Nasutoceratops
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''Nasutoceratops'' 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 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
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 ...
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 ...
. It is a basal
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 A ...
which lived 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'', ...
Period (late
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. Campani ...
, about 76.0-75.5 Ma). Fossils have been found in southern
Utah Utah ( , ) is a state in the Mountain West subregion of the Western United States. Utah is a landlocked U.S. state bordered to its east by Colorado, to its northeast by Wyoming, to its north by Idaho, to its south by Arizona, and to its ...
, United States. ''Nasutoceratops'' was a large, ground-dwelling,
quadrupedal Quadrupedalism is a form of locomotion where four limbs are used to bear weight and move around. An animal or machine that usually maintains a four-legged posture and moves using all four limbs is said to be a quadruped (from Latin ''quattuor ...
herbivore A herbivore is an animal anatomically and physiologically adapted to eating plant material, for example foliage or marine algae, for the main component of its diet. As a result of their plant diet, herbivorous animals typically have mouthpar ...
with a short snout and unique rounded horns above its eyes that have been likened to those of modern cattle. Extending almost to the tip of its snout, these horns are the longest of all the members of the centrosaurine
subfamily In biological classification, a subfamily (Latin: ', plural ') is an auxiliary (intermediate) taxonomic rank, next below family but more inclusive than genus Genus ( plural genera ) is a taxonomic rank used in the biological classifica ...
. The presence of pneumatic elements in the nasal bones of ''Nasutoceratops'' are a unique trait and are unknown in any other ceratopsid.


Discovery and naming

''Nasutoceratops'' is known from 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 seve ...
UMNH VP 16800, a partially associated nearly complete
skull The skull is a bone protective cavity for the brain. The skull is composed of four types of bone i.e., cranial bones, facial bones, ear ossicles and hyoid bone. However two parts are more prominent: the cranium and the mandible. In humans, th ...
, a coronoid process, a syncervical, three partial anterior dorsal vertebrae, a shoulder girdle, an associated left
forelimb A forelimb or front limb is one of the paired articulated appendages (limbs) attached on the cranial ( anterior) end of a terrestrial tetrapod vertebrate's torso. With reference to quadrupeds, the term foreleg or front leg is often used inste ...
, parts of the right forelimb and skin impressions. Two specimens were referred: UMNH VP 19466, a disarticulated adult
skull The skull is a bone protective cavity for the brain. The skull is composed of four types of bone i.e., cranial bones, facial bones, ear ossicles and hyoid bone. However two parts are more prominent: the cranium and the mandible. In humans, th ...
consisting of an incomplete premaxilla, maxilla and nasal, and UMNH VP 19469, an isolated
squamosal 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 ...
of a subadult. The holotype was discovered and collected in 2006 during the Kaiparowits Basin Project, initiated by the
University of Utah The University of Utah (U of U, UofU, or simply The U) is a public research university in Salt Lake City, Utah. It is the flagship institution of the Utah System of Higher Education. The university was established in 1850 as the University of De ...
in 2000. It was recovered from channel sandstone from the middle unit of the upper
Kaiparowits Formation The Kaiparowits Formation is a sedimentary rock formation found in the Kaiparowits Plateau in Grand Staircase–Escalante National Monument, in the southern part of Utah in the western United States. It is over 2800 feet (850 m) thick, and is ...
within the Grand Staircase–Escalante National Monument, in sediment that dates to the late
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. Campani ...
stage of the
Cretaceous The Cretaceous ( ) is a geological period that lasted from about 145 to 66 million years ago (Mya). It is the third and final period of the Mesozoic Era, as well as the longest. At around 79 million years, it is the longest geological period of ...
period, approximately 75 million years ago. It was first named and described in a thesis by its discoverer Eric Karl Lund in 2010 as ''Nasutuceratops titusi'', remaining at first an invalid ''nomen ex dissertatione''. Scott D. Sampson, Lund, Mark A. Loewen, Andrew A. Farke and Katherine E. Clayton validly named it in
2013 File:2013 Events Collage V2.png, From left, clockwise: Edward Snowden becomes internationally famous for leaking classified NSA wiretapping information; Typhoon Haiyan kills over 6,000 in the Philippines and Southeast Asia; The Dhaka garment fa ...
, emending the generic name to ''Nasutoceratops''. 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 specim ...
is ''Nasutoceratops titusi''. The generic name comes from ''nasutus'' in
Latin Latin (, or , ) is a classical language belonging to the Italic languages, 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 ...
meaning "large-nosed", and ''ceratops'', "horned-face" in
Greek Greek may refer to: Greece 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 ...
. The specific name honors Alan L. Titus for recovering fossils of ''Nasutoceratops'' from the GSENM.


