Trimucrodon
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''Trimucrodon'' is a
genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of living and fossil organisms as well as viruses. In the hierarchy of biological classification, genus comes above species and below family. In binomial nom ...
of ornithischian
dinosaur Dinosaurs are a diverse group of reptiles of the clade Dinosauria. They first appeared during the Triassic period, between 243 and 233.23 million years ago (mya), although the exact origin and timing of the evolution of dinosaurs is t ...
from the
Late Jurassic The Late Jurassic is the third epoch of the Jurassic Period, and it spans the geologic time from 163.5 ± 1.0 to 145.0 ± 0.8 million years ago (Ma), which is preserved in Upper Jurassic strata.Owen 1987. In European lithostratigraphy, the name ...
of
Portugal Portugal, officially the Portuguese Republic ( pt, República Portuguesa, links=yes ), is a country whose mainland is located on the Iberian Peninsula of Southwestern Europe, and whose territory also includes the Atlantic archipelagos of ...
. Three isolated teeth found at the Porto Dinheiro (or Pinhiero) locality in the Lisboa District of Portugal were given the name in
1973 Events January * January 1 - The United Kingdom, the Republic of Ireland and Denmark 1973 enlargement of the European Communities, enter the European Economic Community, which later becomes the European Union. * January 15 – Vietnam War: ...
by Richard A. Thulborn, derived from the
Latin Latin (, or , ) is a classical language belonging to the Italic branch of the Indo-European languages. Latin was originally a dialect spoken in the lower Tiber area (then known as Latium) around present-day Rome, but through the power of the ...
words for "three" and a dagger point, and , and the
Ancient Greek Ancient Greek includes the forms of the Greek language used in ancient Greece and the ancient world from around 1500 BC to 300 BC. It is often roughly divided into the following periods: Mycenaean Greek (), Dark Ages (), the Archaic p ...
word for "tooth". The only species in the taxon is ''Trimucrodon cuneatus'', taken from the wedge shape of the teeth. Though the unit the specimens came from was originally unnamed, it was referred to the Alcobaça, and then
Lourinhã Formation The Lourinhã Formation () is a fossil rich geological formation in western Portugal, named for the municipality of Lourinhã. The formation is mostly Late Jurassic in age (Kimmeridgian/Tithonian), with the top of the formation extending into the ...
s, specifically the late
Kimmeridgian In the geologic timescale, the Kimmeridgian is an age in the Late Jurassic Epoch and a stage in the Upper Jurassic Series. It spans the time between 157.3 ± 1.0 Ma and 152.1 ± 0.9 Ma (million years ago). The Kimmeridgian follows the Oxford ...
Amoreira–Porto Novo Member. The type specimen, uncovered between 1962 and 1967 by German
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 kingdom, including the structure, embryology, evolution, classification, habits, and d ...
and
paleontologist Paleontology (), also spelled palaeontology or palæontology, is the scientific study of life that existed prior to, and sometimes including, the start of the Holocene epoch (roughly 11,700 years before present). It includes the study of fossi ...
Georg Krusat, is distinguished by prominent denticles at the front and rear ends of the crown, and comes from an individual under long.G. Krusat, 1969, "Ein Panthotheria-Molar mit dreispitzigem Talonid aus dem Kimmeridge von Portugal", ''Paläontologische Zeitschrift'' 43(1/2): 52-56 It is currently kept in the collections of the Museu Geológico do Instituto Geológico e Mineiro in Lisbon, Portugal, formerly having been kept in the collections of the
Free University of Berlin The Free University of Berlin (, often abbreviated as FU Berlin or simply FU) is a public research university in Berlin, Germany. It is consistently ranked among Germany's best universities, with particular strengths in political science and t ...
. The holotype tooth of ''Trimucrodon'' is wide, similar to some of the referred specimens, with the crown about as tall as wide. Other specimens have crowns as small as wide and only half as tall as wide. The crowns are of varying amounts of asymmetry, though each edge bears the same number of tapering denticles.
Tooth enamel Tooth enamel is one of the four major tissues that make up the tooth in humans and many other animals, including some species of fish. It makes up the normally visible part of the tooth, covering the crown. The other major tissues are dentin, ...
is present of an equal thickness on both sides of the tooth crown, and both faces are smooth without ridges. The denticles increase in size away from the apex of the crown, ending with two widely divergent and sharply-pointed denticles at the front and rear ends of the base. ''Trimucrodon'' was originally referred to as a member of the
ornithopod Ornithopoda () is a clade of ornithischian dinosaurs, called ornithopods (), that started out as small, bipedal running grazers and grew in size and numbers until they became one of the most successful groups of herbivores in the Cretaceous wo ...
family
Fabrosauridae Ornithischia () is an extinct order of mainly herbivorous dinosaurs characterized by a pelvic structure superficially similar to that of birds. The name ''Ornithischia'', or "bird-hipped", reflects this similarity and is derived from the Greek ...
by Thulborn in 1973, closest to ''
Echinodon ''Echinodon'' is a genus of heterodontosaurid dinosaur that lived during the earliest Cretaceous of southern England and possibly western France in the Berriasian epoch. The first specimens were jaw bones named ''Echinodon becklesii'' by Sir Ric ...
