Iguanodon
Iguanodon | |
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I. bernissartensis mounted in modern quadrupedal posture, Royal Belgian Institute of Natural Sciences , Brussels
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Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Clade: | Dinosauria |
Clade: | †Ornithischia |
Clade: | †Ornithopoda |
Family: | †Iguanodontidae |
Genus: | †Iguanodon Mantell, 1825[1] |
Type species | |
†Iguanodon bernissartensis Boulenger in Beneden, 1881
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Other species | |
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Synonyms | |
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Iguanodon (
The genus was named in 1825 by English geologist Gideon Mantell but discovered by William Harding Bensted, 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. In 2015, a second valid species, I. galvensis, was named, based on fossils found in the Iberian Peninsula.
Scientific understanding of Iguanodon has evolved over time as new information has been obtained from
Discovery and history
Gideon Mantell, Sir Richard Owen, and the discovery of dinosaurs
The discovery of Iguanodon has long been accompanied by a popular
In May 1822 he first presented the herbivorous teeth to the
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
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".[14] Mantell formally published his findings on 10 February 1825, when he presented a paper on the remains to the Royal Society of London.[1][5]
A more complete specimen of a similar animal was discovered in a quarry in
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.[21][22]
In 1849, a few years before his death in 1852, Mantell realised that iguanodonts were not heavy, pachyderm-like animals,[23] 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.[21] 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.[24][25][26]
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
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 "
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.[34][35] His reconstruction would prevail for a long period of time, but would later be discounted.[27]
Excavations at the quarry were stopped in 1881, although it was not exhausted of fossils, as recent drilling operations have shown.[36] 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.[27][37]
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.[38]
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,[39] and David B. Norman's work on numerous aspects of the genus has made it one of the best-known dinosaurs.[28][27][40][41] 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).[29][42] 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., 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.[43] In 2007, Gregory S. Paul split I. atherfieldensis into a new, separate genus, Mantellisaurus which has been generally accepted.[44] 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.[45]
In 2010, David Norman split the Valanginian species I. dawsoni and I. fittoni into Barilium and Hypselospinus respectively.[46] 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.[20]
In 2011, a new genus Delapparentia was named for a specimen in Spain that was originally thought to belong to I. bernissartensis.
Description
Iguanodon were bulky herbivores that could shift from
The arms of I. bernissartensis were long (up to 75% the length of the legs) and robust,[11] with rather inflexible hands built so that the three central fingers could bear weight.[28] The thumbs were conical spikes that stuck out away from the three main digits. In early restorations, the spike was placed on the animal's nose. Later fossils revealed the true nature of the thumb spikes,[27] although their exact function is still debated. They could have been used for defense, or for foraging for food. The little finger was elongated and dextrous, and could have been used to manipulate objects. The phalangeal formula is 2-3-3-2-4, meaning that the innermost finger (phalange) has two bones, the next has three, etc.[54] The legs were powerful, but not built for running, and each foot had three toes. The backbone and tail were supported and stiffened by ossified tendons, which were tendons that turned to bone during life (these rod-like bones are usually omitted from skeletal mounts and drawings).[28]
These animals had large, tall but narrow skulls, with toothless beaks probably covered with keratin, and teeth like those of iguanas, as the name suggests, but much larger and more closely packed.[28] Unlike hadrosaurids, which had columns of replacement teeth, Iguanodon only had one replacement tooth at a time for each position. The upper jaw held up to 29 teeth per side, with none at the front of the jaw, and the lower jaw 25; the numbers differ because teeth in the lower jaw are broader than those in the upper.[55] Because the tooth rows are deeply inset from the outside of the jaws, and because of other anatomical details, it is believed that, as with most other ornithischians, Iguanodon had some sort of cheek-like structure, muscular or non-muscular, to retain food in the mouth.[56][57]
Classification and evolution
Iguanodon gives its name to the
With the advent of
The cladogram below follows an analysis by Andrew McDonald, 2012.[61]
Iguanodontia |
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Species
Because Iguanodon is one of the first dinosaur genera to have been named, numerous species have been assigned to it. While never becoming the wastebasket taxon several other early genera of dinosaurs (such as Megalosaurus) became, Iguanodon has had a complicated history, and its taxonomy continues to undergo revisions.[62][42][63][64] Although Gregory Paul recommended restricting I. bernissartensis to the famous sample from Bernissart, ornithopod workers like Norman and McDonald have disagreed with Paul's recommendations, except exercising caution when accepting records of Iguanodon from France and Spain as valid.[42][2][65]
I. anglicus was the original
Species currently accepted as valid
Only two species assigned to Iguanodon are still considered to be valid.[28][42]
- I. bernissartensis, described by George Albert Boulenger in 1881, is the type species for the genus. This species is best known for the many skeletons discovered in the Sainte-Barbe Clays Formation at Bernissart, but is also known from remains across Europe.
