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''Erectopus'' (meaning "upright foot") is an extinct genus of basal allosauroid
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 ...
from 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 Pr ...
La Penthiève Beds Formation of
France France (), officially the French Republic ( ), is a country primarily located in Western Europe. It also comprises of overseas regions and territories in the Americas and the Atlantic, Pacific and Indian Oceans. Its metropolitan ar ...
and also possibly the Cernavoda Formation of southern
Romania Romania ( ; ro, România ) is a country located at the crossroads of Central, Eastern, and Southeastern Europe. It borders Bulgaria to the south, Ukraine to the north, Hungary to the west, Serbia to the southwest, Moldova to the east, a ...
. The type species is ''E. superbus'', which was initially known as a species of '' Megalosaurus''.


Discovery and naming

The holotype, specimen MNHN 2001-4, was discovered between the early 1870s and before 1882 in the Phosphate-bearing beds of La Penthiève (''Mammilatum'' Zone; lower Albian) at
Louppy-le-Château Louppy-le-Château () is a commune in the Meuse department in Grand Est in north-eastern France France (), officially the French Republic ( ), is a country primarily located in Western Europe. It also comprises of Overseas France, ov ...
in eastern France,Allain, R. (2005). The enigmatic theropod dinosaur ''Erectopus superbus'' (Sauvage, 1882) from the Lower Albian of Louppy-le-Château (Meuse, France). in Carpenter, K. 2005. ''The Carnivorous Dinosaurs'', Indiana University Press: 72-86. which have also produced remains of plesiosaurs, ichthyosaurs, and
crocodile Crocodiles (family Crocodylidae) or true crocodiles are large semiaquatic reptiles that live throughout the tropics in Africa, Asia, the Americas and Australia. The term crocodile is sometimes used even more loosely to include all extant ...
s. The fossils originally resided in the private collection of Louis Pierson. The first, two teeth and a vertebra, were first described by
Charles Barrois Charles Eugene Barrois (21 August 18515 November 1939) was a French geologist and palaeontologist. Life Barrois was born at Lille and educated at the Jesuit College of St Joseph in that town, where he studied geology under Professor Jules ...
in 1875. After more remains had been found, in 1882 Henri-Émile Sauvage made them the basis for a new
taxon In biology, a taxon ( back-formation from '' taxonomy''; plural taxa) is a group of one or more populations of an organism or organisms seen by taxonomists to form a unit. Although neither is required, a taxon is usually known by a particular n ...
, '' Megalosaurus superbus.''Sauvage, H.-É., (1882), "Recherches sur les reptiles trouvés dans le Gault de l'est du bassin de Paris", ''Mémoires de la Société Géologique de France, série 3'' 2(4): 1-42 In 1923, the material was redescribed by Friedrich von Huene, who argued that it could not be included within the
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 n ...
''Megalosaurus'' and created for the Pierson theropod a separate genus, naming the species ''Erectopus superbus''.Huene, F. von, (1923), "Carnivorous Saurischia in Europe since the Triassic",  ''Bulletin of the Geological Society of America''34: 449-458 In 1932 von Huene concluded that the original fossils described by Barrois were not necessarily of the same species as the later finds.Huene, F. von, (1932). Die fossile Reptil-Ordnung Saurischia, ihre Entwicklung und Geschichte. ''Monographien zur Geologie und Palaeontologie'', ser. 1: 1-361. Assuming that Sauvage had used the former as the
holotype A holotype is a single physical example (or illustration) of an organism, known to have been used when the species (or lower-ranked taxon) was formally described. It is either the single such physical example (or illustration) or one of seve ...
of ''Megalosaurus superbus'', he therefore created another species: ''Erectopus sauvagei''. Von Huene even declined to use the generic name ''Erectopus'' for the first species, indicating it as "Gen. indeterm. ''superbus''", which however does not constitute a valid name. Subsequently, the Pierson collection was dispersed after the death of its owner and the holotype was long believed lost to science after
World War II World War II or the Second World War, often abbreviated as WWII or WW2, was a world war that lasted from 1939 to 1945. It involved the World War II by country, vast majority of the world's countries—including all of the great power ...
. However, casts of some of the bones have been located in the Muséum National d'Histoire Naturelle (MNHN,
Paris Paris () is the capital and most populous city of France, with an estimated population of 2,165,423 residents in 2019 in an area of more than 105 km² (41 sq mi), making it the 30th most densely populated city in the world in 2020. ...
), and the anterior part of a left
maxilla The maxilla (plural: ''maxillae'' ) in vertebrates is the upper fixed (not fixed in Neopterygii) bone of the jaw formed from the fusion of two maxillary bones. In humans, the upper jaw includes the hard palate in the front of the mouth. The ...
, described by Sauvage in 1882, was found through a Parisian fossil dealer in the late 20th century and purchased by Christian de Muison, a
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 foss ...
at the MNHN. The casts and the incomplete maxilla allowed for a reevaluation of ''Erectopus'' by Ronan Allain in 2005, which determined that the correct taxonomic name for the material is ''E. superbus'', as nothing indicated that Sauvage had limited the holotype to the original teeth. The recovered maxilla has been designated a lectotype, and the casts of Pierson's specimen has become the plastotype for the taxon. The plastotype includes a partial right manus, the left femur, left calcaneum, the
proximal Standard anatomical terms of location are used to unambiguously describe the anatomy of animals, including humans. The terms, typically derived from Latin or Greek roots, describe something in its standard anatomical position. This position ...
and distal halves of the left tibia, and right metatarsal II. The combined inventory number for the syntypes is MNHN 2001-4. A tooth, specimen UAIC (SCM1) 615, discovered sometime between 1900 and 1913 in the Cernavoda Formation in Romania was listed as belonging to ''Megalosaurus'' (''Erectopus'') ''cf. superbus'' by Simonescu (1913) and later by Csiki-Sava ''et al.'' (2016), although it most likely belonged to Carcharodontosauridae.


