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''Bagaceratops'' (meaning "small-horned face") is a
genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of living and fossil organisms as well as viruses. In the hierarchy of biological classification, genus comes above species and below family. In binomial nom ...
of small protoceratopsid
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 ...
s that lived in
Asia Asia (, ) is one of the world's most notable geographical regions, which is either considered a continent in its own right or a subcontinent of Eurasia, which shares the continental landmass of Afro-Eurasia with Africa. Asia covers an are ...
during 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'', ...
, around 72 to 71 million years ago. ''Bagaceratops'' remains have been reported from the Barun Goyot Formation and Bayan Mandahu Formation. One specimen may argue the possible presence of ''Bagaceratops'' in the Djadochta Formation. ''Bagaceratops'' was among the smallest ceratopsians, growing up to in length, with a weight about . Although emerging late in the reign of the dinosaurs, ''Bagaceratops'' had a fairly primitive anatomy—when compared to the much derived
ceratopsids 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 ar ...
—and kept the small body size that characterized early ceratopsians. Unlike its close relative, '' Protoceratops'', ''Bagaceratops'' lacked premaxillary teeth (cylindrical, blunt teeth near the tip of the upper jaw).


History of discovery

During the large field work of the Polish-Mongolian Palaeontological Expeditions in the 1970s, abundant protoceratopsid specimens were discovered on eroded surfaces of the Hermiin Tsav locality of the Barun Goyot Formation,
Gobi Desert The Gobi Desert ( Chinese: 戈壁 (沙漠), Mongolian: Говь (ᠭᠣᠪᠢ)) () is a large desert or brushland region in East Asia, and is the sixth largest desert in the world. Geography The Gobi measures from southwest to northeast a ...
. This newly collected and rich fossil material is stored in the Institute of Paleobiology of the
Polish Academy of Sciences The Polish Academy of Sciences ( pl, Polska Akademia Nauk, PAN) is a Polish state-sponsored institution of higher learning. Headquartered in Warsaw, it is responsible for spearheading the development of science across the country by a society o ...
(
Poland Poland, officially the Republic of Poland, is a country in Central Europe. It is divided into 16 administrative provinces called voivodeships, covering an area of . Poland has a population of over 38 million and is the fifth-most populou ...
). In 1975, two of the expedition's leading scientists, namely Polish paleontologists Teresa Maryańska and Halszka Osmólska, published a large
monograph A monograph is a specialist work of writing (in contrast to reference works) or exhibition on a single subject or an aspect of a subject, often by a single author or artist, and usually on a scholarly subject. In library cataloging, ''monogra ...
dedicated to describe this material where they named the new genus and type species of protoceratopsid ''Bagaceratops rozhdestvenskyi''. The selected
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 seve ...
is ZPAL MgD-I/126, which consists of a relatively medium-sized skull, and a vast majority of the specimens collected by the expeditions were assigned to ''Bagaceratops'', including juvenile and sub-adult skulls. The generic name, ''Bagaceratops'', means "small-horned face" and is derived from the Mongolian ''baga'' = meaning small; and Greek ''ceratops'' = meaning horn face. The specific name, ''B. rozhdestvenskyi'', is named in honor of the Russian
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 ...
Anatoly Konstantinovich Rozhdestvensky for his notorious work on dinosaurs.


Additional specimens

In 1993, the Japan-Mongolia Joint Paleontological Expedition collected an articulated and nearly complete ''Bagaceratops'' skeleton (MPC-D 100/535) from the Barun Goyot Formation at the Hermiin Tsav locality. In 2010 and 2011 this specimen was examined to analyze several borings (tunnel-like holes) left by invertebrae scavengers on joint areas. As of 2019, MPC-D 100/535 remains largely undescribed. In 2019 a partial skeleton (specimen KID 196) of ''Bagaceratops'' was described by Bitnara Kim and colleagues, who noted no significant diifferences between the skeleton of ''Protoceratops'' and the former. This specimen was discovered in 2007 also from the Hermiin Tsav locality of the Barun Goyot Formation, and includes a partially preserved skull with partial skeleton of an adult individual. In 2020, Czepiński described new specimens of ''Bagaceratops'' and ''Protoceratops'' from the Udyn Sayr and Zamyn Khond localities, respectively, of the Djadochta Formation, and evaluated the implications of these specimens for correlation of fossil sites of the latter formation. He considered one of these specimens in particular, MPC-D 100/551B, as a potential evidence of an
anagenetic Anagenesis is the gradual evolution of a species that continues to exist as an interbreeding population. This contrasts with cladogenesis, which occurs when there is branching or splitting, leading to two or more lineages and resulting in separate ...
transition from ''Protoceratops andrewsi'' to ''Bagaceratops rozhdestvenskyi''.


