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''Cedarpelta'' is a extinct genus of basal
ankylosaurid Ankylosauridae () is a family of armored dinosaurs within Ankylosauria, and is the sister group to Nodosauridae. The oldest known Ankylosaurids date to around 122 million years ago and went extinct 66 million years ago during the Cretaceous–Pal ...
dinosaur from Utah that lived during the Late Cretaceous period ( Cenomanian to lower Turonian stage, 98.2 to 93 Ma) in what is now the Mussentuchit Member of the
Cedar Mountain Formation The Cedar Mountain Formation is the name given to a distinctive sedimentary geologic formation in eastern Utah, spanning most of the early and mid-Cretaceous. The formation was named for Cedar Mountain in northern Emery County, Utah, where Willia ...
. The type and only species, ''Cedarpelta bilbeyhallorum'', is known from multiple specimens including partial skulls and
postcrania Postcrania (postcranium, adjective: postcranial) in zoology and vertebrate paleontology is all or part of the skeleton apart from the skull. Frequently, fossil remains, e.g. of dinosaurs or other extinct tetrapods, consist of partial or isolated sk ...
l material. It was named in 2001 by
Kenneth Carpenter Kenneth Carpenter (born September 21, 1949, in Tokyo, Japan) is a paleontologist. He is the former director of the USU Eastern Prehistoric Museum and author or co-author of books on dinosaurs and Mesozoic life. His main research interests ...
, James Kirkland, Don Burge, and John Bird. ''Cedarpelta'' has an estimated length of 7 metres (23 feet) and weight of 5 tonnes (11,023 lbs). The skull of ''Cedarpelta'' lacks extensive
cranial Standard anatomical terms of location are used to unambiguously describe the anatomy of animals, including humans. The terms, typically derived from Latin or Greek roots, describe something in its standard anatomical position. This position prov ...
ornamentation and is one of the only known ankylosaurs with individual skull bones that are not completely fused together.


