Thecodontosaurus Dubius
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''Thecodontosaurus'' ("socket-tooth lizard") 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
herbivorous A herbivore is an animal anatomically and physiologically evolved to feed on plants, especially upon vascular tissues such as foliage, fruits or seeds, as the main component of its diet. These more broadly also encompass animals that eat n ...
basal
sauropodomorph Sauropodomorpha ( ; from Greek, meaning "lizard-footed forms") is an extinct clade of long-necked, herbivorous, saurischian dinosaurs that includes the sauropods and their ancestral relatives. Sauropods generally grew to very large sizes, had lo ...
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 ...
that lived during the late
Triassic period The Triassic ( ; sometimes symbolized 🝈) is a geologic period and system which spans 50.5 million years from the end of the Permian Period 251.902 million years ago ( Mya), to the beginning of the Jurassic Period 201.4 Mya. The Triassic is the ...
(
Carnian The Carnian (less commonly, Karnian) is the lowermost stage (stratigraphy), stage of the Upper Triassic series (stratigraphy), Series (or earliest age (geology), age of the Late Triassic Epoch (reference date), Epoch). It lasted from 237 to 227.3 ...
? age). Its remains are known mostly from Triassic "fissure fillings" in
South England Southern England, also known as the South of England or the South, is a sub-national part of England. Officially, it is made up of the southern, south-western and part of the eastern parts of England, consisting of the statistical regions of ...
. ''Thecodontosaurus'' was a small bipedal animal, about 2 m (6.5 ft) long. It is one of the first dinosaurs to be discovered and is one of the oldest that existed. Many species have been named in the genus, but only the
type species In International_Code_of_Zoological_Nomenclature, 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 spe ...
''Thecodontosaurus antiquus'' is seen as valid today.


Discovery and naming


''Thecodontosaurus antiquus''

