''Brachychampsa'' is an extinct

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 ...

alligatoroid Alligatoroidea is one of three superfamilies of crocodylians, the other two being Crocodyloidea and Gavialoidea. Alligatoroidea evolved in the Late Cretaceous period, and consists of the alligators and caimans, as well as extinct members more c ...

, possibly a basal

caiman A caiman (also cayman as a variant spelling) is an alligatorid belonging to the subfamily Caimaninae, one of two primary lineages within the Alligatoridae family, the other being alligators. Caimans inhabit Mexico, Central and South America fro ...

. Specimens have been reported from

New Mexico ) , population_demonym = New Mexican ( es, Neomexicano, Neomejicano, Nuevo Mexicano) , seat = Santa Fe , LargestCity = Albuquerque , LargestMetro = Tiguex , OfficialLang = None , Languages = English, Spanish ( New Mexican), Navajo, Ker ...

Colorado Colorado (, other variants) is a state in the Mountain West subregion of the Western United States. It encompasses most of the Southern Rocky Mountains, as well as the northeastern portion of the Colorado Plateau and the western edge of t ...

Wyoming Wyoming () is a U.S. state, state in the Mountain states, Mountain West subregion of the Western United States. It is bordered by Montana to the north and northwest, South Dakota and Nebraska to the east, Idaho to the west, Utah to the south ...

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 ...

North North is one of the four compass points or cardinal directions. It is the opposite of south and is perpendicular to east and west. ''North'' is a noun, adjective, or adverb indicating Direction (geometry), direction or geography. Etymology T ...

and

South Dakota South Dakota (; Sioux language, Sioux: , ) is a U.S. state in the West North Central states, North Central region of the United States. It is also part of the Great Plains. South Dakota is named after the Lakota people, Lakota and Dakota peo ...

New Jersey New Jersey is a state in the Mid-Atlantic and Northeastern regions of the United States. It is bordered on the north and east by the state of New York; on the east, southeast, and south by the Atlantic Ocean; on the west by the Delaware ...

, and

Saskatchewan Saskatchewan ( ; ) is a Provinces and territories of Canada, province in Western Canada, western Canada, bordered on the west by Alberta, on the north by the Northwest Territories, on the east by Manitoba, to the northeast by Nunavut, and on t ...

, though only those from Montana, Utah, and New Mexico are based on material sufficient to justify the referral. One specimen has been reported from the Darbasa Formation of

Kazakhstan Kazakhstan, officially the Republic of Kazakhstan, is a transcontinental country located mainly in Central Asia and partly in Eastern Europe. It borders Russia to the north and west, China to the east, Kyrgyzstan to the southeast, Uzbeki ...

, although the species status is indeterminate for the

fossil A fossil (from Classical Latin , ) is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserved ...

. The genus first appeared 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 ...

stage of 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 ...

(

Judithian The Judithian was a North American faunal stage lasting from 83.5 to 70.6 million years ago. It overlaps with the Campanian global stage. Fauna Dinosaur faunas of the Judithian age may represent the peak of dinosaur evolution in North America. H ...

North American stage) and became extinct during the late

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 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 ...

(

Lancian The Lancian was a North American faunal stage of the Late Cretaceous. It was the final stage of the Cretaceous period in North America, lasting from approximately 70.6 to 66 million years ago. Geology Terrestrial sedimentary strata from the Judit ...

