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Nodosaurids Temporal range: Late Jurassic–Late Cretaceous, | |
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Gargoyleosaurus skeleton cast | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Clade: | Dinosauria |
Clade: | † Ornithischia |
Clade: | † Thyreophora |
Clade: | † Ankylosauria |
Clade: | † Euankylosauria |
Family: | † Nodosauridae Marsh, 1890 |
Subgroups | |
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Synonyms | |
Acanthopholididae Nopcsa, 1902 Contents |
Nodosauridae is a family of ankylosaurian dinosaurs known from the Late Jurassic to the Late Cretaceous periods in what is now Asia, Europe, North America, and possibly South America. While traditionally regarded as a monophyletic clade as the sister taxon to the Ankylosauridae, some analyses recover it as a paraphyletic grade leading to the ankylosaurids.
Nodosaurids, like their sister group the ankylosaurids, were heavily armored dinosaurs adorned with rows of bony armor nodules and spines (osteoderms), which were covered in keratin sheaths. Nodosaurids, like other ankylosaurians, were small- to large-sized, heavily built, quadrupedal, herbivorous dinosaurs, possessing small, leaf-shaped teeth. Unlike ankylosaurids, nodosaurids lacked mace-like tail clubs, instead having more flexible tail tips. Many nodosaurids had spikes projecting outward from their shoulders. One particularly well-preserved nodosaurid "mummy", the holotype of Borealopelta markmitchelli , preserves a nearly complete set of armor in life position, as well as the keratin covering and mineralized remains of the underlying skin, which indicate reddish pigments in a countershading pattern. [1] [2]
The family Nodosauridae was erected by Othniel Charles Marsh in 1890, and anchored on the genus Nodosaurus . [3] [4]
The clade Nodosauridae was first informally defined by Paul Sereno in 1998 as "all ankylosaurs closer to Panoplosaurus than to Ankylosaurus ," a definition followed by Vickaryous, Teresa Maryańska, and Weishampel in 2004. Vickaryous et al. considered two genera of nodosaurids to be of uncertain placement (incertae sedis): Struthiosaurus and Animantarx, and considered the most primitive member of the Nodosauridae to be Cedarpelta. [5]
Following the publication of the PhyloCode, Nodosauridae needed to be formally defined following certain parameters, including that the type genus Nodosaurus was required as an internal specifier. In formally defining Nodosauridae, Madzia and colleagues followed the previously established use for the clade, defining it as the largest clade including Nodosaurus textilis but not Ankylosaurus magniventris . As all phylogenies referenced included both Panoplosaurus and Nodosaurus within the same group relative to Ankylosaurus, the addition of another internal specifier was deemed unnecessary.
Nodosauridae is traditionally composed of the basal clade Polacanthinae (sometimes recovered outside of the Nodosauridae), as well as the Panoplosaurini and Struthiosaurini within the Nodosaurinae. Nodosaurinae is defined in the PhyloCode as "the largest clade containing Nodosaurus textilis , but not Hylaeosaurus armatus , Mymoorapelta maysi , and Polacanthus foxii ". [6] Panoplosaurini is defined in the PhyloCode as "the largest clade containing Panoplosaurus mirus , but not Nodosaurus textilis and Struthiosaurus austriacus " while Struthiosaurini has a similar definition of "the largest clade containing Struthiosaurus austriacus , but not Nodosaurus textilis and Panoplosaurus mirus ". [6] Topology A below demonstrates these relationships, following the phylogenetic analyses of Rivera-Sylva and colleagues (2018), with clade names added by definition from Madzia et al. (2021). [7] [6] However, in 2023, Raven and colleagues proposed an alternate phylogeny for nodosaurids; instead of the traditional dichotomous split between nodosaurids and ankylosaurids, their analyses resulted in a paraphyletic grade of these taxa comprising the monophyletic clades Panoplosauridae, Polacanthidae and Struthiosauridae. These results are displayed in Topology B below. [8] [9] Corresponding clades are shown in matching colors for clarity, and ⊞ buttons can be clicked to expand nodes:
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Nodosaurinae is defined as the largest clade containing Nodosaurus textilis but not Hylaeosaurus armatus , Mymoorapelta maysi , or Polacanthus foxii , and was formally defined in 2021 by Madzia and colleagues, who utilized the name of Othenio Abel in 1919, who created the term to unite Ankylosaurus , Hierosaurus and Stegopelta . [6] [10] The name has been significantly refined in content since Abel first used it to unite all quadrupedal, plate-armoured ornithischians, [10] now including a significant number of taxa from the Early Cretaceous through Maastrichtian of Europe, North America, and Argentina. Previous informal definitions of the group described the clade as all taxa closer to Panoplosaurus , or Panoplosaurus and Nodosaurus, than to the early ankylosaurs Sarcolestes , Hylaeosaurus, Mymoorapelta or Polacanthus, which was reflected in the specifiers chosen by Madzia et al. when formalizing the clade following the PhyloCode. The 2018 phylogenetic analysis of Rivera-Sylva and colleagues was used as the primary reference for Panoplosaurini by Madzia et al., in addition to the supplemental analyses of Thompson et al. (2012), Arbour and Currie (2016), Arbour et al. (2016), and Brown et al. (2017). [6] [11] [12] [7] [13] [1]
Panoplosaurini is defined as the largest clade containing Panoplosaurus mirus , but not Nodosaurus textilis or Struthiosaurus austriacus , and was named in 2021 by Madzia and colleagues for the group found in many previous analyses, both morphological and phylogenetic. Panoplosaurini includes not only the Late Cretaceous Panoplosaurus, Denversaurus and Edmontonia , but also the mid Cretaceous Animantarx and Texasetes , as well as Patagopelta . However, in the study describing it, its authors only placed it as a nodosaurine outside Panoplosaurini. [14] The approximately equivalent clade Panoplosaurinae, named in 1929 by Franz Nopcsa, but was not significantly used until Robert Bakker reused the name in 1988, alongside the new clades Edmontoniinae and Edmontoniidae, which were considered to unite Panoplosaurus, Denversaurus and Edmontonia to the exclusion of other ankylosaurs. As none of the clades were commonly used, or formally named following the PhyloCode, Madzia et al. named Panoplosaurini instead, as the group of taxa fell within the clade Nodosaurinae, and having the same -inae suffix on both parent and child taxon could be confusing in future. [6] The 2018 phylogenetic analysis of Rivera-Sylva and colleagues was used as the primary reference for Panoplosaurini by Madzia et al., in addition to the supplemental analyses of Arbour et al. (2016), Brown et al. (2017), and Zheng et al. (2018). [6] [7] [13] [1] [15]
Struthiosaurini is defined as the largest clade containing Struthiosaurus austriacus , but not Nodosaurus textilis or Panoplosaurus mirus , and was named in 2021 by Madzia and colleagues for the relatively stable group found in many previous analyses. Struthiosaurini includes not only the Late Cretaceous European Struthiosaurus, but also the Early Cretaceous European Europelta , the Late Cretaceous European Hungarosaurus , and Stegopelta and Pawpawsaurus from the mid Cretaceous of North America. The approximately equivalent clade Struthiosaurinae, named in 1923 by Franz Nopcsa was previously used to include European nodosaurids, but was never formally named following the PhyloCode, so Madzia et al. named Struthiosaurini instead, as the group of taxa fell within the clade Nodosaurinae, and having the same -inae suffix on both parent and child taxon could be confusing in future. [6] The 2018 phylogenetic analysis of Rivera-Sylva and colleagues was used as the primary reference for Struthiosaurini by Madzia et al., in addition to the supplemental analyses of Arbour et al. (2016), Brown et al. (2017), and Zheng et al. (2018). [6] [7] [13] [1] [15]
Nodosaurids are known from diverse remains throughout Europe, Asia, and North America. [16]
Some Gondwanan ankylosaurs, including the Antarctican Antarctopelta and Argentinian Patagopelta , were originally regarded as belonging to the Nodosauridae, but later analyses provided support for them belonging to the Parankylosauria, a separate lineage of basal ankylosaurs restricted to the Southern Hemisphere. [17] [18] [19]
Ankylosauria is a group of herbivorous dinosaurs of the clade Ornithischia. It includes the great majority of dinosaurs with armor in the form of bony osteoderms, similar to turtles. Ankylosaurs were bulky quadrupeds, with short, powerful limbs. They are known to have first appeared in North Africa during the Middle Jurassic, and persisted until the end of the Late Cretaceous. The two main families of ankylosaurians, Nodosauridae and Ankylosauridae are primarily known from the Northern Hemisphere, but the more basal Parankylosauria are known from southern Gondwana during the Cretaceous.