'' but also related to ''
Alocodon ''Alocodon'' is a genus of ornithischian dinosaur known from multiple teeth from the Middle or Late Jurassic. The taxon was first described in 1973 by Richard A. Thulborn for an assemblage of teeth from the Pedrógão locality of Portugal, dist ...
'' and ''
Fabrosaurus ''Fabrosaurus'' ( ) is an extinct genus of ornithischian dinosaur that lived during the Early Jurassic during the Hettangian to Sinemurian stages 199 - 189 mya.Ginsburg, L., (1964), "Decouverte d’un Scelidosaurien (Dinosaure ornithischien) ...
''. Peter M. Galton retained it in the family in 1978, though he noted that there were significant differences between ''Trimucrodon'' and ''Echinodon'', and the Middle to Late Jurassic fabrosaurids ''Alocodon'', ''Trimucrodon'' and ''Echinodon'' were representative of three independent and only distantly related branches of the family, with ''
Nanosaurus ''Nanosaurus'' ("small or dwarf lizard") is the name given to a genus of neornithischian dinosaur that lived about 155 to 148 million years ago, during the Late Jurassic-age. Its fossils are known from the Morrison Formation of the south-weste ...
'' not preserving enough material to determine its relationships. Given that the species was only represented by teeth, ''Trimucrodon'' was designated as a '' nomen dubium'' in
1990 File:1990 Events Collage.png, From left, clockwise: The 1990 FIFA World Cup is played in Italy; The Human Genome Project is launched; Voyager I takes the famous Pale Blue Dot image- speaking on the fragility of humanity on Earth, astrophysicist ...
by David B. Weishampel and Lawrence M. Witmer, as an indeterminate member of Ornithischia outside Ornithopoda. While a basal ornithischian position outside Ornithopoda was retained by
Paul Sereno Paul Callistus Sereno (born October 11, 1957) is a professor of paleontology at the University of Chicago and a National Geographic "explorer-in-residence" who has discovered several new dinosaur species on several continents, including at sites ...
in
1991 File:1991 Events Collage.png, From left, clockwise: Boris Yeltsin, elected as Russia's first president, waves the new flag of Russia after the 1991 Soviet coup d'état attempt, orchestrated by Soviet hardliners; Mount Pinatubo erupts in the Phi ...
, it was considered a possibly valid taxon based on its prominent anterior and posteriormost denticles. Galton revised his classification of the taxon in
1994 File:1994 Events Collage.png, From left, clockwise: The 1994 Winter Olympics are held in Lillehammer, Norway; The Kaiser Permanente building after the 1994 Northridge earthquake; A model of the MS Estonia, which sank in the Baltic Sea; Nelson ...
, considering it only referrable to Ornithopoda, and not closely related to ''Echinodon''. ''Trimucrodon'' was then compared favourable by Galton in 1996 to the also Portuguese taxon '' Taveirosaurus'', previously considered a
pachycephalosaur Pachycephalosauria (; from Greek παχυκεφαλόσαυρος for 'thick headed lizards') is a clade of ornithischian dinosaurs. Along with Ceratopsia, it makes up the clade Marginocephalia. With the exception of two species, most pachyc ...
. With the later reclassifications of ''Taveirosaurus'' as more similar to
nodosaurid Nodosauridae is a family of ankylosaurian dinosaurs, from the Late Jurassic to the Late Cretaceous period in what is now North America, South America, Europe, and Asia. Description Nodosaurids, like their close relatives the ankylosaurids, we ...
s, ''Trimucrodon'' was identified by José Ruiz-Omeñaca in
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to either be a member of
Heterodontosauridae Heterodontosauridae is a family of ornithischian dinosaurs that were likely among the most basal (primitive) members of the group. Their phylogenetic placement is uncertain but they are most commonly found to be primitive, outside of the group ...
alongside ''Echinodon'', or a member of Nodosauridae related to ''Taveirosaurus''. While the identity as a possible heterodontosaurid was upheld by Ruiz-Omeñaca and José Canudo in 2004, in the same year Weishampel, Witmer and colleague David B. Norman followed their 1990 opinion on ''Trimucrodon'', placing it as a dubious ornithischian, though they noted that further study could potentially support the validity of the taxon. ''Trimucrodon'' was found at the top of a cliff south of Porto Dinheiro, in the lower beds of the Lournihã Formation alongside a tooth referred to the ornithischian ''
Hypsilophodon ''Hypsilophodon'' (; meaning "''Hypsilophus''-tooth") is a neornithischian dinosaur genus from the Early Cretaceous period of England. It has traditionally been considered an early member of the group Ornithopoda, but recent research has put thi ...
