Sahitisuchus
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''Sahitisuchus'' 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
sebecid Sebecidae is an extinct family of prehistoric terrestrial sebecosuchian crocodylomorphs. The oldest known member of the group is ''Ogresuchus furatus'' known from the Upper Cretaceous (Maastrichtian) Tremp Formation (Spain). Sebecids were di ...
mesoeucrocodylia Mesoeucrocodylia is the clade that includes Eusuchia and crocodyliforms formerly placed in the paraphyletic group Mesosuchia. The group appeared during the Early Jurassic, and continues to the present day. Diagnosis It was long known that M ...
n known from
Rio de Janeiro State Rio de Janeiro () is one of the 27 federative units of Brazil. It has the second largest economy of Brazil, with the largest being that of the state of São Paulo. The state, which has 8.2% of the Brazilian population, is responsible for 9.2% of ...
of southeastern
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 ...
.''Sahitisuchus''
at
Fossilworks Fossilworks is a portal which provides query, download, and analysis tools to facilitate access to the Paleobiology Database The Paleobiology Database is an online resource for information on the distribution and classification of fossil animals ...
.org
It contains a single species, ''Sahitisuchus fluminensis''. It is a terrestrial sebecid, however also adopted to a semi-aquatic lifestyle to some degree, most probably coexisting with the semi-aquatic
alligatorid The family (biology), family Alligatoridae of crocodylians includes alligators, caimans and their extinct relatives. Phylogeny The superfamily Alligatoroidea includes all crocodilians (fossil and extant) that are more closely related to the Am ...
''
Eocaiman itaboraiensis ''Eocaiman'' is an extinct genus of caiman containing species living from the Early Paleocene to Miocene in what is now Argentina (Salamanca and Sarmiento Formations), Itaboraí Formation of Brazil and Colombia ( Honda Group).
''.