Discovery and naming

The partial remains of an
ankylosaur Ankylosauria is a group of herbivorous dinosaurs of the order Ornithischia. It includes the great majority of dinosaurs with armor in the form of bony osteoderms, similar to turtles. Ankylosaurs were bulky quadrupeds, with short, powerful limbs. ...
were discovered by Evan Hall and Sue Ann Bilbey at the CEM site near the
Price River The Price River is a U.S. Geological Survey. National Hydrography Dataset high-resolution flowline dataThe National Map accessed October 30, 2020 southeastward flowing river in Carbon, Utah and Emery counties in eastern Utah. It is a tributary to ...
in
Carbon County, Utah Carbon County is a county in the U.S. state of Utah. As of the 2010 United States Census, the population was 21,403. Its county seat and largest city is Price. The Price, UT Micropolitan Statistical Area includes all of Carbon County. Histor ...
while they were visiting an excavation in the surrounding area.Carpenter, K., Kirkland, J. I., Burge, D., and Bird, J. 2001. Disarticulated skull of a new primitive ankylosaurid from the Lower Cretaceous of Utah. in Carpenter, K. (editor) 2001. The Armored Dinosaurs. Indiana University Press The site was originally interpreted as being from the top of the Ruby Ranch Member of the
Cedar Mountain Formation The Cedar Mountain Formation is the name given to a distinctive sedimentary geologic formation in eastern Utah, spanning most of the early and mid-Cretaceous. The formation was named for Cedar Mountain in northern Emery County, Utah, where Willia ...
, but was later interpreted as being from the bottom of the Mussentuchit Member. The age of the layer was originally thought to have been 104.46 ± 0.95 Ma, but more recent estimates date it to 98.2 ± 0.6 to 93 Ma. In 1998, the discovery was reported by
Kenneth Carpenter Kenneth Carpenter (born September 21, 1949, in Tokyo, Japan) is a paleontologist. He is the former director of the USU Eastern Prehistoric Museum and author or co-author of books on dinosaurs and Mesozoic life. His main research interests ...
and James Kirkland.Carpenter K., Kirkland J.I., 1998, "Review of Lower and middle Cretaceous ankylosaurs from North America", ''New Mexico Museum of Natural History and Science Bulletin'' 14: 249-270 In 2001, it was subsequently described, along with other material, by Kenneth Carpenter, James Kirkland, Don Burge, and John Bird. The holotype specimen, CEUM 12360, consists of a partial skull that is missing the snout and lower jaws. Numerous
osteoderm Osteoderms are bony deposits forming scales, plates, or other structures based in the dermis. Osteoderms are found in many groups of extant and extinct reptiles and amphibians, including lizards, crocodilians, frogs, temnospondyls (extinct amp ...
s, postcranial material and a disarticulated skull were designated as
paratype In zoology and botany, a paratype is a specimen of an organism that helps define what the scientific name of a species and other taxon actually represents, but it is not the holotype (and in botany is also neither an isotype nor a syntype). Of ...
specimens. Both holotype and paratype specimens represent at least three individuals and are currently housed at the College of Eastern Utah, Prehistoric Museum, Utah. The generic name, ''Cedarpelta'', is derived from the Cedar Mountain Formation and the Greek word "''pelte''" (small shield). The specific name, ''bilbeyhallorum'', honours Sue Ann Bilbey and Evan Hall, who discovered the remains of ''Cedarpelta''. In 2008, additional specimens were referred to ''Cedarpelta'' from the Price River II Quarry, which is about 24.5 km southeast of Price River, Utah and at the base of the Mussentuchit Member. The quarry also produced specimens pertaining to four individuals of a
brachiosaurid The Brachiosauridae ("arm lizards", from Greek ''brachion'' (βραχίων) = "arm" and ''sauros'' = "lizard") are a family or clade of herbivorous, quadrupedal sauropod dinosaurs. Brachiosaurids had long necks that enabled them to access the le ...
, an iguanodontian, a turtle, a
pterosaur Pterosaurs (; from Greek ''pteron'' and ''sauros'', meaning "wing lizard") is an extinct clade of flying reptiles in the order, Pterosauria. They existed during most of the Mesozoic: from the Late Triassic to the end of the Cretaceous (228 to ...
, and specimens of the
nodosaurid Nodosauridae is a family of ankylosaurian dinosaurs, from the Late Jurassic to the Late Cretaceous period in what is now North America, South America, Europe, and Asia. Description Nodosaurids, like their close relatives the ankylosaurids, we ...
''
Peloroplites ''Peloroplites'' (meaning “monstrous heavy one”) is a monospecific genus of nodosaurid dinosaur from Utah that lived during the Late Cretaceous ( Cenomanian to lower Turonian stage, 98.2 to 93 Ma) in what is now the Mussentuchit Member ...
''. The referred material includes: CEUM 10396, a
cervical vertebra In tetrapods, cervical vertebrae (singular: vertebra) are the vertebrae of the neck, immediately below the skull. Truncal vertebrae (divided into thoracic and lumbar vertebrae in mammals) lie caudal (toward the tail) of cervical vertebrae. In sau ...
; CEUM 10412, CEUM 10404, caudal
vertebra The spinal column, a defining synapomorphy shared by nearly all vertebrates,Hagfish are believed to have secondarily lost their spinal column is a moderately flexible series of vertebrae (singular vertebra), each constituting a characteristic ...
e; CEUM 10371, a
coracoid A coracoid (from Greek κόραξ, ''koraks'', raven) is a paired bone which is part of the shoulder assembly in all vertebrates except therian mammals (marsupials and placentals). In therian mammals (including humans), a coracoid process is prese ...
; CEUM 10256, CEUM 11629, humeri; CEUM 10266, an ischium; CEUM 11334, a femur; and CEUM 11640, a tibia.


