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'''''Liaoningosaurus''''' is an unusual genus of [[ankylosauria]]n [[dinosaur]]s from the [[Early Cretaceous]] period of [[China]]. It contains a single species, '''''Liaoningosaurus paradoxus''''', and is represented by two fossil specimens collected from the [[Yixian Formation]] ([[Aptian]] age) of [[Liaoning]] Province. ''L. paradoxus'' was unusual among advanced [[ornithischian]] dinosaurs in that it is speculated to have hunted or scavenged, with preserved gut contents showing that it may have eaten fish. Additionally, some features of its skeleton may suggest that it was partially [[Aquatic animal|aquatic]].<ref name=liaoningosaurus_fish>Ji Q., Wu X., Cheng Y., Ten F., Wang X., and Ji Y. 2016. Fish-hunting ankylosaurs (Dinosauria, Ornithischia) from the Cretaceous of China. ''Journal of Geology'', '''40'''(2).</ref>
'''''Liaoningosaurus''''' is an unusual genus of [[ankylosauria]]n [[dinosaur]]s from the [[Early Cretaceous]] period of [[China]]. It contains a single species, '''''Liaoningosaurus paradoxus''''', and is represented by two fossil specimens collected from the [[Yixian Formation]] ([[Aptian]] age) of [[Liaoning]] Province. ''L. paradoxus'' was unusual among advanced [[ornithischian]] dinosaurs in that it is speculated to have hunted or scavenged, with preserved gut contents showing that it may have eaten fish. Additionally, some features of its skeleton may suggest that it was partially [[Aquatic animal|aquatic]].<ref name=liaoningosaurus_fish>Ji Q., Wu X., Cheng Y., Ten F., Wang X., and Ji Y. 2016. Fish-hunting ankylosaurs (Dinosauria, Ornithischia) from the Cretaceous of China. ''Journal of Geology'', '''40'''(2).</ref>


The [[type species]] ''L. paradoxus'' was named by [[Xu Xing (paleontologist)|Xu Xing]], Wang Xiaolin and You Hailu in 2001. The generic name refers to Liaoning. The [[specific name (zoology)|specific name]] refers to the confusing mix of nodosaurid and ankylosaurid features shown by the specimen.
The [[type species]] ''L. paradoxus'' was named in 2001. The generic name refers to Liaoning. The [[specific name (zoology)|specific name]] refers to the confusing mix of nodosaurid and ankylosaurid features shown by the specimen.


== Discovery ==
== Discovery ==
The [[holotype]] [[Institute of Vertebrate Paleontology and Paleoanthropology|IVPP]] V12560 is an articulated skeleton measuring approximately {{convert|34|cm|ftin}} in length. It was discovered in the [[Early Cretaceous]] [[Yixian Formation]] in [[Liaoning]] Province, China, and was formally described as ''Liaoningosaurus paradoxus'' in 2001. The [[generic name (biology)|generic name]] is after Liaoning Province, and the [[specific name (zoology)|specific name]] in reference to a suite of highly unusual traits for an [[ankylosaur]]. It was originally interpreted as a juvenile in based on the unfused [[vertebra]]l centra and neural arches.<ref name=holotype/>
The [[holotype]] [[Institute of Vertebrate Paleontology and Paleoanthropology|IVPP]] V12560 is an articulated skeleton measuring approximately {{convert|34|cm|ftin}} in length. It was discovered in the [[Early Cretaceous]] [[Yixian Formation]] in [[Liaoning]] Province, China, and was formally described as ''Liaoningosaurus paradoxus'' in 2001 by Chinese paleontologists [[Xu Xing (paleontologist)|Xu Xing]], Wang Xiaolin, and You Hai-lu. The [[generic name (biology)|generic name]] is after Liaoning Province, and the [[specific name (zoology)|specific name]] in reference to a suite of highly unusual traits for an [[ankylosaur]]. It was originally interpreted as a juvenile in based on the unfused [[vertebra]]l centra and neural arches.<ref name=holotype/>


