Timeline of ankylosaur research

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.[1] 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.[2] 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.[3]

The first scientifically documented ankylosaur remains were recovered from Early Cretaceous rocks in England and named Hylaeosaurus armatus by Gideon Mantell in 1833.[4] However, the Ankylosauria itself would not be named until Henry Fairfield Osborn did so in 1923 nearly a hundred years later.[5] Prior to this, the ankylosaurs had been considered members of the Stegosauria, which included all armored dinosaurs when Othniel Charles Marsh named the group in 1877. It was not until 1927 that Alfred Sherwood Romer implemented the modern use of the name Stegosauria as specifically pertaining to the plate-backed and spike-tailed dinosaurs of the Jurassic that form the ankylosaurs' nearest relatives.[6] The next major revision to ankylosaur taxonomy would not come until Walter Coombs divided the group into the two main families paleontologists still recognize today; the nodosaurids and ankylosaurids.[5] Since then, many new ankylosaur genera and species have been discovered from all over the world and continue to come to light. Many fossil ankylosaur trackways have also been recognized.[7]

Euoplocephalus tutus
Skeletal mounts of the ankylosaur Scolosaurus

Prescientific

Osteoderms of Ankylosaurus
Osteoderms of Ankylosaurus
  • The Delaware people of what is now New Jersey or Pennsylvania had a tradition regarding a hunting party that returned with a piece of an ancient bone supposedly belonging to a monster that killed humans. One of the village's wise men instructed people to burn bits of the bone in clay spoons with tobacco and make a wish while the concoction was still smoking. This ritual could bestow such favors as success in hunting, long life, and health for one's children. This tale might be inspired by local fossils, which include ankylosaurs, Coelosaurus, Dryptosaurus, and Hadrosaurus.[1]
  • Traditional Navajo creation mythology portrays modern Earth as the most recent of a series of worlds. They believe that the earlier worlds were inhabited by monsters that were killed with lightning bolts wielded by the heroic Monster Slayers.[8] The most terrifying monster of the old worlds was the Big Gray Monster, Yeitso.[9] The Navajo of Arizona feared fossil remains, attributing them to his corpse. They believe that Yeitso's ghost still haunts his remains.[10] Yeitso's flint-like scales may have been inspired by the fossilized armored plates of various prehistoric creatures that once lived in what is now the western US. Ankylosaurs like Ankylosaurus are one such potential candidate for the source of Yeitso's armored hide. Others include non-ankylosaurs like the Permian amphibian Eryops, Triassic phytosaurs and Desmatosuchus, as well as other armored dinosaurs like Scutellosaurus or Stegosaurus.[2]
  • The Piegan people of Alberta attributed the fossils of dinosaurs to the "grandfather of the buffalo" they left offerings of cloth and tobacco to this mythical creature near the Red Deer River. Ankylosaur remains are among those preserved in the area that helped inspire this legend and associated practice, as are the remains of ceratopsians, hadrosaurs, and carnivorous theropods.[3]

19th century

Hylaeosaurus
Early artistic restoration of Hylaeosaurus armatus

1830s

1832

  • Quarry workers discovered a fossilized partial skeleton. The remains were sent to paleontologist Gideon Mantell, who recognized that they represented a significant scientific discovery.[11]
  • Mantell reported the specimen discovered by quarry workers that would later be formally named Hylaeosaurus to the Geological Society.[11]

1833

1840s

1842

Early April

  • Sir Richard Owen published his second report on British fossil reptiles, wherein he formally named the Dinosauria.[14] Hylaeosaurus was included as a founding member and was the third dinosaur to be named.[15]

1843

1844

1850s

1856

1858

  • Sir Richard Owen published a study on Hylaeosaurus.[17]

1860s

Polacanthus skeleton
Early skeletal reconstruction of Polacanthus foxii skeletal restoration by Franz Nopcsa von Felső-Szilvás

1865

  • Reverend William Fox discovered the Polacanthus type specimen.[18]

1867

1869

1870s

1871

1875

1879

1880s

Nodosaurus textilis
Early restoration of a Nodosaurus textilis skeleton

1881

1882

  • The British Museum of Natural History bought a large number of fossils from Rev. Fox, including the Polacanthus type specimen.[20]

1888

Palaeoscincus costatus
Illustration of the Palaeoscincus costatus holotype tooth

1889

1890s

1890

  • Marsh named the Nodosauridae.[21] He regarded them as relatives of the stegosaurs due to the shared presence of bony plates embedded in the skin.[5]

