Aussiedraco

Aussiedraco
Temporal range: Early Cretaceous,
Holotype mandible in top down (top) and oblique views
Scientific classification Edit this classification
Kingdom: Animalia
Phylum: Chordata
Class: Reptilia
Order: †Pterosauria
Suborder: †Pterodactyloidea
Clade: †Ornithocheiriformes
Clade: †Ornithocheirae
Clade: †Targaryendraconia
Family: †Targaryendraconidae
Genus: †Aussiedraco
Kellner, Rodrigues & Costa, 2011
Type species
†Aussiedraco molnari
Kellner, Rodrigues & Costa, 2011

Aussiedraco (meaning "Australian dragon") is a genus of pterosaur, an order of flying reptiles, that lived during the Early Cretaceous in what is now Australia. The genus belongs to the family Targaryendraconidae, a group of toothed pterodactyloids. It is known from a single dentary (lower jawbone) fragment that was unearthed in a limestone outcrop of the Albian-aged Toolebuc Formation. This dentary was first discovered in the 1970s in a site in western Queensland. It was initially described in 1980 by Australian paleontologists Ralph E. Molnar and Richard A. Thulborn as belonging to Ornithocheirus. Until it was recognized as coming from its own genus and species, this dentary was assigned to Ornithocheirus, Anhanguera, and Lonchodectes. In 2011, Brazilian paleontologists Alexander Kellner, Taissa Rodrigues, and Fabiana R. Costa scientifically described it as Aussiedraco molnari.

Aussiedraco was among the first pterosaurs recognized from Australia. It was assigned to the family Ornithocheiridae and as an indeterminate pteranodontoid until in 2019, Brazilian paleontologist Rubi Pêgas and colleagues moved it to Targaryendraconidae. The dentary of Aussiedraco is very unique. It lacks any crests, bears a convex surface, and has a flat ventral (bottom) surface. Based on the anatomy of its alveoli (tooth sockets) the bottom and top teeth could interlock. Aussiedraco lived alongside the other pterosaurs Mythunga, Ferrodraco, and Haliskia in the Toolebuc Formation. During the Albian the Toolebuc Formation was a near-land open ocean that had low oxygenated waters. It preserves a diverse array of plesiosaur, ichthyosaur, turtle, dinosaur, fish, shark, and invertebrate fossils.

Discovery and classification

Geologic map of Queensland, showing Bouila and outcrops of the Toolebuc Formation where QM F10613 was found

Prior to 1980, no pterosaur material had been reported form Australia in scientific literature.[1] Fossils of Aussiedraco were first unearthed in the 1970s from a flaggy limestone outcrop of the Toolebuc Formation around 70 kilometres (43 mi) east of Bouila and 13 kilometres (8.1 mi) south of the Hamilton Hotel in western Queensland, Australia. The Toolebuc Formation, a part of the larger Eromanga Basin, is composed of crystalline limestones dotted with concretions, as well as minor shale bands. This formation derives from the upper Albian stage of the Lower Cretaceous period, which dates to about 105-104 million years ago.[2][3] The remains known of Aussiedraco consist of a single, incomplete mandibular symphysis (the area of the lower jaw where the left and right mandibles meet), which is cataloged at the Queensland Museum under specimen number QM F10613. This specimen was discovered in the same block as a pterosaur vertebra. QM F10613 was the first pterosaur cranial material found in Australia.[4]

In 1980, Australian paleontologists Ralph E. Molnar and Richard A. Thulborn described QM F10613 in an article in the journal Nature. They assigned QM F10613 to Ornithocheiridae indet. aff. Ornithocheirus at a time when Ornithocheirus was a wastebasket taxon for Cretaceous toothed pterodactyloid fossils.[4] In 2000, German researcher David Unwin and colleagues reassigned QM F10613 to the anhanguerid Anhanguera? cuvieri,[5][6] which is now in the genus Cimoliopterus.[4][7] In contrast, Molnar and Thunborn (2007) opted to refer the specimen to the lonchodectid Lonchodectes.[8] In 2010, American researcher Timothy Myers followed Molnar and Thunborn (1980) by assigning the dentary to aff. Ornithocherius.[9] The constant reassignment of QM F10613 was rooted in its fragmentary nature and the wider taxonomic issues plaguing Cambridge Greensand pterosaur genera,[10][4] such as Ornithocheirus and Lonchodectes.[11]

