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'''Dyoplosaurus''' (meaning “double-armoured lizard”) is a monospecific [[genus]] of [[ankylosauridae|ankylosaurid]] [[dinosaur]] from [[Alberta]] that lived during the [[Late Cretaceous]] (middle [[Campanian]], ~76.5–75 Ma) in what is now the [[Dinosaur Park Formation]].<ref name=PWA24/> ''Dyoplosaurus'' represents a close relative of ''[[Scolosaurus]]'' and ''[[Anodontosaurus]]'', two ankylosaurids known from the [[Horseshoe Canyon Formation|Horseshoe Canyon]] and Dinosaur Park Formation.<ref name=PWA24>{{cite journal |last=Parks |first=W. A. |year=1924 |title=''Dyoplosaurus acutosquameus'', a new genus and species of armored dinosaur; and notes on a skeleton of ''Prosaurolophus maximus'' |journal=University of Toronto Studies Geological Series |volume=18 |pages=1–35}}</ref><ref name=ABS09>{{Cite journal | last1 = Arbour | first1 = V. M. | last2 = Burns | first2 = M. E. | last3 = Sissons | first3 = R. L. | doi = 10.1671/039.029.0405 | title = A redescription of the ankylosaurid dinosaur ''Dyoplosaurus acutosquameus'' Parks, 1924 (Ornithischia: Ankylosauria) and a revision of the genus | journal = Journal of Vertebrate Paleontology | volume = 29 | issue = 4 | pages = 1117 | year = 2009 }}</ref>
'''''Dyoplosaurus''''' (meaning "double-armored lizard") is an [[extinct]] [[genus]] of [[ankylosaurid]] [[dinosaur]]s 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'''''.<ref name=ABS09/>


==History of discovery==
==History of discovery==
[[File:Dinosaur_Park_Fm.jpg|thumb|left|Dinosaur Park Formation in Alberta, Canada]]
''Dyoplosaurus'' was named by [[William Parks (paleontologist)|William Parks]] in 1924, based on [[holotype]] ROM 784, a partial [[skeleton]] including parts of the [[skull]] and [[lower jaw]]s. It was collected from the bottom 10 m of the [[Dinosaur Park Formation]], near what is now the [[Red Deer River]] in [[Alberta]], [[Canada]].
The [[holotype]] specimen was obtained in 1919 from the bottom 10 metres of the [[Dinosaur Park Formation]], near what is now the [[Red Deer River]] in [[Alberta]], [[Canada]].<ref name=PWA24/><ref name=ABS09/> The holotype specimen, '''ROM 784''', consists of a partial skull roof, mandible fragments with teeth, osteoderms, skin impressions, articulated post-thoracic vertebrae, partial thoracic ribs, a partial ilium, both ischia, tail club, associated radius, metacarpal, femur, tibia, fibula, and pes.<ref name=PWA24/><ref name=ABS09/> The holotype is currently housed at the [[Royal Ontario Museum]].<ref name=PWA24/><ref name=ABS09/> Two specimens were referred to ''Dyoplosaurus'', ROM 7761 and UA 47273, and both consist of partial tail clubs.<ref name=Platypelta>{{Cite journal|author=Paul Penkalski |year=2018 |title=Revised systematics of the armoured dinosaur ''Euoplocephalus'' and its allies |journal=Neues Jahrbuch für Geologie und Paläontologie - Abhandlungen |volume=287 |issue=3 |pages=261–306 |doi=10.1127/njgpa/2018/0717 }}</ref>

The generic name, ''Dyoplosaurus'', is derived from the [[Ancient Greek|Greek]] words “dyo” (double), “hoplon” (weapon, shield, armour) and “sauros” (lizard).<ref name=PWA24/> The specific name, ''acutosquameus'', is derived from the [[Latin]] words “acutus” (sharp) and “squama” (scale).<ref name=PWA24/>

