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Australovenator

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Australovenator
Temporal range: Late Cretaceous, 95 Ma Possible Albian record
Reconstructed skeleton, Australian Age of Dinosaurs Museum, Winton, Australia
Scientific classification Edit this classification
Kingdom: Animalia
Phylum: Chordata
Class: Reptilia
Clade: Dinosauria
Clade: Saurischia
Clade: Theropoda
Clade: †Megaraptora
Family: †Megaraptoridae
Genus: †Australovenator
Hocknull et al. 2009
Species:
†A. wintonensis
Binomial name
†Australovenator wintonensis
Hocknull et al. 2009

Australovenator (meaning "southern hunter") is a genus of megaraptoran theropod dinosaur from the Late Cretaceous Winton Formation (Cenomanian, 95 million years ago[1]) of Australia. Some specimens from the Albian-aged Eumeralla Formation and the Wonthaggi Formation may belong to Australovenator. 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. It has been suggested that Australovenator is a sister taxon to Fukuiraptor, although some phylogenetic analyses find it to be a more derived member of the Megaraptora, possibly being part of the main Megaraptoridae family itself.

History of discovery

[edit]

Initial description

[edit]
Map of the Winton Formation depicting Queensland Province and the Winton Formation.
Map of the location and distribution of the Winton Formation (green), where fossils of Australovenator are known

Prior to the discovery of Australovenator, Australian theropods were only represented by taxa known from fragmentary and/or isolated material.[2][3][4] Beginning in 2006, the Australian Age of Dinosaurs Museum of Natural History (AODF) and the Queensland Museum began a series of expeditions in the "Matilda Site" at Elderslie Station. Fossils at the Station were first identified by the owners of property, with the first discovery being sauropod bone fragments.[5]: 68  This locale is located 60 kilometres (37 mi) north-west of Winton, central Queensland, Australia.[6][7][5]: 46-47  In 2006, an incomplete skeleton of a large theropod dinosaur was discovered at the "Matilda Site" in a layer of the Winton Formation, which dates to the Cenomanian stage of the Late Cretaceous period, around 95 million years ago. Strata containing the theropod fossils was made up of black soils that lacked recognizable facies and plant fossil-rich claystones, though all are known to derive from the Winton Formation.[1][6] This skeleton was found intermingled with the remains of a sauropod, later named Diamantinasaurus matildae.[7][5]: 69  After an initial three years of excavation from 2006 to 2009, the theropod skeleton was deposited at the AODF under specimen number AODF 604. Due to the material being encased in a phosphatic concretion, many fossils remained unidentified for years.[1][6][8][5]: 69 

Silhouette with known skeletal elements.

The first wave of preparation uncovered nine teeth, a left dentary (lower jaw bone), several ribs, an incomplete ilium (upper pelvic bone), parts of both hindlimbs and forelimbs, and some pedal (toe bones) and ungual phalanges (finger bones).[7] This included a giant, recurved manual ungual (hand claw), a trait characteristic of megaraptorans, a clade of theropods.[9] In 2009, Australian paleontologist Scott Hocknull and colleagues published a paper in the journal PLOS One in which they described several recently discovered specimens from the Winton Formation. Hocknull and colleagues scientifically described AODF 604 as the holotype (the specimen that forms the basis and bears the name of a species) of a new genus and species of theropod, Australovenator wintonensis. The generic name Australovenator derives from the Latin australis "southern", in reference to being unearthed in the Southern Hemisphere in Australia, and venator "hunter", in reference to its carnivorous diet. The specific name witonensis derives from Winton, the township the holotype was collected in. In the same paper, Hocknull and colleagues named Wintonotitan and Diamantinasaurus, two sauropods that were found in the "Matilda site".[7][10][11] Although the holotype was first discovered in 2006 and first described in 2009, the processing of excavating the "Matilda site" is still ongoing.[7] This included the preparation of additional fossils from the concretion containing the holotype, including elements of the arms and hands (which were described in 2012),[12] hindlimbs (which were described in 2013),[8] and the right dentary (which was described in 2015).[6] In total, the holotype is represented by both dentaries, many teeth, three dorsal ribs, some rib shaft pieces, humeri (upper arm bones), radii (upper forearm bones), ulnae (lower forearm bones), radialae (a carpal bone), both first metacarpals, incomplete manii (hands), an ilium fragment, left femur (thigh bone), tibiae (shin bones), fibulae, left astragalus, incomplete peses (feet), and nine gastralia fragments.[6][13][12][8]

Rapator and assigned material

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A manual ungual of cf. Australovenator in several views

