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''Mantellisaurus'' 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 n ...
of iguanodontian
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 ...
that lived in the
Barremian The Barremian is an age in the geologic timescale (or a chronostratigraphic stage) between 129.4 ± 1.5 Ma ( million years ago) and 121.4 ± 1.0 Ma). It is a subdivision of the Early Cretaceous Epoch (or Lower Cretaceous Series). It is pre ...
and early
Aptian The Aptian is an age in the geologic timescale or a stage in the stratigraphic column. It is a subdivision of the Early or Lower Cretaceous Epoch or Series and encompasses the time from 121.4 ± 1.0 Ma to 113.0 ± 1.0 Ma (million years ag ...
ages of 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 ...
Period of
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 ...
. Its remains are known from
Belgium Belgium, ; french: Belgique ; german: Belgien officially the Kingdom of Belgium, is a country in Northwestern Europe. The country is bordered by the Netherlands to the north, Germany to the east, Luxembourg to the southeast, France to ...
( Bernissart),
England England is a country that is part of the United Kingdom. It shares land borders with Wales to its west and Scotland to its north. The Irish Sea lies northwest and the Celtic Sea to the southwest. It is separated from continental Europe ...
,
Spain , image_flag = Bandera de España.svg , image_coat = Escudo de España (mazonado).svg , national_motto = '' Plus ultra'' ( Latin)(English: "Further Beyond") , national_anthem = (English: "Royal March") , ...
and
Germany Germany, officially the Federal Republic of Germany (FRG),, is a country in Central Europe. It is the most populous member state of the European Union. Germany lies between the Baltic and North Sea to the north and the Alps to the sou ...
. The
type Type may refer to: Science and technology Computing * Typing, producing text via a keyboard, typewriter, etc. * Data type, collection of values used for computations. * File type * TYPE (DOS command), a command to display contents of a file. * Ty ...
and only species is ''M. atherfieldensis''. Formerly known as '' Iguanodon atherfieldensis'', the new genus ''Mantellisaurus'' was erected for the species by Gregory Paul in 2007. According to Paul, ''Mantellisaurus'' was more lightly built than ''Iguanodon'' and more closely related to ''
Ouranosaurus ''Ouranosaurus'' is a genus of herbivorous basal hadrosauriform dinosaur that lived during the Aptian stage of the Early Cretaceous of modern-day Niger and Cameroon. ''Ouranosaurus'' measured about long. Two rather complete fossils were found i ...
'', making ''Iguanodon'' in its traditional sense
paraphyletic In taxonomy (general), taxonomy, a group is paraphyletic if it consists of the group's most recent common ancestor, last common ancestor and most of its descendants, excluding a few Monophyly, monophyletic subgroups. The group is said to be pa ...
. It is known from many complete and almost complete skeletons. The genus name honours Gideon Mantell, the discoverer of ''Iguanodon''.


History of discovery


Discovery and naming of the holotype

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 ...
fossil, NHMUK R5764, was originally discovered by
Reginald Walter Hooley Reginald Walter Hooley (5 September 1865 – 5 May 1923) was a businessman and amateur paleontologist, collecting on the Isle of Wight. He is probably best remembered for describing the dinosaur ''Iguanodon atherfieldensis'', now ''Mantellisaurus ...
in 1914 in the upper Vectis Formation of southern England and reported upon in 1917. He posthumously named it ''Iguanodon atherfieldensis'' in 1925.
Atherfield Atherfield is a rural location in the south west of the Isle of Wight, UK. It includes the small settlements of Atherfield Green and Little Atherfield, as well as several farms, and is set in largely open farmland. To the south west it is boun ...
is the name of a village on the southwest shore of 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 ...
where the fossil was found.


The Maidstone specimen and "''Mantellodon''"