In 1956, [[Evgeny Maleev]] named a second [[species]] of ''Dyoplosaurus'': ''D. giganteus''.<ref name=Arbour2014>Victoria M. Arbour, Philip J. Currie and Demchig Badamgarav, 2014, "The ankylosaurid dinosaurs of the Upper Cretaceous Baruungoyot and Nemegt formations of Mongolia", Zoological Journal of the Linnean Society 172(3): 631–652|doi=https://doi.org/10.1111/zoj12185</ref> The species was based on PIN 551/29, which consists of a series of caudal vertebrae, metatarsals, phalanges and osteoderms including tail club knob from the [[Nemegt Formation]] of [[Mongolia]].<ref name=Arbour2014/> The species was diagnosed based on the short anterior caudal vertebrae; large chevrons that were fused to the caudal vertebrae; low, long distal caudal vertebrae; short, wide metatarsals; thick, hoofed-shaped unguals; sharp, thin-walled osteoderms with numerous pits and channels on the external surface.<ref name=Arbour2014/> However, a 2014 study by [[Victoria Arbour|Arbour]] and colleagues considered that the holotype lacked diagnostic traits, as such traits are present in all [[ankylosaurinae|ankylosaurines]], and considered the species as a [[nomen dubium]].<ref name=Arbour2014/>


==Description==
==Description==
{{multiple image
[[File:Ossified tendons in Ankylosaurus.png|thumb|left|Tail of the holotype specimen]]
|align = left
The type specimen of ''Dyoplosaurus'' was about {{convert|4|-|4.5|m|ft|abbr=on}} long<ref name="2017desc">{{cite journal |first1=V.M. |last1=Arbour |first2=J.C. |last2=Mallon |year=2017 |title=Unusual cranial and postcranial anatomy in the archetypal ankylosaur ''Ankylosaurus magniventris'' |journal=FACETS |volume=2 |issue=2 |pages=764–794 |doi=10.1139/facets-2017-0063 |doi-access=free }}</ref> and about 5.8 feet (1.7 m) wide, with a skull measuring 14&nbsp;inches (35 centimeters) across. Like most other ankylosaurids, it was quadrupedal, ground-dwelling and herbivorous, with an armored body and a bony club on the end of its tail. The club was composed of several armor plates fused together.<ref name=PWA24>{{cite journal |last=Parks |first=W. A. |year=1924 |title=''Dyoplosaurus acutosquameus'', a new genus and species of armored dinosaur; and notes on a skeleton of ''Prosaurolophus maximus'' |journal=University of Toronto Studies Geological Series |volume=18 |pages=1–35}}</ref> The tail club possessed by ''Dyoplosaurus'' is relatively small and narrow, when compared to that of other ankylosaurids.
|perrow=1/1
|total_width = 230

|image1 = Ossified_tendons_in_Ankylosaurus.png
|alt1 =

|image2 = Ankylosaurus_tail_terminology.png
|alt2 =
|caption2 = Tail of the holotype specimen and reconstruction of the tail and pelvis.
}}
Dyoplosaurus has an estimated body length of 3.58 to 4.16 metres (13 to 15 feet) and weight of 1500 to 2500 kg (3300 to 5500 lbs).<ref name="Miyashita2011">{{cite journal|vauthors=Miyashita T, Arbour VM, Witmer LM, Currie PJ |year=2011|title=The internal cranial morphology of an armoured dinosaur ''Euoplocephalus'' corroborated by X-ray computed tomographic reconstruction|journal=Journal of Anatomy|volume=219|issue=6|pages=661–75|doi=10.1111/j.1469-7580.2011.01427.x|pmid=21954840|pmc=3237876}}</ref> All referred specimens represent almost fully mature individuals.<ref name=Arbor2009>{{cite journal|author=Arbour, V. M.|year=2009|title=Estimating Impact Forces of Tail Club Strikes by Ankylosaurid Dinosaurs|journal=PLOS ONE|volume=4|issue=8|page=e6738|doi=10.1371/journal.pone.0006738|pmid=19707581|pmc=2726940|bibcode=2009PLoSO...4.6738A|doi-access=free}}.</ref>