In 1932, German paleontologist Friedrich von Huene named Rapator ornitholestoides, a genus now recognized to be a megaraptoran, on the basis of a single metacarpal.[14] This fossil was found in Lightning Ridge,[15] an outcrop of the Cenomanian-aged Griman Creek Formation.[16] Following the discovery of Australovenator, comparisons were made of the two.[1][9][15] While Hocknull and colleagues (2009) noted some differences in the anatomy based on a poorly preserved metacarpal I from Australovenator, their relationships were indeterminate.[7] In 2010, Argentine researcher Federico Agnolín and colleagues classified Rapator as a megaraptoran, possibly even Australovenator's sister taxon (closest relative). They followed up by stating that Megaraptor was less similar to Rapator than Australovenator, and that no definite differences between the latter two could be identified.[15] In 2013, Australian researcher Matt White and colleagues stated that the two were both morphologically and temporally different from one another, as the Griman Creek Formation is 10 million years older than the Winton Formation. This suggests that the two are not synonyms (the same taxon).[1]

In 1981, Australian paleontologist Ralph E. Molnar described an isolated left astragalus (NMVP 150070) that was discovered in an outcrop of the Aptian-aged Wonthaggi Formation in Victoria, Australia. He assigned the fossil to a species of pygmy Allosaurus, calling it Allosaurus sp.[17] Although some studies later considered it an allosauroid,[18] allosaurid,[19][20] or abelisauroid,[21] the consensus is that it belongs to a megaraptoran.[7][22][23][24] In a 2019 paper, Australian paleontologist Stephen Poropat and colleagues described several isolated megaraptoran fossils from the upper Aptian to lower Albian-aged Eumeralla Formation of Victoria, Australia. This material included two isolated teeth, two right manual unguals (hand claws), a right astragalus, and fragments possibly from a right pedal phalanx. The authors provisionally assigned the fossils to cf. Australovenator.[25] In 2020, a heavily eroded specimen was described, belonging to an indeterminate megaraptoran found near the type locality. The fossil material comprises "two fragmentary vertebrae, three partial metatarsals and the distal end of a pedal phalanx" as well as other indeterminate bone fragments. The animal was slightly larger than the holotype individual of Australovenator.[26]

Description

[edit]
Size of Australovenator compared to a human

Megaraptorans were bipedal, carnivorous theropods with elongated snouts, large arms, and relatively enlarged claws.[27][28] Australovenator was a relatively lightweight predator, leading Hocknull to coin it the "cheetah of its time".[10] South American megaraptorans attained very large sizes, exceeding 7 m (23 ft) in length in genera like Megaraptor.[28] In contrast, in a 2009 interview Hocknull stated that Australovenator was 2 metres (6.6 ft) tall at the hip and 6 metres (20 ft).[10] In 2016, American paleontologist Gregory S. Paul placed Australovenator at 6 metres (20 ft) long and weighed 500 kilograms (1,100 lb).[29] However, a 2014 study by American researcher Roger Benson and colleagues claimed its body mass was about 310 kilograms (680 lb).[30] Fossils of a megaraptoran, possibly Australovenator itself, from the Eumeralla Formation come from a slightly smaller individual than the holotype. However, morphologically they are very similar.[25]

Dentary and teeth

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Dentary in several views

Overall, the dentary is gracile and long, measuring 342.6 millimetres (13.49 in) in length as preserved.[6] The end of the symphysis (the area in which the dentaries meet) is rounded and lacks a "chin", unlike those of allosauroids like Allosaurus.[7] Each dentary bears 19 alveoli (tooth sockets), all of which touch interdental plates. These plates connect on their anterior (front) and posterior (back) ends, creating an elongated interdental ridge that trails across the whole tooth row. On the medial (inner) face of the dentary is a shallow paradental groove which runs below the alveolar margin and makes up the ventral (bottom) side of a protruding band of bone. This band is also known from Neovenator, but is absent in Eotyrannus.[6] Although foramina (small pits of bone) on the lateral (outside) side of the dentary were described by Hocknull and colleagues (2009),[7] later analysis has determined that Australovenator lacked these foramina. The absence of this feature may be an autapomorphy (characteristic unique to a species) of Australovenator.[6]

Its teeth are recurved with fine serrations, though the location of the serrations differs between the anterior and posterior teeth. Those with circular to quadrangular bases and two series of serrations, one on the lingual (facing towards the back of the jaw) carina (cutting edges) and the other on the distal carina, come from the anteriormost section of the dentary or premaxilla. In contrast, teeth that are thinner with serrations on the mesial (towards the front midline of the jaw) margin and the distal carina come from the posterior region of the dentary. Unlike the wrinkly teeth of carcharodontosaurids and Neovenator, Australovenator's and Fukuiraptor's, a megaraptoran from Japan, are smooth at the base.[7] Uniquely compared to most theropods, the dentition of megaraptorans like Australovenator displays some heterodonty, as the anatomy of the mesial and distal teeth are different. This suggests a functional difference between the two kinds of teeth.[6]

