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''Bravoceratops'' is a
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 large
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 ...
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 ...
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 ...
that lived approximately 70
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). ...
, and is 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 ...
Javelina Formation The Javelina Formation is a geological formation in Texas. Dating has shown that the strata date to the Maastrichtian stage of the Late Cretaceous, approximately 70 to 66.5 million years old. The middle part of the formation has been dated to abo ...
in what is now
Texas Texas (, ; Spanish language, Spanish: ''Texas'', ''Tejas'') is a state in the South Central United States, South Central region of the United States. At 268,596 square miles (695,662 km2), and with more than 29.1 million residents in 2 ...
, United States.


Discovery and naming

''Bravoceratops'' is only 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 several ...
specimen TMM 46015-1; it is housed in the collection of the Texas Memorial Museum,
Austin, Texas Austin is the capital city of the U.S. state of Texas, as well as the county seat, seat and largest city of Travis County, Texas, Travis County, with portions extending into Hays County, Texas, Hays and Williamson County, Texas, Williamson co ...
. The skull consists of a number of fragments, altogether including: the , parts of each brow horn, the rear end of the left , assorted parts of the nasal area and horn, the , and quadratojugals from each side, a section of the and , multiple parts of the dentary, and some of the right . It was recovered from the lowermost rocks of the
Javelina Formation The Javelina Formation is a geological formation in Texas. Dating has shown that the strata date to the Maastrichtian stage of the Late Cretaceous, approximately 70 to 66.5 million years old. The middle part of the formation has been dated to abo ...
, in
Big Bend National Park Big Bend National Park is an American national park located in West Texas, bordering Mexico. The park has national significance as the largest protected area of Chihuahuan Desert topography and ecology in the United States, and was named after ...
; ceratopsid fossils are uncommon in this formation, giving the discovery of ''Bravoceratops'' importance in clarifying the groups' diversity. The specimen was found in sandy conglomerate sediment at the Hippiewalk geologic locality. Signs of erosion are presented in the larger bone fragments, which were found disorganized over an area of ten square metres; the site is thought to have been a lag deposit of a river, only most durable extremities able to survive preservation. The animal would have lived during 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. Campanian s ...
or early
Maastrichtian The Maastrichtian () is, in the ICS geologic timescale, the latest age (uppermost stage) of the Late Cretaceous Epoch or Upper Cretaceous Series, the Cretaceous Period or System, and of the Mesozoic Era or Erathem. It spanned the interval from ...
stage Stage or stages may refer to: Acting * Stage (theatre), a space for the performance of theatrical productions * Theatre, a branch of the performing arts, often referred to as "the stage" * ''The Stage'', a weekly British theatre newspaper * Sta ...
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 th ...
period, approximately 70 million years ago. The genus name means "wild horn-face", and is derived from the Mexican name for the Rio Grande, "
Rio Bravo del Norte The Rio Grande ( and ), known in Mexico as the Río Bravo del Norte or simply the Río Bravo, is one of the principal rivers (along with the Colorado River) in the southwestern United States and in northern Mexico. The length of the Rio ...
" (wild river of the north), and the
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 ...
words "keras" (κέρας) meaning "horn" and "ops" (ὤψ) referring to the "face". The specific name ''polyphemus'', refers to the giant cyclops
Polyphemus Polyphemus (; grc-gre, Πολύφημος, Polyphēmos, ; la, Polyphēmus ) is the one-eyed giant son of Poseidon and Thoosa in Greek mythology, one of the Cyclopes described in Homer's ''Odyssey''. His name means "abounding in songs and lege ...
confronted by Odysseus in the Greek epic poem,
The Odyssey The ''Odyssey'' (; grc, Ὀδύσσεια, Odýsseia, ) is one of two major ancient Greek epic poems attributed to Homer. It is one of the oldest extant works of literature still widely read by modern audiences. As with the ''Iliad'', the ...
. This is in reference to the presumption that it had a single central horn on the surface of the frill, recalling the placement of a Cyclops' eye. The genus was described and named by Steven L. Wick and Thomas M. Lehman 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 fact ...
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 ''Bravoceratops polyphemus''.


