Stephen F. Poropat

New Australian sauropods shed light on Cretaceous dinosaur palaeobiogeography

Australian dinosaurs have played a rare but controversial role in the debate surrounding the effect of Gondwanan break-up on Cretaceous dinosaur distribution. Major spatiotemporal gaps in the Gondwanan Cretaceous fossil record, coupled with taxon incompleteness, have hindered research on this effect, especially in Australia. Here we report on two new sauropod specimens from the early Late Cretaceous of Queensland, Australia, that have important implications for Cretaceous dinosaur palaeobiogeography. Savannasaurus elliottorum gen. et sp. nov. comprises one of the most complete Cretaceous sauropod skeletons ever found in Australia, whereas a new specimen of Diamantinasaurus matildae includes the first ever cranial remains of an Australian sauropod. The results of a new phylogenetic analysis, in which both Savannasaurus and Diamantinasaurus are recovered within Titanosauria, were used as the basis for a quantitative palaeobiogeographical analysis of macronarian sauropods. Titanosaurs achieved a worldwide distribution by at least 125 million years ago, suggesting that mid-Cretaceous Australian sauropods represent remnants of clades which were widespread during the Early Cretaceous. These lineages would have entered Australasia via dispersal from South America, presumably across Antarctica. High latitude sauropod dispersal might have been facilitated by Albian–Turonian warming that lifted a palaeoclimatic dispersal barrier between Antarctica and South America.

New Australovenator Hind Limb Elements Pertaining to the Holotype Reveal the Most Complete Neovenatorid Leg

We report new skeletal elements pertaining to the same individual which represents the holotype of Australovenator wintonensis, from the ‘Matilda Site’ in the Winton Formation (Upper Cretaceous) of western Queensland. The discovery of these new elements means that the hind limb of Australovenator is now the most completely understood hind limb among Neovenatoridae. The new hind limb elements include: the left fibula; left metatarsal IV; left pedal phalanges I-2, II-1, III-4, IV-2, IV-3; and right pedal phalanges, II-2 and III-1. The detailed descriptions are supported with three dimensional figures. These coupled with the completeness of the hind limb will increase the utility of Australovenator in comparisons with less complete neovenatorid genera. These specimens and the previously described hind limb elements of Australovenator are compared with other theropods classified as neovenatorids (including Neovenator, Chilantaisaurus, Fukuiraptor, Orkoraptor and Megaraptor). Hind limb length proportion comparisons indicate that the smaller neovenatorids Australovenator and Fukuiraptor possess more elongate and gracile hind limb elements than the larger Neovenator and Chilantaisaurus. Greater stride lengths to body size exist in both Fukuiraptor and Australovenator with the femur discovered to be proportionally shorter the rest of the hind limb length. Additionally Australovenator is identified as possessing the most elongate metatarsus. The metatarsus morphology varies with body size. The larger neoventorids possess a metatarsus with greater width but shorter length compared to smaller forms.

Photographic Atlas and three-dimensional reconstruction of the holotype skull of Euhelopus zdanskyi with description of additional cranial elements.

Background Euhelopus zdanskyi is one of relatively few sauropod taxa known from an almost complete skull and mandible. Recent phylogenetic analyses suggest that Euhelopus is a somphospondylan titanosauriform, and that it is a member of the clade (Euhelopodidae) which is the sister taxon to the hugely successful, dominantly Cretaceous sauropod group Titanosauria. Methodology/Principal Findings The skull elements of Euhelopus were CT scanned at Uppsala Akademiska Sjukhuset. Three-dimensional models of the elements were constructed from the DICOM data using Mimics 14.0, InVesalius 3.0, and GeoMagic Studio 2012, the skull was rearticulated in Rhinoceros 4.0, and the final version was rendered in GeoMagic Studio 2012. Conclusions/Significance The fact that relatively complete sauropod skulls are so rare in the fossil record, particularly among titanosauriforms, means that the skulls that are known should be as thoroughly described and well-illustrated as possible. This contribution supplements previous descriptions of the cranial elements of Euhelopus, one of the few euhelopodid taxa for which cranial material is known, by presenting a comprehensive photographic atlas of the skull elements to facilitate a better understanding of their morphology. We describe several elements which have been overlooked in past studies of Euhelopus, and also provide as accurate a reconstruction of the skull as possible (in the absence of the braincase), the most significant components of which are the articulations of the palate and the mandible.