In 2011, Brazilian paleontologists Alexander Kellner, Taissa Rodrigues, and Fabiana R. Costa scientifically described QM F10613 and identified it as belonging to a new genus and species of pternodontoid pterosaur, Aussiedraco molnari. The generic name Aussiedraco derives from Aussie, a shortened name for Australia, and the Latin word draco "dragon". The specific name is in honor of Molnar, who contributed significantly to the field of Australian vertebrate paleontology. QM F10613 was selected as the holotype (name-bearing) specimen and remains the only material assigned to the species.[11] Aussiedraco remains the only confirmed non-anhanguerian pterosaur discovered from Australia.[3][12]

Classification

Prior to the description of Aussiedraco, QM F10613 was ascribed to Ornithocheirus,[9] Anhanguera,[5] Lonchodectes,[8] and as an indeterminate ornithocheirid.[1] In their description, Kellner and colleagues assigned Aussiedraco to the Pteranodontoidea, a clade roughly containing the same species as the Ornithocheiroidea sensu Unwin. They further stated that it would be more closely allied with Anhangueridae, but would not itself be a member.[11]

Snout fossils of Ferrodraco, Mythunga, and Aussiedraco (bottom left) compared


In 2019, Brazilian paleontologist Rubi Pêgas and colleagues published a study in which they recognized a new cosmopolitan group of toothed pterodactyloids, which they termed Targaryendraconia. This order is united by the presence of a dentary symphysis with subparallel anteriorlateral (front side) margins, a laterally compressed (thin from sides) anterior (front) dentary tip, jaws widest at the third pair of alveoli (tooth sockets), teeth on over 70% of the total jaw length, a bony bar bifurcating (dividing) the first pair of alveoli, and a deep and narrow median mandibular groove (a groove in the middle of the lower jaw).[13] Outside of Targaryendraconia, the group is part of the larger clade Ornithocheiromorpha, which includes a multitude of toothed pterodactyloid families such as Anhangueridae, Istiodactylidae, and Hamipteridae.[14][15][16][17] Targaryendraconians have been reported from sites in Germany, Brazil, England, Australia, and the United States, with a range dating from the Hauterivian to the Cenomanian stages of the Cretaceous period.[13][18][19] This clade includes two families: Cimoliopteridae, which includes genera like Cimoliopterus and Aetodactylus, and Targaryendraconidae, which includes Aussiedraco itself, Targaryendraco, and Barbosania. Targaryendraconids are primarily known from fragmentary rostral or mandibular remains. The genus Saratovia may also a targaryendraconid as well, but this has not been confirmed.[19] In their phylogenetic analysis, Pêgas and colleagues (2019) recovered Aussiedraco, Barbosania, and Targaryendraco as sister taxa. This result has been replicated by other phylogenetic analyses.[20][21] Below is a cladogram following Pêgas et al. (2019). In this analysis, they recovered Aussiedraco within the family Targaryendraconidae within the larger clade Targaryendraconia.[13]

Ornithocheirae

Description

In 2019, Pêgas and colleagues abstained from giving an Aussiedraco wingspan estimate, but they estimated the wingspan of the closely related Barbosania at 2.9 to 4 meters (9.5 to 13.1 ft)

Skull reconstruction with known elements in white and unknown in grey (based on Barbosania)

.[13] Many isolated pterosaur postcranial remains have been found from the Toolebuc Formation as well, such as a scapulocoracoid (shoulder girdle), vertebrae (backbones), and wing remains. These could belong to Aussiedraco, but the lack of overlap means this is impossible to confirm until more material is known.[22][23]

