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''Mantellisaurus'' 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
iguanodontia Ornithopoda () is a clade of ornithischian dinosaurs, called ornithopods (). They represent one of the most successful groups of herbivorous dinosaurs during the Cretaceous. The most primitive members of the group were bipedal and relatively sm ...
n
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 in the
Barremian The Barremian is an age in the geologic timescale (or a chronostratigraphic stage) between 125.77 Ma (million years ago) and 121.4 ± 1.0 Ma (Historically, this stage was placed at 129.4 million to approximately 125 million years ago) It is a ...
and early
Aptian The Aptian is an age (geology), age in the geologic timescale or a stage (stratigraphy), stage in the stratigraphic column. It is a subdivision of the Early Cretaceous, Early or Lower Cretaceous epoch (geology), Epoch or series (stratigraphy), S ...
ages of the
Early Cretaceous The Early Cretaceous (geochronology, geochronological name) or the Lower Cretaceous (chronostratigraphy, chronostratigraphic name) is the earlier or lower of the two major divisions of the Cretaceous. It is usually considered to stretch from 143.1 ...
Period of
Europe Europe is a continent located entirely in the Northern Hemisphere and mostly in the Eastern Hemisphere. It is bordered by the Arctic Ocean to the north, the Atlantic Ocean to the west, the Mediterranean Sea to the south, and Asia to the east ...
. Its remains are known from
Belgium Belgium, officially the Kingdom of Belgium, is a country in Northwestern Europe. Situated in a coastal lowland region known as the Low Countries, it is bordered by the Netherlands to the north, Germany to the east, Luxembourg to the southeas ...
(
Bernissart Bernissart (; ) is a municipality of Wallonia located in the province of Hainaut, Belgium. On January 1, 2024, Bernissart had a total population of 11,878. The total area is 43.81 km2, which gives a population density of 271 inhabitants p ...
),
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 List of islands of England, more than 100 smaller adjacent islands. It ...
,
Spain Spain, or the Kingdom of Spain, is a country in Southern Europe, Southern and Western Europe with territories in North Africa. Featuring the Punta de Tarifa, southernmost point of continental Europe, it is the largest country in Southern Eur ...
and
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 ...
. 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. * ...
and only species is ''M. atherfieldensis''. Formerly known as ''
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 ...
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 and weighed . Two rather complete fossils ...
'', making ''Iguanodon'' in its traditional sense
paraphyletic Paraphyly is a taxonomic term describing a grouping that consists of the grouping's last common ancestor and some but not all of its descendant lineages. The grouping is said to be paraphyletic ''with respect to'' the excluded subgroups. In co ...
. It is known from many complete and almost complete skeletons. The genus name honours
Gideon Mantell Gideon Algernon Mantell Membership of the Royal College of Surgeons, MRCS Fellow of the Royal Society, FRS (3 February 1790 – 10 November 1852) was an English obstetrician, geologist and paleontology, palaeontologist. His attempts to reconstr ...
, the discoverer of ''Iguanodon''.


