Aniksosaurus
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''Aniksosaurus'' (meaning "spring lizard", from
Modern Greek Modern Greek (, or , ), 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 language sometimes referred to ...
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 (; : genera ) is a taxonomic rank above species and below family (taxonomy), family as used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In bino ...
of avetheropod
dinosaur Dinosaurs are a diverse group of reptiles of the clade Dinosauria. They first appeared during the Triassic Geological period, period, between 243 and 233.23 million years ago (mya), although the exact origin and timing of the #Evolutio ...
from what is now
Chubut Province Chubut ( from Tehuelche language, Tehuelche 'transparent'; ) is a provinces of Argentina, province in southern Argentina, situated between the 42nd parallel south (the border with Río Negro Province), the 46th parallel south (bordering Santa ...
,
Argentina Argentina, officially the Argentine Republic, is a country in the southern half of South America. It covers an area of , making it the List of South American countries by area, second-largest country in South America after Brazil, the fourt ...
. It lived during the
Cenomanian The Cenomanian is, in the International Commission on Stratigraphy's (ICS) geological timescale, the oldest or earliest age (geology), age of the Late Cretaceous epoch (geology), Epoch or the lowest stage (stratigraphy), stage of the Upper Cretace ...
to
Turonian The Turonian is, in the International Commission on Stratigraphy, ICS' geologic timescale, the second age (geology), age in the Late Cretaceous epoch (geology), Epoch, or a stage (stratigraphy), stage in the Upper Cretaceous series (stratigraphy), ...
of the
Cretaceous The Cretaceous ( ) is a geological period that lasted from about 143.1 to 66 mya (unit), million years ago (Mya). It is the third and final period of the Mesozoic Era (geology), Era, as well as the longest. At around 77.1 million years, it is the ...
period Period may refer to: Common uses * Period (punctuation) * Era, a length or span of time *Menstruation, commonly referred to as a "period" Arts, entertainment, and media * Period (music), a concept in musical composition * Periodic sentence (o ...
, between 96-91 million years ago. The
type species In International_Code_of_Zoological_Nomenclature, 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 spe ...
, ''Aniksosaurus darwini'', was formally described from the
Bajo Barreal Formation The Bajo Barreal Formation is a geological Formation (geology), formation in the Golfo San Jorge Basin of Chubut Province, Chubut and Santa Cruz Province, Argentina, Santa Cruz, Argentina whose strata date back to the Cenomanian, Middle Cenomanian ...
of the
Golfo San Jorge Basin The Golfo San Jorge Basin () 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 Cruz Province and the other i ...
by Rubén Dario Martínez and
Fernando Emilio Novas Fernando Emilio Novas (born 1960) is an Argentina, Argentine Paleontology, 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 Natural history#Before 1900, naturalist, geologist, and biologist, widely known for his contributions to evolutionary biology. His proposition that all speci ...
who visited
Patagonia Patagonia () is a geographical region that includes parts of Argentina and Chile at the southern end of South America. The region includes the southern section of the Andes mountain chain with lakes, fjords, temperate rainforests, and glaciers ...
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 (geology), formation in the Golfo San Jorge Basin of Chubut Province, Chubut and Santa Cruz Province, Argentina, Santa Cruz, Argentina whose strata date back to the Cenomanian, Middle Cenomanian ...
, consists of an articulated right hindlimb, which includes the
femur The femur (; : femurs or femora ), or thigh bone is the only long bone, bone in the thigh — the region of the lower limb between the hip and the knee. In many quadrupeds, four-legged animals the femur is the upper bone of the hindleg. The Femo ...
,
fibula The fibula (: fibulae or fibulas) 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. ...
,
tibia The tibia (; : tibiae or tibias), also known as the shinbone or shankbone, is the larger, stronger, and anterior (frontal) of the two Leg bones, bones in the leg below the knee in vertebrates (the other being the fibula, behind and to the outsi ...
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. 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, tyra ...
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). "Biostratigrap ...
'', and coelurids", but less advanced than later coelurosaurs such as ''
Tyrannosaurus ''Tyrannosaurus'' () is a genus of large theropod dinosaur. The type species ''Tyrannosaurus rex'' ( meaning 'king' in Latin), often shortened to ''T. rex'' or colloquially t-rex, is one of the best represented theropods. It lived througho ...
'' 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. Choiniere et al. (2010) demonstrated that
phylogenetically In biology, phylogenetics () is the study of the evolutionary history of life using observable characteristics of organisms (or genes), which is known as phylogenetic inference. It infers the relationship among organisms based on empirical data ...
, ''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 Period (geology), Periods. The bird-like features of these species, a ...