The Maidstone specimen was discovered in a quarry in
Maidstone Maidstone is the largest Town status in the United Kingdom, town in Kent, England, of which it is the county town. Maidstone is historically important and lies 32 miles (51 km) east-south-east of London. The River Medway runs through the c ...
,
Kent Kent is a county in South East England and one of the home counties. It borders Greater London to the north-west, Surrey to the west and East Sussex to the south-west, and Essex to the north across the estuary of the River Thames; it faces ...
, owned by William Harding Bensted, in February 1834 (lower Lower Greensand Formation). In June 1834 it was acquired for £25 by scientist Gideon Mantell. He was led to identify it as an '' Iguanodon'' based on its distinctive teeth. The Maidstone slab was utilized in the first skeletal reconstructions and artistic renderings of ''Iguanodon'', but due to its incompleteness, Mantell made some mistakes, the most famous of which was the placement of what he thought was a horn on the nose. Shortly after the discovery, tension began to build between Mantell and
Richard Owen Sir Richard Owen (20 July 1804 – 18 December 1892) was an English biologist, comparative anatomist and paleontologist. Owen is generally considered to have been an outstanding naturalist with a remarkable gift for interpreting fossils. Ow ...
, an ambitious scientist with much better funding and society connections in the turbulent worlds of Reform Act–era British politics and science. Owen, at the time a firm
creationist Creationism is the religious belief that nature, and aspects such as the universe, Earth, life, and humans, originated with supernatural acts of divine creation. Gunn 2004, p. 9, "The ''Concise Oxford Dictionary'' says that creationism is 'th ...
, opposed the early versions of evolutionary science (" transmutationism") then being debated and used what he would soon coin as dinosaurs as a weapon in this conflict. With the paper describing Dinosauria, he scaled down dinosaurs from lengths of over 61 metres (200 ft), determined that they were not simply giant lizards, and put forward that they were advanced and mammal-like, characteristics given to them by
God In monotheistic thought, God is usually viewed as the supreme being, creator, and principal object of faith. Swinburne, R.G. "God" in Honderich, Ted. (ed)''The Oxford Companion to Philosophy'', Oxford University Press, 1995. God is typically ...
; according to the understanding of the time, they could not have been "transmuted" from reptiles to mammal-like creatures.Torrens, Hugh. "Politics and Paleontology". ''The Complete Dinosaur,'' 175–190. In 1849, a few years before his death in 1852, Mantell realised that the genus today known as ''Mantellodon'' was not a heavy,
pachyderm Pachyderm may refer to: * Any of the Pachydermata, an obsolete 19th-century taxonomic order of mammals that included elephants, rhinoceroses, tapirs and hippopotami. * Pachyderm Studios, a recording studio in Cannon Falls, Minnesota. * Pachyderma ...
-like animal, as Owen was putting forward, but had slender forelimbs; however, his passing left him unable to participate in the creation of the Crystal Palace dinosaur sculptures, and so Owen's vision of the dinosaurs became that seen by the public for decades. With Benjamin Waterhouse Hawkins, Owen had nearly two dozen lifesize
sculpture Sculpture is the branch of the visual arts that operates in three dimensions. Sculpture is the three-dimensional art work which is physically presented in the dimensions of height, width and depth. It is one of the plastic arts. Durable sc ...
s of various prehistoric animals built out of
concrete Concrete is a composite material composed of fine and coarse aggregate bonded together with a fluid cement (cement paste) that hardens (cures) over time. Concrete is the second-most-used substance in the world after water, and is the most ...
sculpted over a steel and brick framework; two ''Mantellodon'', one standing and one resting on its belly, were included. Before the sculpture of the standing ''Mantellodon'' was completed, a banquet for twenty was held inside it.Norman, David B. ''The Illustrated Encyclopedia of Dinosaurs.'' p. 11. The discovery of much better specimens of ''Iguanodon bernissartensis'' in later years revealed that the horn was actually a modified thumb. Still encased in rock, the Maidstone skeleton is currently displayed at the
Natural History Museum A natural history museum or museum of natural history is a scientific institution with natural history collections that include current and historical records of animals, plants, fungi, ecosystems, geology, paleontology, climatology, and more ...
in
London London is the capital and List of urban areas in the United Kingdom, largest city of England and the United Kingdom, with a population of just under 9 million. It stands on the River Thames in south-east England at the head of a estuary dow ...
. The borough of Maidstone commemorated this find by adding an ''Iguanodon'' as a
supporter In heraldry, supporters, sometimes referred to as ''attendants'', are figures or objects usually placed on either side of the shield and depicted holding it up. Early forms of supporters are found in medieval seals. However, unlike the co ...
to their
coat of arms A coat of arms is a heraldic visual design on an escutcheon (i.e., shield), surcoat, or tabard (the latter two being outer garments). The coat of arms on an escutcheon forms the central element of the full heraldic achievement, which in it ...
in 1949. This specimen has become linked with the name ''I. mantelli'', a species named in 1832 by Christian Erich Hermann von Meyer in place of ''I. anglicus'', but it actually comes from a different
formation Formation may refer to: Linguistics * Back-formation, the process of creating a new lexeme by removing or affixes * Word formation, the creation of a new word by adding affixes Mathematics and science * Cave formation or speleothem, a secondary ...
than the original ''I. mantelli''/''I. anglicus'' material. The Maidstone specimen, also known as Gideon Mantell's "Mantel-piece", and formally labelled NHMUK 3741 was subsequently excluded from ''Iguanodon''. It was classified as ''cf.'' ''Mantellisaurus'' by McDonald (2012), as cf. ''Mantellisaurus atherfieldensis'' by David Bruce Norman (2012), and made the holotype of a separate genus and species ''Mantellodon carpenteri'' by Gregory S. Paul (2012). The generic name combines Mantell's name with a Greek ''odon'', "tooth", analogous to ''Iguanodon''. The
specific name Specific name may refer to: * in Database management systems, a system-assigned name that is unique within a particular database In taxonomy, either of these two meanings, each with its own set of rules: * Specific name (botany), the two-part (bino ...
honours Kenneth Carpenter for his work on dinosaurs in general and iguandonts in particular. David Norman, in 2013, considered Paul's description of ''Mantellodon'' to be inadequate, identical to that given by Paul of ''
Darwinsaurus ''Hypselospinus'' is a genus of iguanodontian dinosaur which was first described as a species of '' Iguanodon'' (''I. fittoni'') by Richard Lydekker in 1889, the specific name honouring William Henry Fitton. History In May 2010 the fossils c ...
'' and entirely incorrect, noting that no dentary is preserved in the holotype specimen, and that the preserved forelimb elements "are gracile, carpals are not preserved, the metacarpals are elongate and slender, and a thumb spike is not preserved". Norman considered the holotype specimen of ''Mantellodon carpenteri'' to be referable to the species ''Mantellisaurus atherfieldensis''. In 2021, a sculpture nicknamed Iggy the ''Iguanodon'' based on the Maidstone specimen was unveiled at the
Maidstone East station Maidstone East railway station is one of three stations in the central area of Maidstone, Kent, England, but currently the only one with a regular direct service to London. The station is on the Maidstone line, from , and is served by trains ...
. The dinosaur is also featured in Maidstone's
coat of arms A coat of arms is a heraldic visual design on an escutcheon (i.e., shield), surcoat, or tabard (the latter two being outer garments). The coat of arms on an escutcheon forms the central element of the full heraldic achievement, which in it ...
, and is the only dinosaur featured in a borough's coat of arms in the UK.


