''Rukwasuchus'' is an

extinct Extinction is the termination of an organism by the death of its Endling, last member. A taxon may become Functional extinction, functionally extinct before the death of its last member if it loses the capacity to Reproduction, reproduce and ...

genus Genus (; : genera ) is a taxonomic rank above species and below family (taxonomy), family as used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In bino ...

peirosaurid Peirosauridae is a Gondwanan family of mesoeucrocodylians that lived during the Cretaceous period. It was a clade of terrestrial crocodyliforms that evolved a rather dog-like skull, and were terrestrial carnivores. It was phylogenetically defined ...

crocodyliform Crocodyliformes is a clade of Crurotarsi, 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 ...

s known from the middle

Cretaceous The Cretaceous ( ) is a geological period that lasted from about 143.1 to 66 mya (unit), million years ago (Mya). It is the third and final period of the Mesozoic Era (geology), Era, as well as the longest. At around 77.1 million years, it is 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, ...

of southwestern

Tanzania Tanzania, officially the United Republic of Tanzania, is a country in East Africa within the African Great Lakes region. It is bordered by Uganda to the northwest; Kenya to the northeast; the Indian Ocean to the east; Mozambique and Malawi to t ...

. It contains a single species, ''Rukwasuchus yajabalijekundu''.

Discovery

''Rukwasuchus'' is known from its

holotype A holotype (Latin: ''holotypus'') is a single physical example (or illustration) of an organism used when the species (or lower-ranked taxon) was formally described. It is either the single such physical example (or illustration) or one of s ...

, RRBP 08630, a well-preserved rear part of the

skull The skull, or cranium, is typically a bony enclosure around the brain of a vertebrate. In some fish, and amphibians, the skull is of cartilage. The skull is at the head end of the vertebrate. In the human, the skull comprises two prominent ...

including the cranial table,

braincase In human anatomy, the neurocranium, also known as the braincase, brainpan, brain-pan, or brainbox, is the upper and back part of the skull, which forms a protective case around the brain. In the human skull, the neurocranium includes the calv ...

, and interorbital region lacking the rostrum, the front portion of the

palate The palate () is the roof of the mouth in humans and other mammals. It separates the oral cavity from the nasal cavity. A similar structure is found in crocodilians, but in most other tetrapods, the oral and nasal cavities are not truly sep ...

, both

lacrimals The lacrimal bones are two small and fragile bones of the facial skeleton; they are roughly the size of the little fingernail and situated at the front part of the medial wall of the orbit. They each have two surfaces and four borders. Several bon ...

, jugals, and quadratojugals, as well as the

mandible In jawed vertebrates, the mandible (from the Latin ''mandibula'', 'for chewing'), lower jaw, or jawbone is a bone that makes up the lowerand typically more mobilecomponent of the mouth (the upper jaw being known as the maxilla). The jawbone i ...

. RRBP 08630 was collected during 2008 at Namba 2 locality (also known as RRBP 2007-02), together with the

titanosaurian Titanosaurs (or titanosaurians; members of the group Titanosauria) were a diverse group of Sauropoda, sauropod dinosaurs, including genera from all seven continents. The titanosaurs were the last surviving group of long-necked sauropods, with tax ...

'' Rukwatitan bisepultus'' which is exclusive to this locality. Material referred to ''Rukwasuchus'' includes four isolated

teeth A tooth (: teeth) is a hard, calcified structure found in the jaws (or mouths) of many vertebrates and used to break down food. Some animals, particularly carnivores and omnivores, also use teeth to help with capturing or wounding prey, tear ...

, which came from the neighboring localities RRBP 2007-01 yielding the 3 teeth RRBP 07351, 07369, 09362, and RRBP 2009-01 yielding the tooth RRBP 09367. All specimens came from approximately 25 km south of

Lake Rukwa Lake Rukwa is an endorheic lake located in the Rukwa Valley of Rukwa Region, Songwe Region and Katavi Region in southwestern Tanzania. The lake is the third largest inland body of water in the country. Geography The alkaline Lake Rukwa lies ...

in the Galula Study Area, Rukwa Rift Basin of southwestern Tanzania, belonging to the Namba Member of the

which dates to the late

Aptian The Aptian is an age (geology), age in the geologic timescale or a stage (stratigraphy), stage in the stratigraphic column. It is a subdivision of the Early Cretaceous, Early or Lower Cretaceous epoch (geology), Epoch or series (stratigraphy), S ...

or possibly early

Cenomanian The Cenomanian is, in the International Commission on Stratigraphy's (ICS) geological timescale, the oldest or earliest age (geology), age of the Late Cretaceous epoch (geology), Epoch or the lowest stage (stratigraphy), stage of the Upper Cretace ...

stage of the middle

, approximately 100 mya.

Etymology

''Rukwasuchus'' was named by Joseph J. W. Sertich and Patrick M. O’Connor in

2014 The year 2014 was marked by the surge of the Western African Ebola epidemic, West African Ebola epidemic, which began in 2013, becoming the List of Ebola outbreaks, most widespread outbreak of the Ebola, Ebola virus in human history, resul ...

and the

type species In International_Code_of_Zoological_Nomenclature, 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 spe ...

is ''Rukwasuchus yajabalijekundu''. The generic name refers to

and the Rukwa Rift Basin, located in southwestern Tanzania, where the holotype of ''Rukwasuchus'' and other vertebrates were collected by the Rukwa Rift Basin Project, and ''suchus'',

Latin Latin ( or ) is a classical language belonging to the Italic languages, Italic branch of the Indo-European languages. Latin was originally spoken by the Latins (Italic tribe), Latins in Latium (now known as Lazio), the lower Tiber area aroun ...

ized from the

Greek Greek may refer to: Anything of, from, or related to Greece, a country in Southern Europe: *Greeks, an ethnic group *Greek language, a branch of the Indo-European language family **Proto-Greek language, the assumed last common ancestor of all kno ...