'', teeth from
rhamphorhynchoid The Rhamphorhynchoidea forms one of the two suborders of pterosaurs and represents an evolutionary grade of primitive members of flying reptiles. This suborder is paraphyletic unlike the Pterodactyloidea, which arose from within the Rhamphorhync ...
pterosaurs, the
crocodylomorphs Crocodylomorpha is a group of pseudosuchian archosaurs that includes the crocodilians and their extinct relatives. They were the only members of Pseudosuchia to survive the end-Triassic extinction. During Mesozoic and early Cenozoic times, cr ...
'' Lusitanisuchus'' and ''
Goniopholis ''Goniopholis'' (meaning "angled scale") is an extinct genus of goniopholidid crocodyliform that lived in Europe and Africa during the Late Jurassic and Early Cretaceous. Being semi-aquatic it is very similar to modern crocodiles. It ranged from ...
'', the choristoderan ''
Cteniogenys ''Cteniogenys'' is a genus of choristodere, a morphologically diverse group of aquatic reptiles. It is part of the monotypic family Cteniogenidae. The type and only named species, ''C. antiquus'', was named in 1928 by Charles W. Gilmore. The ho ...
'', the lizard '' Saurillus'', and over 800 teeth of various groups of mammals. Other deposits from the Amoreira–Porto Novo Member at Porto Dinheiro have contained material from the sauropods ''
Zby ''Zby'' is an extinct genus of turiasaurian sauropod dinosaur known from the Late Jurassic (late Kimmeridgian stage) of the Lourinhã Formation, central west Portugal. It contains a single species, ''Zby atlanticus''. It is named after Georges Z ...
'' and ''
Dinheirosaurus ''Dinheirosaurus'' is a genus of diplodocid sauropod dinosaur that is known from fossils uncovered in modern-day Portugal. It may represent a species of ''Supersaurus''. The only species is ''Dinheirosaurus lourinhanensis'', first described by J ...
'', the
theropod Theropoda (; ), whose members are known as theropods, is a dinosaur clade that is characterized by hollow bones and three toes and claws on each limb. Theropods are generally classed as a group of saurischian dinosaurs. They were ancestrally c ...
s '' Lourinhanosaurus'', ''
Torvosaurus ''Torvosaurus'' () is a genus of carnivorous megalosaurid theropod dinosaur that lived approximately 165 to 148 million years ago during the late Middle and Late Jurassic period (Callovian to Tithonian stages) in what is now Colorado, Portuga ...
'' and ''
Ceratosaurus ''Ceratosaurus'' (from Greek κέρας/κέρατος, ' meaning "horn" and σαῦρος ' meaning "lizard") was a carnivorous theropod dinosaur in the Late Jurassic period ( Kimmeridgian to Tithonian). The genus was first described in 1 ...
'', ornithischian remains from the stegosaur '' Miragaia'' and an intermediate member of
Iguanodontia Iguanodontia (the iguanodonts) is a clade of herbivorous dinosaurs that lived from the Middle Jurassic to Late Cretaceous. Some members include ''Camptosaurus'', ''Dryosaurus'', ''Iguanodon'', '' Tenontosaurus'', and the hadrosaurids or "duck-bil ...
, a possible pleurosternid
turtle Turtles are an order of reptiles known as Testudines, characterized by a special shell developed mainly from their ribs. Modern turtles are divided into two major groups, the Pleurodira (side necked turtles) and Cryptodira (hidden necked t ...
, and scales from the fish ''
Lepidotes ''Lepidotes'' (from el, λεπιδωτός , 'covered with scales') (previously known as ''Lepidotus'') is an extinct genus of Mesozoic ray-finned fish. It has been considered a wastebasket taxon, characterised by "general features, such as thi ...
''.


References

Ornithischian genera Kimmeridgian life Late Jurassic dinosaurs of Europe Jurassic Portugal Fossils of Portugal Fossil taxa described in 1975 Nomina dubia {{ornithischian-stub