Postcrania

[edit]
Photos of the tibiae of Australovenator in several perspectives
Photos of the femora of Australovenator in several perspectives
Tibiae (left) and femora in several views

The humerus is S-shaped in anterior and lateral views, as in theropods like Allosaurus, Megaraptor, and Ornitholestes, and bears a prominent deltopectoral crest.[9] It has a bowed shaft, similar to that of Fukuiraptor.[12] A large furrow runs down the medial side of the shaft, as in Fukuiraptor. On the distal (away from body) end, both condyles (protuberances of bone for articulation) are pronounced and significantly rounded anteriorly to a greater extent than seen in Allosaurus and Xuanhanosaurus. These condyles are bifurcated by deep grooves, which are not found in non-coelurosaurian tetanurans like allosaurids but are similar to those of coelurosaurs like Guanlong. Although the radial condyle is taller and wider than that of the ulnar condyle, but not larger than it. This is unlike the distal humeral condyles of Fukuiraptor, where the radial condyle is twice as large as the ulnar condyle. On the posterior face of the humeral shaft is an enlarged tubercule, which is autapomorphic of the taxon. Australovenator preserves giant, sickle-shaped manual unguals, the largest on the hand being on the first finger.[7][25] The proximal end of the unguals are tall, and strongly taper to a sharp tip. These unguals are also strongly compressed mediolaterally (flat) with large vascular grooves running down their lateral (side) faces.[25]

Like other megaraptorans, Australovenator has a high humerus/femur length ratio (0.56).[9] This is around the same as that of Suchomimus (0.54), but higher than that of Chilantaisaurus (0.46).[31] Unlike Chilantaisaurus and Neovenator, Australovenator's femora were marked by large grooves that tapered disto-laterally (down to the left) on their posterior sides. Its femoral condyles were closely spaced from eachother and weakly divided, though not to the extent seen in Neovenator. The left medial condyle points distally, a trait characteristic of megaraptorans or neovenatorids. In contrast to the robustly built fibulae of Neovenator, Australovenator's are elongate and lithe. The metatarsus is gracile compared to those of Neovenator, Chilantaisaurus, and Allosaurus. A distinct ridge is on the posterior face of Australovenator's astragali, which runs from the center of the bone to the lateral edge. This trait is absent in relatives like Fukuiraptor. More pedal phalanges are known from Australovenator than most other megaraptorans.[8] Similar to the other megaraptoran Aerosteon,[32] Australovenator's dorsal ribs contain pneumatic cavities (air sacs).[7]

Classification

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Australovenator belonged to the clade Megaraptora, a group of theropods that existed during the Cretaceous period in Asia, South America, and Oceania,[33] and possibly Africa,[34] North America,[35] and Europe.[36] Most megaraptorans are only known from fragme vntary or isolated remains, leading to their relationships to be poorly known. Although they are always considered tetanurans, whether they are carcharodontosaurs,[37] basal coelurosaurs,[23][38] or tyrannosauroids is hotly debated.[39] Megaraptora was not recognized as a group until 2010, when it was named by American paleontologist Roger Benson and collagues. They considered it a subgroup of Neovenatoridae,[37] but later studies have generally opposed this.[23][40][41] When initially described, Australovenator was considered Theropoda incertae sedis,[7] although at the time few megaraptorans were known.[28][23]

In their description of Megaraptora, Benson and colleagues found Australovenator was a megaraptoran in their phylogenetic analysis. However, Australovenator was sister taxon to Fukuiraptor and the two were outside of Megaraptoridae itself.[37] The same results were recovered by Coria and Currie (2016)[42] and Novas and colleagues (2013).[43] In 2016, Argentine researcher Sebastian Apesteguía and colleagues supported the allosauroid affinities of Megaraptora, including Australovenator. However, they also said they could be basal coelurosaurs.[34] Below are the results of Coria and Currie (2016):

Allosauria

Most recent studies have found Megaraptora as a coelurosaurian clade, often sister to or within Tyrannosauroidea. Australovenator itself is typically a basal megaraptorid.[23][38][39][44][45] In their phylogenetic analyses, Ibiricu and colleagues (2025) recovered Australovenator as a basal member of Megaraptoridae, at a similar grade to Orkoraptor. Their analysis recovered megaraptorans as part of the theropod clade Coelurosauria, as the sister group to the Tyrannosauroidea. These results are displayed in the cladogram below (strict consensus of most parsimonious trees, after pruning Aoniraptor):[46]

Palaeobiology

[edit]
Bones of the left hand

With comprehensive and well-preserved remains of its hand and foot, Australovenator has been made a topic of various research papers studying the dynamics of theropod appendages.

