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''Antaeusuchus taouzensis'' is a species of
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 form, and were terrestrial carnivores. It was phylogenetically defined ...
notosuchian Notosuchia is a suborder of primarily Gondwanan mesoeucrocodylian crocodylomorphs that lived during the Jurassic and Cretaceous. Some phylogenies recover Sebecosuchia as a clade within Notosuchia, others as a sister group (see below); if Sebecosu ...
from the Late
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 ...
Kem Kem Group 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, ...
of
Morocco Morocco (),, ) officially the Kingdom of Morocco, is the westernmost country in the Maghreb region of North Africa. It overlooks the Mediterranean Sea to the north and the Atlantic Ocean to the west, and has land borders with Algeria to ...
. It was described in 2021, and it is the only species in the genus ''Antaeusuchus''. It is the fourth
notosuchian Notosuchia is a suborder of primarily Gondwanan mesoeucrocodylian crocodylomorphs that lived during the Jurassic and Cretaceous. Some phylogenies recover Sebecosuchia as a clade within Notosuchia, others as a sister group (see below); if Sebecosu ...
described from the region and the second Kem Kem peirosaurid after ''
Hamadasuchus ''Hamadasuchus'' is an extinct genus of sebecian 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 Cretaceous. It was first ...
''.


Discovery and naming

''Antaeusuchus'' is known from two specimens: the
holotype A holotype is a single physical example (or illustration) of an organism, known to have been used when the species (or lower-ranked taxon) was formally described. It is either the single such physical example (or illustration) or one of several ...
NHMUK PV R36829, consisting of paired mandibles, and the
paratype In zoology and botany, a paratype is a specimen of an organism that helps define what the scientific name of a species and other taxon actually represents, but it is not the holotype (and in botany is also neither an isotype nor a syntype). Of ...
NHMUK PV R36874, a partial right mandible. Both specimens were commercially collected and recovered from unspecified beds of the Kem Kem Group near
Jebel Beg'aa Jabal, Jabel, Jebel or Jibal may refer to: People * Jabal (name), a male Arabic given name * Jabal (Bible), mentioned in the Hebrew Bible Places In Arabic, ''jabal'' or ''jebel'' (spelling variants of the same word) means 'mountain'. * Dzhebel, ...
, Taouz township in the Errachidia Province of Morocco. Both fossils are preserved without distortion and in good condition, with damage mostly restricted to the teeth. The generic name derives from
Antaeus Antaeus (; Ancient Greek: Ἀνταῖος ''Antaîos'', "opponent", derived from , ''antao'' – 'I face, I oppose'), known to the Berbers as Anti, was a figure in Berber and Greek mythology. He was famed for his defeat by Heracles as part o ...
, a giant from Greek and Berber mythology said to be buried in northern Morocco and the Ancient Greek σοῦχος, soukhos meaning crocodile. The species name refers to the township Taouz where both specimens had been found.


Description

The anterior region of the
mandible In anatomy, the mandible, lower jaw or jawbone is the largest, strongest and lowest bone in the human facial skeleton. It forms the lower jaw and holds the lower tooth, teeth in place. The mandible sits beneath the maxilla. It is the only movabl ...
is characterized by a broad ’U’-shaped symphysis. Each mandibular ramus diverges at an 22° angle from the midline and remains mostly straight for most its preserved length, curving slightly medially towards the fossil's posterior. The anterior of the dorsal margin of the mandible is characterized by two 'waves', the first of which spanning the
dentary In anatomy, the mandible, lower jaw or jawbone is the largest, strongest and lowest bone in the human facial skeleton. It forms the lower jaw and holds the lower tooth, teeth in place. The mandible sits beneath the maxilla. It is the only movabl ...
teeth 1 to 6. The second wave rises from the 9th to 15th tooth. These 'waves' reach their peak at the 4th and 13th dentary tooth respectively. Other than a series of foramina adjacent to the tooth row, this part of the bone shows little ornamentation. The holotype dentary preserves 18
alveoli Alveolus (; pl. alveoli, adj. alveolar) is a general anatomical term for a concave cavity or pit. Uses in anatomy and zoology * Pulmonary alveolus, an air sac in the lungs ** Alveolar cell or pneumocyte ** Alveolar duct ** Alveolar macrophage * ...
, the first 16 of which preserving partial or complete teeth, all of which are closely spaced. The posterior half of the dorsal margin remains largely straight and is strongly sloping. The mandibular fenestra is not fully preserved in either specimen, but can be inferred to be large in size and anteroventrally-posterodorsally elongated. The first alveoli are circular in shape, suggesting the anterior teeth to be roughly conical while from the 10th dentary alveoli onward become more labiolingually compressed, more extreme flattening is present at the anterior and posterior margin of the preserved teeth. These margins are adorned with denticulated
carinae ''Carinae'' was an area of ancient Rome. It was one of its most exclusive neighborhoods, where many of the senatorial class lived. Florus described the ''Carinae'' as the "''most celebrated part of the city''" (''celeberrima pars urbis''). Des ...
which form the cutting edge of the teeth. These denticles are small in size and show no significant variation in size. In ''Antaeusuchus,'' the mandibular rami diverge at a much broader angle (43 to 44°) than in ''Hamadasuchus'' (between 20 and 30° depending on the specimen), meaning that it had a broader rostrum than its relative. The surface of the mandible in ''Antaeusuchus'' is covered in shallow pits and groves different from the deeper ornamentation of ''Hamadasuchus''. The posterior teeth of the ''Hamadasuchus'' holotype are
lanceolate The following is a list of terms which are used to describe leaf morphology in the description and taxonomy of plants. Leaves may be simple (a single leaf blade or lamina) or compound (with several leaflets). The edge of the leaf may be regular o ...
in shape, which is not seen in ''Antaeusuchus''. Furthermore the teeth of the ''Antaeusuchus'' holotype, while slightly pointed at their
apices The apex is the highest point of something. The word may also refer to: Arts and media Fictional entities * Apex (comics), a teenaged super villainess in the Marvel Universe * Ape-X, a super-intelligent ape in the Squadron Supreme universe *Apex ...
, they are not comparable to the almost triangular teeth seen in the holotype of ''Hamadasuchus''. The 10th to 14th teeth in referred specimen NHMUK PV R36874 are angular, but their anterior and posterior margin run parallel for most of their length and only converge to a point at the apex of the crown. ''Antaeusuchus'' was significantly larger than ''Hamadasuchus'', almost twice the size of the later. While size is not a distinguishing character on its own, it adds to the already present morphological differences between the two genera. Furthermore, Nicholl ''et al.'' argue that the features present in ''Antaeusuchus'' are not compatible with the
ontogenetic Ontogeny (also ontogenesis) is the origination and development of an organism (both physical and psychological, e.g., moral development), usually from the time of fertilization of the egg to adult. The term can also be used to refer to the st ...
series known for ''Hamadasuchus'', which already includes specimen presumed to be 'adult'.