Discovery

''Sahitisuchus'' was named by Alexander W. A. Kellner, André E. P. Pinheiro and Diogenes A. Campos in
2014 File:2014 Events Collage.png, From top left, clockwise: Stocking up supplies and personal protective equipment (PPE) for the Western African Ebola virus epidemic; Citizens examining the ruins after the Chibok schoolgirls kidnapping; Bundles of wat ...
and 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 ''Sahitisuchus fluminensis''. The generic name honors the
Xavante people The Xavante (also Shavante, Chavante, Akuen, A'uwe, Akwe, Awen, or Akwen) are an indigenous people, comprising 15,315 individuals within the territory of eastern Mato Grosso state in Brazil. They speak the Xavante language, part of the Jê langu ...
, one of the indigenous Brazilian inhabitants. It is derived from ''sahi ti'', meaning "to be angry" or "to be brave" in
Xavante language The Xavante language is an Akuwẽ (Central Jê) language ( Jê, Macro-Jê) spoken by the Xavante people in the area surrounding Eastern Mato Grosso, Brazil. The Xavante language is unusual in its phonology, its ergative object–agent–verb ...
, alluding at
warrior A warrior is a person specializing in combat or warfare, especially within the context of a tribal or clan-based warrior culture society that recognizes a separate warrior aristocracies, class, or caste. History Warriors seem to have been p ...
s, and ''suchus'',
Latin Latin (, or , ) is a classical language belonging to the Italic branch of the Indo-European languages. Latin was originally a dialect spoken in the lower Tiber area (then known as Latium) around present-day Rome, but through the power of the ...
ized from the
Greek Greek may refer to: Greece 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 ...
''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 ...
, ''fluminensis'', is a latinization of fluminense, a designation of citizens born in the
Rio de Janeiro State Rio de Janeiro () is one of the 27 federative units of Brazil. It has the second largest economy of Brazil, with the largest being that of the state of São Paulo. The state, which has 8.2% of the Brazilian population, is responsible for 9.2% of ...
. ''Sahitisuchus'' is known solely from 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 ...
MCT 1730-R, a nearly complete
skull The skull is a bone protective cavity for the brain. The skull is composed of four types of bone i.e., cranial bones, facial bones, ear ossicles and hyoid bone. However two parts are more prominent: the cranium and the mandible. In humans, the ...
and
lower jaw 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 teeth in place. The mandible sits beneath the maxilla. It is the only movable bone ...
, proatlas, an intercentrum, the axis and the third
cervical vertebra In tetrapods, cervical vertebrae (singular: vertebra) are the vertebrae of the neck, immediately below the skull. Truncal vertebrae (divided into thoracic and lumbar vertebrae in mammals) lie caudal (toward the tail) of cervical vertebrae. In sau ...
, housed at the
Museu de Ciências da Terra The Earth Sciences Museum (''Museu de Ciências da Terra'') is a geological museum in Rio de Janeiro, Brazil Brazil ( pt, Brasil; ), officially the Federative Republic of Brazil (Portuguese: ), is the largest country in both South Americ ...
, Companhia de Pesquisas de Recursos Minerais, in Rio de Janeiro, Brazil. A cast of the holotype, MN 4711-V, is housed at the
National Museum of Brazil The National Museum of Brazil ( pt, Museu Nacional) is the oldest scientific institution of Brazil. It is located in the city of Rio de Janeiro, where it is installed in the Paço de São Cristóvão (Saint Christopher's Palace), which is in ...
. MCT 1730-R was discovered during the exploration of the São José de Itaboraí Basin, located in Rio de Janeiro State, that lasted for about five decades and ended in 1984. It was collected from São José Farm, of São José de Itaboraí Municipal District, in the S2 sequence of the
Itaboraí Formation The Itaboraí Formation ( pt, Formação Itaboraí) is a highly list of fossiliferous stratigraphic units in Brazil, fossiliferous geologic Formation (geology), formation and LagerstätteKellner & Campos, 1999, p.399 of the Itaboraí Basin in Rio ...
in the
Itaboraí Basin Itaboraí (, ) is a city in the state of Rio de Janeiro in Brazil, that belongs to the Rio de Janeiro metropolitan area. It was founded in 1672. In 2020, it had a population of 242,543. Location Culturally, its closest municipalities are São ...
, dating to the late
Itaboraian The Itaboraian ( pt, Itaboraiense) age is a period within the Early Eocene geologic time (53.0–50.0 Mya (unit), Ma) epoch of the Paleogene, used more specifically with South American land mammal ages (SALMA). It follows the Riochican and precedes ...
South American land mammals age of the
Early Eocene In the geologic timescale the Ypresian is the oldest age or lowest stratigraphic stage of the Eocene. It spans the time between , is preceded by the Thanetian Age (part of the Paleocene) and is followed by the Eocene Lutetian Age. The Ypresian i ...
, about 53-50
million years ago The abbreviation Myr, "million years", is a unit of a quantity of (i.e. ) years, or 31.556926 teraseconds. Usage Myr (million years) is in common use in fields such as Earth science and cosmology. Myr is also used with Mya (million years ago). ...
. MCT 1730-R was briefly mentioned in the literature by Price and Paula-Couto (1946) and later in 1991 and 1993, but had never been figured or described before Kellner ''et al.'' (2014) assigned it to a new genus and species. Apart from ''Sahitisuchus fluminensis'', ''
Eocaiman itaboraiensis ''Eocaiman'' is an extinct genus of caiman containing species living from the Early Paleocene to Miocene in what is now Argentina (Salamanca and Sarmiento Formations), Itaboraí Formation of Brazil and Colombia ( Honda Group).
'' is the only other species formally described based on remains collected at São José de Itaboraí Basin. To date, this is the only Paleocene deposit where the
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 ...
fauna was composed by rather primitive (''Sahitisuchus'', a
sebecosuchia Sebecosuchia 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 baurusuchids and become extinct in the ...
n) and more derived (
Alligatoridae The family Alligatoridae of crocodylians includes alligators, caimans and their extinct relatives. Phylogeny The superfamily Alligatoroidea includes all crocodilians (fossil and extant) that are more closely related to the American alligator t ...
) post- K-Pg taxa. ''Sahitisuchus'' and ''E. itaboraiensis'' were either set apart for a comparatively short geological timespan or, most likely, co-occurred.


