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Iguanodon

Genus of ornithopod dinosaurs

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Iguanodon de Bernissart (Iguanodon bernissartensis) présenté en position quadrupède, Institut Royal des Sciences naturelles de Belgique, Bruxelles
Ben2 · CC BY-SA 3.0

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Through Tooth, Taxonomy (biology) and Belgium

The genus was named in 1825 by English geologist Gideon Mantell, based on fossil specimens found in England, and was given the species name I. anglicus. Iguanodon was the second type of dinosaur formally named based on fossil specimens, after Megalosaurus. Together with Megalosaurus and Hylaeosaurus, it was one of the three genera originally used to define Dinosauria. The genus Iguanodon belongs to the larger group Iguanodontia, along with the duck-billed hadrosaurs. The taxonomy of this genus continues to be a topic of study as new species are named or long-standing ones reassigned to other genera.

In 1878, new, far more complete remains of Iguanodon were discovered in Belgium and studied by Louis Dollo. These were given the new species I. bernissartensis. In the early 21st century it became understood that the remains referred to as Iguanodon in England belonged to four different species (including I. bernissartensis) that were not closely related to each other, which were subsequently split off into Mantellisaurus, Barilium and Hypselospinus. It was also found that the originally described type species of Iguanodon, I. anglicus is now a nomen dubium, and not valid. Thus, the name "Iguanodon" became fixed around the well-known species based primarily on the Belgian specimens. Another species, Iguanodon galvensis, was named in 2015 based on fossils found in the Iberian Peninsula.

Scientific understanding of Iguanodon has evolved over time as new information has been obtained from fossils. The numerous specimens of this genus, including nearly complete skeletons from two well-known bone beds, have allowed researchers to make informed hypotheses regarding many aspects of the living animal, including feeding, movement, and social behaviour. As one of the first scientifically well-known dinosaurs, Iguanodon has occupied a small but notable place in the public's perception of dinosaurs, its artistic representation changing significantly in response to new interpretations of its remains.

Discovery and history

Gideon Mantell, Sir Richard Owen, and the discovery of dinosaurs

The discovery of Iguanodon has long been accompanied by a popular legend. The story posits that Gideon Mantell's wife, Mary Ann, discovered the first teeth of an Iguanodon in the strata of Tilgate Forest in Whitemans Green, Sussex, England, in 1822 while her husband was visiting a patient. However, there is no evidence that Mantell took his wife with him while seeing patients. Furthermore, he admitted in 1851 that he himself had found the teeth, although he had previously stated in 1827 and 1833 that Mrs. Mantell had indeed found the first of the teeth later named Iguanodon. Other later authors agree that the story is not certainly false. It is known from his notebooks that Mantell first acquired large fossil bones from the quarry at Whitemans Green in 1820. Because also theropod teeth were found, thus belonging to carnivores, he at first interpreted these bones, which he tried to combine into a partial skeleton, as those of a giant crocodile. In 1821 Mantell mentioned the find of herbivorous teeth and began to consider the possibility that a large herbivorous reptile was present in the strata. However, in his 1822 publication Fossils of the South Downs he as yet did not dare to suggest a connection between the teeth and his very incomplete skeleton, presuming that his finds presented two large forms, one carnivorous ("an animal of the Lizard Tribe of enormous magnitude"), the other herbivorous.

In May 1822 he first presented the herbivorous teeth to the Royal Society of London but the members, among them William Buckland, dismissed them as fish teeth or the incisors of a rhinoceros from a Tertiary stratum. On 23 June 1823 Charles Lyell showed some to Georges Cuvier, during a soiree in Paris, but the famous French naturalist at once dismissed them as those of a rhinoceros. Though the very next day Cuvier retracted, Lyell reported only the dismissal to Mantell, who became rather diffident about the issue. In 1824 Buckland described Megalosaurus and was on that occasion invited to visit Mantell's collection. Seeing the bones on 6 March he agreed that these were of some giant saurian—though still denying it was a herbivore. Emboldened nevertheless, Mantell again sent some teeth to Cuvier, who answered on 22 June 1824 that he had determined that they were reptilian and quite possibly belonged to a giant herbivore. In a new edition that year of his Recherches sur les Ossemens Fossiles Cuvier admitted his earlier mistake, leading to an immediate acceptance of Mantell, and his new saurian, in scientific circles. Mantell tried to corroborate his theory further by finding a modern-day parallel among extant reptiles. In September 1824 he visited the Royal College of Surgeons but at first failed to find comparable teeth. However, assistant-curator Samuel Stutchbury recognised that they resembled those of an iguana he had recently prepared, albeit twenty times longer.