History of discovery


Discovery and naming of the holotype

The
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 ...
fossil, NHMUK PV R5764, was originally discovered by Reginald Walter Hooley in 1914 in the upper
Vectis Formation The Vectis Formation is a geological Formation (geology), formation on the Isle of Wight and Swanage, England whose strata were formed in the Barremian, approximately 125 million years ago."Magnetostratigraphy of the Lower Cretaceous Vectis Forma ...
of southern England and reported upon in 1917. He posthumously named it ''Iguanodon atherfieldensis'' in 1925. Atherfield is the name of a village on the southwest shore of the
Isle of Wight The Isle of Wight (Help:IPA/English, /waɪt/ Help:Pronunciation respelling key, ''WYTE'') is an island off the south coast of England which, together with its surrounding uninhabited islets and Skerry, skerries, is also a ceremonial county. T ...
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 east-south-east of London. The River Medway runs through the centre of the town, l ...
,
Kent Kent is a Ceremonial counties of England, ceremonial county in South East England. It is bordered by Essex across the Thames Estuary to the north, the Strait of Dover to the south-east, East Sussex to the south-west, Surrey to the west, and Gr ...
, owned by William Harding Bensted, in February 1834 (lower Lower Greensand Formation). In June 1834 it was acquired for £25 by scientist
Gideon Mantell Gideon Algernon Mantell Membership of the Royal College of Surgeons, MRCS Fellow of the Royal Society, FRS (3 February 1790 – 10 November 1852) was an English obstetrician, geologist and paleontology, palaeontologist. His attempts to reconstr ...
. He was led to identify it as an ''
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 ...
'' 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 Horn may refer to: Common uses * Horn (acoustic), a tapered sound guide ** Horn antenna ** Horn loudspeaker ** Vehicle horn ** Train horn *Horn (anatomy), a pointed, bony projection on the head of various animals * Horn (instrument), a family ...
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 anatomy, comparative anatomist and paleontology, palaeontologist. Owen is generally considered to have been an outstanding naturalist with a remarkabl ...
, an ambitious scientist with much better funding and society connections in the turbulent worlds of
Reform Act The Reform Acts (or Reform Bills, before they were passed) are legislation enacted in the United Kingdom in the 19th and 20th century to enfranchise new groups of voters and to redistribute seats in the House of Commons of the Parliament of the U ...
–era British politics and science. Owen, at the time a firm creationist, 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 belief systems, God is usually viewed as the supreme being, creator, and principal object of faith. In polytheistic belief systems, a god is "a spirit or being believed to have created, or for controlling some part of the un ...
; 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-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 aggregate bound together with a fluid cement that cures to a solid over time. It is the second-most-used substance (after water), the most–widely used building material, and the most-manufactur ...
sculpted over a
steel Steel is an alloy of iron and carbon that demonstrates improved mechanical properties compared to the pure form of iron. Due to steel's high Young's modulus, elastic modulus, Yield (engineering), yield strength, Fracture, fracture strength a ...
and
brick A brick is a type of construction material used to build walls, pavements and other elements in masonry construction. Properly, the term ''brick'' denotes a unit primarily composed of clay. But is now also used informally to denote building un ...
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 scientific collection, collections that include current and historical records of animals, plants, Fungus, fungi, ecosystems, geology, paleo ...
in
London London is the Capital city, capital and List of urban areas in the United Kingdom, largest city of both England and the United Kingdom, with a population of in . London metropolitan area, Its wider metropolitan area is the largest in Wester ...
. 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. Historically, supporters were left to an individual's free choice and were assu ...
to their
coat of arms A coat of arms is a heraldry, heraldic communication design, visual design on an escutcheon (heraldry), escutcheon (i.e., shield), surcoat, or tabard (the last two being outer garments), originating in Europe. The coat of arms on an escutcheon f ...
in 1949. This specimen has become linked with the name ''I. mantelli'', a species named in 1832 by
Christian Erich 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 ...
in place of ''I. anglicus'', but it actually comes from a different formation 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 Gregory Scott Paul (born December 24, 1954) is an American freelance researcher, author and illustrator who works in paleontology. He is best known for his work and research on theropoda, theropod dinosaurs and his detailed illustrations, both l ...
(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'' 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. The dinosaur is also featured in Maidstone's
coat of arms A coat of arms is a heraldry, heraldic communication design, visual design on an escutcheon (heraldry), escutcheon (i.e., shield), surcoat, or tabard (the last two being outer garments), originating in Europe. The coat of arms on an escutcheon f ...
, 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, officially the Kingdom of Belgium, is a country in Northwestern Europe. Situated in a coastal lowland region known as the Low Countries, it is bordered by the Netherlands to the north, Germany to the east, Luxembourg to the southeas ...
, became the second mounted skeleton of a non-avian dinosaur made primarily out of actual bone when put on display by
Louis Dollo Louis Antoine Marie Joseph Dollo (; 7 December 1857 – 19 April 1931) was a Belgian palaeontologist, known for his work on dinosaurs. He also posited that evolution is not reversible, known as Dollo's law. Together with the Austrian Othenio Ab ...
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 botani ...
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 Gregory Scott Paul (born December 24, 1954) is an American freelance researcher, author and illustrator who works in paleontology. He is best known for his work and research on theropoda, theropod dinosaurs and his detailed illustrations, both l ...
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 (anatomy), rostrum, beak or proboscis. The wet furless surface around the nostrils of the n ...
and trunk 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'', a species based on BMNH R 28685 from
Wessex Formation The Wessex Formation is a fossil-rich England, English geological formation that dates from the Berriasian to Barremian Stage (stratigraphy), stages of the Early Cretaceous. It forms part of the Wealden Group and underlies the younger Vectis Form ...
,
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 List of islands of England, more than 100 smaller adjacent islands. It ...
. Likewise, David B. Norman and Andrew McDonald do not consider ''Dollodon'' a valid
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 ...
or species and instead include it with ''Mantellisaurus''.