Description

The holotype of ''Sahitisuchus'' is well preserved, although during preservation the skull became slightly compressed dorsoventrally. This effect is mainly observable on some elements of the rear portion of the skull, particularly the
supraoccipital The occipital bone () is a cranial dermal bone and the main bone of the occiput (back and lower part of the skull). It is trapezoidal in shape and curved on itself like a shallow dish. The occipital bone overlies the occipital lobes of the cereb ...
that is displaced towards the
foramen In anatomy and osteology, a foramen (;Entry "foramen"
in
magnum. The tip of the snout and 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 ...
e are broken off. ''Sahitisuchus'' possesses the two following
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 t ...
among
sebecid Sebecidae is an extinct family of prehistoric terrestrial sebecosuchian crocodylomorphs. The oldest known member of the group is ''Ogresuchus furatus'' known from the Upper Cretaceous (Maastrichtian) Tremp Formation (Spain). Sebecids were di ...
s: the odontoid process is fused to the axis with vertical anterior surface, lacking medial processes, and its mandible lacks an external mandibular fenestra. ''Sahitisuchus'' can be also distinguished from other sebecids by a combination of characters. As in ''
Lorosuchus ''Lorosuchus'' is an extinct genus of sebecid mesoeucrocodylian known from the Río Loro Formation in Tucumán Province of northwestern Argentina. Discovery ''Lorosuchus'' is known from the holotype PVL 6219, a nearly complete skull fou ...
'', there is a shallow ventrolateral depression in its infraorbital
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 ...
region. A shallow elliptical depression is present near the cranio-mandibular articulation on the posterior surface of the quadrate, and the dorsal edge of supratemporal
fossa Fossa may refer to: Animals * Fossa (animal), the common name of a carnivoran mammal of genus ''Cryptoprocta'' endemic to Madagascar * ''Fossa'', the Latin genus name of the Malagasy civet, a related but smaller mammal endemic to Madagascar Pla ...
is rough and rugose, as seen in ''
Sebecus ''Sebecus'' (meaning " Sebek" in Latin) is an extinct genus of sebecid crocodylomorph from Eocene of South America. Like other sebecosuchians, it was entirely terrestrial and carnivorous. The genus is currently represented by two species, the ty ...
''. The exoccipital posterior processes are sharp, half moon-shaped and directed medially, as in ''
Ayllusuchus ''Ayllusuchus'' is an extinct genus of sebecid mesoeucrocodylian. Fossils have been found in the Lumbrera Formation of Argentina (Eocene age, Casamayoran).Bretesuchus ''Bretesuchus'' is an extinct genus of sebecosuchian mesoeucrocodylian within the family Sebecidae known from northwestern Argentina.
'', its rear ramus is higher than the anterior process and expands laterally. As in ''Bretesuchus'' and ''Sebecus'', it has a rough longitudinal ridge on the lateroventral edge of
angular bone The angular is a large bone in the lower jaw (mandible) of amphibians and reptiles (birds included), which is connected to all other lower jaw bones: the dentary (which is the entire lower jaw in mammals), the splenial, the suprangular, and the art ...
and
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 ...
, that ends near the mandibular symphysis level. The frontal bone is broad and triangular and ornamented similarly to 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 ...
. As in many basal mesoeucrocodylian taxa, which includes some sebecosuchians like ''Lorosuchus'', ''Sebecus'', ''
