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''Medusaceratops'' is an
extinct Extinction is the termination of an organism by the death of its Endling, last member. A taxon may become Functional extinction, functionally extinct before the death of its last member if it loses the capacity to Reproduction, reproduce and ...
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 ...
of
centrosaurine Centrosaurinae (from the Greek, meaning "pointed lizards") is a subfamily of ceratopsid, a group of large quadrupedal ornithischian dinosaur. Centrosaurine fossil remains are known primarily from the northern region of Laramidia (modern day Alber ...
ceratopsia Ceratopsia or Ceratopia ( or ; Ancient Greek, Greek: "horned faces") is a group of herbivore, herbivorous, beaked dinosaurs that thrived in what are now North America, Asia and Europe, during the Cretaceous Period (geology), Period, although ance ...
n
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 ...
known from the
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 ...
Judith River Formation The Judith River Formation is a fossil-bearing geologic formation in Montana, and is part of the Montana Group. It dates to the Late Cretaceous, between 79 and 75.3 million years ago, corresponding to the "Judithian" land vertebrate age. It was ...
(middle
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. Campa ...
stage) of
Montana Montana ( ) is a landlocked U.S. state, state in the Mountain states, Mountain West subregion of the Western United States. It is bordered by Idaho to the west, North Dakota to the east, South Dakota to the southeast, Wyoming to the south, an ...
, northern
United States The United States of America (USA), also known as the United States (U.S.) or America, is a country primarily located in North America. It is a federal republic of 50 U.S. state, states and a federal capital district, Washington, D.C. The 48 ...
. It contains a single species, ''Medusaceratops lokii''.Ryan, Michael J.; Russell, Anthony P., and Hartman, Scott. (2010). "A New Chasmosaurine Ceratopsid from the Judith River Formation, Montana", In: Michael J. Ryan, Brenda J. Chinnery-Allgeier, and David A. Eberth (eds), ''New Perspectives on Horned Dinosaurs: The Royal Tyrrell Museum Ceratopsian Symposium'', Indiana University Press, 656 pp. .


