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''Denversaurus'' (meaning "Denver lizard") is a genus of panoplosaurin
nodosaurid Nodosauridae is a family of ankylosaurian dinosaurs known from the Late Jurassic to the Late Cretaceous periods in what is now Asia, Europe, North America, and possibly South America. While traditionally regarded as a monophyletic clade as the s ...
dinosaur Dinosaurs are a diverse group of reptiles of the clade Dinosauria. They first appeared during the Triassic Geological period, period, between 243 and 233.23 million years ago (mya), although the exact origin and timing of the #Evolutio ...
from the late
Maastrichtian The Maastrichtian ( ) is, in the International Commission on Stratigraphy (ICS) geologic timescale, the latest age (geology), age (uppermost stage (stratigraphy), stage) of the Late Cretaceous epoch (geology), Epoch or Upper Cretaceous series (s ...
of
Late Cretaceous The Late Cretaceous (100.5–66 Ma) is the more recent of two epochs into which the Cretaceous Period is divided in the geologic time scale. Rock strata from this epoch form the Upper Cretaceous Series. The Cretaceous is named after ''cre ...
Western North America. Although at one point treated as a
junior synonym In taxonomy, the scientific classification of living organisms, a synonym is an alternative scientific name for the accepted scientific name of a taxon. The botanical and zoological codes of nomenclature treat the concept of synonymy differently. ...
of '' Edmontonia'' by some
taxonomists In biology, taxonomy () is the scientific study of naming, defining ( circumscribing) and classifying groups of biological organisms based on shared characteristics. Organisms are grouped into taxa (singular: taxon), and these groups are given ...
, current research indicates that it is its own distinct nodosaurid
genus Genus (; : genera ) is a taxonomic rank above species and below family (taxonomy), family as used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In bino ...
.


