Cerrejonisuchus
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''Cerrejonisuchus'' 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 ...
of
dyrosaurid Dyrosauridae is a family of extinct neosuchian crocodyliforms that lived from the Late Cretaceous (Maastrichtian) to the Eocene. Dyrosaurid fossils are globally distributed, having been found in Africa, Asia, Europe, North America and South Amer ...
crocodylomorph Crocodylomorpha is a group of pseudosuchian archosaurs that includes the crocodilians and their extinct relatives. They were the only members of Pseudosuchia to survive the end-Triassic extinction. During Mesozoic and early Cenozoic times, cro ...
. It is known from a complete skull and mandible from the
Cerrejón Formation The Cerrejón Formation is a geologic formation in Colombia dating back to the Middle-Late Paleocene. It is found in the El Cerrejón sub-basin of the Cesar-Ranchería Basin of La Guajira and Cesar. The formation consists of bituminous coal field ...
in northeastern
Colombia Colombia (, ; ), officially the Republic of Colombia, is a country in South America with insular regions in North America—near Nicaragua's Caribbean coast—as well as in the Pacific Ocean. The Colombian mainland is bordered by the Car ...
, which is
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 ...
in age. Specimens belonging to ''Cerrejonisuchus'' and to several other dyrosaurids have been found from the
Cerrejón Cerrejón is a large open-pit coal mine in Northern Colombia owned by Glencore. At Cerrejón, low-ash, low-sulphur bituminous coal from the Cerrejón Formation is excavated. At over the mine is one of the largest of its type, the largest in Lati ...
open-pit coal mine in
La Guajira La Guajira () is a department of Colombia. It occupies most of the Guajira Peninsula in the northeast region of the country, on the Caribbean Sea and bordering Venezuela, at the northernmost tip of South America. The capital city of the departm ...
. The length of the rostrum is only 54-59% of the total length of the skull, making the snout of ''Cerrejonisuchus'' the shortest of all dyrosaurids.