Discovery

The known material of ''Medusaceratops'' came from a bonebed in the badlands on the west side of Kennedy Coulee adjacent to the Milk River, in the Milk River Natural Area, near Havre,
Hill County Hill County is the name of two counties in the United States: * Hill County, Montana Hill County is a County (United States), county located in the U.S. state of Montana. As of the 2020 United States census, 2020 census, the population was 16, ...
of Montana. The material was first reported by Sweeney and Boyden (1993), who considered it to represent the southernmost occurrence of ''
Styracosaurus albertensis ''Styracosaurus'' ( ; meaning "spiked lizard" from the Ancient Greek / "spike at the butt-end of a spear-shaft" and / "lizard") is an extinct genus of herbivorous ceratopsian dinosaur from the Late Cretaceous (Campanian stage) of North America. ...
'', based on misidentified frill spikes. Trexler and Sweeney (1995) reinterpreted the spikes as eye-socket horncores and noted their similarity to those of the ''
nomen dubium In binomial nomenclature, a ''nomen dubium'' (Latin for "doubtful name", plural ''nomina dubia'') is a scientific name that is of unknown or doubtful application. Zoology In case of a ''nomen dubium,'' it may be impossible to determine whether a ...
'' ''
Ceratops montanus ''Ceratops'' (meaning "horn face") is a dubious genus of herbivorous ceratopsian dinosaur which lived during the Late Cretaceous. Its fossils have been found in the Judith River Formation in Montana. Although poorly known, ''Ceratops'' is import ...
'' from a nearby area, however, could not refer the bonebed material to any valid existing taxon. The bonebed, known as the Mansfield Bonebed honoring its landowner, is located on private land and historically has been excavated by several commercial companies. The type material of ''Medusaceratops'' and other specimens were excavated more recently and have been purchased by the
Wyoming Dinosaur Center The Wyoming Dinosaur Center is located in Thermopolis, Wyoming and is one of the few dinosaur museums in the world to have excavation sites within driving distance. The museum displays the Thermopolis Specimen of ''Archaeopteryx'', which is one of ...
from Canada Fossils, Ltd., of Calgary, Alberta. Additional material from the same excavation was purchased and accessioned by the
Royal Tyrrell Museum of Palaeontology The Royal Tyrrell Museum of Palaeontology (RTMP; 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 situate ...
. Canada Fossils, Ltd., also assembled two composite skeletons using the Mansfield Bonebed material which are in the collections of the Wyoming Dinosaur Center and the
Fukui Prefectural Dinosaur Museum The , located in Katsuyama, Fukui, Japan, is one of the leading dinosaur museums in Asia that is renowned for its exhibits of fossil specimens of dinosaurs and paleontological research. It is sited in the Nagaoyama Park ( Katsuyama Dinosaur Fore ...
, however neither of the casts has an exact reconstruction of ''Medusaceratops'' as it was later described. The name ''Medusaceratops'' was coined by
Canadian Canadians () are people identified with the country of Canada. This connection may be residential, legal, historical or cultural. For most Canadians, many (or all) of these connections exist and are collectively the source of their being ''C ...
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 ...
Michael J. Ryan of the Cleveland Museum of Natural History in 2003 in a dissertation. Its fossils were subsequently confused with those of '' Albertaceratops'', another
centrosaurine Centrosaurinae (from the Greek, meaning "pointed lizards") is a subfamily of ceratopsid, a group of large quadrupedal ornithischian dinosaur. Centrosaurine fossil remains are known primarily from the northern region of Laramidia (modern day Alber ...
ceratopsian from
Alberta Alberta is a Provinces and territories of Canada, province in Canada. It is a part of Western Canada and is one of the three Canadian Prairies, prairie provinces. Alberta is bordered by British Columbia to its west, Saskatchewan to its east, t ...
which had been described by Ryan in 2007. Later, Ryan realized that the Mansfield Bonebed fossils did not belong to ''Albertaceratops''. ''Medusaceratops'' was formally described and name by Michael J. Ryan, Anthony P. Russell and Scott Hartman in
2010 The year saw a multitude of natural and environmental disasters such as the 2010 Haiti earthquake, the Deepwater Horizon oil spill, and the 2010 Chile earthquake. The 2009 swine flu pandemic, swine flu pandemic which began the previous year ...
and 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 ...
is ''Medusaceratops lokii''. The generic name refers to
Medusa In Greek mythology, Medusa (; ), also called Gorgo () or the Gorgon, was one of the three Gorgons. Medusa is generally described as a woman with living snakes in place of hair; her appearance was so hideous that anyone who looked upon her wa ...
, a monster from
Greek mythology Greek mythology is the body of myths originally told by the Ancient Greece, ancient Greeks, and a genre of ancient Greek folklore, today absorbed alongside Roman mythology into the broader designation of classical mythology. These stories conc ...
whose "hair" consists of snakes and its gaze could turn men to stone, alluding to a unique trait of this genus - the large, thick snake-like spikes that extend to the sides of the frill, in combination with
Latin Latin ( or ) is a classical language belonging to the Italic languages, Italic branch of the Indo-European languages. Latin was originally spoken by the Latins (Italic tribe), Latins in Latium (now known as Lazio), the lower Tiber area aroun ...
ized
Greek Greek may refer to: Anything of, from, or related to Greece, a country in Southern Europe: *Greeks, an ethnic group *Greek language, a branch of the Indo-European language family **Proto-Greek language, the assumed last common ancestor of all kno ...
''ceratops'', meaning "horned-face", which is a common suffix for
ceratopsia Ceratopsia or Ceratopia ( or ; Ancient Greek, Greek: "horned faces") is a group of herbivore, herbivorous, beaked dinosaurs that thrived in what are now North America, Asia and Europe, during the Cretaceous Period (geology), Period, although ance ...
n genera names. 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 ...
''lokii'' honors
Loki Loki is a Æsir, god in Norse mythology. He is the son of Fárbauti (a jötunn) and Laufey (mythology), Laufey (a goddess), and the brother of Helblindi and Býleistr. Loki is married to the goddess Sigyn and they have two sons, Narfi (son of Lo ...
, a troublemaking god in the
Norse mythology Norse, Nordic, or Scandinavian mythology, is the body of myths belonging to the North Germanic peoples, stemming from Old Norse religion and continuing after the Christianization of Scandinavia as the Nordic folklore of the modern period. The ...
, in reference to the years confusion that surrounded the taxonomic designations of the Mansfield Bonebed material before it was given its own name. The Mansfield Bonebed material was collected from the upper part of the
Judith River Formation The Judith River Formation is a fossil-bearing geologic formation in Montana, and is part of the Montana Group. It dates to the Late Cretaceous, between 79 and 75.3 million years ago, corresponding to the "Judithian" land vertebrate age. It was ...
, in a region where it is
lithologically The lithology of a rock unit is a description of its physical characteristics visible at outcrop, in hand or core samples, or with low magnification microscopy. Physical characteristics include colour, texture, grain size, and composition. Lith ...
equivalent to 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 ...
of
Canada Canada is a country in North America. Its Provinces and territories of Canada, ten provinces and three territories extend from the Atlantic Ocean to the Pacific Ocean and northward into the Arctic Ocean, making it the world's List of coun ...
. The bonebed is located at approximately the same level as the holotype of '' Albertaceratops'', dating to 77.5
million years ago Million years ago, abbreviated as Mya, Myr (megayear) or Ma (megaannum), is a unit of time equal to (i.e. years), or approximately 31.6 teraseconds. Usage Myr is in common use in fields such as Earth science and cosmology. Myr is also used w ...
, to the middle
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. Campa ...
stage of the Late Cretaceous. Thus, ''Medusaceratops'' was considered to represent the oldest known
chasmosaurine 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 ...
, until the naming of ''
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'' ...
'' by Longrich (2013), also from the Judith River Formation, but from an area equivalent to the lower Oldman Formation or upper
Foremost Formation The Foremost Formation is a stratigraphic unit of Late Cretaceous (Campanian) age that underlies much of southern Alberta, Canada. It was named for outcrops in Chin Coulee near the town of ForemostGlass, D.J. (editor) 1997. Lexicon of Canadian ...
. However, Ryan had already indicated that part of the Mansfield material represented not a chasmosaurine, but a centrosaurine. Chiba ''et al.'' in 2017 described new material of ''Medusaceratops'' from the Mansfield Bonebed, found in 2011 and 2012 by David Trexler, indicating the presence of traits that were characteristic of Centrosaurinae in the skeleton of ''M. lokii''. They concluded that all the material could be referred to a single species. The phylogenetic analysis conducted by the authors indicated that ''Medusaceratops'' was not a member of Chasmosaurinae after all, but rather an early centrosaurine ceratopsid that was more closely related to Centrosaurini and
Pachyrhinosaurini Centrosaurinae (from the Greek, meaning "pointed lizards") is a subfamily of ceratopsid, a group of large quadrupedal ornithischian dinosaur. Centrosaurine fossil remains are known primarily from the northern region of Laramidia (modern day Alber ...
than
Nasutoceratopsini Centrosaurinae (from the Greek, meaning "pointed lizards") is a subfamily of ceratopsid, a group of large quadrupedal ornithischian dinosaur. Centrosaurine fossil remains are known primarily from the northern region of Laramidia (modern day Alber ...
.