The dentary of Australovenator wintonensis (Theropoda, Megaraptoridae); implications for megaraptorid dentition

Megaraptorid theropods were an enigmatic group of medium-sized predatory dinosaurs, infamous for the hypertrophied claw on the first manual digit. Megaraptorid dentition is largely restricted to isolated teeth found in association with skeletal parts; however, the in situ maxillary dentition of Megaraptor was recently described. A newly discovered right dentary pertaining to the Australovenator holotype preserves in situ dentition, permitting unambiguous characterisation of the dentary tooth morphology. The new jaw is virtually complete, with an overall elongate, shallow profile, and fifteen visible in situ teeth at varying stages of eruption. In situ teeth confirm Australovenator exhibited modest pseudoheterodonty, recurved lateral teeth with a serrate distal carina and reduced mesial carina, similar to other megaraptorids. Australovenator also combines of figure-of-eight basal cross-section with a lanceolate shape due to the presence of labial and lingual depressions and the lingual twist of the distal carina. Computed tomography and three-dimensional imagery provided superior characterisation of the dentary morphology and enabled an accurate reconstruction to a pre-fossilised state. The newly established dental morphology also afforded re-evaluation of isolated theropod teeth discovered at the Australovenator holotype locality and from several additional Winton Formation localities. The isolated Winton teeth are qualitatively and quantitatively similar to the in situ dentary teeth of Australovenator, but are also morphometrically similar to Abelisauridae, Allosauridae, Coelophysoidea, Megalosauridae and basal Tyrannosauroidea. Qualitative characters, however, clearly distinguish the teeth of Australovenator and the isolated Winton teeth from all other theropods. Evidence from teeth suggests megaraptorids were the dominant predators in the Winton Formation, which contrasts with other penecontemporaneous Gondwanan ecosystems.

Late Triassic capitosaurian remains from Svalbard and the palaeobiogeographical context of Scandinavian Arctic temnospondyls

Abstract The Norwegian Arctic Svalbard archipelago is famous for its prolific record of Early–Middle Triassic vertebrate fossils. These represent mainly marine amniotes, together with sharks, bony fishes and temnospondyl amphibians, the latter providing an important faunal correlate with coeval assemblages from the Danish autonomous region of Greenland. However, substantial biostratigraphical gaps exist in the Upper Triassic strata of Svalbard, which are marked by pronounced facies shifts from marine to deltaic systems and intermittent depositional hiatuses. These are accompanied by a dearth of documented vertebrate remains, a notable exception being the partial skull of the capitosaurian Capitosaurus polaris and a few isolated stereospondylian intercentra probably from the middle–late Carnian De Geerdalen Formation of Spitsbergen. Reassessment of this material, which incorporates the only undisputed capitosaurian fossil from Svalbard, indicates affinity with Cyclotosaurus, known elsewhere from the late Norian–early Rhaetian Fleming Fjord Formation of Greenland. The Scandinavian Arctic temnospondyls constituted components of sympatric assemblages that inhabited the Boreal margin of Pangaea throughout the Triassic.

Late Cretaceous dinosaurian remains from the Kristianstad Basin of southern Sweden

Abstract Mesozoic dinosaur fossils are exceptionally rare in Scandinavia. The Swedish record is typically depauperate, with the Kristianstad Basin of Skåne (Scania) yielding all of the known fossils from Swedish Cretaceous strata. Although highly fragmentary, these body remnants are important because they provide evidence of a relatively diverse fauna, including previously recognized hesperornithiform birds and leptoceratopsid ceratopsians, as well as indeterminate ornithopods that are confirmed here for the first time. In this paper, we describe three phalanges (from Åsen) and an incomplete right tibia (from Ugnsmunnarna) from the Kristianstad Basin. One of the phalanges appears to pertain to a leptoceratopsid ceratopsian, providing further evidence of these small ornithischians in the Cretaceous sediments of Sweden. The other two phalanges are interpreted as deriving from small ornithopods similar to Thescelosaurus and Parksosaurus. The tibia appears to represent the first evidence of a non-avian theropod dinosaur in the Cretaceous of Sweden, with a previous report of theropod remains based on fish teeth having been corrected by other authors. The remains described herein provide important additions to the enigmatic dinosaurian fauna that inhabited the Fennoscandian archipelago during the latest Cretaceous.