''Dyoplosaurus'' can be distinguished from all other ankylosaurids in having sacral ribs that are anterolaterally-directed, triangular unguals in dorsal view and a tail club knob that is longer than wide.<ref name=ABS09/> It differs from ''Scolosaurus'' in having a proportionately shorter postacetabular process of the ilium and triangular osteoderms on the lateral sides of the anterior portion of the tail.<ref name=ABS09/> ''Dyoplosaurus'' also differs from ''Euoplocephalus'' in the pelvis as it has anterolaterally projecting, ventrally directed sacral transverse processes on the third sacral vertebra, forming a butterfly-like arrangement of the sacral fenestrae, and in having ischia that articulate with the ilia at right angles.<ref name=ABS09/>
[[File:Ankylosaurid_tail_clubs.png|170px|thumb|right|Tail club of Dyoplosaurus compared to other ankylosaurid tail clubs]]
The holotype specimen of ''Dyoplosaurus'' preserves ossified tendons on the tail club.<ref name=Arbor2009/><ref name=PWA24/> The tail club preserves three series of tendons on the dorsolateral sides of the handle, and four on the distal, ventral side of the tail.<ref name=Arbor2009/><ref name=PWA24/> These tendons are more readily grouped into two sets on the dorsolateral sides.<ref name=Arbor2009/><ref name=PWA24/> Arbour, 2009 conducted a study to determine the impact force of ankylosaurids and used ROM 784. Arbour found that ''Dyoplosaurus'' could generate an impact force of 797–1127 N and a more realistic tensions of 571 N, an impact force that isn’t enough to puncture bone.<ref name=Arbor2009/> This is mainly due to the knob being smaller in comparison to that of other ankylosaurids.<ref name=Arbor2009/> The small size of the tail club of suggests that ankylosaurid knobs were not primarily used as defensive weapons, as a weapon that is not functional until very late in life would probably not have a selective advantage over a weapon that is of use earlier in life.<ref name=Arbor2009/>

The holotype specimen preserves remnants of fossilized skin and osteoderms on the left side.<ref name=ABS09/> These skin impressions are punctuated by a nearly unbroken mosaic of small (0.50–1.0 cm), sub-angular to subrounded osteoderms.<ref name=ABS09/> The skin imperssions on the right side have been lost via erosion.<ref name=ABS09/>


==Classification==
==Classification==
In 1971, Walter Coombs synonymized ''Dyoplosaurus'', along with ''[[Scolosaurus]]'' and ''[[Anodontosaurus]]'', into ''[[Euoplocephalus]]'' as one of the four mandibles assigned to ''Dyoplosaurus'' was identical to those of other ''Euoplocephalus'' specimens, but did not offer any other characteristics to support the synonymization.<ref>{{Cite journal|last1=Coombs|first1=Walter|date=1971|title=The Ankylosauria|journal=Columbia University}}</ref> However, a re-description of ''Dyoplosaurus'' published in 2009 by [[Victoria Arbour]], Michael Burns and Robin Sissons considered it as a valid [[taxon]] and proposed that the synonymy was due to the fragmentary nature of the holotype and other referred specimens of ''Euoplocephalus''.<ref name=ABS09/> Thompson et al., 2011 confirmed its separation and recovered it as sister taxon to [[Pinacosaurus|Pinacosaurus mephistocephalus]].<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> A cladistic analysis conducted by Arbour and [[Phillip J. Currie|Currie]], 2015 recovered ''Dyoplosaurus'' as sister taxon to a clade containing ''[[Ankylosaurus]]'', ''Euoplocephalus'', ''Anodontosaurus'' and ''Scolosaurus'', while an analysis conducted by Arbour and Evans, 2017 recovered it as sister taxon to ''[[Zuul]]''.<ref name="systematics ankylosaurid">{{cite journal|author1=Arbour, V. M. |author2=Currie, P. J. |year=2015|title=Systematics, phylogeny and palaeobiogeography of the ankylosaurid dinosaurs|journal=Journal of Systematic Palaeontology|volume=14 |issue=5 |pages=1–60|doi=10.1080/14772019.2015.1059985}}</ref><ref name="Arbour2017">{{cite journal|last1=Arbour|first1=Victoria M.|last2=Evans|first2=David C.|year=2017|title=A new ankylosaurine dinosaur from the Judith River Formation of Montana, USA, based on an exceptional skeleton with soft tissue preservation|journal=Royal Society Open Science|volume=4|issue=5|pages=161086|doi=10.1098/rsos.161086|pmid=28573004|bibcode=2017RSOS....461086A|pmc=5451805}}</ref>
[[File:Ankylosaurus tail terminology.png|thumb|left|Reconstructed tail and pelvis]]