Synonyms

Juvenile remains, at first tentatively named ''Protoceratops kozlowskii'', and then renamed '' Breviceratops kozlowskii'' by Kurzanov in 1990, were considered to be juvenile ''Bagaceratops''.
Paul Sereno Paul Callistus Sereno (born October 11, 1957) is a professor of paleontology at the University of Chicago and a National Geographic "explorer-in-residence" who has discovered several new dinosaur species on several continents, including at si ...
in 2000 explained this by extrapolating that the juvenile ''Breviceratops'' would grow into a mature ''Bagaceratops''. In 2003 Russian paleontologist Vladimir R. Alifanov named the new taxa ''Lamaceratops tereschenkoi'' and ''Platyceratops tatarinovi'' from the Barun Goyot Formation. Material assigned by Alifanov corresponds to the holotype of ''Lamaceratops'' (PIN 4487/26; a partial small-sized skull), recovered from the Khulsan locality, and the holotype of ''Platyceratops'' (PIN 3142/4; an almost complete medium-sized skull), found in the
red beds Red beds (or redbeds) are sedimentary rocks, typically consisting of sandstone, siltstone, and shale, that are predominantly red in color due to the presence of ferric oxides. Frequently, these red-colored sedimentary strata locally contain t ...
of the Hermiin Tsav locality. He also coined the family Bagaceratopidae in order to contain these new taxa and ''Bagaceratops''. Also during 2003,
You Hailu In Modern English, ''you'' is the second-person pronoun. It is grammatically plural, and was historically used only for the dative case, but in most modern dialects is used for all cases and numbers. History ''You'' comes from the Proto- ...
and Dong Zhiming described and named the new genus and species of protoceratopsid ''Magnirostris dodsoni'' from red beds at the Bayan Mandahu locality of the Bayan Mandahu Formation,
Inner Mongolia Inner Mongolia, officially the Inner Mongolia Autonomous Region, is an autonomous region of the People's Republic of China. Its border includes most of the length of China's border with the country of Mongolia. Inner Mongolia also accounts for a ...
(
China China, officially the People's Republic of China (PRC), is a country in East Asia. It is the world's List of countries and dependencies by population, most populous country, with a Population of China, population exceeding 1.4 billion, slig ...
). The holotype of ''Magnirostris'', IVPP V12513, represents a nearly complete skull lacking the frill region of a large individual and was collected during expeditions lead by the Sino-Canadian Dinosaur Project. In 2006 Mackoviky regarded all of these ceratopsians as
junior synonym The Botanical and Zoological Codes of nomenclature treat the concept of synonymy differently. * In botanical nomenclature, a synonym is a scientific name that applies to a taxon that (now) goes by a different scientific name. For example, Linn ...
s of ''Bagaceratops'' based on the reasoning that all exhibit anatomical traits already seen on other specimens of this protoceratopsid, and some of them are likely products of preservation. In 2008 Alifanov described and named another ceratopsian taxon from the Barun Goyot Formation, ''Gobiceratops minitus''. Its holotype (PIN 3142/299) is represented by a very small and juvenile skull that was collected from the Hermiin Tsav locality near the end of the 1970s by the Joint Soviet–Mongolian Paleontological Expedition. Though Alifanov used this skull to erect the new ''Gobiceratops'', it had already been displayed for several years at the Moscow Paleontological Museum under the name ''Bagaceratops rozhdestvenskyi''. A comprehensive study on the intraspecific variation in morphology of ''B. rozhdestvenskyi'' was conducted by Polish paleontologist Łukasz Czepiński in
2019 File:2019 collage v1.png, From top left, clockwise: Hong Kong protests turn to widespread riots and civil disobedience; House of Representatives votes to adopt articles of impeachment against Donald Trump; CRISPR gene editing first used to experim ...
, where he concluded that the previously named ''Gobiceratops minutus'', ''Lamaceratops tereschenkoi'', ''Platyceratops tatarinovi'' and ''Magnirostris dodsoni'' represent additional specimens and growth stages of ''B. rozhdestvenskyi'' and therefore, junior synonyms. Czepiński rexamined many of the specimens originally described by Maryańska and Osmólska, as well as the respective holotypes of these taxa, providing evidence that all traits used to separate them are, in fact, indistinguishably present on ''Bagaceratops'' and they fall within the large intraspecific variation of this taxon. He also considered ''Breviceratops'' to be a distinct and separate genus of protoceratopsid, from both ''Bagaceratops'' and ''Protoceratops'', as it features a combination of basal (primitive) and derived (advanced) traits.