Reappraisal of Austrosaurus mckillopi Longman, 1933 from the Allaru Mudstone of Queensland, Australia’s first named Cretaceous sauropod dinosaur

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. XX.XXXX. 2017. Reappraisal of Austrosaurus mckillopi Longman, 1933 from the Allaru Mudstone of Queensland, Australia’s first named Cretaceous sauropod dinosaur. Alcheringa 41, 543–580. ISSN 0311-5518 Austrosaurus mckillopi was the first Cretaceous sauropod reported from Australia, and the first Cretaceous dinosaur reported from Queensland (northeast Australia). This sauropod taxon was established on the basis of several fragmentary presacral vertebrae (QM F2316) derived from the uppermost Lower Cretaceous (upper Albian) Allaru Mudstone, at a locality situated 77 km west-northwest of Richmond, Queensland. Prior to its rediscovery in 2014, the type site was considered lost after failed attempts to relocate it in the 1970s. Excavations at the site in 2014 and 2015 led to the recovery of several partial dorsal ribs and fragments of presacral vertebrae, all of which clearly pertained to a single sauropod dinosaur. The discovery of new material of the type individual of Austrosaurus mckillopi, in tandem with a reassessment of the material collected in the 1930s, has facilitated the rearticulation of the specimen. The resultant vertebral series comprises six presacral vertebrae—the posteriormost cervical and five anteriormost dorsals—in association with five left dorsal ribs and one right one. The fragmentary nature of the type specimen has historically hindered assessments of the phylogenetic affinities of Austrosaurus, as has the fact that these evaluations were often based on a subset of the type material. The reappraisal of the type series of Austrosaurus presented herein, on the basis of both external morphology and internal morphology visualized through CT data, validates it as a diagnostic titanosauriform taxon, tentatively placed in Somphospondyli, and characterized by the possession of an accessory lateral pneumatic foramen on dorsal vertebra I (a feature that appears to be autapomorphic) and by the presence of a robust ventral mid-line ridge on the centra of dorsal vertebrae I and II. The interpretation of the anteriormost preserved vertebra in Austrosaurus as a posterior cervical has also prompted the re-evaluation of an isolated, partial, posterior cervical vertebra (QM F6142, the ‘Hughenden sauropod’) from the upper Albian Toolebuc Formation (which underlies the Allaru Mudstone). Although this vertebra preserves an apparent unique character of its own (a spinopostzygapophyseal lamina fossa), it is not able to be referred unequivocally to Austrosaurus and is retained as Titanosauriformes indet. Austrosaurus mckillopi is one of the oldest known sauropods from the Australian Cretaceous based on skeletal remains and potentially provides phylogenetic and/or palaeobiogeographic context for later taxa such as Wintonotitan wattsi, Diamantinasaurus matildae and Savannasaurus elliottorum. Stephen F. Poropat* [sporopat@swin.edu.au; stephenfporopat@gmail.com] Department of Chemistry and Biotechnology, Swinburne University of Technology, John St, Hawthorn, Victoria 3122, Australia; Jay P. Nair [j.nair@uq.edu.au; jayraptor@gmail.com] School of Biological Sciences, The University of Queensland, St Lucia, Queensland 4072, Australia; Caitlin E. Syme [caitlin.syme@uqconnect.edu.au] School of Biological Sciences, The University of Queensland, St Lucia, Queensland 4072, Australia; Philip D. Mannion [philipdmannion@gmail.com] Department of Earth Science and Engineering, Imperial College London, South Kensington Campus, London SW7 2AZ, UK; Paul Upchurch [p.upchurch@ucl.ac.uk] Department of Earth Sciences, University College London, Gower Street, London WC1E 6BT, UK; Scott A. Hocknull [scott.hocknull@qm.qld.gov.au] Geosciences, Queensland Museum, 122 Gerler Rd, Hendra, Queensland 4011, Australia; Alex G. Cook [alex.cook@y7mail.com] School of Earth Sciences, The University of Queensland, St Lucia, Queensland 4072, Australia; Travis R. Tischler [travisr.tischler@outlook.com] Australian Age of Dinosaurs Museum of Natural History, Lot 1 Dinosaur Drive, PO Box 408, Winton, Queensland 4735, Australia; Timothy Holland [drtimothyholland@gmail.com] Kronosaurus Korner, 91 Goldring St, Richmond, Queensland 4822, Australia. *Also affiliated with: Australian Age of Dinosaurs Museum of Natural History, Lot 1 Dinosaur Drive, PO Box 408, Winton, Queensland 4735, Australia.