Description

Allain (2005) diagnoses ''Erectopus superbus'' as follows: "Rounded anterior ramus of
maxilla The maxilla (plural: ''maxillae'' ) in vertebrates is the upper fixed (not fixed in Neopterygii) bone of the jaw formed from the fusion of two maxillary bones. In humans, the upper jaw includes the hard palate in the front of the mouth. The ...
; slender neck of femur; posterior curvature of proximal half of femur; anterodorsal edge of calcaneum dorsally projected; calcaneum twice as long as deep vertically; posteromedial process for tibia on articular surface of astragalus; length of second metatarsal equal to half the length of femur; lateral margin of proximal end of second metatarsal regularly concave." Allain estimated the weight at . In 2016 it was estimated to be 5 meters (16 ft) in length and 315 kg (694 lbs) in weight.


Classification

Based on the morphology of the distal end of the tibia and the inferred morphology of the astragalus, Allain (2005) placed ''Erectopus superbus'' within the Allosauroidea (a concept equivalent to Carnosauria, a term some researchers prefer). It is the third youngest carnosaur known from the
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 subcontinent of Eurasia and it is located enti ...
an Lower Cretaceous, after specimen MM-2-21, the "Montmirat theropod" ( Valanginian) of southern France and ''
Neovenator ''Neovenator'' (nee-o-ven-a-tor meaning "new hunter") is a genus of carcharodontosaurian theropod dinosaur. It is known from several skeletons found in the Early Cretaceous ( Barremian~130-125 million years ago) Wessex Formation on the sout ...
salerii'' ( Barremian) from the
Isle of Wight The Isle of Wight ( ) is a Counties of England, county in the English Channel, off the coast of Hampshire, from which it is separated by the Solent. It is the List of islands of England#Largest islands, largest and List of islands of England#Mo ...
. Carrano ''et al.'' (2012) commented that ''Erectopus'' may be a metricanthosaurid, but didn't list shared characters.


References

* {{Taxonbar, from=Q2342517 Carnosaurs Early Cretaceous dinosaurs of Europe Fossil taxa described in 1923 Taxa named by Friedrich von Huene