- Delapparentia turolensis, named in 2011[3] based on a specimen previously assigned to Iguanodon bernissartensis,[67] was argued to be distinct from the latter based on the relative height of its neural spines.[68] However, a 2017 study noted that this is easily within the range of individual variation, and that the difference may also arise from D. turolensis being an adult older than other specimens of I. bernissartensis.[47]
- I. seelyi (also incorrectly spelled I. seeleyi), described by John Hulke in 1882, has also been synonymised with Iguanodon bernissartensis, though this is not universally accepted. It was discovered in Brook, on the Isle of Wight, and named after Charles Seely MP, Liberal politician and philanthropist, on whose estate it was found.[42][69]
- David Norman has suggested that I. bernissartensis includes the dubious Mongolian I. orientalis (see also below),[70] but this has not been followed by other researchers.[42]
- I. galvensis, described in 2015, is based on adult and juvenile remains found in Barremian-age deposits in Teruel, Spain.[2]
Reassigned species of Iguanodon
- I. albinus (or Albisaurus scutifer), described by
- I. atherfieldensis, described by R.W. Hooley in 1925,[38] was smaller and less robust than I. bernissartensis, with longer neural spines. It was renamed Mantellisaurus atherfieldensis in 2007.[62] The Bernissart specimen RBINS 1551 was described as Dollodon bampingi in 2008, but McDonald and Norman returned Dollodon to synonymy with Mantellisaurus.[46][72]
- I. dawsoni, described by Lydekker in 1888,
- I. exogyrarum was described by Fritsch in 1878. It is a nomen dubium based on very poor material and was renamed Ponerosteus in 2000.[74]
- I. fittoni was described by Lydekker in 1889.[75] Like I. dawsoni, this species was described from the Wadhurst Clay[42] of East Sussex.[28] It is now the type species of Hypselospinus.[46]
- I. hilli, coined by Edwin Tully Newton in 1892 for a tooth from the early hadrosaurid of some sort.[76] However, recent work places it as indeterminate beyond Hadrosauroidea outside Hadrosauridae.[77]
- I. hoggi (also spelled I. boggii or hoggii), named by Owen for a lower jaw from the
- I. hollingtoniensis (also spelled I. hollingtonensis), described by Lydekker in 1889, has variously been considered a synonym of Hypselospinus fittoniDarwinsaurus evolutionis.[18]
- I. lakotaensis was described by David B. Weishampel and Philip R. Bjork in 1989.[79] The only well-accepted North American species of Iguanodon, I. lakotaensis was described from a partial skull from the Barremian-age Lower Cretaceous Lakota Formation of South Dakota. Its assignment has been controversial. Some researchers suggest that it was more basal than I. bernissartensis, and related to Theiophytalia,[80] but David Norman has suggested that it was a synonym of I. bernissartensis.[63] Gregory S. Paul has since given the species its own genus, Dakotadon.[42]
- I. mantelli described by Mantellodon, but also the dubious hadrosauroid Trachodon cantabrigiensis the hypsilophodont Hypsilophodon, and Valdosaurus, were previously mis-assigned to I. mantelli.
- "I. mongolensis" is a nomen nudum from a photo caption in a book by Whitfield in 1992[82] of remains that would later be named Altirhinus.[83]
- I. orientalis, described by A. K. Rozhdestvensky in 1952,[84] was based on poor material, but a skull with a distinctive arched snout that had been assigned to it was renamed Altirhinus kurzanovi in 1998.[63] At the same time, I. orientalis was considered to be a nomen dubium because it cannot be compared to I. bernissartensis.[63][70]
- I. phillipsi was described by Harry Seeley in 1869,[85] but he later reassigned it to Priodontognathus.[86]
- I. praecursor (also spelled I. precursor), described by E. Sauvage in 1876 from teeth from an unnamed Kimmeridgian (Late Jurassic) formation in Pas-de-Calais, France, is actually a sauropod, sometimes assigned to Neosodon,[87] although the two come from different formations.[88]
- I. prestwichii (also spelled I. prestwichi), described by John Hulke in 1880, has been reassigned to Camptosaurus prestwichii or to its own genus Cumnoria.