Oldest meiolaniid turtle remains from Australia: evidence from the Eocene Kerosene Creek Member of the Rundle Formation, Queensland

Poropat, S.F., Kool, L., Vickers-Rich, P. & Rich, T.H., September 2016. Oldest meiolaniid turtle remains from Australia: evidence from the Eocene Kerosene Creek Member of the Rundle Formation, Queensland. Alcheringa 41, XX–XX. ISSN 0311-5518. Fossil meiolaniid turtles are known only from South America and Australasia. The South American record is restricted to the Eocene, and comprises two genera: Niolamia and Gaffneylania. The Australasian meiolaniid record is more diverse, with three genera known (Ninjemys, Warkalania and Meiolania); however, the oldest known specimens from this continent are significantly younger than those from South America, deriving from upper Oligocene sediments in South Australia and Queensland. Herein, we describe the oldest meiolaniid remains found in Australasia to date. The specimens comprise a posterior peripheral, a caudal ring, and an osteoderm, all of which derive from the middle–upper Eocene Rundle Formation of The Narrows Graben, Gladstone, eastern Queensland. Despite their fragmentary nature, each of these specimens can be assigned to Meiolaniidae with a high level of confidence. This is particularly true of the partial caudal ring, which is strongly similar to those of Niolamia, Ninjemys and Meiolania. The extension of the Australasian meiolaniid record to the Eocene lends strong support to the hypothesis that these turtles arose before South America and Australia detached from Antarctica, and that they were consequently able to spread across all three continents. Stephen F. Poropat*† [stephenfporopat@gmail.com], Australian Age of Dinosaurs Natural History Museum, The Jump-Up, Winton, Queensland 4735, Australia; Lesley Kool*† [koollesley@gmail.com] and Thomas H. Rich [trich@museum.vic.gov.au], Melbourne Museum, 11 Nicholson St, Carlton, Victoria 3053, Australia; Patricia Vickers-Rich [pat.rich@monash.edu], Monash University, Wellington Rd, Clayton, Victoria 3800, Australia. *These authors contributed equally to this work. †Also affiliated with Monash University, Wellington Rd, Clayton, Victoria 3800, Australia.

Three-Dimensional Musculoskeletal Modeling of the Sauropodomorph Hind Limb: The Effect of Postural Change on Muscle Leverage: Muscle modeling of sauropodomorph hind limbs

The biomechanical constraints for life at massive size can be directly observed in the evolutionary history of sauropodomorph dinosaurs. Members of this lineage underwent a number of major postural transitions as they increased in size from relatively small bipedal dinosaurs to massive titanosaurs that include the largest terrestrial animals of all time. To better understand the impact of gigantic size on the biomechanics of sauropods, we used three‐dimensional musculoskeletal modeling to investigate how hind limb musculature was affected, first by the development of a quadrupedal stance from a bipedal one, and later in the transition from a narrow‐gauge to a wide‐gauge stance. Muscle moment arms were measured in four sauropodomorph taxa: the bipedal basal sauropodomorph Plateosaurus engelhardti, the narrow‐gauge diplodocid Diplodocus carnegii, the titanosauriform Giraffatitan brancai, and the wide‐gauge titanosaur Diamantinasaurus matildae. In Plateosaurus, low moment arm leverage in the hip extensors and knee flexors and extensors was observed suggesting high‐velocity movement for fast locomotion. A reduction in hip extensor leverage in Diamantinasaurus was found which suggests a reduced role for the hind limb in forward propulsion in titanosaurs. An increase in overall hip adductor leverage and leverage of adductors 1 and 2 in Diamantinasaurus, compared with other taxa studied, might relate to the development of a wide‐gauge stance. High knee flexor–extensor leverage in Giraffatitan but not Diamantinasaurus partially refutes the idea that broader femoral condyles in titanosauriforms increased knee torque production capabilities. Sauropodomorph postural changes clearly had an impact on the function and leverage of hind limb muscles. Anat Rec, 301:2145–2163, 2018.