Specimen IRSNB 1551 and "''Dollodon''"

Specimen IRSNB 1551 from the Sainte-Barbe Clays,
Belgium Belgium, ; french: Belgique ; german: Belgien officially the Kingdom of Belgium, is a country in Northwestern Europe. The country is bordered by the Netherlands to the north, Germany to the east, Luxembourg to the southeast, France to ...
, became the second mounted skeleton of a non-avian dinosaur made primarily out of actual bone when put on display by Louis Dollo in 1884. This specimen was originally assigned to '' Iguanodon mantelli'' by
George Albert Boulenger George Albert Boulenger (19 October 1858 – 23 November 1937) was a Belgian-British zoologist who described and gave scientific names to over 2,000 new animal species, chiefly fish, reptiles, and amphibians. Boulenger was also an active botan ...
in 1881, but was in 1986 thought to pertain to ''Iguanodon atherfieldensis'' by David Bruce Norman. The specimen was assigned to its own genus and species, ''Dollodon bampingi'', by Gregory S. Paul in 2008. The genus was named after Dollo, who first described the remains, and the
specific name Specific name may refer to: * in Database management systems, a system-assigned name that is unique within a particular database In taxonomy, either of these two meanings, each with its own set of rules: * Specific name (botany), the two-part (bino ...
was in honour of popular science writer Daniel Bamping, who assisted Paul in his investigations. Paul noted several differences between the ''Mantellisaurus'' type (NHMUK R5764) and IRSNB 1551. The ''Mantellisaurus'' type had proportionally shorter forelimbs with a larger pelvis and he argued it was probably more bipedal, whereas IRSNB 1551 was more likely to be quadrupedal. Paul also noted that the
snout A snout is the protruding portion of an animal's face, consisting of its nose, mouth, and jaw. In many animals, the structure is called a muzzle, rostrum, or proboscis. The wet furless surface around the nostrils of the nose of many mammals is ...
and
trunk Trunk may refer to: Biology * Trunk (anatomy), synonym for torso * Trunk (botany), a tree's central superstructure * Trunk of corpus callosum, in neuroanatomy * Elephant trunk, the proboscis of an elephant Computing * Trunk (software), in rev ...
of IRSNB 1551 were proportionally longer than the ''Mantellisaurus'' type specimen. The validity of ''Dollodon'' has since been disputed. In 2010, Kenneth Carpenter and Yusuke Ishida synonymized ''Dollodon bampingi'' with ''
Iguanodon seelyi ''Iguanodon'' ( ; meaning 'iguana-tooth'), named in 1825, is a genus of iguanodontian dinosaur. While many species have been classified in the genus ''Iguanodon'', dating from the late Jurassic Period to the early Cretaceous Period of Asia, ...
'', a species based on BMNH R 28685 from Wessex Formation,
England England is a country that is part of the United Kingdom. It shares land borders with Wales to its west and Scotland to its north. The Irish Sea lies northwest and the Celtic Sea to the southwest. It is separated from continental Europe ...
. Likewise, David B. Norman and Andrew McDonald do not consider ''Dollodon'' a valid
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 ...
or species and instead include it with ''Mantellisaurus''.


