Thescelosauridae: Difference between revisions
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==Classification== |
==Classification== |
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{{Cladogram|align=left|caption=This [[cladistics|cladogram]] is from Brown ''et al.'', (2013).<ref name=Albertadromeus>{{Cite journal | last1 = Brown | first1 = C. M. | last2 = Evans | first2 = D. C. | last3 = Ryan | first3 = M. J. | last4 = Russell | first4 = A. P. | title = New data on the diversity and abundance of small-bodied ornithopods (Dinosauria, Ornithischia) from the Belly River Group (Campanian) of Alberta | doi = 10.1080/02724634.2013.746229 | journal = Journal of Vertebrate Paleontology | volume = 33 | issue = 3 | pages = 495–520 | year = 2013 }}</ref> |
{{Cladogram|align=left|caption=This [[cladistics|cladogram]] is from Brown ''et al.'', (2013).<ref name=Albertadromeus>{{Cite journal | last1 = Brown | first1 = C. M. | last2 = Evans | first2 = D. C. | last3 = Ryan | first3 = M. J. | last4 = Russell | first4 = A. P. | title = New data on the diversity and abundance of small-bodied ornithopods (Dinosauria, Ornithischia) from the Belly River Group (Campanian) of Alberta | doi = 10.1080/02724634.2013.746229 | journal = Journal of Vertebrate Paleontology | volume = 33 | issue = 3 | pages = 495–520 | year = 2013 | s2cid = 129160518 }}</ref> |
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|clades={{clade| style=font-size:85%;line-height:85% |
|clades={{clade| style=font-size:85%;line-height:85% |
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|label1='''Parksosauridae/[[Thescelosauridae]]''' |
|label1='''Parksosauridae/[[Thescelosauridae]]''' |
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|3=''[[Jeholosaurus]]'' |
|3=''[[Jeholosaurus]]'' |
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|4=''[[Thescelosaurus]]'' }} }} }} }} }} |
|4=''[[Thescelosaurus]]'' }} }} }} }} }} |
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Although [[Hypsilophodontidae]] was interpreted as a natural group in the early 1990s,<ref name=SN90>{{cite book |last=Sues |first=Hans-Dieter |author2=Norman, David B. |editor= Weishampel, David B. |editor2=Dodson, Peter |editor3=Osmólska, Halszka |title=The Dinosauria |edition=1st |year=1990 |publisher=University of California Press |location=Berkeley |isbn=978-0-520-06727-1 |pages=498–509 |chapter=Hypsilophodontidae, ''Tenontosaurus'', Dryosauridae}}</ref><ref name=WH92>{{cite journal |last=Weishampel |first=David B. |author-link=David B. Weishampel |author2=Heinrich, Ronald E. |year=1992 |title=Systematics of Hypsilophodontidae and Basal Iguanodontia (Dinosauria: Ornithopoda) |journal=Historical Biology |volume=6 |pages=159–184 |url=http://www.tandf.co.uk/journals/pdf/ghbi_06_01_01.pdf |access-date=2007-03-10 |doi=10.1080/10292389209380426 |issue=3 }}</ref> this hypothesis has fallen out of favor and Hypsilophodontidae has been found to be an unnatural family composed of a variety of animals more or less closely related to Iguanodontia ([[paraphyletic]]), with various small [[clade]]s of closely related taxa.<ref name=CABetal09>{{cite journal|last=Boyd|first=Clint A.