Description

''Bagaceratops'' was a small-sized protoceratopsid, reaching adult dimensions of about in length and in body mass based on ''Maginostris''. It had a smaller frill, about ten grinding teeth per jaw, and more triangular skull than its close relative, '' Protoceratops''. Although both ''Bagaceratops'' and ''Protoceratops'' were very similar (mostly in the postcranial skeleton), the former had a much derived (advanced) skull morphology. ''Bagaceratops'' lacked primitive
premaxilla The premaxilla (or praemaxilla) is one of a pair of small cranial bones at the very tip of the upper jaw of many animals, usually, but not always, bearing teeth. In humans, they are fused with the maxilla. The "premaxilla" of therian mammal has ...
ry teeth, had paired (fused) nasal bones, and an oval-shaped fenestra (hole) was developed in 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. T ...
—otherwise known as accessory antorbital fenestra.


Classification

''Bagaceratops'' belonged to
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 Jurass ...
, a group of herbivorous dinosaurs with
parrot Parrots, also known as psittacines (), are birds of the roughly 398 species in 92 genera comprising the order Psittaciformes (), found mostly in tropical and subtropical regions. The order is subdivided into three superfamilies: the Psittacoide ...
-like beaks which thrived in
North America North America is a continent in the Northern Hemisphere and almost entirely within the Western Hemisphere. It is bordered to the north by the Arctic Ocean, to the east by the Atlantic Ocean, to the southeast by South America and th ...
and
Asia Asia (, ) is one of the world's most notable geographical regions, which is either considered a continent in its own right or a subcontinent of Eurasia, which shares the continental landmass of Afro-Eurasia with Africa. Asia covers an are ...
during the Cretaceous Period, which ended roughly 66 million years ago. In 2019 Czepiński analyzed a vast majority of referred specimens to the ceratopsians ''Bagaceratops'' and ''Breviceratops'', and concluded that most were in fact specimens of the former. Although the genera ''Gobiceratops'', ''Lamaceratops'', ''Magnirostris'', and ''Platyceratops'', were long considered valid and distinct taxa, and sometimes placed within Protoceratopsidae, Czepiński found the diagnostic features used to distinguish these taxa to be largely present in ''Bagaceratops'' and thus becoming synonyms of this genus. Under this reasoning, Protoceratopsidae consists of ''Bagaceratops'', ''Breviceratops'', and ''Protoceratops''. Based on cranial characters such as presence or absence of premaxillary teeth and an antorbital fenestra, ''P. andrewsi'' is the basal-most protoceratopsid and ''Bagaceratops'' the derived-most one. Below are the proposed phylogenetic relationships within Protoceratopsidae by Czepiński:


