Medusaceratops Lokii
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''Medusaceratops'' is an
extinct Extinction is the termination of a kind of organism or of a group of kinds (taxon), usually a species. The moment of extinction is generally considered to be the death of the last individual of the species, although the capacity to breed and ...
genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In the hierarchy of biological classification, genus com ...
of
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 ...
ceratopsia 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. ...
n
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 ...
known 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 ...
Judith River Formation (middle Campanian stage) of
Montana Montana () is a state in the Mountain West division of the Western United States. It is bordered by Idaho to the west, North Dakota and South Dakota to the east, Wyoming to the south, and the Canadian provinces of Alberta, British Columbi ...
, northern
United States The United States of America (U.S.A. or USA), commonly known as the United States (U.S. or US) or America, is a country primarily located in North America. It consists of 50 states, a federal district, five major unincorporated territorie ...
. 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 Milk River Natural Area is a provincially designated protected area in the southeastern corner of the province of Alberta, Canada, approximately 160 km south of the city of Medicine Hat. It lies primarily in the County of Forty Mile with a s ...
, near Havre, Hill County of Montana. The material was first reported by Sweeney and Boyden (1993), who considered it to represent the southernmost occurrence of '' Styracosaurus albertensis'', 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 s ...
'' ''
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 importa ...
'' 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 the onl ...
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, 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 situ ...
. 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 For ...
, however neither of the casts has an exact reconstruction of ''Medusaceratops'' as it was later described. The name ''Medusaceratops'' was coined by
Canadian Canadians (french: Canadiens) 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 ...
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 fossi ...
Michael J. Ryan of the Cleveland Museum of Natural History in 2003 in a dissertation. Its fossils were subsequently confused with those of ''
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 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 from
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 ...
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 File:2010 Events Collage New.png, From top left, clockwise: The 2010 Chile earthquake was one of the strongest recorded in history; The Eruption of Eyjafjallajökull in Iceland disrupts air travel in Europe; A scene from the opening ceremony of ...
and 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 ...
is ''Medusaceratops lokii''. The generic name refers to Medusa, a monster from
Greek mythology A major branch of classical mythology, Greek mythology is the body of myths originally told by the Ancient Greece, ancient Greeks, and a genre of Ancient Greek folklore. These stories concern the Cosmogony, origin and Cosmology#Metaphysical co ...
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 branch of the Indo-European languages. Latin was originally a dialect spoken in the lower Tiber area (then known as Latium) around present-day Rome, but through the power of the ...
ized Greek ''ceratops'', meaning "horned-face", which is a common suffix for
ceratopsia 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. ...
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 god in Norse mythology. According to some sources, Loki is the son of Fárbauti (a jötunn) and Laufey (mentioned as a goddess), and the brother of Helblindi and Býleistr. Loki is married to Sigyn and they have two sons, Narfi or Na ...
, 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, and into the Nordic folklore of the modern period ...
, 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, in a region where it is lithologically equivalent to the Oldman Formation 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 ...
. The bonebed is located at approximately the same level as the holotype of ''
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 ...
'', dating to 77.5
million years ago The abbreviation Myr, "million years", is a unit of a quantity of (i.e. ) years, or 31.556926 teraseconds. Usage Myr (million years) is in common use in fields such as Earth science and cosmology. Myr is also used with Mya (million years ago). ...
, to the middle Campanian 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 t ...
, 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 formation (stratigraphy), 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 Foremost, Alberta, ForemostGlas ...
. 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 than
Nasutoceratopsini 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 ...
.


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 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 ...
WDC DJR 001 and the paratype 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 t ...
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 t ...
. 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, during ...
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 highly broadened and curved first frill spike pair of ''Medusaceratops'' closely resembles the third pair of the frill ornamentation of ''
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 ...
'', 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 dinosaurs, a group of large quadrupedal ornithischians. Centrosaurine fossil remains are known primarily from the northern region of Laramidia (modern day Al ...
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 ''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 '' Wendiceratops'', 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, 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 k ...
s from the Judith River Formation, ''Medusaceratops'' can be directly distinguished from centrosaurine '' Avaceratops'', 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, 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 presented below follows a phylogenetic analysis by Chiba ''et al.'' (2017) who redescribed ''Medusaceratops'' as a centrosaurine:


See also

* Timeline of ceratopsian research


References

{{Taxonbar, from=Q2272018 Centrosaurines Late Cretaceous dinosaurs of North America Fossil taxa described in 2010 Paleontology in Montana Campanian genus first appearances Campanian genus extinctions Ornithischian genera