Early Cretaceous polar biotas of Victoria, southeastern Australia - an overview of research to date

Poropat, S.F., Martin, S.K., Tosolini, A.-M.P., Wagstaff, B.E, Bean, L.B., Kear, B.P., Vickers-Rich, P. & Rich, T.H., May 2018. Early Cretaceous polar biotas of Victoria, southeastern Australia—an overview of research to date. Alcheringa 42, 158–230. ISSN 0311-5518. Although Cretaceous fossils (coal excluded) from Victoria, Australia, were first reported in the 1850s, it was not until the 1950s that detailed studies of these fossils were undertaken. Numerous fossil localities have been identified in Victoria since the 1960s, including the Koonwarra Fossil Bed (Strzelecki Group) near Leongatha, the Dinosaur Cove and Eric the Red West sites (Otway Group) at Cape Otway, and the Flat Rocks site (Strzelecki Group) near Cape Paterson. Systematic exploration over the past five decades has resulted in the collection of thousands of fossils representing various plants, invertebrates and vertebrates. Some of the best-preserved and most diverse Hauterivian–Barremian floral assemblages in Australia derive from outcrops of the lower Strzelecki Group in the Gippsland Basin. The slightly younger Koonwarra Fossil Bed (Aptian) is a Konservat-Lagerstätte that also preserves abundant plants, including one of the oldest known flowers. In addition, insects, crustaceans (including the only syncaridans known from Australia between the Triassic and the present), arachnids (including Australia’s only known opilione), the stratigraphically youngest xiphosurans from Australia, bryozoans, unionoid molluscs and a rich assemblage of actinopterygian fish are known from the Koonwarra Fossil Bed. The oldest known—and only Mesozoic—fossil feathers from the Australian continent constitute the only evidence for tetrapods at Koonwarra. By contrast, the Barremian–Aptian-aged deposits at the Flat Rocks site, and the Aptian–Albian-aged strata at the Dinosaur Cove and Eric the Red West sites, are all dominated by tetrapod fossils, with actinopterygians and dipnoans relatively rare. Small ornithopod (=basal neornithischian) dinosaurs are numerically common, known from four partial skeletons and a multitude of isolated bones. Aquatic meiolaniform turtles constitute another prominent faunal element, represented by numerous isolated bones and articulated carapaces and plastrons. More than 50 specimens—mostly lower jaws—evince a high diversity of mammals, including monotremes, a multituberculate and several enigmatic ausktribosphenids. Relatively minor components of these fossil assemblages are diverse theropods (including birds), rare ankylosaurs and ceratopsians, pterosaurs, non-marine plesiosaurs and a lepidosaur. In the older strata of the upper Strzelecki Group, temnospondyl amphibians—the youngest known worldwide—are a conspicuous component of the fauna, whereas crocodylomorphs appear to be present only in up-sequence deposits of the Otway Group. Invertebrates are uncommon, although decapod crustaceans and unionoid bivalves have been described. Collectively, the Early Cretaceous biota of Victoria provides insights into a unique Mesozoic high-latitude palaeoenvironment and elucidates both palaeoclimatic and palaeobiogeographic changes throughout more than 25 million years of geological time. Stephen F. Poropat*† [sporopat@swin.edu.au; stephenfporopat@gmail.com], Faculty of Science, Engineering and Technology, Swinburne University of Technology, John St, Hawthorn, Victoria 3122, Australia; Sarah K. Martin*‡ [sarah.martin@dmirs.wa.gov.au; martin.sarahk@gmail.com] Geological Survey of Western Australia, 100 Plain St, East Perth, Western Australia 6004, Australia; Anne-Marie P. Tosolini [a.tosolini@unimelb.edu.au] and Barbara E. Wagstaff [wagstaff@unimelb.edu.au] School of Earth Sciences, The University of Melbourne, Melbourne, Victoria 3010, Australia; Lynne B. Bean [lynne.bean@anu.edu.au] Research School of Earth Sciences, Australian National University, Acton, Canberra, Australian Capital Territory 2001, Australia; Benjamin P. Kear [benjamin.kear@em.uu.se] Museum of Evolution, Uppsala University, Norbyvägen 16, Uppsala SE-752 36, Sweden; Patricia Vickers-Rich§ [prich@swin.edu.au; pat.rich@monash.edu] Faculty of Science, Engineering and Technology, Swinburne University of Technology, John St, Hawthorn, Victoria 3122, Australia; Thomas H. Rich [trich@museum.vic.gov.au] Museum Victoria, PO Box 666, Melbourne, Victoria 3001, Australia. *These authors contributed equally to this work. †Also affiliated with: Australian Age of Dinosaurs Museum of Natural History, Lot 1 Dinosaur Drive, PO Box 408, Winton, Queensland 4735, Australia. ‡Also affiliated with: Earth and Planetary Sciences, Western Australian Museum, Welshpool, Western Australia 6101, Australia. §Also affiliated with: School of Earth, Atmosphere and Environment, Monash University, Melbourne, Victoria 3800, Australia.