Description

At an estimated length of to , ''Cerrejonisuchus'' was small for a dyrosaur. This size estimate is based on the dorsal skull lengths of specimens UF/IGM 29 and UF/IGM 31. ''Cerrejonisuchus'' has the shortest body length of any known dyrosaur, much smaller than that of the longest dyrosaur, ''
Phosphatosaurus gavialoides ''Phosphatosaurus'' is an extinct genus of dyrosaurid crocodylomorph. It existed during the early Eocene, with fossils having been found from North Africa in Tunisia and Mali. Named in 1955, ''Phosphatosaurus'' is a monotypic genus; the type spe ...
'', which was to in length. Currently the only known specimens of ''Cerrejonisuchus'' are UF/IGM 29 (the
type specimen In biology, a type is a particular wiktionary:en:specimen, specimen (or in some cases a group of specimens) of an organism to which the scientific name of that organism is formally attached. In other words, a type is an example that serves to a ...
), UF/IGM 30, UF/IGM 31, and UF/IGM 32. Of these, UF/IGM 29 and UF/IGM 31 are thought to represent fully mature individuals while UF/IGM 32 is thought to represent a less mature individual. In UF/IGM 31, the neurocentral sutures of the anterior dorsal
vertebrae The spinal column, a defining synapomorphy shared by nearly all vertebrates,Hagfish are believed to have secondarily lost their spinal column is a moderately flexible series of vertebrae (singular vertebra), each constituting a characteristic i ...
are closed, an indication of morphological maturity. Additionally, the presence of well-developed
osteoderm Osteoderms are bony deposits forming scales, plates, or other structures based in the dermis. Osteoderms are found in many groups of extant and extinct reptiles and amphibians, including lizards, crocodilians, frogs, temnospondyls (extinct amp ...
s is likely to be an indication that the animal was mature because in living crocodylians, the osteoderms begin calcification after 1 year and grow to articulate with other osteoderms to form a dermal shield at maturity. Also, the sutures that separate the bones of the skull in both specimens are fully fused, suggesting that the individuals have reached a late
ontogenic Ontogeny (also ontogenesis) is the origination and development of an organism (both physical and psychological, e.g., moral development), usually from the time of fertilization of the egg to adult. The term can also be used to refer to the stu ...
stage. In contrast, UF/IGM 32 has an unfused nasal suture, suggesting that it was less mature than the other individuals. UF/IGM 32 is also noticeably smaller than the other specimens. Relative to the entire skull length, the rostrum of ''Cerrejonisuchus'' is the shortest of any dyrosaurid. It, along with '' Chenanisuchus'', are the only short-snouted dyrosaurids. The snout of ''Cerrejonisuchus'' is narrow and consistent in width from the external nares, or
nostril A nostril (or naris , plural ''nares'' ) is either of the two orifices of the nose. They enable the entry and exit of air and other gasses through the nasal cavities. In birds and mammals, they contain branched bones or cartilages called turbi ...
openings, to the
orbit In celestial mechanics, an orbit is the curved trajectory of an object such as the trajectory of a planet around a star, or of a natural satellite around a planet, or of an artificial satellite around an object or position in space such as a p ...
s, or eye sockets. The margin of the snout, unlike that of many long-snouted dyrosaurids, is smooth rather than festooned. "Festooned" refers to the lateral undulations in the maxillae and premaxillae that form around the tooth sockets, or alveoli. The external nares are positioned extremely anteriorly at the very tip of the snout. The orbits are oriented anterodorsally, facing upward and slightly forward. The dentition of ''Cerrejonisuchus'' is generally
homodont In anatomy, a heterodont (from Greek, meaning 'different teeth') is an animal which possesses more than a single tooth morphology. In vertebrates, heterodont pertains to animals where teeth are differentiated into different forms. For example ...
, although the third
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 is enlarged and the fourth is somewhat smaller than the rest. They are conical, labiolingually compressed, each having a relatively rounded apex. The carinae, or tooth edges, are strongly developed both anteriorly and posteriorly. The
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 ...
ry teeth are generally thinner and longer than the maxillary teeth. Like ''Chenanisuchus'', ''Cerrejonisuchus'' visibly lacks striations on the tooth surfaces. Unlike many other dyrosaurids, including '' Dyrosaurus maghribensis'', '' Atlantosuchus coupatezi'', '' Guarinisuchus munizi'', ''Phosphatosaurus gavialoides'', and '' Sokotosuchus ianwilsoni'', the teeth of ''Cerrejonisuchus'' are not curved.


Classification

A
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 ...
analysis of dyrosaurids by Hastings ''et al.'' (2010) placed ''Cerrejonisuchus'' relatively basally in the dyrosaur clade between ''Phosphatosaurus gavialoides'' and ''
Arambourgisuchus khouribgaensis ''Arambourgisuchus'' ("Prof. Camille Arambourg's crocodile") is an extinct genus of dyrosaurid crocodylomorph from the late Palaeocene of Morocco, found in the region of Sidi Chenane in 2000, following collaboration by French and Moroccan instit ...
''. ''Cerrejonisuchus'' was not found to be closely related to the other short-snouted dyrosaur ''Chenanisuchus'', which was placed at the base of the clade. Although it might be expected that ''Chenanisuchus'' and ''Cerrejonisuchus'' are closely related because they are the only dyrosaurids with short snouts, the results of the analysis show that snout proportions alone are not indicative of phylogenetic relatedness in dyrosaurs. Below is 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 ...
from Hastings ''et al.'' (2010) showing the phylogenetic relationship of ''Cerrejonisuchus'' within Dyrosauridae:


Paleobiology


Feeding strategy

''Cerrejonisuchus'' likely had a diet consisting of fish, invertebrates, frogs, lizards, small snakes, and possibly mammals. The short snout of ''Cerrejonisuchus'' is thought to be an adaptation to such a generalized diet. Other long-snouted marine dyrosaurs are presumed to have had a strongly piscivorous diet consisting solely of fish. With its short snout, ''Cerrejonisuchus'' would have been able to occupy a new
ecological niche In ecology, a niche is the match of a species to a specific environmental condition. Three variants of ecological niche are described by It describes how an organism or population responds to the distribution of resources and competitors (for ...
in the neotropical rainforest environment of Paleocene Colombia. The presence of homodont dentition with compressed teeth similar to those of
sebecia Sebecia is an extinct clade of mesoeucrocodylian crocodyliforms that includes peirosaurids and sebecids. It was first constructed in 2007 to include ''Hamadasuchus'', Peirosauridae, and ''Sebecus''. It was initially considered to be the sister ta ...
ns and other terrestrial crocdylomorphs has been used as an argument for a terrestrial predatory lifetyle, though unlike other terrestrial crocodylomorphs it has a flat skull.


Locomotion

Compared to other dyrosaurids, ''Cerrejonisuchus'' has stronger limbs and a cylindrical torso, suggesting a primarily terrestrial ecology as opposed to the semi-aquatic habits of modern crocodilians and the fully marine habits of other dyrosaurids.


Paleoenvironment

''Cerrejonisuchus'' is known from the Middle to Late Paleocene Cerrejón Formation. All known specimens have been found from the Cerrejón open-pit coal mine at the La Puente Pit below Coal Seam 90. The mine has also yielded remains of '' Titanoboa cerrejonensis'', a recently described long extinct
boid The Boidae, commonly known as boas or boids, are a family of nonvenomous snakes primarily found in the Americas, as well as Africa, Europe, Asia, and some Pacific Islands. Boas include some of the world's largest snakes, with the green anaconda ...
that is the largest known snake to have ever existed. Like ''Cerrejonisuchus'', fossils of ''Titanoboa'' were found in a gray
claystone Mudrocks are a class of fine-grained siliciclastic sedimentary rocks. The varying types of mudrocks include siltstone, claystone, mudstone, slate, and shale. Most of the particles of which the stone is composed are less than and are too sm ...
layer directly underlying Coal Seam 90. Additional dyrosaurid material has been found from the Cerrejón Formation alongside that of ''Cerrejonisuchus'', and is thought to represent at least two different taxa. The age of the Cerrejón Formation has been dated as Middle-Late Paleocene based on
carbon Carbon () is a chemical element with the symbol C and atomic number 6. It is nonmetallic and tetravalent In chemistry, the valence (US spelling) or valency (British spelling) of an element is the measure of its combining capacity with o ...
isotope Isotopes are two or more types of atoms that have the same atomic number (number of protons in their nuclei) and position in the periodic table (and hence belong to the same chemical element), and that differ in nucleon numbers (mass numbers) ...
s,
pollen Pollen is a powdery substance produced by seed plants. It consists of pollen grains (highly reduced microgametophytes), which produce male gametes (sperm cells). Pollen grains have a hard coat made of sporopollenin that protects the gametophyt ...
,
spore In biology, a spore is a unit of sexual or asexual reproduction that may be adapted for dispersal and for survival, often for extended periods of time, in unfavourable conditions. Spores form part of the life cycles of many plants, algae, f ...
s, and
dinoflagellate The dinoflagellates (Greek δῖνος ''dinos'' "whirling" and Latin ''flagellum'' "whip, scourge") are a monophyletic group of single-celled eukaryotes constituting the phylum Dinoflagellata and are usually considered algae. Dinoflagellates are ...
cysts. The section of the Cerrejón Formation from which fossils of ''Cerrejonisuchus'' have been found was likely deposited in a transitional environment, probably
brackish water Brackish water, sometimes termed brack water, is water occurring in a natural environment that has more salinity than freshwater, but not as much as seawater. It may result from mixing seawater (salt water) and fresh water together, as in estua ...
in a
river A river is a natural flowing watercourse, usually freshwater, flowing towards an ocean, sea, lake or another river. In some cases, a river flows into the ground and becomes dry at the end of its course without reaching another body of wate ...
-to-
lagoon A lagoon is a shallow body of water separated from a larger body of water by a narrow landform, such as reefs, barrier islands, barrier peninsulas, or isthmuses. Lagoons are commonly divided into ''coastal lagoons'' (or ''barrier lagoons'') a ...
al setting. Large freshwater podocnemidid
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 and dipnoan and
elopomorph The superorder Elopomorpha contains a variety of types of fishes that range from typical silvery-colored species, such as the tarpons and ladyfishes of the Elopiformes and the bonefishes of the Albuliformes, to the long and slender, smooth-bodi ...
fish Fish are aquatic, craniate, gill-bearing animals that lack limbs with digits. Included in this definition are the living hagfish, lampreys, and cartilaginous and bony fish as well as various extinct related groups. Approximately 95% of li ...
es have also been found from this part of the formation. ''Cerrejonisuchus'' may have been a food source for ''Titanoboa'', which would have lived in the same brackish water environment. The vertebrate paleofauna of the Cerrejón Formation was similar to modern neotropical riverine vertebrate faunas. During the Paleocene, river systems would have incised a
coastal plain A coastal plain is flat, low-lying land adjacent to a sea coast. A fall line commonly marks the border between a coastal plain and a piedmont area. Some of the largest coastal plains are in Alaska and the southeastern United States. The Gulf Coa ...
covered by a wet
neotropical The Neotropical realm is one of the eight biogeographic realms constituting Earth's land surface. Physically, it includes the tropical terrestrial ecoregions of the Americas and the entire South American temperate zone. Definition In bioge ...
rainforest Rainforests are characterized by a closed and continuous tree canopy, moisture-dependent vegetation, the presence of epiphytes and lianas and the absence of wildfire. Rainforest can be classified as tropical rainforest or temperate rainfores ...
. The global temperature was much warmer than it is today, based on paleoclimate models. The latitudinal temperature gradient between the equator and mid-latitudes of South America was similar to the gradient that exists today. Elevated levels of
carbon dioxide Carbon dioxide (chemical formula ) is a chemical compound made up of molecules that each have one carbon atom covalently double bonded to two oxygen atoms. It is found in the gas state at room temperature. In the air, carbon dioxide is transpar ...
in the atmosphere are thought to have caused the global greenhouse temperature. The high rainfall estimates and increased pCO2 would have maintained the rainforest floras during the Paleocene greenhouse.