1892

1893

20th century

1900s

Ankylosaurus
Artistic skeletal reconstruction of Ankylosaurus (AMNH 5895) by Barnum Brown, 1908, before the tail club was known
Ankylosaurus magniventris reconstruction
Artistic restoration of Ankylosaurus magniventris

1901

1902

1905

1908

1909

1910s

Struthiosaurus Nopsca 1915
Early illustration of Struthiosaurus by Nopsca from 1915

1914

  • While collecting fossils in Dinosaur Provincial Park, William Edmund Cutler discovered the type specimen of an ankylosaur taxon that would later be named Scolosaurus cutleri in his honor. However, while undercutting the specimen it collapsed on him "resulting in serious upper body injuries."[23]

1915

1918

1919

1920s

Edmontonia restoration
Life restoration of two Edmontonia from 1922, based on the 1915 AMNH specimen

1923

1924

1927

  • Alfred Sherwood Romer published the first formal diagnosis for the Ankylosauria.[5] He observed that the anatomy of the stegosaur pelvis and hindlimb as well as their primarily Jurassic age distinguished them from the mainly Cretaceous ankylosaurs. As the Stegosauria originally included all armored dinosaurs, Romer's distinction marked the beginning of the modern use of the name to refer to the plate-backed and spike-tailed dinosaurs.[6]
Scolosaurus cutleri
Type specimen of Scolosaurus

1928

1929

1930s

Pinacosaurus
Skeletal reconstruction of Pinacosaurus

1930

1932

  • Sternberg described the new ichnogenus and species Tetrapodosaurus borealis from the Early Cretaceous Gething Formation of British Columbia, Canada. He attributed the tracks to ceratopsians, but they would later be attributed to ankylosaurs.[24]

1933

1934

1935

1936

1940s

1940

  • Russell concluded that ankylosaurs chewed with a simple straight-up-and-down movement of the jaws and only fed on soft vegetation based on aspects of their skull and tooth anatomy.[7]

1950s

Talarurus plicatospineus
Skeletal reconstruction of Talarurus plicatospineus

1952

1953

1955

1956

1960s

1960

1963

  • F. H. Khakimov discovered a new dinosaur track site in Shirkent National Park, Tajikistan.[25]
  • Zakharov and Khakimov reported the dinosaur track site discovered by the latter to the scientific literature.[25]

1964

  • Zakharov described the new ichnogenus and species Macropodosaurus gravis. He attributed it to a theropod, but these tracks are more likely to have been produced by ankylosaurs.[25]

1969

  • Haas interpreted the ankylosaur diet and consisting of soft plants that ankylosaurs chewed with a simple straight-up-and-down movement of the jaws based on their skull and tooth anatomy.[7]

1970s

Euoplocephalus Royal Tyrell
Skeletal mount of Scolosaurus thronus

1970

1971

  • Walter Coombs published landmark research into ankylosaur taxonomy, bringing order to a once "chaotic and confused" field of study. He recognized two main groups of ankylosaurs, the Ankylosauridae and Nodosauridae.[5] Coombs interpreted the ankylosaur diet as consisting of soft plants that ankylosaurs chewed with a simple straight-up-and-down movement of the jaws based on their skull and tooth anatomy.[7]
  • Haubold reported the presence of the ichnospecies Metatetrapous valdensis from the Buckeburg Formation of Germany. This ichnospecies is attributed to ankylosaurs.[26]

1972

  • Coombs observed that Euoplocephalus was so thoroughly armored that there was even a bony plate protecting its eyelids.[27]

1977

  • Teresa Maryanska described the new genus Tarchia for the species "Dyoplosaurus" giganteus. She also named the new species Tarchia kielanae and the new genus and species Saichania chulsanensis.[22] She followed the scheme proposed by Coombs earlier that decade dividing the ankylosaurs into ankylosaurids and nodosaurids.[5] She also made observations regarding ankylosaur limb posture, noting that while the hind limb was nearly straight up and down, the humerus was oriented at an angle downward and toward the rear of the animal. When studying the ankylosaur tail she noted that the centra of the vertebrae near its tip are fused, which would make it hard for the animal to raise the tail club very high.[7]

1978

  • Kurzanov and Tumanova described the new genus and species Amtosaurus magnus.[19]
  • Coombs published more work on ankylosaur taxonomy.[5] He noted that ankylosaurs were probably completely unable to walk on their hind legs and published further remarks on ankylosaur limb posture. He argued that while some researchers interpreted some aspects of ankylosaur forelimb anatomy as adaptations for digging, their hoof-like toe nails made this interpretation unlikely.[28]

1979

  • Coombs interpreted the bony tendons near the tip of the ankylosaur tail as a means to convey the forces generated by the tail musculature closer to the animal's body all the way down to its club.[7]

1980s

Sauropelta jconway
Life restoration of Sauropelta edwardsorum.

