Ceratosuchus
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''Ceratosuchus'' ("horned crocodile") is an
extinct Extinction is the termination of an organism by the death of its Endling, last member. A taxon may become Functional extinction, functionally extinct before the death of its last member if it loses the capacity to Reproduction, reproduce and ...
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 alligatorine
crocodylia Crocodilia () is an Order (biology), order of semiaquatic, predatory reptiles that are known as crocodilians. They first appeared during the Late Cretaceous and are the closest living relatives of birds. Crocodilians are a type of crocodylomorp ...
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), ...
rocks of
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 ...
'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 ( ) is a geological epoch (geology), epoch that lasted from about 56 to 33.9 million years ago (Ma). It is the second epoch of the Paleogene Period (geology), Period in the modern Cenozoic Era (geology), Era. The name ''Eocene'' comes ...
rocks of
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 ...
'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 Bigho ...
in
North America North America is a continent in the Northern Hemisphere, Northern and Western Hemisphere, Western hemispheres. North America is bordered to the north by the Arctic Ocean, to the east by the Atlantic Ocean, to the southeast by South Ameri ...
, a slice of time known as the Clarkforkian
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 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 ...
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 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 i ...
s, and cervical
armor Armour (Commonwealth English) or armor (American English; see American and British English spelling differences#-our, -or, spelling differences) is a covering used to protect an object, individual, or vehicle from physical injury or damage, e ...
, was recovered from Wyoming by
University of Michigan The University of Michigan (U-M, U of M, or Michigan) is a public university, public research university in Ann Arbor, Michigan, United States. Founded in 1817, it is the oldest institution of higher education in the state. The University of Mi ...
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 bones form the cheek series of the skull. The bone forms an ancestra ...
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 mammals h ...
), fifteen in the paired
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 ...
e that formed the sides of the upper jaw, and twenty in both dentaries 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 ...
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''. 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 () is an order of semiaquatic, predatory reptiles that are known as crocodilians. They first appeared during the Late Cretaceous and are the closest living relatives of birds. Crocodilians are a type of crocodylomorph pseudosuchia ...
ns; similar structures are known on two other species: '' Voay robustus'' and '' Crocodylus rhombifer''. 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.Blatt, H., and R.J. Tracy, 1996, ''Petrology.'' New York, New York, ...
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 natural science of the relationships among living organisms and their environment. Ecology considers organisms at the individual, population, community, ecosystem, and biosphere levels. Ecology overlaps with the closely re ...
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 a phylum of protostomic invertebrate animals, whose members are known as molluscs or mollusks (). Around 76,000 extant species of molluscs are recognized, making it the second-largest animal phylum after Arthropoda. The num ...
or
turtle Turtles are reptiles of the order (biology), order Testudines, characterized by a special turtle shell, shell developed mainly from their ribs. Modern turtles are divided into two major groups, the Pleurodira (side necked turtles) and Crypt ...
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 a shelled gastropod. The name is most often applied to land snails, terrestrial molluscs, terrestrial pulmonate gastropod molluscs. However, the common name ''snail'' is also used for most of the members of the molluscan class Gas ...
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 in which one organism, the predator, kills and eats another organism, its prey. It is one of a family of common List of feeding behaviours, feeding behaviours that includes parasitism and micropredation ...
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 m ...
is disputed. Some studies have shown it as a basal member of
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 ...
, 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, 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 studies have recovered ''Ceratosuchus'' outside of
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 Alligatorinae, 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.


References

{{Taxonbar, from=Q3664998 Alligatorinae Paleocene crocodylomorphs Eocene crocodylomorphs Paleogene reptiles of North America Fossil taxa described in 1938 Prehistoric pseudosuchian genera Paleontology in Colorado Fossils of Wyoming