The holotype consists of a three-dimensionally preserved fragment measuring about 88 millimetres (3.5 in) in length. Internally it is lightly built with a thin bone cortex, as in other pterosaurs. As far as can be judged from the empty elliptical alveoli, the lower jaws carry at least five pairs of teeth, which are rather large and become more outwards inclining and procumbent towards the front. Based on the scalloping of the alveoli, the upper and lower teeth were likely able to interlock, especially at the anterior (front) margin of the symphysis. The first through fourth alveoli are placed more dorsoventrally (up and forward), causing them to project outwards. It is distinguished from all other pterosaurs by the presence of a convex dorsal (top) mandibular margin and a straight ventral (bottom) margin. These two traits are autapomorphies (characteristics not found in other species), but it has a unique combination of synapomorphies (traits shared with other species) too such as a symphysis that is somewhat deeper than wide. The distal (away from body) end of the symphysis lacks any dorsal or lateral expansions, like a crest, unlike those of anhanguerids, ornithocheirids, and lonchodectids.[11]

The occlusal (inside) surface of the mandible was relatively curved and lacked a crest, unlike the contemporaneous Haliskia.[24] Compared to Ferrodraco, Aussiedraco's mandible is less distally expanded and less robust, with a relatively straighter left lateral surface. Unlike Ferrodraco, the anterior margin of the mandibular groove is markedly prominent and appears pronounced from the posterior boundary of the third alveolus until the posterior end of the fourth alveolus. Furthermore, Aussiedraco is distinguished from both Ferrodraco and Mythunga by its lack of raised alveolar borders (ridges around the alveoli). The mesiodistal (length of tooth going from the midline of the face to the end of the face) lengths of the frontmost two pairs of alveoli are also greater than those of Ferrodraco, indicating that the front teeth of Aussiedraco were larger than Ferrodraco's. However, this might also be an artifact of erosion. The most notably difference between the two genera is that Aussiedraco bears two sucli (folds) on its rostral mandibular tip. This characteristic is absent in all other pterosaurs, but may have been caused by over preparation of the fossils with acetic acid or may represent a split mandibular groove.[25] The only other pterosaur this is known from is Cearadactylus, a synonym of Brasileodactylus, suggesting that they may be synonyms (the same taxon).[26] However, this is unlikely as this trait has not been confirmed and the two come from different terminals, continents, and families.[25]

Paleoecology

Diagram showing the fauna of the Toolebuc Formation and other units of the Eromanga Basin

The ecology and diet of ornithocheiromorphs like Aussiedraco has been debated.[27][28][29] Although, no studies directly analyzing the diet or ecology of targaryendraconids have been published.[13] Historically, most ornithocheiromorphs were regarded as purely piscivorous, with a diet made up of small and medium fish. However, some genera like Istiodactylus possibly fed on terrestrial fauna and carrion,[30] while Coloborhynchus consumed terrestrial vertebrates and invertebrates.[31] In a 2006 study, Dutch paleontologist André J. Veldmeijer and colleagues stated ornithocheiromorphs likely had piscivorous diets due to their prevalence in aquatic habitats, interlocking teeth, and anteriorly-placed eyes. Furthermore, Veldmeijer and colleagues hypothesized that the small bodies and large heads of ornithocheiromorphs suggest that they may have lacked a capacity for giant fish. Instead, they potentially hunted small fish or pre-digested them prior to swallowing, as their teeth lacked adaptations for chewing.[32] In 2012, British paleontologist Mark Witton pointed out that the protruded, sharp teeth at the anterior tips of ornithocheiromorph jaws could have been specialized for catching prey from the water without landing. This would have lowered the risk of crashing into the water.[28]

The Toolebuc Formation was deposited in the southern edge of a late Albian Toolebuc Sea, which featured an anaerobic, relatively deep-water marine environment to the north and an aerobic, shallow-water marine and brackish environment to the south.[33] Its paleofauna of pelagic (open ocean) invertebrates like ammonites, belemnites, and coleoids and the presence of benthic invertebrates suggests that most fossils derive from a shallow near-shore marine setting with poorly oxygenated water.[2] This Toolebuc Sea was part of what now composes the Great Artesian Basin complex in the southern Tethys Ocean,[22] an ancient sea that dissipated in the early-mid Cenozoic era.[34][35] The Toolebuc Formation preserves most of Australia's pterosaur fossils, including those of the anhanguerians Haliskia, Mythunga, Thapunngaka, and an indeterminate taxon. Being an oceanic deposit, the Toolebuc Formation bears an array of marine reptile fossils including from the plesiosaurs Kronosaurus and Eromangasaurus, ichthyosaur Platypterygius, and the sea turtles Bouilachelys, Cratochelone, and Notochelone. Three dinosaurs, the parankylosaur Kunbarrasaurus,[36] an unnamed sauropod,[37] and the enantiornithean bird Nanantius have been reported as well.[38] Many fish fossils have been discovered, such as those of the pachycormiform Australopachycormus, the ichthyodectid Cooyoo, the aspidorhyhnchid Richmondichthys, and the shark Pseudocorax.[39]

