Late Cretaceous sauropod dinosaur undertracks in floodplain deposits of Sanfranciscana Basin, Brazil, and their paleoecological significance

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 0311-5518Austrosaurus 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.

We describe a giant titanosaurid sauropod dinosaur discovered in coastal deposits in the Upper Cretaceous Bahariya Formation of Egypt, a unit that has produced three Tyrannosaurus-sized theropods and numerous other vertebrate taxa. Paralititan stromeri is the first tetrapod reported from Bahariya since 1935. Its 1.69-meter-long humerus is longer than that of any known Cretaceous sauropod. The autochthonous scavenged skeleton was preserved in mangrove deposits, raising the possibility that titanosaurids and their predators habitually entered such environments.

New observations on various sauropod postcranial elements from Queensland provide insights into the taxonomic composition of northern Australia's sauropod fauna and the structure of sauropod vertebrae. An incomplete sauropod humerus from a site near Blackall, Queensland, represents the southernmost occurrence of sauropod fossils in the Eromanga Basin, and indicates a possibly new taxon. The internal architecture of at least one of the vertebral centra of Austrosaurus mckillopi comprises bony disks parallel to the posterior articular face and bony lamellae perpendicular to the anterior articular face reinforcing the structure against axial forces. The lack of pneumaticity proximally in dorsal ribs indicates that A. mckillopi may not be a titanosauriform. Material (QM F6737) from the Winton Formation includes probable osteoderms, the first known from Australian sauropods, and some of the oldest known. Comparison of this specimen with named Winton Formation sauropods suggests that it represents a distinct taxon.

The discovery of deep-sea hydrothermal vents in 1977 challenged our views of ecosystem functioning and yet, the research conducted at these extreme and logistically challenging environments still continues to reveal unique biological processes. Here, we report for the first time, a unique behavior where the deep-sea skate, Bathyraja spinosissima, appears to be actively using the elevated temperature of a hydrothermal vent environment to naturally “incubate” developing egg-cases. We hypothesize that this behavior is directly targeted to accelerate embryo development time given that deep-sea skates have some of the longest egg incubation times reported for the animal kingdom. Similar egg incubating behavior, where eggs are incubated in volcanically heated nesting grounds, have been recorded in Cretaceous sauropod dinosaurs and the rare avian megapode. To our knowledge, this is the first time incubating behavior using a volcanic source is recorded for the marine environment.

A new gigantic sauropod, Huabeisaurus allocotus gen. et sp. nov., about 20 m in length and 5 m in height, was discovered in the Upper Cretaceous Huiquanpu Formation, Tianzhen County, Shanxi Province. It is notably different from Diplodocidae, Titanosauridae and Nemegtosauridae in the following aspects: the teeth are strong, peglike with a length ratio of the tooth crown to tooth root at about 3 to 1; the cervical vertebrae are long with forked spines; the spines in dorsal vertebrae are relatively high, unbifurcated; the caudal vertebrae are amphicoelous, with anterior neural spines and unbifurcated spines and chevrons; the femur is straight and long, narrow and flat and the tibia and fibula are long and flat. These characters show that the described genus should represent a new family, Huabeisauridae fam. nov. The discovery enriches the sauropod dinosaur record in China, and is quite significant to the study of the taxonomy, evolution, migration, extinction and palaeobiogeographic provincialism of the Late Cretaceous sauropod dinosaurs.

New dinosaur remains from the Quiricó Formation, Sanfranciscana Basin (Lower Cretaceous), Southwestern Brazil

Franz Baron Nopcsa recognised that the Late Cretaceous titanosaurid sauropod from Transylvania, Magyarosaurus dacus, was much smaller than the better known sauropods from the Upper Jurassic of the United States and even from Late Cretaceous deposits elsewhere in the world. In keeping with his biogeographic interpretations, Nopcsa viewed this difference as a consequence of body-size effects via island habitation. We present a preliminary re-evaluation of Nopcsa's claims and their heterochronic consequences using two approaches, viz. (1) regression analysis of humeral data as a means of establishing patterns in body size among titanosaurids, and (2) optimisation of humeral data onto titanosaurid cladograms to evaluate evolutionary trends within the clade. Our regression analysis is based on twenty species distributed among fourteen genera of neosauropods and uses length and mid-shaft mediolateral width of the humerus of presumed fully adult forms, and of growth samples that consist of postnatal to adult individuals. Linear regression analysis suggests that, among adult neosauropods, M. dacus appears to be represented by the smallest individuals; that M. dacus humeri appear to be more similar to those of subadults than to adults of other taxa; and that this ‘juvenile’ morphology may constitute dwarfing in M. dacus by paedomorphosis. In order for these regressions to reflect their evolutionary context more fully, we also present optimisation analyses of humeral form within Titanosauroidea. Although many aspects of the phylogeny of this clade are relatively poorly resolved, preliminary results of our analyses are consistent with the conclusion that M. dacus was a heterochronic dwarf.