Discovery and naming

In 1986, the paleontologists Kenneth Carpenter and Brent Breithaupt described DMNH 468 which was a specimen of a late
Maastrichtian The Maastrichtian ( ) is, in the International Commission on Stratigraphy (ICS) geologic timescale, the latest age (geology), age (uppermost stage (stratigraphy), stage) of the Late Cretaceous epoch (geology), Epoch or Upper Cretaceous series (s ...
nodosaurid Nodosauridae is a family of ankylosaurian dinosaurs known from the Late Jurassic to the Late Cretaceous periods in what is now Asia, Europe, North America, and possibly South America. While traditionally regarded as a monophyletic clade as the s ...
, tentatively assigned to '' Edmontonia'' sp., discovered from the lower
Hell Creek Formation The Hell Creek Formation is an intensively studied division of mostly Upper Cretaceous and some lower Paleocene rocks in North America, named for exposures studied along Hell Creek, near Jordan, Montana. The Formation (stratigraphy), formation s ...
of
South Dakota South Dakota (; Sioux language, Sioux: , ) is a U.S. state, state in the West North Central states, North Central region of the United States. It is also part of the Great Plains. South Dakota is named after the Dakota people, Dakota Sioux ...
. In 1988, Robert Thomas Bakker decided to split the genus ''Edmontonia''. The species ''Edmontonia rugosidens'' was made into a separate genus named ''Chassternbergia'' and DMNH 468 was designated as a holotype of a new genus and species. The
type species In International_Code_of_Zoological_Nomenclature, zoological nomenclature, a type species (''species typica'') is the species name with which the name of a genus or subgenus is considered to be permanently taxonomically associated, i.e., the spe ...
of this genus was ''Denversaurus schlessmani''. The generic name referred to the Denver Museum of Natural History at
Denver, Colorado Denver ( ) is a List of municipalities in Colorado#Consolidated city and county, consolidated city and county, the List of capitals in the United States, capital and List of municipalities in Colorado, most populous city of the U.S. state of ...
. The
specific name Specific name may refer to: * in Database management systems, a system-assigned name that is unique within a particular database In taxonomy, either of these two meanings, each with its own set of rules: * Specific name (botany), the two-part (bino ...
honoured Lee E. Schlessman, a major benefactor of the museum and the founder of the Schlessman Family Foundation.Bakker, R.T. (1988). "Review of the Late Cretaceous nodosauroid Dinosauria: ''Denversaurus schlessmani'', a new armor-plated dinosaur from the Latest Cretaceous of South Dakota, the last survivor of the nodosaurians, with comments on Stegosaur-Nodosaur relationships". ''Hunteria'' 1(3): 1-23.(1988). The holotype consists of a skull without the lower jaws and a number of
osteoderms Osteoderms are bony deposits forming scales, plates, or other structures based in the dermis. Osteoderms are found in many groups of Extant taxon, extant and extinct reptiles and amphibians, including lizards, crocodilians, frogs, Temnospondyli, ...
of the body armour. It is part of the collection of the Denver Museum of Nature and Science after which the genus was named. Bakker referred a second fossil to the species, specimen AMNH 3076, a skull found by Brown and
American Museum of Natural History The American Museum of Natural History (AMNH) is a natural history museum on the Upper West Side of Manhattan in New York City. Located in Theodore Roosevelt Park, across the street from Central Park, the museum complex comprises 21 interconn ...
paleontologist Roland T. Bird at the Tornillo Creek in
Brewster County, Texas Brewster County is a county (United States), county located in the U.S. state of Texas. It is in West Texas and its county seat (and only city) is Alpine, Texas, Alpine. It is one of the nine counties that comprise the Trans-Pecos region, and b ...
, in a layer of the poorly dated
Upper Cretaceous The Late Cretaceous (100.5–66 Ma) is the more recent of two epochs into which the Cretaceous Period is divided in the geologic time scale. Rock strata from this epoch form the Upper Cretaceous Series. The Cretaceous is named after ''cret ...
Aguja Formation, possibly from the Maastrichtian too. However, subsequent studies reassigned AMNH 3076 to '' Edmontonia'', probably ''E. longicpes''. The validity of ''Denversaurus'' was disputed in a 1990 paper on ankylosaurian systematics by Kenneth Carpenter, who noted that Bakker's diagnosis of ''Denversaurus'' was based primarily on Bakker's artistic restoration of the holotype in an uncrushed state. Since DMNH 468 was found crushed, Carpenter assigned ''Denversaurus'' back to ''Edmontonia'' sp., even though he noted its similarity to ''Edmontonia rugosidens''.Carpenter, K. 1990. "Ankylosaur systematics: example using Panoplosaurus and Edmontonia (Ankylosauria: Nodosauridae)", In: Carpenter, K. & Currie, P.J. (eds) Dinosaur Systematics: Approaches and Perspectives, Cambridge University Press, Cambridge, pp. 281-298 A number of workers treated ''Denversaurus'' as synonymous with either ''E. rugosidens'' or ''E. longiceps'', or alternatively as a valid species of ''Edmontonia'': ''E. schlessmani''.Paul, G.S. (2010). The Princeton Field Guide to Dinosaurs, Princeton University Press. In 1988, fossil hunters found a partial nodosaurid skeleton consisting of the fragmentary skull, parts of postcranial skeleton and seventy-five osteoderms from the
Lance Formation The Lance (Creek) Formation is a division of Late Cretaceous (dating to about 69–66 Ma) rocks in the western United States. Named after Lance Creek, Wyoming, the microvertebrate fossils and dinosaurs represent important components of the lates ...
of
Niobrara County, Wyoming Niobrara County is a county in the U.S. state of Wyoming. As of the 2020 United States census, the population was 2,467, making it the least populous county in Wyoming. Its county seat is Lusk. Its eastern boundary abuts the west lines of the ...
, nicknamed "Tank", which has been tentatively assigned to as ''Edmontonia'' sp. in 1994. It was part of the collection of the Rocky Mountain Dinosaur Resource Center under inventory number BHI 127327, referred to as ''Edmontonia schlessmani'' by Carpenter et al. (2013), and is now under the inventory number FPDM-V9673. In the 2015 thesis, Michael Burns revisited the systematics of latest Cretaceous nodosaurids from the Western Interior. According to Burns, ''Denversaurus'' is likely a valid taxon based on its
phylogenetic In biology, phylogenetics () is the study of the evolutionary history of life using observable characteristics of organisms (or genes), which is known as phylogenetic inference. It infers the relationship among organisms based on empirical dat ...
position. Recent phylogenetic analyses included ''Denversaurus'' as a valid genus closely related to ''Edmontonia''.