In recognition of the resemblance of the teeth to those of the iguana, Mantell decided to name his new animal Iguanodon or 'iguana-tooth', from iguana and the Greek word ὀδών (odon, odontos or 'tooth'). Based on isometric scaling, he estimated that the creature might have been up to 18 metres (59 feet) long, more than the 12-metre (39 ft) length of Megalosaurus. His initial idea for a name was Iguana-saurus ('Iguana lizard'), but his friend William Daniel Conybeare suggested that that name was more applicable to the iguana itself, so a better name would be Iguanoides ('Iguana-like') or Iguanodon. He neglected to add a specific name to form a proper binomial, but one was supplied in 1829 by Friedrich Holl: I. anglicum, which was later emended to I. anglicus.

Mantell sent a letter detailing his discovery to the local Portsmouth Philosophical Society in December 1824, several weeks after settling on a name for the fossil creature. The letter was read to members of the Society at a meeting on 17 December, and a report was published in the Hampshire Telegraph the following Monday, 20 December, which announced the name, misspelled as "Iguanadon". Mantell formally published his findings on 10 February 1825, when he presented a paper on the remains to the Royal Society of London.

A more complete specimen of a similar animal was discovered in a quarry in Maidstone, Kent, in 1834 (lower Lower Greensand Formation), which Mantell soon acquired. He was led to identify it as an Iguanodon based on its distinctive teeth. The Maidstone slab (NHMUK PV OR 3791) was used 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. The discovery of much better specimens 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 in London. The borough of Maidstone commemorated this find by adding an Iguanodon as a supporter to their coat of arms 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 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 is classified as cf. Mantellisaurus by McDonald (2012); as cf. Mantellisaurus atherfieldensis by Norman (2012); and made the holotype of a separate species Mantellodon carpenteri by Paul (2012), but this is considered dubious and it is generally considered a specimen of Mantellisaurus.

At the same time, tension began to build between Mantell and Richard Owen, an ambitious scientist with much better funding and society connections in the turbulent worlds of Reform Act-era British politics and science. Owen, 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 feet), determined that they were not simply giant lizards, and put forward that they were advanced and mammal-like, characteristics given to them by God; according to the understanding of the time, they could not have been "transmuted" from reptiles to mammal-like creatures.

In 1849, a few years before his death in 1852, Mantell realised that iguanodonts were not heavy, pachyderm-like animals, as Owen was putting forward, but had slender forelimbs. However, since his passing left him unable to participate in the creation of the Crystal Palace dinosaur sculptures, Owen's vision of the dinosaurs became that seen by the public for decades. With Benjamin Waterhouse Hawkins, he had nearly two dozen lifesize sculptures of various prehistoric animals built out of concrete sculpted over a steel and brick framework; two iguanodonts (based on the Maidstone specimen), one standing and one resting on its belly, were included. Before the sculpture of the standing iguanodont was completed, he held a banquet for twenty inside it.