''souchos'', an Egyptian crocodile god. 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 ...

''yajabalijekundu'' is derived from the

Swahili language Swahili, also known as as it is referred to endonym and exonym, in the Swahili language, is a Bantu languages, Bantu language originally spoken by the Swahili people, who are found primarily in Tanzania, Kenya, and Mozambique (along the East Af ...

meaning "of/from the red outcrop", in reference to the Red Sandstone Group deposits exposed at the basin.

Description

The skull of ''Rukwasuchus'' possesses several

autapomorphies In phylogenetics, an autapomorphy is a distinctive feature, known as a derived trait, that is unique to a given taxon. That is, it is found only in one taxon, but not found in any others or outgroup taxa, not even those most closely related to ...

which distinguished it from other

s. These include the presence of a mediolaterally narrow, elongate, and septate internal narial fenestra located anteriorly on the

pterygoid Pterygoid, from the Greek for 'winglike', may refer to: * Pterygoid bone, a bone of the palate of many vertebrates * Pterygoid processes of the sphenoid bone ** Lateral pterygoid plate ** Medial pterygoid plate * Lateral pterygoid muscle * Medial ...

, the presence of a markedly depressed posterior border of the parietal that excludes the supraoccipital from the dorsal cranial table, and a ventrally directed descending process of the postorbital with a well-developed posteroventral process. The morphology of the skull and isolated teeth suggests close relations to the

Hamadasuchus rebouli ''Hamadasuchus'' is an extinct genus of sebecian Crocodylomorpha, crocodylomorph. Fossils have been found from the Kem Kem Formation outcropping in southeastern Morocco. These beds date back to the Albian and Cenomanian stages of the Late Cretace ...

'' from the middle Cretaceous

Kem Kem Beds The Kem Kem Group (commonly known as the Kem Kem beds) is a geological group in the Kem Kem region of eastern Morocco, whose strata date back to the Cenomanian stage of the Late Cretaceous. Its strata are subdivided into two geological formations ...

Morocco Morocco, officially the Kingdom of Morocco, is a country in the Maghreb region of North Africa. It has coastlines on the Mediterranean Sea to the north and the Atlantic Ocean to the west, and has land borders with Algeria to Algeria–Morocc ...

. ''Rukwasuchus'' is the only known sub-

Sahara The Sahara (, ) is a desert spanning across North Africa. With an area of , it is the largest hot desert in the world and the list of deserts by area, third-largest desert overall, smaller only than the deserts of Antarctica and the northern Ar ...

from Africa, thus representing the only link between middle Cretaceous southern vertebrate faunas and much more abundant and taxonomically diverse faunas from northern Africa.

Phylogeny

phylogenetic In biology, phylogenetics () is the study of the evolutionary history of life using observable characteristics of organisms (or genes), which is known as phylogenetic inference. It infers the relationship among organisms based on empirical dat ...

analysis of

Crocodyliformes 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 pseu ...

supports a close relationship between ''Rukwasuchus'' and other African members of

Trematochampsidae Trematochampsidae is an extinct family of mesoeucrocodylian crocodyliforms. Fossils are present from Madagascar, Morocco, Niger, Argentina, and Brazil (in the case of '' Caririsuchus'', where some specimens have been found in the Romualdo Membe ...

, e.g. ''Hamadasuchus'' and ''

Trematochampsa ''Trematochampsa'' is a dubious extinct genus of crocodyliform from the Late Cretaceous (Coniacian-Santonian age) In Beceten Formation of Niger. Taxonomy The type species, ''T. taqueti'', was described by Eric Buffetaut in 1974. A second specie ...

'', that is positioned within a larger clade of

Gondwana Gondwana ( ; ) was a large landmass, sometimes referred to as a supercontinent. The remnants of Gondwana make up around two-thirds of today's continental area, including South America, Africa, Antarctica, Australia (continent), Australia, Zea ...

s. The following

cladogram A cladogram (from Greek language, Greek ''clados'' "branch" and ''gramma'' "character") is a diagram used in cladistics to show relations among organisms. A cladogram is not, however, an Phylogenetic tree, evolutionary tree because it does not s ...

is simplified after the analysis of Sertich & O’Connor (2014); the relationships within

Sebecosuchia Sebecosuchia (meaning "Sobek crocodiles") is an extinct group of mesoeucrocodylian crocodyliforms that includes the families Sebecidae and Baurusuchidae. The group was long thought to have first appeared in the Late Cretaceous with the baurusu ...

and advanced

notosuchia Notosuchia is a clade of primarily Gondwanan mesoeucrocodylian Crocodylomorpha, crocodylomorphs that lived during the Jurassic and Cretaceous. Some phylogeny, phylogenies recover Sebecosuchia as a clade within Notosuchia, others as a sister group ...

ns are not shown.

References

{{Taxonbar, from=Q16992356 Peirosauridae Cretaceous crocodylomorphs of Africa Fossils of Tanzania Fossil taxa described in 2014 Prehistoric pseudosuchian genera