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''Protorosaurus'' (from , 'earlier' and , 'lizard') is an extinct genus of reptile. Members of the genus lived during the late
Permian The Permian ( ) is a geologic period and System (stratigraphy), stratigraphic system which spans 47 million years, from the end of the Carboniferous Period million years ago (Mya), to the beginning of the Triassic Period 251.902 Mya. It is the s ...
period in what is now
Germany Germany, officially the Federal Republic of Germany, is a country in Central Europe. It lies between the Baltic Sea and the North Sea to the north and the Alps to the south. Its sixteen States of Germany, constituent states have a total popu ...
and
Great Britain Great Britain is an island in the North Atlantic Ocean off the north-west coast of continental Europe, consisting of the countries England, Scotland, and Wales. With an area of , it is the largest of the British Isles, the List of European ...
. Once believed to have been an ancestor to
lizard Lizard is the common name used for all Squamata, squamate reptiles other than snakes (and to a lesser extent amphisbaenians), encompassing over 7,000 species, ranging across all continents except Antarctica, as well as most Island#Oceanic isla ...
s, ''Protorosaurus'' is now known to be one of the oldest and most primitive members of
Archosauromorpha Archosauromorpha ( Greek for "ruling lizard forms") is a clade of diapsid reptiles containing all reptiles more closely related to archosaurs (such as crocodilians and dinosaurs, including birds) than to lepidosaurs (such as tuataras, lizards, ...
, the group that would eventually lead to
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 such as
crocodilian Crocodilia () is an Order (biology), 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 crocodylomorp ...
s and
dinosaur Dinosaurs are a diverse group of reptiles of the clade Dinosauria. They first appeared during the Triassic Geological period, period, between 243 and 233.23 million years ago (mya), although the exact origin and timing of the #Evolutio ...
s.


Description

''Protorosaurus'' grew up to in length, and was a slender, lizard-like animal, vaguely resembling a
monitor lizard Monitor lizards are lizards in the genus ''Varanus,'' the only extant genus in the family Varanidae. They are native to Africa, Asia, and Oceania, and West African Nile monitor, one species is also found in south America as an invasive species. A ...
, with long legs and a long neck.