A 2015 study tested the range of motion of Australovenator's arms using computer models and found that it had flexible arms, with the forearms capable of making an angle of 144 to 66 degrees with the humerus, an elbow range of motion similar to that of maniraptoriforms. Unusually, its radius could slide independently of the ulna when its arm was flexed, similar to that of birds but unlike most non-avian dinosaurs. However, the study also found that Australovenator's fingers were capable of extension far beyond those of any other sampled theropod, with only Dilophosaurus having capabilities even near it. This study concluded that Australovenator's flexibility, facilitated by a combination of traits in both primitive and advanced theropods, played a role in prey capture, giving it the ability to grasp prey towards its chest to make it easier for its weak jaws to disembowel food.[13] The gracile morphology of the skull also concludes that this genus had a specialization towards prey capture using its arms and hands.[47]

A 2016 study used CT scans of an emu foot to digitally reconstruct the musculature and soft tissue of an Australovenator foot, as well as determine how soft tissue affects flexibility. The study determined that muscular range of motion is often overestimated when not accounting for soft tissue and that soft tissue reconstruction is vital for making future analyses of theropod flexibility more accurate. A review of hindlimb elements described in 2013 re-identified several phalanges which were initially positioned incorrectly. In addition, it noted that Australovenator's phalanx II-3 was splayed, a pathology that may have resulted from the impacts of kicking motions. Some modern birds, such as the cassowary, are known to use their second toe as weapons in defensive or territorial fights.[48]

Life restoration

A 2017 follow-up to the 2016 study used a 3-D printed model of the reconstructed foot to make footprints in a matrix of clay and sand in an effort to understand the creation of dinosaur footprints. The study specifically was designed to clarify the identity of particular controversial footprints from Lark Quarry, which may have been left from either a large theropod (like Australovenator) or an ornithopod (like Muttaburrasaurus). The study found that the artificial Australovenator footprints were similar to those at Lark Quarry, concluding that the trackways in question were likely those of a theropod. The writers of the study expressed interest in creating a reconstruction of a Muttaburrasaurus foot as an extension of the study, although no Muttaburrasaurus pedal material is known.[49]

The right fourth pedal phalanx from the third digit features an unusual tuberosity on the lateral side of the proximal (towards body) articular facet (area where the phalanx would articulate with another). This may be a result of a pathology, such as an infection or arthritis. However, this could not be confirmed.[8]

Palaeoecology

[edit]

AODL 604 was found about 60 kilometres (37 mi) northwest of Winton, near Elderslie Station. It was recovered from the lower part of the Winton Formation, dated to the late Cenomanian. AODL 604 was found in a clay layer between sandstone layers. This has been interpreted as an oxbow lake, or billabong, deposit that was located at the eastern boundary of the Eromanga Sea. This formation was laid down at a paleolatitude of about 51 degrees South, which places it at around the same latitude as modern-day Patagonia and New Zealand.[6][50] Also found at the site were the type specimen of the sauropod Diamantinasaurus, bivalves, fish, turtles, crocodilians, and plant fossils. The Winton Formation had a faunal assemblage including bivalves, gastropods, insects, the lungfish Metaceratodus, turtles, the crocodilian Isisfordia, pterosaurs, and several types of dinosaurs, such as the sauropods Diamantinasaurus and Wintonotitan, and unnamed ankylosaurians and hypsilophodonts. Plants known from the formation include ferns, ginkgoes, gymnosperms, and angiosperms.[7]