. Hartman et al. (2019) instead find it to be a maniraptoromorph outside compsognathids and maniraptorans, while Naish and Cau (2022) find it to be a
megaraptora Megaraptora is a clade of carnivorous theropod dinosaurs. Its derived members, the Megaraptoridae are noted for their large hand claws and powerfully-built forelimbs, which are usually reduced in size in other large theropods. Although undoubt ...
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 (: 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 saurop ...
have dorsoventrally deep
neural arch Each vertebra (: vertebrae) is an irregular bone with a complex structure composed of bone and some hyaline cartilage, that make up the vertebral column or spine, of vertebrates. The proportions of the vertebrae differ according to their spinal ...
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 (; : femurs or femora ), or thigh bone is the only long bone, bone in the thigh — the region of the lower limb between the hip and the knee. In many quadrupeds, four-legged animals the femur is the upper bone of the hindleg. The Femo ...
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 that have been interpreted as gregarious are ''
Coelophysis ''Coelophysis'' ( Traditional English pronunciation of Latin, traditionally; or , as heard more commonly in recent decades) is a genus of coelophysid Theropoda, theropod dinosaur that lived Approximation, approximately 215 to 201.4 million y ...
bauri'', '' Syntarsus rhodiensensis'', ''
Albertosaurus ''Albertosaurus'' (; meaning "Alberta lizard") is a genus of large tyrannosaurid theropod dinosaur that lived in northwestern North America during the early to middle Maastrichtian age of the Late Cretaceous period, about 71 million yea ...
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 (geology), formation in the Golfo San Jorge Basin of Chubut Province, Chubut and Santa Cruz Province, Argentina, Santa Cruz, Argentina whose strata date back to the Cenomanian, Middle Cenomanian ...
of
Patagonia Patagonia () is a geographical region that includes parts of Argentina and Chile at the southern end of South America. The region includes the southern section of the Andes mountain chain with lakes, fjords, temperate rainforests, and glaciers ...
in Chubut,
Argentina Argentina, officially the Argentine Republic, is a country in the southern half of South America. It covers an area of , making it the List of South American countries by area, second-largest country in South America after Brazil, the fourt ...
. The specimen was collected by Martínez in 1995 in green sandstone that was deposited between the
Cenomanian The Cenomanian is, in the International Commission on Stratigraphy's (ICS) geological timescale, the oldest or earliest age (geology), age of the Late Cretaceous epoch (geology), Epoch or the lowest stage (stratigraphy), stage of the Upper Cretace ...
stage Stage, stages, or staging may refer to: Arts and media 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 Brit ...
and the
Turonian The Turonian is, in the International Commission on Stratigraphy, ICS' geologic timescale, the second age (geology), age in the Late Cretaceous epoch (geology), Epoch, or a stage (stratigraphy), stage in the Upper Cretaceous series (stratigraphy), ...
stage of the
Cretaceous The Cretaceous ( ) is a geological period that lasted from about 143.1 to 66 mya (unit), million years ago (Mya). It is the third and final period of the Mesozoic Era (geology), Era, as well as the longest. At around 77.1 million years, it is the ...
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 ch ...
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 (biology), isotype ...
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 secondary ...
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 foun ...
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 b ...
'' Drusilasaura'', ''
Epachthosaurus ''Epachthosaurus'' (meaning "heavy lizard") was a genus of titanosaurian sauropod dinosaur from the Late Cretaceous. It was a basal Lithostrotia, lithostrotian titanosaur. Its fossils have been found in Central and Northern Patagonia in South Ame ...
'' 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 me ...
'', the
hadrosaur Hadrosaurids (), also hadrosaurs 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 fami ...
''
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 and
dromaeosaurids Dromaeosauridae () is a family of feathered coelurosaurian theropod dinosaurs. They were generally small to medium-sized feathered carnivores that flourished in the Cretaceous Period. The name Dromaeosauridae means 'running lizards', from Gree ...
.


Taphonomy

The nature of the lithology at the ''Aniksosaurus'' site suggests that the deposits were made by a low-energy, unidirectional, overbank
fluvial A river is a natural stream of fresh water that flows on land or inside caves towards another body of water at a lower elevation, such as an ocean, lake, or another river. A river may run dry before reaching the end of its course if it ru ...
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 co ...
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, sunlight, and biological organisms. It occurs '' in situ'' (on-site, with little or no move ...
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 Compsognathidae Dinosaur genera Cenomanian dinosaurs Bajo Barreal Formation Fossil taxa described in 2006 Taxa named by Fernando Novas Dinosaurs of Argentina