In 2016, paleontologist Ji Qiang suggested that the holotype was actually fully grown, and unusual characteristics are aquatic adaptions. If correct, this would make ''Liaoningosaurus'' the smallest known [[ornithischia]]n dinosaur and the only known aquatic or semi-aquatic armoured dinosaur. In 2018, a [[histology|microanatomical]] study by Chinese palaeontologist Wenjie Zheng instead concluded that the holotype was no older than 12 months. In 2019, Li Xiaobo and Robert R. Reisz investigated three more specimens and also concluded they represent juveniles. About 20 specimens of ''Liaoningosaurus'' have been collected from four different localities of the Yixian Formation, and one locality from the nearby [[Jiufotang Formation]]. Many of them remain unstudied.<ref name=Li2019>{{cite conference|last1=Li|first1=X.|last2=Reisz|first2=R. R.|year=2019|title=The early Cretaceous ankylosaur ''Liaoningosaurus'' from Western Liaoning, China; Progress and problems|conference=7th Annual meeting Canadian Society of Vertebrate Palaeontology|date=May 10-13, 2019|pages=31–32}}</ref>
In 2016, paleontologist Ji Qiang suggested that the holotype was actually fully grown, and unusual characteristics are aquatic adaptions. If correct, this would make ''Liaoningosaurus'' the smallest known [[ornithischia]]n dinosaur and the only known aquatic or semi-aquatic armoured dinosaur. In 2018, a [[histology|microanatomical]] study by Chinese palaeontologist Wenjie Zheng instead concluded that the holotype was no older than 12 months. In 2019, Li Xiaobo and Robert R. Reisz investigated three more specimens and also concluded they represent juveniles. About 20 specimens of ''Liaoningosaurus'' have been collected from four different localities of the Yixian Formation, and one locality from the nearby [[Jiufotang Formation]]. Many of them remain unstudied.<ref name=Li2019>{{cite conference|last1=Li|first1=X.|last2=Reisz|first2=R. R.|year=2019|title=The early Cretaceous ankylosaur ''Liaoningosaurus'' from Western Liaoning, China; Progress and problems|conference=7th Annual meeting Canadian Society of Vertebrate Palaeontology|date=May 10-13, 2019|pages=31–32}}</ref>


==Description==
==Description==
The specimen is unique among all known ankylosaur fossils in the retention of the external [[mandibular fenestra]]. [[Antorbital fenestra]]e may also be present. It has relatively large teeth, including teeth in the praemaxilla (primitive or possibly juvenile trait). The points on the tooth crown were unusually long and sharp, giving each tooth a fork-like shape. It had long feet, long lower legs, and long, sharp claws on the hands and feet, unlike the blunt claws of other ankylosaurians. All of these were initially interpreted by the original describers as juvenile features, given the specimen's small size and lack of fusion between the spine and hip bones.<ref name=holotype>Xu X., Wang X.-L., and You H.-L. (2001). "A juvenile ankylosaur from China". ''Naturwissenschaften'', '''88'''(7): 297-300.</ref>
The specimen is unique among all known ankylosaur fossils in the retention of the external [[mandibular fenestra]]. [[Antorbital fenestra]]e may also be present. It has relatively large teeth, including teeth in the praemaxilla (primitive or possibly juvenile trait). The points on the tooth crown were unusually long and sharp, giving each tooth a fork-like shape. It had long feet, long lower legs, and long, sharp claws on the hands and feet, unlike the blunt claws of other ankylosaurians. All of these were initially interpreted by the original describers as juvenile features, given the specimen's small size and lack of fusion between the spine and hip bones.<ref name=holotype>Xu X., Wang X.-L., and You H.-L. (2001). "A juvenile ankylosaur from China". ''Naturwissenschaften'', '''88'''(7): 297-300. {{doi|10.1007/s001140100233}}.</ref>