Paleoenvironment


Barun Goyot Formation

The Barun Goyot Formation, based on
sediment Sediment is a naturally occurring material that is broken down by processes of weathering and erosion, and is subsequently transported by the action of wind, water, or ice or by the force of gravity acting on the particles. For example, sand ...
s, is regarded as
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'', ...
in age (Middle-Upper Campanian) and has virtually yielded the bulk of material for which ''Bagaceratops'' is known. This formation is mostly characterized by series of
red beds Red beds (or redbeds) are sedimentary rocks, typically consisting of sandstone, siltstone, and shale, that are predominantly red in color due to the presence of ferric oxides. Frequently, these red-colored sedimentary strata locally contain t ...
, mostly light-coloured
sand Sand is a granular material composed of finely divided mineral particles. Sand has various compositions but is defined by its grain size. Sand grains are smaller than gravel and coarser than silt. Sand can also refer to a textural class ...
s (yellowish, grey-brown, and rarely reddish) that are locally cemented. Sandy
claystone Mudrocks are a class of fine-grained siliciclastic sedimentary rocks. The varying types of mudrocks include siltstone, claystone, mudstone, slate, and shale. Most of the particles of which the stone is composed are less than and are too ...
s (often red-coloured),
siltstone Siltstone, also known as aleurolite, is a clastic sedimentary rock that is composed mostly of silt. It is a form of mudrock with a low clay mineral content, which can be distinguished from shale by its lack of fissility.Blatt ''et al.'' 1980, ...
s,
conglomerate Conglomerate or conglomeration may refer to: * Conglomerate (company) * Conglomerate (geology) * Conglomerate (mathematics) In popular culture: * The Conglomerate (American group), a production crew and musical group founded by Busta Rhymes ** ...
s, and large-scale trough cross-stratification in sands are also common across the unit. In addition, structureless, medium-grained, fine-grained and very fine-grained
sandstone Sandstone is a clastic sedimentary rock composed mainly of sand-sized (0.0625 to 2 mm) silicate grains. Sandstones comprise about 20–25% of all sedimentary rocks. Most sandstone is composed of quartz or feldspar (both silicat ...
s predominate in sediments of the Barun Goyot Formation. The sediments of this formation were deposited in
alluvial plain An alluvial plain is a largely flat landform created by the deposition of sediment over a long period of time by one or more rivers coming from highland regions, from which alluvial soil forms. A floodplain is part of the process, being the smal ...
(flat land consisting of sediments deposited by highland
river A river is a natural flowing watercourse, usually freshwater, flowing towards an ocean, sea, lake or another river. In some cases, a river flows into the ground and becomes dry at the end of its course without reaching another body of ...
s), lacustrine, and aeolian paleoenvironments, under relatively
arid A region is arid when it severely lacks available water, to the extent of hindering or preventing the growth and development of plant and animal life. Regions with arid climates tend to lack vegetation and are called xeric or desertic. Most ...
to semiarid climates. ''Bagaceratops'' is the most common taxon across the Barun Goyot Formation, which was also home to many other
vertebrate Vertebrates () comprise all animal taxa within the subphylum Vertebrata () ( chordates with backbones), including all mammals, birds, reptiles, amphibians, and fish. Vertebrates represent the overwhelming majority of the phylum Chordata, with ...
s, including the ankylosaurids '' Saichania'', '' Tarchia'' and '' Zaraapelta''; alvarezsaurids ''
Khulsanurus ''Khulsanurus'' (meaning "tail from Khulsan") is an extinct genus of alvarezsaurid theropod dinosaur from the Late Cretaceous Barungoyot Formation of the Khulsan Locality in the Gobi Desert region of Mongolia. The type and only species is ' ...
'' and '' Parvicursor'';
bird Birds are a group of warm-blooded vertebrates constituting the class Aves (), characterised by feathers, toothless beaked jaws, the laying of hard-shelled eggs, a high metabolic rate, a four-chambered heart, and a strong yet lightweig ...
s '' Gobipipus'', ''
Gobipteryx ''Gobipteryx'' (from Gobi eferring to the Gobi Desert where it was first discovered and Greek pteryx “wing”) is a genus of prehistoric bird from the Campanian Age of the Late Cretaceous Period.Elżanowski, A. (1974): Preliminary note on t ...
'' and '' Hollanda''; fellow protoceratopsid '' Breviceratops'';
dromaeosaurid Dromaeosauridae () is a family of feathered theropod dinosaurs. They were generally small to medium-sized feathered carnivores that flourished in the Cretaceous Period. The name Dromaeosauridae means 'running lizards', from Greek ('), meaning ...
s ''
Kuru Kuru may refer to: Anthropology and history * Kuru (disease), a type of transmissible spongiform encephalopathy associated with the cannibalistic funeral practices of the Fore people * Kuru (mythology), part of Meithei mythology * Kuru Kingdom, ...
'' and ''
Shri Shri (; , ) is a Sanskrit term denoting resplendence, wealth and prosperity, primarily used as an honorific. The word is widely used in South and Southeast Asian languages such as Marathi, Malay (including Indonesian and Malaysian), Javan ...
'';
halszkaraptorine Halszkaraptorinae is a basal ("primitive") subfamily of Dromaeosauridae (or possibly Unenlagiidae) that includes the enigmatic genera ''Halszkaraptor'', ''Natovenator'', ''Mahakala'', and ''Hulsanpes''. Halszkaraptorines are definitively known o ...
'' Hulsanpes''; pachycephalosaurid '' Tylocephale''; and oviraptorids '' Conchoraptor'', ''
Heyuannia ''Heyuannia'' ("from Heyuan") is a genus of oviraptorid dinosaur that lived in Asia during the Late Cretaceous epoch, in what is now China and Mongolia. It was the first oviraptorid found in China; most others were found in neighbouring Mongoli ...
'' and '' Nemegtomaia''. Other taxa are represented by the
large Large means of great size. Large may also refer to: Mathematics * Arbitrarily large, a phrase in mathematics * Large cardinal, a property of certain transfinite numbers * Large category, a category with a proper class of objects and morphisms ...
titanosaur ''
Quaesitosaurus ''Quaesitosaurus'' (meaning "extraordinary lizard") is a genus of nemegtosaurid sauropod containing only the type species, ''Q. orientalis'', described in 1983. It lived from 72 to 71 million years ago during the Late Cretaceous epoch in the Barun ...
'', and a wide diversity of
mammal Mammals () are a group of vertebrate animals constituting the class Mammalia (), characterized by the presence of mammary glands which in females produce milk for feeding (nursing) their young, a neocortex (a region of the brain), fur ...
s and squamates.