Description

Two partial parietals (frills) that are housed at the Wyoming Dinosaur Center were chosen as the type material of ''Medusaceratops'', including the
holotype A holotype (Latin: ''holotypus'') is a single physical example (or illustration) of an organism used when the species (or lower-ranked taxon) was formally described. It is either the single such physical example (or illustration) or one of s ...
WDC DJR 001 and the
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 (biology), isotype ...
WDC DJR 002. Although all the
chasmosaurine 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 ...
material from the Mansfield Bonebed was thought to be referable to ''Medusaceratops'', which totals at several hundred individual elements, upon its original description only these two partial frills have been scientifically described while the rest of the material was being reexamined. Many of the other elements are not diagnosable to a genus level and can only be confidently referable to
Ceratopsidae Ceratopsidae (sometimes spelled Ceratopidae) is a family of ceratopsian dinosaurs including ''Triceratops'', ''Centrosaurus'', and ''Styracosaurus''. All known species were quadrupedal herbivores from the Upper Cretaceous. All but one species are k ...
. Upon its original description ''Medusaceratops'' was thought to represent a
chasmosaurine 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 ...
. It was suggested that based on its type material alone, ''Medusaceratops'' is unique among
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, durin ...
in having only three epiparietals (frill spikes) on each side of the frill. The first frill spike pair is large and the second is smaller, and both are uniquely widened at base, pachyostotic, curve down the sides of the frill, and are depressed down the front of the frill. The third frill spike pair is small and triangular, unmodified in comparison to early chasmosaurines, but similarly depressed, and borders 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 ...
. The highly broadened and curved first frill spike pair of ''Medusaceratops'' closely resembles the third pair of the frill ornamentation of '' Albertaceratops'', however ''Medusaceratops'' was thought to differ (like all chasmosaurines) in lacking tab-shaped, frequently overlapping fourth to seventh pairs of the frill ornamentation of
centrosaurine Centrosaurinae (from the Greek, meaning "pointed lizards") is a subfamily of ceratopsid, a group of large quadrupedal ornithischian dinosaur. Centrosaurine fossil remains are known primarily from the northern region of Laramidia (modern day Alber ...
s. This was challenged in 2018, by the description of additional Mansfield Bonebed material assignable to ''Medusaceratops''. It became apparent that the 1-3 frill spikes mentioned above are in fact spikes 2–4. The first epiparietal is small and variably procurving and thus was misinterpreted before. At least one more epiparietal pair was also identified (after the fourth), resulting in a total of at least 5 pairs, consistent with centrosaurines like '' Albertaceratops'' and ''
Wendiceratops ''Wendiceratops'' is a genus of herbivorous centrosaurine ceratopsian dinosaur from the Late Cretaceous of Canada. Discovery In 2010, Canadian fossil hunter Wendy Sloboda found a centrosaurine bonebed in the Pinhorn Provincial Grazing Reserve ...
'', but not with chasmosaurines. The midline ramus of ''Medusaceratops'', a bone separating the two sides of the frill, was also among the newly described material. It is broad, resulting in rounder and smaller frill fenestrae (holes) like in other centrosaurines. Thus, the new study reassigned ''Medusaceratops'' to
Centrosaurinae Centrosaurinae (from the Greek, meaning "pointed lizards") is a subfamily of ceratopsid, a group of large quadrupedal ornithischian dinosaur. Centrosaurine fossil remains are known primarily from the northern region of Laramidia (modern day Alber ...
, among which it is most similar to ''Albertaceratops'' and ''Wendiceratops''. In 2010, a length of roughly 6 meters (~20 feet) was estimated for ''Medusaceratops lokii''.