|author2=Brown, Caleb M. |author3=Scheetz, Rodney D. |author4= Clarke, Julia A. |year=2009|title=Taxonomic revision of the basal neornithischian taxa ''Thescelosaurus'' and ''Bugenasaura''|journal=Journal of Vertebrate Paleontology|volume=29|issue=3|pages=758–770|doi=10.1671/039.029.0328}}</ref><ref name=Hanetal12>{{cite journal |last=Han |first=Feng-Lu |author2=Paul M. Barrett |author3=Richard J. Butler |author4=Xing Xu |year=2012 |title=Postcranial anatomy of ''Jeholosaurus shangyuanensis'' (Dinosauria, Ornithischia) from the Lower Cretaceous Yixian Formation of China |journal=Journal of Vertebrate Paleontology |volume=32 |issue=6 |pages=1370–1395 |doi=10.1080/02724634.2012.694385 }}</ref><ref name=NSWC04>{{cite book |last=Norman |first=David B. |author-link=David B. Norman |author2=Sues, Hans-Dieter |author3=Witmer, Larry M. |author4= Coria, Rodolfo A. |editor=Weishampel, David B. |editor2=Dodson, Peter |editor3=Osmólska, Halszka |title=The Dinosauria |url=https://archive.org/details/dinosauriandedit00weis |url-access=limited |edition=2nd |year= 2004|publisher=University of California Press |location=Berkeley |isbn=978-0-520-24209-8 |pages=[https://archive.org/details/dinosauriandedit00weis/page/n411 393]–412 |chapter=Basal Ornithopoda}}</ref><ref name=WJCN03/><ref name=VMK07>{{cite journal |last=Varricchio |first=David J. |author2=Martin, Anthony J. |author3= Katsura, Yoshihiro |year=2007 |title=First trace and body fossil evidence of a burrowing, denning dinosaur |journal=Proceedings of the Royal Society B: Biological Sciences |pmid=17374596 |volume=274 |issue=1616 |pmc=2176205 |pages=1361–1368 |doi=10.1098/rspb.2006.0443 |url=}}</ref> "Hypsilophodontidae" and "hypsilophodont" are better understood as informal terms for an [[evolutionary grade]], not a true clade. ''Thescelosaurus'' has been regarded as both very basal<ref name=WH92/> and very [[Synapomorphy|derived]]<ref name=NSWC04/> among the hypsilophodonts. One issue that has potentially interfered with classifying ''Thescelosaurus'' is that not all of the remains assigned to ''T. neglectus'' necessarily belong to it.<ref name=BUN08>{{cite journal |last=Butler |first=Richard J. |author2=Upchurch, Paul |author3=Norman, David B. |author3-link=David B. Norman |title=The phylogeny of the ornithischian dinosaurs|journal=Journal of Systematic Palaeontology |volume=6 |issue=1 |year=2008 |pages=1–40 |doi=10.1017/S1477201907002271}}</ref> Clint Boyd and colleagues found that while the clade ''Thescelosaurus'' included the genus ''Bugenasaura'' and the species that had been assigned to that genus, there were at least two and possibly three species within ''Thescelosaurus'', and several specimens previously assigned to ''T. neglectus'' could not yet be assigned to a species within the genus.<ref name=CABetal09/> It appears to be closely related to ''[[Parksosaurus]]'',<ref name=CABetal09/><ref name=NSWC04/><ref name=WJCN03>{{cite journal |last=Weishampel |first= David B. |author-link=David B. Weishampel |author2=Jianu, Coralia-Maria |author3=Csiki, Z. |author4= Norman, David B. |year=2003 |title=Osteology and phylogeny of ''Zalmoxes'' (n.g.), an unusual euornithopod dinosaur from the latest Cretaceous of Romania |journal=Journal of Systematic Palaeontology |volume=1 |issue=2 |pages=1–56 |doi=10.1017/S1477201903001032 }}</ref><ref name=CMBetal2011>{{cite journal |authors=Brown; Caleb M.; Boyd, Clint A.; and Russell, Anthony P. |year=2011 |title=A new basal ornithopod dinosaur (Frenchman Formation, Saskatchewan, Canada), and implications for late Maastrichtian ornithischian diversity in North America |journal=Zoological Journal of the Linnean Society |volume=163 |issue=4 |pages=1157–1198 |doi=10.1111/j.1096-3642.2011.00735.x |doi-access=free }}</ref><ref name=CMBetal13>{{cite journal |last=Brown |first=Caleb Marshall |author2=Evans, David C. |author3=Ryan, Michael J. |author4= Russell, Anthony P. |year=2013 |title=New data on the diversity and abundance of small-bodied ornithopods (Dinosauria, Ornithischia) from the Belly River Group (Campanian) of Alberta |journal=Journal of Vertebrate Paleontology |volume=33 |issue=3 |pages=495–520 |doi=10.1080/02724634.2013.746229}}</ref> although this relationship has been called into question.<ref name=herne2019>{{cite journal |
