Wendiceratops Restoration
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''Wendiceratops'' is a genus of herbivorous
centrosaurine Centrosaurinae (from the Greek, meaning "pointed lizards") is a subfamily of ceratopsid dinosaurs, a group of large quadrupedal ornithischians. Centrosaurine fossil remains are known primarily from the northern region of Laramidia (modern day Al ...
ceratopsian Ceratopsia or Ceratopia ( or ; Greek: "horned faces") is a group of herbivorous, beaked dinosaurs that thrived in what are now North America, Europe, and Asia, during the Cretaceous Period, although ancestral forms lived earlier, in the Jurassic. ...
dinosaur Dinosaurs are a diverse group of reptiles of the clade Dinosauria. They first appeared during the Triassic period, between 243 and 233.23 million years ago (mya), although the exact origin and timing of the evolution of dinosaurs is t ...
from the
Late Cretaceous The Late Cretaceous (100.5–66 Ma) is the younger 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 ''creta'', the ...
of
Canada Canada is a country in North America. Its ten provinces and three territories extend from the Atlantic Ocean to the Pacific Ocean and northward into the Arctic Ocean, covering over , making it the world's second-largest country by tot ...
.


Discovery

In 2010, Canadian fossil hunter Wendy Sloboda found a centrosaurine
bonebed A bone bed is any geological stratum or deposit that contains bones of whatever kind. Inevitably, such deposits are sedimentary in nature. Not a formal term, it tends to be used more to describe especially dense collections such as Lagerstätte. ...
in the Pinhorn Provincial Grazing Reserve south of the Milk River,
County of Forty Mile No. 8 The County of Forty Mile No. 8 is a municipal district in south eastern Alberta, Canada. It is located in Census Division 1, southwest of Medicine Hat. Its municipal office is located in the Village of Foremost. Geography Communities and ...
,
Alberta Alberta ( ) is one of the thirteen provinces and territories of Canada. It is part of Western Canada and is one of the three prairie provinces. Alberta is bordered by British Columbia to the west, Saskatchewan to the east, the Northwest Ter ...
. In 2011, a team of the
Royal Tyrrell Museum The Royal Tyrrell Museum of Palaeontology (RTMP, and often referred to as the Royal Tyrrell Museum) is a palaeontology museum and research facility in Drumheller, Alberta, Canada. The museum was named in honour of Joseph Burr Tyrrell, and is situa ...
explored the site and began excavating fossils. Next year, a deep overburden was removed. In 2013 and 2014, numerous fossils were discovered. In 2015, paleontologists David Evans and Michael Ryan named and described the
type species In 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 species that contains the biological type specimen ...
''Wendiceratops pinhornensis''. The generic name combines a reference to Wendy Sloboda with a Latinised Greek ~ceratops, "horn face". 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 ...
refers to the provenance from the Pinhorn Reserve. ''Wendiceratops'' was the second ceratopsian discovered and named in 2015. The discovery of this new horned dinosaur species - starting from the ground in Alberta, through scientific study and restoration, all the way to its eventual reconstruction and public display in The
Royal Ontario Museum The Royal Ontario Museum (ROM) is a museum of art, world culture and natural history in Toronto, Ontario, Canada. It is one of the largest museums in North America and the largest in Canada. It attracts more than one million visitors every year ...
(ROM) - was captured in the HISTORY series and companion website Dino Hunt Canada.http://dinohuntcanada.history.ca/ The
holotype A holotype is a single physical example (or illustration) of an organism, known to have been used when the species (or lower-ranked taxon) was formally described. It is either the single such physical example (or illustration) or one of several ...
, TMP 2011.051.0009, was found in a layer of the
Oldman Formation The Oldman Formation is a stratigraphic unit of Late Cretaceous (Campanian stage) age that underlies much of southern Alberta, Canada. It consists primarily of sandstones that were deposited in fluvial channel and floodplain environments. It was ...
, dating from the
Campanian The Campanian is the fifth of six ages of the Late Cretaceous Epoch on the geologic timescale of the International Commission on Stratigraphy (ICS). In chronostratigraphy, it is the fifth of six stages in the Upper Cretaceous Series. Campanian s ...
. The layer of mudstone, thick and probably deposited in a single event, has an upper possible age of 79 million years and a lower age of 78.7 million years. The holotype consists of a right
parietal bone The parietal bones () are two bones in the Human skull, skull which, when joined at a fibrous joint, form the sides and roof of the Human skull, cranium. In humans, each bone is roughly quadrilateral in form, and has two surfaces, four borders, an ...
. Under the assumption that the bonebed contained only a single centrosaurine species, all additional centrosaurine material was referred to ''Wendiceratops pinhornensis'', for a total of 184 specimens representing several individuals, including juveniles. The bones comprise elements from the skull, lower jaws, vertebral column, shoulder girdle, pelvis and limbs. They are largely disarticulated. In addition to the ''Wendiceratops'' bones, remains were found of
tyrannosaurid Tyrannosauridae (or tyrannosaurids, meaning "tyrant lizards") is a family of coelurosaurian theropod dinosaurs that comprises two subfamilies containing up to thirteen genera, including the eponymous ''Tyrannosaurus''. The exact number of genera ...
theropods Theropoda (; ), whose members are known as theropods, is a dinosaur clade that is characterized by hollow bones and three toes and claws on each limb. Theropods are generally classed as a group of saurischian dinosaurs. They were ancestrally c ...
, crocodiles,
gar Gars are members of the family Lepisosteidae, which are the only surviving members of the Ginglymodi, an ancient holosteian group of ray-finned fish, which first appeared during the Triassic, over 240 million years ago. Gars comprise seven livin ...
s and plants.