Nodosaurus is a genus of herbivorous nodosaurid ankylosaurian dinosaur from the Late Cretaceous, the fossils of which are found exclusively in the Frontier Formation in Wyoming.
Minmi is a genus of small herbivorous ankylosaurian dinosaur that lived during the early Cretaceous Period of Australia, about 120 to 112 million years ago.
Edmontonia is a genus of panoplosaurin nodosaurid dinosaur from the Late Cretaceous Period. It is part of the Nodosauridae, a family within Ankylosauria. It is named after the Edmonton Formation, the unit of rock where it was found.
Ankylosauridae is a family of armored dinosaurs within Ankylosauria, and is the sister group to Nodosauridae. The oldest known ankylosaurids date to around 122 million years ago and went extinct 66 million years ago during the Cretaceous–Paleogene extinction event. These animals were mainly herbivorous and were obligate quadrupeds, with leaf-shaped teeth and robust, scute-covered bodies. Ankylosaurids possess a distinctly domed and short snout, wedge-shaped osteoderms on their skull, scutes along their torso, and a tail club.
Struthiosaurus is a genus of nodosaurid dinosaurs, from the Late Cretaceous period (Santonian-Maastrichtian) of Austria, Romania, France and Hungary in Europe. It was a small dinosaur, measuring 2–3 m (6.6–9.8 ft) in length and weighing 300–400 kg (660–880 lb).
Aletopelta is a monospecific genus of basal ankylosaurid dinosaur from Southern California that lived during the Late Cretaceous in what is now the Point Loma Formation. The type and only species, Aletopelta coombsi, is known from a partial skeleton preserving osteoderms. It was originally described in 1996 by W. P. Coombs, Jr. and T.A. Deméré before being named in 2001 by Tracy Ford and James Kirkland. Aletopelta has an estimated size of 5 metres and weight of 2 tonnes. The holotype formed a miniature reef and was scavenged upon by invertebrates and sharks.
Texasetes is a genus of ankylosaurian dinosaurs from the late Lower Cretaceous of North America. This poorly known genus has been recovered from the Paw Paw Formation near Haslet, Tarrant County, Texas, which has also produced the nodosaurid ankylosaur Pawpawsaurus.
Cedarpelta is an extinct genus of basal ankylosaurid dinosaur from Utah that lived during the Late Cretaceous period in what is now the Mussentuchit Member of the Cedar Mountain Formation. The type and only species, Cedarpelta bilbeyhallorum, is known from multiple specimens including partial skulls and postcranial material. It was named in 2001 by Kenneth Carpenter, James Kirkland, Don Burge, and John Bird. Cedarpelta has an estimated length of 7 metres and weight of 5 tonnes (11,023 lbs). The skull of Cedarpelta lacks extensive cranial ornamentation and is one of the only known ankylosaurs with individual skull bones that are not completely fused together.
Mymoorapelta is a nodosaurid ankylosaur from the Late Jurassic Morrison Formation of western Colorado and central Utah, USA. The animal is known from a single species, Mymoorapelta maysi, and few specimens are known. The most complete specimen is the holotype individual from the Mygatt-Moore Quarry, which includes osteoderms, a partial skull, vertebrae, and other bones. It was initially described by James Kirkland and Kenneth Carpenter in 1994. Along with Gargoyleosaurus, it is one of the earliest known nodosaurids.
Panoplosaurus is a genus of armoured dinosaur from the Late Cretaceous of Alberta, Canada. Few specimens of the genus are known, all from the middle Campanian of the Dinosaur Park Formation, roughly 76 to 75 million years ago. It was first discovered in 1917, and named in 1919 by Lawrence Lambe, named for its extensive armour, meaning "well-armoured lizard". Panoplosaurus has at times been considered the proper name for material otherwise referred to as Edmontonia, complicating its phylogenetic and ecological interpretations, at one point being considered to have existed across Alberta, New Mexico and Texas, with specimens in institutions from Canada and the United States. The skull and skeleton of Panoplosaurus are similar to its relatives, but have a few significant differences, such as the lumpy form of the skull osteoderms, a completely fused shoulder blade, and regularly shaped plates on its neck and body lacking prominent spines. It was a quadrupedal animal, roughly 5 m (16 ft) long and 1,600 kg (3,500 lb) in weight. The skull has a short snout, with a very domed surface, and bony plates directly covering the cheek. The neck had circular groups of plates arranged around the top surface, both the forelimb and hindlimb were about the same length, and the hand may have only included three fingers. Almost the entire surface of the body was covered in plates, osteoderms and scutes of varying sizes, ranging from large elements along the skull and neck, to smaller, round bones underneath the chin and body, to small ossicles that filled in the spaces between other, larger osteoderms.
