Iguanodontids Temporal range: Early Cretaceous, | |
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Iguanodon bernissartensis mounted in modern quadrupedal posture, Royal Belgian Institute of Natural Sciences, Brussels | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Clade: | Dinosauria |
Clade: | † Ornithischia |
Clade: | † Neornithischia |
Clade: | † Ornithopoda |
Clade: | † Hadrosauriformes |
Family: | † Iguanodontidae Bonaparte, 1850 |
Subgroups | |
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Iguanodontidae is a family of iguanodontians belonging to Styracosterna, a derived clade within Ankylopollexia. The clade is formally defined in the PhyloCode by Daniel Madzia and colleagues in 2021 as "the largest clade containing Iguanodon bernissartensis , but not Hadrosaurus foulkii ". [2]
Characterized by their elongated maxillae, they were herbivorous and typically large in size. This family exhibited locomotive dynamism; there exists evidence for both bipedalism and quadrupedalism within iguanodontid species, supporting the idea that individual organisms were capable of both locomoting exclusively with their hind limbs and locomoting quadrupedally. [3] Iguanodontids possess hoof-like second, third, and fourth digits, and in some cases, a specialized thumb spike and an opposable fifth digit. [4] Their skull construction allows for a strong chewing mechanism called a transverse power stroke. [5] This, paired with their bilateral dental occlusion, made them extremely effective as herbivores. [6] Members of Iguanodontidae are thought to have had a diet that consisted of both gymnosperms and angiosperms, the latter of which co-evolved with the iguanodontids in the Cretaceous period. [7]
There is no consensus on the phylogeny of the group. Iguanodontidae is most frequently characterized as paraphyletic with respect to Hadrosauridae, [8] [9] although some researchers advocate for a monophyletic view of the family. [10] [11]
The upper surface of a typical iguanodontid skull has a convex curve that extends from the snout to just past the orbit, where the skull flattens out to form a roughly level plane directly above the braincase. [5] The antorbital fenestra, an opening in the skull anterior to the eye sockets, is reduced in size in iguanodontids. Their maxillae are roughly triangular, fairly flat, and sport thickened bony walls. An elongated maxilla is characteristic of the family. [12] Iguanodontid dentaries are very long as well, and become increasingly thick towards the back of the skull. A pair of bony processes extending from the maxilla insert into the jugal and lacrimal, respectively. The iguanodontid jugal has particularly deep crevices that serve to mediate this contact. The lacrimal process constitutes the rostral margin of the reduced antorbital fenestra. [5]
Iguanodontids are generally limited to the possession of single replacement tooth at each position, although exceptions exist. The most primitive example bears positions for 13 maxillary and 14 dentary teeth. More derived forms have a larger number of positions per row. For example, I. bernissartensis is able to accommodate up to 29 maxillary and 25 dentary teeth. Iguanodontids exhibit contact between maxillary and dentary teeth upon closure of the jaw. [6] They have a thick layer of enamel over the lip-facing (labial) surface of the crown, a robust primary ridge beginning at the base of the crown, and a denticulate margin. Most members of the family have maxillary tooth crowns lanceolate in shape. The labial surface of the teeth has some grooves, while the tongue-facing (lingual) surface is smooth. Iguanodontids have lost their premaxillary teeth. [5]
The second, third, and fourth digits of the iguanodontid forelimb are close together. In some cases, it is possible that digits three and four were bound into a single structure by layers of skin, a specialized adaptation for quadruped locomotion. [4] In addition, the wrist bones are fused into a block, and the thumb bones are fused into a spike-like point. In Iguanodon , the fifth digit is long, flexible, and opposable. On the hind limb, digits two, three, and four are wide and short, with blunt claws that resemble hooves. [5]
All of the cervical vertebrae have ribs attached. The initial set are linear; the rest are two-headed. Tendons along the neural arches were ossified, limiting mobility in the backbone in exchange for reinforcement. A similar ossification is seen in the tail. [12] Iguanodontids have a rod-shaped pubis that extends parallel to the ischium. The paired sternal bones are often hatchet-shaped. The humerus has a shallow curve, in contrast to the straight ulna and radius. The ilium is thinner at the anterior end than it is at the posterior. Evidence suggests that these dinosaurs do not have plated, armored skin. [5]