Although [[Hypsilophodontidae]] was interpreted as a natural group in the early 1990s,<ref name=SN90>{{cite book |last=Sues |first=Hans-Dieter |author2=Norman, David B. |editor= Weishampel, David B. |editor2=Dodson, Peter |editor3=Osmólska, Halszka |title=The Dinosauria |edition=1st |year=1990 |publisher=University of California Press |location=Berkeley |isbn=978-0-520-06727-1 |pages=498–509 |chapter=Hypsilophodontidae, ''Tenontosaurus'', Dryosauridae}}</ref><ref name=WH92>{{cite journal |last=Weishampel |first=David B. |author-link=David B. Weishampel |author2=Heinrich, Ronald E. |year=1992 |title=Systematics of Hypsilophodontidae and Basal Iguanodontia (Dinosauria: Ornithopoda) |journal=Historical Biology |volume=6 |pages=159–184 |url=http://www.tandf.co.uk/journals/pdf/ghbi_06_01_01.pdf |access-date=2007-03-10 |doi=10.1080/10292389209380426 |issue=3 }}</ref> this hypothesis has fallen out of favor and Hypsilophodontidae has been found to be an unnatural family composed of a variety of animals more or less closely related to Iguanodontia ([[paraphyletic]]), with various small [[clade]]s of closely related taxa.<ref name=CABetal09>{{cite journal|last=Boyd|first=Clint A.|author2=Brown, Caleb M. |author3=Scheetz, Rodney D. |author4= Clarke, Julia A. |year=2009|title=Taxonomic revision of the basal neornithischian taxa ''Thescelosaurus'' and ''Bugenasaura''|journal=Journal of Vertebrate Paleontology|volume=29|issue=3|pages=758–770|doi=10.1671/039.029.0328|s2cid=84273584}}</ref><ref name=Hanetal12>{{cite journal |last=Han |first=Feng-Lu |author2=Paul M. Barrett |author3=Richard J. Butler |author4=Xing Xu |year=2012 |title=Postcranial anatomy of ''Jeholosaurus shangyuanensis'' (Dinosauria, Ornithischia) from the Lower Cretaceous Yixian Formation of China |journal=Journal of Vertebrate Paleontology |volume=32 |issue=6 |pages=1370–1395 |doi=10.1080/02724634.2012.694385 |s2cid=86754247 }}</ref><ref name=NSWC04>{{cite book |last=Norman |first=David B. |author-link=David B. Norman |author2=Sues, Hans-Dieter |author3=Witmer, Larry M. |author4= Coria, Rodolfo A. |editor=Weishampel, David B. |editor2=Dodson, Peter |editor3=Osmólska, Halszka |title=The Dinosauria |url=https://archive.org/details/dinosauriandedit00weis |url-access=limited |edition=2nd |year= 2004|publisher=University of California Press |location=Berkeley |isbn=978-0-520-24209-8 |pages=[https://archive.org/details/dinosauriandedit00weis/page/n411 393]–412 |chapter=Basal Ornithopoda}}</ref><ref name=WJCN03/><ref name=VMK07>{{cite journal |last=Varricchio |first=David J. |author2=Martin, Anthony J. |author3= Katsura, Yoshihiro |year=2007 |title=First trace and body fossil evidence of a burrowing, denning dinosaur |journal=Proceedings of the Royal Society B: Biological Sciences |pmid=17374596 |volume=274 |issue=1616 |pmc=2176205 |pages=1361–1368 |doi=10.1098/rspb.2006.0443 |url=}}</ref> "Hypsilophodontidae" and "hypsilophodont" are better understood as informal terms for an [[evolutionary grade]], not a true clade. ''Thescelosaurus'' has been regarded as both very basal<ref name=WH92/> and very [[Synapomorphy|derived]]<ref name=NSWC04/> among the hypsilophodonts. One issue that has potentially interfered with classifying ''Thescelosaurus'' is that not all of the remains assigned to ''T. neglectus'' necessarily belong to it.