In 1971, [[Walter Coombs]] proposed that ''Dyoplosaurus'' was a [[junior synonym]] of ''[[Euoplocephalus]]''. The two were since generally assumed to be the same species and thus, former ''Dyoplosaurus'' specimens have been identified as ''Euoplocephalus tutus''. However, a redescription of the genus published in 2009 in the ''[[Journal of Vertebrate Paleontology]]'' claims that the genus is a valid taxon, suggesting that the proposed synonymy was due to the fragmentary nature of the holotype and other specimens of ''E. tutus''.<ref name=ABS09>{{Cite journal | last1 = Arbour | first1 = V. M. | last2 = Burns | first2 = M. E. | last3 = Sissons | first3 = R. L. | doi = 10.1671/039.029.0405 | title = A redescription of the ankylosaurid dinosaur ''Dyoplosaurus acutosquameus'' Parks, 1924 (Ornithischia: Ankylosauria) and a revision of the genus | journal = Journal of Vertebrate Paleontology | volume = 29 | issue = 4 | pages = 1117 | year = 2009 }}</ref> The [[cladistic analysis]] performed by Thompson ''et al.'', 2011 confirmed this separation.<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=in press |issue= 2|pages= 301–312|doi=10.1080/14772019.2011.569091 }}</ref>
A phylogenetic analysis conducted by Arbour & Evans, 2017 is reproduced below.<ref name="Arbour2017"/>
The following [[cladogram]] is based on a 2015 [[phylogenetic analysis]] of the Ankylosaurinae conducted by [[Victoria Arbour|Arbour]] and [[Philip J. Currie|Currie]] where it is recovered as a member of the subfamily Ankylosaurini:<ref name="systematics ankylosaurid">{{cite journal|author1=Arbour, V. M. |author2=Currie, P. J. |year=2015|title=Systematics, phylogeny and palaeobiogeography of the ankylosaurid dinosaurs|journal=Journal of Systematic Palaeontology|volume=14 |issue=5 |pages=1–60|doi=10.1080/14772019.2015.1059985}}</ref>

{{clade| style=font-size:85%; line-height:85%
|label1=[[Ankylosaurinae]]
|1={{clade
|1=''[[Zhejiangosaurus|Zhejiangosaurus luoyangensis]]''
|2={{clade
|1={{clade
|1=''[[Pinacosaurus|Pinacosaurus grangeri]]''
|2=''[[Pinacosaurus|Pinacosaurus mephistocephalus]]''}}
|2={{clade
|1={{clade
|1=''[[Tsagantegia|Tsagantegia longicranialis]]''
|2={{clade
|1=''[[Talarurus|Talarurus plicatospineus]]''
|2=''[[Nodocephalosaurus|Nodocephalosaurus kirtlandensis]]''}}}}
|2={{clade
|label2=[[Ankylosaurini]]
|1={{clade
|1=''[[Saichania|Saichania chulsanensis]]''
|2={{clade
|1=''[[Zaraapelta|Zaraapelta nomadis]]''
|2=''[[Tarchia|Tarchia kielanae]]''}}}}
|2={{clade
|1=''[[Ziapelta|Ziapelta sanjuanensis]]''
|2={{clade
|1=''[[Euoplocephalus|Euoplocephalus tutus]]''
|2={{clade
|1=''[[Ankylosaurus|Ankylosaurus magniventris]]''
|2={{clade
|1=''[[Anodontosaurus|Anodontosaurus lambei]]''
|2={{clade
|1=''[[Scolosaurus|Scolosaurus cutleri]]''
|2={{clade
|1=''[[Zuul crurivastator]]''
|2='''Dyoplosaurus'''}}}}}}}}}}}}}}}}}}}}}}

The results of an earlier analysis by Arbour & Currie, 2015 is reproduced below.<ref name="systematics ankylosaurid"/>


{{clade| style=font-size:85%; line-height:85%
{{clade| style=font-size:85%; line-height:85%
Line 54: Line 113:
|1=''[[Scolosaurus]]''
|1=''[[Scolosaurus]]''
|2=''[[Ziapelta]]'' }} }} }} }} }} }} }} }}
|2=''[[Ziapelta]]'' }} }} }} }} }} }} }} }}

==Paleoenvironment==
The holotype specimen of ''Dyoplosaurus'' was recovered from the base of the Dinosaur Park Formation, which dates to the middle [[Campanian]] stage of the [[Late Cretaceous]].<ref name=ABS09/> The lower Dinosaur Park Formation consist primarily of dryland habitats with modified channel-fills that experienced impeded drainage in the lower horizons and were subject to frequent flooding while more distal reaches of the floodplain, flooding was less frequent.<ref>{{cite journal|last=Matson|first=C.C.|title=Paleoenvironments of the Upper Cretaceous Dinosaur Park Formation in southern Alberta, Canada. (Master Thesis)|date=2010|institution=University of Calgary, Calgary, AB.|doi=http://dx.doi.org/10.11575/PRISM/18677}}</ref>