1980

1982

1983

  • Tumanova described the new genus and species Shamosaurus scutatus.[22]
  • Campbell reported the presence of dinosaur footprints in the Toro Toro Formation of Bolivia which he attributed to sauropods.[29]

1984

  • Kenneth Carpenter attributed the ichnogenus Tetrapodosaurus reported by Sternberg from British Columbia in the 1930s to ankylosaurs rather than ceratopsians.[24] He argued that the most likely trackmaker was Sauropelta.[30]
  • Leonardi described the dinosaur footprints reported by Campbell the previous year in detail and named them Ligabuichnium bolivianum. Rather than sauropods, Leonardi argued that these tracks were produced by ankylosaurs or ceratopsians although it was difficult ascertain which of these taxa were responsible due to the poor preservation of the tracks.[29]

1986

  • Galton interpreted the ankylosaur diet and consisting of soft plants that ankylosaurs chewed with a simple straight-up-and-down movement of the jaws based on their skull and tooth anatomy.[7]

1987

  • Paul Ensom described dinosaur footprints from the Purbeck Beds of England once thought to have been left by sauropods. They are now thought to have been left by ankylosaurs.[31]
  • Tumanova erected the new genus Maleevus to house the species Syrmosaurus disparoserratus.[16] Tumanova followed the scheme proposed by Coombs earlier that decade dividing the ankylosaurs into ankylosaurids and nodosaurids.[5]
  • Gasparini and others reported ankylosaur remains from Antarctica.[5]

1988

1989

  • Currie reported the discovery of a Tetrapodosaurus track from British Columbia. Although he could not confidently identify its stratigraphic origin, the rock preserving the tracks has since been attributed to the Dunvegan Formation.[32]
  • A worker at the Smoky River Coal Mine near Grande Cache, Alberta alerted the Royal Tyrell Museum to the presence of dinosaur footprints in the area. This site would come to be recognized as the most important ankylosaur track site in the world.[33]

1990s

Mymoorapelta maysi
Skeletal reconstruction of Mymoorapelta at the Wyoming Dinosaur Center

1990

  • Coombs and Maryanska remarked that the boney secondary palate of the ankylosaur skull would have strengthened it by acting as a brace.[7]
  • A well-preserved skeleton of Minmi was excavated from the Allaru Formation in Queensland, Australia by the Queensland Museum and catalogued as QM F18101. The skeleton was mostly articulated, including its armor.[34] Since most ankylosaur specimens do not preserve the life arrangement of their armor, QM F18101 represented a rare find.[35] The specimen also preserved the animal's gut contents, the first to be discovered in any armored dinosaur.[36]

1991

  • Lockley argued that the supposed sauropod tracks reported by Ensom from the Purbeck Beds of Dorset, England were actually made by ankylosaurs.[37]
  • Frank DeCourten discovered dinosaur tracks preserved in the Cedar Mountain Formation of Utah that were likely produced by ankylosaurs.[38]

1993

  • Tumanova described the new genus and species Tsagantegia longicranialis.[22]
  • Jerzykiewicz and others reported the presence of borings of unknown cause on the bones of some juvenile Pinacosaurus grangeri from Bayan Mandahu, Inner Mongolia, China.[39]
  • Z. Dong described the new genus and species Tianchisaurus nedegoaperferima.[16]
  • Thulborn proposed that the tail club of ankylosaurs may actually have functioned as a "false head" meant to distract predators. However, this hypothesis has not received much support from the paleontological community, and has been criticized as "dubiou[s]".[7]
  • Grady published an illustration of an ankylosaur trackway from the "Mine" site in the Smoky River Coal Mine at Grande Cache, Alberta.[40]