See also

References

  1. 1 2 Molnar, Ralph E.; Thulborn, Richard A. (1980). "First pterosaur from Australia". Nature. 288 (5789): 361–363. Bibcode:1980Natur.288..361M. doi:10.1038/288361a0. ISSN 1476-4687.
  2. 1 2 Frauenfelder, Timothy G.; Bell, Phil R.; Brougham, Tom; Bevitt, Joseph J.; Bicknell, Russell D. C.; Kear, Benjamin P.; Wroe, Stephen; Campione, Nicolás E. (2022-03-28). "New Ankylosaurian Cranial Remains From the Lower Cretaceous (Upper Albian) Toolebuc Formation of Queensland, Australia". Frontiers in Earth Science. 10 803505. Bibcode:2022FrEaS..1003505F. doi:10.3389/feart.2022.803505. ISSN 2296-6463.
  3. 1 2 Richards, Timothy M.; Stumkat, Paul E.; Salisbury, Steven W. (2021). "A New Species of Crested Pterosaur (pterodactyloidea, Anhangueridae) from the Lower Cretaceous (upper Albian) of Richmond, North West Queensland, Australia". Journal of Vertebrate Paleontology. 41 (3) e1946068: 1–14. Bibcode:2021JVPal..41E6068R. doi:10.1080/02724634.2021.1946068. ISSN 0272-4634. JSTOR 27122723.
  4. 1 2 3 4 Rodrigues, Taissa; Kellner, Alexander Wilhelm Armin (2013). "Taxonomic review of the Ornithocheirus complex (Pterosauria) from the Cretaceous of England". ZooKeys (308): 1–112. Bibcode:2013ZooK..308....1R. doi:10.3897/zookeys.308.5559. ISSN 1313-2989. PMC 3689139. PMID 23794925.
  5. 1 2 Unwin, D. M.; Lü, J.; Bakhurina, N. N. (2000-01-01). "On the systematic and stratigraphic significance of pterosaurs from the Lower Cretaceous Yixian Formation (Jehol Group) of Liaoning, China". Fossil Record. 3 (1): 181–206. Bibcode:2000FossR...3..181U. doi:10.1002/mmng.20000030109. ISSN 2193-0066.
  6. ↑ Unwin, David M. (2008-04-22). "An overview of the pterosaur assemblage from the Cambridge Greensand (Cretaceous) of Eastern England". Fossil Record. 4 (1): 189–221. doi:10.1002/mmng.20010040112.
  7. ↑ Myers, Timothy S. (2015-11-02). "First North American occurrence of the toothed pteranodontoid pterosaur Cimoliopterus". Journal of Vertebrate Paleontology. 35 (6) e1014904. Bibcode:2015JVPal..35E4904M. doi:10.1080/02724634.2015.1014904. ISSN 0272-4634.
  8. 1 2 Molnar, Ralph E.; Thulborn, Richard A. (2007). "An incomplete pterosaur skull from the Cretaceous of North-Central Queensland, Australia". Archivos do Museu Nacional do Rio de Janeiro (in Portuguese). 65 (4). Archived from the original on 2025-08-05.
  9. 1 2 Myers, Timothy S. (2010-01-29). "A new ornithocheirid pterosaur from the Upper Cretaceous (Cenomanian–Turonian) Eagle Ford Group of Texas". Journal of Vertebrate Paleontology. 30 (1): 280–287. Bibcode:2010JVPal..30..280M. doi:10.1080/02724630903413099. ISSN 0272-4634.
  10. ↑ Unwin, David M. (2001). "An overview of the pterosaur assemblage from the Cambridge Greensand (Cretaceous) of Eastern England". Fossil Record. 4 (1): 189–221. Bibcode:2001FossR...4..189U. doi:10.1002/mmng.20010040112. ISSN 1860-1014.
  11. 1 2 3 4 Kellner, Alexander W.A.; Taissa Rodrigues; Fabiana R. Costa (2011). "Short note on a pteranodontoid pterosaur (Pterodactyloidea) from western Queensland, Australia" (PDF). Anais da Academia Brasileira de Ciências. 83 (1): 301–308. Bibcode:2011AnABC..83..301K. doi:10.1590/S0001-37652011000100018. PMID 21437387.