Description


Size and distinguishing traits

''Wendiceratops'' reached in length and in body mass. The describing authors indicated two unique traits, or
autapomorphies In phylogenetics, an autapomorphy is a distinctive feature, known as a derived trait, that is unique to a given taxon. That is, it is found only in one taxon, but not found in any others or outgroup taxa, not even those most closely related to t ...
: On the rear rim of the neck frill, the second and third epiparietals, which are skin ossifications attached to the rim, have a wide base, are vertically thick and curve obliquely upwards to the front, overhanging the rear and outer side branches of the parietal bone. The
ischium The ischium () form ...
has an expanded rectangular lower end. ''Wendiceratops'' is further distinguished by the presence of an erect nose horn, and the lack of a vertical spike on the rear frill. The Chinese ''Sinoceratops'' has similar epiparietals, but in addition a central midline epiparietal and bumps below the epiparietals, which are lacking in ''Wendiceratops''.


Skeleton

The
maxilla The maxilla (plural: ''maxillae'' ) in vertebrates is the upper fixed (not fixed in Neopterygii) bone of the jaw formed from the fusion of two maxillary bones. In humans, the upper jaw includes the hard palate in the front of the mouth. The t ...
, the upper jaw bone, of ''Wendiceratops'' had at least twenty-six tooth positions, in each of which several teeth were stacked to form a tooth battery. No orbital horns have been found. The
nasal bone The nasal bones are two small oblong bones, varying in size and form in different individuals; they are placed side by side at the middle and upper part of the face and by their junction, form the bridge of the upper one third of the nose. Eac ...
bore a vertical nasal horn. The exact size and profile of this horn are unknown but one broken specimen has a height of and a base length of . Contact facets with the
prefrontal bone The prefrontal bone is a bone separating the lacrimal and frontal bones in many tetrapod skulls. It first evolved in the sarcopterygian clade Rhipidistia, which includes lungfish and the Tetrapodomorpha. The prefrontal is found in most modern and ...
suggest that the horn was located just in front of the eye sockets. The skull frill of ''Wendiceratops'' was broad. The front side of the frill was formed by the
squamosal bone The squamosal is a skull bone found in most reptiles, amphibians, and birds. In fishes, it is also called the pterotic bone. In most tetrapods, the squamosal and quadratojugal bones form the cheek series of the skull. The bone forms an ancestral co ...
which had a rectangular shape. To its edge four skin ossifications or
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 were attached, the episquamosals. They had an asymmetrical triangular shape. Uniquely, they curled upwards. From the position of the fourth, frontmost, episquamosal a relatively high ridge was running to the inside, adorned with three substantial bumps. In cross-section, this ridge gave the squamosal the typical "stepped" centrosaurine profile. The rear of the frill consisted of the rounded
parietal bone The parietal bones () are two bones in the Human skull, skull which, when joined at a fibrous joint, form the sides and roof of the Human skull, cranium. In humans, each bone is roughly quadrilateral in form, and has two surfaces, four borders, an ...
s. They too had osteoderms attached to their edges, in this case called epiparietals. Each parietal had five of them, conventionally numbered "p1" to "p5". The fifth and fourth epiparietals, the ones in front, resembled the episquamosals. However, more to the back the epiparietals became increasingly wider, longer and thicker, curving upwards and to the front. This resulted in the first epiparietal becoming tongue-shaped and overhanging a large part of the frill, slightly pointing outwards. In this area the frill edge continued the curvature of these osteoderms, concavely curling upwards. Close to the edge two paired large openings were present, the parietal fenestrae. As no complete frill has been found, their exact form is uncertain but they probably were shaped like transversely oriented ovals. These openings were separated by a broad bone bar which featured a midline ridge that was rather smooth, not adorned by bumps. The bar ended at a gentle embayment of the frill rear edge. This lacked a central epiparietal or "p0". Whereas centrosaurine species each possessed a unique pattern of frill ornamentation, their postcranial skeleton, the parts of the body behind the skull, was very conservative, i.e. it showed little variation. Accordingly, the describers could find little of distinction among the ''Wendiceratops'' postcranial bones. An exception was the
ischium The ischium () form ...
. In other centrosaurines this pelvic bone had, in side view, the widest point of its shaft at midlength. The ischium of ''Wendiceratops'' however, grew wider towards its lower end, resulting in a generally rectangular shape. Though most specimens were robust, apparently of adult individuals, some bones were of juveniles. Two shinbones were found with a length of just twenty centimetres.


