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Chasmosaurinae is a
subfamily In biological classification, a subfamily (Latin: ', plural ') is an auxiliary (intermediate) taxonomic rank, next below family but more inclusive than genus. Standard nomenclature rules end botanical subfamily names with "-oideae", and zo ...
of
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 ...
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 ...
s. They were one of the most successful groups of
herbivores A herbivore is an animal anatomically and physiologically evolved to feed on plants, especially upon vascular tissues such as foliage, fruits or seeds, as the main component of its diet. These more broadly also encompass animals that eat ...
of their time. Chasmosaurines appeared in the early
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 ...
, and became extinct, along with all other non- avian dinosaurs, during the
Cretaceous–Paleogene extinction event The Cretaceous–Paleogene (K–Pg) extinction event, also known as the K–T extinction, was the extinction event, mass extinction of three-quarters of the plant and animal species on Earth approximately 66 million years ago. The event cau ...
. Broadly, the most distinguishing features of chasmosaurines are prominent brow horns and long frills lacking long spines;
centrosaurines Centrosaurinae (from the Greek language, 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 ( ...
generally had short brow horns and relatively shorter frills, and often had long spines projecting from their frills. Chasmosaurines evolved in western North America (Laramidia). They are currently known definitively from rocks in western Canada, the western United States, and northern Mexico. They were highly diverse and among the most species-rich groups of dinosaurs, with new species frequently described. This high diversity of named species is likely a result of the frill. The distinctive shape of the frill with the hornlets on its edges (epoccipitals) make it possible to recognize species from incomplete or fragmentary remains.


Classification

Chasmosaurinae is defined officially in the ''
PhyloCode The ''International Code of Phylogenetic Nomenclature'', known as the ''PhyloCode'' for short, is a formal set of rules governing phylogenetic nomenclature. Its current version is specifically designed to regulate the naming of clades, leaving the ...
'' by Daniel Madzia and colleagues in 2021 as "the largest clade containing '' Chasmosaurus belli'' and ''
Triceratops horridus ''Triceratops'' ( ; ) is a genus of Chasmosaurinae, chasmosaurine Ceratopsia, ceratopsian dinosaur that lived during the late Maastrichtian age of the Late Cretaceous Period (geology), period, about 68 to 66 million years ago on the island ...
'', but not '' Centrosaurus apertus''". Below is the result of a
phylogenetic analysis In biology, phylogenetics () is the study of the evolutionary history of life using observable characteristics of organisms (or genes), which is known as phylogenetic inference. It infers the relationship among organisms based on empirical data ...
by Mallon et al., following the traditional epiparietal homology scheme from their description of '' Spiclypeus shipporum''. ''
Bravoceratops ''Bravoceratops'' is a genus of large chasmosaurine ceratopsid dinosaur that lived approximately 70 million years ago, and is known from the Late Cretaceous Javelina Formation in what is now Texas, United States. Discovery and naming ''Bravocer ...
'' and ''
Eotriceratops ''Eotriceratops'' (meaning "dawn three-horned face") is a genus of herbivorous ceratopsian dinosaurs which lived in the area of North America during the late Cretaceous period. The only named species is ''Eotriceratops xerinsularis''. Discovery ...
'' were removed because it was found that they decrease resolution in the analysis because of the authors' new interpretation of epiparietal configurations. '' Regaliceratops'' was not resolved as a member of the Triceratopsini. The findings of a phylogenetic study done by Fowler and Freedman in 2020 are given below. The authors proposed that two distinct and roughly contemporaneous lineages of chasmosaurines existed in the late Cretaceous: a northern "''Chasmosaurus''" lineage with a heart-shaped frill margin that flattens and curls over onto itself, and a southern "''Pentaceratops''" lineage with a pinched shut indentation in the heart-shaped frill margin. According to the cladogram below, the Triceratopsini may have derived from this later lineage. To improve resolution, certain species based on partial or immature remains (''Bravoceratops'' and ''Agujaceratops'') were excluded, as in the Mallon et al. study above. While this new study did not yield a single "''Pentaceratops'' lineage," was not recovered by this revised analysis as they had predicted based on frill shape, the authors speculated that this may be due to some specimens included as ''Pentaceratops sternbergii'' being misclassified, and possibly referable to other species pending further study. The authors also noted that some newer species included in the previous analysis by Mallon et al. (''Spiclypeus'', ''Regaliceratops'', etc.) had yet to be coded into their revised dataset. Triceratopsini was named by Nicholas R. Longrich in 2011 for the description of '' Titanoceratops'', which he defined as "all species closer to ''
Triceratops horridus ''Triceratops'' ( ; ) is a genus of Chasmosaurinae, chasmosaurine Ceratopsia, ceratopsian dinosaur that lived during the late Maastrichtian age of the Late Cretaceous Period (geology), period, about 68 to 66 million years ago on the island ...
'' than to '' Anchiceratops ornatus'' or '' Arrhinoceratops brachyops''". Triceratopsins were the largest of the chasmosaurines; suggesting that gigantism had evolved in the Ceratopsidae once. In addition there is an evolutionary trend in the solidification of the frills, the most extreme being in ''Triceratops''.


See also

* Timeline of ceratopsian research


References


External links

*1.https://www.paleodb.org {{Taxonbar, from=Q137441 Dinosaur subfamilies Late Cretaceous dinosaurs Dinosaurs of North America