Titanosaurs were a globally distributed clade of Cretaceous sauropods. Historically regarded as a primarily Gondwanan radiation, there is a growing number of Eurasian taxa, with several putative titanosaurs contemporaneous with, or even pre-dating, the oldest known Southern Hemisphere remains. The early Late Cretaceous Jinhua Formation, in Zhejiang Province, China, has yielded two putative titanosaurs, Jiangshanosaurus lixianensis and Dongyangosaurus sinensis. Here, we provide a detailed re-description and diagnosis of Jiangshanosaurus, as well as new anatomical information on Dongyangosaurus. Previously, a ‘derived’ titanosaurian placement for Jiangshanosaurus was primarily based on the presence of procoelous anterior caudal centra. We show that this taxon had amphicoelous anterior-middle caudal centra. Its only titanosaurian synapomorphy is that the dorsal margins of the scapula and coracoid are approximately level with one another. Dongyangosaurus can clearly be differentiated from Jiangshanosaurus, and displays features that indicate a closer relationship to the titanosaur radiation. Revised scores for both taxa are incorporated into an expanded phylogenetic data matrix, comprising 124 taxa scored for 548 characters. Under equal weights parsimony, Jiangshanosaurus is recovered as a member of the non-titanosaurian East Asian somphospondylan clade Euhelopodidae, and Dongyangosaurus lies just outside of Titanosauria. However, when extended implied weighting is applied, both taxa are placed within Titanosauria. Most other ‘middle’ Cretaceous East Asian sauropods are probably non-titanosaurian somphospondylans, but at least Xianshanosaurus appears to belong to the titanosaur radiation. Our analyses also recover the Early Cretaceous European sauropod Normanniasaurus genceyi as a ‘derived’ titanosaur, clustering with Gondwanan taxa. These results provide further support for a widespread diversification of titanosaurs by at least the Early Cretaceous.

South American titanosaurians have been central to the study of the evolution of Cretaceous sauropod dinosaurs. Despite their remarkable diversity, the fragmentary condition of several taxa and the scarcity of records outside Patagonia and southwestern Brazil have hindered the study of continental-scale paleobiogeographic relationships. We describe two new Late Cretaceous titanosaurians from Quebrada de Santo Domingo (La Rioja, Argentina), which help to fill a gap between these main areas of the continent. Our phylogenetic analysis recovers both new species, and several Brazilian taxa, within Rinconsauria. The data suggest that, towards the end of the Cretaceous, this clade spread throughout southern South America. At the same locality, we discovered numerous accumulations of titanosaurian eggs, likely related to the new taxa. With eggs distributed in three levels along three kilometres, the new site is one of the largest ever found and provides further evidence of nesting site philopatry among Titanosauria.

Agustinia ligabuei is an Early Cretaceous sauropod dinosaur from the northwest of Patagonia that is currently the topic of debate with respect to its phylogenetic position and atypical dermal armor. The presence of four morphotypes of laminar and transversely elongated putative osteoderms was used to consider Agustinia as an armored sauropod. Regarding the different hypotheses about the identity of the bony structures of Agustinia (e.g., osteoderms, cervical or dorsal ribs, hypertrophied elements), a comparative histological analysis has been carried out. Histological evidence is presented herein and reveals that none of the morphotypes of Agustinia shows a primary bone tissue formed by structural fiber bundles as in other sauropod dinosaur osteoderms. Furthermore, on the basis of their gross morphology and microstructure, the bony structures originally classified as types 1+4 and 3 are more comparable respectively with dorsal and cervical ribs than any other kind of dermal or bony element. Due to poor preservation, the nature of the type 2 cannot be assessed but is here tentatively assigned to a pelvic girdle element. Although a phylogenetic reassessment of Agustinia is not the purpose of this paper, our paleohistological analyses have broader implications: by not supporting the dermal armor hypothesis for Agustinia, its inclusion in Lithostrotia is not justified in the absence of other diagnostic features.

Review of Cretaceous sauropod dinosaurs from Central Asia

Reappraisal of the Early Cretaceous sauropod dinosaur Amargatitanis macni (Apesteguía, 2007), from northwestern Patagonia, Argentina

Lower Cretaceous fossils from central Niger document the succession of sauropod dinosaurs on Africa as it drifted into geographic isolation. A new broad-toothed genus of Neocomian age ( approximately 135 million years ago) shows few of the specializations of other Cretaceous sauropods. A new small-bodied sauropod of Aptian-Albian age ( approximately 110 million years ago), in contrast, reveals the highly modified cranial form of rebbachisaurid diplodocoids. Rates of skeletal change in sauropods and other major groups of dinosaurs are estimated quantitatively and shown to be highly variable.

Skolithos serratus differs from other simple, vertical, unbranched trace fossils by its distinctive horizontal scratch marks arranged in a screw-like pattern attributed to tiger beetle larvae (Cicindelidae). Despite its diagnostic morphology and established tracemaker, this ichnospecies remains poorly documented, with only sparse occurrences in the geological record. In this study, we present a detailed analysis of Skolithos serratus from floodplain deposits of the Quiricó Formation (Sanfranciscana Basin, Brazil). Our results demonstrate colonization in low-energy environments characterized by periods of non-deposition and subaerial exposure. This indicates a tracemaker preference for exposed, well-drained fine sandy substrates associated with lowered water tables, conditions necessary for the complete development of larval stages. Furthermore, Skolithos serratus colonization occurred predominantly during late-stage substrate drainage, facilitated by organisms with appendages adapted for burrowing into firm substrates developed between aggradation cycles driven by autocyclic processes. We also highlight the striking disparity in the documentation of S. serratus compared to other Skolithos ichnospecies in fluvial environments. This discrepancy reflects niche specialization by the S. serratus tracemaker and the strong control exerted by substrate properties, as all recorded occurrences are restricted to fine-grained heterolithic deposits composed predominantly of fine sand and mud. Finally, we expand the paleobiogeographic distribution of Cicindelidae in South America, expanding the record of their Mesozoic occurrences.

A new narrow-gauge sauropod trackway from the Cenomanian Candeleros Formation, northern Patagonia, Argentina