Description

Carepnter et al. (2001) originally gave Cedarpelta an estimated length of 7.5-8.5 metres (24.6-27.9 feet). However, Gregory S. Paul gave a lower estimate of 7 metres (23 feet) and a weight of 5 tonnes (11,023 lbs), while
Thomas Holtz Thomas Richard Holtz Jr. (born September 13, 1965) is an American vertebrate palaeontologist, author, and principal lecturer at the University of Maryland's Department of Geology. He has published extensively on the phylogeny, morphology, ecomor ...
gave a higher estimation at 9 meters suggesting that it was rivalling ''
Ankylosaurus ''Ankylosaurus'' is a genus of armored dinosaur. Its fossils have been found in geological formations dating to the very end of the Cretaceous Period, about 68–66 million years ago, in western North America, making it among the last of th ...
''.Paul, G.S., 2010, ''The Princeton Field Guide to Dinosaurs'', Princeton University Press p. 231Paul, G.S., 2016, ''The Princeton Field Guide to Dinosaurs 2nd Edition'', Princeton University Press Carpenter ''et al.'' (2001) established several distinguishing traits of ''Cedarpelta''. The body of the praemaxilla, the front snout bone, is short in front of its nasal branch. The outer sides of the two praemaxillae run more parallel compared to the snouts of later forms which are strongly diverging to behind. The cutting edge of the bone core of the upper beak is limited to the front of the praemaxilla. Each praemaxilla has six (conical)
teeth A tooth ( : teeth) is a hard, calcified structure found in the jaws (or mouths) of many vertebrates and used to break down food. Some animals, particularly carnivores and omnivores, also use teeth to help with capturing or wounding prey, tea ...
. The quadrate, and with it the entire back of the skull, is inclined to the front. The head of the quadrate is not fused with the paroccipital process, contrary to the situation in ''
Shamosaurus ''Shamosaurus'' is an extinct genus of herbivorous basal ankylosaurid ankylosaur from Early Cretaceous (Aptian to Albian stage) deposits of Höövör, Mongolia. Discovery and naming In 1977, a Soviet-Mongolian expedition discovered the skeleto ...
''. The neck of the
occipital condyle The occipital condyles are undersurface protuberances of the occipital bone in vertebrates, which function in articulation with the superior facets of the atlas vertebra. The condyles are oval or reniform (kidney-shaped) in shape, and their anteri ...
is long and sticking out to behind, like with nodosaurids, not obliquely to below as in typical ankylosaurids. The ''tubera basilaria'', appending processes of the rear lower braincase, form a large wedge directed to below. The
pterygoid Pterygoid, from the Greek for 'winglike', may refer to: * Pterygoid bone, a bone of the palate of many vertebrates * Pterygoid processes of the sphenoid bone ** Lateral pterygoid plate ** Medial pterygoid plate * Lateral pterygoid muscle * Medi ...
is elongated from the front to the rear and has a saddle-shaped process on its outer edge oriented to behind and sideways. The coronoid process of the rear lower jaw has an oval process at the inside. The straight ischium has a knob-shaped boss at the inside near the pubic pedicle. ''Cedarpelta'' shows a mix of basal and derived traits. The presence of premaxillary teeth is a plesiomorphic character because it is inherited from earlier Ornithischia. In contrast, closure of the opening on the side of the skull behind the orbit, the lateral temporal fenestra, is an advanced, derived (
apomorphic In phylogenetics, an apomorphy (or derived trait) is a novel character or character state that has evolved from its ancestral form (or plesiomorphy). A synapomorphy is an apomorphy shared by two or more taxa and is therefore hypothesized to have ...
) character only known in ankylosaurid ankylosaurians. Two skulls are known, and the skull length for ''Cedarpelta'' is estimated to have been roughly . One of the ''Cedarpelta'' skulls was found disarticulated, a first for an ankylosaur skull, allowing
paleontologist Paleontology (), also spelled palaeontology or palæontology, is the scientific study of life that existed prior to, and sometimes including, the start of the Holocene epoch (roughly 11,700 years before present). It includes the study of foss ...
s a unique opportunity to examine the individual bones instead of being limited to an ossified unit. The skull is relatively elongated and does not show a strongly appending beak. Of the conical premaxillary teeth, the first is the largest. The maxilla bears eighteen teeth. The eye socket is surrounded by the lacrimal, a single supraorbital and a large
postorbital The ''postorbital'' is one of the bones in vertebrate skulls which forms a portion of the dermal skull roof and, sometimes, a ring about the orbit. Generally, it is located behind the postfrontal and posteriorly to the orbital fenestra. In some ve ...
, excluding the prefrontal and the
jugal The jugal is a skull bone found in most reptiles, amphibians and birds. In mammals, the jugal is often called the malar or zygomatic. It is connected to the quadratojugal and maxilla, as well as other bones, which may vary by species. Anatomy ...
from the orbital rim. The postcranial skeleton was in 2001 not described in any detail. The skulls, though of large and thus not juvenile individuals, do not show a distinctive pattern of fused ''caputegulae'', head tiles. This inspired Carpenter to propose an alternative hypothesis of ankylosaur skull
osteoderm Osteoderms are bony deposits forming scales, plates, or other structures based in the dermis. Osteoderms are found in many groups of extant and extinct reptiles and amphibians, including lizards, crocodilians, frogs, temnospondyls (extinct amp ...
formation. Formerly, it had been assumed that such armour plates were either formed by direct skin ossification into distinct scutes which later fused to the skull (the more popular theory), or by a reaction of the skull bones to the pattern of overlying scales. The lack of a clear pattern in ''Cedarpelta'' suggested to Carpenter that the ossification took place in an intermediate layer between the scales and the skull roof itself, which he surmised to have been the
periosteum The periosteum is a membrane that covers the outer surface of all bones, except at the articular surfaces (i.e. the parts within a joint space) of long bones. Endosteum lines the inner surface of the medullary cavity of all long bones. Structu ...
.