1994

  • Kirkland and Carpenter described the new genus and species Mymoorapelta maysi.[12]
  • Psihoyos and Knoebber described a dinosaur track site in the Smoky Hill Coal Mine of Grande Cache, Alberta and reported that the site had been destroyed in a rock slide.[41]
  • Whyte and Romano described the new ichnogenus and species Deltapodus brodericki for dinosaur footprints discovered in the Aalenian-Bajocian Saltwick Formation of Yorkshire, England. The authors attributed the tracks to sauropods, but they may actually have been made by ankylosaurs.[42]
  • Leonardi concluded that the Bolivian Ligabuichnium tracks were made by an ankylosaur after all.[29]

1995

  • Carpenter, Dilkes, and Weishampel erected the new genus Niobrarasaurus to house the species Hierosaurus coleii.[19]
  • Coombs studied the anatomy of the tail of Euoplocephalus and concluded that its club was held just slightly off the ground rather than dragging or held and a significant height. He reiterated observations previously made in 1977 by Maryanska that the fusion of the vertebral centra near the tip of the animal's tail would make it difficult to raise very high.[7]
  • Coombs described the new genus and species Texasetes pleurohalio.[19]

1996

  • Lee described the new genus and species Pawpawsaurus campbelli.[21]
  • Molnar reported the existence of a second species of Minmi but did not name it.[22]
Gastoniasaur
Skeletal reconstructions of Gastonia at the North American Museum of Ancient Life

1997

  • Witmer studied archosaur "craniofacial pneumaticity". He concluded that rather than performing a biological function, paranasal sinuses in archosaurs "are best explained as an optimization of skull architecture". This cast doubt on various researchers' interpretations of the sometimes complex nasal cavities and sinus systems possessed by ankylosaurs. Past workers had thought that these cavities and sinuses may have given ankylosaurs an improved sense of smell, housed glands, acted as a resonating chamber for loud vocalizations, or helped conserve body heat and moisture.[7]

1998

Tianzhenosaurus
Skeletal reconstruction of Tianzhenosaurus
  • Pang and Cheng described the new genus and species Tianzhenosaurus youngi.[22]
  • Barret and others described the new genus and species Shanxia tianzhensis.[16]
  • Sereno defined the ankylosaurs as all eurypods more closely related to Ankylosaurus than to Stegosaurus.[43]
  • McCrea and Currie described the dinosaur tracks discovered in the Smoky River Coal Mine at Grand Cache, Alberta. They noted that this was the most important ankylosaur track site ever discovered.[33]
  • McCrea and others reported the first known ankylosaur skin impression to be preserved in a footprint tracks preserved in the Dunvegan Formation near Pouce Coupe, Alberta.[44]

1999

21st century

2000s

2000

2001

  • Rybczynsky and Vickaryous studied the jaws and teeth of Euoplocephalus. Contrary to decades of support for ankylosaurs chewing with a simple straight-up-and-down movement, they noticed visible wear facets and microscopic grooves that could only be explained by relatively complex jaw movements.[7]
  • Barrett reported wear facets on the teeth of Tarchia.[7]
  • Molnar and Clifford described the gut contents preserved in a specimen of the Australian ankylosaur Minmi.[47] This specimen is catalogued by the Queensland Museum as QM F18101 and was excavated by the museum from near the Flinders River in 1990.[48] The stomach contents consisted of plant vascular tissue, fruiting bodies, seeds, and possible fern spores.[49] Molnar and Clifford described it as the most reliable evidence for the diet of an herbivorous dinosaur ever discovered.[50]
  • McCrea, Lockley, and Meyer observed that by this point in the history of ankylosaur research, ankylosaurs track fossils had been reported from North America, South America, Asia, and Europe. Most of these trackways were preserved in moist floodplain habitats where plant life was abundant.[7] They attributed the Metatetrapous valdensis tracks from the Buckeburg Formation of Germany reported by Haubold in 1971 to ankylosaurs.[26] They similarly argued that Macropodosaurus gravis of Tajikistan was produced by an ankylosaur rather than a theropod.[25] The authors reported a single possible ankylosaur footprint from the Dakota Group of Baca County, Colorado.[51] They also proposed that several footprint specimens collected from the Blackhawk Formation of Utah may have been ankylosaurian.[52]
  • Vickaryous and others described the new genus and species Gobisaurus domoculus.[22]
  • Carpenter and others described the new genus and species Cedarpelta bilbeyhallorum.[21]
  • Xu, Wang, and You described the new genus and species Liaoningosaurus paradoxus.[19]
Aletopelta coombsi
Artistic restoration of Aletopelta coombsi

