Australovenator

File:Australovenator wintonensis.png

Australovenator is based on a theropod specimen (AODF 604), affectionately nicknamed "Banjo" after Banjo Paterson, which was found intermingled with the remains of the sauropod Diamantinasaurus matildae at the "Matilda site" (AODL 85). The parts of the holotype as it was initially described, which are held at the Australian Age of Dinosaurs Museum of Natural History, consists of a left dentary, teeth, partial forelimbs and hindlimbs, a partial right ilium, ribs, and gastralia. Australovenator was described in 2009 by paleontologist Scott Hocknull of the Queensland Museum and colleagues. The type species is A. wintonensis, in reference to nearby Winton. Although the holotype was first discovered in 2006 and first described in 2009, the process of excavating the "Matilda site" is still ongoing, and papers describing new elements of the holotype are still being published.

Additional arm elements of the holotype were described in 2012,{{Cite journal | last1 = White | first1 = M. A. | last2 = Cook | first2 = A. G. | last3 = Hocknull | first3 = S. A. | last4 = Sloan | first4 = T. | last5 = Sinapius | first5 = G. H. K. | last6 = Elliott | first6 = D. A. | editor1-last = Dodson | editor1-first = Peter | title = New Forearm Elements Discovered of Holotype Specimen Australovenator wintonensis from Winton, Queensland, Australia | doi = 10.1371/journal.pone.0039364 | journal = PLOS ONE | volume = 7 | issue = 6 | pages = e39364 | year = 2012 | pmid = 22761772| pmc = 3384666|bibcode = 2012PLoSO...739364W | doi-access = free }} more leg elements were described in 2013,{{Cite journal|last1=White|first1=Matt A.|last2=Benson|first2=Roger B. J.|last3=Tischler|first3=Travis R.|last4=Hocknull|first4=Scott A.|last5=Cook|first5=Alex G.|last6=Barnes|first6=David G.|last7=Poropat|first7=Stephen F.|last8=Wooldridge|first8=Sarah J.|last9=Sloan|first9=Trish|date=2013-07-24|title=New Australovenator Hind Limb Elements Pertaining to the Holotype Reveal the Most Complete Neovenatorid Leg|journal=PLOS ONE|volume=8|issue=7|pages=e68649|doi=10.1371/journal.pone.0068649|pmid=23894328|pmc=3722220|issn=1932-6203|bibcode=2013PLoSO...868649W|doi-access=free}} and a right dentary was described in 2015.{{Cite journal|last1=White|first1=Matt A.|last2=Bell|first2=Phil R.|last3=Cook|first3=Alex G.|last4=Poropat|first4=Stephen F.|last5=Elliott|first5=David A.|date=2015-12-15|title=The dentary ofAustralovenator wintonensis(Theropoda, Megaraptoridae); implications for megaraptorid dentition|journal=PeerJ|language=en|volume=3|pages=e1512|doi=10.7717/peerj.1512|pmid=26713256|issn=2167-8359|pmc=4690360 |doi-access=free }}

File:Australovenator.jpg]]

For some time, Australovenator was considered to be within the same genus as another Australian Megaraptorid called Rapator. Rapator is based on a metacarpal first described by Friedrich von Huene during the early 1900s and prior to the recognition of Megaraptora, it was considered to be either an alvarezsaurid or an intermediate theropod.{{Cite book|title=Dinosaurs: The Most Complete, Up-to-Date Encyclopedia for Dinosaur Lovers of All Ages|publisher=Random House|year=2007|isbn=9780375824197}} After the discovery of Australovenator, some scientists noted the similarity between Rapator and Australovenator. While Hocknull et al. (2009) identified a few distinguishing characters between the two taxa, based on a poorly preserved metacarpal I from the holotype of Australovenator. Agnolin et al. (2010) reclassified Rapator, instead finding it a megaraptoran, potentially a sister taxon to Australovenator. They mentioned that Megaraptor, the only other taxon also preserving metacarpal I, was less similar to Rapator than Australovenator. However, there were no clear differences between the two latter taxa. The metacarpals of both taxa were redescribed in White et al. (2014), who determined that they were not synonymous, adding multiple features to the potential characters identified by Hocknull et al. and Angolan et al.