References

[edit]
  1. 1 2 3 4 5 White, M. A.; Falkingham, P. L.; Cook, A. G.; Hocknull, S. A.; Elliott, D. A. (2013). "Morphological comparisons of metacarpal I for Australovenator wintonensis and Rapator ornitholestoides: Implications for their taxonomic relationships". Alcheringa: An Australasian Journal of Palaeontology. 37 (4): 435–441. Bibcode:2013Alch...37..435W. doi:10.1080/03115518.2013.770221. S2CID 82672110.
  2. ↑ Molnar, R.E.; Pledge, N.S. (1980-01-01). "A new theropod dinosaur from South Australia". Alcheringa: An Australasian Journal of Palaeontology. 4 (4): 281–287. Bibcode:1980Alch....4..281M. doi:10.1080/03115518008558972. ISSN 0311-5518.
  3. ↑ Long, J.A. (1998). Dinosaurs of Australia and New Zealand and other animals of the Mesozoic Era. Harvard University Press, UNSW Press, pp 94–96. ISBN 0-86840-448-9
  4. ↑ Benson, Roger B. J.; Rich, Thomas H.; Vickers-Rich, Patricia; Hall, Mike (2012-05-16). "Theropod Fauna from Southern Australia Indicates High Polar Diversity and Climate-Driven Dinosaur Provinciality". PLOS ONE. 7 (5) e37122. Bibcode:2012PLoSO...737122B. doi:10.1371/journal.pone.0037122. ISSN 1932-6203. PMC 3353904. PMID 22615916.
  5. 1 2 3 4 White, Matt (2017). "The revised skeletal description and biological range of motion analysis of Australia's most complete theropod dinosaur Australovenator wintonensis (Theropoda: Megaraptoridae)". University of Newcastle, Australia; Thesis.
  6. 1 2 3 4 5 6 7 8 9 10 White, Matt A.; Bell, Phil R.; Cook, Alex G.; Poropat, Stephen F.; Elliott, David A. (2015-12-15). "The dentary of Australovenator wintonensis (Theropoda, Megaraptoridae); implications for megaraptorid dentition". PeerJ. 3 e1512. Bibcode:2015PeerJ...3e1512W. doi:10.7717/peerj.1512. ISSN 2167-8359. PMC 4690360. PMID 26713256.
  7. 1 2 3 4 5 6 7 8 9 10 11 12 13 14 Hocknull, Scott A.; White, Matt A.; Tischler, Travis R.; Cook, Alex G.; Calleja, Naomi D.; Sloan, Trish; Elliott, David A. (2009). Sereno, Paul (ed.). "New Mid-Cretaceous (Latest Albian) Dinosaurs from Winton, Queensland, Australia". PLOS ONE. 4 (7) e6190. Bibcode:2009PLoSO...4.6190H. doi:10.1371/journal.pone.0006190. PMC 2703565. PMID 19584929.
  8. 1 2 3 4 5 White, Matt A.; Benson, Roger B. J.; Tischler, Travis R.; Hocknull, Scott A.; Cook, Alex G.; Barnes, David G.; Poropat, Stephen F.; Wooldridge, Sarah J.; Sloan, Trish (2013-07-24). "New Australovenator Hind Limb Elements Pertaining to the Holotype Reveal the Most Complete Neovenatorid Leg". PLOS ONE. 8 (7) e68649. Bibcode:2013PLoSO...868649W. doi:10.1371/journal.pone.0068649. ISSN 1932-6203. PMC 3722220. PMID 23894328.
  9. 1 2 3 4 Novas, Fernando E.; Rolando, Alexis M. Aranciaga; Agnolin, Federico L. (2016). "Phylogenetic relationships of the Cretaceous Gondwanan theropods 'Megaraptor' and 'Australovenator': The evidence afforded by their manual anatomy". Memoirs of Museum Victoria. 74: 49–61. doi:10.24199/j.mmv.2016.74.05.
  10. 1 2 3 Phillips, Nicky (2009-07-03). "Scientists discover 3 new Aussie dinosaurs". ABC Science Online. ABC News. Retrieved 2018-01-11.{{cite news}}: CS1 maint: deprecated archival service (link)
  11. ↑ Poropat, Stephen F.; Upchurch, Paul; Mannion, Philip D.; Hocknull, Scott A.; Kear, Benjamin P.; Sloan, Trish; Sinapius, George H. K.; Elliott, David A. (2015-04-01). "Revision of the sauropod dinosaur Diamantinasaurus matildae Hocknull et al. 2009 from the mid-Cretaceous of Australia: Implications for Gondwanan titanosauriform dispersal". Gondwana Research. 27 (3): 995–1033. Bibcode:2015GondR..27..995P. doi:10.1016/j.gr.2014.03.014. ISSN 1342-937X.
  12. 1 2 3 White, M. A.; Cook, A. G.; Hocknull, S. A.; Sloan, T.; Sinapius, G. H. K.; Elliott, D. A. (2012). Dodson, Peter (ed.). "New Forearm Elements Discovered of Holotype Specimen Australovenator wintonensis from Winton, Queensland, Australia". PLOS ONE. 7 (6) e39364. Bibcode:2012PLoSO...739364W. doi:10.1371/journal.pone.0039364. PMC 3384666. PMID 22761772.
