Ceratosuchus
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''Ceratosuchus'' 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 extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In the hierarchy of biological classification, genus com ...
of
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 ...
crocodylia 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 living ...
n from latest
Paleocene The Paleocene, ( ) or Palaeocene, is a geological epoch (geology), epoch that lasted from about 66 to 56 mya (unit), million years ago (mya). It is the first epoch of the Paleogene Period (geology), Period in the modern Cenozoic Era (geology), E ...
rocks of
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 ...
's
Piceance Basin The Piceance Basin is a geologic structural basin in northwestern Colorado, in the United States. It includes geologic formations from Cambrian to Holocene in age, but the thickest section is made up of rocks from the Cretaceous Period. The basi ...
and earliest
Eocene The Eocene ( ) Epoch is a geological epoch (geology), epoch that lasted from about 56 to 33.9 million years ago (mya). It is the second epoch of the Paleogene Period (geology), Period in the modern Cenozoic Era (geology), Era. The name ''Eocene' ...
rocks of
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 ...
's
Bighorn Basin The Bighorn Basin is a plateau region and intermontane basin, approximately 100 miles (160 km) wide, in north-central Wyoming in the United States. It is bounded by the Absaroka Range on the west, the Pryor Mountains on the north, the Bighor ...
in
North America North America is a continent in the Northern Hemisphere and almost entirely within the Western Hemisphere. It is bordered to the north by the Arctic Ocean, to the east by the Atlantic Ocean, to the southeast by South America and the Car ...
, a slice of time known as the
Clarkforkian The Clarkforkian North American Stage, on the geologic timescale, is the North American faunal stage according to the North American Land Mammal Ages chronology (NALMA), typically set from 56,800,000 to 55,400,000 years BP lasting . Considered t ...
North American Land Mammal Age The North American land mammal ages (NALMA) establishes a geologic timescale for North American fauna beginning during the Late Cretaceous and continuing through to the present. These periods are referred to as ages or intervals (or stages when ref ...
. Like its modern relatives, ''Ceratosuchus'' was a swamp-dwelling predator. It is named for the pair of flattened, triangular bony plates that extend from the back of its head. 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 ''C. burdoshi'', a name chosen by the Field Museum after Theodore Burdosh discovered a nearly complete skull on an expedition to Western Colorado in 1937.
"Fortunately, a knob of bone projecting from an otherwise undistinguished piece of rock had caught the eye of Mr. Burdosh, and the block had been broughtto the Museum. When the rock was chipped away, the insignificant external lump proved to belong to a fairly complete skull of a fossil crocodilian allied to the alligators; and on one posterior corner it bore a triangualr horn-like knob which proved to be identical with the mysterious separate fragments."


Description

''Ceratosuchus'' was named in 1938 by K. P. Schmidt for a skull from Colorado. Further remains, including additional skulls,
mandible In anatomy, the mandible, lower jaw or jawbone is the largest, strongest and lowest bone in the human facial skeleton. It forms the lower jaw and holds the lower tooth, teeth in place. The mandible sits beneath the maxilla. It is the only movabl ...
s, and
cervical In anatomy, cervical is an adjective that has two meanings: # of or pertaining to any neck. # of or pertaining to the female cervix: i.e., the ''neck'' of the uterus. *Commonly used medical phrases involving the neck are **cervical collar **cerv ...
armor Armour (British English) or armor (American English; see spelling differences) is a covering used to protect an object, individual, or vehicle from physical injury or damage, especially direct contact weapons or projectiles during combat, or fr ...
, was recovered from Wyoming by
University of Michigan , mottoeng = "Arts, Knowledge, Truth" , former_names = Catholepistemiad, or University of Michigania (1817–1821) , budget = $10.3 billion (2021) , endowment = $17 billion (2021)As o ...
paleontologists and described by William Bartels in 1984. The skull, of a moderately-size alligatorine, is most notable for its horns, formed by expansion of the bones (
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 The quadratojugal is a skull bone present in many vertebrates, including ...
s) that formed the rear corners of the skull roof. These horns were bulbous and pointed up. There were five teeth in both of the bones that made up the tip of the snout (
premaxilla The premaxilla (or praemaxilla) is one of a pair of small cranial bones at the very tip of the upper jaw of many animals, usually, but not always, bearing teeth. In humans, they are fused with the maxilla. The "premaxilla" of therian mammal has b ...
), fifteen in the paired
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 ...
e that formed the sides of the upper jaw, and twenty in both
dentaries In anatomy, the mandible, lower jaw or jawbone is the largest, strongest and lowest bone in the human facial skeleton. It forms the lower jaw and holds the lower teeth in place. The mandible sits beneath the maxilla. It is the only movable bone ...
of the lower jaw. The front of the lower jaw had a flattened shape, and the teeth located here pointed partially forward, with a
spade A spade is a tool primarily for digging consisting of a long handle and blade, typically with the blade narrower and flatter than the common shovel. Early spades were made of riven wood or of animal bones (often shoulder blades). After the a ...
-like form. The teeth had variable shapes; the first thirteen teeth in the lower jaw were pointed, while the last seven graded from a
spatula A spatula is a broad, flat, flexible blade used to mix, spread and lift material including foods, drugs, plaster and paints. In medical applications, "spatula" may also be used synonymously with tongue depressor. The word ''spatula'' derives f ...
te shape to a large globular shape. Aside from the horns, the skull and particularly the lower jaw of ''Ceratosuchus'' were very similar to that of its contemporary ''
Allognathosuchus ''Allognathosuchus'' (meaning "other jaw crocodile") is an extinct genus of alligatorine crocodylian with a complicated taxonomic history. It was named in 1921. Description ''Allognathosuchus'' was a medium-sized predator up to 1.5 m in leng ...
''. The neck armor had blade-like keels that may have been aligned with the skull horns.