In the autumn of 1834,
surgeon In medicine, a surgeon is a medical doctor who performs surgery. Even though there are different traditions in different times and places, a modern surgeon is a licensed physician and received the same medical training as physicians before spec ...
Henry Riley (1797–1848) and the
curator A curator (from , meaning 'to take care') is a manager or overseer. When working with cultural organizations, a curator is typically a "collections curator" or an "exhibitions curator", and has multifaceted tasks dependent on the particular ins ...
of the Bristol Institution,
Samuel Stutchbury Samuel Stutchbury (15 January 1798 – 12 February 1859) was an English people, English natural history, naturalist and geology, geologist. Alongside Henry Riley (scientist), Henry Riley, Stutchbury was the co-discoverer of ''Thecodontosaurus'', ...
, began to excavate "saurian remains" at the quarry of
Durdham Down Durdham Down is an area of public open space in Bristol, England England is a Countries of the United Kingdom, country that is part of the United Kingdom. It is located on the island of Great Britain, of which it covers about 62%, and ...
, at Clifton, presently a part of
Bristol Bristol () is a City status in the United Kingdom, cathedral city, unitary authority area and ceremonial county in South West England, the most populous city in the region. Built around the River Avon, Bristol, River Avon, it is bordered by t ...
, which is part of the
Magnesian Conglomerate The Magnesian Conglomerate is a geological Formation (geology), formation in Clifton, Bristol in England (originally Avon (county), Avon), Gloucestershire and southern Wales, present in Tytherington, Gloucestershire, Tytherington, Durdham Down, S ...
. In 1834 and 1835, they briefly reported on the finds. They provided their initial description in 1836, naming a new genus: ''Thecodontosaurus''. The name is derived from Greek θήκή, ', "socket", and οδους, ', "tooth", a reference to the fact that the roots of the teeth were not fused with the jaw bone, as in present lizards, but positioned in separate tooth sockets. ''Thecodontosaurus'' was the fifth dinosaur named, after ''
Megalosaurus ''Megalosaurus'' (meaning "great lizard", from Ancient Greek, Greek , ', meaning 'big', 'tall' or 'great' and , ', meaning 'lizard') is an extinct genus of large carnivorous theropod dinosaurs of the Middle Jurassic Epoch (Bathonian stage, 166 ...
'', ''
Iguanodon ''Iguanodon'' ( ; meaning 'iguana-tooth'), named in 1825, is a genus of iguanodontian dinosaur. While many species found worldwide have been classified in the genus ''Iguanodon'', dating from the Late Jurassic to Early Cretaceous, Taxonomy (bi ...
'', ''
Streptospondylus ''Streptospondylus'', from Ancient Greek στρεπτός (''streptós''), meaning "twisted", and σπόνδυλος (''spóndulos''), meaning "vertebra", is a genus of tetanuran theropod dinosaur known from the Late Jurassic period of France, 1 ...
'' and ''
Hylaeosaurus ''Hylaeosaurus'' ( ; Greek: / "belonging to the forest" and / "lizard") is a herbivorous ankylosaurian dinosaur that lived about 136 million years ago, in the late Valanginian stage of the early Cretaceous period of England. It was found i ...
'', though Riley and Stutchbury were not aware of this, the very concept of Dinosauria only being created in 1842. In 1843, in his catalogue of British fossils, John Morris provided a complete
species name In taxonomy, binomial nomenclature ("two-term naming system"), also called binary nomenclature, is a formal system of naming species of living things by giving each a name composed of two parts, both of which use Latin grammatical forms, altho ...
: ''Thecodontosaurus antiquus''. The
specific epithet In Taxonomy (biology), taxonomy, binomial nomenclature ("two-term naming system"), also called binary nomenclature, is a formal system of naming species of living things by giving each a name composed of two parts, both of which use Latin gramm ...
, ''"antiquus"'', means "ancient" in
Latin Latin ( or ) is a classical language belonging to the Italic languages, Italic branch of the Indo-European languages. Latin was originally spoken by the Latins (Italic tribe), Latins in Latium (now known as Lazio), the lower Tiber area aroun ...
. The original
type specimen In biology, a type is a particular wikt:en:specimen, specimen (or in some cases a group of specimens) of an organism to which the scientific name of that organism is formally associated. In other words, a type is an example that serves to ancho ...
or
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 ''Thecodontosaurus'', BCM 1, a lower jaw, fell victim to heavy
World War II World War II or the Second World War (1 September 1939 – 2 September 1945) was a World war, global conflict between two coalitions: the Allies of World War II, Allies and the Axis powers. World War II by country, Nearly all of the wo ...
bombings. Many remains of this dinosaur and other material related to it were destroyed in November 1940 during the
Bristol Blitz The Bristol Blitz was the heavy bombing of Bristol, England by the Nazi German ''Luftwaffe'' during the Second World War. Due to the presence of Bristol Harbour and the Bristol Aeroplane Company, the city was a target for bombing and was ea ...
. However, most bones were salvaged: today 184 fossil bones are part of the collection of the
Bristol City Museum and Art Gallery Bristol Museum & Art Gallery is a large museum and art gallery in Bristol, England. The museum is situated in Clifton, about from the city centre. As part of Bristol Culture and Creative Industries it is run by the Bristol City Council with no ...
. Later, more remains were found near
Bristol Bristol () is a City status in the United Kingdom, cathedral city, unitary authority area and ceremonial county in South West England, the most populous city in the region. Built around the River Avon, Bristol, River Avon, it is bordered by t ...
at Tytherington. Currently about 245 fragmentary specimens are known, representing numerous individuals. In 1985,
Peter Galton Peter Malcolm Galton (born 14 March 1942 in London) is a British vertebrate paleontologist who has to date written or co-written about 190 papers in scientific journals or chapters in paleontology textbooks, especially on ornithischian and prosau ...
designated another lower jaw, a right dentary, as the
neotype In biology, a type is a particular specimen (or in some cases a group of specimens) of an organism to which the scientific name of that organism is formally associated. In other words, a type is an example that serves to anchor or centralizes ...
, BCM 2. The remains had been found in
chalkstone Chalk is a soft, white, porous, sedimentary carbonate rock. It is a form of limestone composed of the mineral calcite and originally formed deep under the sea by the compression of microscopic plankton that had settled to the sea floor. Chalk ...
infillings,
breccia Breccia ( , ; ) is a rock composed of large angular broken fragments of minerals or Rock (geology), rocks cementation (geology), cemented together by a fine-grained matrix (geology), matrix. The word has its origins in the Italian language ...
deposited in fissures in older rocks. The age of these deposits was once estimated as old as the late
Carnian The Carnian (less commonly, Karnian) is the lowermost stage (stratigraphy), stage of the Upper Triassic series (stratigraphy), Series (or earliest age (geology), age of the Late Triassic Epoch (reference date), Epoch). It lasted from 237 to 227.3 ...
, but recent studies indicate that they date from the
Rhaetian The Rhaetian is the latest age (geology), age of the Triassic period (geology), Period (in geochronology) or the uppermost stage (stratigraphy), stage of the Triassic system (stratigraphy), System (in chronostratigraphy). It was preceded by the N ...
.