- I. suessii, described by Emanuel Bunzel in 1871, has been reassigned to Mochlodon suessi.[28]
Species reassigned to Iguanodon
- I. foxii (also spelled I. foxi) was originally described by Thomas Henry Huxley in 1869 as the type species of Hypsilophodon; Owen (1873 or 1874) reassigned it to Iguanodon, but his assignment was soon overturned.[89]
- I. gracilis, named by Lydekker in 1888 as the type species of Sphenospondylus and assigned to Iguanodon in 1969 by Rodney Steel, has been suggested to be a synonym of Mantellisaurus atherfieldensis,[28] but is considered dubious nowadays.[41][72]
- I. major, a species named by Justin Delair in 1966,[90] based on vertebrae from the Isle of Wight and Sussex originally described by Owen in 1842 as a species of Streptospondylus, S. major, is a nomen dubium.[69]
- I. valdensis, a renaming of Vectisaurus valdensis by Ernst van den Broeck in 1900.[91] Originally named by Hulke as a distinct genus in 1879 based on vertebral and pelvic remains, it was from the Barremian stage of the Isle of Wight.[92] It was considered a juvenile specimen of Mantellisaurus atherfieldensis,[93] or an undetermined species of Mantellisaurus,[42] but is indeterminate beyond Iguanodontia.[72]
- The nomen nudum "Proiguanodon" (van den Broeck, 1900) also belongs here.[94]
Dubious species
- I. anglicus, described by Friedrich Holl in 1829,junior objective synonym, a later name for the material of I. anglicus.
- I. ottingeri, described by Peter Galton and James A. Jensen in 1979, is a nomen dubium based on teeth from the possibly Aptian-age lower Cedar Mountain Formation of Utah.[97]
Palaeobiology
Feeding
One of the first details noted about Iguanodon was that it had the teeth of a
The skull was structured in such a way that as it closed, the bones holding the teeth in the upper jaw would bow out. This would cause the lower surfaces of the upper jaw teeth to rub against the upper surface of the lower jaw's teeth, grinding anything caught in between and providing an action that is the rough equivalent of mammalian chewing.[39] Because the teeth were always replaced, the animal could have used this mechanism throughout its life, and could eat tough plant material.[100] Additionally, the front ends of the animal's jaws were toothless and tipped with bony nodes, both upper and lower,[28] providing a rough margin that was likely covered and lengthened by a keratinous material to form a cropping beak for biting off twigs and shoots.[27] Its food gathering would have been aided by its flexible little finger, which could have been used to manipulate objects, unlike the other fingers.[28]
Exactly what Iguanodon ate with its well-developed jaws is not known. The size of the larger species, such as I. bernissartensis, would have allowed them access to food from ground level to
Posture and movement
Early fossil remains were fragmentary, which led to much speculation on the posture and nature of Iguanodon. Iguanodon was initially portrayed as a quadrupedal horn-nosed beast. However, as more bones were discovered, Mantell observed that the forelimbs were much smaller than the hindlimbs. His rival Owen was of the opinion it was a stumpy creature with four pillar-like legs. The job of overseeing the first lifesize reconstruction of dinosaurs was initially offered to Mantell, who declined due to poor health, and Owen's vision subsequently formed the basis on which the sculptures took shape. Its bipedal nature was revealed with the discovery of the Bernissart skeletons. However, it was depicted in an upright posture, with the tail dragging along the ground, acting as the third leg of a tripod.[104]
During his re-examination of Iguanodon, David Norman was able to show that this posture was unlikely, because the long tail was stiffened with ossified tendons.
