Aerosteon

Aerosteon is a genus of megaraptoran dinosaur from the Late Cretaceous period of Argentina. Its remains were discovered in 1996 in the Anacleto Formation, dating to the Santonian stage (about 84 million years ago).[1] The type and only known species is A. riocoloradense. Its specific name indicates that its remains were found 1 km (0.6 miles) north of the Río Colorado, in Mendoza Province, Argentina.

They show evidence of a bird-like respiratory system.[2] Aerosteon's name can be translated as air bone and derives from Greek ἀήρ (aer, "air") and ὀστέον (osteon, "bone"). Though the species name was originally published as "riocoloradensis", Greek ὀστέον is neuter gender, so according to the ICZN the species name must be riocoloradense to match.

Aerosteon
Temporal range: Santonian
~84 Ma
Aerosteon riocoloradensis
Skeletal diagram illustrating air-filled bones
Scientific classification
Kingdom: Animalia
Phylum: Chordata
Clade: Dinosauria
Order: Saurischia
Suborder: Theropoda
Clade: Megaraptora
Family: Megaraptoridae
Genus: Aerosteon
Sereno et al. 2009
Type species
Aerosteon riocoloradense
Sereno et al. 2009

Description

Aerosteon was an approximately 9 metres (30 ft) long, 2 metric ton bipedal carnivorous dinosaur that lived approximately 84 million years ago during the Santonian stage.

The holotype specimen, MCNA-PV-3137 consists of a single tooth (which may instead belong to an abelisaurid), some cranial bones, a number of partial or complete vertebrae from the neck, back, and sacrum, several cervical and dorsal ribs, gastralia, furcula (wishbone), left scapulocoracoid, left ilium, and left and right pubes. The incomplete fusion of some of its bones indicate that it was not quite fully mature.

Aerosteon did not initially appear to belong to any of the three groups of large theropods that were known to have inhabited the southern continents during this time (namely the Abelisauridae, Carcharodontosauridae or Spinosauridae). Sereno suggested that it might be related to the allosauroid radiation of the Jurassic period, and this was supported in subsequent studies that recognized a clade of late-surviving, lightly built, advanced allosauroids with large hand claws similar to the spinosaurs, called the Megaraptora, within the allosaur family Neovenatoridae.[3] A later analysis has placed Megaraptora, including Aerosteon, within the Tyrannosauroidea.[4] Megaraptorans have since been also considered as non-tyrannosauroid basal coelurosaurs in some analyses.[5][6]

Physiology

Furcula of the theropod Aerosteon riocoloradensis and magpie goose Anseranas semiplamata
Stereo images of the furculae of A. riocoloradense (A) and the Magpie-goose, Anseranas semipalmata (B). Scale bars are 10 cm in (A) and 2 cm in (B).

Some of Aerosteon's bones show pneumatisation (air-filled spaces), including pneumatic hollowing of the furcula and ilium, and pneumatisation of several gastralia, suggesting that it may have had a respiratory air-sac system similar to that of modern birds. These air sacs would have acted like bellows, moving air into and out of the animal's relatively inflexible lungs, instead of the lungs themselves being expanded and contracted as occurs with mammals. See avian respiratory system for more detailed information on this.

Sereno et al. theorize that this respiratory system may have developed to assist with regulating body temperature and was later co-opted for breathing.[2]

Classification and naming

Aerosteon was first described by Sereno et al. in a paper which appeared in the online journal PLoS ONE in September 2008. However, at the time, the International Code of Zoological Nomenclature did not recognize online publication of names for new species as valid unless print copies were also produced and distributed to several libraries, and that this action is noted in the paper itself. PLoS ONE initially failed to meet this requirement for Aerosteon. On 21 May 2009, the journal's managing editor coordinated with the ICZN to correct this oversight, publishing a comment to the original paper with an addendum stating that the requirements had been met as of that date. Consequently, though the description appeared in 2008, Aerosteon was not a valid name until 2009.[7]

A very close relative of Aerosteon, Murusraptor, was described in 2016 which preserved some bones with a lesser level of pneumaticity. However, the Murusraptor holotype also preserved several teeth which were very dissimilar to the one tooth observed in Aerosteon's holotype. The authors of the description noted that this tooth closely resembled that of abelisaurids and was probably incorrectly referred to Aerosteon. Murusraptor and Aerosteon are practically identical in the structure of their cranial bones and vertebrae, only noticeably differing in the proportions of the ilium, with Aerosteon's ilium being taller than that of Murusraptor.[8]