History

''Protorosaurus'' was one of the first fossil reptiles to be described, being initially described in Latin in 1710 by from a specimen found in
Thuringia Thuringia (; officially the Free State of Thuringia, ) is one of Germany, Germany's 16 States of Germany, states. With 2.1 million people, it is 12th-largest by population, and with 16,171 square kilometers, it is 11th-largest in area. Er ...
in Germany, who considered the animal to be a crocodile, and most similar to the
Nile crocodile The Nile crocodile (''Crocodylus niloticus'') is a large crocodilian native to freshwater habitats in Africa, where it is present in 26 countries. It is widely distributed in sub-Saharan Africa, occurring mostly in the eastern, southern, and ce ...
(''C. niloticus''). Over a century later, in publications in 1830 and 1832
Hermann von Meyer Christian Erich Hermann von Meyer (3 September 1801 – 2 April 1869), known as Hermann von Meyer, was a German palaeontologist. He was awarded the 1858 Wollaston medal by the Geological Society of London. Life He was born in Frankfurt am ...
recognised ''Protorosaurus'' as distinct extinct reptile and gave it a formal species description, dedicating an extensive
monograph A monograph is generally a long-form work on one (usually scholarly) subject, or one aspect of a subject, typically created by a single author or artist (or, sometimes, by two or more authors). Traditionally it is in written form and published a ...
to it in 1856. In 1871
Thomas Henry Huxley Thomas Henry Huxley (4 May 1825 – 29 June 1895) was an English biologist and anthropologist who specialized in comparative anatomy. He has become known as "Darwin's Bulldog" for his advocacy of Charles Darwin's theory of evolution. The stor ...
erected the clade
Protorosauria Protorosauria is an extinct, likely paraphyletic group of basal archosauromorph reptiles from the latest Middle Permian ( Capitanian stage) to the end of the Late Triassic ( Rhaetian stage) of Asia, Europe and North America. It was named by t ...
, with ''Protorosaurus'' as the only member. The German specimens were found in the
Kupferschiefer The Kupferschiefer (German for Copper Shale, Copper Schist or Copper Slate) or Kupfermergel (Copper Marl), (T1 or Z1) is an extensive and remarkable sedimentary unit in Central Europe. The relatively monotonous succession is typically and maximu ...
a widespread unit of Late Permian (likely
Wuchiapingian In the geologic timescale, the Wuchiapingian or Wujiapingian (from in the Liangshan area of Hanzhong, Shaanxi Province ) is an age or stage of the Permian. It is also the lower or earlier of two subdivisions of the Lopingian Epoch or S ...
) age. In 1914, a new
ceratopsian Ceratopsia or Ceratopia ( or ; Greek: "horned faces") is a group of herbivorous, beaked dinosaurs that thrived in what are now North America, Asia and Europe, during the Cretaceous Period, although ancestral forms lived earlier, in the Late Ju ...
dinosaur found by
Lawrence Lambe Lawrence Morris Lambe (August 27, 1863 – March 12, 1919) was a Canadian geologist, palaeontologist, and ecologist from the Geological Survey of Canada (GSC). His published work, describing the diverse and plentiful dinosaur discoveries from t ...
was again given the name ''Protorosaurus'' (in this sense meaning "before ''
Torosaurus ''Torosaurus'' (meaning "perforated lizard", in reference to the large openings in its frill) is a genus of herbivorous Chasmosaurinae, chasmosaurine Ceratopsia, ceratopsian dinosaur that lived during the late Maastrichtian age of the Late Cret ...
''"). When Lambe found that the name had already been used for the early archosauromorph, he renamed his ceratopsian ''
Chasmosaurus ''Chasmosaurus'' ( ) is a genus of ceratopsid dinosaur from the Late Cretaceous Period in North America. Its given name means 'opening lizard', referring to the large openings ( fenestrae) in its frill (Greek ''chasma'', meaning 'opening', 'hol ...
''. In 1870, a new species, ''Protorosaurus huxleyi'' was described from a
railway cutting Rail transport (also known as train transport) is a means of transport using wheeled vehicles running in tracks, which usually consist of two parallel steel rails. Rail transport is one of the two primary means of land transport, next to road ...
near
Middridge Middridge is a village and civil parish in County Durham, England. It is situated east of Shildon and north-west of Newton Aycliffe. The village is situated near a quarry that was mined by the people many generations ago. There is one public h ...
in
County Durham County Durham, officially simply Durham, is a ceremonial county in North East England.UK General Acts 1997 c. 23Lieutenancies Act 1997 Schedule 1(3). From legislation.gov.uk, retrieved 6 April 2022. The county borders Northumberland and Tyne an ...
in northern England, in sediments belonging to the
Marl Slate The Marl Slate Formation is a geological formation in England. Despite its name, it is mostly dolomite rock. The Marl Slate Formation was formed about 273 to 259 million years ago, during the Guadalupian and Lopingian epochs of the late Permian ...
, a unit that is equivalent to the Kupferschiefer, however it was later shown that this specimen was not ''Protorosaurus'', and it was subsequently placed in the new genus '' Adelosaurus''. In 1993, a specimen of ''Protorosaurus'' was described from Quarrington Quarry also in County Durham, in sediments of the Marl Slate. This skeleton included fragments of the skull that had not previously been known. A thorough redescription of ''Protorosaurus'' was published in 2009, including a specimen with a complete skull. The complete skull ( NMK S 180) was redescribed again in 2025 according to new data from micro-CT scans.


Paleobiology

Stomach contents of a specimen of ''Protorosaurus'' show the presence of numerous ovules of the locally common voltzialean conifer '' Ullmannia frumentaria'', which was unexpected given its unspecialised dentiton of conical and unserrated teeth, which were assumed to represent a predatory lifestyle. Along with the coniferous ovules, several rounded pebbles were found in the body cavity, these may have functioned as
gastroliths A gastrolith, also called a stomach stone or gizzard stone, is a rock held inside a gastrointestinal tract. Gastroliths in some species are retained in the muscular gizzard and used to grind food in animals lacking suitable grinding teeth. In oth ...
.


Classification

''Protorosaurus'' has been suggested to be closely related to '' Czatkowiella'' from the Early Triassic of Poland, but the remains of ''Czatkowiella'' are fragmentary and mixed in with those of other reptiles, so it is possible that the taxon is a
chimaera Chimaeras are Chondrichthyes, cartilaginous fish in the order (biology), order Chimaeriformes (), known informally as ghost sharks, rat fish (not to be confused with rattails), spookfish, or rabbit fish; the last two names are also applied to B ...
. In a comprehensive analysis of "Protorosauria" in 2021, ''Protorosaurus'' was consistently recovered as the earliest diverging member of
Archosauromorpha Archosauromorpha ( Greek for "ruling lizard forms") is a clade of diapsid reptiles containing all reptiles more closely related to archosaurs (such as crocodilians and dinosaurs, including birds) than to lepidosaurs (such as tuataras, lizards, ...
. The cladogram below follows an analysis by Ezcurra (2016), and highlights the position of ''Protorosaurus'' among other early archosauromorph reptiles. Cladogram after Spiekman et al. in 2024.