Based on a second specimen, Ji and colleagues instead argued that these traits are adaptations to a semiaquatic lifestyle. The large, fork-like teeth and sharp claws would have been adapted to catch fish and other small animals. Stomach contents recovered with the second ''L. paradoxus'' specimen show that fish may have formed part of the animal's diet.<ref name="liaoningosaurus_fish"/> This indicates that ''Liaoningosaurus'' could have been either carnivorous or at least omnivorous, making it the first non-herbivorous ornithischian ever discovered, though the behavior has also been suspected in more basal ornithischians like [[heterodontosaurs]]<ref>{{Cite web | url=http://www.eartharchives.org/articles/armored-dinosaur-was-a-fish-eating-turtle-mimic/ | title=Armored dinosaur was a fish-eating turtle-mimic}}</ref> and [[Pachycephalosauria|pachycephalosaurids]]
Based on a second specimen, Ji and colleagues instead argued that these traits are adaptations to a semiaquatic lifestyle. The large, fork-like teeth and sharp claws would have been adapted to catch fish and other small animals. Stomach contents recovered with the second ''L. paradoxus'' specimen show that fish may have formed part of the animal's diet.<ref name="liaoningosaurus_fish"/> This indicates that ''Liaoningosaurus'' could have been either carnivorous or at least omnivorous, making it the first non-herbivorous ornithischian ever discovered, though the behavior has also been suspected in more basal ornithischians like [[heterodontosaurs]]<ref>{{Cite web | url=http://www.eartharchives.org/articles/armored-dinosaur-was-a-fish-eating-turtle-mimic/ | title=Armored dinosaur was a fish-eating turtle-mimic}}</ref> and [[Pachycephalosauria|pachycephalosaurids]]
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[[File:Liaoningosaurus.jpg|thumb|Reconstruction of ''Liaoningosaurus'' as a semi-aquatic, turtle-like dinosaur]]
[[File:Liaoningosaurus.jpg|thumb|Reconstruction of ''Liaoningosaurus'' as a semi-aquatic, turtle-like dinosaur]]
Triangular osteoderms were found on the shoulder, including a small shoulder spike.<ref name=holotype/> Ji and colleagues also identified armor plating on the belly, specifically a large segment of what they identified as a flat [[osteoderm]] as it is dotted with [[hexagonal]] and [[rhombus|rhombic]] bumps. Belly plates had never been found with other [[ankylosauria]]ns, so they believed the evolution of this trait [[convergent evolution|converged]] with [[turtle]]s to protect the underside when swimming.<ref name=liaoningosaurus_fish/> In 2013, paleontologist [[Victoria Arbour]] and colleagues analyzed the supposed osteoderm and interpreted it as normal dinosaur scales (that is, ''Liaoningosaurus'' did not have belly plates).<ref name=ankylosaur_skin>Arbour, V. M., Burns, M. E., Bell, P. R., & Currie, P. J. (2014). [https://www.researchgate.net/profile/Phil_Bell/publication/257535382_Epidermal_and_dermal_integumentary_structures_of_ankylosaurian_dinosaurs/links/00b7d52547876e446b000000.pdf Epidermal and dermal integumentary structures of ankylosaurian dinosaurs]. ''Journal of Morphology'', '''275'''(1): 39-50. {{doi|10.1002/jmor.20194}}</ref>
Triangular osteoderms were found on the shoulder, including a small shoulder spike. Xu identified an expansive belly plate ([[osteoderm]]), unlike any other ankylosaurian; those with belly armor have unfused, isolated osteoderms instead of this shield-like structure.<ref name=holotype/> The [osteoderm as it is dotted with [[hexagonal]] and [[rhombus|rhombic]] bumps. Ji and colleagues believed the evolution of this plate [[convergent evolution|converges]] with [[turtle]]s to protect the underside when swimming.<ref name=liaoningosaurus_fish/> In 2013, paleontologist [[Victoria Arbour]] and colleagues analyzed the supposed osteoderm and interpreted it as normal dinosaur scales (that is, ''Liaoningosaurus'' did not have a belly plate).<ref name=ankylosaur_skin>Arbour, V. M., Burns, M. E., Bell, P. R., & Currie, P. J. (2014). [https://www.researchgate.net/profile/Phil_Bell/publication/257535382_Epidermal_and_dermal_integumentary_structures_of_ankylosaurian_dinosaurs/links/00b7d52547876e446b000000.pdf Epidermal and dermal integumentary structures of ankylosaurian dinosaurs]. ''Journal of Morphology'', '''275'''(1): 39-50. {{doi|10.1002/jmor.20194}}</ref>