Pneumatopores on the left ilium of the theropod Aerosteon riocoloradensis
Pneumatopores on the left ilium of A. riocoloradense (A to D)

The cladogram below follows the 2010 analysis by Benson, Carrano and Brusatte, which considered megaraptorans as tetanurans.[3]

Neovenatoridae

Neovenator

unnamed

Chilantaisaurus

Megaraptora
unnamed

Siats meekerorum[9]

Australovenator

?Rapator

Fukuiraptor

unnamed

?Orkoraptor

Aerosteon

Megaraptor

The cladogram shown below follows an analysis by Porfiri et al., 2014, which recovered megaraptorans as tyrannosauroids.[10]

Megaraptora

Fukuiraptor

Megaraptoridae

Australovenator

Aerosteon

unnamed

Orkoraptor

Eotyrannus

Megaraptor

References

  1. ^ Novas, F.E.; Agnolin, F.L.; Ezcurra, M.D.; Porfiri, J.; Canale, J.I. (2013-10-01). "Evolution of the carnivorous dinosaurs during the Cretaceous: The evidence from Patagonia". Cretaceous Research. 45: 174–215. doi:10.1016/j.cretres.2013.04.001. ISSN 0195-6671.
  2. ^ a b Sereno, P.C., Martinez,R.N., Wilson, J.A., Varricchio, D.J., Alcober, O.A., and Larsson, H.C.E. (2008). Kemp, Tom (ed.). "Evidence for Avian Intrathoracic Air Sacs in a New Predatory Dinosaur from Argentina". PLoS ONE. 3 (9): e3303. Bibcode:2008PLoSO...3.3303S. doi:10.1371/journal.pone.0003303. PMC 2553519. PMID 18825273.CS1 maint: Uses authors parameter (link)
  3. ^ a b Benson R.B.J.; Carrano M.T; Brusatte S.L. (2010). "A new clade of archaic large-bodied predatory dinosaurs (Theropoda: Allosauroidea) that survived to the latest Mesozoic". Naturwissenschaften. 97 (1): 71–78. Bibcode:2010NW.....97...71B. doi:10.1007/s00114-009-0614-x. PMID 19826771.
  4. ^ F. E. Novas; F. L. Agnolín; M. D. Ezcurra; J. I. Canale; J. D. Porfiri (2012). "Megaraptorans as members of an unexpected evolutionary radiation of tyrant-reptiles in Gondwana". Ameghiniana. 49 (Suppl.): R33.
  5. ^ Apesteguía, S; Smith, N.D.; Valieri, R.J.; Makovicky, P.J. (2016). "An Unusual New Theropod with a Didactyl Manus from the Upper Cretaceous of Patagonia, Argentina". PLoS ONE. 11 (7): e0157793. Bibcode:2016PLoSO..1157793A. doi:10.1371/journal.pone.0157793. PMC 4943716. PMID 27410683.
  6. ^ Novas, F.E.; Aranciaga Rolando, A.M.; Agnolín, F.L. (2016). "Phylogenetic relationships of the Cretaceous Gondwanan theropods Megaraptor and Australovenator: the evidence afforded by their manual anatomy" (PDF). Memoirs of Museum Victoria. 74: 49–61. Archived from the original (PDF) on 2016-08-18.
  7. ^ PLoS ONE Group (2009). "Steps taken to meet the requirements of the ICZN to make new taxonomic names nomenclaturally available." Comment on Original Article: "Evidence for Avian Intrathoracic Air Sacs in a New Predatory Dinosaur from Argentina." PLoS ONE, 21 May 2009.
  8. ^ Coria, Rodolfo A.; Currie, Philip J. (2016-07-20). "A New Megaraptoran Dinosaur (Dinosauria, Theropoda, Megaraptoridae) from the Late Cretaceous of Patagonia". PLOS ONE. 11 (7): e0157973. Bibcode:2016PLoSO..1157973C. doi:10.1371/journal.pone.0157973. ISSN 1932-6203. PMC 4954680.
  9. ^ Zanno, L. E.; Makovicky, P. J. (2013). "Neovenatorid theropods are apex predators in the Late Cretaceous of North America". Nature Communications. 4: 2827. Bibcode:2013NatCo...4E2827Z. doi:10.1038/ncomms3827. PMID 24264527.
  10. ^ Juan D. Porfiri; Fernando E. Novas; Jorge O. Calvo; Federico L. Agnolín; Martín D. Ezcurra; Ignacio A. Cerda (2014). "Juvenile specimen of Megaraptor (Dinosauria, Theropoda) sheds light about tyrannosauroid radiation". Cretaceous Research. 51: 35–55. doi:10.1016/j.cretres.2014.04.007.

