Aplestosuchus With Prey
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''Aplestosuchus'' 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
baurusuchid Baurusuchidae is a Gondwanan Family (biology), family of mesoeucrocodylians that lived during the Late Cretaceous. It is a group of terrestrial hypercarnivorous crocodilians from South America (Argentina and Brazil) and possibly Pakistan. Baurusu ...
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 the
Late Cretaceous The Late Cretaceous (100.5–66 Ma) is the younger 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 ''creta'', the ...
Adamantina Formation The Adamantina Formation is a geological formation in the Bauru Basin of western São Paulo state, in southeastern Brazil. Its strata date back to the Late Cretaceous epoch of the Cretaceous Period, during the Mesozoic Era. The formation is p ...
of
São Paulo São Paulo (, ; Portuguese for 'Saint Paul') is the most populous city in Brazil, and is the capital of the state of São Paulo, the most populous and wealthiest Brazilian state, located in the country's Southeast Region. Listed by the GaWC a ...
, southern
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 ...
. It contains a single species, ''Aplestosuchus sordidus''. ''A. sordidus'' is represented by a single articulated and nearly complete
skeleton A skeleton is the structural frame that supports the body of an animal. There are several types of skeletons, including the exoskeleton, which is the stable outer shell of an organism, the endoskeleton, which forms the support structure inside ...
, preserving the remains of an unidentified
sphagesaurid Sphagesauridae is a Gondwanan family of mesoeucrocodylians that lived during the Late Cretaceous. It was a clade of terrestrial crocodilians that evolved very mammal-like teeth and jaws.MARINHO, T.S. & CARVALHO, I.S. Revision of the Sphagesaurida ...
crocodyliform in its
abdominal cavity The abdominal cavity is a large body cavity in humans and many other animals that contains many organs. It is a part of the abdominopelvic cavity. It is located below the thoracic cavity, and above the pelvic cavity. Its dome-shaped roof is the th ...
. The specimen represents direct evidence of
predation Predation is a biological interaction where one organism, the predator, kills and eats another organism, its prey. It is one of a family of common feeding behaviours that includes parasitism and micropredation (which usually do not kill the ...
between different taxa of
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 in the
fossil A fossil (from Classical Latin , ) is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserved ...
record.