''Dyoplosaurus'' would have coexisted with the [[ankylosauria|ankylosaurs]] ''[[Edmontonia]]''<ref name=ABS09/>, ''Euoplocephalus''<ref name=ABS09/>, and ''Scolosaurus''<ref name=ABS09/>, the [[ceratopsidae|ceratopsid]] ''[[Chasmosaurus]]''<ref name=ABS09/> , the [[hadrosauridae|hadrosaurids]] ''[[Corythosaurus]]''<ref name=ABS09/> , ''[[Gryposaurus]]'' and ''[[Parasaurolophus]]''<ref name=ABS09/> , the [[tyrannosauridae|tyrannosaurid]] ''[[Gorgosaurus]]''<ref name=ABS09/> , the [[dromaeosauridae|dromaeosaurid]] ''[[Hesperonychus]]''<ref name="longrich&currie2009">Longrich, N.R. and Currie, P.J. (2009). "A microraptorine (Dinosauria–Dromaeosauridae) from the Late Cretaceous of North America." ''Proceedings of the National Academy of Sciences'' '''106'''(13): 5002–5007. {{doi| 10.1073/pnas.0811664106}}</ref>, the [[troodontidae|troodontid]] ''[[Latenivenatrix]]''<ref name="desc">{{cite journal|last1=van der Reest|first1=A. J.|last2=Currie|first2=P. J.|date=2017|title=Troodontids (Theropoda) from the Dinosaur Park Formation, Alberta, with a description of a unique new taxon: implications for deinonychosaur diversity in North America|journal=Canadian Journal of Earth Sciences|volume=54|issue=9|pages=919−935|doi=10.1139/cjes-2017-0031|hdl=1807/78296|hdl-access=free}}</ref>, and the [[caenagnathidae|caenagnathids]] ''[[Caenagnathus]]''<ref name="Citipes">{{Cite journal|last=Funston|first=Gregory|date=2020-07-27|title=Caenagnathids of the Dinosaur Park Formation (Campanian) of Alberta, Canada: anatomy, osteohistology, taxonomy, and evolution|journal=Vertebrate Anatomy Morphology Palaeontology|volume=8|pages=105–153|doi=10.18435/vamp29362|issn=2292-1389|doi-access=free}}</ref> and ''[[Chirostenotes]]''.<ref name="Citipes"/>


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

Revision as of 07:33, 20 November 2021

Dyoplosaurus
Temporal range: Late Cretaceous, 76.5 Ma
↓
Holotype skull
Scientific classification Edit this classification
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Clade: Dinosauria
Clade: †Ornithischia
Clade: †Thyreophora
Clade: †Ankylosauria
Family: †Ankylosauridae
Subfamily: †Ankylosaurinae
Tribe: †Ankylosaurini
Genus: †Dyoplosaurus
Parks, 1924
Species:
†D. acutosquameus
Binomial name
†Dyoplosaurus acutosquameus
Parks, 1924

Dyoplosaurus (meaning “double-armoured lizard”) is a monospecific genus of ankylosaurid dinosaur from Alberta that lived during the Late Cretaceous (middle Campanian, ~76.5–75 Ma) in what is now the Dinosaur Park Formation.[1] Dyoplosaurus represents a close relative of Scolosaurus and Anodontosaurus, two ankylosaurids known from the Horseshoe Canyon and Dinosaur Park Formation.[1][2]

History of discovery

Dinosaur Park Formation in Alberta, Canada

The holotype specimen was obtained in 1919 from the bottom 10 metres of the Dinosaur Park Formation, near what is now the Red Deer River in Alberta, Canada.[1][2] The holotype specimen, ROM 784, consists of a partial skull roof, mandible fragments with teeth, osteoderms, skin impressions, articulated post-thoracic vertebrae, partial thoracic ribs, a partial ilium, both ischia, tail club, associated radius, metacarpal, femur, tibia, fibula, and pes.[1][2] The holotype is currently housed at the Royal Ontario Museum.[1][2] Two specimens were referred to Dyoplosaurus, ROM 7761 and UA 47273, and both consist of partial tail clubs.[3]