Paleobiology

Although ''Ceratosuchus'' is the only known horned alligatorine, horns are not unknown in
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 living ...
ns; similar structures are known on two other species: ''
Voay robustus ''Voay'' is an extinct genus of crocodile from Madagascar that lived during the Late Pleistocene to Holocene, containing only one species, ''V. robustus''. Numerous subfossils have been found, including complete skulls, noted for their distin ...
'' and ''
Crocodylus rhombifer The Cuban crocodile (''Crocodylus rhombifer'') is a small-medium species of crocodile endemic to Cuba. Typical length is and typical weight . Large males can reach as much as in length and weigh more than . Despite its smaller size, it is a hig ...
''. Bartels proposed that the horns' small size and bluntness made them unlikely weapons, and their small size also made use in a threat display unlikely. Instead, he favored their use as signals for species recognition: in this case, the horns would allow ''Ceratosuchus'' and species of ''Allognathosuchus'' to tell each other apart. ''Ceratosuchus'' is known from overbank
mudstone Mudstone, a type of mudrock, is a fine-grained sedimentary rock whose original constituents were clays or muds. Mudstone is distinguished from '' shale'' by its lack of fissility (parallel layering).Blatt, H., and R.J. Tracy, 1996, ''Petrology. ...
deposits. It so far has been a rare find compared to ''Allognathosuchus'' from the same rocks. Bartels reported that only 5% of Clarkforkian crocodilians collected by the University of Michigan can be certainly assigned to ''Ceratosuchus'' (although this may be partially artificial because fragmentary specimens of ''Ceratosuchus'' could be confused with ''Allognathosuchus''). ''Ceratosuchus'' may have been
ecologically Ecology () is the study of the relationships between living organisms, including humans, and their biophysical environment, physical environment. Ecology considers organisms at the individual, population, community (ecology), community, ecosy ...
restricted in comparison to ''Allognathosuchus''. Although the blunt posterior teeth of ''Allognathosuchus'' and ''Ceratosuchus'' have been traditionally interpreted as for feeding on
mollusks Mollusca is the second-largest phylum of invertebrate animals after the Arthropoda, the members of which are known as molluscs or mollusks (). Around 85,000  extant species of molluscs are recognized. The number of fossil species is esti ...
or
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 tu ...
s, Bartels noted that these crocodilians were too small to feed on large bivalves or non-juvenile turtles, and that modern crocodilians usually swallow
snail A snail is, in loose terms, a shelled gastropod. The name is most often applied to land snails, terrestrial pulmonate gastropod molluscs. However, the common name ''snail'' is also used for most of the members of the molluscan class Gastro ...
s whole. Instead, he proposed that skull
morphology Morphology, from the Greek and meaning "study of shape", may refer to: Disciplines * Morphology (archaeology), study of the shapes or forms of artifacts * Morphology (astronomy), study of the shape of astronomical objects such as nebulae, galaxies ...
and tooth wear better fit generalized
predation Predation is a biological interaction where one organism, the predator, kills and eats another organism, its prey. It is one of a family of common feeding behaviours that includes parasitism and micropredation (which usually do not kill the ...
on a variety of invertebrates and vertebrates.


Classification

The precise placement of ''Ceratosuchus'' 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 ...
is disputed. Some studies have shown it as a basal member of
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 ...
, within the
stem group In phylogenetics, the crown group or crown assemblage is a collection of species composed of the living representatives of the collection, the most recent common ancestor of the collection, and all descendants of the most recent common ancestor. ...
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 ' ...
, 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 studies have recovered ''Ceratosuchus'' outside of
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
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 ' ...
, 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 more c ...
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.


References

{{Taxonbar, from=Q3664998 Alligatoridae Paleocene crocodylomorphs Eocene crocodylomorphs Paleogene reptiles of North America Fossil taxa described in 1938 Prehistoric pseudosuchian genera