In the past, Iguanodontidae became a waste-basket for any ornithopod that did not belong in either Hadrosauridae, or the now defunct Hypsilophodontidae. A number of studies suggest that Iguanodontidae as traditionally defined is paraphyletic with respect to Hadrosauridae. [13] That is, iguanodontids represent successive steps in the acquisition of advanced hadrosaurian characteristics, and in this view cannot be defined as a single distinct clade. [14] Nevertheless, some researchers have found support for a monophyletic Iguanodontidae consisting of a handful of genera. [10] [11] Some other studies, however, fail to recover the group. [8] The left cladogram was recovered in a 2015 analysis that supports a monophyletic Iguanodontidae, [11] whereas the right cladogram from 2012 study finds the group to be paraphyletic: [9]
Fossilized footprints provide evidence for both quadrupedality and bipedality within iguanodontids. It is thought that iguanodontids were primarily quadrupedal but could optionally walk on two limbs. The ossification of tendons along the neural arches may have played a role in facilitating the dynamic pedality of iguanodontids, as the ossified tendons could help withstand the additional stress incurred on the backbone by standing upright. [12] Some research suggests that organism size plays a role in the determination of pedality, where larger organisms are more likely to choose to walk on all fours than their smaller counterparts. [3]
Iguanodontids are low-browsing herbivores that fed extensively on gymnosperms like ferns and horsetails, especially during the early Cretaceous period. These dinosaurs were very effective as herbivores due in part to their combination of bilateral dental occlusion with the transverse power stroke of their chewing mechanism. Additionally, iguanodontids lack a rigid secondary palate, which helps to mitigate torsional stresses during occlusion, a feature that enhanced their ability to break down plant matter. [5] Additionally, the iguanodontids co-evolved with the radiation of angiosperms in the Cretaceous period. Angiosperms typically develop more rapidly and lower to the ground than gymnosperms; their proliferation provided a wealth of easily accessible food for the members of Iguanodontidae. [7]
Iguanodon, 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, taxonomic revision in the early 21st century has defined Iguanodon to be based on one well-substantiated species: I. bernissartensis, which lived during the Barremian to early Aptian ages of the Early Cretaceous in Belgium, Germany, England, and Spain, between about 126 and 122 million years ago. Iguanodon was a large, bulky herbivore, measuring up to 9–11 metres (30–36 ft) in length and 4.5 metric tons in body mass. Distinctive features include large thumb spikes, which were possibly used for defense against predators, combined with long prehensile fifth fingers able to forage for food.
Hypsilophodon is a neornithischian dinosaur genus from the Early Cretaceous period of England. It has traditionally been considered an early member of the group Ornithopoda, but recent research has put this into question.
Muttaburrasaurus was a genus of herbivorous iguanodontian ornithopod dinosaur, which lived in what is now northeastern Australia sometime between 112 and 103 million years ago during the early Cretaceous period. It has been recovered in some analyses as a member of the iguanodontian clade Rhabdodontomorpha. After Kunbarrasaurus, it is Australia's most completely known dinosaur from skeletal remains. It was named after Muttaburra, the site in Queensland, Australia, where it was found.
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 small-sized, while advanced members of the subgroup Iguanodontia became quadrupedal and developed large body size. Their major evolutionary advantage was the progressive development of a chewing apparatus that became the most sophisticated ever developed by a non-avian dinosaur, rivaling that of modern mammals such as the domestic cow. They reached their apex of diversity and ecological dominance in the hadrosaurids, before they were wiped out by the Cretaceous–Paleogene extinction event along with all other non-avian dinosaurs. Members are known worldwide.