Discovery

''Aplestosuchus'' 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 ...
LPRP/USP 0229a, an articulated and nearly complete
skeleton A skeleton is the structural frame that supports the body of an animal. There are several types of skeletons, including the exoskeleton, which is the stable outer shell of an organism, the endoskeleton, which forms the support structure inside ...
including the
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 ...
, housed at the Laboratório de Paleontologia,
Universidade de São Paulo The Universiade is an international multi-sport event, organized for university athletes by the International University Sports Federation (FISU). The name is a portmanteau of the words "University" and "Olympiad". The Universiade is referred t ...
. Additionally, 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 ...
and skull bones of an unidentified
sphagesaurid Sphagesauridae is a Gondwanan family of mesoeucrocodylians that lived during the Late Cretaceous. It was a clade of terrestrial crocodilians that evolved very mammal-like teeth and jaws.MARINHO, T.S. & CARVALHO, I.S. Revision of the Sphagesaurida ...
crocodyliform were preserved in the abdominal cavity of LPRP/USP 0229a, and assigned to the specimen number LPRP/USP 0229b. The find represents direct evidence of
predation Predation is a biological interaction where one organism, the predator, kills and eats another organism, its prey. It is one of a family of common feeding behaviours that includes parasitism and micropredation (which usually do not kill the ...
between different taxa of
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 in the
fossil A fossil (from Classical Latin , ) is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserved ...
record. LPRP/USP 0229 was found in the Buruti creek area, of the General Salgado municipality,
São Paulo São Paulo (, ; Portuguese for 'Saint Paul') is the most populous city in Brazil, and is the capital of the state of São Paulo, the most populous and wealthiest Brazilian state, located in the country's Southeast Region. Listed by the GaWC a ...
, in southern Brazil. To date, the locality yielded the type specimens of four other crocodyliforms, namely, '' Baurusuchus albertoi'', '' Baurusuchus salgadoensis'', ''
Armadillosuchus arrudai ''Armadillosuchus'' is an extinct genus of sphagesaurid crocodylomorph. It was described in February 2009 from the late Campanian to early Maastrichtian Adamantina Formation of the Bauru Basin in Brazil, dating to approximately 70 Ma. ''Armadill ...
'' and '' Gondwanasuchus scabrosus''. The specimens were collected from the
Adamantina Formation The Adamantina Formation is a geological formation in the Bauru Basin of western São Paulo state, in southeastern Brazil. Its strata date back to the Late Cretaceous epoch of the Cretaceous Period, during the Mesozoic Era. The formation is p ...
, Bauru Group of Paraná Basin, dating probably to the
Turonian The Turonian is, in the ICS' geologic timescale, the second age in the Late Cretaceous Epoch, or a stage in the Upper Cretaceous Series. It spans the time between 93.9 ± 0.8 Ma and 89.8 ± 1 Ma (million years ago). The Turonian is preceded by t ...
or the
Santonian The Santonian is an age in the geologic timescale or a chronostratigraphic stage. It is a subdivision of the Late Cretaceous Epoch or Upper Cretaceous Series. It spans the time between 86.3 ± 0.7 mya (million years ago) and 83.6 ± 0.7 mya. The ...
stage of 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 ...
, about 93.5-83.5
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). ...
. ''Aplestosuchus'' was first described and named by Pedro L. Godoy (best brother ever), Felipe C. Montefeltro, Mark A. Norell and Max C. Langer 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 ''Aplestosuchus sordidus''. The generic name is derived from the Greek ''aplestos'', meaning "insatiate", "gluttonous", 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 ''souchos'', an Egyptian crocodile god
Sebek Sobek (also called Sebek or Sobki, cop, Ⲥⲟⲩⲕ, Souk) was an ancient Egyptian deity with a complex and elastic history and nature. He is associated with the Nile crocodile or the West African crocodile and is represented either in its fo ...
. 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 ...
is derived from the
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 ...
''sordidus'', meaning "filthy", in reference to the greedy behavior of the animal, demonstrated by the
predation Predation is a biological interaction where one organism, the predator, kills and eats another organism, its prey. It is one of a family of common feeding behaviours that includes parasitism and micropredation (which usually do not kill the ...
of another closely related
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 ...
, an unidentified
sphagesaurid Sphagesauridae is a Gondwanan family of mesoeucrocodylians that lived during the Late Cretaceous. It was a clade of terrestrial crocodilians that evolved very mammal-like teeth and jaws.MARINHO, T.S. & CARVALHO, I.S. Revision of the Sphagesaurida ...
.