In 2019, material from the Eumeralla Formation (Albian age) was referred to cf. Australovenator.{{Cite journal|last1=Poropat|first1=Stephen F.|last2=White|first2=Matt A.|last3=Vickers-Rich|first3=Patricia|last4=Rich|first4=Thomas H.|date=2019-07-04|title=New megaraptorid (Dinosauria: Theropoda) remains from the Lower Cretaceous Eumeralla Formation of Cape Otway, Victoria, Australia|url=https://doi.org/10.1080/02724634.2019.1666273|journal=Journal of Vertebrate Paleontology|volume=39|issue=4|pages=e1666273|doi=10.1080/02724634.2019.1666273|s2cid=208603798|issn=0272-4634}} 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.{{Cite journal|last1=White|first1=Matt A.|last2=Bell|first2=Phil R.|last3=Poropat|first3=Stephen F.|last4=Pentland|first4=Adele H.|last5=Rigby|first5=Samantha L.|last6=Cook|first6=Alex G.|last7=Sloan|first7=Trish|last8=Elliott|first8=David A.|title=New theropod remains and implications for megaraptorid diversity in the Winton Formation (lower Upper Cretaceous), Queensland, Australia|journal=Royal Society Open Science|volume=7|issue=1|pages=191462|doi=10.1098/rsos.191462|doi-access=free|pmid=32218963|pmc=7029900|year=2020|bibcode=2020RSOS....791462W}}

File:Australovenator Scale V2.svg

According to Gregory S. Paul, it was estimated at {{convert|6|m|ft|abbr=on}} long, with a body mass of {{convert|500|kg|lb}}.{{Cite book|last=Paul|first=Gregory S.|url=http://worldcat.org/oclc/985402380|title=The Princeton Field Guide to Dinosaurs|year=2016|publisher=Princeton University Press|isbn=978-1-78684-190-2|oclc=985402380|pages=107}} A 2014 study estimated its body mass to be around {{convert|310|kg|lb}}.{{Cite journal |last1=Benson |first1=Roger B. J. |last2=Campione |first2=Nicolás E. |last3=Carrano |first3=Matthew T. |last4=Mannion |first4=Philip D. |last5=Sullivan |first5=Corwin |last6=Upchurch |first6=Paul |last7=Evans |first7=David C. |date=2014-05-06 |title=Rates of Dinosaur Body Mass Evolution Indicate 170 Million Years of Sustained Ecological Innovation on the Avian Stem Lineage |journal=PLOS Biology |language=en |volume=12 |issue=5 |pages=e1001853 |doi=10.1371/journal.pbio.1001853 |issn=1545-7885 |pmc=4011683 |pmid=24802911 |doi-access=free }} [https://journals.plos.org/plosbiology/article?id=10.1371/journal.pbio.1001853#s5 Supporting Information] Because it was a relatively lightweight predator, Hocknull coined it as the "cheetah of its time".{{Cite news |url=http://www.abc.net.au/news/stories/2009/07/03/2615874.htm |archive-url=https://archive.today/20120720191633/http://www.abc.net.au/news/stories/2009/07/03/2615874.htm |url-status=dead |archive-date=July 20, 2012 |title=Scientists discover 3 new Aussie dinosaurs |last=Phillips |first=Nicky |date=2009-07-03 |work=ABC News |access-date=2018-01-11 |department=ABC Science Online |language=en-AU}} Like other megaraptorans, Australovenator would have been a bipedal carnivore.{{Cite book |title=The Dinosauria |last1=Holtz |first1=Thomas R. Jr. |last2=Molnar |first2=Ralph E. |last3=Currie |first3=Philip J. |publisher=University of California Press |year=2004 |isbn=0-520-24209-2 |editor-last=Weishampel |editor-first=David B. |edition=2nd |location=Berkeley |pages=71–110 |author-link=Thomas R. Holtz, Jr. |editor-last2=Dodson |editor-first2=Peter |editor-last3=Halszka |editor-first3=Osmólska }}