== Relationships ==
== Classification ==
Determining the relations of ''Liaoningosaurus'' is difficult because of the young age of the individual specimen and the presence of traits both typical of ankylosaurids and nodosaurids. Xu ''et al.'' published a [[cladistic]] analysis in 2001, placing ''Liaoningosaurus'' as an early branch within the [[Nodosauridae]],<ref name=holotype/> while Vickaryous ''et al.'' provisionally assigned it to the [[Ankylosauria]] ''[[incertae sedis]]'' in 2004.<ref>Vickaryous, [[Teresa Maryańska|Maryanska]], and Weishampel (2004). "Ankylosauria". in ''The Dinosauria'' (2nd edition), Weishampel, D. B., Dodson, P., and Osmólska, H., editors. University of California Press.</ref> A second [[cladistic analysis]] performed by Thompson ''et al.'' in 2011 suggested that ''Liaoningosaurus'' is a very [[Basal (phylogenetics)|basal]] [[ankylosaurid]].<ref name=ThompsonEtAl>{{Cite journal |author1=Richard S. Thompson |author2=Jolyon C. Parish |author3=Susannah C. R. Maidment |author4=Paul M. Barrett |year=2011 |title=Phylogeny of the ankylosaurian dinosaurs (Ornithischia: Thyreophora) |journal=Journal of Systematic Palaeontology |volume=10 |issue=2 |pages= 301–312|doi=10.1080/14772019.2011.569091 |s2cid=86002282 }}</ref> In 2018, Zheng suggested that ''Liaoningosaurus'' could be the juvenile form of the much larger ''[[Chuanqilong]]'' discovered in 2014 from the Jiufotang Formation.<ref name=Li2019/>
Given the animal's unusual anatomy, Xu and colleagues had trouble identifying its relationship to other ankylosaurs. Their phylogenic analysis loosely placed it into [[Nodosauridae]] (as opposed to [[Ankylosauridae]]). They conceded this binary division of Ankylosauria may not be entirely sound, meaning ''Liaoningosaurus'' could represent some third lineage.<ref name=holotype/> In 2004, paleontologist Matthew Vickaryous and colleagues left it under ''[[incertae sedis]]''.<ref>Vickaryous, [[Teresa Maryańska|Maryanska]], and Weishampel (2004). "Ankylosauria". in ''The Dinosauria'' (2nd edition), Weishampel, D. B., Dodson, P., and Osmólska, H., editors. University of California Press.</ref> A second [[cladistic analysis]] performed by Richard Thompson and colleagues in 2011 suggested that ''Liaoningosaurus'' is a [[Basal (phylogenetics)|basal]] [[ankylosaurid]].<ref name=ThompsonEtAl>{{Cite journal |author1=Richard S. Thompson |author2=Jolyon C. Parish |author3=Susannah C. R. Maidment |author4=Paul M. Barrett |year=2011 |title=Phylogeny of the ankylosaurian dinosaurs (Ornithischia: Thyreophora) |journal=Journal of Systematic Palaeontology |volume=10 |issue=2 |pages= 301–312|doi=10.1080/14772019.2011.569091 |s2cid=86002282 }}</ref> In 2018, Zheng suggested that ''Liaoningosaurus'' could be the juvenile form of the much larger ''[[Chuanqilong]]'' discovered in 2014 from the Jiufotang Formation.<ref name=Li2019/>


== See also ==
== See also ==

Revision as of 05:00, 24 August 2021

Liaoningosaurus
Temporal range: Early Aptian, 122 Ma
Liaoningosaurus holotype
Scientific classification Edit this classification
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Clade: Dinosauria
Clade: Ornithischia
Clade: Thyreophora
Clade: Ankylosauria
Clade: Euankylosauria
Family: Ankylosauridae
Genus: Liaoningosaurus
Xu et al., 2001
Species:
L. paradoxus
Binomial name
Liaoningosaurus paradoxus
Xu et al., 2001

Liaoningosaurus is an unusual genus of ankylosaurian dinosaurs from the Early Cretaceous period of China. It contains a single species, Liaoningosaurus paradoxus, and is represented by two fossil specimens collected from the Yixian Formation (Aptian age) of Liaoning Province. L. paradoxus was unusual among advanced ornithischian dinosaurs in that it is speculated to have hunted or scavenged, with preserved gut contents showing that it may have eaten fish. Additionally, some features of its skeleton may suggest that it was partially aquatic.[1]

The type species L. paradoxus was named in 2001. The generic name refers to Liaoning. The specific name refers to the confusing mix of nodosaurid and ankylosaurid features shown by the specimen.

Discovery

The holotype IVPP V12560 is an articulated skeleton measuring approximately 34 centimetres (1 ft 1 in) in length. It was discovered in the Early Cretaceous Yixian Formation in Liaoning Province, China, and was formally described as Liaoningosaurus paradoxus in 2001 by Chinese paleontologists Xu Xing, Wang Xiaolin, and You Hai-lu. The generic name is after Liaoning Province, and the specific name in reference to a suite of highly unusual traits for an ankylosaur. It was originally interpreted as a juvenile in based on the unfused vertebral centra and neural arches.[2]

In 2016, paleontologist Ji Qiang suggested that the holotype was actually fully grown, and unusual characteristics are aquatic adaptions. If correct, this would make Liaoningosaurus the smallest known ornithischian dinosaur and the only known aquatic or semi-aquatic armoured dinosaur. In 2018, a microanatomical study by Chinese palaeontologist Wenjie Zheng instead concluded that the holotype was no older than 12 months. In 2019, Li Xiaobo and Robert R. Reisz investigated three more specimens and also concluded they represent juveniles. About 20 specimens of Liaoningosaurus have been collected from four different localities of the Yixian Formation, and one locality from the nearby Jiufotang Formation. Many of them remain unstudied.[3]