Description

In 2010, American
paleontologist Paleontology, also spelled as palaeontology or palæontology, is the scientific study of the life of the past, mainly but not exclusively through the study of fossils. Paleontologists use fossils as a means to classify organisms, measure geolo ...
Gregory S. Paul Gregory Scott Paul (born December 24, 1954) is an American freelance researcher, author and illustrator who works in paleontology. He is best known for his work and research on theropoda, theropod dinosaurs and his detailed illustrations, both l ...
estimated the length of ''Denversaurus'' at and its body mass at . Robert T. Bakker considered ''Denversaurus'' distinct from ''Edmontonia'' and ''Chassternbergia'' in having a skull that was wide at the rear and a more rearward position of the eye sockets. The holotype skull has a length of 496 millimetres and a rear width of 346 millimetres. In the referred specimen AMNH 3076, these proportions are less extreme, measuring 395 millimetres long with a rear width of 220 millimetres. According to American paleontologist Kenneth Carpenter, the greater width of both the holotype and the referred specimen was due to crushing. In 2015, vertebrate anatomist and paleontologist Michael Burns published an abstract concluding that ''Denversaurus'' was different from ''Edmontonia'', but similar to '' Panoplosaurus'' in having inflated, convex, cranial sculpturing with visible sulci, or troughs, between individual top skull armour elements, but is distinct from ''Panoplosaurus'' in having a relatively wider snout.


Classification

In 1988, Bakker placed ''Denversaurus'' within Edmontoniidae, the presumed
sister group In phylogenetics, a sister group or sister taxon, also called an adelphotaxon, comprises the closest relative(s) of another given unit in an evolutionary tree. Definition The expression is most easily illustrated by a cladogram: Taxon A and ...
of
Nodosauridae Nodosauridae is a family of ankylosaurian dinosaurs known from the Late Jurassic to the Late Cretaceous periods in what is now Asia, Europe, North America, and possibly South America. While traditionally regarded as a monophyletic clade as the ...
within Nodosauroidea that would not have been
Ankylosauria Ankylosauria is a group of herbivorous dinosaurs of the clade 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 ...
, but the last surviving
Stegosauria Stegosauria is a group of Herbivore, herbivorous ornithischian dinosaurs that lived during the Jurassic and early Cretaceous Period (geology), periods. Stegosaurian fossils have been found mostly in the Northern Hemisphere (North America, Europe a ...
. However, these hypotheses have not been confirmed by modern
cladistic Cladistics ( ; from Ancient Greek 'branch') is an approach to biological classification in which organisms are categorized in groups ("clades") based on hypotheses of most recent common ancestry. The evidence for hypothesized relationships is ...
analysis. Whether it presents a separate species or is identical to ''E. rugosidens'' or ''E. longiceps'', ''Denversaurus'' material is considered nodosaurid and ankylosaurian. Paul suggested that it was the direct descendant of ''E. longiceps''. Burns recovered ''Denversaurus'' as the
sister species In phylogenetics, a sister group or sister taxon, also called an adelphotaxon, comprises the closest relative(s) of another given unit in an evolutionary tree. Definition The expression is most easily illustrated by a cladogram: Taxon A and ...
of ''Panoplosaurus''. ''Denversaurus'' is the latest known member of
Thyreophora Thyreophora ("shield bearers", often known simply as "armored dinosaurs") is a group of armored ornithischian dinosaurs that lived from the Early Jurassic until the end of the Cretaceous. Thyreophorans are characterized by the presence of bod ...
.


References

{{Taxonbar, from=Q21012043 Nodosauridae Dinosaur genera Maastrichtian dinosaurs Hell Creek Formation Lance Formation Fossil taxa described in 1988 Taxa named by Robert T. Bakker Dinosaurs of the United States