Pakasuchus Dentition
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''Pakasuchus'' is a genus of notosuchian
crocodyliform Crocodyliformes is a clade of crurotarsan archosaurs, the group often traditionally referred to as "crocodilians". They are the first members of Crocodylomorpha to possess many of the features that define later relatives. They are the only pseudo ...
distinguished by its unusual mammal-like appearance, including mammal-like teeth that would have given the animal the ability to chew. It also had long, slender legs and a doglike nose. ''Pakasuchus'' lived approximately 105 million years ago, in the mid-
Cretaceous The Cretaceous ( ) is a geological period that lasted from about 145 to 66 million years ago (Mya). It is the third and final period of the Mesozoic Era, as well as the longest. At around 79 million years, it is the longest geological period of th ...
. Fossils have been found in the
Galula Formation The Galula Formation is a geological formation located south of Lake Rukwa in Tanzania, part of the Red Sandstone Group of the Rukwa Rift Basin. Along with the unconformably overlying Oligocene Nsungwe Formation. It is divided into two members, t ...
of Rukwa Rift Basin of southwestern
Tanzania Tanzania (; ), officially the United Republic of Tanzania ( sw, Jamhuri ya Muungano wa Tanzania), is a country in East Africa within the African Great Lakes region. It borders Uganda to the north; Kenya to the northeast; Comoro Islands and ...
, and were described in 2010 in the journal ''
Nature Nature, in the broadest sense, is the physics, physical world or universe. "Nature" can refer to the phenomenon, phenomena of the physical world, and also to life in general. The study of nature is a large, if not the only, part of science. ...
''. 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 ''P. kapilimai''. ''Pakasuchus'' means "cat crocodile" (''paka'' meaning "cat" in Kiswahili) in reference to its catlike appearance and probable behavior.


Description

''Pakasuchus'' was around long. Like all notosuchians, it was an active terrestrial animal. It probably hunted small prey such as insects. It had a short, broad skull somewhat similar in shape to a cat. Unlike living
crocodilia Crocodilia (or Crocodylia, both ) is an order of mostly large, predatory, semiaquatic reptiles, known as crocodilians. They first appeared 95 million years ago in the Late Cretaceous period ( Cenomanian stage) and are the closest living ...
ns, ''Pakasuchus'' had distinctive
heterodont 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, ...
teeth that varied in shape throughout its jaws. There are large, sharp teeth near the front of the jaws, and broad molar-like teeth at the back of the mouth. While
multicuspid The molars or molar teeth are large, flat teeth at the back of the mouth. They are more developed in mammals. They are used primarily to grind food during chewing. The name ''molar'' derives from Latin, ''molaris dens'', meaning "millstone to ...
teeth are seen in many other notosuchians such as ''
Simosuchus ''Simosuchus'' (meaning "pug-nosed crocodile" in Greek, referring to the animal's blunt snout) is an extinct genus of notosuchian crocodylomorphs from the Late Cretaceous of Madagascar. It is named for its unusually short skull. Fully grown indi ...
'' and ''
Yacarerani ''Yacarerani'' (meaning "first Yacare" in Guaraní) is an extinct genus of Late Cretaceous notosuchian crocodylomorph. Fossils have been found in 2002 in the Cajones Formation in Amboró National Park of central Bolivia and are Turonian-San ...
'', they are most complex in ''Pakasuchus''. The molar-like teeth show a level of complexity that matches that of mammals, being able to occlude, or fit with one another, and provide sharp shearing edges for slicing food. ''Pakasuchus'' also differed from modern crocodilians, and many other crocodyliforms, in the reduction of
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 covering the body. Small, reduced osteoderms overlie the dorsal vertebrae, but fewer in number and are not as large as those of other notosuchians. However, caudal osteoderms still covered the tail. The loss of osteoderms on the body and retention of them on the tail is unique among crocodyliforms.


Discovery

A complete skeleton of ''Pakasuchus'' was found in 2008 in southwestern Tanzania by an international research team funded by the
National Science Foundation The National Science Foundation (NSF) is an independent agency of the United States government that supports fundamental research and education in all the non-medical fields of science and engineering. Its medical counterpart is the National I ...
and the
National Geographic Society The National Geographic Society (NGS), headquartered in Washington, D.C., United States, is one of the largest non-profit scientific and educational organizations in the world. Founded in 1888, its interests include geography, archaeology, and ...
as part of the Rukwa Rift Basin Project. Remains from six other individuals were later uncovered. The specific name honors Saidi Kapilima, one of the leaders of the Rukwa Rift Basin Project and who helped in the excavation of the specimens. The most complete specimen includes a nearly complete skull. Because the jaws were closed in this specimen, some of the teeth were obscured. The describers of the specimens used X-ray computed tomography scanning, or CT scanning, to image the teeth. This provided a detailed view of the animal's dentition that could not normally be observed in the specimen.


Phylogeny

Below is a cladogram modified from O’Connor ''et al.'', 2010:


Paleobiology

''Pakasuchus'' was probably very mammal-like in behavior as well as appearance. The molariform teeth are well suited to shearing food like modern mammalian carnivores. In fact, the teeth are so similar to those of mammals that it has led paleontologist Greg Buckley to state "If only isolated teeth had been discovered, without the skull, it is very likely that some of the molariform teeth would have been mistaken for a mammal's." The relative lack of osteoderms on the body may have been an adaptation for an active terrestrial lifestyle, as it would have lightened the animal. However, the retention of osteoderms on the tail is hard to explain, as they would have been quite heavy. The long legs and slender build would also have made ''Pakasuchus'' more agile. ''Pakasuchus'' was probably an herbivore; its dentition was at least as complex as those of modern herbivorous
lizard Lizards are a widespread group of squamate reptiles, with over 7,000 species, ranging across all continents except Antarctica, as well as most oceanic island chains. The group is paraphyletic since it excludes the snakes and Amphisbaenia alt ...
s. Notosuchians were widespread across
Gondwana Gondwana () was a large landmass, often referred to as a supercontinent, that formed during the late Neoproterozoic (about 550 million years ago) and began to break up during the Jurassic period (about 180 million years ago). The final stages ...
throughout the Cretaceous. ''Pakasuchus'', as well as many other notosuchians, would have filled
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 ...
s in these areas that were otherwise occupied by mammals in the northern continents. Mammals were relatively uncommon in Gondwana at the time, making it possible for notosuchians to occupy a similar niche. The Rukwa Rift Basin is known for a rich vertebrate fauna that existed during the Cretaceous. During the Early Cretaceous, the basin was part of a large river system with braided channels and low-lying vegetated floodplains. Several dinosaurs are known from the basin that would have lived alongside ''Pakasuchus'', including large
sauropod Sauropoda (), whose members are known as sauropods (; from '' sauro-'' + '' -pod'', 'lizard-footed'), is a clade of saurischian ('lizard-hipped') dinosaurs. Sauropods had very long necks, long tails, small heads (relative to the rest of their bo ...
s and theropods. Aquatic crocodyliforms inhabited the rivers along with
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
fishes 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 ...
.


References


External links


"These Crocs Are Made for Biting!" - video discussing ''Pakasuchus'' from ''National Geographic Daily News''
{{Taxonbar, from=Q1959540 Early Cretaceous crocodylomorphs of Africa Fossil taxa described in 2010 Ziphosuchians Prehistoric pseudosuchian genera Albian genera