Zulmasuchus ''Zulmasuchus'' (meaning "Zulma Gasparini's crocodile") is an extinct genus of sebecid sebecosuchian mesoeucrocodylian. Its fossils have been found in Early Paleocene-age rocks (Danian stage) of the Santa Lucía Formation in Bolivia. ''Zulmasuch ...
'', ''
Iberosuchus ''Iberosuchus'' (meaning "Iberian crocodile") is a genus of extinct sebecosuchian mesoeucrocodylian found in Western Europe from the Eocene. Remains from Portugal was described in 1975 by Antunes as a sebecosuchian crocodilian. This genus has one ...
'' and ''
Pepesuchus ''Pepesuchus'' is an extinct genus of carnivorous metasuchian from the Late Cretaceous period. It is a peirosaurid which lived during the Campanian and Maastrichtian stages of the Late Cretaceous in what is now state of São Paulo, Brazil.
'', a longitudinal frontal crest is present. In ''Sahitisuchus'', it is low and smooth, running from the middle part to the posterior portion the frontal. As in all sebecid species and some other not closely related taxa, the quadratojugal forms a "double articulation" by participating in the cranio-mandibular articulation. As in the peirosaurids ''
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 ...
'' and ''
Lomasuchus ''Lomasuchus'' is an extinct genus of peirosaurid notosuchian known from the Late Cretaceous (Santonian stage) of Neuquén Province, western central Argentina. It contains a single species, ''Lomasuchus palpebrosus''. Gasparini, Z., Chiappe, L. M ...
'', ''Sahitisuchus'' possesses a
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 The quadratojugal is a skull bone present in many vertebrates, including ...
prong which is directed backwards and does not form a horn. Such sculptured dorsal posteriorly pointed lobe is also present in ''Sebecus'', but with a more squared-shape rear end. The portion of the
tympanic cavity The tympanic cavity is a small cavity surrounding the bones of the middle ear. Within it sit the ossicles, three small bones that transmit vibrations used in the detection of sound. Structure On its lateral surface, it abuts the external auditory ...
formed by the quadrate is not multi-
fenestra A fenestra (fenestration; plural fenestrae or fenestrations) is any small opening or pore, commonly used as a term in the biological sciences. It is the Latin word for "window", and is used in various fields to describe a pore in an anatomical st ...
ted like in
protosuchia Protosuchia is a group of extinct Mesozoic crocodyliforms. They were small in size (~1 meter in length) and terrestrial. In phylogenetic terms, Protosuchia is considered an informal group because it is a grade of basal crocodyliforms, not a true ...
ns,
notosuchia 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 ...
ns and baurusuchids, and lacks the characteristic oblong concavity of baurusuchids. In ''Sahitisuchus'' it shows only two openings, as seen in ''Sebecus'', ''Hamadasuchus'' and recent species. As in ''Sebecus'', the only other sebecid known from this part of the skeleton, th
diapophysis
is divided by the neurocentral suture and in lateral view, the
centrum (Latin for ''center'') may refer to: Places In Greenland * Nuuk Centrum, a district of Nuuk, Greenland * Centrum Lake, Greenland In the Netherlands * Amsterdam-Centrum, the inner-most borough of Amsterdam, Netherlands * Rotterdam Centrum, a borou ...
shows a medial constriction and a
trapezoid A quadrilateral with at least one pair of parallel sides is called a trapezoid () in American and Canadian English. In British and other forms of English, it is called a trapezium (). A trapezoid is necessarily a Convex polygon, convex quadri ...
al shape, with anterior and posterior articulations inclined anteriorly. In comparison to ''Sebecus'', ''Sahitisuchus'' has more robust and broader diapophyses, and the length of the third cervical centrum is nearly equal compared to the axis.