Bernissart mine discoveries and Dollo's new reconstruction

The largest find of Iguanodon remains to that date occurred on 28 February 1878 in a coal mine at Bernissart in Belgium, at a depth of 322 m (1,056 ft), when two mineworkers, Jules Créteur and Alphonse Blanchard, accidentally hit on a skeleton that they initially took for petrified wood. With the encouragement of Alphonse Briart, supervisor of mines at nearby Morlanwelz, Louis de Pauw on 15 May 1878 started to excavate the skeletons and in 1882 Louis Dollo reconstructed them. At least 38 Iguanodon individuals were uncovered, most of which were adults. In 1882, the holotype specimen of I. bernissartensis became one of the first ever dinosaur skeletons mounted for display. It was put together in a chapel at the Palace of Charles of Lorraine using a series of adjustable ropes attached to scaffolding so that a lifelike pose could be achieved during the mounting process. This specimen, along with several others, first opened for public viewing in an inner courtyard of the palace in July 1883. In 1891 they were moved to the Royal Museum of Natural History, where they are still on display; nine are displayed as standing mounts, and nineteen more are still in the Museum's basement. The exhibit makes an impressive display in the Royal Belgian Institute of Natural Sciences, in Brussels. A replica of one of these is on display at the Oxford University Museum of Natural History and at the Sedgwick Museum in Cambridge. Most of the remains were referred to a new species, I. bernissartensis, a larger and much more robust animal than the English remains had yet revealed. One specimen, IRSNB 1551, was at first referred to the nebulous, gracile I. mantelli, but is currently referred to Mantellisaurus atherfieldensis. The skeletons were some of the first complete dinosaur skeletons known. Found with the dinosaur skeletons were the remains of plants, fish, and other reptiles, including the crocodyliform Bernissartia.

The science of conserving fossil remains was in its infancy, and new techniques had to be improvised to deal with what soon became known as "pyrite disease". Crystalline pyrite in the bones was being oxidized to iron sulphate, accompanied by an increase in volume that caused the remains to crack and crumble. When in the ground, the bones were isolated by anoxic moist clay that prevented this from happening, but when removed into the drier open air, the natural chemical conversion began to occur. To limit this effect, De Pauw immediately, in the mine-gallery, re-covered the dug-out fossils with wet clay, sealing them with paper and plaster reinforced by iron rings, forming in total about six hundred transportable blocks with a combined weight of a hundred and thirty tons. In Brussels after opening the plaster he impregnated the bones with boiling gelatine mixed with oil of cloves as a preservative. Removing most of the visible pyrite he then hardened them with hide glue, finishing with a final layer of tin foil. Damage was repaired with papier-mâché. This treatment had the unintended effect of sealing in moisture and extending the period of damage. In 1932 museum director Victor van Straelen decided that the specimens had to be completely restored again to safeguard their preservation. From December 1935 to August 1936 the staff at the museum in Brussels treated the problem with a combination of alcohol, arsenic, and 390 kilograms of shellac. This combination was intended to simultaneously penetrate the fossils (with alcohol), prevent the development of mould (with arsenic), and harden them (with shellac). The fossils entered a third round of conservation from 2003 until May 2007, when the shellac, hide glue and gelatine were removed and impregnated with polyvinyl acetate and cyanoacrylate and epoxy glues. Modern treatments of this problem typically involve either monitoring the humidity of fossil storage, or, for fresh specimens, preparing a special coating of polyethylene glycol that is then heated in a vacuum pump, so that moisture is immediately removed and pore spaces are infiltrated with polyethylene glycol to seal and strengthen the fossil.

Dollo's specimens allowed him to show that Owen's prehistoric pachyderms were not correct for Iguanodon. He instead modelled the skeletal mounts after the cassowary and wallaby, and put the spike that had been on the nose firmly on the thumb. His reconstruction would prevail for a long period of time, but would later be discounted.

Excavations at the quarry were stopped in 1881, although it was not exhausted of fossils, as recent drilling operations have shown. During World War I, when the town was occupied by German forces, preparations were made to reopen the mine for palaeontology, and Otto Jaekel was sent from Berlin to supervise. Just as the first fossiliferous layer was about to be uncovered, however, the German army surrendered and had to withdraw. Further attempts to reopen the mine were hindered by financial problems and were stopped altogether in 1921 when the mine flooded.