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 7–8.3 metres (23–27 ft) long and weighed 2.2 metric tons. Two rather complete fossils were found in the Elrhaz Formation, Gadoufaoua deposits, Agadez, Niger, in 1965 and 1970, with a third indeterminate specimen known from the Koum Formation of Cameroon. The animal was named in 1976 by French paleontologist Philippe Taquet; the type species being Ouranosaurus nigeriensis. The generic name is a combination of ourane, a word with multiple meanings, and sauros, the Greek word for lizard. The specific epithet nigeriensis alludes to Niger, its country of discovery. And so, Ouranosaurus nigeriensis could be interpreted as "brave lizard originating from Niger".
Hypsilophodontidae is a traditionally used family of ornithopod dinosaurs, generally considered invalid today. It historically included many small bodied bipedal neornithischian taxa from around the world, and spanning from the Middle Jurassic until the Late Cretaceous. This inclusive status was supported by some phylogenetic analyses from the 1990s and mid 2000s, although there have also been many finding that the family is an unnatural grouping which should only include the type genus, Hypsilophodon, with the other genera being within clades like Thescelosauridae and Elasmaria. A 2014 analysis by Norman recovered a grouping of Hypsilophodon, Rhabdodontidae and Tenontosaurus, which he referred to as Hypsilophodontia. That clade is formally defined in the PhyloCode as "the smallest clade within Ornithopoda containing Hypsilophodon foxii and Tenontosaurus tilletti provided it does not include Iguanodon bernissartensis". All other analyses from around the same time have instead found these latter taxa to be within Iguanodontia. The family Hypsilophodontidae is formally defined in the PhyloCode by Daniel Madzia and colleagues in 2021 as "the largest clade containing Hypsilophodon foxii, but not Iguanodon bernissartensis and Rhabdodon priscus".
Altirhinus is a genus of hadrosauroid ornithopod dinosaur from the Early Cretaceous period of Mongolia.
Orodromeus is a genus of herbivorous orodromine thescelosaurid dinosaur from the Late Cretaceous of North America. Only one species is known, the type species Orodromeus makelai.
Lurdusaurus is a genus of massive and unusually shaped iguanodont dinosaur from the Elrhaz Formation in Niger. It contains one species, L. arenatus. The formation dates to the Early Cretaceous, roughly 112 million years ago. Lurdusaurus has a highly atypical body plan for an iguanodont, with a small skull, long neck, rotund torso, and powerful forelimbs and claws, somewhat reminiscent of a ground sloth. Its metacarpals are fused and reinforced into a large block, and the thumb spike is remarkably enormous. These would have allowed the hand to have functioned almost like a ball-and-chain flail. Lurdusaurus is estimated to have been 7–9 m (23–30 ft) long and 2 m high when on all-fours, but its stomach would have been only 70 cm off the ground. It may have weighed 2.5–5.5 t, conspicuously heavy for an iguanodontid this size.
Probactrosaurus is an early herbivorous hadrosauroid iguanodont dinosaur. It lived in China during the Early Cretaceous period.
Gasparinisaura is a genus of herbivorous ornithopod dinosaur from the Late Cretaceous.
Hadrosauroidea is a clade or superfamily of ornithischian dinosaurs that includes the "duck-billed" dinosaurs, or Hadrosauridae, and all dinosaurs more closely related to them than to Iguanodon. Their remains have been recovered in Asia, Europe, Africa and the Americas. Many primitive hadrosauroids, such as the Asian Probactrosaurus and Altirhinus, have traditionally been included in a paraphyletic "Iguanodontidae". With cladistic analysis, the traditional Iguanodontidae has been largely disbanded, and probably includes only Iguanodon and perhaps its closest relatives.
Cedrorestes is a genus of iguanodontian dinosaur from the Early Cretaceous of Utah. It is based on an incomplete skeleton which was found in the Valanginian-age Yellow Cat Member of the Cedar Mountain Formation.