  13. 1 2 White, Matt A.; Bell, Phil R.; Cook, Alex G.; Barnes, David G.; Tischler, Travis R.; Bassam, Brant J.; Elliott, David A. (2015-09-14). "Forearm Range of Motion in Australovenator wintonensis (Theropoda, Megaraptoridae)". PLOS ONE. 10 (9) e0137709. Bibcode:2015PLoSO..1037709W. doi:10.1371/journal.pone.0137709. ISSN 1932-6203. PMC 4569425. PMID 26368529.
  14. ↑ von Huene, Friedrich (1932). "Die fossile Reptil-Ordnung Saurischia, ihre Entwicklung und Geschichte" [The fossil reptile order Saurischia, its evolution and history]. Monographien zur Geologie und Paläontologie, Series. 1 and 2. 4 (1): 1–361.
  15. 1 2 3 Agnolin, Federico L.; Ezcurra, Martín D.; Pais, Diego F.; Salisbury, Steven W. (2010-05-24). "A reappraisal of the Cretaceous non-avian dinosaur faunas from Australia and New Zealand: evidence for their Gondwanan affinities". Journal of Systematic Palaeontology. 8 (2): 257–300. doi:10.1080/14772011003594870. ISSN 1477-2019.
  16. ↑ Bell, Phil R.; Fanti, Federico; Hart, Lachlan J.; Milan, Luke A.; Craven, Stephen J.; Birch, Sienna A.; Smith, Elizabeth (2019-01-15). "Revised geology, age, and vertebrate diversity of the dinosaur-bearing Griman Creek Formation (Cenomanian), Lightning Ridge, New South Wales, Australia". Palaeogeography, Palaeoclimatology, Palaeoecology. 514: 655–671. doi:10.1016/j.palaeo.2018.11.020. ISSN 0031-0182.
  17. ↑ Molnar, Ralph E.; Flannery, Timothy F.; Rich, Thomas H.V. (1981). "An allosaurid theropod dinosaur from the Early Cretaceous of Victoria, Australia". Alcheringa. 5 (2): 141–146. Bibcode:1981Alch....5..141M. doi:10.1080/03115518108565427.
  18. ↑ Currie, Philip J.; Zhao, Xi-Jin (1993). "A new carnosaur (Dinosauria, Theropoda) from the Jurassic of Xinjiang, People's Republic of China". Canadian Journal of Earth Sciences. 30 (10): 2037–2081. doi:10.1139/e93-179. ISSN 0008-4077.
  19. ↑ PéRez-Moreno, B. P.; Chure, D. J.; Pires, C.; Marques Da Silva, C.; Dos Santos, V.; Dantas, P.; PóVoas, L.; CachãO, M.; Sanz, J. L.; Galopim De Carvalho, A. M. (1999). "On the presence of Allosaurus fragilis (Theropoda: Carnosauria) in the Upper Jurassic of Portugal: first evidence of an intercontinental dinosaur species". Journal of the Geological Society. 156 (3): 449–452. doi:10.1144/gsjgs.156.3.0449. ISSN 0016-7649.
  20. ↑ Chure, Daniel (1998). "A reassessment of the Australian Allosaurus and its implications for the Australian refugium concept" (PDF). Fifty-Eighth Annual Meeting, Society of Vertebrate Paleontology. 18: 34A.
  21. ↑ Agnolin, F. L.; Ezcurra, M. D.; Pais, D. F.; Salisbury, S. W. (2010). "A reappraisal of the Cretaceous non-avian dinosaur faunas from Australia and New Zealand: Evidence for their Gondwanan affinities". Journal of Systematic Palaeontology. 8 (2): 257–300. Bibcode:2010JSPal...8..257A. doi:10.1080/14772011003594870. S2CID 130568551.
  22. ↑ Samathi, Adun; Chanthasit, Phornphen; Sander, Paul (2019). "Two new basal coelurosaurian theropod dinosaurs from the Early Cretaceous Sao Khua Formation of Thailand". Acta Palaeontologica Polonica. 64. doi:10.4202/app.00540.2018. ISSN 0567-7920.
  23. 1 2 3 4 5 Aranciaga Rolando, Alexis M.; Motta, Matias J.; Agnolín, Federico L.; Manabe, Makoto; Tsuihiji, Takanobu; Novas, Fernando E. (2022-04-26). "A large Megaraptoridae (Theropoda: Coelurosauria) from Upper Cretaceous (Maastrichtian) of Patagonia, Argentina". Scientific Reports. 12 (1): 6318. doi:10.1038/s41598-022-09272-z. ISSN 2045-2322. PMC 9042913. PMID 35474310.
  24. ↑ Kotevski, Jake; Duncan, Ruairidh J.; Ziegler, Tim; Bevitt, Joseph J.; Vickers-Rich, Patricia; Rich, Thomas H.; Evans, Alistair R.; Poropat, Stephen F. (2024-07-03). "Evolutionary and paleobiogeographic implications of new carcharodontosaurian, megaraptorid, and unenlagiine theropod remains from the upper Lower Cretaceous of Victoria, southeast Australia". Journal of Vertebrate Paleontology. 44 (4): e2441903. doi:10.1080/02724634.2024.2441903. ISSN 0272-4634.{{cite journal}}: CS1 maint: article number as page number (link)
