Caiman Brevirostris
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''Caiman brevirostris'' 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 ...
species In biology, a species is the basic unit of classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of the appropriate s ...
of
caiman A caiman (also cayman as a variant spelling) is an alligatorid belonging to the subfamily Caimaninae, one of two primary lineages within the Alligatoridae family, the other being alligators. Caimans inhabit Mexico, Central and South America fro ...
that lived during the
Late Miocene The Late Miocene (also known as Upper Miocene) is a sub-epoch of the Miocene epoch (geology), Epoch made up of two faunal stage, stages. The Tortonian and Messinian stages comprise the Late Miocene sub-epoch, which lasted from 11.63 Ma (million ye ...
, around 11.6 million years ago, to the end of the Miocene 5.3 million years ago in
Acre The acre is a unit of land area used in the imperial Imperial is that which relates to an empire, emperor, or imperialism. Imperial or The Imperial may also refer to: Places United States * Imperial, California * Imperial, Missouri * Imp ...
and
Amazonas, Brazil Amazonas () is a state of Brazil, located in the North Region in the northwestern corner of the country. It is the largest Brazilian state by area and the 9th largest country subdivision in the world, and the largest in South America, being g ...
as well as Urumaco, Venezuela. Several specimens have been referred to the species, but only 3 of them are confidently placed in the species. ''C. brevirostris'' was originally named in 1987 on the basis of a single, incomplete rostrum with an associated mandibular ramus that had been found in Acre, Brazil. ''C. brevirostris'' is very distinct among ''Caiman'' species and caimaninae overall in that it preserves a characteristically short and robust skull that bears blunt posterior teeth that were built to break down harder foods. This was an adaption for
durophagy Durophagy is the eating behavior of animals that consume hard-shelled or exoskeleton bearing organisms, such as corals, shelled mollusks, or crabs. It is mostly used to describe fish, but is also used when describing reptiles, including fossil tu ...
(the diet of hard-shelled organisms), likely to crush shells of mollusks and clams which were common in the wetlands that ''C. brevirostris'' resided in.


Discovery and naming

''Caiman brevirostris'' was described in 1987 by Brazilian paleontologist J. D. Souzha Filho on the basis of an incomplete rostrum and associated right mandibular ramus that were collected from the
Miocene The Miocene ( ) is the first geological epoch of the Neogene Period and extends from about (Ma). The Miocene was named by Scottish geologist Charles Lyell; the name comes from the Greek words (', "less") and (', "new") and means "less recen ...
age strata of the Solimões Formation in the municipality of
Sena Madureira Sena Madureira () is a Municipalities of Brazil, municipality located in the center of the States of Brazil, Brazilian state of Acre (state), Acre. Its population is 46,511 and its area is , making it the largest municipality in the state. It ha ...
in Acre, Brazil. The species name “''brevirostris''” comes from the Greek roots ''brevis-'' meaning “short” and ''-rostrum'' meaning “snout” after the species’ distinct short and robust snout.Souza-Filho, J. D. (1987, July). Caiman brevirostris sp. nov., um novo Alligatoridae da Formação Solimões (Pleistoceno) do Estado do Acre, Brasil. In ''Anais X Congresso Brasileiro de Paleontologia, Rio de Janeiro'' (Vol. 1, pp. 173-180). Several additional specimens from the
Urumaco Formation The Urumaco Formation is a formation in Venezuela that includes deposits from the Late Miocene. It is the site of several "giant forms": the turtles, crocodiles, sloths and rodents of Urumaco are among the largest of their groups. Location Th ...
of
Venezuela Venezuela (; ), officially the Bolivarian Republic of Venezuela ( es, link=no, República Bolivariana de Venezuela), is a country on the northern coast of South America, consisting of a continental landmass and many islands and islets in th ...
were incorrectly referred to the species in the 2000s and 2010s, many of these specimens actually being from other caimanines. In 2003 during a joint expedition between the Universidade Federal do Acre and the
Universidade Federal de Rondônia The Universiade is an international multi-sport event, organized for university athletes by the International University Sports Federation (FISU). The name is a portmanteau of the words "University" and "Olympiad". The Universiade is referred t ...
, a complete skull and associated postcranial material of ''Caiman brevirostris'' were collected from the
Upper Miocene The Late Miocene (also known as Upper Miocene) is a sub-epoch of the Miocene Epoch made up of two stages. The Tortonian and Messinian stages comprise the Late Miocene sub-epoch, which lasted from 11.63 Ma (million years ago) to 5.333 Ma. The ev ...
strata of the Solimões Formation in Talismã locality in Manuel Urban municipality in Amazonas, Brazil. The specimen was deposited in the collections of the Universidade Federal do Acre and described in 2014 in a study that found only the Talismã specimen and the holotype could be referred to the species. However, a single specimen, an articulated skull and mandible, from the
Urumaco Formation The Urumaco Formation is a formation in Venezuela that includes deposits from the Late Miocene. It is the site of several "giant forms": the turtles, crocodiles, sloths and rodents of Urumaco are among the largest of their groups. Location Th ...
was also referred to ''C. brevirostris'' in 2016.


