Pampadromaeus
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''Pampadromaeus'' 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 living and fossil organisms as well as viruses. In the hierarchy of biological classification, genus comes above species and below family. In binomial nom ...
of basal
sauropodomorph Sauropodomorpha ( ; from Greek, meaning "lizard-footed forms") is an extinct clade of long-necked, herbivorous, saurischian dinosaurs that includes the sauropods and their ancestral relatives. Sauropods generally grew to very large sizes, had lon ...
dinosaur Dinosaurs are a diverse group of reptiles of the clade Dinosauria. They first appeared during the Triassic period, between 243 and 233.23 million years ago (mya), although the exact origin and timing of the evolution of dinosaurs is t ...
s known from the
Late Triassic The Late Triassic is the third and final epoch of the Triassic Period in the geologic time scale, spanning the time between Ma and Ma (million years ago). It is preceded by the Middle Triassic Epoch and followed by the Early Jurassic Epoch ...
(
Carnian The Carnian (less commonly, Karnian) is the lowermost stage (stratigraphy), stage of the Upper Triassic series (stratigraphy), Series (or earliest age (geology), age of the Late Triassic Epoch (reference date), Epoch). It lasted from 237 to 227 m ...
)
Santa Maria Formation The Santa Maria Formation is a sedimentary rock formation found in Rio Grande do Sul, Brazil. It is primarily Carnian in age (Late Triassic), and is notable for its fossils of cynodonts, "rauisuchian" pseudosuchians, and early dinosaurs and other ...
of the
Paraná Basin The Paraná Basin ( pt, Bacia do Paraná, es, Cuenca del Paraná) is a large cratonic sedimentary basin situated in the central-eastern part of South America. About 75% of its areal distribution occurs in Brazil, from Mato Grosso to Rio Grande d ...
in
Rio Grande do Sul Rio Grande do Sul (, , ; "Great River of the South") is a state in the southern region of Brazil. It is the fifth-most-populous state and the ninth largest by area. Located in the southernmost part of the country, Rio Grande do Sul is border ...
,
southern Brazil The South Region of Brazil (; ) is one of the five regions of Brazil. It includes the states of Paraná, Rio Grande do Sul, and Santa Catarina, and covers , being the smallest region of the country, occupying only about 6.76% of the territory of ...
.Cabreira et al., 2011


Discovery

''Pampadromaeus'' is known only 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 seve ...
specimen
ULBRA The Lutheran University of Brazil (Portuguese: or ULBRA) is a university which covers several states throughout Brazil. History The university has its roots in the parish school of St. Paul's Lutheran Church of Canoas (CELSP), in Canoas, Rio ...
-PVT016, a disarticulated, partial but well preserved
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 ...
from a single individual which includes most of 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, th ...
bones and the
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 ...
s;
dorsal Dorsal (from Latin ''dorsum'' ‘back’) may refer to: * Dorsal (anatomy), an anatomical term of location referring to the back or upper side of an organism or parts of an organism * Dorsal, positioned on top of an aircraft's fuselage * Dorsal c ...
, sacral and
caudal 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 ...
; elements of the
shoulder girdle The shoulder girdle or pectoral girdle is the set of bones in the appendicular skeleton which connects to the arm on each side. In humans it consists of the clavicle and scapula; in those species with three bones in the shoulder, it consists ...
and the
forelimb A forelimb or front limb is one of the paired articulated appendages (limbs) attached on the cranial ( anterior) end of a terrestrial tetrapod vertebrate's torso. With reference to quadrupeds, the term foreleg or front leg is often used inste ...
s, an ilium and elements 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-Hin ...
s. It was collected in the upper '' Hyperodapedon'' biozone from the Alemoa Member of the
Santa Maria Formation The Santa Maria Formation is a sedimentary rock formation found in Rio Grande do Sul, Brazil. It is primarily Carnian in age (Late Triassic), and is notable for its fossils of cynodonts, "rauisuchian" pseudosuchians, and early dinosaurs and other ...
( Rosário do Sul Group) in the "Janner" (also known as "Várzea do Agudo") locality,
geopark A geopark is a protected area with internationally significant geology within which sustainable development is sought and which includes tourism, conservation, education and research concerning not just geology but other relevant sciences. In 20 ...
of Paleorrota, dating to the
Carnian The Carnian (less commonly, Karnian) is the lowermost stage (stratigraphy), stage of the Upper Triassic series (stratigraphy), Series (or earliest age (geology), age of the Late Triassic Epoch (reference date), Epoch). It lasted from 237 to 227 m ...
faunal stage In chronostratigraphy, a stage is a succession of rock strata laid down in a single age on the geologic timescale, which usually represents millions of years of deposition. A given stage of rock and the corresponding age of time will by convent ...
of the early
Late Triassic The Late Triassic is the third and final epoch of the Triassic Period in the geologic time scale, spanning the time between Ma and Ma (million years ago). It is preceded by the Middle Triassic Epoch and followed by the Early Jurassic Epoch ...
, about 230–228 
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 ...
. A U-Pb (
uranium Uranium is a chemical element with the symbol U and atomic number 92. It is a silvery-grey metal in the actinide series of the periodic table. A uranium atom has 92 protons and 92 electrons, of which 6 are valence electrons. Uranium is weak ...
decay) dating found that the Santa Maria Formation dated around 233.23 million years ago, putting it 1.5 million years older than the
Ischigualasto Formation The Ischigualasto Formation is a Late Triassic fossiliferous formation and Lagerstätte in the Ischigualasto-Villa Unión Basin of the southwestern La Rioja Province and northeastern San Juan Province in northwestern Argentina. The formation ...
, and making the two formations approximately equal as the earliest dinosaur localities.Langer et al., 2018


Description

''Pampadromaeus'' was a small bipedal animal. It shows a mosaic of basal and derived traits. It differs from other sauropodomorphs by a combination of characters. Some of these are shared with members of the
Theropoda Theropoda (; ), whose members are known as theropods, is a dinosaur clade that is characterized by hollow bones and three toes and claws on each limb. Theropods are generally classed as a group of saurischian dinosaurs. They were ancestrally ...
: 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 ...
is pointed downwards forming a subnarial gap with 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. T ...
and the anterior-most teeth are unserrated; in the location where with theropods the ''fenestra promaxillaris'' is positioned, a small depression is present. Basal traits consist of a large skull, a short thighbone, the possession of just two sacral vertebrae and the presence of fifteen teeth in 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 * Medi ...
. There were four teeth in the premaxilla and about twenty in both the maxilla and the lower jaw for a total of eighty-eight. The teeth were large, elongated, lanceolate, slightly recurved, sharply pointed and coarsely serrated. The lower leg was much longer than the thighbone, indicating ''Pampadromaeus'' was a good runner.


Etymology

''Pampadromaeus'' was first named by Sergio F. Cabreira, Cesar L. Schultz, Jonathas S. Bittencourt, Marina B. Soares, Daniel C. Fortier, Lúcio R. Silva and Max C. Langer in
2011 File:2011 Events Collage.png, From top left, clockwise: a protester partaking in Occupy Wall Street heralds the beginning of the Occupy movement; protests against Libyan dictator Muammar Gaddafi, who was killed that October; a young man celebrates ...
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 specim ...
is ''Pampadromaeus barberenai''. The generic name is derived from
Quechua Quechua may refer to: *Quechua people, several indigenous ethnic groups in South America, especially in Peru *Quechuan languages, a Native South American language family spoken primarily in the Andes, derived from a common ancestral language **So ...
''
pampa The Pampas (from the qu, pampa, meaning "plain") are fertile South American low grasslands that cover more than and include the Argentine provinces of Buenos Aires, La Pampa, Santa Fe, Entre Ríos, and Córdoba; all of Uruguay; and Brazi ...
'', "plain", in reference to the present landscape of the site, and Greek δρομεύς, ''dromeus'', "runner", referring to the cursorial habits; the Latinised spelling variant ''dromaeus'' is used. The specific name honours the Brazilian
paleontologist Paleontology (), also spelled palaeontology or palæontology, is the scientific study of life that existed prior to, and sometimes including, the start of the Holocene epoch (roughly 11,700 years before present). It includes the study of fossi ...
Mário Costa Barberena.


Phylogeny

''Pampadromaeus'' was found to be a basal sauropodomorph in four different
cladistic Cladistics (; ) is an approach to biological classification in which organisms are categorized in groups ("clades") based on hypotheses of most recent common ancestry. The evidence for hypothesized relationships is typically shared derived char ...
analyses. The describers emphasized however, that this position was not strongly supported, showing the difficulties of determining the affinities of such early forms with the basal
Dinosauromorpha Dinosauromorpha is a clade of avemetatarsalian archosaurs (reptiles closer to birds than to crocodilians) that includes the Dinosauria (dinosaurs) and some of their close relatives. It was originally defined to include dinosauriforms and lage ...
,
Saurischia Saurischia ( , meaning "reptile-hipped" from the Greek ' () meaning 'lizard' and ' () meaning 'hip joint') is one of the two basic divisions of dinosaurs (the other being Ornithischia), classified by their hip structure. Saurischia and Ornithis ...
, Sauropodomorpha and Theropoda.


References


Bibliography

* * * {{Taxonbar, from=Q1705580 Sauropodomorphs Prehistoric tetrapod genera Carnian genera Late Triassic dinosaurs of South America Triassic Brazil Fossils of Brazil Santa Maria Formation Fossil taxa described in 2011 Paraná Basin