External links

Air sac

Air sacs are spaces within an organism where there is the constant presence of air. Among modern animals, birds possess the most air sacs (9–11), with their extinct dinosaurian relatives showing a great increase in the pneumatization (presence of air) in their bones. Theropods, like Aerosteon, have many air sacs in the body that are not just in bones, and they can be identified as the more primitive form of modern bird airways. Sauropods are well known for the amount of air pockets in their bones (especially vertebra), although one theropod, Deinocheirus, shows a rivalling amount of air pockets.

Anacleto Formation

The Anacleto Formation is a geologic formation with outcrops in the Argentine Patagonian provinces of Mendoza, Río Negro, and Neuquén. It is the youngest formation within the Neuquén Group and belongs to the Río Colorado Subgroup. Formerly that subgroup was treated as a formation, and the Anacleto Formation was known as the Anacleto Member.The type locality of this formation lies 40 kilometres (25 mi) west of the city of Neuquén. At its base, the Anacleto Formation conformably overlies the Bajo de la Carpa Formation, also of the Río Colorado Subgroup, and it is in turn unconformably overlain by the Allen Formation of the younger Malargüe Group.The Anacleto Formation varies between 60 and 90 metres (200 and 300 ft) thick, and consists mainly of claystones and mudstones, purple

and dark red in color, deposited in fluvial, lacustrine and floodplain environments. Geodes are often found scattered throughout this formation.

Australovenator

Australovenator (meaning "southern hunter") is a genus of megaraptorid theropod dinosaur from Cenomanian (Late Cretaceous)-age Winton Formation (dated to 95 million years ago) of Australia. It is known from partial cranial and postcranial remains which were described in 2009 by Scott Hocknull and colleagues, although additional descriptions and analyses continue to be published. It is the most complete predatory dinosaur discovered in Australia.

Dinosaur

Dinosaurs were a diverse group of reptiles of the clade Dinosauria. They first appeared during the Triassic period, between 243 and 233.23 million years ago, although the exact origin and timing of the evolution of dinosaurs is the subject of active research. They became the dominant terrestrial vertebrates after the Triassic–Jurassic extinction event 201 million years ago; their dominance continued through the Jurassic and Cretaceous periods. The fossil record demonstrates that birds are modern feathered dinosaurs, having evolved from earlier theropods during the late Jurassic Period. As such, birds were the only dinosaur lineage to survive the Cretaceous–Paleogene extinction event 66 million years ago. Dinosaurs can therefore be divided into avian dinosaurs, or birds; and non-avian dinosaurs, which are all dinosaurs other than birds. This article deals primarily with non-avian dinosaurs.

Dinosaurs are a varied group of animals from taxonomic, morphological and ecological standpoints. Birds, at over 10,000 living species, are the most diverse group of vertebrates besides perciform fish. Using fossil evidence, paleontologists have identified over 500 distinct genera and more than 1,000 different species of non-avian dinosaurs. Dinosaurs are represented on every continent by both extant species (birds) and fossil remains. Through the first half of the 20th century, before birds were recognized to be dinosaurs, most of the scientific community believed dinosaurs to have been sluggish and cold-blooded. Most research conducted since the 1970s, however, has indicated that all dinosaurs were active animals with elevated metabolisms and numerous adaptations for social interaction. Some were herbivorous, others carnivorous. Evidence suggests that all dinosaurs were egg-laying; and nest-building was a trait shared by many dinosaurs, both avian and non-avian.

While dinosaurs were ancestrally bipedal, many extinct groups included quadrupedal species, and some were able to shift between these stances. Elaborate display structures such as horns or crests are common to all dinosaur groups, and some extinct groups developed skeletal modifications such as bony armor and spines. While the dinosaurs' modern-day surviving avian lineage (birds) are generally small due to the constraints of flight, many prehistoric dinosaurs (non-avian and avian) were large-bodied—the largest sauropod dinosaurs are estimated to have reached lengths of 39.7 meters (130 feet) and heights of 18 meters (59 feet) and were the largest land animals of all time. Still, the idea that non-avian dinosaurs were uniformly gigantic is a misconception based in part on preservation bias, as large, sturdy bones are more likely to last until they are fossilized. Many dinosaurs were quite small: Xixianykus, for example, was only about 50 cm (20 in) long.