File:Australovenator dentary.png

A phylogenetic analysis found Australovenator to be an allosauroid carnosaurian, with similarities to Fukuiraptor and carcharodontosaurids. In the initial analysis, it was shown to be the sister taxon of the Carcharodontosauridae.{{Cite journal |last1=Hocknull |first1=Scott A. |last2=White |first2=Matt A. |last3=Tischler |first3=Travis R. |last4=Cook |first4=Alex G. |last5=Calleja |first5=Naomi D. |last6=Sloan |first6=Trish |last7=Elliott |first7=David A. |year=2009 |editor-last=Sereno |editor-first=Paul |editor-link=Paul Sereno |title=New Mid-Cretaceous (Latest Albian) Dinosaurs from Winton, Queensland, Australia |journal=PLOS ONE |volume=4 |issue=7 |pages=e6190 |bibcode=2009PLoSO...4.6190H |doi=10.1371/journal.pone.0006190 |pmc=2703565 |pmid=19584929|doi-access=free }} More detailed studies found that it formed a clade with several other carcharodontosaurid-like allosaurs, the Neovenatoridae.{{Cite journal |last1=Benson |first1=R.B.J. |last2=Carrano |first2=M.T |last3=Brusatte |first3=S.L. |year=2010 |title=A new clade of archaic large-bodied predatory dinosaurs (Theropoda: Allosauroidea) that survived to the latest Mesozoic |journal=Naturwissenschaften |volume=97 |issue=1 |pages=71–78 |bibcode=2010NW.....97...71B |doi=10.1007/s00114-009-0614-x |pmid=19826771|s2cid=22646156 }} Recent phylogenetic analysis suggests Australovenator is a tyrannosauroid, like with all other megaraptorans.{{Cite journal |last1=Novas |first1=F. E. |last2=Agnolín |first2=F. L. |last3=Ezcurra |first3=M. D. |last4=Canale |first4=J. I. |last5=Porfiri |first5=J. D. |year=2012 |title=Megaraptorans as members of an unexpected evolutionary radiation of tyrant-reptiles in Gondwana |url=http://www.ameghiniana.org.ar/index.php/ameghiniana/article/view/868/1618 |journal=Ameghiniana |volume=49 |issue=Suppl |pages=R33}} A phylogenetic analysis in 2016 focusing on the new neovenatorid Gualicho found that Australovenator and other megaraptorids were either allosauroids or basal coelurosaurs as opposed to being tyrannosauroids.{{Cite journal |last1=Apesteguía |first1=Sebastián |last2=Smith |first2=Nathan D. |last3=Valieri |first3=Rubén Juárez |last4=Makovicky |first4=Peter J. |date=2016-07-13 |title=An Unusual New Theropod with a Didactyl Manus from the Upper Cretaceous of Patagonia, Argentina |journal=PLOS ONE |volume=11 |issue=7 |pages=e0157793 |doi=10.1371/journal.pone.0157793 |pmid=27410683 |pmc=4943716 |issn=1932-6203|bibcode=2016PLoSO..1157793A |doi-access=free }}

The ankles of Australovenator and Fukuiraptor are similar to the Australian talus bone known as NMVP 150070 that had previously been identified as belonging to Allosaurus sp., and this bone likely represents Australovenator or a close relative of it.{{cite journal |last=Molnar |first=Ralph E. |author2=Flannery, Timothy F.|author3= Rich, Thomas H.V. |year=1981 |title=An allosaurid theropod dinosaur from the Early Cretaceous of Victoria, Australia |journal=Alcheringa |volume=5 |pages=141–146 |doi=10.1080/03115518108565427 |issue=2 }} Alternatively, this bone could belong to an abelisaur.{{Cite journal |last1=Agnolin |first1=F. L. |last2=Ezcurra |first2=M. D. |last3=Pais |first3=D. F. |last4=Salisbury |first4=S. W. |year=2010 |title=A reappraisal of the Cretaceous non-avian dinosaur faunas from Australia and New Zealand: Evidence for their Gondwanan affinities |journal=Journal of Systematic Palaeontology |volume=8 |issue=2 |pages=257–300 |doi=10.1080/14772011003594870|s2cid=130568551 }}

The cladogram below follows the 2010 analysis by Benson, Carrano, and Brusatte. Another study published later in 2010 also found the Australian theropod Rapator to be a megaraptoran extremely similar to Australovenator.{{Cite journal | last1 = Agnolin | first1 = Ezcurra | last2 = Pais | last3 = Salisbury | year = 2010 | title = A reappraisal of the Cretaceous non-avian dinosaur faunas from Australia and New Zealand: Evidence for their Gondwanan affinities | journal = Journal of Systematic Palaeontology | volume = 8 | issue = 2| pages = 257–300 | doi=10.1080/14772011003594870| s2cid = 130568551 }}