Other species

Apart from the original
type species In International_Code_of_Zoological_Nomenclature, 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 spe ...
, ''Thecodontosaurus antiquus'', Riley and Stutchbury also found some teeth of carnivorous
phytosauria Phytosaurs (Φυτόσαυροι in greek language, Greek, meaning 'plant lizard') are an extinct group of large, mostly semiaquatic Late Triassic archosauriform or Basal (phylogenetics), basal archosaurian reptiles. Phytosaurs belong to the ord ...
ns that they named ''
Palaeosaurus ''Palaeosaurus'' (or ''Paleosaurus'') is a genus of indeterminate archosaur known from two teeth found in the Bromsgrove Sandstone Formation and also either the Magnesian Conglomerate or the Avon Fissure Fill of Clifton, Bristol, England (or ...
cylindrodon'' and ''P. platyodon''. In the late nineteenth century, the theory became popular that such remains belonged to carnivorous
prosauropods Sauropodomorpha ( ; from Greek, meaning "lizard-footed forms") is an extinct clade of long-necked, herbivorous, saurischian dinosaurs that includes the sauropods and their ancestral relatives. Sauropods generally grew to very large sizes, had lo ...
: animals with the body of ''Thecodontosaurus'', but with slicing teeth. In 1890,
Arthur Smith Woodward Sir Arthur Smith Woodward, FRS (23 May 1864 – 2 September 1944) was an English palaeontologist, known as a world expert in fossil fish. He also described the Piltdown Man fossils, which were later determined to be fraudulent. He is not rel ...
accordingly named a ''Thecodontosaurus platyodon,'' and in 1908
Friedrich von Huene Baron Friedrich Richard von Hoyningen-Huene (22 March 1875 – 4 April 1969) was a German nobleman paleontologist who described a large number of dinosaurs, more than anyone else in 20th-century Europe. He studied a range of Permo-Carbonife ...
named a ''Thecodontosaurus cylindrodon''. Though still defended by Michael Cooper in 1981, the hypothesis that such creatures existed has now been totally discredited. On one occasion, material of ''Thecodontosaurus'' was, by mistake, described as a separate genus. In 1891,
Harry Govier Seeley Harry Govier Seeley (18 February 1839 – 8 January 1909) was a British paleontologist. Early life Seeley was born in London on 18 February 1839, the second son of Richard Hovill Seeley, a goldsmith, and his second wife Mary Govier. When his fa ...
named '' Agrosaurus macgillivrayi'', assuming the remains had been collected in 1844 by the crew of HMS ''Fly'' on the northeast coast of
Australia Australia, officially the Commonwealth of Australia, is a country comprising mainland Australia, the mainland of the Australia (continent), Australian continent, the island of Tasmania and list of islands of Australia, numerous smaller isl ...
. It was long considered the first dinosaur found in Australia, but in 1999 it was discovered that the bones probably belonged to a lot sent by Riley and Stutchbury to the
British Museum of Natural History The Natural History Museum in London is a museum that exhibits a vast range of specimens from various segments of natural history. It is one of three major museums on Exhibition Road in South Kensington, the others being the Science Museum (Lo ...
and then mislabelled. In 1906, von Huene had already noted the close resemblance and renamed the species ''Thecodontosaurus macgillivrayi''. It is thus a
junior synonym In taxonomy, the scientific classification of living organisms, a synonym is an alternative scientific name for the accepted scientific name of a taxon. The botanical and zoological codes of nomenclature treat the concept of synonymy differently. ...
of ''Thecodontosaurus antiquus''. Presently, the only valid species is thus ''T. antiquus''.