  25. 1 2 3 4 Poropat, Stephen F.; White, Matt A.; Vickers-Rich, Patricia; Rich, Thomas H. (2019-07-04). "New megaraptorid (Dinosauria: Theropoda) remains from the Lower Cretaceous Eumeralla Formation of Cape Otway, Victoria, Australia". Journal of Vertebrate Paleontology. 39 (4) e1666273. Bibcode:2019JVPal..39E6273P. doi:10.1080/02724634.2019.1666273. ISSN 0272-4634. S2CID 208603798.
  26. ↑ White, Matt A.; Bell, Phil R.; Poropat, Stephen F.; Pentland, Adele H.; Rigby, Samantha L.; Cook, Alex G.; Sloan, Trish; Elliott, David A. (2020). "New theropod remains and implications for megaraptorid diversity in the Winton Formation (lower Upper Cretaceous), Queensland, Australia". Royal Society Open Science. 7 (1) 191462. Bibcode:2020RSOS....791462W. doi:10.1098/rsos.191462. PMC 7029900. PMID 32218963.
  27. ↑ Holtz, Thomas R. Jr.; Molnar, Ralph E.; Currie, Philip J. (2004). Weishampel, David B.; Dodson, Peter; Halszka, Osmólska (eds.). The Dinosauria (2nd ed.). Berkeley: University of California Press. pp. 71–110. ISBN 0-520-24209-2.
  28. 1 2 3 Lamanna, Matthew C.; Casal, Gabriel A.; Martínez, Rubén D. F.; Ibiricu, Lucio M. (2020-11-30). "Megaraptorid (Theropoda: Tetanurae) Partial Skeletons from the Upper Cretaceous Bajo Barreal Formation of Central Patagonia, Argentina: Implications for the Evolution of Large Body Size in Gondwanan MegaraptoranS". Annals of Carnegie Museum. 86 (3). doi:10.2992/007.086.0302. ISSN 0097-4463.
  29. ↑ Paul, Gregory S. (2016). The Princeton Field Guide to Dinosaurs. Princeton University Press. p. 107. ISBN 978-1-78684-190-2. OCLC 985402380.
  30. ↑ Benson, Roger B. J.; Campione, Nicolás E.; Carrano, Matthew T.; Mannion, Philip D.; Sullivan, Corwin; Upchurch, Paul; Evans, David C. (2014-05-06). "Rates of Dinosaur Body Mass Evolution Indicate 170 Million Years of Sustained Ecological Innovation on the Avian Stem Lineage". PLOS Biology. 12 (5) e1001853. Bibcode:2014PLOSB..1201853B. doi:10.1371/journal.pbio.1001853. ISSN 1545-7885. PMC 4011683. PMID 24802911. Supporting Information
  31. ↑ Benson, Roger B. J.; Xing, Xu (November 2008). "The anatomy and systematic position of the theropod dinosaur Chilantaisaurus tashuikouensis Hu, 1964 from the Early Cretaceous of Alanshan, People's Republic of China". Geological Magazine. 145 (6): 778–789. Bibcode:2008GeoM..145..778B. doi:10.1017/S0016756808005475. ISSN 1469-5081.
  32. ↑ Sereno, Paul C.; Martinez, Ricardo N.; Wilson, Jeffrey A.; Varricchio, David J.; Alcober, Oscar A.; Larsson, Hans C. E. (2008-09-30). "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. ISSN 1932-6203. PMC 2553519. PMID 18825273.
  33. ↑ Agnolin, Federico L.; Ezcurra, Martín D.; Pais, Diego F.; Salisbury, Steven W. (2010-05-24). "A reappraisal of the Cretaceous non-avian dinosaur faunas from Australia and New Zealand: evidence for their Gondwanan affinities". Journal of Systematic Palaeontology. 8 (2): 257–300. doi:10.1080/14772011003594870. ISSN 1477-2019.
  34. 1 2 Apesteguía, Sebastián; Smith, Nathan D.; Valieri, Rubén Juárez; Makovicky, Peter J. (2016-07-13). "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. ISSN 1932-6203. PMC 4943716. PMID 27410683.
  35. ↑ Zanno, Lindsay E.; Makovicky, Peter J. (2013-11-22). "Neovenatorid theropods are apex predators in the Late Cretaceous of North America". Nature Communications. 4 (1). Nature Publishing Group: 2827. doi:10.1038/ncomms3827. ISSN 2041-1723.
  36. ↑ Barker, Chris T.; Naish, Darren; Clarkin, Claire E.; Farrell, Paul; Hullmann, Gabriel; Lockyer, James; Schneider, Philipp; Ward, Robin K. C.; Gostling, Neil J. (2020). "A highly pneumatic middle Cretaceous theropod from the British Lower Greensand". Papers in Palaeontology. 6 (4): 661–679. doi:10.1002/spp2.1338. ISSN 2056-2802.