2002

2003

  • Garcia and Pereda-Superbiola described the new species Struthiosaurus languedocensis.[12]
  • Vickaryous and Russell described the common ways ankylosaur skulls were distorted after death that could potentially confound anatomical interpretation. They noted a relationship between this tendency to suffer distortion and their unusual cranial traits like the fusion of the skull bones and their "embossing" cranial ornamentation.[39]

2004

Minotaurasaurus BW
Artistic restoration of Minotaurasaurus ramachandrani

2005

2006

2007

2008

Skeletal reconstruction of Europelta
Skeletal reconstruction of Europelta carbonensis

2009

2010s

2011

Ziapelta
Artistic restoration of Ziapelta sanjuanensis

2013

2014

2015

2017

Jinyunpelta NT
Life restoration of Jinyunpelta sinensis

2018

2019

See also

Footnotes

  1. ^ a b Mayor (2005); "Smoking the Monster's Bone: An Ancient Delaware Fossil Legend," pages 68–69.
  2. ^ a b Mayor (2005); "The Monsters," page 122.
  3. ^ a b Mayor (2005); "Blackfeet and Ojibwe Fossil Discoveries," page 292.
  4. ^ Sarjeant (1999); "Further Finds in England," pages 9–10.
  5. ^ a b c d e f g h i j Vickaryous, Maryanska, and Weishampel (2004); "Introduction", page 363.
  6. ^ a b Galton and Upchurch (2004); "Introduction", page 343.
  7. ^ a b c d e f g h i j k l m n o p q Vickaryous, Maryanska, and Weishampel (2004); "Paleoecology and Behavior", page 392.
  8. ^ Mayor (2005); "Fossils in Navajo Land," page 119.
  9. ^ Mayor (2005); "The Monsters," page 119.
  10. ^ Mayor (2005); "Fossils in Navajo Land," page 117.
  11. ^ a b Moore (2014); "1832" (1), page 31.
  12. ^ a b c d e f g h i j k l m n o p Vickaryous, Maryanska, and Weishampel (2004); "Table 17.1: Ankylosauria", page 366.
  13. ^ For Hylaeosaurus as the first ankylosaur, see Sarjeant (1999); "Further Finds in England," pages 9–10. For the date of the description of Ankylosauria, see Vickaryous, Maryanska, and Weishampel (2004); "Introduction", page 363.
  14. ^ Torrens (1999); "Politics and Paleontology", page 182.
  15. ^ Torrens (1999); "Politics and Paleontology", page 184.
  16. ^ a b c d e f g h i j k l m n o p q Vickaryous, Maryanska, and Weishampel (2004); "Table 17.1: Ankylosauria", page 368.
  17. ^ Moore (2014); "1858" (3), page 53.
  18. ^ Moore (2014); "1865" (3), page 61.
  19. ^ a b c d e f g h i j k l m n o p q r s t u v w Vickaryous, Maryanska, and Weishampel (2004); "Table 17.1: Ankylosauria", page 367.
  20. ^ Moore (2014); "1882" (1), page 91.
  21. ^ a b c d e f g h i j k l m n o Vickaryous, Maryanska, and Weishampel (2004); "Table 17.1: Ankylosauria", page 365.
  22. ^ a b c d e f g h i j k l m n o p Vickaryous, Maryanska, and Weishampel (2004); "Table 17.1: Ankylosauria", page 364.
  23. ^ Tanke (2010); "Background and Collection History," page 542.
  24. ^ a b McCrea, Lockley, and Meyer (2001); "Gething Formation, British Columbia (Aptian-Albian)", page 422.
  25. ^ a b c d McCrea, Lockley, and Meyer (2001); "Shirabad Suite, Tadjikistan (Albian)", page 433.
  26. ^ a b McCrea, Lockley, and Meyer (2001); "Wealden Beds, Germany (Berriasian)", pages 421-422.
  27. ^ Vickaryous, Russell, and Currie (2001); "Testing the Hypothesis", page 327.
  28. ^ Vickaryous, Maryanska, and Weishampel (2004); "Paleoecology and Behavior", pages 391–392.
  29. ^ a b c McCrea, Lockley, and Meyer (2001); "Torotoro Formation, Bolivia (Campanian)", page 442.
  30. ^ McCrea (2000); "Tetrapodosaurus borealis Sternberg, 1932", page 41.
  31. ^ McCrea, Lockley, and Meyer (2001); "Purbeck Beds, England (Berriasian)", page 421.
  32. ^ McCrea, Lockley, and Meyer (2001); "Dunvegan Formation, Alberta and Northeast British Columbia (Cenomanian)", page 437.
  33. ^ a b McCrea (2000); "1.2 Previous work on the Gates Formation", page 2.