Skull

The skull of ''Sahitisuchus'' is elongated in dorsal view, showing two slight constrictions at about the level of the fifth and eleventh maxillary alveoli, being comparatively shorter than the skull of ''Sebecus'' and ''
Stolokrosuchus ''Stolokrosuchus'' is an extinct genus of neosuchian crocodylomorph that lived during the Early Cretaceous. Its fossils, including a skull with a long thin snout and bony knobs on the prefrontal, have been found in Niger. ''Stolokrosuchus'' wa ...
'', but not as short as ''Lorosuchus''. The snout of ''Sahitisuchus'' is not as tall and domed (oreinirostral) as seen in ''Sebecus'', ''
Barinasuchus ''Barinasuchus'' (meaning " Barinas crocodile," in reference to where the type material was found) is an extinct genus of sebecid mesoeucrocodylian. Its fossils have been found in middle Eocene-age rocks of the Divisadero Largo Formation of Arge ...
'', ''Bretesuchus'', ''Zulmasuchus'' and ''
Langstonia ''Langstonia'' (meaning " rocodileof Langston", in honor of paleontologist Wann Langston, Jr.) is an extinct genus of notosuchian crocodylomorph of the family Sebecidae. It lived in the middle Miocene (specifically in the Laventan land-mammal a ...
''. The
skull roof The skull roof, or the roofing bones of the skull, are a set of bones covering the brain, eyes and nostrils in bony fishes and all land-living vertebrates. The bones are derived from dermal bone and are part of the dermatocranium. In comparati ...
of ''Sahitisuchus'' is flat and rectangular, being wider than long. The supratemporal
fossa Fossa may refer to: Animals * Fossa (animal), the common name of a carnivoran mammal of genus ''Cryptoprocta'' endemic to Madagascar * ''Fossa'', the Latin genus name of the Malagasy civet, a related but smaller mammal endemic to Madagascar Pla ...
is much larger than the supratemporal fenestra and is about three times smaller than the orbits. The dorsal border of this fossa is surrounded by rugosities, forming an elevation that is more developed in the medial and lateral borders, a condition that is similar only to ''Sebecus'' and the
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 ...
''Hamadasuchus'' among
mesoeucrocodylia Mesoeucrocodylia is the clade that includes Eusuchia and crocodyliforms formerly placed in the paraphyletic group Mesosuchia. The group appeared during the Early Jurassic, and continues to the present day. Diagnosis It was long known that M ...
ns. The
eye socket In anatomy, the orbit is the cavity or socket of the skull in which the eye and its appendages are situated. "Orbit" can refer to the bony socket, or it can also be used to imply the contents. In the adult human, the volume of the orbit is , of ...
is placed rather laterally, as in terrestrial
sebecid Sebecidae is an extinct family of prehistoric terrestrial sebecosuchian crocodylomorphs. The oldest known member of the group is ''Ogresuchus furatus'' known from the Upper Cretaceous (Maastrichtian) Tremp Formation (Spain). Sebecids were di ...
s, and not laterodorsally as in semi-aquatic
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 ...
s, like ''Stolokrosuchus'', ''Lorosuchus'' and extant species of
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 ...
. ''Sahitisuchus'' possesses 12
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 ...
ry teeth, similarly to 10 maxillary teeth in other Paleogene sebecosuchians (''Lorosuchus'', ''Bretesuchus'' and ''Zulmasuchus''), as well as blunt posterior teeth, unlike Cretaceous sebecosuchians. Cretaceous sebecosuchians, like ''
Baurusuchus ''Baurusuchus'' is an extinct member of the ancestral crocodilian lineage, which lived in Brazil from 90 to 83.5 million years ago, in the Late Cretaceous period. Technically, it is a genus of baurusuchid mesoeucrocodylian. It was a terres ...
'' and ''
Stratiotosuchus ''Stratiotosuchus'' (from Greek, (stratiōtēs, "soldier") and (suchos, "crocodile")) is an extinct genus of baurusuchid mesoeucrocodylian from the Adamantina Formation in Brazil. It lived during the Late Cretaceous. The first fossils were ...
'' showing only five maxillary teeth, and more recent sebecosuchians (e.g. the Eocene ''Sebecus'' exhibiting nine maxillary teeth, and the Upper Miocene ''Langstonia''), show higher skulls and a specialized, and quite reduced, dentition suggesting that only the less specialized crocodyliforms survived the
Cretaceous–Paleogene extinction event The Cretaceous–Paleogene (K–Pg) extinction event (also known as the Cretaceous–Tertiary extinction) was a sudden mass extinction of three-quarters of the plant and animal species on Earth, approximately 66 million years ago. With the ...
(except for the marine
dyrosaurid Dyrosauridae is a family of extinct neosuchian crocodyliforms that lived from the Late Cretaceous (Maastrichtian) to the Eocene. Dyrosaurid fossils are globally distributed, having been found in Africa, Asia, Europe, North America and South Amer ...
s).


Classification

Below is a
cladogram A cladogram (from Greek ''clados'' "branch" and ''gramma'' "character") is a diagram used in cladistics to show relations among organisms. A cladogram is not, however, an evolutionary tree because it does not show how ancestors are related to d ...
from Kellner ''et al.'' (2014) showing the placement of ''Sahitisuchus'' within
Sebecosuchia Sebecosuchia 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 baurusuchids and become extinct in the ...
.


References

{{Taxonbar, from=Q15630807 Sebecids Eocene crocodylomorphs Eocene reptiles of South America Itaboraian Paleogene Brazil Fossils of Brazil Fossil taxa described in 2014 Taxa named by Alexander Kellner Prehistoric pseudosuchian genera