Kaprosuchus
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''Kaprosuchus'' is an extinct
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 ...
of mahajangasuchid
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 ...
. It is known from a single nearly complete skull collected from the
Upper Cretaceous The Late Cretaceous (100.5–66 Ma) is the more recent 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 ''cret ...
Echkar Formation The Echkar Formation is a geological Formation (geology), formation comprising sandstones and claystones in the Agadez Region of Niger, central Africa. Description Its strata date back to the Albian, Late Albian to Late Cretaceous (Cenomanian s ...
of
Niger Niger, officially the Republic of the Niger, is a landlocked country in West Africa. It is a unitary state Geography of Niger#Political geography, bordered by Libya to the Libya–Niger border, north-east, Chad to the Chad–Niger border, east ...
. The name means "boar crocodile" 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 ...
, ''kapros'' ("boar") and , ''soukhos'' ("crocodile") in reference to its unusually large caniniform teeth which resemble those of a
boar The wild boar (''Sus scrofa''), also known as the wild swine, common wild pig, Eurasian wild pig, or simply wild pig, is a Suidae, suid native to much of Eurasia and North Africa, and has been introduced to the Americas and Oceania. The speci ...
. It has been nicknamed "BoarCroc" by
Paul Sereno Paul Callistus Sereno (born October 11, 1957) is a professor of paleontology at the University of Chicago who has discovered several new dinosaur species on several continents, including at sites in Inner Mongolia, Argentina, Morocco and Niger. ...
and Hans Larsson, who first described the genus in a monograph published in ''
ZooKeys ''ZooKeys'' is a peer-reviewed open access scientific journal covering zoological taxonomy, phylogeny, and biogeography. It was established in 2008 and the founding editor-in-chief was Terry Erwin (Smithsonian Institution The Smithsonian ...
'' in 2009 along with other Saharan crocodyliformes such as ''
Anatosuchus ''Anatosuchus'' ("duck crocodile", the name from the Latin ''anas'' ("duck") and the Greek ''souchos'' ("crocodile"), for the broad, duck-like snout) is an extinct genus of notosuchian crocodyliforms discovered in Gadoufaoua, Niger, and described ...
'' and ''
Laganosuchus ''Laganosuchus'' is an extinct genus of stomatosuchid crocodyliform. Fossils have been found from Niger and Morocco and date back to the Upper Cretaceous. Discovery The name means "pancake crocodile" from the Greek , ("pancake") and , ("croc ...
''. 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 ''K. saharicus''.


Description

''Kaprosuchus'' is known from a nearly complete skull about 507 mm in length in which the lower jaw measured 603 mm long. Its total body length is estimated to be around . It possesses three sets of tusk-like
caniniform In mammalian oral anatomy, the canine teeth, also called cuspids, dogteeth, eye teeth, vampire teeth, or fangs, are the relatively long, pointed tooth, teeth. In the context of the upper jaw, they are also known as ''fangs''. They can appear mo ...
teeth that project above and below the skull, one of which in the lower jaw fits into notches in the upper jaw. This type of dentition is not seen in any other known crocodyliform. Another unique characteristic of ''Kaprosuchus'' is the presence of large, rugose horns formed from the
squamosal The squamosal is a skull bone found in most reptiles, amphibians, and birds. In fishes, it is also called the pterotic bone. In most tetrapods, the squamosal and quadratojugal bones form the cheek series of the skull. The bone forms an ancestra ...
and
parietal bone The parietal bones ( ) are two bones in the skull which, when joined at a fibrous joint known as a cranial suture, form the sides and roof of the neurocranium. In humans, each bone is roughly quadrilateral in form, and has two surfaces, four bord ...
s that project posteriorly from the skull. Smaller projections are also seen in the closely related ''
Mahajangasuchus ''Mahajangasuchus'' is an extinct genus of crocodyliform which had blunt, laterally compressed and serrated teeth. The type species, ''M. insignis'', lived during the Late Cretaceous; its fossils have been found in the Maevarano Formation in nort ...
''. The snout of ''Kaprosuchus'' shows generalized proportions and the
naris A nostril (or naris , : 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 turbinates, ...
is positioned dorsally. In ''Kaprosuchus'' many teeth are hypertrophied and labiolingually (laterally) compressed, unlike those of other crocodyliforms with similarly shallow snouts, which are usually subconical and of moderate length. Another difference between the skull of ''Kaprosuchus'' and those of other crocodyliforms that also possess dorsoventrally compressed snouts is the great depth of the posterior portion of the skull. In ''Kaprosuchus'', the
orbits In celestial mechanics, an orbit (also known as orbital revolution) is the curved trajectory of an physical body, object such as the trajectory of a planet around a star, or of a natural satellite around a planet, or of an satellite, artificia ...
(i.e., eye sockets) open laterally and are angled slightly forward rather than upward. The orbits turned forward suggest that there was somewhat
stereoscopic vision Binocular vision is seeing with two eyes, which increases the size of the visual field. If the visual fields of the two eyes overlap, binocular depth can be seen. This allows objects to be recognized more quickly, camouflage to be detected, spa ...
, i.e., an overlap in the visual field of the animal.National Geographic. November 2009. p. 140-141 The surfaces of 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 mammals h ...
e are rugose with the edges elevated above the body of the bone, suggesting that a
keratinous Keratin () is one of a family of structural fibrous proteins also known as ''scleroproteins''. It is the key structural material making up Scale (anatomy), scales, hair, Nail (anatomy), nails, feathers, horn (anatomy), horns, claws, Hoof, hoove ...
shield would have been supported by the rugosities at the tip of the snout. Along the interpremaxillary suture, the area where the two premaxillae meet, the surface is smooth, giving the paired rugosity of the premaxillae the resemblance of a moustache in anterior view.