Classification

Carpenter (2001) placed ''Cedarpelta'' within the family
Ankylosauridae Ankylosauridae () is a family of armored dinosaurs within Ankylosauria, and is the sister group to Nodosauridae. The oldest known Ankylosaurids date to around 122 million years ago and went extinct 66 million years ago during the Cretaceous–Pal ...
and offered two interpretations of its position. The first was that it could be the basalmost known ankylosaurid, i.e. the first discovered branch to split off from the ankylosaurid stem line. This would be in line with its
plesiomorphic In phylogenetics, a plesiomorphy ("near form") and symplesiomorphy are synonyms for an ancestral character shared by all members of a clade, which does not distinguish the clade from other clades. Plesiomorphy, symplesiomorphy, apomorphy, and ...
traits and the fact that the in 2001 supposed
Barremian The Barremian is an age in the geologic timescale (or a chronostratigraphic stage) between 129.4 ± 1.5 Ma (million years ago) and 121.4 ± 1.0 Ma). It is a subdivision of the Early Cretaceous Epoch (or Lower Cretaceous Series). It is preceded ...
age made it one of the oldest known ankylosaurids. The second was that it formed an early ankylosaurid branch, or clade, Shamosaurinae together with ''
Gobisaurus ''Gobisaurus'' is an extinct genus of herbivorous basal ankylosaurid ankylosaur from the Upper Cretaceous (and possibly also the Lower Cretaceous) of China ('' Nei Mongol Zizhiqu''). The genus is monotypic, containing only the species ''Gobisaur ...
'' of north-central China and the eponymous ''
Shamosaurus ''Shamosaurus'' is an extinct genus of herbivorous basal ankylosaurid ankylosaur from Early Cretaceous (Aptian to Albian stage) deposits of Höövör, Mongolia. Discovery and naming In 1977, a Soviet-Mongolian expedition discovered the skeleto ...
'' of Mongolia.Carpenter, K. 2001. Phylogenetic analysis of the Ankylosauria. in Carpenter, K. (editor) 2001. The Armored Dinosaurs. Indiana University Press Thompson et al. (2012),Richard S. Thompson, Jolyon C. Parish, Susannah C. R. Maidment and Paul M. Barrett, 2012, "Phylogeny of the ankylosaurian dinosaurs (Ornithischia: Thyreophora)", ''Journal of Systematic Palaeontology'' 10(2): 301–312 Chen et al. (2013), Yang et al. (2013), Han et al. (2014), Arbour & Currie (2015), Arbour et al. (2016), Arbour & Evans (2017), Yang et al. (2017), Zheng et al. (2018), Rivera-Sylva et al. (2018), Park et al. (2019) and Frauenfelder et al. (2022) have all found ''Cedarpelta'' to be within Ankylosauridae, as either within a polytomy with ''
Liaoningosaurus ''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 Form ...
'', '' Aletopelta'', '' Chuanqilong'', ''Gobisaurus'' and ''Shamosaurus'' or as sister taxon to ''Chuanqilong''. The results of Arbour & Currie (2015) are reproduced below. Vickaryous et al. (2004) interpreted ''Cedarpelta'' as the basalmost member of the family
Nodosauridae Nodosauridae is a family of ankylosaurian dinosaurs, from the Late Jurassic to the Late Cretaceous period in what is now North America, South America, Europe, and Asia. Description Nodosaurids, like their close relatives the ankylosaurids ...
, positioned even below the nodosaurids ''
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 co ...
'', ''
Silvisaurus ''Silvisaurus'', from the Latin silva "woodland" and Greek sauros "lizard", is a nodosaurid ankylosaur from the Early to Late Cretaceous period. Discovery and species A fossil of the species was discovered in the fifties by rancher Warren ...
'', and ''
Sauropelta ''Sauropelta'' ( ; meaning 'lizard shield') is a genus of nodosaurid dinosaur that existed in the Early Cretaceous Period of North America. One species (''S. edwardsorum'') has been named although others may have existed. Anatomically, ''Saurope ...
''.Vickaryous, M. K., Maryanska, T., and Weishampel, D. B. (2004). Chapter Seventeen: Ankylosauria. in The Dinosauria (2nd edition), Weishampel, D. B., Dodson, P., and Osmólska, H., editors. University of California Press. Wiersma & Irmis (2018) also interpreted Cedarpelta as a nodosaurid. The results of Vickaryous et al. (2004) are reproduced below.


See also

*
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 club ...


References

{{Taxonbar, from=Q135487 Early Cretaceous dinosaurs of North America Ankylosaurids Fossil taxa described in 2001 Taxa named by James I. Kirkland Taxa named by Kenneth Carpenter Ornithischian genera