Caninemys Skull
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''Caninemys'' 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 large freshwater
side-necked turtle The Pleurodira are one of the two living suborders of turtles, the other being the Cryptodira. The division between these two suborders represents a very deep evolutionary divide between two very different types of turtles. The physical differen ...
, belonging to the family
Podocnemididae Podocnemididae is a family of pleurodire (side-necked) turtles, once widely distributed. Most of its 20 genera and 30 species are now extinct. Seven of its eight surviving species are native to South America: the genus ''Peltocephalus'', with onl ...
. Its
fossil A fossil (from Classical Latin , ) is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserved ...
s have been found in
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
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 ...
, in rocks dating back from the middle to late
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 ...
.


Discovery and naming

The type specimen of ''Caninemys'', a well preserved skull (DNPM-MCT 1496-R), was found in 1962 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 ...
by
Llewellyn Ivor Price Llewellyn Ivor Price (October 9, 1905 – June 9, 1980) was one of the first Brazilian paleontologists. His work contributed not only to the development of Brazilian but also to global paleontology. He collected '' Staurikosaurus'', the first ...
. Originally mentioned by Lapparent de Broin (1993) alongside several shell fragments similar to ''
Stupendemys ''Stupendemys'' is an extinct genus of freshwater side-necked turtle, belonging to the family Podocnemididae. It is the largest freshwater turtle known to have existed, with a carapace over 2 meters long. Its fossils have been found in norther ...
'', it was concluded that the data was not sufficient enough to refer the skull to the massive podocnemid, especially in the light of other similarly sized turtle remains from Miocene South America. In 2009 Meylan, Gaffney and De Almeida Campos described ''Caninemys'' on the basis of the skull alone, not including the shell fragments mentioned by de Broin due to the possibility of a chimeric combination. They do however consider it very probable that LACM 141498, a lower jaw, belongs to the same taxon. Both the holotype skull and the afforementioned lower jaw were used as a basis for the skull of the restored ''Stupendemys'' skeleton exhibited at the
American Museum of Natural History The American Museum of Natural History (abbreviated as AMNH) is a natural history museum on the Upper West Side of Manhattan in New York City. In Theodore Roosevelt Park, across the street from Central Park, the museum complex comprises 26 inter ...
. However, for the remains of the enormous ''Stupendemys'' to match the skull material of ''Caninemys'', the later had to be scaled up to twice its original size. In 2020 Cadena and colleagues described new specimens of '' Stupendemys geographica'', in the process synonymizing both ''S. souzai'' and ''Caninemys'' with this genus on the basis of lower jaws found in the same area as ''Stupendemys'' fossils. Later finds of additional ''Caninemys'' material (shells and skulls) as well as new undisputed ''Stupendemys'' skull material helped re-establish ''Caninemys'' as a valid genus. The new material was discovered in the
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 ...
n
La Victoria Formation The Honda Group ( es, Grupo Honda, Tsh, Ngh) is a geological group of the Upper and Middle Magdalena Basins and the adjacent Central and Eastern Ranges of the Colombian Andes. The group, in older literature also defined as formation, is in its ...
in the
Tatacoa Desert The Tatacoa Desert is the second largest arid zone in Colombia after the Guajira Peninsula. It occupies more than 330 square kilometers. This region is located north of Huila Department, 38 km from the city of Neiva in Colombia and from Nata ...
. ''Caninemys'' is named for its bulldog-like appearance, with the maxillary processes found in the taxon mirroring the placement of the canine (tooth) in mammals. The species name derives from the tridentate ("three toothed") appearance of the skull, as most easily observed when viewed from the front.