  37. 1 2 3 Benson, Roger B. J.; Carrano, Matthew T.; Brusatte, Stephen L. (2010-01-01). "A new clade of archaic large-bodied predatory dinosaurs (Theropoda: Allosauroidea) that survived to the latest Mesozoic". Naturwissenschaften. 97 (1): 71–78. doi:10.1007/s00114-009-0614-x. ISSN 1432-1904.
  38. 1 2 Ibiricu, Lucio M.; Lamanna, Matthew C.; Alvarez, Bruno N.; Cerda, Ignacio A.; Caglianone, Julieta L.; Cardozo, Noelia V.; Luna, Marcelo; Martínez, Rubén D. (2025-09-23). "Latest Cretaceous megaraptorid theropod dinosaur sheds light on megaraptoran evolution and palaeobiology". Nature Communications. 16 (1): 8298. doi:10.1038/s41467-025-63793-5. ISSN 2041-1723. PMC 12457595. PMID 40987777.
  39. 1 2 Naish, Darren; Cau, Andrea (2022-07-07). "The osteology and affinities of Eotyrannus lengi , a tyrannosauroid theropod from the Wealden Supergroup of southern England". PeerJ. 10: e12727. doi:10.7717/peerj.12727. ISSN 2167-8359. PMC 9271276. PMID 35821895.{{cite journal}}: CS1 maint: article number as page number (link)
  40. ↑ Porfiri, Juan D.; Juárez Valieri, Rubén D.; Santos, Domenica D. D.; Lamanna, Matthew C. (2018-09-01). "A new megaraptoran theropod dinosaur from the Upper Cretaceous Bajo de la Carpa Formation of northwestern Patagonia". Cretaceous Research. 89: 302–319. doi:10.1016/j.cretres.2018.03.014. ISSN 0195-6671.
  41. ↑ Aranciaga Rolando, Alexis M.; Novas, Fernando E.; Agnolín, Federico L. (2019-07-01). "A reanalysis of Murusraptor barrosaensis Coria & Currie (2016) affords new evidence about the phylogenetical relationships of Megaraptora". Cretaceous Research. 99: 104–127. doi:10.1016/j.cretres.2019.02.021. ISSN 0195-6671.
  42. ↑ Coria, Rodolfo A.; Currie, Philip J. (20 July 2016). "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. PMID 27439002.
  43. ↑ Méndez, Ariel H.; Novas, Fernando E.; Iori, Fabiano V. (May–June 2012). "Premier enregistrement de Megaraptora (Theropoda, Neovenatoridae) au Brésil" [First record of Megaraptora (Theropoda, Neovenatoridae) from Brazil]. Comptes Rendus Palevol. 11 (4): 251–256. doi:10.1016/j.crpv.2011.12.007. hdl:11336/19445.
  44. ↑ Porfiri, Juan D.; Juárez Valieri, Rubén D.; Santos, Domenica D. D.; Lamanna, Matthew C. (2018-09-01). "A new megaraptoran theropod dinosaur from the Upper Cretaceous Bajo de la Carpa Formation of northwestern Patagonia". Cretaceous Research. 89: 302–319. doi:10.1016/j.cretres.2018.03.014. ISSN 0195-6671.
  45. ↑ 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. Bibcode:2014CrRes..51...35P. doi:10.1016/j.cretres.2014.04.007. hdl:11336/12129.
  46. ↑ Ibiricu, Lucio M.; Lamanna, Matthew C.; Alvarez, Bruno N.; Cerda, Ignacio A.; Caglianone, Julieta L.; Cardozo, Noelia V.; Luna, Marcelo; Martínez, Rubén D. (2025). "Latest Cretaceous megaraptorid theropod dinosaur sheds light on megaraptoran evolution and palaeobiology". Nature Communications. 16 8298. doi:10.1038/s41467-025-63793-5. PMC 12457595.
  47. ↑ "Australian Age of Dinosaurs | Australovenator wintonensis". www.australianageofdinosaurs.com. Retrieved 2021-07-06.
  48. ↑ White, Matt A.; Cook, Alex G.; Klinkhamer, Ada J.; Elliott, David A. (2016-08-03). "The pes ofAustralovenator wintonensis(Theropoda: Megaraptoridae): analysis of the pedal range of motion and biological restoration". PeerJ. 4 e2312. Bibcode:2016PeerJ...4e2312W. doi:10.7717/peerj.2312. ISSN 2167-8359. PMC 4975041. PMID 27547591.
  49. ↑ White, Matt A.; Cook, Alex G.; Rumbold, Steven J. (2017-06-06). "A methodology of theropod print replication utilising the pedal reconstruction of Australovenator and a simulated paleo-sediment". PeerJ. 5 e3427. Bibcode:2017PeerJ...5e3427W. doi:10.7717/peerj.3427. ISSN 2167-8359. PMC 5463970. PMID 28603673.
  50. ↑ Seton, M.; Müller, R.D.; Zahirovic, S.; Gaina, C.; Torsvik, T.; Shephard, G.; Talsma, A.; Gurnis, M.; Turner, M.; Maus, S.; Chandler, M. (2012). "Global continental and ocean basin reconstructions since 200Ma". Earth-Science Reviews. 113 (3–4): 212–270. Bibcode:2012ESRv..113..212S. doi:10.1016/j.earscirev.2012.03.002.
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