Paleobiogeography

The presence of dyrosaurids such as ''Cerrejonisuchus'' in the Paleocene of Colombia suggests that there was a radiation of dyrosaurids in South America following the
Cretaceous–Paleogene boundary The Cretaceous–Paleogene (K–Pg) boundary, formerly known as the Cretaceous–Tertiary (K–T) boundary, is a geological signature, usually a thin band of rock containing much more iridium than other bands. The K–Pg boundary marks the end of ...
(K–T boundary) and the
Cretaceous–Paleogene extinction event The Cretaceous–Paleogene (K–Pg) extinction event (also known as the Cretaceous–Tertiary extinction) was a sudden mass extinction of three-quarters of the plant and animal species on Earth, approximately 66 million years ago. With the ...
. There may have been a dispersal from
Africa Africa is the world's second-largest and second-most populous continent, after Asia in both cases. At about 30.3 million km2 (11.7 million square miles) including adjacent islands, it covers 6% of Earth's total surface area ...
to
Brazil Brazil ( pt, Brasil; ), officially the Federative Republic of Brazil (Portuguese: ), is the largest country in both South America and Latin America. At and with over 217 million people, Brazil is the world's fifth-largest country by area ...
, and continued immigration into
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 ...
. Colombia can be seen as a transitional route from Brazil to North America, and also to
Bolivia , image_flag = Bandera de Bolivia (Estado).svg , flag_alt = Horizontal tricolor (red, yellow, and green from top to bottom) with the coat of arms of Bolivia in the center , flag_alt2 = 7 × 7 square p ...
, assuming that it was coastal.


References

{{Taxonbar, from=Q2128842 Dyrosaurids Prehistoric pseudosuchian genera Prehistoric marine crocodylomorphs Paleocene crocodylomorphs Paleocene reptiles of South America Peligran Itaboraian Paleogene Colombia Fossils of Colombia Fossil taxa described in 2010