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''Caipirasuchus'' 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 sphagesaurid notosuchians known from 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 ...
of northern
São Paulo State SAO or Sao may refer to: Places * Sao civilisation, in Middle Africa from 6th century BC to 16th century AD * Sao, a town in Boussé Department, Burkina Faso * Saco Transportation Center (station code SAO), a train station in Saco, Maine, U.S ...
, southeastern
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 ...
. 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 ...
, ''C. paulistanus'', was named in 2011. A second species, ''C. montealtensis'', was referred to ''Caipirasuchus'' in 2013 after having been named in 2008 as a species of ''
Sphagesaurus ''Sphagesaurus'' is an extinct genus of sphagesaurid notosuchian crocodylomorph from the Late Cretaceous of southwest São Paulo, southern Brazil. Species and discovery ''Sphagesaurus'' was first described and named by Llewellyn Ivor Price in ...
''. A third species, ''C. stenognathus'', was described in 2014. A fourth species, ''C. mineirus'', was described in 2018. A fifth species, ''C. attenboroughi'', was named in 2021 in honour of
David Attenborough Sir David Frederick Attenborough (; born 8 May 1926) is an English broadcaster, biologist, natural historian and author. He is best known for writing and presenting, in conjunction with the BBC Natural History Unit, the nine natural histor ...
.


Discovery and naming

''Caipirasuchus'' was first named by Fabiano V. Iori and Ismar S. Carvalho in
2011 File:2011 Events Collage.png, From top left, clockwise: a protester partaking in Occupy Wall Street heralds the beginning of the Occupy movement; protests against Libyan dictator Muammar Gaddafi, who was killed that October; a young man celebrate ...
and 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. paulistanus''. The generic name is derived from Portuguese word Caipira which refers to the rural inhabitants of the Brazilian states, covering almost all of the Bauru Basin, and Greek ''souchus'' meaning "crocodile". The
specific name Specific name may refer to: * in Database management systems, a system-assigned name that is unique within a particular database In taxonomy, either of these two meanings, each with its own set of rules: * Specific name (botany), the two-part (bino ...
is derived from "
paulista Paulista is a municipality in Pernambuco, Brazil, with a population of 334,376 as of 2020. It has the highest Human Development Index (HDI) of the Recife metropolitan area. It is the birthplace of footballer Rivaldo and is also famous for its beac ...
", the designation for the residents of São Paulo State, where the holotype was found. ''C. paulistanus'' is known only from 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 MPMA 67-0001/00, an almost complete and undeformed
skull The skull is a bone protective cavity for the brain. The skull is composed of four types of bone i.e., cranial bones, facial bones, ear ossicles and hyoid bone. However two parts are more prominent: the cranium and the mandible. In humans, the ...
and
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 ...
with teeth (only three teeth are missing) and partial
postcranium Postcrania (postcranium, adjective: postcranial) in zoology and vertebrate paleontology is all or part of the skeleton apart from the skull. Frequently, fossil remains, e.g. of dinosaurs or other extinct tetrapods, consist of partial or isolated sk ...
from a single individual. It was found on the São Francisco Farm, in Homem de Mello, the rural area of Monte Alto County of northern São Paulo. It was collected from the
Adamantina Formation The Adamantina Formation is a geological formation in the Bauru Basin of western São Paulo state, in southeastern Brazil. Its strata date back to the Late Cretaceous epoch of the Cretaceous Period, during the Mesozoic Era. The formation is p ...
of the
Bauru Basin The Bauru Group is a geological group of the Bauru Sub-basin, Paraná Basin in Minas Gerais, São Paulo, General Salgado, Itapecuru-Mirim, Mato Grosso, Brazil whose strata date back to the Late Cretaceous. Dinosaur remains are among the fossils ...
, which dates to the
Turonian The Turonian is, in the ICS' geologic timescale, the second age in the Late Cretaceous Epoch, or a stage in the Upper Cretaceous Series. It spans the time between 93.9 ± 0.8 Ma and 89.8 ± 1 Ma (million years ago). The Turonian is preceded by t ...
and
Santonian The Santonian is an age in the geologic timescale or a chronostratigraphic stage. It is a subdivision of the Late Cretaceous Epoch or Upper Cretaceous Series. It spans the time between 86.3 ± 0.7 mya (million years ago) and 83.6 ± 0.7 mya. The ...
stages of the Late Cretaceous. A second species, ''C. montealtensis'', was named by Marco Brandalise de Andrade and Reinaldo J. Bertini in
2008 File:2008 Events Collage.png, From left, clockwise: Lehman Brothers went bankrupt following the Subprime mortgage crisis; Cyclone Nargis killed more than 138,000 in Myanmar; A scene from the opening ceremony of the 2008 Summer Olympics in Beijing; ...
as a species of the closely related notosuchian genus ''
Sphagesaurus ''Sphagesaurus'' is an extinct genus of sphagesaurid notosuchian crocodylomorph from the Late Cretaceous of southwest São Paulo, southern Brazil. Species and discovery ''Sphagesaurus'' was first described and named by Llewellyn Ivor Price in ...
'', on the basis of MPMA 15-001/90, a nearly complete skull and mandible preserved in occlusion. It was collected from the Bairro Cachoeira locality at the base of the Serra da
Água Limpa Água Limpa is a municipality in south Goiás state, Brazil. The population was 1,830 in 2020 and the municipal area was 454.3 km2. Geography Água Limpa is located in the Meia Ponte Microregion (SeCitybrazilfor all the regions) in the extre ...
, about 8 km northwest of Monte Alto, from the Adamantina Formation. In 2013, this specimen was reassigned to ''Caipirasuchus'' on the basis of a newly discovered specimen referrable to it. MPMA 68-0003/12 consists of a nearly complete cranium and mandible, and a posterior portion of the post-cranium. It was discovered in the municipality of
Catanduva Catanduva is a municipality in the state of São Paulo, Brazil. The population is 122,497 (2020 est.) in an area of 290.59 km2. Is the second largest city in the Northern part of the state, after São José do Rio Preto. The city has a divers ...
, northern São Paulo, from the Adamantina Formation. At least five
synapomorphies In phylogenetics, an apomorphy (or derived trait) is a novel character or character state that has evolved from its ancestral form (or plesiomorphy). A synapomorphy is an apomorphy shared by two or more taxa and is therefore hypothesized to have ...
unite the species, and 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 found them to be sister taxa.