Classification

''Kaprosuchus'' is a member of the
family Family (from ) is a Social group, group of people related either by consanguinity (by recognized birth) or Affinity (law), affinity (by marriage or other relationship). It forms the basis for social order. Ideally, families offer predictabili ...
Mahajangasuchidae along with closely related ''Mahajangasuchus insignis'' from the Upper Cretaceous of
Madagascar Madagascar, officially the Republic of Madagascar, is an island country that includes the island of Madagascar and numerous smaller peripheral islands. Lying off the southeastern coast of Africa, it is the world's List of islands by area, f ...
. Although it differs greatly from any other known crocodyliform, ''Kaprosuchus'' shares several characteristics with ''Mahajangasuchus''. These include the obliteration of all but the posterior portion of the
internasal suture 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 ...
; a laterally facing rugose external articular fossa; the positioning of the jaw joint below the posterior
maxilla In vertebrates, the maxilla (: maxillae ) 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 two maxil ...
ry teeth; a deep, anterodorsally oriented
mandibular symphysis In human anatomy, the facial skeleton of the skull the external surface of the mandible is marked in the median line by a faint ridge, indicating the mandibular symphysis (Latin: ''symphysis menti'') or line of junction where the two lateral ha ...
; a vertically descending ectopterygoid that is slightly inset from the lateral margin of the
jugal The jugal is a skull bone found in most reptiles, amphibians and birds. In mammals, the jugal is often called the malar or zygomatic. It is connected to the quadratojugal and maxilla, as well as other bones, which may vary by species. Anatomy ...
; a flared
choana The choanae (: choana), posterior nasal apertures or internal nostrils are two openings found at the back of the nasal passage between the nasal cavity and the pharynx, in humans and other mammals (as well as crocodilians and most skinks). They ...
l septum forming an articular foot for the
palatine A palatine or palatinus (Latin; : ''palatini''; cf. derivative spellings below) is a high-level official attached to imperial or royal courts in Europe since Roman Empire, Roman times.
; and the hornlike dorsal projection of the external rim of the squamosal (although this is much more developed in ''Kaprosuchus'' than ''Mahajangasuchus''). At the time of ''Kaprosuchus description, Sereno and Larrson considered mahajangasuchids to be a family of
Neosuchian Neosuchia is a clade within Mesoeucrocodylia that includes all modern extant crocodilians and their closest fossil relatives. It is defined as the most inclusive clade containing all crocodylomorphs more closely related to ''Crocodylus niloticus' ...
crocodyliforms. However further studies on the relationship of this family have repeatedly found them to form a sister clade to
peirosaurids Peirosauridae is a Gondwanan family of mesoeucrocodylians that lived during the Cretaceous period (geology), period. It was a clade of terrestrial crocodyliforms that evolved a rather dog-like skull, and were terrestrial carnivores. It was phylog ...
, forming a clade that in turn groups together with
uruguaysuchids Uruguaysuchidae is a family (biology), family of notosuchian crocodyliforms that lived in South America and Africa during the Cretaceous period. It was formally defined under the PhyloCode in 2024 as "the most inclusive clade containing ''Uruguay ...
such as ''Anatosuchus'' and ''Araripesuchus'' as an early diverging branch of
notosuchians 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 ...
.


Paleobiology

''Kaprosuchus'' was once thought to have been a primarily if not exclusively terrestrial predator. Evidence for this behavior includes the positioning of the orbits laterally and somewhat anteriorly, which suggests an overlap in vision. This is unlike many other neosuchians, including extant
crocodilian Crocodilia () is an Order (biology), order of semiaquatic, predatory reptiles that are known as crocodilians. They first appeared during the Late Cretaceous and are the closest living relatives of birds. Crocodilians are a type of crocodylomorp ...
s, in which the orbits are positioned dorsally as an adaptation to aquatic predation where the head can be held underwater while the eyes remain above the surface. Additional support for terrestrial predation can be found in the teeth and jaws. The enlarged caniniforms are sharp-edged and relatively straight, unlike the fluted, subconical, recurved teeth of aquatic crocodyliforms. Because the retroarticular process of the lower jaw is long, it is likely that the jaws were able to open relatively quickly with a large gape to allow for the opposing caniniforms to clear one another. The fused nasal bones are thought to have provided reinforcement for the jaws against compression associated with a powerful bite. The telescoped, dorsally positioned external nares are seen as protection against impact if the animal rammed prey with its robust snout. The keratinous shield thought to have covered the tip of the snout would have provided further protection. However, ''Kaprosuchus'' is now thought to have been semiaquatic, as the relative ''Mahajangasuchus'' has been suggested to be a primarily aquatic predator, and that the specimens of both genera show cranial adaptations usually found in "definitively semiaquatic" crocodylomorphs, "such as elongate platyrostral or tube-like snouts, orbits located dorsally on the skull, and/or dorsally-facing external nares."


References


External links


''Kaprosuchus''
in the
Paleobiology Database The Paleobiology Database (PBDB) is an online resource for information on the distribution and classification of fossil animals, plants, and microorganisms. History The Paleobiology Database originated in the NCEAS-funded Phanerozoic Marine Pale ...
{{Taxonbar, from=Q137176 Terrestrial crocodylomorphs Late Cretaceous crocodylomorphs of Africa Fossil taxa described in 2009 Notosuchia Prehistoric pseudosuchian genera Cenomanian genera