Furthermore, it appears that Iguanodon became more quadrupedal as it got older and heavier;
Large three-toed footprints are known in Early Cretaceous rocks of England, particularly Wealden beds on the Isle of Wight, and these trace fossils were originally difficult to interpret. Some authors associated them with dinosaurs early on. In 1846, E. Tagert went so far as to assign them to an ichnogenus he named Iguanodon,[108] and Samuel Beckles noted in 1854 that they looked like bird tracks, but might have come from dinosaurs.[109] The identity of the trackmakers was greatly clarified upon the discovery in 1857 of the hind leg of a young Iguanodon, with distinctly three-toed feet, showing that such dinosaurs could have made the tracks.[110][111] Despite the lack of direct evidence, these tracks are often attributed to Iguanodon.[27] A trackway in England shows what may be an Iguanodon moving on all fours, but the foot prints are poor, making a direct connection difficult.[40] Tracks assigned to the ichnogenus Iguanodon are known from locations including places in Europe where the body fossil Iguanodon is known, to Spitsbergen, Svalbard, Norway.[112][113]
Thumb spike
The thumb spike is one of the best-known features of Iguanodon. Although it was originally placed on the animal's nose by Mantell, the complete Bernissart specimens allowed Dollo to place it correctly on the hand, as a modified thumb.[104] (This would not be the last time a dinosaur's modified thumb claw would be misinterpreted; Noasaurus, Baryonyx, and Megaraptor are examples since the 1980s where an enlarged thumb claw was first put on the foot, as in dromaeosaurids.[114][115])
This thumb is typically interpreted as a close-quarter stiletto-like weapon against predators,[28][27] although it could also have been used to break into seeds and fruits,[28] or against other Iguanodon.[11] One author has suggested that the spike was attached to a venom gland,[116][117] but this has not been accepted, as the spike was not hollow,[11] nor were there any grooves on the spike for conducting venom.[118][nb 1]
Possible social behaviour
Although sometimes interpreted as the result of a single catastrophe, the Bernissart finds instead are now interpreted as recording multiple events. According to this interpretation, at least three occasions of mortality are recorded, and though numerous individuals would have died in a geologically short time span (?10–100 years),[29] this does not necessarily mean these Iguanodon were herding animals.[28]
An argument against herding is that juvenile remains are very uncommon at this site, unlike modern cases with herd mortality. They more likely were the periodic victims of flash floods whose carcasses accumulated in a lake or marshy setting.[29] The Nehden find, however, with its greater span of individual ages, more even mix of Dollodon or Mantellisaurus to Iguanodon bernissartensis, and confined geographic nature, may record mortality of herding animals migrating through rivers.[29]
There is no evidence that Iguanodon was sexually dimorphic (with one sex appreciably different from the other).[47] At one time, it was suggested that the Bernissart I. "mantelli", or I. atherfieldensis (Dollodon and Mantellisaurus, respectively) represented a sex, possibly female, of the larger and more robust, possibly male, I. bernissartensis.[119] However, this is not supported today.[27][40][62] A 2017 analysis showed that I. bernissartensis does exhibit a large level of individual variation in both its limbs (scapula, humerus, thumb claw, ilium, ischium, femur, tibia) and spinal column (axis, sacrum, tail vertebrae). Additionally, this analysis found that individuals of I. bernissartensis generally seemed to fall into two categories based on whether their tail vertebrae bore a furrow on the bottom, and whether their thumb claws were large or small.[47]
Paleopathology
Evidence of a fractured hip bone was found in a specimen of Iguanodon, which had an injury to its ischium. Two other individuals were observed with signs of osteoarthritis as evidenced by bone overgrowths in their anklebones which are called osteophytes.[120]
In popular culture
Since its description in 1825, Iguanodon has been a feature of worldwide
Several
Aside from appearances in movies, Iguanodon has also been featured on the
Because it is both one of the first dinosaurs described and one of the best-known dinosaurs, Iguanodon has been well-placed as a barometer of changing public and scientific perceptions on dinosaurs. Its reconstructions have gone through three stages: the elephantine quadrupedal horn-snouted reptile satisfied the Victorians, then a bipedal but still fundamentally reptilian animal using its tail to prop itself up dominated the early 20th century, but was slowly overturned during the 1960s by its current, more agile and dynamic representation, able to shift from two legs to all fours.[126]
Notes
- ^ Naish's works cite Tweedie (1977) as the source of the venomous spike proposal, but this work does not explicitly attribute venom to Iguanodon, only noting on page 69 that there is no hard-tissue evidence for venom in stingray tails and platypus spurs.[117] Elting and Goodman (1987:46) explicitly report a venom spike proposal, attributing it to "one scientist".[116]
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External links
- The Bernissart Iguanodons (Iguanodon herd found in Belgium).
- Mantell's Iguanodon tooth in the collection of the Museum of New Zealand Te Papa Tongarewa