Description

''C. paulistanus'' has about ten autapomorphies or unique characteristics. The external
naris 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 ...
(the nostril opening in the skull) is bordered only by 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 ...
e bones. Each premaxilla has four teeth set into it. There is a gap called a
diastema A diastema (plural diastemata, from Greek διάστημα, space) is a space or gap between two teeth. Many species of mammals have diastemata as a normal feature, most commonly between the incisors and molars. More colloquially, the condition ...
in the premaxillary tooth row, and a diastema in between the premaxillary and
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 teeth. The
dentary bone 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 has ten teeth on either side, with two diastemata separating them. The forward-most teeth are suited for capturing food, while the back teeth are adapted for food processing. The
palatine bone In anatomy, the palatine bones () are two irregular bones of the facial skeleton in many animal species, located above the uvula in the throat. Together with the maxillae, they comprise the hard palate. (''Palate'' is derived from the Latin ''pa ...
in the roof of the mouth connects to the maxilla bone of the snout by a region of bone called the cuneiform process. ''Caipirasuchus'' also has large pterygoid and ectopterygoid bones and a well-developed hole in front of the eye sockets called the antorbital fenestra.


References

{{Taxonbar, from=Q5017641 Sphagesaurids Terrestrial crocodylomorphs Late Cretaceous crocodylomorphs of South America Adamantina Formation Fossil taxa described in 2011 Prehistoric pseudosuchian genera