Sauerland specimens

In 1971, a fossiliferous
karst Karst () is a topography formed from the dissolution of soluble carbonate rocks such as limestone and Dolomite (rock), dolomite. It is characterized by features like poljes above and drainage systems with sinkholes and caves underground. Ther ...
ic
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 also known as shakeholes, and to openings where surface water ...
clay deposit was found at a quarry just south-west of the village of Nehden near
Brilon Brilon (; ) is a town in North Rhine-Westphalia, Germany, which belongs to the Hochsauerlandkreis. Geography Brilon is on the Brilon Heights, at an altitude of about 450m, and the upper reaches of the river Möhne. The town lies between the Ar ...
in
Sauerland The Sauerland () is a rural, hilly area spreading across most of the south-eastern part of the States of Germany, German federal state of North Rhine-Westphalia, in parts heavily forested and, apart from the major valleys, sparsely inhabited. ...
, Germany containing numerous disarticulated iguanodontid remains predominantly of ''Mantellisaurus'' with lesser quantities of ''
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 ...
'', 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.


Description

''Mantellisaurus'' was a lightly constructed
iguanodont Ornithopoda () is a clade of ornithischian dinosaurs, called ornithopods (). They represent one of the most successful groups of herbivore, herbivorous dinosaurs during the Cretaceous. The most primitive members of the group were bipedal and rel ...
. 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 Bipedalism is a form of terrestrial locomotion where an animal moves by means of its two rear (or lower) limbs or legs. An animal or machine that usually moves in a bipedal manner is known as a biped , meaning 'two feet' (from Latin ''bis'' ...
, only going on all fours when standing still or moving slowly.


Classification

The
cladogram A cladogram (from Greek language, Greek ''clados'' "branch" and ''gramma'' "character") is a diagram used in cladistics to show relations among organisms. A cladogram is not, however, an Phylogenetic tree, evolutionary tree because it does not s ...
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. * Bonsor, J. A., Lockwood, J. A. F., Leite, J. V., Scott-Murray, A., & Maidment, S. C. R. (2023) The Osteology of the Holotype of the British Iguanodontian Dinosaur ''Mantellisaurus atherfieldensis'', ''Monographs of the Palaeontographical Society'', 177:665, 1-63, DOI: 10.1080/02693445.2023.2234156 https://www.tandfonline.com/doi/abs/10.1080/02693445.2023.2234156 {{Portal bar, Paleontology, Dinosaurs, Spain Ankylopollexia Dinosaur genera Barremian dinosaurs Aptian dinosaurs Fossil taxa described in 1925 Fossil taxa described in 2008 Taxa named by Gregory S. Paul