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''Alnashetri'' 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
alvarezsaurid Alvarezsauridae is a family of small, long-legged dinosaurs. Although originally thought to represent the earliest known flightless birds, they are now thought to be an early diverging branch of maniraptoran theropods. Alvarezsaurids were highly ...
coelurosauria Coelurosauria (; from Greek, meaning "hollow tailed lizards") is the clade containing all theropod dinosaurs more closely related to birds than to carnosaurs. Coelurosauria is a subgroup of theropod dinosaurs that includes compsognathids, tyrann ...
n
theropod 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 c ...
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 ...
known from the early
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 ...
(
Cenomanian The Cenomanian is, in the ICS' geological timescale, the oldest or earliest age of the Late Cretaceous Epoch or the lowest stage of the Upper Cretaceous Series. An age is a unit of geochronology; it is a unit of time; the stage is a unit in the s ...
stage) of La Buitrera,
Río Negro Province Río Negro (, ''Black River'') is a province of Argentina, located in northern Patagonia. Neighboring provinces are from the south clockwise Chubut, Neuquén, Mendoza, La Pampa and Buenos Aires. To the east lies the Atlantic Ocean. Its capit ...
,
Argentina Argentina (), officially the Argentine Republic ( es, link=no, República Argentina), is a country in the southern half of South America. Argentina covers an area of , making it the second-largest country in South America after Brazil, th ...
. It contains one known species, ''Alnashetri cerropoliciensis''. Its remains were discovered in
Candeleros Formation The Candeleros Formation (formerly known as the Candeleros Member of the "Río Limay Formation") is a geologic formation that crops out in the Río Negro, Neuquén, and Mendoza provinces of northern Patagonia, Argentina. It is the oldest formati ...
.


Discovery

''Alnashetri'' 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 several ...
MPCA-477, articulated
hind limb 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 of a single individual. The holotype consists of a partial left
femur The femur (; ), or thigh bone, is the proximal bone of the hindlimb in tetrapod vertebrates. The head of the femur articulates with the acetabulum in the pelvic bone forming the hip joint, while the distal part of the femur articulates with ...
, distal portions of left and right
tibia The tibia (; ), also known as the shinbone or shankbone, is the larger, stronger, and anterior (frontal) of the two bones in the leg below the knee in vertebrates (the other being the fibula, behind and to the outside of the tibia); it connects ...
e, right
fibula The fibula or calf bone is a leg bone on the lateral side of the tibia, to which it is connected above and below. It is the smaller of the two bones and, in proportion to its length, the most slender of all the long bones. Its upper extremity is ...
, proximal tarsals of both ankles, a nearly complete right
metatarsus The metatarsal bones, or metatarsus, are a group of five long bones in the foot, located between the tarsal bones of the hind- and mid-foot and the phalanges of the toes. Lacking individual names, the metatarsal bones are numbered from the med ...
, proximal and distal parts of the left
metapodial Metapodials are long bones of the hand (metacarpals) and feet (metatarsal The metatarsal bones, or metatarsus, are a group of five long bones in the foot, located between the tarsal bones of the hind- and mid-foot and the phalanges of the toes. ...
s and left pedal digit III. The specimen was collected at the "Hoyada de los esfenodontes" sublocality, part of the main fossiliferous locality of La Buitrera, about 30 km south of the village of El Chocόn in 2005. It was recovered from massive red sandstones that form the upper part of the
Candeleros Formation The Candeleros Formation (formerly known as the Candeleros Member of the "Río Limay Formation") is a geologic formation that crops out in the Río Negro, Neuquén, and Mendoza provinces of northern Patagonia, Argentina. It is the oldest formati ...
of the
Rio Limay Subgroup Rio or Río is the Portuguese, Spanish, Italian, and Maltese word for "river". When spoken on its own, the word often means Rio de Janeiro, a major city in Brazil. Rio or Río may also refer to: Geography Brazil * Rio de Janeiro * Rio do Sul, a ...
(
Cenomanian The Cenomanian is, in the ICS' geological timescale, the oldest or earliest age of the Late Cretaceous Epoch or the lowest stage of the Upper Cretaceous Series. An age is a unit of geochronology; it is a unit of time; the stage is a unit in the s ...
to
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 ...
stage), close to its border with the
Huincul Formation The Huincul Formation is a geologic formation of Late Cretaceous ( Early Cenomanian to Late Turonian) age of the Neuquén Basin that outcrops in the Mendoza, Río Negro and Neuquén Provinces of northern Patagonia, Argentina.
. It comes from the lower levels of the outcrops in this region, which produces a rich fauna of small vertebrates, including specimens of the large rhynchocephalian '' Priosphenodon avelesi'', the basal snake ''
Najash rionegrina ''Najash'' is an extinct genus of basal snake from the Late Cretaceous Candeleros Formation of Patagonia. Like a number of other Cretaceous and living snakes it retained hindlimbs, but ''Najash'' is unusual in having well-developed legs that ...
'', the notosuchian '' Araripesuchus buitreraensis'', the unenlagiine dromaeosaurid ''
Buitreraptor gonzalezorum ''Buitreraptor'' (meaning "La Buitrera seizer") is a genus of dromaeosaurid dinosaurs that lived during the Late Cretaceous of Argentina at the Candeleros Formation. ''Buitreraptor'' was described in 2005 and the type species is ''Buitreraptor ...
'', and also mammals and pterosaurs. It probably dates to the middle
Cenomanian The Cenomanian is, in the ICS' geological timescale, the oldest or earliest age of the Late Cretaceous Epoch or the lowest stage of the Upper Cretaceous Series. An age is a unit of geochronology; it is a unit of time; the stage is a unit in the s ...
stage of 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 ...
period Period may refer to: Common uses * Era, a length or span of time * Full stop (or period), a punctuation mark Arts, entertainment, and media * Period (music), a concept in musical composition * Periodic sentence (or rhetorical period), a concept ...
, about 97 
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). ...
.