The generic name, Dyoplosaurus, is derived from the Greek words “dyo” (double), “hoplon” (weapon, shield, armour) and “sauros” (lizard).[1] The specific name, acutosquameus, is derived from the Latin words “acutus” (sharp) and “squama” (scale).[1]

In 1956, Evgeny Maleev named a second species of Dyoplosaurus: D. giganteus.[4] The species was based on PIN 551/29, which consists of a series of caudal vertebrae, metatarsals, phalanges and osteoderms including tail club knob from the Nemegt Formation of Mongolia.[4] The species was diagnosed based on the short anterior caudal vertebrae; large chevrons that were fused to the caudal vertebrae; low, long distal caudal vertebrae; short, wide metatarsals; thick, hoofed-shaped unguals; sharp, thin-walled osteoderms with numerous pits and channels on the external surface.[4] However, a 2014 study by Arbour and colleagues considered that the holotype lacked diagnostic traits, as such traits are present in all ankylosaurines, and considered the species as a nomen dubium.[4]

Description

Tail of the holotype specimen and reconstruction of the tail and pelvis.

Dyoplosaurus has an estimated body length of 3.58 to 4.16 metres (13 to 15 feet) and weight of 1500 to 2500 kg (3300 to 5500 lbs).[5] All referred specimens represent almost fully mature individuals.[6]

Dyoplosaurus can be distinguished from all other ankylosaurids in having sacral ribs that are anterolaterally-directed, triangular unguals in dorsal view and a tail club knob that is longer than wide.[2] It differs from Scolosaurus in having a proportionately shorter postacetabular process of the ilium and triangular osteoderms on the lateral sides of the anterior portion of the tail.[2] Dyoplosaurus also differs from Euoplocephalus in the pelvis as it has anterolaterally projecting, ventrally directed sacral transverse processes on the third sacral vertebra, forming a butterfly-like arrangement of the sacral fenestrae, and in having ischia that articulate with the ilia at right angles.[2]

Tail club of Dyoplosaurus compared to other ankylosaurid tail clubs

The holotype specimen of Dyoplosaurus preserves ossified tendons on the tail club.[6][1] The tail club preserves three series of tendons on the dorsolateral sides of the handle, and four on the distal, ventral side of the tail.[6][1] These tendons are more readily grouped into two sets on the dorsolateral sides.[6][1] Arbour, 2009 conducted a study to determine the impact force of ankylosaurids and used ROM 784. Arbour found that Dyoplosaurus could generate an impact force of 797–1127 N and a more realistic tensions of 571 N, an impact force that isn’t enough to puncture bone.[6] This is mainly due to the knob being smaller in comparison to that of other ankylosaurids.[6] The small size of the tail club of suggests that ankylosaurid knobs were not primarily used as defensive weapons, as a weapon that is not functional until very late in life would probably not have a selective advantage over a weapon that is of use earlier in life.[6]

The holotype specimen preserves remnants of fossilized skin and osteoderms on the left side.[2] These skin impressions are punctuated by a nearly unbroken mosaic of small (0.50–1.0 cm), sub-angular to subrounded osteoderms.[2] The skin imperssions on the right side have been lost via erosion.[2]

Classification

In 1971, Walter Coombs synonymized Dyoplosaurus, along with Scolosaurus and Anodontosaurus, into Euoplocephalus as one of the four mandibles assigned to Dyoplosaurus was identical to those of other Euoplocephalus specimens, but did not offer any other characteristics to support the synonymization.[7] However, a re-description of Dyoplosaurus published in 2009 by Victoria Arbour, Michael Burns and Robin Sissons considered it as a valid taxon and proposed that the synonymy was due to the fragmentary nature of the holotype and other referred specimens of Euoplocephalus.[2] Thompson et al., 2011 confirmed its separation and recovered it as sister taxon to Pinacosaurus mephistocephalus.[8] A cladistic analysis conducted by Arbour and Currie, 2015 recovered Dyoplosaurus as sister taxon to a clade containing Ankylosaurus, Euoplocephalus, Anodontosaurus and Scolosaurus, while an analysis conducted by Arbour and Evans, 2017 recovered it as sister taxon to Zuul.[9][10]

A phylogenetic analysis conducted by Arbour & Evans, 2017 is reproduced below.[10]