  12. ↑ Poropat, Stephen F., Bell, Phil R., Hart, Lachlan J., Salisbury, Steven W., & Kear, Benjamin P. (2023). An annotated checklist of Australian Mesozoic tetrapods. Alcheringa: An Australasian Journal of Palaeontology, 47(2), 129–205. https://doi.org/10.1080/03115518.2023.2228367
  13. 1 2 3 4 5 Rubi V. Pêgas, Borja Holgado & Maria Eduarda C. Leal (2019) On Targaryendraco wiedenrothi gen. nov. (Pterodactyloidea, Pteranodontoidea, Lanceodontia) and recognition of a new cosmopolitan lineage of Cretaceous toothed pterodactyloids, Historical Biology, doi:10.1080/08912963.2019.1690482
  14. ↑ Kellner, Alexander W. A.; Caldwell, Michael W.; Holgado, Borja; Vecchia, Fabio M. Dalla; Nohra, Roy; Sayão, Juliana M.; Currie, Philip J. (2019). "First complete pterosaur from the Afro-Arabian continent: Insight into pterodactyloid diversity". Scientific Reports. 9 (1): 17875. Bibcode:2019NatSR...917875K. doi:10.1038/s41598-019-54042-z. PMC 6884559. PMID 31784545.
  15. ↑ Andres, B. (2021) Phylogenetic systematics of Quetzalcoatlus Lawson 1975 (Pterodactyloidea: Azhdarchoidea). Journal of Vertebrate Paleontology, 41:sup1, 203-217. DOI: 10.1080/02724634.2020.1801703 https://www.tandfonline.com/doi/full/10.1080/02724634.2020.1801703
  16. ↑ Longrich, N.R., Martill, D.M., and Andres, B. (2018). "Late Maastrichtian pterosaurs from North Africa and mass extinction of Pterosauria at the Cretaceous-Paleogene boundary." PLoS Biology, 16(3): e2001663. doi:10.1371/journal.pbio.2001663
  17. ↑ Holgado, B.; Pêgas, R.V. (2020). "A taxonomic and phylogenetic review of the anhanguerid pterosaur group Coloborhynchinae and the new clade Tropeognathinae". Acta Palaeontologica Polonica. 65. doi:10.4202/app.00751.2020. hdl:2072/444907.
  18. ↑ Pêgas, R. V. (2025-06-18). "On the systematics and phylogenetic nomenclature of the Ornithocheiriformes (Pterosauria, Pteranodontoidea)". Palaeontologia Electronica. 25 (2): 1–21. doi:10.26879/20. ISSN 1094-8074.
  19. 1 2 Averianov, Alexander O. (2025). "A new ornithocheiran pterosaur from the Upper Cretaceous (Cenomanian) of Saratov, Russia". Anais da Academia Brasileira de Ciências. 97 (Suppl. 1) e20241063. doi:10.1590/0001-3765202520241063. ISSN 0001-3765. PMID 40298668.
  20. ↑ Pentland, Adele H.; Poropat, Stephen F.; White, Matt A.; Rigby, Samantha L.; Bevitt, Joseph J.; Duncan, Ruairidh J.; Sloan, Trish; Elliott, Robert A.; Elliott, Harry A.; Elliott, Judy A.; Elliott, David A. (2021-09-01). "The osteology of Ferrodraco lentoni, an anhanguerid pterosaur from the mid-Cretaceous of Australia". Journal of Vertebrate Paleontology. 41 (5) e2038182: 1. Bibcode:2021JVPal..41E8182P. doi:10.1080/02724634.2021.2038182. ISSN 0272-4634.
  21. ↑ Richards, Timothy M.; Stumkat, Paul E.; Salisbury, Steven W. (2024-02-01). "A second specimen of the pterosaur Thapunngaka shawi from the Lower Cretaceous (upper Albian) Toolebuc Formation of North West Queensland, Australia". Cretaceous Research. 154 105740. Bibcode:2024CrRes.15405740R. doi:10.1016/j.cretres.2023.105740. ISSN 0195-6671.