Classification

''Wendiceratops'' was placed in the
Centrosaurinae Centrosaurinae (from the Greek, meaning "pointed lizards") is a subfamily of ceratopsid dinosaurs, a group of large quadrupedal ornithischians. Centrosaurine fossil remains are known primarily from the northern region of Laramidia (modern day Al ...
, as a
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 t ...
to ''
Sinoceratops ''Sinoceratops'' is an extinct genus of ceratopsian dinosaur that lived approximately 73 million years ago during the latter part of the Cretaceous Period in what is now Shandong province in China. It was named in 2010 by Xu Xing ''et al.'' fo ...
''. Their
clade A clade (), also known as a monophyletic group or natural group, is a group of organisms that are monophyletic – that is, composed of a common ancestor and all its lineal descendants – on a phylogenetic tree. Rather than the English term, ...
formed a "comb" or
polytomy An internal node of a phylogenetic tree is described as a polytomy or multifurcation if (i) it is in a rooted tree and is linked to three or more child subtrees or (ii) it is in an unrooted tree and is attached to four or more branches. A tr ...
with ''
Albertaceratops ''Albertaceratops'' (meaning "Alberta horned face") was a genus of centrosaurine horned dinosaur from the middle Campanian-age Upper Cretaceous Oldman Formation of Alberta, Canada. Description ''Albertaceratops'' is unusual in combining long ...
'', another centrosaurine from the same formation, and the clade of the more derived centrosaurines. ''Wendiceratops'' and ''Sinoceratops'' were more derived, higher in the evolutionary tree, than ''
Xenoceratops ''Xenoceratops'' (meaning "alien horned face") is a genus of centrosaurine ceratopsid dinosaur known from the Late Cretaceous (middle Campanian stage), and is known to have lived in what is currently Alberta, Canada. The genus has one known speci ...
''. Their nevertheless in general rather basal position is in accordance with the high absolute age of ''Wendiceratops'' living only a million years later than the oldest known centrosaurines ''Xenoceratops'' and ''
Diabloceratops ''Diabloceratops'' is an extinct genus of centrosaurine ceratopsian dinosaur that lived approximately 81.4-81 million years ago during the latter part of the Cretaceous Period in what is now Utah, in the United States. ''Diabloceratops'' was a ...
''. The latest phylogenetic analysis including ''Wendiceratops'' is reproduced below, after Chiba ''et al.'' (2017): The nose horn of ''Wendiceratops'', erect but probably of moderate size, was by the describing authors seen as a transition between the low horns of earlier forms like ''Diabloceratops'', ''
Nasutoceratops ''Nasutoceratops'' is an extinct genus of ceratopsian dinosaur. It is a basal centrosaurine which lived during the Late Cretaceous Period (late Campanian, about 76.0-75.5 Ma). Fossils have been found in southern Utah, United States. ''Nasutocera ...
'' and ''
Albertaceratops ''Albertaceratops'' (meaning "Alberta horned face") was a genus of centrosaurine horned dinosaur from the middle Campanian-age Upper Cretaceous Oldman Formation of Alberta, Canada. Description ''Albertaceratops'' is unusual in combining long ...
'' and the much taller horns of derived centrosaurines such as ''
Coronosaurus ''Coronosaurus'' is a genus of centrosaurine ceratopsian dinosaurs which lived in the Late Cretaceous, in the middle Campanian stage. Its remains, two bone beds, were discovered by Phillip J. Currie in the Oldman Formation of Alberta, Canada, and ...
'', ''
Centrosaurus ''Centrosaurus'' ( ; ) is a genus of herbivorous ceratopsian dinosaur from the Late Cretaceous of Canada. Their remains have been found in the Dinosaur Park Formation, dating from 76.5 to 75.5 million years ago. Discovery and naming The firs ...
'' and ''
Styracosaurus ''Styracosaurus'' ( ; meaning "spiked lizard" from the Ancient Greek / "spike at the butt-end of a spear-shaft" and / "lizard") is a genus of herbivorous ceratopsian dinosaur from the Cretaceous Period (Campanian stage), about 75.5 to 74.5&nbs ...
''. This would imply that the also long vertical nasal horn of the
Chasmosaurinae Chasmosaurinae is a subfamily of ceratopsid dinosaurs. They were one of the most successful groups of herbivores of their time. Chasmosaurines appeared in the early Campanian, and became extinct, along with all other non-avian dinosaurs, during ...
would have separately been developed in a process of
parallel evolution Parallel evolution is the similar development of a trait in distinct species that are not closely related, but share a similar original trait in response to similar evolutionary pressure.Zhang, J. and Kumar, S. 1997Detection of convergent and paral ...
, the second time such a horn would have evolved among the ceratopsids. Remarkably, the curled epiparietals of ''Wendiceratops'' resemble the osteoderms of two chasmosaurines: ''
Vagaceratops ''Vagaceratops'' (meaning "wandering (''vagus'', Latin) horned face", in reference to its close relationship with ''Kosmoceratops'' from Utah) is a genus of herbivorous ceratopsian dinosaur. It is a chasmosaurine ceratopsian which lived during ...
'' and ''
Kosmoceratops ''Kosmoceratops'' () is a genus of ceratopsid dinosaur that lived in North America about 76–75.9 million years ago during the Late Cretaceous period. Specimens were discovered in Utah in the Kaiparowits Formation of the Grand Staircase–Esc ...
'', though they differ from the latter in their saddle-shaped interspaces.