Since the first dinosaur fossils were recognized in the early 19th century, mounted fossil dinosaur skeletons have been major attractions at museums around the world, and dinosaurs have become an enduring part of world culture. The large sizes of some dinosaur groups, as well as their seemingly monstrous and fantastic nature, have ensured dinosaurs' regular appearance in best-selling books and films, such as Jurassic Park. Persistent public enthusiasm for the animals has resulted in significant funding for dinosaur science, and new discoveries are regularly covered by the media.

Eotyrannus

Eotyrannus (meaning "dawn tyrant") is a genus of tyrannosauroid theropod dinosaur hailing from the Early Cretaceous Wessex Formation beds, included in Wealden Group, located in the southwest coast of the Isle of Wight, United Kingdom. The remains (MIWG1997.550), consisting of assorted skull, axial skeleton and appendicular skeleton elements, from a juvenile or subadult, found in a plant debris clay bed, were described by Hutt et al. in early 2001. The etymology of the generic name refers to the animals classification as an early tyrannosaur or "tyrant lizard", while the specific name honors the discoverer of the fossil.

Gualicho

Gualicho (named in reference to the gualichu) is a genus of theropod dinosaur. The type species is Gualicho shinyae. Gualicho lived in what is now northern Patagonia, on what was then a South American island continent split off from the supercontinent Gondwana. The fossils were found in the Huincul Formation, dating to the late Cenomanian-early Turonian age of the upper Cretaceous Period, around 93 million years ago.

John Ruben

John A. Ruben is a researcher in Zoology and Vertebrate paleontology at the Oregon State University in Corvallis. Much of his published research is focused on studying the respiratory system in birds, in order to contradict the theory of theropodan ancestry of birds, as well as their metabolism.

List of South American dinosaurs

This is a list of dinosaurs whose remains have been recovered from South America.

List of dinosaur genera

This list of dinosaurs is a comprehensive listing of all genera that have ever been included in the superorder Dinosauria, excluding class Aves (birds, both living and those known only from fossils) and purely vernacular terms.

The list includes all commonly accepted genera, but also genera that are now considered invalid, doubtful (nomen dubium), or were not formally published (nomen nudum), as well as junior synonyms of more established names, and genera that are no longer considered dinosaurs. Many listed names have been reclassified as everything from birds to crocodilians to petrified wood. The list contains 1559 names, of which approximately 1192 are considered either valid dinosaur genera or nomina dubia.

Megaraptor

Megaraptor ("giant thief") is a genus of large theropod dinosaur that lived in the Turonian to Coniacian ages of the Late Cretaceous. Its fossils have been discovered in the Patagonian Portezuelo Formation of Argentina. Initially thought to have been a giant dromaeosaur-like coelurosaur, it was classified as a neovenatorid allosauroid in previous phylogenies, but more recent phylogeny and discoveries of related megaraptoran genera has placed it as either a basal tyrannosauroid or a basal coelurosaur.

Megaraptora

Megaraptora is a clade of carnivorous theropod dinosaurs with elongated hand claws and controversial relations to other theropods.Megaraptorans are incompletely known, and no complete megaraptoran skeleton has been found. However, they still possessed a number of unique features. Their forelimbs were large and strongly built, and the ulna bone had a unique shape in members of the family Megaraptoridae, a subset of megaraptorans which excludes Fukuiraptor. The first two fingers were elongated, with massive curved claws, while the third finger was small. Megaraptoran skull material is very incomplete, but a juvenile Megaraptor described in 2014 preserved a portion of the snout, which was long and slender. Leg bones referred to megaraptorans were also quite slender and similar to those of coelurosaurs adapted for running. Although megaraptorans were thick-bodied theropods, their bones were heavily pneumatized, or filled with air pockets. The vertebrae, ribs, and the ilium bone of the hip were pneumatized to an extent which was very rare among theropods, only seen elsewhere in taxa such as Neovenator. Other characteristic features include opisthocoelous neck vertebrae and compsognathid-like teeth.The clade was originally named in 2010 as a subset of the family Neovenatoridae, a group of lightly-built allosauroids related to the massive carcharodontosaurids such as Giganotosaurus and Carcharodontosaurus. A 2013 phylogenetic analysis by Fernando Novas and his colleagues disagreed with this classification scheme, and instead argued that the megaraptorans evolved deep within Tyrannosauroidea, a superfamily of basal coelurosaurs including the famous Tyrannosaurus. Subsequent refinements to Novas's data and methodologies have supported a third position for the group, at the base of Coelurosauria among other controversial theropods such as Gualicho, but not within the Tyrannosauroidea. Regardless of their position, it is clear that megaraptorans experienced a large amount of convergent evolution with either Neovenator-like allosauroids or basal coelurosaurs.Megaraptorans were most diverse in the early Late Cretaceous of South America, particularly Patagonia. However, they had a widespread distribution. Fukuiraptor, the most basal ("primitive") known member of the group, lived in Japan. Megaraptoran material is also common in Australia, and the largest known predatory dinosaur from the continent, Australovenator, was a megaraptoran.