Description

The specimen is unique among all known ankylosaur fossils in the retention of the external mandibular fenestra. Antorbital fenestrae may also be present. It has relatively large teeth, including teeth in the praemaxilla (primitive or possibly juvenile trait). The points on the tooth crown were unusually long and sharp, giving each tooth a fork-like shape. It had long feet, long lower legs, and long, sharp claws on the hands and feet, unlike the blunt claws of other ankylosaurians. All of these were initially interpreted by the original describers as juvenile features, given the specimen's small size and lack of fusion between the spine and hip bones.[2]

Based on a second specimen, Ji and colleagues instead argued that these traits are adaptations to a semiaquatic lifestyle. The large, fork-like teeth and sharp claws would have been adapted to catch fish and other small animals. Stomach contents recovered with the second L. paradoxus specimen show that fish may have formed part of the animal's diet.[1] This indicates that Liaoningosaurus could have been either carnivorous or at least omnivorous, making it the first non-herbivorous ornithischian ever discovered, though the behavior has also been suspected in more basal ornithischians like heterodontosaurs[4] and pachycephalosaurids [5]

Reconstruction of Liaoningosaurus as a semi-aquatic, turtle-like dinosaur

Triangular osteoderms were found on the shoulder, including a small shoulder spike. Xu identified an expansive belly plate (osteoderm), unlike any other ankylosaurian; those with belly armor have unfused, isolated osteoderms instead of this shield-like structure.[2] The [osteoderm as it is dotted with hexagonal and rhombic bumps. Ji and colleagues believed the evolution of this plate converges with turtles to protect the underside when swimming.[1] In 2013, paleontologist Victoria Arbour and colleagues analyzed the supposed osteoderm and interpreted it as normal dinosaur scales (that is, Liaoningosaurus did not have a belly plate).[6]

Classification

Given the animal's unusual anatomy, Xu and colleagues had trouble identifying its relationship to other ankylosaurs. Their phylogenic analysis loosely placed it into Nodosauridae (as opposed to Ankylosauridae). They conceded this binary division of Ankylosauria may not be entirely sound, meaning Liaoningosaurus could represent some third lineage.[2] In 2004, paleontologist Matthew Vickaryous and colleagues left it under incertae sedis.[7] A second cladistic analysis performed by Richard Thompson and colleagues in 2011 suggested that Liaoningosaurus is a basal ankylosaurid.[8] In 2018, Zheng suggested that Liaoningosaurus could be the juvenile form of the much larger Chuanqilong discovered in 2014 from the Jiufotang Formation.[3]

See also

References

  1. ^ a b c Ji Q., Wu X., Cheng Y., Ten F., Wang X., and Ji Y. 2016. Fish-hunting ankylosaurs (Dinosauria, Ornithischia) from the Cretaceous of China. Journal of Geology, 40(2).
  2. ^ a b c d Xu X., Wang X.-L., and You H.-L. (2001). "A juvenile ankylosaur from China". Naturwissenschaften, 88(7): 297-300. doi:10.1007/s001140100233.
  3. ^ a b Li, X.; Reisz, R. R. (May 10–13, 2019). The early Cretaceous ankylosaur Liaoningosaurus from Western Liaoning, China; Progress and problems. 7th Annual meeting Canadian Society of Vertebrate Palaeontology. pp. 31–32.{{cite conference}}: CS1 maint: date and year (link) CS1 maint: date format (link)
  4. ^ "Armored dinosaur was a fish-eating turtle-mimic".
  5. ^ Pickrell, John (2018-08-24). "Vegetarian dinosaur may have actually eaten meat, skull suggests". National Geographic. Retrieved 2018-09-08.
  6. ^ Arbour, V. M., Burns, M. E., Bell, P. R., & Currie, P. J. (2014). Epidermal and dermal integumentary structures of ankylosaurian dinosaurs. Journal of Morphology, 275(1): 39-50. doi:10.1002/jmor.20194
  7. ^ Vickaryous, Maryanska, and Weishampel (2004). "Ankylosauria". in The Dinosauria (2nd edition), Weishampel, D. B., Dodson, P., and Osmólska, H., editors. University of California Press.
  8. ^ Richard S. Thompson; Jolyon C. Parish; Susannah C. R. Maidment; Paul M. Barrett (2011). "Phylogeny of the ankylosaurian dinosaurs (Ornithischia: Thyreophora)". Journal of Systematic Palaeontology. 10 (2): 301–312. doi:10.1080/14772019.2011.569091. S2CID 86002282.