Description

Two
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 ...
(unique
derived Derive may refer to: * Derive (computer algebra system), a commercial system made by Texas Instruments * ''Dérive'' (magazine), an Austrian science magazine on urbanism *Dérive, a psychogeographical concept See also * *Derivation (disambiguatio ...
traits) were originally proposed. Firstly, the median bar (which runs down the middle of the frill between its large holes) at mid-length splays out to the rear like a fan and its rear edge is not notched or embayed. Secondly, the upper surface of the bar is, at the midline, hollowed out by a symmetrical depression. It is this hollow in the form of an inverted tear that occasioned the specific name as it resembled the single eye of a cyclops, thus the allusion with Polyphemus. The authors assumed it formed the base of an epiparietal, or small horn, unpreserved in the fossil. In 2020, paleontologists Denver W. Fowler and Elizabeth A. Freedman Fowler suggested that the parietal median bar of the specimen had been reconstructed upside down, and that the genus therefore did not have any distinguishing features among chasmosaurines, making it a ''
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 ...
'', or invalid genus. However, a number of authors considered the genus valid (see classification below) and it was included as part of a southwest clade of chasmosaurines by Dalman et al., (2021). Unlike Fowler and Freedman-Folwer, Sebastian G. Dalman and colleagues, in a 2021 paper naming the related genus ''
Sierraceratops ''Sierraceratops'' (meaning " Sierra horned face") is a genus of chasmosaurine ceratopsian from the Late Cretaceous Hall Lake Formation of New Mexico, United States. The genus contains a single species, ''Sierraceratops turneri'', known from a p ...
'', treated ''Bravoceratops'' as valid, noting both distinct traits and those unique to it and ''Sierraceratops''. A flange in front of its near the bones is present, large as in ''Sierraceratops'' while small or not present in other chasmosaurs. The epijugal, a small horn on the jugals, is longer than tall and sharp on its tip, as in ''Sierraceratops'' and unlike other derived chasmosaurines. The two brow horns are compressed, being taller than wide rather than conical, a trait shared with ''Sierraceratops'' and the distantly related ''Judiceratops''. The cross section of the lower bar of its parietal bar has a D-shape shared with ''Sierraceratops'' and similar but not identical to that of '' Coahuilaceratops'', but unlike other chasmosaurines. ''Bravoceratops'' is distinguishable from ''Coahuilaceratops'' in lacking enormous brow horns; it is distinguishable from ''Sierraceratops'' in having larger brow horns and a smaller epijugal horn.


Classification

''Bravoceratops'' is assigned to 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 ...
. It is an "intermediate" form of chasmosaurine; it, along with genera like ''
Pentaceratops ''Pentaceratops'' ("five-horned face") is a genus of herbivorous ceratopsid dinosaur from the late Cretaceous Period of what is now North America. Fossils of this animal were first discovered in 1921, but the genus was named in 1923 when its typ ...
'' and ''
Anchiceratops ''Anchiceratops'' ( ) is an extinct genus of chasmosaurine ceratopsid dinosaur that lived approximately 72 to 71 million years ago during the latter part of the Cretaceous Period in what is now Alberta, Canada. ''Anchiceratops'' was a medium-size ...
'', was distinctly more derived ("advanced") than forms like ''
Chasmosaurus ''Chasmosaurus'' ( ) is a genus of ceratopsid dinosaur from the Upper Cretaceous Period of North America. Its name means 'opening lizard', referring to the large openings ( fenestrae) in its frill (Greek ''chasma'' meaning 'opening' or 'hollow' ...
'' but less so than
taxa In biology, a taxon (back-formation from ''taxonomy''; plural taxa) is a group of one or more populations of an organism or organisms seen by taxonomists to form a unit. Although neither is required, a taxon is usually known by a particular nam ...
like ''
Triceratops ''Triceratops'' ( ; ) is a genus of herbivore, herbivorous Chasmosaurinae, chasmosaurine Ceratopsidae, ceratopsid dinosaur that first appeared during the late Maastrichtian stage of the Late Cretaceous Period (geology), period, about 68 m ...
''. Its position in 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 o ...
analysis performed by Wick and Lehman (2013), who first named the taxon. It was found that ''Bravoceratops'' may 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 t ...
of '' Coahuilaceratops'' (a relationship later supported by Dalman et al., 2021). The first analysis, with unmodified characters from a previous study, found the pair to be relatively basal . The second, with two added characters, instead found ''Bravoceratops'' to be a relatively derived taxon. Both positions were considered to be somewhat unintuitive; several of its derived characters would be odd for a taxon in the basal position, but the genus also possessed primitive characters would require evolutionary reversals if the more derived position was correct. Regardless, the more derived placement was considered to be somewhat more likely. The first tree is reproduced on the left, and the second on the right: Future studies found trees showing the derived position, with ''Bravoceratops'' found related to ''
Arrhinoceratops ''Arrhinoceratops'' (meaning "no nose-horn face", derived from the Ancient Greek "a-/α-" "no", rhis/ῥίς "nose" "keras/κέρας" "horn", "-ops/ὤψ" "face") is a genus of herbivorous ceratopsian dinosaur. The name was coined as its origina ...
'' and
Triceratopsini Triceratopsini is a tribe of herbivorous chasmosaurine dinosaurs that lived between the late Campanian to the late Maastrichtian stages of the Cretaceous period, between 74.73 and 66 million years ago. Fossils of these animals have been found in ...
. However, the fragmentary nature of ''Bravoceratops'' caused it to generally be removed from phylogenetic analyses, as it was a wildcard taxon (meaning it was unstable and caused poorer results). Fowler and Freedman Fowler (2020) recovered ''Bravoceratops'' in the more basal position, and, similar to Wick and Lehman (2013), found it close to ''Pentaceratops'', and once again sister to ''Coahuilaceratops''. However, they also decided to remove it from their analysis due to their unsubstantiated conclusion the taxon was invalid. However, Dalman et al. (2021) convincingly validated the genus. They demonstrated that ''Bravoceratops'' shared a number morphological traits with other chasmosaurines from the same time and region. As a result, ''Bravoceratops'' was determined to be part of a clade endemic to the Southwestern United states. Sister taxa include ''Coahuilaceratops'' from Mexico and ''Sierraceratops'' from New Mexico.


See also

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


References

{{Taxonbar, from=Q13410268 Chasmosaurines Dinosaurs of North America Fossil taxa described in 2013 Maastrichtian life Paleontology in Texas Ornithischian genera