North American Land Mammal "Age"). ''Brachychampsa'' is distinguished by an enlarged fifth maxillary tooth in the upper jaw. Brachychampsa NT small

Species

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 ...

of ''Brachychampsa'' is ''B. montana'', first discovered from the

Hell Creek Formation The Hell Creek Formation is an intensively studied division of mostly Upper Cretaceous and some lower Paleocene rocks in North America, named for exposures studied along Hell Creek, near Jordan, Montana. The formation stretches over portions of ...

of Montana and described by

Charles W. Gilmore Charles Whitney Gilmore (March 11, 1874 – September 27, 1945) was an American paleontologist who gained renown in the early 20th century for his work on vertebrate fossils during his career at the United States National Museum (now the N ...

in a paper in 1911. In that same paper, Gilmore recombined '' Bottosaurus perrugosus'' as a new species of ''Brachychampsa'', called ''B. perrugosus''. 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 of ''B. perrugosus'' went missing as the paper was being written, but it was later rediscovered and soon afterward designated as 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 ...

due to a lack of diagnostic features that distinguish it from other alligatorids discovered since the paper was published. Another species from the Allison Member of the

Menefee Formation The Menefee Formation is a lower Campanian geologic formation found in Colorado and New Mexico, United States. Description The Menefee Formation consists of fluvial sandstone, shale, and coal. Based on ammonite biostratigraphy, the age of the Me ...

of the

San Juan Basin The San Juan Basin is a geologic structural basin located near the Four Corners region of the Southwestern United States. The basin covers 7,500 square miles and resides in northwestern New Mexico, southwestern Colorado, and parts of Utah a ...

, ''B. sealeyi'', was discovered in 1996, but was later argued to be

synonymous A synonym is a word, morpheme, or phrase that means exactly or nearly the same as another word, morpheme, or phrase in a given language. For example, in the English language, the words ''begin'', ''start'', ''commence'', and ''initiate'' are all ...

with ''B. montana'' by interpreting it as an immature specimen of the latter species. However, other studies have shown that some of the variation seen between the two species, such as the orientation of 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. The t ...

ry tooth row, are not ontogenic, thus making ''B. sealeyi'' a valid taxon.

Phylogeny

''Brachychampsas position within

Alligatoroidea Alligatoroidea is one of three superfamilies of crocodylians, the other two being Crocodyloidea and Gavialoidea. Alligatoroidea evolved in the Late Cretaceous period, and consists of the alligators and caimans, as well as extinct members more c ...

has undergone many revisions since it was first named. Originally it was placed within

Alligatoridae The family Alligatoridae of crocodylians includes alligators, caimans and their extinct relatives. Phylogeny The superfamily Alligatoroidea includes all crocodilians (fossil and extant) that are more closely related to the American alligator t ...

, and was later refined to the

Alligatorinae Alligatorinae is a subfamily within the family Alligatoridae that contains the alligators and their closest extinct relatives, and is the sister taxon to Caimaninae (the caimans). Many genera in Alligatorinae are described, but only the genus ' ...

in 1964, only to be placed outside both Alligatorinae and Alligatoridae (but still within Alligatoroidea) in 1994. Accordingly, studies have shown ''Brachychampsa'' as a basal member of

, within the

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 term, ...

Globidonta Globidonta is a clade of alligatoroids that includes alligators, caimans, and closely related extinct forms. It is defined as a stem-based clade including ''Alligator mississippiensis'' (the American Alligator) and all forms more closely relate ...

, as shown in 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 d ...

below. Alternatively, other

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 ...

studies have recovered ''Brachychampsa'' as an alligatorid, specifically as a

stem Stem or STEM may refer to: Plant structures * Plant stem, a plant's aboveground axis, made of vascular tissue, off which leaves and flowers hang * Stipe (botany), a stalk to support some other structure * Stipe (mycology), the stem of a mushro ...

, as shown in the cladogram below.

References

External links

''Brachychampsa''
in the

Paleobiology Database The Paleobiology Database is an online resource for information on the distribution and classification of fossil animals, plants, and microorganisms. History The Paleobiology Database (PBDB) originated in the NCEAS-funded Phanerozoic Marine Pale ...

{{Taxonbar, from=Q4953372 Crocodilians Alligatoridae Late Cretaceous crocodylomorphs of North America Paleogene crocodylomorphs Late Cretaceous reptiles of North America Paleocene reptiles of North America Hell Creek fauna Fossil taxa described in 1911 Taxa named by Charles W. Gilmore Prehistoric pseudosuchian genera