{{clade| style=font-size:100%;line-height:80%

|label1=Neovenatoridae

|1={{clade

|1=Neovenator 80px

|label2=unnamed

|2={{clade

|1=Chilantaisaurus 80px

|label2=Megaraptora

|2={{clade

|1={{clade

|1=Australovenator 80px

|2=Fukuiraptor 80px

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|2=?Orkoraptor 80px

|3={{clade

|1=Aerosteon 80px

|2=Megaraptor 80px

}}

}}

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The cladogram below follows the 2014 analysis by Porfiri et al. that finds megaraptorans to be tyrannosauroids.{{Cite journal |last1=Porfiri |first1=Juan D. |last2=Novas |first2=Fernando E. |last3=Calvo |first3=Jorge O. |last4=Agnolín |first4=Federico L. |last5=Ezcurra |first5=Martín D. |last6=Cerda |first6=Ignacio A. |year=2014 |title=Juvenile specimen of Megaraptor (Dinosauria, Theropoda) sheds light about tyrannosauroid radiation |journal=Cretaceous Research |volume=51 |pages=35–55 |doi=10.1016/j.cretres.2014.04.007|hdl=11336/12129 |hdl-access=free }}

{{clade| style=font-size:100%;line-height:80%

|label1=Megaraptora

|1={{clade

|1=Fukuiraptor 80px

|label2=Megaraptoridae

|2={{clade

|label3=unnamed

|1=Australovenator 80px

|2=Aerosteon 80px

|3=Orkoraptor 80px

|4=Eotyrannus 80px

|5=Megaraptor 80px

}}

}}

}}

File:Australovenator left manus.jpg

With very comprehensive and well-preserved hand and foot remains, 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.{{Cite journal|last1=White|first1=Matt A.|last2=Bell|first2=Phil R.|last3=Cook|first3=Alex G.|last4=Barnes|first4=David G.|last5=Tischler|first5=Travis R.|last6=Bassam|first6=Brant J.|last7=Elliott|first7=David A.|date=2015-09-14|title=Forearm Range of Motion in Australovenator wintonensis (Theropoda, Megaraptoridae)|journal=PLOS ONE|volume=10|issue=9|pages=e0137709|doi=10.1371/journal.pone.0137709|pmid=26368529|pmc=4569425|issn=1932-6203|bibcode=2015PLoSO..1037709W|doi-access=free}} The gracile morphology of the skull also concludes that this genus had a specialisation towards prey capture using its arms and hands.{{Cite web|title=Australian Age of Dinosaurs {{!}} Australovenator wintonensis|url=https://www.australianageofdinosaurs.com/page/48/australian-age-of-dinosaurs-australovenator-wintonensis|access-date=2021-07-06|website=www.australianageofdinosaurs.com}}

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.{{Cite journal|last1=White|first1=Matt A.|last2=Cook|first2=Alex G.|last3=Klinkhamer|first3=Ada J.|last4=Elliott|first4=David A.|date=2016-08-03|title=The pes ofAustralovenator wintonensis(Theropoda: Megaraptoridae): analysis of the pedal range of motion and biological restoration|journal=PeerJ|language=en|volume=4|pages=e2312|doi=10.7717/peerj.2312|pmid=27547591|issn=2167-8359|pmc=4975041 |doi-access=free }}

File:Australovenator reconstruction.jpg

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.{{Cite journal|last1=White|first1=Matt A.|last2=Cook|first2=Alex G.|last3=Rumbold|first3=Steven J.|date=2017-06-06|title=A methodology of theropod print replication utilising the pedal reconstruction of Australovenator and a simulated paleo-sediment|journal=PeerJ|language=en|volume=5|pages=e3427|doi=10.7717/peerj.3427|pmid=28603673|issn=2167-8359|pmc=5463970 |doi-access=free }}

AODL 604 was found about {{convert|60|km|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, interpreted as an oxbow lake, or billabong, deposit. 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.

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Category:Megaraptora

Category:Dinosaur genera

Category:Cenomanian dinosaurs

Category:Winton Formation

Category:Fossil taxa described in 2009

Category:Taxa named by Scott Hocknull

Category:Dinosaurs of Australia