Brachychampsa
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''Brachychampsa'' is an extinct
genus Genus (; : genera ) is a taxonomic rank above species and below family (taxonomy), family as used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In bino ...
of
alligatorid 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 ...
, possibly a basal
caiman A caiman ( (also spelled cayman) from Taíno language, Taíno ''kaiman'') is an alligatorid belonging to the subfamily Caimaninae, one of two primary lineages within the Alligatoridae family (biology), family, the other being alligators. ...
. Specimens have been reported from
New Mexico New Mexico is a state in the Southwestern United States, Southwestern region of the United States. It is one of the Mountain States of the southern Rocky Mountains, sharing the Four Corners region with Utah, Colorado, and Arizona. It also ...
,
Colorado Colorado is a U.S. state, state in the Western United States. It is one of the Mountain states, sharing the Four Corners region with Arizona, New Mexico, and Utah. It is also bordered by Wyoming to the north, Nebraska to the northeast, Kansas ...
,
Wyoming Wyoming ( ) is a landlocked U.S. state, state in the Mountain states, Mountain West subregion of the Western United States, Western United States. It borders Montana to the north and northwest, South Dakota and Nebraska to the east, Idaho t ...
,
Montana Montana ( ) is a landlocked U.S. state, state in the Mountain states, Mountain West subregion of the Western United States. It is bordered by Idaho to the west, North Dakota to the east, South Dakota to the southeast, Wyoming to the south, an ...
,
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, 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 Dakota people, Dakota Sioux ...
,
New Jersey New Jersey is a U.S. state, state located in both the Mid-Atlantic States, Mid-Atlantic and Northeastern United States, Northeastern regions of the United States. Located at the geographic hub of the urban area, heavily urbanized Northeas ...
, and
Saskatchewan Saskatchewan is a Provinces and territories of Canada, province in Western Canada. It is bordered on the west by Alberta, on the north by the Northwest Territories, on the east by Manitoba, to the northeast by Nunavut, and to the south by the ...
, though only those from Montana, Utah, and New Mexico are based on material sufficient to justify the referral. Some specimens have been reported from the
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 ...
-aged deposits of
Central Asia Central Asia is a region of Asia consisting of Kazakhstan, Kyrgyzstan, Tajikistan, Turkmenistan, and Uzbekistan. The countries as a group are also colloquially referred to as the "-stans" as all have names ending with the Persian language, Pers ...
(
Chimkent Shymkent (, ; ) is a city in southern Kazakhstan, located near the border with Uzbekistan. It holds the status of a city of republican significance, one of only three cities in Kazakhstan with this distinction, alongside Almaty and Astana. As of ...
of
Kazakhstan Kazakhstan, officially the Republic of Kazakhstan, is a landlocked country primarily in Central Asia, with a European Kazakhstan, small portion in Eastern Europe. It borders Russia to the Kazakhstan–Russia border, north and west, China to th ...
and Kirkuduk of
Tajikistan Tajikistan, officially the Republic of Tajikistan, is a landlocked country in Central Asia. Dushanbe is the capital city, capital and most populous city. Tajikistan borders Afghanistan to the Afghanistan–Tajikistan border, south, Uzbekistan to ...
), although the species status is indeterminate for these
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 preserve ...
s. 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. Campa ...
stage of the
Late Cretaceous The Late Cretaceous (100.5–66 Ma) is the more recent 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 ''cre ...
(
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 International Commission on Stratigraphy (ICS) geologic timescale, the latest age (geology), age (uppermost stage (stratigraphy), stage) of the Late Cretaceous epoch (geology), Epoch or Upper Cretaceous series (s ...
stage of the
Cretaceous The Cretaceous ( ) is a geological period that lasted from about 143.1 to 66 mya (unit), million years ago (Mya). It is the third and final period of the Mesozoic Era (geology), Era, as well as the longest. At around 77.1 million years, it is the ...
(
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.