Sauerland specimens

In 1971, a fossiliferous karstic
sinkhole A sinkhole is a depression or hole in the ground caused by some form of collapse of the surface layer. The term is sometimes used to refer to doline, enclosed depressions that are locally also known as ''vrtače'' and shakeholes, and to openi ...
clay deposit was found at a quarry just south-west of the village of Nehden near Brilon in
Sauerland The Sauerland () is a rural, hilly area spreading across most of the south-eastern part of North Rhine-Westphalia, in parts heavily forested and, apart from the major valleys, sparsely inhabited. The Sauerland is the largest tourist region in ...
, Germany containing numerous disarticulated iguanodontid remains predominantly of ''Mantellisaurus'' with lesser quantities of '' Iguanodon'', alongside other fragmentary dinosaur and crocodylian material.


Iberian specimens

''Mantellisaurus'' is known from several localities in Spain. With an articulated hindlimb known from Las Hoyas A specimen is known from the Rubielos de Mora 1 locality in Spain. Three specimens are known from the
Arcillas de Morella Formation The Arcillas de Morella Formation is a geological formation in Spain whose strata date back to the Barremian stage of the Early Cretaceous. Dinosaur remains are among the fossils that have been recovered from the formation.Weishampel, David B; et a ...
.


Description

''Mantellisaurus'' was a lightly constructed iguanodont. Compared to ''Iguanodon bernissartensis'', ''Mantellisaurus '' was smaller, estimated at in weight. Its forelimbs were proportionally shorter than those of ''I. bernissartensis''. In ''Mantellisaurus '' the forelimbs were about half the length of the hindlimbs whereas they were about 70 percent the length of the hindlimbs in ''I. bernissartensis''. Due to the short length of its forelimbs and the shortness of its body, Paul proposed that it was primarily bipedal, only going on all fours when standing still or moving slowly.


Classification

The
cladogram A cladogram (from Greek ''clados'' "branch" and ''gramma'' "character") is a diagram used in cladistics to show relations among organisms. A cladogram is not, however, an evolutionary tree because it does not show how ancestors are related to ...
below follows an analysis by Andrew McDonald, 2012.


References


Literature

* Cornuel, M., 1850, Note sur des ossements fossiles decouvertes dans le calcaire neocomien de Wassy (Haute-Marne): Bulletin de la societie geologiques de France, 2nd series, v. 7, p. 702-704. * Hooley, W., 1925, On the skeleton of Iguanodon atherfieldensis sp. nov., from the Wealden Shales of Atherfield (Isle of Wight): Quarterly Journal of the Geological Society of London, v. 81, p. 1-61. * Hulke, J. W., 1879, Vectisaurus valdensis, a new Wealden Dinosaur: Quarterly Journal of the Geological Society of London, v. 35, p. 421-424. * Owen, R., 1842, Report on British Fossil Reptiles. Part II: Report of the British Association for the Advancement of Science, v. 11, p. 60-204. * Lydekker, R., 1888, Catalogue of the Fossil Reptilia and Amphibia in the British Museum (Natural History), Cromwell Road, S.W., Part 1. Containing the Orders Ornithosauria, Crocodilia, Dinosauria, Squamta, Rhynchocephalia, and Proterosauria: British Museum of Natural History, London, 309pp. * Norman, D.B., 2012. "Iguanodontian Taxa (Dinosauria: Ornithischia) from the Lower Cretaceous of England and Belgium". In: Pascal Godefroit (ed.), Bernissart Dinosaurs and Early Cretaceous Terrestrial Ecosystems. Indiana University Press. 464 pp. http://www.iupress.indiana.edu/product_info.php?products_id=800408 * Paul, G.S. 2007. Turning the old into the new: a separate genus for the gracile iguanodont from the Wealden of England; pp. 69–77 in K. Carpenter (ed.), ''Horns and Beaks: Ceratopsian and Ornithopod Dinosaurs''. Indiana University Press, Bloomington. {{Portal bar, Paleontology, Dinosaurs, Cretaceous Iguanodonts Early Cretaceous dinosaurs of Europe Cretaceous Spain Fossils of Spain La Huérguina Formation Fossil taxa described in 2007 Ornithischian genera Taxa named by Gregory S. Paul