  34. ^ Molnar (2001); "Introduction", page 342.
  35. ^ Molnar (2001); "Introduction", page 341.
  36. ^ Molnar and Clifford (2001); "Introduction", pages 399-400.
  37. ^ Lockley and Meyer (2000); "The First Ankylosaur Tracks," pages 182-183.
  38. ^ McCrea, Lockley, and Meyer (2001); "Cedar Mountain Formation, Utah (Albian-Cenomanian)", page 433.
  39. ^ a b c Vickaryous, Maryanska, and Weishampel (2004); "Taphonomy", page 391.
  40. ^ McCrea, Lockley, and Meyer (2001); "Gates Formation, Grande Cache, Alberta (Lower Albian)", page 429.
  41. ^ McCrea, Lockley, and Meyer (2001); "Gates Formation, Grande Cache, Alberta (Lower Albian)", page 423.
  42. ^ McCrea, Lockley, and Meyer (2001); "Saltwick Formation, England (Aalenian-Bajocian)", page 421.
  43. ^ Vickaryous, Maryanska, and Weishampel (2004); "Definition and Diagnosis", page 363.
  44. ^ McCrea, Lockley, and Meyer (2001); "Dunvegan Formation, Alberta and Northeast British Columbia (Cenomanian)", page 440.
  45. ^ McCrea, Lockley, and Meyer (2001); "Cedar Mountain Formation, Utah (Albian-Cenomanian)", pages 433-434.
  46. ^ McCrea, Lockley, and Meyer (2001); "Djadokhta Formation, Mongolia (Campanian)", page 441.
  47. ^ Molnar and Clifford (2001); "Abstract", page 399.
  48. ^ For date and location of discovery, see Molnar (2001); "Introduction", page 342. For catalogue number and stomach contents, see Molnar and Clifford (2001); "Introduction", page 399.
  49. ^ Molnar and Clifford (2001); "Description", page 401.
  50. ^ Molnar and Clifford (2001); "Introduction", page 400.
  51. ^ McCrea, Lockley, and Meyer (2001); "Dakota Group (Albian-Cenomanian)", page 435.
  52. ^ McCrea, Lockley, and Meyer (2001); "Blackhawk Formation, Utah (Campanian)", page 440.
  53. ^ Parish and Barrett (2004); "Abstract", page 299.
  54. ^ Ősi (2005); "Abstract", page 370.
  55. ^ Salgado and Gasparini (2006); "Abstract", page 199.
  56. ^ Lü et al. (2007); "Abstract", page 344.
  57. ^ Xu et al. (2007); "Abstract", page 433.
  58. ^ Carpenter et al. (2008); "Abstract", page 1089.
  59. ^ Burns, Michael E. (2008-12-12). "Taxonomic utility of ankylosaur (Dinosauria, Ornithischia) osteoderms: Glyptodontopelta mimus Ford, 2000: a test case". Journal of Vertebrate Paleontology. 28 (4): 1102–1109. doi:10.1671/0272-4634-28.4.1102. ISSN 0272-4634.
  60. ^ Miles and Miles (2009); "Abstract", page 65.
  61. ^ Parsons and Parsons (2009); "Abstract", page 721.
  62. ^ Arbour, Victoria M.; Burns, Michael E.; Sissons, Robin L. (2009-12-12). "A redescription of the ankylosaurid dinosaur Dyoplosaurus acutosquameus Parks, 1924 (Ornithischia: Ankylosauria) and a revision of the genus". Journal of Vertebrate Paleontology. 29 (4): 1117–1135. doi:10.1671/039.029.0405. ISSN 0272-4634.
  63. ^ Burns and Sullivan (2011); "Abstract", page 169.
  64. ^ Burns, Michael E.; Currie, Philip J.; Sissons, Robin L.; Arbour, Victoria M. (2011). "Juvenile specimens of Pinacosaurus grangeri Gilmore, 1933 (Ornithischia: Ankylosauria) from the Late Cretaceous of China, with comments on the specific taxonomy of Pinacosaurus". Cretaceous Research. 32 (2): 174–186. doi:10.1016/j.cretres.2010.11.007.
  65. ^ Stanford, Weishampel, and DeLeon (2011); "Abstract", page 916.
  66. ^ Chen et al. (2013); "Abstract", page 658.
  67. ^ Kirkland et al. (2013); "Abstract", page 1.
  68. ^ Penkalski (2013); "Abstract", page 617.
  69. ^ Yang et al. (2013); "Abstract", page 265.
  70. ^ Han et al. (2014); "Abstract", page 1.
  71. ^ Arbour and Currie (2015); "Abstract".
  72. ^ Arbour, Currie, and Badamgarav (2014); "Abstract", page 631.
  73. ^ Arbour et al. (2014); "Abstract", page 1.
  74. ^ Blows (2015); in passim.
  75. ^ Leahey et al. (2015); in passim.
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References