  22. 1 2 Kellner, Alexander W.A.; Rich, Thomas H.; Costa, Fabiana R.; Vickers-Rich, Patricia; Kear, Benjamin P.; Walters, Mary; Kool, Lesley (2010-09-01). "New isolated pterodactyloid bones from the Albian Toolebuc Formation (western Queensland, Australia) with comments on the Australian pterosaur fauna". Alcheringa: An Australasian Journal of Palaeontology. 34 (3): 219–230. Bibcode:2010Alch...34..219K. doi:10.1080/03115511003656552. ISSN 0311-5518.
  23. ↑ Pentland, Adele H.; Poropat, Stephen F.; White, Matt A.; Rigby, Samantha L.; Vickers-Rich, Patricia; Rich, Thomas H.; Elliott, David A. (2022-04-03). "New anhanguerian pterosaur remains from the Lower Cretaceous of Queensland, Australia". Alcheringa: An Australasian Journal of Palaeontology. 46 (2): 188–197. Bibcode:2022Alch...46..188P. doi:10.1080/03115518.2022.2065028. ISSN 0311-5518.
  24. ↑ Pentland, Adele H.; Poropat, Stephen F.; Duncan, Ruairidh J.; Kellner, Alexander W. A.; Bantim, Renan A. M.; Bevitt, Joseph J.; Tait, Alan M.; Grice, Kliti (2024-06-12). "Haliskia peterseni, a new anhanguerian pterosaur from the late Early Cretaceous of Australia". Scientific Reports. 14 (1): 11789. Bibcode:2024NatSR..1411789P. doi:10.1038/s41598-024-60889-8. ISSN 2045-2322. PMC 11169243. PMID 38866826.
  25. 1 2 Pêgas, R. V.; Nascimento, Anderson S.; Piazentin, Lucas C.; Pinheiro, Felipe L.; Zaher, Hussam; Costa, Fabiana R. (2026-08-03). "Untangling the identity of Romualdo pterosaurs: 'Cearadactylus atrox' as a junior synonym of Brasileodactylus araripensis (Pterosauria, Anhangueridae)". Historical Biology. 38 (8): 2587–2597. Bibcode:2026HBio...38.2587P. doi:10.1080/08912963.2025.2582068. ISSN 0891-2963.
  26. ↑ Pentland, Adele H.; Poropat, Stephen F.; Tischler, Travis R.; Sloan, Trish; Elliott, Robert A.; Elliott, Harry A.; Elliott, Judy A.; Elliott, David A. (2019-10-03). "Ferrodraco lentoni gen. et sp. nov., a new ornithocheirid pterosaur from the Winton Formation (Cenomanian–lower Turonian) of Queensland, Australia". Scientific Reports. 9 (1): 13454. Bibcode:2019NatSR...913454P. doi:10.1038/s41598-019-49789-4. ISSN 2045-2322. PMC 6776501. PMID 31582757.
  27. ↑ Wellnhofer, P. (1991). The Illustrated Encyclopedia of Pterosaurs. New York: Crescent Books. pp. 114–116. ISBN 978-0-517-03701-0.
  28. 1 2 Veldmeijer, A. J.; Witton, M.; Nieuwland, I. (2012). Pterosaurs: Flying Contemporaries of the Dinosaurs. Leiden: Sidestone Press. pp. 81–83, 98. ISBN 978-90-8890-093-8.
  29. ↑ Ősi, A. (2011). "Feeding-related characters in basal pterosaurs: implications for jaw mechanism, dental function and diet" (PDF). Lethaia. 44 (2): 136–152. Bibcode:2011Letha..44..136O. doi:10.1111/j.1502-3931.2010.00230.x. hdl:10831/74599. Archived (PDF) from the original on 10 November 2020. Retrieved 6 August 2018.