Neovenator

Neovenator (nee-o-ven-a-tor) which means "new hunter" is a genus of allosauroid dinosaur. At the time of its discovery on the Isle of Wight, United Kingdom, it was the best-known large carnivorous dinosaur from the Early Cretaceous (Hauterivian-Barremian) of Europe.

Orkoraptor

Orkoraptor is a genus of medium-sized theropod dinosaur from the late Cretaceous Period of Argentina. It is known from incomplete fossil remains including parts of the skull, teeth, tail vertebrae, and a partial tibia. The specialized teeth resemble those of some maniraptoriform theropods, namely the deinonychosaurs and compsognathids. This and other anatomical features led the authors who described it (Novas, Ezcurra, and Lecuona) to suggest that it was a maniraptoran coelurosaur. However, subsequent studies found it to be a megaraptoran. Found in the Pari Aike Formation of Southern Patagonia, it is one of the southernmost carnivorous dinosaurs known from South America.

Paul Sereno

Paul Callistus Sereno (born October 11, 1957) is a professor of paleontology at the University of Chicago and a National Geographic "explorer-in-residence" who has discovered several new dinosaur species on several continents, including at sites in Inner Mongolia, Argentina, Morocco and Niger. One of his most widely publicized discoveries is that of a nearly complete specimen of Sarcosuchus imperator — popularly known as SuperCroc — at Gadoufaoua in the Tenere desert of Niger.

Planet Dinosaur

Planet Dinosaur, is a six-part documentary television series created by Nigel Paterson and Phil Dobree, produced by the BBC, and narrated by John Hurt. It first aired in the United Kingdom in 2011, with VFX studio Jellyfish Pictures as its producer. It is the first major dinosaur-related series for BBC One since Walking with Dinosaurs. There are more than 50 different prehistoric species featured, and they and their environments were created entirely as computer-generated images, for around a third of the production cost that was needed a decade earlier for Walking with Dinosaurs. Much of the series' plot is based on scientific discoveries made since Walking with Dinosaurs. The companion book to Planet Dinosaur was released on 8 September 2011, and the DVD and Blu-ray were released on 24 October 2011.

Siats

Siats is an extinct genus of large neovenatorid theropod dinosaur known from the Late Cretaceous Cedar Mountain Formation of Utah, US. It contains a single species, Siats meekerorum. S. meekerorum could be the first neovenatorid discovered in North America and the geologically youngest allosauroid yet discovered from the continent. It was initially classified as a megaraptoran, a clade of large theropods with very controversial relationships. This group may be examples of tyrannosauroids, neovenatorid allosauroids, or basal coelurosaurs.

Skeletal pneumaticity

Skeletal pneumaticity is the presence of air spaces within bones. It is generally produced during development by excavation of bone by pneumatic diverticula (air sacs) from an air-filled space, such as the lungs or nasal cavity. Pneumatization is highly variable between individuals, and bones not normally pneumatized can become pneumatized in pathological development.

Tratayenia

Tratayenia is an extinct genus of megaraptoran theropod dinosaurs known from remains found in the Santonian-age Bajo de la Carpa Formation of Argentina. The type and only species, Tratayenia rosalesi, was described in March 2018.Tratayenia can be distinguished from other megaraptorans on the basis of three autapomorphies (unique derived features) of the front portion of each dorsal vertebra, as well as a single autapomorphy of the sacrum. Tratayenia is the youngest known genus of megaraptoran, having lived only about 83 million years ago.

Yangchuanosaurus

Yangchuanosaurus is an extinct genus of metriacanthosaurid theropod dinosaur that lived in China during the Bathonian and Callovian stages of the Middle Jurassic, and was similar in size and appearance to its North American and European relative, Allosaurus. It hails from the Upper Shaximiao Formation and was the largest predator in a landscape that included the sauropods Mamenchisaurus and Omeisaurus and the stegosaurs Chialingosaurus, Tuojiangosaurus and Chungkingosaurus. It was named after the area in which was discovered, Yongchuan, in China.

Coelurosauria

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