Paleoenvironment

The Kupferschiefer and equivalent Marl Slate is a marine unit that forms part of the
Zechstein The Zechstein ( German either from ''mine stone'' or ''tough stone'') is a unit of sedimentary rock layers of Late Permian ( Lopingian) age located in the European Permian Basin which stretches from the east coast of England to northern Poland. T ...
, a sequence of rocks formed on the edge of the Zechstein Sea, a large inland shallow sea that existed in Northern Europe during the Late Permian. The environment at the time of deposition is considered to have been
semi-arid A semi-arid climate, semi-desert climate, or steppe climate is a aridity, dry climate sub-type. It is located on regions that receive precipitation below Evapotranspiration#Potential evapotranspiration, potential evapotranspiration, but not as l ...
. The terrestrial flora of the Zechstein is dominated by conifers, with
seed ferns Pteridospermatophyta, also called pteridosperms or seed ferns, are a polyphyletic grouping of extinct Spermatophyte, seed-producing plants. The earliest fossil evidence for plants of this type are the Lyginopteridales, lyginopterids of late Devon ...
also being common, while taeniopterids, ginkgophytes and sphenophytes are rare. Other terrestrial vertebrates found in the Kupfershiefer and lower Zechstein include the gliding weigeltisaurid reptiles ''
Weigeltisaurus ''Weigeltisaurus'' is an extinct genus of weigeltisaurid reptile from the Late Permian Kupferschiefer of Germany and Marl Slate of England. It has a single species, originally named as ''Palaechamaeleo jaekeli'' in 1930 and later assigned the na ...
'' and ''
Glaurung J. R. R. Tolkien's Middle-earth legendarium features dragons based on those of European legend, but going beyond them in having personalities of their own, such as the wily Smaug, who has features of both Fafnir and the ''Beowulf'' dragon. ...
,'' the
pareiasaur Pareiasaurs (meaning "cheek lizards") are an extinct clade of large, herbivorous parareptiles. Members of the group were armoured with osteoderms which covered large areas of the body. They first appeared in southern Pangea during the Middle Per ...
''
Parasaurus ''Parasaurus'' (meaning "near lizard") is a genus of pareiasaur known from fossils collected in the Kupferschiefer in Germany (Hesse, Thuringia and Lower Saxony), dating to the Late Permian (Wuchiapingian). The type species, ''Parasaurus geinitz ...
,'' the
cynodont Cynodontia () is a clade of eutheriodont therapsids that first appeared in the Late Permian (approximately 260 Megaannum, mya), and extensively diversified after the Permian–Triassic extinction event. Mammals are cynodonts, as are their extin ...
''
Procynosuchus ''Procynosuchus'' (Greek: "Before dog crocodile") is an extinct genus of cynodonts from the Late Permian. It is considered to be one of the earliest and most basal (phylogenetics), basal cynodonts. It was 60 cm (2 ft) long. Remains of ...
,'' and indeterminate
captorhinids Captorhinidae is an extinct family of tetrapods, traditionally considered primitive reptiles, known from the late Carboniferous to the Late Permian. They had a cosmopolitan distribution across Pangea. Description Captorhinids are a clade of sm ...
,
dicynodont Dicynodontia is an extinct clade of anomodonts, an extinct type of non-mammalian therapsid. Dicynodonts were herbivores that typically bore a pair of tusks, hence their name, which means 'two dog tooth'. Members of the group possessed a horny, t ...
s, and dissorophid
temnospondyls Temnospondyli (from Greek τέμνειν, ''temnein'' 'to cut' and σπόνδυλος, ''spondylos'' 'vertebra') or temnospondyls is a diverse ancient order of small to giant tetrapods—often considered primitive amphibians—that flourished wo ...
.


References

{{Taxonbar, from=Q134702 Archosauromorpha Prehistoric reptile genera Wuchiapingian life Lopingian reptiles of Europe Permian Germany Fossils of Germany Kupferschiefer Fossil taxa described in 1830 Taxa named by Christian Erich Hermann von Meyer