  30. ↑ Witton, M. P. (2012). "New Insights into the Skull of Istiodactylus latidens (Ornithocheiroidea, Pterodactyloidea)". PLOS ONE. 7 (3) e33170. Bibcode:2012PLoSO...733170W. doi:10.1371/journal.pone.0033170. PMC 3310040. PMID 22470442.
  31. ↑ Myers, Timothy S. (2017). "Diet of ornithocheiroid pterosaurs inferred from stable Carbon isotope analysis of tooth enamel". Geological Society of America Annual Meeting in Seattle, Washington, USA. 49 (6). doi:10.1130/abs/2017AM-305496.
  32. ↑ Veldmeijer, A. J.; Signore, M.; Bucci, E. (2006). "Predator-prey interaction of Brazilian Cretaceous toothed pterosaurs: A case example". In Elewa, Ashraf M. T. (ed.). Predation in Organisms: A Distinct Phenomenon. pp. 295–308. doi:10.1007/978-3-540-46046-6_13. ISBN 978-3-540-46044-2.
  33. ↑ Ozimic, S. (1982). "Depositional environment of the Toolebuc Formation and its equivalents, Eromanga Basin, Australia". Petroleum Exploration Society of Australia (PESA). Eromanga Basin Symposium (1982): 200–203.
  34. ↑ Bialik, Or M.; Frank, Martin; Betzler, Christian; Zammit, Ray; Waldmann, Nicolas D. (2019-06-20). "Two-step closure of the Miocene Indian Ocean Gateway to the Mediterranean". Scientific Reports. 9 (1): 8842. Bibcode:2019NatSR...9.8842B. doi:10.1038/s41598-019-45308-7. ISSN 2045-2322. PMC 6586870. PMID 31222018.
  35. ↑ Steininger, F.F.; Wessely, G. (2000). "From the Tethyan Ocean to the Paratethys Sea: Oligocene to Neogene stratigraphy, paleogeography and paleobiogeography of the circum-Mediterranean region and the Oligocene to Neogene Basin evolution in Austria". Mitteilungen der Österreichischen Geologischen Gesellschaft. 92: 95–116.
  36. ↑ Frauenfelder, Timothy G.; Bell, Phil R.; Brougham, Tom; Bevitt, Joseph J.; Bicknell, Russell D. C.; Kear, Benjamin P.; Wroe, Stephen; Campione, Nicolás E. (2022-03-28). "New Ankylosaurian Cranial Remains From the Lower Cretaceous (Upper Albian) Toolebuc Formation of Queensland, Australia". Frontiers in Earth Science. 10 803505. Bibcode:2022FrEaS..1003505F. doi:10.3389/feart.2022.803505. ISSN 2296-6463.
  37. ↑ Poropat, S.F.; Nair, J.P.; Syme, C.E.; Mannion, P.D.; Upchurch, P.; Hocknull, S.A.; Cook, A.G.; Tischler, T.R.; Holland, T. (2017). "Reappraisal of Austrosaurus mckillopi Longman, 1933 from the Allaru Mudstone of Queensland, Australia's first named Cretaceous sauropod dinosaur" (PDF). Alcheringa. 41 (4): 543–580. Bibcode:2017Alch...41..543P. doi:10.1080/03115518.2017.1334826. hdl:10044/1/48659. S2CID 134237391.
  38. ↑ Kurochkin, Evgeny N.; Molnar, Ralph E. (1997-01-01). "New material of enantiornithine birds from the Early Cretaceous of Australia". Alcheringa: An Australasian Journal of Palaeontology. 21 (4): 291–297. Bibcode:1997Alch...21..291K. doi:10.1080/03115519708619169. ISSN 0311-5518.
  39. ↑ Rozefelds, Andrew C. (1993-01-01). "Lower Cretaceous Anacoracidae? (Lamniformes: Neoselachii); vertebrae and associated dermal scales from Australia". Alcheringa: An Australasian Journal of Palaeontology. 17 (3): 199–210. Bibcode:1993Alch...17..199R. doi:10.1080/03115519308619604. ISSN 0311-5518.
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