Taxonomy

The
type species In International_Code_of_Zoological_Nomenclature, 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 spe ...
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 (stratigraphy), formation s ...
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 (Latin: ''holotypus'') is a single physical example (or illustration) of an organism used when the species (or lower-ranked taxon) was formally described. It is either the single such physical example (or illustration) or one of s ...
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 ...
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 an upper Santonian to lower Campanian geologic Formation (geology), formation found in Colorado and New Mexico, United States. Description The Menefee Formation consists of fluvial sandstone, shale, and coal. Based on am ...
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 precisely 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 a ...
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 In vertebrates, the maxilla (: maxillae ) 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 two maxil ...
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 m ...
has undergone many revisions since it was first named. Originally it was placed within
Alligatoridae The family (biology), 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 Am ...
, 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
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 m ...
, within the
clade In biology, a clade (), also known as a Monophyly, monophyletic group or natural group, is a group of organisms that is composed of a common ancestor and all of its descendants. Clades are the fundamental unit of cladistics, a modern approach t ...
Globidonta Globidonta is a clade of alligatoroids that includes alligators, caimans, and closely related extinct forms. It is defined as a Stem-based taxon, stem-based clade including ''Alligator mississippiensis'' (the American Alligator) and all forms mor ...
, as shown in the
cladogram A cladogram (from Greek language, Greek ''clados'' "branch" and ''gramma'' "character") is a diagram used in cladistics to show relations among organisms. A cladogram is not, however, an Phylogenetic tree, evolutionary tree because it does not s ...
below. Alternatively, other
phylogenetic 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 dat ...
studies have recovered ''Brachychampsa'' as an alligatorid, specifically as a
stem Stem or STEM most commonly refers to: * Plant stem, a structural axis of a vascular plant * Stem group * Science, technology, engineering, and mathematics Stem or STEM can also refer to: Language and writing * Word stem, part of a word respon ...
-
caiman A caiman ( (also spelled cayman) from Taíno language, Taíno ''kaiman'') is an alligatorid belonging to the subfamily Caimaninae, one of two primary lineages within the Alligatoridae family (biology), family, the other being alligators. ...
, as shown in the cladogram below. However, Walter et al. (2022) recovered ''Brachychampsa'', ''
Stangerochampsa ''Stangerochampsa'' is an extinct genus of alligatorid, possibly an alligatorine or a stem-caiman, from the Late Cretaceous of Alberta. It is based on RTMP.86.61.1, a skull, partial lower jaws, and partial postcranial skeleton discovered in the ...
'' and '' Albertochampsa'' as the basalmost
alligatorine 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 ''A ...
s based on phylogenetic analysis and claimed that the earliest definitive stem-group caimans are known from the earliest Paleocene.


Palaeoecology

''Brachychampsa'''s
dentition Dentition pertains to the development of teeth and their arrangement in the mouth. In particular, it is the characteristic arrangement, kind, and number of teeth in a given species at a given age. That is, the number, type, and morpho-physiology ...
was
heterodont In anatomy, a heterodont (from Greek, meaning 'different teeth') is an animal which possesses more than a single tooth morphology. Human dentition is heterodont and diphyodont as an example. In vertebrates, heterodont pertains to animals wher ...
, with conical teeth at the front and bulbous teeth at the back. It has been regarded as a specialist hunter of turtles, based on its rostral shape, massive
dentary In jawed vertebrates, the mandible (from the Latin ''mandibula'', 'for chewing'), lower jaw, or jawbone is a bone that makes up the lowerand typically more mobilecomponent of the mouth (the upper jaw being known as the maxilla). The jawbone ...
, and dental morphology, though this has been challenged over the years. A more generalist diet of small invertebrates and vertebrates has also been proposed. In 2003, Robert M. Sullivan and Spencer G. Lucas observed that most late Cretaceous turtles were probably too big for consumption, and that if they did represent a large portion of ''Brachychampsa'''s diet, it would have had to hunt either juveniles or smaller turtle species.


References


External links


''Brachychampsa''
in the
Paleobiology Database The Paleobiology Database (PBDB) is an online resource for information on the distribution and classification of fossil animals, plants, and microorganisms. History The Paleobiology Database originated in the NCEAS-funded Phanerozoic Marine Pale ...
{{Taxonbar, from=Q4953372 Alligatoroidea 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