Paleobiology

Being found in the Oldman Formation, ''Wendiceratops'' shared its habitat with other dinosaurs such as ''
Daspletosaurus ''Daspletosaurus'' ( ; meaning "frightful lizard") is a genus of tyrannosaurid dinosaur that lived in Laramidia between about 79.5 and 74 million years ago, during the Late Cretaceous Period. The genus ''Daspletosaurus'' contains three species ...
'', ''
Paronychodon ''Paronychodon'' (meaning "beside claw tooth") was a theropod dinosaur genus. It is a tooth taxon, often considered dubious because of the fragmentary nature of the fossils, which include "buckets" of teeth from many disparate times and places bu ...
'', ''
Troodon ''Troodon'' ( ; ''Troödon'' in older sources) is a wastebasket taxon and a dubious genus of relatively small, bird-like dinosaurs known definitively from the Campanian age of the Late Cretaceous period (about 77  mya). It includes at least ...
'', ''
Albertaceratops ''Albertaceratops'' (meaning "Alberta horned face") was a genus of centrosaurine horned dinosaur from the middle Campanian-age Upper Cretaceous Oldman Formation of Alberta, Canada. Description ''Albertaceratops'' is unusual in combining long ...
'', and ''
Parasaurolophus ''Parasaurolophus'' (; meaning "near crested lizard" in reference to '' Saurolophus)'' is a genus of herbivorous hadrosaurid ornithopod dinosaur that lived in what is now North America and possibly Asia during the Late Cretaceous Period, abou ...
''. The authors pointed out that after the discovery of ''Wendiceratops'', no less than five ceratopids are known from the lower Oldman or coeval North-American formations, in addition to ''Albertaceratops'' also ''
Judiceratops ''Judiceratops'' ( ; meaning " Judith River horned face") is an extinct horned dinosaur. It lived around 78 million years ago, during the Late Cretaceous Period in what is now Montana, United States. Like other horned dinosaurs, ''Judiceratops ...
'', ''
Medusaceratops ''Medusaceratops'' is an extinct genus of centrosaurine ceratopsian dinosaur known from the Late Cretaceous Judith River Formation (middle Campanian stage) of Montana, northern United States. It contains a single species, ''Medusaceratops lokii'' ...
'' and ''
Avaceratops ''Avaceratops'' is a genus of small herbivorous ceratopsian dinosaurs which lived during the late Campanian during the Late Cretaceous Period in what are now the Northwest United States. Most fossils come from the Judith River Formation. Discove ...
''. They assumed this was made possible by ecological specialisations and also indicated a high
faunal turnover An evolutionary radiation is an increase in taxonomic diversity that is caused by elevated rates of speciation, that may or may not be associated with an increase in morphological disparity. Radiations may affect one clade or many, and be rap ...
, a quick succession of related species.


References


External links


Video: Michael Ryan describes ''Wendiceratops''
at
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Video: Fieldwork with David Evans
at
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{{Taxonbar, from=Q20651326 Centrosaurines Late Cretaceous dinosaurs of North America Fossil taxa described in 2015 Oldman fauna Paleontology in Alberta Campanian genus first appearances Campanian genus extinctions Ornithischian genera