<ref name=BUN08>{{cite journal |last=Butler |first=Richard J. |author2=Upchurch, Paul |author3=Norman, David B. |author3-link=David B. Norman |title=The phylogeny of the ornithischian dinosaurs|journal=Journal of Systematic Palaeontology |volume=6 |issue=1 |year=2008 |pages=1–40 |doi=10.1017/S1477201907002271|s2cid=86728076 }}</ref> Clint Boyd and colleagues found that while the clade ''Thescelosaurus'' included the genus ''Bugenasaura'' and the species that had been assigned to that genus, there were at least two and possibly three species within ''Thescelosaurus'', and several specimens previously assigned to ''T. neglectus'' could not yet be assigned to a species within the genus.<ref name=CABetal09/> It appears to be closely related to ''[[Parksosaurus]]'',<ref name=CABetal09/><ref name=NSWC04/><ref name=WJCN03>{{cite journal |last=Weishampel |first= David B. |author-link=David B. Weishampel |author2=Jianu, Coralia-Maria |author3=Csiki, Z. |author4= Norman, David B. |year=2003 |title=Osteology and phylogeny of ''Zalmoxes'' (n.g.), an unusual euornithopod dinosaur from the latest Cretaceous of Romania |journal=Journal of Systematic Palaeontology |volume=1 |issue=2 |pages=1–56 |doi=10.1017/S1477201903001032 |s2cid= 86339025 }}</ref><ref name=CMBetal2011>{{cite journal |authors=Brown; Caleb M.; Boyd, Clint A.; and Russell, Anthony P. |year=2011 |title=A new basal ornithopod dinosaur (Frenchman Formation, Saskatchewan, Canada), and implications for late Maastrichtian ornithischian diversity in North America |journal=Zoological Journal of the Linnean Society |volume=163 |issue=4 |pages=1157–1198 |doi=10.1111/j.1096-3642.2011.00735.x |doi-access=free }}</ref><ref name=CMBetal13>{{cite journal |last=Brown |first=Caleb Marshall |author2=Evans, David C. |author3=Ryan, Michael J. |author4= Russell, Anthony P. |year=2013 |title=New data on the diversity and abundance of small-bodied ornithopods (Dinosauria, Ornithischia) from the Belly River Group (Campanian) of Alberta |journal=Journal of Vertebrate Paleontology |volume=33 |issue=3 |pages=495–520 |doi=10.1080/02724634.2013.746229|s2cid=129160518 }}</ref> although this relationship has been called into question.<ref name=herne2019>{{cite journal | title=New small-bodied ornithopods (Dinosauria, Neornithischia) from the Early Cretaceous Wonthaggi Formation (Strzelecki Group) of the Australian-Antarctic rift system, with revision of Qantassaurus intrepidus Rich and Vickers-Rich, 1999 | last1=Herne | first1=Matthew C. | last2=Nair | first2=Jay P. | last3=Evans | first3=Alistair R. | last4=Tait | first4=Alan M. | journal=Journal of Paleontology | year=2019 | volume=93 | issue=3 | pages=543–584 | doi=10.1017/jpa.2018.95| doi-access=free }}</ref> |
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[[File:Thescelosaurus skeleton.jpg|thumb|left|Head and arms of ''Thescelosaurus'', [[Rocky Mountain Dinosaur Resource Center]]]] |
[[File:Thescelosaurus skeleton.jpg|thumb|left|Head and arms of ''Thescelosaurus'', [[Rocky Mountain Dinosaur Resource Center]]]] |
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Revision as of 10:14, 8 December 2021
| Parksosaurids Temporal range: Early–Late Cretaceous,
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| Mounted specimen of Thescelosaurus, Burpee Museum of Natural History | |
Scientific classification 
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| Domain: | Eukaryota |
| Kingdom: | Animalia |
| Phylum: | Chordata |
| Clade: | Dinosauria |
| Clade: | †Ornithischia |
| Clade: | †Neornithischia |
| Clade: | †Ornithopoda |
| Family: | †Parksosauridae Bucholtz, 2002 |
| Subfamilies[1] | |
 | It has been suggested that Thescelosauridae be merged into this article. (Discuss) Proposed since November 2021. |
Parksosauridae is a clade or family of small ornithischians which have previously been generally allied to hypsilophodontids. Parksosauridae might be a synonym of Thescelosauridae since they possibly contain the same members.[2]