Description and paleobiology

The most distinctive feature of ''Caiman brevirostris'' is its short and broad skull, the shortest skull known from a caimanine, that was built for crushing mollusks like its relative ''
Globidentosuchus ''Globidentosuchus'' is an extinct genus of basal caimanine crocodylian known from the late Middle to Late Miocene of the Middle and the Upper Members of the Urumaco Formation at Urumaco, Venezuela. Its skull was very short and robust, with lar ...
''. This is also shown in the rostral anatomy, which has a short and wide splenial with few alveoli compared to other caimanines. The external nares are enlarged compared to other ''Caiman'' species, though this condition is similar to that of the "La Venta" ''Caiman'' from the Miocene of Colombia. The size of the
orbits 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 ...
is also enlarged compared to other caimanines, but this characteristic is also present in '' Melanosuchus niger''. One of the most unique traits of ''C. brevirostris'' is the blunt & low crowned
posterior teeth {{unreferenced, date=November 2015 In dentistry, the term posterior teeth usually refers as a group to the premolars and molars, as distinguished from the anterior teeth, which are the incisors and canine teeth. The distinction is one of anterior ( ...
that were adapted for eating harder prey than other caimanines. 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 ...
bears a long posterior process despite the short rostral bones. This process extends posteriorly past the 3rd maxillary alveolus, a trait of brevirostry. The prefrontals are separated by the frontal only and do not meet at the midline, in contrast to Venezuelan caimanines whose prefrontals contact at the midline. The frontal anteriorly projects beyond the prefrontal’s anterior tip, which is likely again caused by the skull’s shortening compared to other species. Majority of the diagnostic traits of ''C. brevirostris'' lie in the mandible. ''C. brevirostris''’ mandible is short and wide, though the dentary symphysis is broken in all known specimens, leaving gaps in the knowledge of its anatomy. The
splenial The splenial is a small bone in the lower jaw of reptiles, amphibians and birds, usually located on the lingual side (closest to the tongue) between the angular and surangular The suprangular or surangular is a jaw bone found in most land ver ...
is completely different from those of other caimanines in that it is extremely robust and approaches the symphysis. The splenial contributes to half of the breadth of the mandibular ramus, forming a wide floor posteriorly, similar to that of ''Caiman latirostris'', but is more pronounced and differs greatly from that of narrow-snouted caimanines. The wide posterior part of the splenial in ''C. brevirostris'' and the wide mandibles as a whole may be due to frequent usage of the posterior blunt and crushing teeth, as possibly being used for greater muscle attachment, but the muscles present in the area play only a minor role in jaw mechanics. The dentary has fewer alveoli compared to other ''Caiman'' species, again due to the shortening of the skull and mandible. The largest alveolus bares a blunt and low-crowned tooth, which may used for crushing feeding behavior, as in ''C. latirostris''. ''Caiman brevirostris'' would therefore be an ecological analogue of the extant ''Caiman latirostris'' during the late Miocene in the Solimões Formation and Urumaco Formation.


Classification

''Caiman brevirostris'' has been included in several phylogenetic analyses, but it most frequently comes up in polytomy with other fossil Caiman species. Below is the phylogenetic analysis conducted by Salas-Gismondi ''et al''., 2015:


Paleoenvironment

As the Proto-Amazonian lake system and the Pebas system began to dissipate with the onset of the transcontinental Amazon Drainage, ''Caiman brevirostris'' inhabited the wetlands of the northern
Urumaco Formation The Urumaco Formation is a formation in Venezuela that includes deposits from the Late Miocene. It is the site of several "giant forms": the turtles, crocodiles, sloths and rodents of Urumaco are among the largest of their groups. Location Th ...
and the
Solimões Formation The Pebas Formation is a lithostratigraphic unit of Miocene age, found in western Amazonia. The formation extends over , including parts of Brazil, Peru, Ecuador and Colombia.Wesselingh et al., 2006 It is interpreted as representing the deposits ...
in
Acre State Acre () is a state located in the west of the North Region of Brazil and the Amazonia Legal. Located in the westernmost part of the country, at a two-hour time difference from Brasília, Acre is bordered clockwise by the Brazilian states of Am ...
,
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 ...
, into the Late Miocene before eventually dying out during the Early Pliocene like much of the large crocodilian fauna of the Miocene wetlands. These wetlands provided favorable conditions to the native reptilian fauna, with several lineages of crocodilians reaching enormous sizes during the Mid to Late Miocene and also diversifying in ecology. Some of the enormous crocodilians that coexisted with ''C. brevirostris'' included the enormous caimain ''
Purussaurus ''Purussaurus'' is an extinct genus of giant caiman that lived in South America during the Miocene epoch, from the Friasian to the Huayquerian in the SALMA classification. It is known from skull material found in the Brazilian and Peruvian Amazon ...
'', the bizarre ''
Mourasuchus ''Mourasuchus'' is an extinct genus of giant, aberrant caiman from the Miocene of South America. Its skull has been described as duck-like, being broad, flat, and very elongate, superficially resembling '' Stomatosuchus'' from the Late Cretaceo ...
'' and large-bodied gharials of the genus ''
Gryposuchus ''Gryposuchus'' is an extinct genus of gavialid crocodilian. Fossils have been found from Argentina, Colombia, Venezuela, Brazil and the Peruvian Amazon. The genus existed during the Miocene epoch (Colhuehuapian to Huayquerian). One recently d ...
'', some species of which reaching lengths of over 10 meters. The largest turtle known, ''Stupendemys'', with one specimen preserving a 2.86 meter long carapace, was also present in the region as an omnivore. Other durophagus caimanines inhabited the Urumaco and the Solimões, including the unusual ''Globidentosuchus'' in Urumaco and the "shovel faced" ''Gnatusuchus'' in the Solimões. Besides the aforementioned reptiles the waterways of Late Miocene South America were also inhabited by fish, including
catfish Catfish (or catfishes; order Siluriformes or Nematognathi) are a diverse group of ray-finned fish. Named for their prominent barbels, which resemble a cat's whiskers, catfish range in size and behavior from the three largest species alive, ...
such as ''
Phractocephalus The redtail catfish, ''Phractocephalus hemioliopterus'', is a pimelodid (long-whiskered) catfish. In Venezuela, it is known as cajaro, and in Brazil, it is known as pirarara, stemming from the Tupi language words ''pirá'' and ''arara''. It i ...
'' and
Callichthyidae Callichthyidae is a family of catfishes ( order Siluriformes), called armored catfishes due to the two rows of bony plates (or scutes) along the lengths of their bodies. It contains some of the most popular freshwater aquarium fish, such as many ...
, characids such as '' Acregoliath rancii'' and the
tambaqui The tambaqui (''Colossoma macropomum'') is a large species of freshwater fish in the family Serrasalmidae. It is native to tropical South America, but kept in aquaculture and introduced elsewhere. It is also known by the names black pacu, blac ...
(''Colossoma macropomum''), the
South American lungfish The South American lungfish (''Lepidosiren paradoxa'') is the single species of lungfish found in swamps and slow-moving waters of the Amazon, Paraguay, and lower Paraná River basins in South America. Notable as an obligate air-breather, it is ...
(''Lepidosiren paradoxa''), trahiras (e.g. '' Paleohoplias assisbrasiliensis'') and freshwater
rays Ray may refer to: Fish * Ray (fish), any cartilaginous fish of the superorder Batoidea * Ray (fish fin anatomy), a bony or horny spine on a fin Science and mathematics * Ray (geometry), half of a line proceeding from an initial point * Ray (gra ...
and
sharks Sharks are a group of elasmobranch fish characterized by a cartilaginous skeleton, five to seven gill slits on the sides of the head, and pectoral fins that are not fused to the head. Modern sharks are classified within the clade Selachimorp ...
. Other turtles and
tortoises Tortoises () are reptiles of the family Testudinidae of the order Testudines (Latin: ''tortoise''). Like other turtles, tortoises have a shell to protect from predation and other threats. The shell in tortoises is generally hard, and like other ...
found in the same deposits are '' Chelus columbiana'' (a fossil relative of the
mata mata Matamata () is a town in Waikato, New Zealand. It is located near the base of the Kaimai Ranges, and is a thriving farming area known for Thoroughbred horse breeding and training pursuits. It is part of the Matamata-Piako District, which takes ...
) and ''
Chelonoidis ''Chelonoidis'' is a genus of turtles in the tortoise family erected by Leopold Fitzinger in 1835. They are found in South America and the Galápagos Islands, and formerly had a wide distribution in the West Indies. The multiple subspecies of t ...
''. Further aquatic vertebrates included
river dolphins River dolphins are a polyphyletic group of fully aquatic mammals that reside exclusively in freshwater or brackish water. They are an informal grouping of dolphins, which itself is a paraphyletic group within the infraorder Cetacea. Extant riv ...
and the large
darter The darters, anhingas, or snakebirds are mainly tropical waterbirds in the family Anhingidae, which contains a single genus, ''Anhinga''. There are four living species, three of which are very common and widespread while the fourth is rarer and ...
'' "Anhinga" fraileyi''. At least within the Solimões Formation ''Stupendemys'' would have inhabited a floodplain or lacustrine environment with savannahs and gallery forests.


References

{{Taxonbar, from=Q104830572 Fossil taxa described in 1987 Miocene reptiles of South America Alligatoridae