Phylogeny

For the phylogenetic analysis both specimens were combined as one
operational taxonomic unit An Operational Taxonomic Unit (OTU) is an operational definition used to classify groups of closely related individuals. The term was originally introduced in 1963 by Robert R. Sokal and Peter H. A. Sneath in the context of numerical taxonomy, w ...
and inserted into a character-taxon matrix based on that of Pol ''et al.'' 2014. This matrix was subsequently used as a basis for multiple further studies running parallel to one another, resulting in numerous 'daughter' matrices, which were united for the phylogeny of Nicholl ''et al.'' 2021. The sampling was also expanded by the inclusion of the putative peirosaurids ''
Rukwasuchus yajabalijekundu ''Rukwasuchus'' is an extinct genus of peirosaurid mesoeucrocodylian known from the middle Cretaceous Galula Formation of southwestern Tanzania. It contains a single species, ''Rukwasuchus yajabalijekundu''. Discovery ''Rukwasuchus'' is known fr ...
'' from Tanzania and ''
Miadanasuchus oblita ''Miadanasuchus'' is an extinct genus of peirosaurid which existed in the Maevarano Formation of Madagascar during the late Cretaceous period (Campanian The Campanian is the fifth of six ages of the Late Cretaceous Epoch on the geologic t ...
'' from Madagascar. The resulting strict consensus tree under equal weighting of characters can be seen below: The resulting phylogeny recovered
peirosaurids 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 form, and were terrestrial carnivores. It was phylogenetically defined ...
and mahajangasuchids grouping together, forming a sister clade to
uruguaysuchids Uruguaysuchidae is a family of notosuchian crocodyliforms that lived in South America and Africa during the Cretaceous period. It includes the genera ''Araripesuchus ''Araripesuchus'' is a genus of extinct crocodyliform that existed during the ...
. Internally, ''Antaeusuchus'' was recovered as a sister taxon to ''Hamadasuchus'' from the Kem Kem Group, the two taxa clade together with ''
Bayomesasuchus ''Bayomesasuchus'' is an extinct genus of peirosaurid mesoeucrocodylian known from the early Late Cretaceous Cerro Lisandro Formation of Neuquén Province, western central Argentina. It contains a single species, ''Bayomesasuchus hernandezi''. Ev ...
'' as their sister, which in turn forms a sister clade to all other peirosaurids. ''
Peirosaurus torminni ''Peirosaurus'' is an extinct genus of peirosaurid crocodylomorph known from the Late Cretaceous period (late Maastrichtian stage) of Minas Gerais, southern Brazil. It contains a single species, ''Peirosaurus torminni''. It is the type genus of ...
'' was not included in the analysis, however its close relative ''
Uberabasuchus terrificus ''Uberabasuchus'' ("Uberaba crocodile")''Uberabasuchus''
at Fossil ...
'' has been consistently recovered as a close relative or possible
junior synonym The Botanical and Zoological Codes of nomenclature treat the concept of synonymy differently. * In botanical nomenclature, a synonym is a scientific name that applies to a taxon that (now) goes by a different scientific name. For example, Linna ...
, consequently ''
Uberabasuchus ''Uberabasuchus'' (" Uberaba crocodile")''Uberabasuchus''
at
'' was used to serve as a proxy for ''
Peirosaurus ''Peirosaurus'' is an extinct genus of peirosaurid crocodylomorph known from the Late Cretaceous period (late Maastrichtian stage) of Minas Gerais, southern Brazil. It contains a single species, ''Peirosaurus torminni''. It is the type genus of ...
''.


References

{{Taxonbar, from= Q109252550, from2= Q109252548 Late Cretaceous crocodylomorphs of Africa Terrestrial crocodylomorphs Peirosaurids Prehistoric pseudosuchian genera Fossil taxa described in 2021 Cenomanian genera