Rhabdodontidae is a family of herbivorous iguanodontian ornithopod dinosaurs whose earliest stem members appeared in the middle of the Lower Cretaceous. The oldest dated fossils of these stem members were found in the Barremian Castrillo de la Reina Formation of Spain, dating to approximately 129.4 to 125.0 million years ago. With their deep skulls and jaws, Rhabdodontids were similar to large, robust iguanodonts. The family was first proposed by David B. Weishampel and colleagues in 2003. Rhabdodontid fossils have been mainly found in Europe in formations dating to the Late Cretaceous.
Owenodon is a genus of iguanodontian dinosaur known from a partial lower jaw discovered in Early Cretaceous-age rocks of Dorset, United Kingdom, and possibly also Romania and Spain. The first and only definitive specimen was found in the Lulworth Formation of the Purbeck Limestone Group, dating to the middle Berriasian stage. It was first described by Richard Owen as a species Iguanodon, I. hoggii, honouring naturalist A.J. Hogg who had originally collected the fossil. Owen described the mandible as it was, partially embedded in a limestone block, but it was given to the Natural History Museum, London where it was accessioned as NHMUK PV R 2998 and further prepared. Some damage occurred to a tooth crown and part of the bone while stored in the collections. Redescription of I. hoggii by David Norman and Paul Barrett subsequently transferred the species to Camptosaurus in 2002, as well as tentatively referring other camptosaur-like material from the Purbeck beds to the species. The identity of the species was questioned, with Kenneth Carpenter and Yvonne Wilson, and Greg Paul, separating "C." hoggi from Camptosaurus as an intermediate ornithopod, until Peter Galton named the new genus Owenodon for it in 2009. Galton removed the material assigned by Norman and Barrett from Owenodon, but referred isolated teeth from the Bauxite of Cornet, Romania, and the El Castellar Formation of Spain to O. hoggii. The taxon, believed by Galton to be intermediate between Camptosaurus and Iguanodon, is of uncertain relationships, with the limited material preventing clear understanding of its position within ornithopod evolution. Phylogenetic studies have found Owenodon to be more primitive, equivalent to, or more derived than Camptosaurus, but it is often excluded to improve results.
Bolong is a genus of iguanodontian dinosaur known from the Early Cretaceous-age Yixian Formation of western Liaoning Province, China. It lived about 125 million years ago in the earliest Aptian.
Hippodraco is a genus of iguanodontian ornithopod dinosaur from the Early Cretaceous Cedar Mountain Formation of Utah, United States. The genus contains a single species, H. scutodens, known from a partial skeleton belonging to an immature individual.
Iguanacolossus is a genus of iguanodontian ornithopod dinosaur that lived in North America during the Early Cretaceous period. It is known from UMNH VP 20205, the associated holotype with a large partial skeleton of a single individual.
Ratchasimasaurus is a genus of iguanodontian ornithopod dinosaur from Early Cretaceous Khok Kruat Formation of Nakhon Ratchasima Province in northeastern Thailand. The type and only species is R. suranareae, named after Thao Suranari, a 19th-century war heroine. It was considered by one study to be a nomen dubium, diagnosed with characters widespread in Styracosterna.
Rhabdodontomorpha is a clade of basal iguanodont dinosaurs. This group was named in 2016 in the context of the description, based on Spanish findings of an early member of the Rhabdodontidae. A cladistic analysis was conducted in which it was found that Muttaburrasaurus was the sister species of the Rhabdodontidae sensu Weishampel. Therefore, Paul-Emile Dieudonné, Thierry Tortosa, Fidel Torcida Fernández-Baldor, José Ignacio Canudo and Ignacio Díaz-Martínez defined Rhabdodontomorpha as a nodal clade: the group consisting of the last common ancestor of Rhabdodon priscus Matheron, 1869 and Muttaburrasaurus langdoni Bartholomai and Molnar, 1981; and all its descendants. Within the clade Zalmoxes and Mochlodon are also included. In 2021, Daniel Madzia redefined Rhabdodontomorpha in the PhyloCode as "the largest clade containing Rhabdodon priscus, but not Iguanodon bernissartensis and Hypsilophodon foxii". The clade is characterized by the following synapomorphies: