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''Aniksosaurus'' (meaning "spring lizard", from
Modern Greek Modern Greek (, , or , ''Kiní Neoellinikí Glóssa''), generally referred to by speakers simply as Greek (, ), refers collectively to the dialects of the Greek language spoken in the modern era, including the official standardized form of the ...
Άνοιξη, "Spring", referring to the fact it was found on 21 September 1995, the onset of Spring on the Southern Hemisphere) is a
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
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 ...
from what is now
Chubut Province Chubut ( es, Provincia del Chubut, ; cy, Talaith Chubut) is a province in southern Argentina, situated between the 42nd parallel south (the border with Río Negro Province), the 46th parallel south (bordering Santa Cruz Province), the Andes ra ...
,
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 was a
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 ...
, specifically a
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 ...
, which lived in 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 ...
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 ...
of the
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 ...
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 ...
, between 96-91 million years ago. 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 ...
, ''Aniksosaurus darwini'', was formally described from the
Bajo Barreal Formation The Bajo Barreal Formation is a geological formation in the Golfo San Jorge Basin of Chubut and Santa Cruz, Argentina whose strata date back to the Middle Cenomanian to Late Turonian. The formation was first described by Teruggi & Rossetto in ...
of the
Golfo San Jorge Basin The Golfo San Jorge Basin ( es, Cuenca del Golfo San Jorge) is a hydrocarbon-rich sedimentary basin located in eastern Patagonia, Argentina. The basin covers the entire San Jorge Gulf and an inland area west of it, having one half located in Santa ...
by Rubén Dario Martínez and
Fernando Emilio Novas Fernando Emilio Novas (born 1960) is an Argentine paleontologist working for the Comparative Anatomy Department of the Bernardino Rivadavia Natural Sciences Museum in Buenos Aires, Argentina.
in 2006;Martínez, R.D., and Novas, F.E. (2006)
"Aniksosaurus darwini gen. et sp. nov., a new coelurosaurian theropod from the early Late Cretaceous of central Patagonia, Argentina."
Revista del Museo Argentino de Ciencias Naturales, n.s. 8(2): 243-259.
the name was first coined in 1995 and reported in the literature in 1997. The specific epithet honors
Charles Darwin Charles Robert Darwin ( ; 12 February 1809 – 19 April 1882) was an English naturalist, geologist, and biologist, widely known for his contributions to evolutionary biology. His proposition that all species of life have descended fr ...
who visited
Patagonia Patagonia () refers to a geographical region that encompasses the southern end of South America, governed by Argentina and Chile. The region comprises the southern section of the Andes Mountains with lakes, fjords, temperate rainforests, and gl ...
in 1832/1833 during the Voyage of the Beagle.


Description

The type specimen MDT-PV 1/48, discovered in the
Bajo Barreal Formation The Bajo Barreal Formation is a geological formation in the Golfo San Jorge Basin of Chubut and Santa Cruz, Argentina whose strata date back to the Middle Cenomanian to Late Turonian. The formation was first described by Teruggi & Rossetto in ...
, consists of an articulated right hindlimb, which includes the
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 ...
,
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 ...
,
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 ...
and foot. The longest femur discovered for this genus measures 247mm, and the longest tibia, 270mm. On average, the tibia is 13% longer than the femur in ''Aniksosaurus'', an adaptation which has been strongly correlated with the development of cursorial habits in dinosaurs. Martinez and Novas (2006), based on femoral measurements, originally suggested that ''Aniksosaurus darwini'' was approximately long and tall at the hip, and weighed up to . However, recent estimates suggest a total body length between and and a body weight between and . In a 2013 study by Ibiricu et al., it was concluded that the holotype and four more individuals referable to ''Aniksosaurus'', were juvenile to sub-adult individuals as shown by a
histological analysis Histology, also known as microscopic anatomy or microanatomy, is the branch of biology which studies the microscopic anatomy of biological tissues. Histology is the microscopic counterpart to gross anatomy, which looks at larger structures vis ...
. The two individuals analyzed had histological characteristics that suggested an approximate age of three. The morphological evidence supporting ontogenetic immaturity was as follows: (a) the absence of an outer circumferential layer; (b) the absence of secondary (Haversian)
osteons In osteology, the osteon or haversian system (; named for Clopton Havers) is the fundamental functional unit of much compact bone. Osteons are roughly cylindrical structures that are typically between 0.25 mm and 0.35 mm in diameter. Their ...
; (c) the presence of only a few growth cycles; and (d) the thickness of the zones.


Classification

Martínez and Novas (2006) originally assigned this genus to the clade
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 ...
based on anatomical features present in the hindlimbs. The paleontologists describe ''Aniksosaurus'' as "more derived than some basal coelurosaurians such as compsognathids, ''
Ornitholestes ''Ornitholestes'' (meaning "bird robber") is a small theropod dinosaur of the late Jurassic (Brushy Basin Member of the Morrison Formation, middle Kimmeridgian age, about 154 million years agoTurner, C.E. and Peterson, F., (1999). "Biostratigraph ...
'', and coelurids", but less advanced than later coelurosaurs such as ''
Tyrannosaurus ''Tyrannosaurus'' is a genus of large theropoda, theropod dinosaur. The species ''Tyrannosaurus rex'' (''rex'' meaning "king" in Latin), often called ''T. rex'' or colloquially ''T-Rex'', is one of the best represented theropods. ''Tyrannosa ...
'' and ''
Oviraptor ''Oviraptor'' (; ) is a genus of oviraptorid dinosaur that lived in Asia during the Late Cretaceous period. The first remains were collected from the Djadokhta Formation of Mongolia in 1923 during a paleontological expedition led by Roy Chapman ...
''. ''Aniksosaurus'' exhibits some of the derived features found in Coelurosauria, such as (a) the ilium has a well developed cuppedicus fossa; (b) the femur possesses an anterior trochanter that is proximally projected, almost reaching the level of the articular head; (c) the greater trochanter is craniocaudally expanded; (d) the femoral head is rectangular-shaped in cranial aspect; (e) and the fibular shaft is craniocaudally narrow. However, research by Choiniere ''et al.'' (2010) demonstrated that
phylogenetically 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 ...
, ''Aniksosaurus'' was closer to individuals in the taxon
Compsognathidae Compsognathidae is a family of coelurosaurian theropod dinosaurs. Compsognathids were small carnivores, generally conservative in form, hailing from the Jurassic and Cretaceous Periods. The bird-like features of these species, along with other d ...
, while Hartman ''et al.'' (2019) find it to be a maniraptoromorph outside compsognathids and maniraptorans, and Naish ''et al.'' (2022) find it to be a
megaraptora Megaraptora is a clade of carnivorous Tetanurae, tetanuran theropod dinosaurs with controversial relations to other theropods. Its Derived (phylogenetics), derived members, the Megaraptoridae are noted for their elongated hand claws and proporti ...
n.


Distinguishing anatomical features

A diagnosis is a statement of the anatomical features of an organism (or group) that collectively distinguish it from all other organisms. Some, but not all, of the features in a diagnosis are also autapomorphies. An autapomorphy is a distinctive anatomical feature that is unique to a given organism. According to Martínez and Novas (2006), ''Aniksosaurus'' can be distinguished based on the following characteristics: * cranial
cervical vertebrae 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 ...
have dorsoventrally deep
neural arch 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 ...
es, and are provided with a pair of cavities at their cranial surfaces * the neural canal is wide * cranial caudals feature a ventral sagittal keel, and their transverse processes are triangular-shaped in dorsal view * manual ungual phalanges are robust * the metatarsal and digit IV of the pes are transversely narrow * ilium with an extremely expanded brevis shelf (a bone surface on the ilium that functions as an attachment site for tail muscles) * the
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 ...
possesses a deep notch for the M. iliotrochantericus


Paleobiology

Monospecific bonebeds suggest it was gregarious, and that this behavior gave ''Aniksosaurus'' a selective advantage over competitors in its
paleoenvironment Paleoecology (also spelled palaeoecology) is the study of interactions between organisms and/or interactions between organisms and their environments across geologic timescales. As a discipline, paleoecology interacts with, depends on and informs ...
. The hypothesis suggesting gregarious (social) behavior is supported by the taphonomic association of individuals from the Bajo Barreal Formation and evidence garnered from analysis of its bone histology. The authors of this study, Ibiricu, Martínez, Casal and Cerda (2013) noted that this particular assemblage is only the second known body fossil association of small, coelurosaurian theropods discovered in South America. The theropod taxa have been interpreted as gregarious are ''
Coelophysis ''Coelophysis'' ( traditionally; or , as heard more commonly in recent decades) is an extinct genus of coelophysid theropod dinosaur that lived approximately 228 to 201.3 million years ago during the latter part of the Triassic Period from t ...
bauri'', '' Syntarsus rhodiensensis'', ''
Albertosaurus ''Albertosaurus'' (; meaning "Alberta lizard") is a genus of tyrannosaurid theropod dinosaurs that lived in western North America during the Late Cretaceous Period, about 71 million years ago. The type species, ''A. sarcophagus'', was app ...
sarcophagus'' and ''
Sinornithomimus ''Sinornithomimus'' is a genus of ornithomimid that lived in Asia during the Late Cretaceous period. The first remains were found in 1997, in the Late Cretaceous strata of the Ulansuhai Formation located at Alshanzuo Banner, Inner Mongolia Auto ...
dongi''.Varricchio DJ, Sereno PC, Xijin Z, Lin T, Wilson JA, et al. (2008) Mud-trapped herd captures evidence of distinctive dinosaur sociality. Acta Palaeontologica Polonica 53: 567–578. doi: 10.4202/app.2008.0402. ''Aniksosaurus'' is likely to have exhibited gregariousness only during a specific interval of its life, most likely when it was a sub-adult.


Paleoecology


Provenance and occurrence

The remains of the type specimen MDT-PV 1/48 were discovered in the Laguna Palacios Ranch locality in the lower
Bajo Barreal Formation The Bajo Barreal Formation is a geological formation in the Golfo San Jorge Basin of Chubut and Santa Cruz, Argentina whose strata date back to the Middle Cenomanian to Late Turonian. The formation was first described by Teruggi & Rossetto in ...
of
Patagonia Patagonia () refers to a geographical region that encompasses the southern end of South America, governed by Argentina and Chile. The region comprises the southern section of the Andes Mountains with lakes, fjords, temperate rainforests, and gl ...
in Chubut,
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 ...
. The specimen was collected by Martínez in 1995 in green sandstone that was deposited between 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 Stage or stages may refer to: Acting * Stage (theatre), a space for the performance of theatrical productions * Theatre, a branch of the performing arts, often referred to as "the stage" * ''The Stage'', a weekly British theatre newspaper * Sta ...
and 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 ...
stage of the
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 ...
period, between 96-91 million years ago, as shown by
radiometric Radiometry is a set of techniques for measuring electromagnetic radiation, including visible light. Radiometric techniques in optics characterize the distribution of the radiation's power in space, as opposed to photometric techniques, which cha ...
Ar-Ar dating Arar or Ar-Ar may refer to: Geography and history * Arar, Saudi Arabia, the capital of Al Hudud ash Shamaliyah (The Northern Border) province ** Arar border crossing, a Saudi-Iraqi border crossing near Arar, Saudi Arabia and Nukhayb, Iraq * Arar, ...
. This specimen is housed in the collection of the Museo Regional Desiderio Torres - in Sarmiento, Chubut, Argentina. Remains of at least four more individuals, in 2006 assigned as 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 ...
s, were uncovered at the site.


Fauna and habitat

Dinosaur remains from this
formation Formation may refer to: Linguistics * Back-formation, the process of creating a new lexeme by removing or affixes * Word formation, the creation of a new word by adding affixes Mathematics and science * Cave formation or speleothem, a secondar ...
include those of the
abelisaurid Abelisauridae (meaning "Abel's lizards") is a family (or clade) of ceratosaurian theropod dinosaurs. Abelisaurids thrived during the Cretaceous period, on the ancient southern supercontinent of Gondwana, and today their fossil remains are found ...
theropod ''
Xenotarsosaurus ''Xenotarsosaurus'' is a genus of abelisaurid theropod dinosaur that lived during the Late Cretaceous of Argentina. Discovery In 1980 geologist Juan Carlos Sciutto discovered a rich fossil site six kilometres north of the Ocho Hermanos ranch in ...
bonapartei'', the
sauropods Sauropoda (), whose members are known as sauropods (; from '' sauro-'' + '' -pod'', 'lizard-footed'), is a clade of saurischian ('lizard-hipped') dinosaurs. Sauropods had very long necks, long tails, small heads (relative to the rest of their bo ...
''
Drusilasaura ''Drusilasaura'' is an extinct genus of possible lognkosaurian titanosaur sauropod dinosaur which lived during the late Cretaceous (Cenomanian-Turonian stage) of Santa Cruz Province of southern Patagonia, Argentina. ''Drusilasaura'' is known fr ...
'', ''
Epachthosaurus ''Epachthosaurus'' (meaning "heavy lizard") was a genus of titanosaurian sauropod dinosaur from the Late Cretaceous. It was a basal lithostrotian titanosaur. Its fossils have been found in Central and Northern Patagonia in South America. Discov ...
'' and ''
Campylodoniscus ''Campylodoniscus'' is a genus of titanosaur sauropod dinosaur from the Late Cretaceous Period of what is now Argentina. The type species was first named and described by Friedrich von Huene in 1929 as ''Campylodon ameghinoi'', the genus name ...
'', the
hadrosaur Hadrosaurids (), or duck-billed dinosaurs, are members of the ornithischian family Hadrosauridae. This group is known as the duck-billed dinosaurs for the flat duck-bill appearance of the bones in their snouts. The ornithopod family, which includ ...
''
Secernosaurus ''Secernosaurus'' (meaning "severed lizard") is a genus of herbivorous dinosaur. ''Secernosaurus'' was a hadrosaur, a "duck-billed" dinosaur which lived during the Late Cretaceous. ''Secernosaurus'' and its close relatives lived in South America, ...
'' as well as indeterminate teeth belonging to
carcharodontosaurids Carcharodontosauridae (carcharodontosaurids; from the Greek καρχαροδοντόσαυρος, ''carcharodontósauros'': "shark-toothed lizards") is a group of carnivorous theropod dinosaurs. In 1931, Ernst Stromer named Carcharodontosauridae ...
and
dromaeosaurids Dromaeosauridae () is a family of feathered theropod dinosaurs. They were generally small to medium-sized feathered carnivores that flourished in the Cretaceous Period. The name Dromaeosauridae means 'running lizards', from Greek ('), meaning ...
.


Taphonomy

The nature of the lithology at the ''Aniksosaurus'' site suggests that the deposits were made by a low-energy, unidirectional, overbank
fluvial In geography and geology, fluvial processes are associated with rivers and streams and the deposits and landforms created by them. When the stream or rivers are associated with glaciers, ice sheets, or ice caps, the term glaciofluvial or fluviog ...
floodplain deposit, which was produced by a lateral migrating fluvial channel. The observation that sandstone tabular bodies were deposited with coarse sandstones and
tuff Tuff is a type of rock made of volcanic ash ejected from a vent during a volcanic eruption. Following ejection and deposition, the ash is lithified into a solid rock. Rock that contains greater than 75% ash is considered tuff, while rock cont ...
interclasts also supports that ''Aniksosaurus'' was preserved in a fluvial environment. The soft tissue underwent subaerial
weathering Weathering is the deterioration of rocks, soils and minerals as well as wood and artificial materials through contact with water, atmospheric gases, and biological organisms. Weathering occurs ''in situ'' (on site, with little or no movement), ...
and then began to decompose, from the contributory action of small vertebrae scavengers as evidenced by a series of small grooves in one of the femora recovered, which was then followed by the partial disarticulation of the skeleton. Disarticulation was followed by reorientation of the remains by fluvial currents, which was then followed by differential burial in what appears to have been at least two stages of sedimentation. No evidence was found to support either cannibalism or intra-specific competition.


References


External links


Mickey Mortimer
an

on the genus. {{Portal bar, Dinosaurs, Paleontology, Cretaceous Compsognathids Cenomanian life Turonian life Late Cretaceous dinosaurs of South America Cretaceous Argentina Fossils of Argentina Bajo Barreal Formation Fossil taxa described in 2006 Taxa named by Fernando Novas