Bayan Mandahu Formation

The Bayan Mandahu Formation, which yielded ''Magnirostris'' (now synonym of ''Bagaceratops''), is considered to be Late Cretaceous in age, roughly Campanian. The dominant
lithology 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. Li ...
is reddish-brown, poorly cemented, fine grained sandstone with some conglomerate, and caliche. Other facies include
alluvial Alluvium (from Latin ''alluvius'', from ''alluere'' 'to wash against') is loose clay, silt, sand, or gravel that has been deposited by running water in a stream bed, on a floodplain, in an alluvial fan or beach, or in similar settings. ...
(
stream A stream is a continuous body of surface water flowing within the bed and banks of a channel. Depending on its location or certain characteristics, a stream may be referred to by a variety of local or regional names. Long large streams ...
-deposited) and aeolian (
wind Wind is the natural movement of air or other gases relative to a planet's surface. Winds occur on a range of scales, from thunderstorm flows lasting tens of minutes, to local breezes generated by heating of land surfaces and lasting a few ...
-deposited) sediments. It is likely that sediments at Bayan Mandahu were deposited by short-lived rivers and
lake A lake is an area filled with water, localized in a basin, surrounded by land, and distinct from any river or other outlet that serves to feed or drain the lake. Lakes lie on land and are not part of the ocean, although, like the much large ...
s on an alluvial plain with a combination of
dune A dune is a landform composed of wind- or water-driven sand. It typically takes the form of a mound, ridge, or hill. An area with dunes is called a dune system or a dune complex. A large dune complex is called a dune field, while broad, f ...
field paleoenvironments, under a semi-arid climate. The formation is known for its vertebrate fossils in life-like poses, most of which are preserved in unstructured sandstone, indicating a catastrophic rapid burial. This formation has produced numerous
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 ...
s, such as closely related protoceratopsid '' Protoceratops''; ankylosaurid '' Pinacosaurus''; alvarezsaurid '' Linhenykus''; dromaeosaurids ''
Linheraptor ''Linheraptor'' is a genus of dromaeosaurid dinosaur which lived in what is now China in the Late Cretaceous. It was named by Xu Xing and colleagues in 2010, and contains the species ''Linheraptor exquisitus''. This bird-like dinosaur was less t ...
'' and ''
Velociraptor ''Velociraptor'' (; ) is a genus of small dromaeosaurid dinosaur that lived in Asia during the Late Cretaceous epoch, about 75 million to 71 million years ago. Two species are currently recognized, although others have been assigned in th ...
''; oviraptorids ''
Machairasaurus ''Machairasaurus'' is a genus of oviraptorid dinosaur which was found in the Bayan Mandahu Formation, China, dating to the late Cretaceous period. Discovery During the Sino-Canadian expeditions of 1988 and 1990 some skeletons of unknown o ...
'' and ''
Wulatelong ''Wulatelong'' is an extinct genus of basal oviraptorid dinosaur known from the Late Cretaceous Wulansuhai Formation (Campanian stage) of Bayan Mandahu, Linhe District of Inner Mongolia, northern China. It contains a single species, ''Wul ...
''; and troodontids '' Linhevenator'', ''
Papiliovenator ''Papiliovenator'' (meaning "butterfly hunter", after a butterfly-shaped feature on its first two dorsal vertebrae) is a genus of troodontid theropod dinosaur from the Bayan Mandahu Formation of Inner Mongolia, China. The type and only species ...
'', and '' Philovenator''. Other paleofauna from this unit comprises a variety of squamates and mammals, and nanhsiungchelyids
turtle Turtles are an order of reptiles known as Testudines, characterized by a special shell developed mainly from their ribs. Modern turtles are divided into two major groups, the Pleurodira (side necked turtles) and Cryptodira (hidden necked t ...
s.


See also

* Timeline of ceratopsian research


References


External links


HVNHM on ''Bagaceratops''
{{Taxonbar, from=Q131256 Late Cretaceous dinosaurs of Asia Coronosaurs Fossil taxa described in 1975 Taxa named by Halszka Osmólska Taxa named by Teresa Maryańska Ornithischian genera