Misassigned species

*''Thecodontosaurus latespinatus'' von Huene, 1907-08 = ''
Tanystropheus ''Tanystropheus'' (~ 'long' + 'hinged') is an extinct genus of archosauromorph reptile which lived during the Triassic period, Triassic Period in Europe, Asia, and North America. It is recognisable by its extremely elongated neck, longer than t ...
'' *''Thecodontosaurus primus'' von Huene, 1907-1908 = indeterminate archosauromorph, previously and questionably referred to '' Protanystropheus'' *''Thecodontosaurus elizae'' Sauvage, 1907 *''Thecodontosaurus gibbidens'' Cope, 1878 = ''
Galtonia ''Galtonia'' is a genus of plants in the family Asparagaceae, subfamily Scilloideae. Native to Southern Africa, the genus is named after Sir Francis Galton. According to some authorities it has been subsumed into ''Ornithogalum'' as a subgenus ...
'' *''Thecodontosaurus skirtopodus'' (Seeley, 1894) = ''
Hortalotarsus ''Hortalotarsus'' is a dubious genus of sauropodomorph dinosaur from the Early Jurassic of southern Africa. The only species is ''H. skirtopodus''. ''Hortalotarsus'' was described by Harry Seeley in 1894 based on parts of a hind limb discovere ...
'' *''Thecodontosaurus polyzelus'' (Hitchcock, 1865) von Huene, 1906 *''Thecodontosaurus hermannianus'' von Huene, 1908 *''Thecodontosaurus diagnosticus'' Fraas, 1912 = ''
Efraasia ''Efraasia'' (pronounced "E-FRAHS-ee-A") is a genus of basal (phylogenetics), basal sauropodomorph dinosaur. It was a herbivore which lived during the middle Norian stage of the Late Triassic, around 210 million years ago, in what is now Germ ...
'' *''Thecodontosaurus minor'' Haughton, 1918 *''Thecodontosaurus dubius'' Haughton, 1924 *''Thecodontosaurus browni'' (Seeley, 1895) von Huene, 1932 *''Thecodontosaurus alophos'' Haughton, 1932 = ''
Nyasasaurus ''Nyasasaurus'' (meaning " Lake Nyasa lizard") is an extinct genus of avemetatarsalian archosaur from the putatively Middle Triassic Manda Formation of Tanzania that may be the earliest known dinosaur. The type species ''Nyasasaurus parringtoni'' ...
'' ''Thecodontosaurus caducus'' was named by
Adam Yates Adam Richard Yates (born 7 August 1992) is a British professional road and track racing cyclist who rides for UCI WorldTeam . Yates placed fourth overall at the 2016 Tour de France and became the first British rider to win the Young rider clas ...
in 2003 for a juvenile specimen found in
Wales Wales ( ) is a Countries of the United Kingdom, country that is part of the United Kingdom. It is bordered by the Irish Sea to the north and west, England to the England–Wales border, east, the Bristol Channel to the south, and the Celtic ...
; in 2007 this was made the separate genus ''
Pantydraco ''Pantydraco'' (where "panty-" is short for Pant-y-ffynnon, signifying ''hollow of the spring/well'' in Welsh, referring to the quarry at Bonvilston in South Wales where it was found) is a genus of basal sauropodomorph dinosaur from the Late T ...
''. However, Ballell, Rayfield & Benton (2020) considered ''Pantydraco caducus'' to be a taxon of uncertain validity, and considered it possible that it might represent a juvenile of ''Thecodontosaurus antiquus''.


Description

From the fragmentary remains of ''Thecodontosaurus'', most of the skeleton can be reconstructed, except for the front of the skull. ''Thecodontosaurus'' had a rather short neck supporting a fairly large skull with large eyes. Its jaws contained many small- to medium-sized, serrated, leaf-shaped teeth. This dinosaur's hands and feet each had five digits, and the hands were long and rather narrow, with an extended claw on each. This dinosaur's front limbs were much shorter than the legs, and its tail was much longer than the head, neck and body put together. On average, it was long, or 1 ft. tall, and weighed . The largest individuals had an estimated length of . In 2000,
Michael Benton Michael James Benton (born 8 April 1956) is a British palaeontologist, and professor of vertebrate paleontology, vertebrate palaeontology in the School of Earth Sciences at the University of Bristol. His published work has mostly concentrated on ...
noted the existence of a robust morph in the population, seen by him as a possible second species or, more likely, an instance of
sexual dimorphism Sexual dimorphism is the condition where sexes of the same species exhibit different Morphology (biology), morphological characteristics, including characteristics not directly involved in reproduction. The condition occurs in most dioecy, di ...
. Benton also indicated some unique derived traits, or
autapomorphies In phylogenetics, an autapomorphy is a distinctive feature, known as a derived trait, that is unique to a given taxon. That is, it is found only in one taxon, but not found in any others or outgroup taxa, not even those most closely related to ...
, for the species: a long basipterygoid process on the
braincase In human anatomy, the neurocranium, also known as the braincase, brainpan, brain-pan, or brainbox, is the upper and back part of the skull, which forms a protective case around the brain. In the human skull, the neurocranium includes the calv ...
; a dentary that is short in relation to the total length of the lower jaw; an ilium that has a back end that is subquadrate instead of rounded.M.J. Benton, L. Juul, G.W. Storrs and P.M. Galton, 2000, "Anatomy and systematics of the prosauropod dinosaur ''Thecodontosaurus antiquus'' from the upper Triassic of southwest England", ''Journal of Vertebrate Paleontology'' 20(1): 77-108 The small size has been explained as an instance of
insular dwarfism Insular dwarfism, a form of phyletic dwarfism, is the process and condition of large animals evolving or having a reduced body size when their population's range is limited to a small environment, primarily islands. This natural process is disti ...
.