Classification
| |||
| This cladogram is from Brown et al., (2013).[1] |
Although Hypsilophodontidae was interpreted as a natural group in the early 1990s,[3][4] this hypothesis has fallen out of favor and Hypsilophodontidae has been found to be an unnatural family composed of a variety of animals more or less closely related to Iguanodontia (paraphyletic), with various small clades of closely related taxa.[5][6][7][8][9] "Hypsilophodontidae" and "hypsilophodont" are better understood as informal terms for an evolutionary grade, not a true clade. Thescelosaurus has been regarded as both very basal[4] and very derived[7] among the hypsilophodonts. One issue that has potentially interfered with classifying Thescelosaurus is that not all of the remains assigned to T. neglectus necessarily belong to it.[10] Clint Boyd and colleagues found that while the clade Thescelosaurus included the genus Bugenasaura and the species that had been assigned to that genus, there were at least two and possibly three species within Thescelosaurus, and several specimens previously assigned to T. neglectus could not yet be assigned to a species within the genus.[5] It appears to be closely related to Parksosaurus,[5][7][8][11][12] although this relationship has been called into question.[13]
The dissolution of Hypsilophodontidae has been followed by the recognition of the distinct family Parksosauridae by Buchholz in 2002, defined as the most inclusive clade containing Parksosaurus warreni, but not Hypsilophodon foxii, Dryosaurus altus, or Iguanodon bernissartensis.[14][2] Boyd et al. (2009) and Brown et al. (2011) found North American "hypsilophodonts" of Cretaceous age to sort into two related clusters, one (Orodrominae) consisting of Orodromeus, Oryctodromeus, and Zephyrosaurus, and the other (Thescelosaurinae) consisting of Parksosaurus and Thescelosaurus.[5][11] Brown et al. (2013) recovered similar results, with the addition of the new genus Albertadromeus to the Orodromeus clade (called Orodrominae) and several long-snouted Asian forms (previously described under Jeholosauridae)[6] to the Thescelosaurus clade (called Thescelosaurinae).[12] They also formally defined Thescelosauridae (Thescelosaurus neglectus, Orodromeus makelai, their most recent common ancestor, and all descendants) and the smaller clades Orodrominae and Thescelosaurinae.[12]
A 2015 analysis by Clint Boyd recovered Parksosauridae outside Ornithopoda, as the sister taxon to Cerapoda.[2] However, a 2019 analysis by Herne et al., which placed Parksosaurus as the most basal member of Ornithopoda, would effectively restrict Parksosauridae to only having the one member, Parksosaurus, as well as reviving Thescelosauridae as a separate group.[13] Below is the cladogram by Herne et al. (2019):
References
- ^ a b Brown, C. M.; Evans, D. C.; Ryan, M. J.; Russell, A. P. (2013). "New data on the diversity and abundance of small-bodied ornithopods (Dinosauria, Ornithischia) from the Belly River Group (Campanian) of Alberta". Journal of Vertebrate Paleontology. 33 (3): 495–520. doi:10.1080/02724634.2013.746229. S2CID 129160518.
- ^ a b c Boyd, Clint A. (2015). "The systematic relationships and biogeographic history of ornithischian dinosaurs". PeerJ. 3 (e1523): e1523. doi:10.7717/peerj.1523. PMC 4690359. PMID 26713260. Retrieved 5 January 2016.
{{cite journal}}: CS1 maint: unflagged free DOI (link) - ^ Sues, Hans-Dieter; Norman, David B. (1990). "Hypsilophodontidae, Tenontosaurus, Dryosauridae". In Weishampel, David B.; Dodson, Peter; Osmólska, Halszka (eds.). The Dinosauria (1st ed.). Berkeley: University of California Press. pp. 498–509. ISBN 978-0-520-06727-1.