Description

The holotype skull is approximately in length. Its body length has been estimated at , its weight at .Paul, G.S., 2016, ''The Princeton Field Guide to Dinosaurs 2nd edition'', Princeton University Press p. 287 ''Nasutoceratops'' has several unique derived 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 ...
. The part of the snout around the nostril is strongly developed, representing about three quarters of the skull length in front of the eye sockets. The rear of each nasal bone is hollowed out by a large internal cavity. The contact surface between 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 ...
and the praemaxilla is exceptionally large. The maxilla also has a large internal flange contacting the praemaxilla via two horizontal facets. The brow horns at their bases are pointed forwards and outwards, then curve inwards, and ultimately twist their points upwards.full text
/ref> ''Nasutoceratops'' also shows a unique combination of traits that are in themselves not unique. The horn on the nose is low and narrow, with an elongated base. The
squamosal 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 ...
has a high ridge on its upper surface, running from the direction of the eye socket towards the squamosal rim. The skull frill is more or less circular with its widest point at the middle edge. The osteoderms on the frill edge, the epiparietals and episquamosals, do not have the form of spikes but are shaped like simple low crescents. The rear frill edge is not notched but instead has an epiparietal on the midline. The snout of ''Nasutoceratops'' was short and high; its nasal bones exhibit internal cavities that the authors consider to represent pneumatic excavations, invading the bone from the
nasal cavity The nasal cavity is a large, air-filled space above and behind the nose in the middle of the face. The nasal septum divides the cavity into two cavities, also known as fossae. Each cavity is the continuation of one of the two nostrils. The nasal ...
. This is noteworthy because pneumatic nasals are unknown in any other ceratopsid, which supports that this feature represents a unique derived trait of this genus. ''Nasutoceratops'' had as many as 29 tooth positions in the maxilla, each occupied by several stacked teeth. The skull roof between the eye sockets is vaulted and is markedly higher than the snout region. The curved horizontally projecting brow horn arrangement was likened to that of modern
cattle Cattle (''Bos taurus'') are large, domesticated, cloven-hooved, herbivores. They are a prominent modern member of the subfamily Bovinae and the most widespread species of the genus '' Bos''. Adult females are referred to as cows and adult ...
by paleontologist
David Hone David Jeremy Hone (born 30 June 1946) is an Australian former sportsman who played first-class cricket with Oxford University and Australian rules football for Melbourne in the Victorian Football League (VFL). Hone spent the 1969 VFL season wit ...
. The brow horns span approximately 40% of the total length of the skull, almost reaching the level of the snout tip, and with a bone core length of up to 457 millimetres are the longest known of any centrosaurine, both in absolute and relative terms. The epijugal (cheek horn) has a length of , also the largest known among centrosaurines. The skull frill is moderately long, and pierced by a large kidney-shaped parietal fenestra at each side. Apart from the midline epiparietal, there are seven epiparietals at each side, and about four to five episquamosals. In the forelimb, the
ulna The ulna (''pl''. ulnae or ulnas) is a long bone found in the forearm that stretches from the elbow to the smallest finger, and when in anatomical position, is found on the medial side of the forearm. That is, the ulna is on the same side of t ...
is exceptionally robust with a large olecranon process. Of the three patches with skin impressions found near the left shoulder, one shows a pattern of large, eight to eleven millimetres wide, hexagonal scales surrounded by smaller triangular scales.


Classification

''Nasutoceratops'' was assigned to the Centrosaurinae in 2013, in a relatively basal position. A
phylogenetic In biology, phylogenetics (; from Greek φυλή/ φῦλον [] "tribe, clan, race", and wikt:γενετικός, γενετικός [] "origin, source, birth") is the study of the evolutionary history and relationships among or within groups ...
analysis performed by Sampson ''et al.'' (2013) found ''Nasutoceratops'' to be the
sister taxon 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 '' Avaceratops''. According to this study, the existence of ''Nasutoceratops'' would support the hypothesis of faunal separation between the north and south of Laramidia. Its
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 ter ...
would differ from the northern centrosaurines in the retention of long brow horns and a short nose horn, combined with developing, convergent with 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, durin ...
, low epiparietals. In 2016, this clade was named Nasutoceratopsini; it contains ''Nasutoceratops'' as well as ANSP 15800 (the holotype of '' Avaceratops''), MOR 692 (previously treated as an adult specimen of ''Avaceratops''), the newly described CMN 8804 from the Oldman Formation, and another undescribed ceratopsian found in
Malta, Montana Malta ( ) is a city in, and the county seat of, Phillips County, Montana, United States, located at the intersection of U.S. Routes 2 and 191. The population was 1,860 at the 2020 census. History After James Hill and his partners built th ...
. The
cladogram A cladogram (from Greek ''clados'' "branch" and ''gramma'' "character") is a diagram used in cladistics to show relations among organisms. A cladogram is not, however, an evolutionary tree because it does not show how ancestors are related to ...
presented below follows a phylogenetic analysis of the '' Centrosaurinae'' by Chiba ''et al.'' (2017), which included a systematic re-evaluation of '' Medusaceratops lokii'':