Classification

Riley and Stutchbury originally saw ''Thecodontosaurus'' as a member of the
Squamata Squamata (, Latin ''squamatus'', 'scaly, having scales') is the largest Order (biology), order of reptiles; most members of which are commonly known as Lizard, lizards, with the group also including Snake, snakes. With over 11,991 species, it i ...
, the group containing lizards and snakes. This did not change when
Richard Owen Sir Richard Owen (20 July 1804 – 18 December 1892) was an English biologist, comparative anatomy, comparative anatomist and paleontology, palaeontologist. Owen is generally considered to have been an outstanding naturalist with a remarkabl ...
coined the term Dinosauria in 1842, because Owen did not recognise ''Thecodontosaurus'' as a dinosaur; in 1865, he assigned it to the
Thecodontia Thecodontia (meaning 'socket-teeth'), now considered an obsolete taxonomic grouping, was formerly used to describe a diverse "order" of early archosaurian reptiles that first appeared in the latest Permian period and flourished until the end of t ...
. It was not until 1870 that
Thomas 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 ...
became the first person to understand that it was a dinosaur, though referring it incorrectly to the
Scelidosauridae ''Scelidosaurus'' (; with the intended meaning of "limb lizard", from Greek / meaning 'rib of beef' and ''sauros''/ meaning 'lizard')Liddell & Scott (1980). Greek-English Lexicon, Abridged Edition. Oxford University Press, Oxford, UK. is a gen ...
. Later, it was placed in either the
Anchisauridae ''Anchisaurus'' is a genus of basal sauropodomorph dinosaur. It lived during the Early Jurassic Period, and its fossils have been found in the red sandstone of the Upper Portland Formation, Northeastern United States, which was deposited from ...
or its own
Thecodontosauridae Thecodontosauridae is a family of basal sauropodomorph dinosaurs that are part of the Bagualosauria, known from fossil remains found exclusively in the Magnesian Conglomerate of Bristol, England,Agrosaurus''. Modern exact
cladistic Cladistics ( ; from Ancient Greek 'branch') is an approach to biological classification in which organisms are categorized in groups ("clades") based on hypotheses of most recent common ancestry. The evidence for hypothesized relationships is ...
analyses have not been conclusive. Although not actually the earliest member of the group, ''Thecodontosaurus'' is sometimes placed in a very basal position among the
sauropodomorph Sauropodomorpha ( ; from Greek, meaning "lizard-footed forms") is an extinct clade of long-necked, herbivorous, saurischian dinosaurs that includes the sauropods and their ancestral relatives. Sauropods generally grew to very large sizes, had lo ...
dinosaurs. It was earlier included under the
Prosauropoda Sauropodomorpha ( ; from Greek, meaning "lizard-footed forms") is an extinct clade of long-necked, herbivorous, saurischian dinosaurs that includes the sauropods and their ancestral relatives. Sauropods generally grew to very large sizes, had lo ...
, but more recently it has been suggested that ''Thecodontosaurus'' and its relatives preceded the prosauropod-sauropod split.Yates, A.M. & Kitching, J. W. (2003). "The earliest known sauropod dinosaur and the first steps towards sauropod locomotion". '' Proc. R. Soc. Lond.: B Biol Sci.'' 2003 Aug 22; 270(1525): 1753–8


Paleobiology

Examination of ''Thecodontosaurus'' revealed it was exclusively bipedal. Studies of the muscle attachments in its fore and hindlimbs suggest that it was an extremely fast bipedal runner that relied on its weaker front limbs for grasping vegetation, cutting it up and feeding it into its mouth. Its advanced running capabilities suggest it was well adapted for high-speed sprinting, probably as a means of escaping predators.


References


Further reading

* Moody, Richard. Dinofile. Pg. 23. Octopus Publishing Group Ltd., 2006 {{Taxonbar, from=Q131300 Sauropodomorpha Dinosaur genera Carnian dinosaurs Taxa named by Samuel Stutchbury Fossil taxa described in 1836 Dinosaurs of the United Kingdom