Pawpawsaurus, meaning "Pawpaw Lizard", is a nodosaurid ankylosaur from the Cretaceous of Tarrant County, Texas, discovered in May 1992. The only species yet assigned to this taxon, Pawpawsaurus campbelli, is based on a complete skull from the marine Paw Paw Formation.
Polacanthinae is a subfamily of ankylosaurs, most often nodosaurids, from the Late Jurassic through Early Cretaceous of Europe and potentially North America and Asia. The group is defined as the largest clade closer to Polacanthus foxii than Nodosaurus textilis or Ankylosaurus magniventris, as long as that group nests within either Nodosauridae or Ankylosauridae. If Polacanthus, and by extent Polacanthinae, falls outside either family-level clade, then the -inae suffix would be inappropriate, and the proper name for the group would be the informally defined Polacanthidae.
Zhejiangosaurus is an extinct genus of ankylosaurian dinosaur from the Upper Cretaceous of Zhejiang, eastern China. It was first named by a group of Chinese authors Lü Junchang, Jin Xingsheng, Sheng Yiming and Li Yihong in 2007 and the type species is Zhejiangosaurus lishuiensis. It has no diagnostic features, and thus is a nomen dubium.
Ahshislepelta is a monospecific genus of ankylosaur dinosaur from New Mexico that lived during the Late Cretaceous in what is now the Hunter Wash Member of the Kirtland Formation. The type and only species, Ahshislepelta minor, is known only from an incomplete postcranial skeleton of a small subadult or adult individual. It was named in 2011 by Michael Burns and Robert M. Sullivan. Based on the size of the humerus, Ahshislepelta is larger than Pinacosaurus mephistocephalus but smaller than Talarurus and Pinacosaurus grangeri.
Dongyangopelta is an monospecific genus of nodosaurid dinosaur that lived in China during the Early to Late Cretaceous period in what is now the Chaochuan Formation. The type and only known species, Dongyangopelta yangyanensis, is known from a partial postcranial skeleton preserving osteoderms and ossified tendons. It was named in 2013 by Rongjun Chen, Wenjie Zheng, Yoichi Azuma, Masateru Shibata, Tianling Lou, Qiang Jin and Xinsheng Jin. Dongyangopelta represents one of the only nodosaurids known from Asia, along with Taohelong and Sauroplites.
This timeline of ankylosaur research is a chronological listing of events in the history of paleontology focused on the ankylosaurs, quadrupedal herbivorous dinosaurs who were protected by a covering bony plates and spikes and sometimes by a clubbed tail. Although formally trained scientists did not begin documenting ankylosaur fossils until the early 19th century, Native Americans had a long history of contact with these remains, which were generally interpreted through a mythological lens. The Delaware people have stories about smoking the bones of ancient monsters in a magic ritual to have wishes granted and ankylosaur fossils are among the local fossils that may have been used like this. The Native Americans of the modern southwestern United States tell stories about an armored monster named Yeitso that may have been influenced by local ankylosaur fossils. Likewise, ankylosaur remains are among the dinosaur bones found along the Red Deer River of Alberta, Canada where the Piegan people believe that the Grandfather of the Buffalo once lived.
Parankylosauria is a group of basal ankylosaurian dinosaurs known from the Cretaceous of South America, Antarctica, and Australia. It is thought the group split from other ankylosaurs during the mid-Jurassic period, despite this being unpreserved in the fossil record.
Patagopelta is an extinct genus of ankylosaurian dinosaur from the Late Cretaceous Allen Formation of Argentina. The genus contains a single species, P. cristata, known from a partial skeleton. While originally described as a nodosaurine, later discoveries provided support for parankylosaurian affinities for the taxon. Patagopelta is a very small ankylosaur, comparable in size to the dwarf nodosaurid Struthiosaurus, about 2 m (6.6 ft) long.
Vectipelta is an extinct genus of ankylosaurian dinosaur recovered from the Early Cretaceous Wessex Formation of England. The genus contains a single species, V. barretti, known from a partial skeleton including several osteoderms. It was historically known as the "Spearpoint ankylosaur" prior to its description.