Description

The skull of ''Caninemys'' is robust and roughly triangular in shape with thick and massive premaxilla. The eyes are dorsally oriented, facing slightly upward. No
nasal bones The nasal bones are two small oblong bones, varying in size and form in different individuals; they are placed side by side at the middle and upper part of the face and by their junction, form the bridge of the upper one third of the nose. Eac ...
are present as in other members of
Pelomedusoides The Pleurodira are one of the two living suborders of turtles, the other being the Cryptodira. The division between these two suborders represents a very deep evolutionary divide between two very different types of turtles. The physical differenc ...
. Instead the prefrontals make up most of the anterior portion of the skull, including the area just above the nares. The prefrontals extend past the maxilla, giving it a somewhat stub-nosed appearance. The paired prefrontals, frontals and parietals all contact their respective second bone at the midline. The frontals lack the groove that is characteristic for ''Podocnemis''. Scales are preserved on the skull roof, especially over the parietals. Based on this ''Caninemys'' had triangular interparietal scales, similar to modern ''Podocnemis'' species. 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 ...
are paired, but only make up a small portion of the jaws just below the nares. The triturating (grinding) surface of ''Caninemys'' is notably more complex than in modern podocnemidids, with a sharp labial ridge that forms a small but distinct toothlike process at the midline contact of the premaxilla. The premaxilla also form a large recess to accommodate for a hooked lower jaw, as seen in the modern
Alligator Snapping Turtle The alligator snapping turtle (''Macrochelys temminckii'') is a large species of turtle in the family Chelydridae. The species is native to freshwater habitats in the United States. ''M. temminckii'' is one of the heaviest freshwater turtles in ...
. The
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 ...
of ''Caninemys'' are noticeably enlarged and give the animal a superficially "bulldog-like" appearance. The maxilla also stands out for the large, tooth-like processes formed by the labial ridge and flanking the premaxilla on both sides. These processes, superficially resembling the canine teeth seen in mammals, are very noticeable when the skull is viewed head-on. These two processes, alongside the premaxillary process, account for the tridentate condition that gives ''Caninemys tridentata'' its species name. Even without these processes the labial ridge is very pronounced in ''Caninemys'', being thick and extending downward 2 centimers into a flattened edge. The ridge curves upwards close to the maxilla, creating a u-shaped notch. However, ''Caninemys'' does not possess a
secondary palate The secondary palate is an anatomical structure that divides the nasal cavity from the oral cavity in many vertebrates. In human embryology, it refers to that portion of the hard palate that is formed by the growth of the two palatine shelves medi ...
like in '' Bairdemys'', '' Stereogenys'' and '' Shweboemys''. The lower jaw LACM 141498 may represent the same taxon as the ''Caninemys'' holotype. The lower jaw possesses a pronounced symphyseal hook similar to those seen in modern alligator snapping and musk turtles. Although the jaw is slightly too large to belong to the exact same specimen as the holotype, the jaws line up well. While there is no direct link between the two fossils, they match closely enough to suggest they belong to at least closely related taxa, if not the same species. This would suggest that ''Caninemys'', besides having pronounced maxilla, also had a robust and hooked lower jaw. This combination of features leads Meylan ''et al.'' to compare ''Caninemys'' to a "pleurodiran snapping turtle". Based on the head:shell ratio of extant podocnemidid species (including ''
Podocnemis expansa The Arrau turtle (''Podocnemis expansa''), also known as the South American river turtle, giant South American turtle, giant Amazon River turtle, Arrau sideneck turtle, Amazon River turtle or simply the Arrau, is the largest of the side-neck tu ...
''), ''Caninemys'' was likely much smaller than ''Stupendemys'', reaching an estimated length of 1.2 to 1.5 meters.


Phylogeny

''Caninemys'' shows multiple morphological characters that clearly identify it as a pelomedusoid pleurodire, specifically a podocnemidid. The phylogenetic tree below is based on the initial analysis conducted in 2009. The following phylogenetic tree is a simplified version of the one recovered by Cadena ''et al.'' (2021), recovering ''Stupendemys'' and ''Caninemys'' as members of two separate branches within Podocnemididae. The scoring of this tree was significantly improved through the discovery of additional remains for both taxa. While ''Stupendemys'' claded with the extant Big-headed Amazon River turtle, ''Caninemys'' closest living relative was found to be the
Madagascan big-headed turtle The Madagascan big-headed turtle (''Erymnochelys madagascariensis'') is a turtle native to the waters of permanent slow moving rivers and lakes in western Madagascar. These turtles are critically endangered and have been evaluated to be the most ...
.


Paleobiology

The three tooth-like processes of ''Caninemys'' are similar to those seen in the living '' Claudius angustatus'', the Narrow-bridged musk turtle, a member of a group known to utilize a specialised
hyoid The hyoid bone (lingual bone or tongue-bone) () is a horseshoe-shaped bone situated in the anterior midline of the neck between the chin and the thyroid cartilage. At rest, it lies between the base of the mandible and the third cervical vertebra. ...
apparatus to capture highly mobile prey via vacuum feeding. Such a feeding style in combination with the massive jaws and tooth-like processes may suggests that it would have been a predatory animal with a lifestyle similar to snapping turtles, feeding on fish, crocodilians and snakes respectively. This hypothesis was later contested by Cadena and colleagues in 2020, who instead proposed that ''Caninemys'' (by them treated as a synonym of ''Stupendemys'') had a much broader diet not exclusive to vertebrate prey. However, in their 2021 follow-up paper, Cadena and colleagues once again inferred a more carnivorous, vacuum-feeding strategy for ''Caninemys'' in opposition to a generalist/durophagous ''Stupendemys'', arguing that these dietary differences might explain the co-occurrence of two large-bodied turtles within the same ecosystems.


References

{{Testudines Prehistoric turtles of South America Podocnemididae Prehistoric turtle genera Extinct turtles Miocene turtles Miocene genus first appearances Miocene extinctions Miocene reptiles of South America Neogene Brazil Fossils of Brazil Neogene Colombia Fossils of Colombia Fossil taxa described in 2009