Asia to the east ...

North America North America is a continent in the Northern Hemisphere, Northern and Western Hemisphere, Western hemispheres. North America is bordered to the north by the Arctic Ocean, to the east by the Atlantic Ocean, to the southeast by South Ameri ...

and

South America South America is a continent entirely in the Western Hemisphere and mostly in the Southern Hemisphere, with a considerably smaller portion in the Northern Hemisphere. It can also be described as the southern Subregion#Americas, subregion o ...

. The name Metriorhynchidae was coined by the Austrian

zoologist Zoology ( , ) is the scientific study of animals. Its studies include the structure, embryology, classification, habits, and distribution of all animals, both living and extinct, and how they interact with their ecosystems. Zoology is one ...

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 and the Geosaurinae. 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 () is an order of semiaquatic, predatory reptiles that are known as crocodilians. They first appeared during the Late Cretaceous and are the closest living relatives of birds. Crocodilians are a type of crocodylomorph pseudosuchian ...

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 taxon, extant and extinct reptiles and amphibians, including lizards, crocodilians, frogs, Temnospondyli, ...

("armour scutes"). Their body shape maximised hydrodynamy (swimming efficiency), as they did have a shark-like tail fluke. Like

ichthyosaurs Ichthyosauria is an taxonomy (biology), order of large extinction, extinct marine reptiles sometimes referred to as "ichthyosaurs", although the term is also used for wider clades in which the order resides. Ichthyosaurians thrived during much of ...

and

plesiosaurs The Plesiosauria or plesiosaurs are an Order (biology), order or clade of extinct Mesozoic marine reptiles, belonging to the Sauropterygia. Plesiosaurs first appeared in the latest Triassic Period (geology), Period, possibly in the Rhaetian st ...

, metriorhynchids developed smooth, scaleless skin. However, unlike those marine reptiles, they were

poikilothermic A poikilotherm () is an animal (Greek ''poikilos'' – 'various', 'spotted', and ''therme'' – 'heat') whose internal temperature varies considerably. Poikilotherms have to survive and adapt to environmental stress. One of the most important s ...

with little to no

endothermic An endothermic process is a chemical or physical process that absorbs heat from its surroundings. In terms of thermodynamics, it is a thermodynamic process with an increase in the enthalpy (or internal energy ) of the system.Oxtoby, D. W; Gillis, ...

ability. Metriorhynchids were the only group of

archosaur Archosauria () or archosaurs () is a clade of diapsid sauropsid tetrapods, with birds and crocodilians being the only extant taxon, extant representatives. Although broadly classified as reptiles, which traditionally exclude birds, the cladistics ...

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. The word ''pelagic'' is derived . The pelagic zone can be thought of as an imaginary cylinder or water column between the sur ...

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 that studies the microscopic anatomy of biological tissue (biology), tissues. Histology is the microscopic counterpart to gross anatomy, which looks at large ...

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 In animals, viviparity is development of the embryo inside the body of the mother, with the maternal circulation providing for the metabolic needs of the embryo's development, until the mother gives birth to a fully or partially developed juve ...

'' Keichousaurus'', these characters suggest that metriorhynchids gave live birth. A fossil of a pregnant ''Dakosaurus'' female recovered from the Late Jurassic plattenkalk, 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 ( Cisuralian), roughly 299 to 270 million years ago. Mesosaurs were the first known aquatic reptiles, having apparently returned to a ...

phytosaur Phytosaurs (Φυτόσαυροι in Greek, meaning 'plant lizard') are an extinct group of large, mostly semiaquatic Late Triassic archosauriform or basal archosaurian reptiles. Phytosaurs belong to the order Phytosauria and are sometimes ref ...

s, thalattosaurians, saurosphargids,

ichthyosaur Ichthyosauria is an order of large extinct marine reptiles sometimes referred to as "ichthyosaurs", although the term is also used for wider clades in which the order resides. Ichthyosaurians thrived during much of the Mesozoic era; based on fo ...

iforms,

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

ns, pleurosaurids or

mosasaur Mosasaurs (from Latin ''Mosa'' meaning the 'Meuse', and Ancient Greek, Greek ' meaning 'lizard') are an extinct group of large aquatic reptiles within the family Mosasauridae that lived during the Late Cretaceous. Their first fossil remains wer ...

oids, as well as mammalian

cetacea Cetacea (; , ) is an infraorder of aquatic mammals belonging to the order Artiodactyla that includes whales, dolphins and porpoises. Key characteristics are their fully aquatic lifestyle, streamlined body shape, often large size and exclusively c ...

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 extant Sirenia comprise two distinct famili ...

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

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and

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Classification

Phylogenetic In biology, phylogenetics () is the study of the evolutionary history of life using observable characteristics of organisms (or genes), which is known as phylogenetic inference. It infers the relationship among organisms based on empirical dat ...

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'', ''Dakosaurus'' and '' Cricosaurus'' 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 biological cladistics for the classification of organisms, monophyly is the condition of a taxonomic grouping being a clade – that is, a grouping of organisms which meets these criteria: # the grouping contains its own most recent comm ...

of '' Metriorhynchus'' and '' Teleidosaurus'' 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 taxon, taxa in biology that uses phylogenetics, phylogenetic definitions for taxon names as explained below. This contrasts with Biological classification, the traditional method, by which ...

defined in the ''

PhyloCode The ''International Code of Phylogenetic Nomenclature'', known as the ''PhyloCode'' for short, is a formal set of rules governing phylogenetic nomenclature. Its current version is specifically designed to regulate the naming of clades, leaving the ...

'' by Mark T. Young and colleagues in 2024 as "the smallest clade within Metriorhynchoidea containing '' Thalattosuchus superciliosus'', '' Gracilineustes leedsi'', '' Metriorhynchus brevirostris'', '' Rhacheosaurus gracilis'', and '' Geosaurus giganteus''" The

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

below follows the topology from the 2020 analyses by Young ''et al.'' and reduced to genera only.

List of genera

The

type genus In biological taxonomy, the type genus (''genus typica'') 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-bearin ...

of the family Metriorhynchidae is '' Metriorhynchus'' from the Middle to Late Jurassic. Other genera included within this family are '' Cricosaurus'', '' Geosaurus'', and '' Dakosaurus''. Though once considered a metriorhynchid, '' Teleidosaurus'' has since been found to be slightly more distantly related to these animals within the superfamily Metriorhynchoidea. Within this family, the genus '' Neustosaurus'' and '' Enaliosuchus'' 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 ...

'' ("doubtful name"). The genus '' Capelliniosuchus'' was once thought to be a metriorhynchid similar to ''Dakosaurus''. However, it was later found to be a

Subclades

Summary of the phylogenetic definitions of metriorhynchid subclades shown in the classification section. These are based on Young and colleagues 2024.

References

External links