Paleoecology

The only known specimen of ''Nasutoceratops'' was recovered at the Kaiparowits Formation, in southern Utah. Argon-argon radiometric dating indicates that the Kaiparowits Formation was deposited between 76.1 and 74.0 million years ago, during the Campanian stage of the Late
Cretaceous The Cretaceous ( ) is a geological period that lasted from about 145 to 66 million years ago (Mya). It is the third and final period of the Mesozoic Era, as well as the longest. At around 79 million years, it is the longest geological period of ...
period. During the Late Cretaceous period, the site of the Kaiparowits Formation was located near the western shore of the Western Interior Seaway, a large inland sea that split North America into two landmasses, Laramidia to the west and
Appalachia Appalachia () is a cultural region in the Eastern United States that stretches from the Southern Tier of New York State to northern Alabama and Georgia. While the Appalachian Mountains stretch from Belle Isle in Newfoundland and Labrador, C ...
to the east. The plateau where dinosaurs lived was an ancient floodplain dominated by large channels and abundant wetland
peat Peat (), also known as turf (), is an accumulation of partially Decomposition, decayed vegetation or organic matter. It is unique to natural areas called peatlands, bogs, mires, Moorland, moors, or muskegs. The peatland ecosystem covers and ...
swamps, ponds and lakes, and was bordered by highlands. The climate was wet and humid, and supported an abundant and diverse range of organisms. This formation contains one of the best and most continuous records of Late Cretaceous terrestrial life in the world. ''Nasutoceratops'' shared its
paleoenvironment Paleoecology (also spelled palaeoecology) is the study of interactions between organisms and/or interactions between organisms and their environments across geologic timescales. As a discipline, paleoecology interacts with, depends on and informs ...
with
theropod 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 ...
s such as
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, the troodontid ''
Talos sampsoni ''Talos'' is an extinct genus of carnivorous bird-like theropod dinosaur, an advanced troodontid which lived during the late Cretaceous period (late Campanian, about 76 Ma) in the geographic area that is now Utah, United States. Discovery '' ...
'', ornithomimids like ''
Ornithomimus velox ''Ornithomimus'' (; "bird mimic") is a genus of ornithomimid dinosaurs from the Late Cretaceous Period of what is now North America. ''Ornithomimus'' was a swift bipedal theropod which fossil evidence indicates was covered in feathers, equipped ...
'',
tyrannosaurids 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 ...
like ''
Teratophoneus ''Teratophoneus'' ("monstrous murderer"; Greek: ''teras'', "monster" and ''phoneus'', "murderer") is a genus of tyrannosaurid dinosaur which lived during the late Cretaceous period (late Campanian age, about 77 to 76 million years ago) in what i ...
'', armored ankylosaurids, the duckbilled hadrosaurs '' Parasaurolophus cyrtocristatus'' and ''
Gryposaurus monumentensis ''Gryposaurus'' (meaning "hooked-nosed (Greek ''grypos'') lizard"; sometimes incorrectly translated as "griffin (Latin ''gryphus'') lizard") was a genus of duckbilled dinosaur that lived about 80 to 75 million years ago, in the Late Cretaceo ...
'', the ceratopsians '' Utahceratops gettyi'' and '' Kosmoceratops richardsoni'' and the oviraptorosaurian '' Hagryphus giganteus''. Paleofauna present in the Kaiparowits Formation included chondrichthyans (sharks and rays),
frog A frog is any member of a diverse and largely Carnivore, carnivorous group of short-bodied, tailless amphibians composing the order (biology), order Anura (ανοὐρά, literally ''without tail'' in Ancient Greek). The oldest fossil "proto-f ...
s,
salamander Salamanders are a group of amphibians typically characterized by their lizard-like appearance, with slender bodies, blunt snouts, short limbs projecting at right angles to the body, and the presence of a tail in both larvae and adults. All t ...
s,
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,
lizard Lizards are a widespread group of squamate reptiles, with over 7,000 species, ranging across all continents except Antarctica, as well as most oceanic island chains. The group is paraphyletic since it excludes the snakes and Amphisbaenia altho ...
s and
crocodilia Crocodilia (or Crocodylia, both ) is an order of mostly large, predatory, semiaquatic reptiles, known as crocodilians. They first appeared 95 million years ago in the Late Cretaceous period ( Cenomanian stage) and are the closest livi ...
ns. A variety of early
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 were present including multituberculates,
marsupial Marsupials are any members of the mammalian infraclass Marsupialia. All extant marsupials are endemic to Australasia, Wallacea and the Americas. A distinctive characteristic common to most of these species is that the young are carried in ...
s, and insectivorans.


See also

* Timeline of ceratopsian research


References


External links


Image
{{Taxonbar, from=Q3914791 Centrosaurines Late Cretaceous dinosaurs of North America Fossil taxa described in 2013 Taxa named by Scott D. Sampson Paleontology in Utah Kaiparowits Formation Campanian genus first appearances Campanian genus extinctions Ornithischian genera