Description

''Alnashetri'' is a small non-avian
coelurosaur Coelurosauria (; from Greek, meaning "hollow tailed lizards") is the clade containing all theropod dinosaurs more closely related to birds than to carnosaurs. Coelurosauria is a subgroup of theropod dinosaurs that includes compsognathids, tyrann ...
. It can be differentiated from all other coelurosaurs in the possession of a low ridge on the distal end of the tibia, which separates the rostral surface for articulation with the
astragalus ''Astragalus'' is a large genus of over 3,000 species of herbs and small shrubs, belonging to the legume family Fabaceae and the subfamily Faboideae. It is the largest genus of plants in terms of described species. The genus is native to tempe ...
from the outer face of the lateral malleolus and which extends up the shaft of the tibia dorsal to the tip of the ascending process of the astragalus. Further
autapomorphy 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 ...
(distinctive trait) of ''Alnashetri'' is the presence of small notches extending
ventral Standard anatomical terms of location are used to unambiguously describe the anatomy of animals, including humans. The terms, typically derived from Latin or Greek language, Greek roots, describe something in its standard anatomical position. Th ...
ly from the collateral ligament pits at the base of the distal articular hemicondyles on phalanges III-2 and III-3. ''Alnashetri'' can be easily distinguished from ''Buitreraptor'', the only other known small theropod from La Buitrera.


Phylogeny

Although ''Alnashetri'' was a coelurosaur, as shown by the fragmentary nature of the holotype skeleton, it lacks
synapomorphies In phylogenetics, an apomorphy (or derived trait) is a novel character or character state that has evolved from its ancestral form (or plesiomorphy). A synapomorphy is an apomorphy shared by two or more taxa and is therefore hypothesized to have ...
that would allow unambiguous referral to a particular coelurosaurian clade. Therefore, its phylogenetic position was initially tested in a cladistic matrix used to widely explore coelurosaurian interrelationships. This preliminary analysis found ''Alnashetri'' to nest within
Alvarezsauroidea Alvarezsauroidea is a group of small maniraptoran dinosaurs. Alvarezsauroidea, Alvarezsauridae, and Alvarezsauria are named for the historian Gregorio Álvarez, not the more familiar physicist Luis Alvarez, or his son geologist Walter Alvarez ...
. Eight characters with relevance for ''Alnashetri'' or for alvarezsaurid relationships and four recently described alvarezsauroids were added to the matrix. In this analysis, ''Alnashetri'' was recovered as a basal alvarezsaurid, in a
polytomy An internal node of a phylogenetic tree is described as a polytomy or multifurcation if (i) it is in a rooted tree and is linked to three or more child subtrees or (ii) it is in an unrooted tree and is attached to four or more branches. A tr ...
with other alvarezsaurids from the
Neuquén Basin Neuquén Basin ( es, Cuenca Neuquina) is a sedimentary basin covering most of Neuquén Province in Argentina. The basin originated in the Jurassic and developed through alternating continental and marine conditions well into the Tertiary. The bas ...
. The evidence supporting this relationship originates in the detailed anatomy of the ankle, however, and this concentration of character support within a single anatomical region may create false results. The
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 ...
below shows ''Alnashetri'' phylogenetic position among other alvarezsaurids following Makovicky, Apesteguía and Gianechini (2012).


Etymology

''Alnashetri'' was first described and named by Peter J. Makovicky, , and Federico A. Gianechini in
2012 File:2012 Events Collage V3.png, From left, clockwise: The passenger cruise ship Costa Concordia lies capsized after the Costa Concordia disaster; Damage to Casino Pier in Seaside Heights, New Jersey as a result of Hurricane Sandy; People gather ...
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 ''Alnashetri cerropoliciensis''. The generic name is derived from the Günün-a-kunna dialect of the
Tehuelche language Tehuelche (''Aoniken, Inaquen, Gunua-Kena, Gununa-Kena'') is one of the Chonan languages of Patagonia. Its speakers were nomadic hunters who occupied territory in present-day Chile, north of Tierra del Fuego and south of the Mapuche people. It is ...
, meaning "slender thighs" in reference to long and slender hind limbs of the holotype. The specific name ''cerropoliciensis'' honors the nearby small village of Cerro Policía for the generous assistance from its residents in fieldwork efforts at La Buitrera locality since its discovery in 1999.


References

{{Taxonbar, from=Q128050 Alvarezsaurids Monotypic dinosaur genera Cenomanian life Late Cretaceous dinosaurs of South America Cretaceous Argentina Fossils of Argentina Candeleros Formation Fossil taxa described in 2012