Ankylosaurinae

The results of an earlier analysis by Arbour & Currie, 2015 is reproduced below.[9]

Ankylosaurinae

Paleoenvironment

The holotype specimen of Dyoplosaurus was recovered from the base of the Dinosaur Park Formation, which dates to the middle Campanian stage of the Late Cretaceous.[2] The lower Dinosaur Park Formation consist primarily of dryland habitats with modified channel-fills that experienced impeded drainage in the lower horizons and were subject to frequent flooding while more distal reaches of the floodplain, flooding was less frequent.[11]

Dyoplosaurus would have coexisted with the ankylosaurs Edmontonia[2], Euoplocephalus[2], and Scolosaurus[2], the ceratopsid Chasmosaurus[2] , the hadrosaurids Corythosaurus[2] , Gryposaurus and Parasaurolophus[2] , the tyrannosaurid Gorgosaurus[2] , the dromaeosaurid Hesperonychus[12], the troodontid Latenivenatrix[13], and the caenagnathids Caenagnathus[14] and Chirostenotes.[14]

See also

References

  1. ^ a b c d e f g h i j Parks, W. A. (1924). "Dyoplosaurus acutosquameus, a new genus and species of armored dinosaur; and notes on a skeleton of Prosaurolophus maximus". University of Toronto Studies Geological Series. 18: 1–35.
  2. ^ a b c d e f g h i j k l m n o p q r s Arbour, V. M.; Burns, M. E.; Sissons, R. L. (2009). "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. doi:10.1671/039.029.0405.
  3. ^ Paul Penkalski (2018). "Revised systematics of the armoured dinosaur Euoplocephalus and its allies". Neues Jahrbuch für Geologie und Paläontologie - Abhandlungen. 287 (3): 261–306. doi:10.1127/njgpa/2018/0717.
  4. ^ a b c d Victoria M. Arbour, Philip J. Currie and Demchig Badamgarav, 2014, "The ankylosaurid dinosaurs of the Upper Cretaceous Baruungoyot and Nemegt formations of Mongolia", Zoological Journal of the Linnean Society 172(3): 631–652|doi=https://doi.org/10.1111/zoj12185
  5. ^ Miyashita T, Arbour VM, Witmer LM, Currie PJ (2011). "The internal cranial morphology of an armoured dinosaur Euoplocephalus corroborated by X-ray computed tomographic reconstruction". Journal of Anatomy. 219 (6): 661–75. doi:10.1111/j.1469-7580.2011.01427.x. PMC 3237876. PMID 21954840.
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  10. ^ a b Arbour, Victoria M.; Evans, David C. (2017). "A new ankylosaurine dinosaur from the Judith River Formation of Montana, USA, based on an exceptional skeleton with soft tissue preservation". Royal Society Open Science. 4 (5): 161086. Bibcode:2017RSOS....461086A. doi:10.1098/rsos.161086. PMC 5451805. PMID 28573004.
  11. ^ Matson, C.C. (2010). "Paleoenvironments of the Upper Cretaceous Dinosaur Park Formation in southern Alberta, Canada. (Master Thesis)". University of Calgary, Calgary, AB. doi:http://dx.doi.org/10.11575/PRISM/18677. {{cite journal}}: Check |doi= value (help); Cite journal requires |journal= (help); External link in |doi= (help)
  12. ^ Longrich, N.R. and Currie, P.J. (2009). "A microraptorine (Dinosauria–Dromaeosauridae) from the Late Cretaceous of North America." Proceedings of the National Academy of Sciences 106(13): 5002–5007. doi:10.1073/pnas.0811664106
  13. ^ van der Reest, A. J.; Currie, P. J. (2017). "Troodontids (Theropoda) from the Dinosaur Park Formation, Alberta, with a description of a unique new taxon: implications for deinonychosaur diversity in North America". Canadian Journal of Earth Sciences. 54 (9): 919−935. doi:10.1139/cjes-2017-0031. hdl:1807/78296.
  14. ^ a b Funston, Gregory (2020-07-27). "Caenagnathids of the Dinosaur Park Formation (Campanian) of Alberta, Canada: anatomy, osteohistology, taxonomy, and evolution". Vertebrate Anatomy Morphology Palaeontology. 8: 105–153. doi:10.18435/vamp29362. ISSN 2292-1389.


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