Turn of the century and the Dinosaur Renaissance

Research on Iguanodon decreased during the early part of the 20th century as World Wars and the Great Depression enveloped Europe. A new species that would become the subject of much study and taxonomic controversy, I. atherfieldensis, was named in 1925 by R. W. Hooley, for a specimen collected at Atherfield Point on the Isle of Wight.

Iguanodon was not part of the initial work of the dinosaur renaissance that began with the description of Deinonychus in 1969, but it was not neglected for long. David B. Weishampel's work on ornithopod feeding mechanisms provided a better understanding of how it fed, and David B. Norman's work on numerous aspects of the genus has made it one of the best-known dinosaurs. In addition, a further find of numerous disarticulated Iguanodon bones in Nehden, Nordrhein-Westphalen, Germany, has provided evidence for gregariousness in this genus, as the animals in this areally restricted find appear to have been killed by flash floods. At least 15 individuals, from 2 to 8 metres (6 ft 7 in to 26 ft 3 in) long, have been found here, most of the individuals belong to the related Mantellisaurus (described as I. atherfieldensis, at that time believed to be another species of Iguanodon). but some are of I. bernissartensis.

One major revision to Iguanodon brought by the Renaissance would be another re-thinking of how to reconstruct the animal. A major flaw with Dollo's reconstruction was the bend he introduced into the tail. This organ was more or less straight, as shown by the skeletons he was excavating, and the presence of ossified tendons. In fact, to get the bend in the tail for a more wallaby or kangaroo-like posture, the tail would have had to be broken. With its correct, straight tail and back, the animal would have walked with its body held horizontal to the ground, arms in place to support the body if needed.

21st century research and the splitting of the genus

In the 21st century, Iguanodon material has been used in the search for dinosaur biomolecules. In research by Graham Embery et al. (2003), Iguanodon bones were processed to look for remnant proteins. In this research, identifiable remains of typical bone proteins, such as phosphoproteins and proteoglycans, were found in a rib. In 2007, Gregory S. Paul split I. atherfieldensis into a new, separate genus, Mantellisaurus which has been generally accepted. In 2009 fragmentary iguanodontid material was described from upper Barremian Paris Basin deposits in Auxerre, Burgundy. While not definitively diagnosable to the genus/species level, the specimen shares "obvious morphological and dimensional affinities" with I. bernissartensis.

In 2010, David Norman split the Valanginian species I. dawsoni and I. fittoni into Barilium and Hypselospinus respectively. After Norman 2010, over half a dozen new genera were named off English "Iguanodon" material. Carpenter and Ishida in 2010 named Proplanicoxa, Torilion and Sellacoxa while Gregory S. Paul in 2012 named Darwinsaurus, Huxleysaurus and Mantellodon and Macdonald et al. in 2012 named Kukufeldia. These species named after Norman 2010 are not considered valid and are considered various junior synonyms of Mantellisaurus, Barilium and Hypselospinus.

In 2011, a new genus Delapparentia was named for a specimen in Spain that was originally thought to belong to I. bernissartensis. The previous identification was subsequently reaffirmed in a new analysis of individual variation in the Belgian specimens, finding that the Delapparentia specimen was within the range of I. bernissartensis. In 2015 a new species of Iguanodon, I. galvensis, was named based on material including 13 juvenile (perinate) individuals found in the Camarillas Formation near Galve, Spain. In 2017 a new study was done of I. galvensis, with further evidence of distinctiveness from I. bernissartensis including several new autapomorphies. It was also found that the Delapparentia holotype (which is also from the Camarillas Formation) was not distinguishable from either I. bernissartensis or I. galvensis. In 2025, Franco Sancarlo, Davide Mandorlo and Tracy L. Ford assigned I. galvensis to a new genus, Paulodon, based on putative differences between it and I. bernissartensis; although paleontologists have informally disagreed with the methodology behind the paper. In 2026, Rotatori and colleagues suggested that Delapperantia is a distinct taxon more closely related to Magnamanus and "I." galvensis, the latter of which they suggested that a new genus name would be required.

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