- ^ a b Weishampel, David B.; Heinrich, Ronald E. (1992). "Systematics of Hypsilophodontidae and Basal Iguanodontia (Dinosauria: Ornithopoda)" (PDF). Historical Biology. 6 (3): 159–184. doi:10.1080/10292389209380426. Retrieved 2007-03-10.
- ^ a b c d Boyd, Clint A.; Brown, Caleb M.; Scheetz, Rodney D.; Clarke, Julia A. (2009). "Taxonomic revision of the basal neornithischian taxa Thescelosaurus and Bugenasaura". Journal of Vertebrate Paleontology. 29 (3): 758–770. doi:10.1671/039.029.0328. S2CID 84273584.
- ^ a b Han, Feng-Lu; Paul M. Barrett; Richard J. Butler; Xing Xu (2012). "Postcranial anatomy of Jeholosaurus shangyuanensis (Dinosauria, Ornithischia) from the Lower Cretaceous Yixian Formation of China". Journal of Vertebrate Paleontology. 32 (6): 1370–1395. doi:10.1080/02724634.2012.694385. S2CID 86754247.
- ^ a b c Norman, David B.; Sues, Hans-Dieter; Witmer, Larry M.; Coria, Rodolfo A. (2004). "Basal Ornithopoda". In Weishampel, David B.; Dodson, Peter; Osmólska, Halszka (eds.). The Dinosauria (2nd ed.). Berkeley: University of California Press. pp. 393–412. ISBN 978-0-520-24209-8.
- ^ a b Weishampel, David B.; Jianu, Coralia-Maria; Csiki, Z.; Norman, David B. (2003). "Osteology and phylogeny of Zalmoxes (n.g.), an unusual euornithopod dinosaur from the latest Cretaceous of Romania". Journal of Systematic Palaeontology. 1 (2): 1–56. doi:10.1017/S1477201903001032. S2CID 86339025.
- ^ Varricchio, David J.; Martin, Anthony J.; Katsura, Yoshihiro (2007). "First trace and body fossil evidence of a burrowing, denning dinosaur". Proceedings of the Royal Society B: Biological Sciences. 274 (1616): 1361–1368. doi:10.1098/rspb.2006.0443. PMC 2176205. PMID 17374596.
- ^ Butler, Richard J.; Upchurch, Paul; Norman, David B. (2008). "The phylogeny of the ornithischian dinosaurs". Journal of Systematic Palaeontology. 6 (1): 1–40. doi:10.1017/S1477201907002271. S2CID 86728076.
- ^ a b "A new basal ornithopod dinosaur (Frenchman Formation, Saskatchewan, Canada), and implications for late Maastrichtian ornithischian diversity in North America". Zoological Journal of the Linnean Society. 163 (4): 1157–1198. 2011. doi:10.1111/j.1096-3642.2011.00735.x.
{{cite journal}}: Unknown parameter|authors=ignored (help) - ^ a b c Brown, Caleb Marshall; Evans, David C.; Ryan, Michael J.; Russell, Anthony P. (2013). "New data on the diversity and abundance of small-bodied ornithopods (Dinosauria, Ornithischia) from the Belly River Group (Campanian) of Alberta". Journal of Vertebrate Paleontology. 33 (3): 495–520. doi:10.1080/02724634.2013.746229. S2CID 129160518.
- ^ a b Herne, Matthew C.; Nair, Jay P.; Evans, Alistair R.; Tait, Alan M. (2019). "New small-bodied ornithopods (Dinosauria, Neornithischia) from the Early Cretaceous Wonthaggi Formation (Strzelecki Group) of the Australian-Antarctic rift system, with revision of Qantassaurus intrepidus Rich and Vickers-Rich, 1999". Journal of Paleontology. 93 (3): 543–584. doi:10.1017/jpa.2018.95.
- ^ Buchholz, P. W. (2002). "Phylogeny and biogeography of basal Ornithischia". In Brown, D. E. (ed.). The Mesozoic in Wyoming. Casper: Tate Geological Museum. pp. 18–34.
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