External links

Acantholipan

Acantholipan is a genus of herbivorous nodosaurid dinosaur from Mexico from the early Santonian age of the Late Cretaceous. It includes one species, Acantholipan gonzalezi.

Aletopelta

Aletopelta coombsi is an herbivorous ankylosaurian ornithischian dinosaur that during the Late Cretaceous lived in the area of what is now Southern California.

Bienosaurus

Bienosaurus (meaning "Bien's lizard") is a genus of thyreophoran dinosaur from the Lower Jurassic (probably Sinemurian) Lower Lufeng Formation in Yunnan Province in China.

Bissektipelta

Bissektipelta is a genus of herbivorous ankylosaurid dinosaur from the Upper Cretaceous of Uzbekistan. Bissektipelta is monospecific, containing only the species B. archibaldi.

Dongyangopelta

Dongyangopelta is an extinct genus of nodosaurid ankylosaurian dinosaur known from the "middle" Cretaceous Chaochuan Formation (Albian or Cenomanian stage) of Dongyang, Zhejiang Province, China. Dongyangopelta was first named by Rongjun Chen, Wenjie Zheng, Yoichi Azuma, Masateru Shibata, Tianliang Lou, Qiang Jin and Xingsheng Jin in 2013 and the type species is Dongyangopelta yangyanensis. It differs from Zhejiangosaurus, the second nodosaurid from southeast China, in the characters of presacral rod, ilium, and femur. Donyangopelta is distinguishable from Zhejiangosaurus only on the basis of the morphology of its pelvic shield.

Dyoplosaurus

Dyoplosaurus (meaning "double-armored lizard") is an extinct genus of ankylosaurid dinosaurs within the subfamily Ankylosaurinae. It is known from the lower levels of the Late Cretaceous Dinosaur Park Formation (latest middle Campanian stage, about 76.5 Ma ago) of Alberta, Canada. It contains a single species, Dyoplosaurus acutosquameus.

Invictarx

Invictarx is a genus of herbivorous nodosaurid dinosaur from New Mexico dating from the early Campanian epoch of the Late Cretaceous.

Mymoorapelta

Mymoorapelta ("Shield of Mygatt-Moore") is an ankylosaur from the Late Jurassic (Kimmeridgian-Tithonian) Morrison Formation (Brushy Basin Member) of western Colorado, USA. The taxon is known from portions of a disarticulated skull, parts of three different skeletons and other postcranial remains. It is present in stratigraphic zones 4 and 5 of the Morrison Formation.

Nodocephalosaurus

Nodocephalosaurus is a genus of herbivorous ankylosaurine ankylosaurid dinosaur from Upper Cretaceous (late Campanian stage) deposits of San Juan Basin, New Mexico. The holotype was recovered from the Late Campanian De-na-zin Member of the Kirtland Formation and consists of an incomplete skull. Nodocephalosaurus (Greek nodus = knob, kephale = head and sauros = lizard) is a monotypic genus, including only the type species, Nodocephalosaurus kirtlandensis. Dinosaurs like Nodocephalosaurus resembled Asian forms, and may be evidence for Asian dinosaurs migrating to North America in the Late Cretaceous.

Nodosaurinae

Nodosaurinae is a group of ankylosaurian dinosaurs named in 1919 by Othenio Abel.

Nodosaurus

Nodosaurus (meaning "knobbed lizard") is a genus of herbivorous ankylosaurian dinosaur from the Late Cretaceous, the fossils of which are found in North America.