Description

The only known specimen of ''Aplestosuchus'' is more than one meter long from the tip of the skull to the base of the tail. It was preserved lying on its side, in the same
death pose Non-avian dinosaur and bird fossils are frequently found in a characteristic posture consisting of head thrown back, tail extended, and mouth wide open. The cause of this posture—often called a "death pose"—has been a matter of scientific deba ...
of other
baurusuchid Baurusuchidae is a Gondwanan Family (biology), family of mesoeucrocodylians that lived during the Late Cretaceous. It is a group of terrestrial hypercarnivorous crocodilians from South America (Argentina and Brazil) and possibly Pakistan. Baurusu ...
s collected in the site. LPRP/USP 0229a, preserved in two different blocks, suffered some post-mortem disarticulation, and as a result most of the tail and the distal parts of the
hindlimb A hindlimb or back limb is one of the paired articulated appendages (limbs) attached on the caudal ( posterior) end of a terrestrial tetrapod vertebrate's torso.http://www.merriam-webster.com/medical/hind%20limb, Merriam Webster Dictionary-Hindl ...
s were lost. ''Aplestosuchus'', like other
baurusuchid Baurusuchidae is a Gondwanan Family (biology), family of mesoeucrocodylians that lived during the Late Cretaceous. It is a group of terrestrial hypercarnivorous crocodilians from South America (Argentina and Brazil) and possibly Pakistan. Baurusu ...
s, was a fully terrestrial predator. Reaching up to four meters in length, baurusuchids were the apex predators of the South American Late Cretaceous ecosystems. They were surpassed only by large theropods, however, these are very rare in the Adamantina Formation and represented by isolated and fragmentary teeth mostly attributed to abelisaurids and carcharodontosaurids, and isolated
megaraptorid Megaraptora is a clade of carnivorous tetanuran theropod dinosaurs with controversial relations to other theropods. Its derived members, the Megaraptoridae are noted for their elongated hand claws and proportionally large arms, which are usually ...
and unenlagiine
vertebrae The spinal column, a defining synapomorphy shared by nearly all vertebrates,Hagfish are believed to have secondarily lost their spinal column is a moderately flexible series of vertebrae (singular vertebra), each constituting a characteristic i ...
. The
sphagesaurid Sphagesauridae is a Gondwanan family of mesoeucrocodylians that lived during the Late Cretaceous. It was a clade of terrestrial crocodilians that evolved very mammal-like teeth and jaws.MARINHO, T.S. & CARVALHO, I.S. Revision of the Sphagesaurida ...
material recovered in the
abdominal cavity The abdominal cavity is a large body cavity in humans and many other animals that contains many organs. It is a part of the abdominopelvic cavity. It is located below the thoracic cavity, and above the pelvic cavity. Its dome-shaped roof is the th ...
of ''Aplestosuchus'' was considered more likely derived from
predation Predation is a biological interaction where one organism, the predator, kills and eats another organism, its prey. It is one of a family of common feeding behaviours that includes parasitism and micropredation (which usually do not kill the ...
, not scavenging, given its size relation to the prey, as sphagesaurids were usually much smaller than about one fourth of derived baurusuchids. ''Aplestosuchus'' is distinguished form all other known Crocodyliformes by a unique combination of trait, including four autapomorphies. It possesses an autapomorphic
nasal bone 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 ...
with dorsal midline crest, and an autapomorphic frontal longitudinal ridge that reaches the midline contact between the prefrontals. Its medial supratemporal rim is crest-shaped, and the ridge along the ectopterygoid- jugal suture is notched at its caudal portion. A lateral depression is present on the quadrate bone. The palatine bar is autapomorphically crested on its ventral surface, and cylindrical in its dorsal portion. The choanal septum is also ridged on the ventral surface. An autapomorphic single parachoanal
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 ...
rostrolateral to the parachoanal
fenestrae 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 ...
is present at the base of the pterygoid wing. In the lower jaw of ''Aplestosuchus'', the outer sculpture 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 limited to the dentary, and the occipital surface of the mandibular symphysis lack a peg. Additionally, the ridged border of the angular is not covering the rostral edge of the mandibular fenestra, and a row of foramina is present between the mandibular fenestra and the ectopterygoid-jugal suture. Some of the non-autapomorphic traits of ''A. sordidus'' listed above, such as the quadrate depression, medial approximation of the prefrontals, and ridge on the ectopterygoid-jugal articulation, confirm its placement within Baurusuchidae.


Phylogeny

The
phylogenetic In biology, phylogenetics (; from Greek φυλή/ φῦλον [] "tribe, clan, race", and wikt:γενετικός, γενετικός [] "origin, source, birth") is the study of the evolutionary history and relationships among or within groups o ...
position of ''Aplestosuchus'' within Baurusuchidae was investigated in its original description, Godoy ''et al.'' (2014). ''Aplestosuchus'', along with the recently described baurusuchids ''
Campinasuchus ''Campinasuchus'' is an extinct genus of baurusuchid mesoeucrocodylian from Minas Gerais State of Brazil. Description ''Campinasuchus'', like other baurusuchids, was a fully terrestrial predator. It had a deep and laterally compressed skull wi ...
'' and '' Gondwanasuchus'', were scored into the taxon-character matrix of Montefeltro ''et al.'' (2011), which is focused solely on Baurusuchidae, with the addition of eight new characters. The resultant matrix includes 10 baurusuchid species as well as three outgroup taxa, scored based on 74 characters. Below is a cladogram from Godoy ''et al.'' (2014) showing the placement of ''Aplestosuchus'' based on their analysis.


References

{{Portal bar, Paleontology, Cretaceous Terrestrial crocodylomorphs Late Cretaceous crocodylomorphs of South America Cretaceous Brazil Fossils of Brazil Adamantina Formation Fossil taxa described in 2014 Prehistoric pseudosuchian genera