Classification

Among valid
ceratopsid Ceratopsidae (sometimes spelled Ceratopidae) is a family of ceratopsian dinosaurs including ''Triceratops'', '' Centrosaurus'', and '' Styracosaurus''. All known species were quadrupedal herbivores from the Upper Cretaceous. All but one species are ...
s from the
Judith River Formation The Judith River Formation is a fossil-bearing geologic formation in Montana, and is part of the Montana Group. It dates to the Late Cretaceous, between 79 and 75.3 million years ago, corresponding to the "Judithian" land vertebrate age. It was ...
, ''Medusaceratops'' can be directly distinguished from centrosaurine ''
Avaceratops ''Avaceratops'' is a genus of small herbivorous ceratopsian dinosaurs which lived during the late Campanian in what are now the Northwest United States. Most fossils come from the Judith River Formation. Discovery and naming The first remains ...
'', and chasmosaurines ''
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'' ...
'' and ''
Spiclypeus ''Spiclypeus'' (meaning "spike shield") is an extinct genus of chasmosaurine ceratopsian dinosaur known from the Late Cretaceous Judith River Formation (late Campanian stage) of Montana, United States. Discovery In 2000, Bill D. Shipp, a nuclea ...
'' based on its unique frill ornamentation. It differs from chasmosaurine ''
Mercuriceratops ''Mercuriceratops'' is an extinct genus of herbivorous chasmosaurine ceratopsid dinosaur known from the Late Cretaceous (Campanian stage) of Alberta, Canada and Montana, United States. It contains a single species, ''Mercuriceratops gemini''. D ...
'' based on its less unique
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 ...
, as evident from newly described squamosal bones of ''Medusaceratops''. All material previously assigned to the centrosaurine ''Albertaceratops'' from the formation is now assigned to ''Medusaceratops'' or considered too fragmentary. The
cladogram A cladogram (from Greek language, Greek ''clados'' "branch" and ''gramma'' "character") is a diagram used in cladistics to show relations among organisms. A cladogram is not, however, an Phylogenetic tree, evolutionary tree because it does not s ...
presented below follows a phylogenetic analysis by Chiba ''et al.'' (2017) who redescribed ''Medusaceratops'' as a centrosaurine:


See also

*
Timeline of ceratopsian research This timeline of ceratopsian research is a chronological listing of events in the History of paleontology, history of paleontology focused on the ceratopsians, a group of herbivorous marginocephalian dinosaurs that evolved parrot-like beaks, b ...


References

{{Taxonbar, from=Q2272018 Centrosaurinae Dinosaur genera Campanian dinosaurs Judith River Formation Fossil taxa described in 2010 Dinosaurs of the United States