Pawpawsaurus

Pawpawsaurus, meaning "Pawpaw Lizard", is a nodosaurid ankylosaur from the Cretaceous (late Albian) of Tarrant County, Texas, discovered in May 1992. The only species yet assigned to this taxon, Pawpawsaurus campbelli, is based on a complete skull (lacking mandibles) from the marine Paw Paw Formation (Wachita Group).

Sarcolestes

Sarcolestes (meaning "flesh robber") is an extinct genus of ankylosaurian ornithischian dinosaur from the Oxford Clay of England. The current type and only species is S. leedsi, and the holotype is a single partial left mandible. The genus and species were named in 1893 by Richard Lydekker, who thought they belonged to a theropod.

Shanxia

Shanxia is a genus of ankylosaurid dinosaur that lived during the upper Cretaceous Period. Its fossils were recovered and named after the Shanxi Province of China, and it is known only from scrappy remains found in river deposits. Based on the relative lengths of the femur and other leg bones, it probably reached a length of around 3.6 metres (12 ft).

Barrett et al. (1998) distinguished Shanxia from other ankylosaurs in having long and flattened triangle-shaped horns that project backward from the squamosal bones on either side of the rear portion of its skull at an angle of 145 degrees. However, Sullivan (1999) considered Shanxia a nomen dubium, possibly synonymous with the related ankylosaurid Tianzhenosaurus, arguing that the unique shape of the squamosal horns could be a product of individual variation, but Upchurch and Barrett (2000) reaffirmed the validity of Shanxia. In their systematic review of ankylosaurids, Arbour and Currie (2015) treated Shanxia as a junior synonym of Saichania.

Silvisaurus

Silvisaurus, from the Latin silva "woodland" and Greek sauros "lizard", is a nodosaurid ankylosaur from the middle Cretaceous period.

Texasetes

Texasetes (meaning "Texas resident") 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 (late Albian) near Haslet, Tarrant County, Texas, which has also produced the nodosaurid ankylosaur Pawpawsaurus. Texasetes is estimated to have been 2.5–3 m (8–10 ft) in length. It was named by Coombs in 1995.

Tianchisaurus

Tianchisaurus (meaning "heavenly pool lizard") is a genus of ankylosaurian dinosaur from the middle Jurassic Period (Bajocian–Bathonian stages) of China. If it actually belongs to the family Ankylosauridae as proposed by Dong Zhiming, it would be the earliest member of that family. Unlike other ankylosaurids, it lacked a bony club at the tip of its tail.

The type specimen was informally referred to as "Jurassosaurus" after the 1993 film Jurassic Park, and the species epithet nedegoapeferima is formed from the surnames of the film's main stars: Sam Neill, Laura Dern, Jeff Goldblum, Richard Attenborough, Bob Peck, Martin Ferrero, Ariana Richards, and Joseph Mazzello. Director Steven Spielberg, who has funded Chinese dinosaur research, proposed the name. Dong Zhiming ultimately discarded the genus name "Jurassosaurus" (which is now a nomen nudum) in favor of Tianchisaurus, but retained the species name honoring the actors.

Tianzhenosaurus

Tianzhenosaurus (Tianzhen + Greek sauros="lizard") is a genus of ankylosaurid dinosaurs discovered in Tianzhen County, at Kangdailiang near Zhaojiagou Village, in Shanxi Province, China, in the Late Cretaceous Huiquanpu Formation. Thus far, a virtually complete skull and postcranial skeleton have been assigned to the genus, which is monotypic (T. youngi Pang & Cheng, 1998).

This was a medium-sized ankylosaurian, the skull measuring 28 cm (11 in) in length, with a total body length around 4 m (13 ft).

Vickaryous et al. (2004) placed Tianzhenosaurus within the Ankylosauridae, nested as the sister group to Pinacosaurus. Some authors have suggested that Tianzhenosaurus is actually a junior synonym of Saichania chulsanensis.

Tsagantegia

Tsagantegia (; meaning "of Tsagan-Teg"; Tumanova, 1993) is a genus of medium-sized ankylosaurid dinosaur from the Late Cretaceous of Mongolia, during the Cenomanian stage.

The holotype specimen (GI SPS N 700/17), a complete skull, was recovered from the Bayan Shireh Formation (Cenomanian-Santonian), at the Tsagan-Teg ("White Mountain") locality, Dzun-Bayan, in the southeastern Gobi Desert, Mongolia. The genus is monotypic, including only the type species, T. longicranialis.

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