Biomechanics and morphological comparisons of the caudal region of titanosaurs from the Cretaceous of Brazil: Paleobiology and paleoecology inferences.

Biomechanical studies in sauropod dinosaurs are mainly focused on neck posture and feeding strategy. Few works investigate other aspects such as tail movement and function, especially in the clade Titanosauria, the most diverse within Neosauropoda. This study applied biomechanical concepts of neutral pose (cartilaginous neutral pose) and range of motion to verify the shape and direction of the caudal region of the advanced titanosaurs Adamantisaurus mezzalirai and Baurutitan britoi, in addition to comparing it with other titanosaurs (e.g., Arrudatitan maximus, Lirainosaurus astibiae, and Trigonosaurus pricei). As a result, the tails analyzed have a sigmoidal-convex shape, probably close to the ground (but not touching it). The sigmoidal-convex shape could increase the moment arm for the M. caudofemoralis longus, making it possible to use the tail as a fifth stabilizing member. This work expands our knowledge about sauropod dinosaurs by exposing a diversity of shapes for the tails of titanosaur sauropods and brings new possibilities for how these dinosaurs could use their tails.

Osteohistology of the hyperelongate hemispinous processes of Amargasaurus cazaui (Dinosauria: Sauropoda): Implications for soft tissue reconstruction and functional significance.

Dicraeosaurid sauropods are iconically characterized by the presence of elongate hemispinous processes in presacral vertebrae. These hemispinous processes can show an extreme degree of elongation, such as in the Argentinean forms Amargasaurus cazaui, Pilmatueia faundezi and Bajadasaurus pronuspinax. These hyperelongated hemispinous processes have been variably interpreted as a support structure for a padded crest/sail as a display, a bison-like hump or as the internal osseous cores of cervical horns. With the purpose to test these hypotheses, here we analyze, for the first time, the external morphology, internal microanatomy and bone microstructure of the hemispinous processes from the holotype of Amargasaurus, in addition to a second dicraeosaurid indet. (also from the La Amarga Formatin; Lower Cretaceous, Argentina). Transverse thin-sections sampled from the proximal, mid and distal portions of both cervical and dorsal hemispinous processes reveal that the cortical bone is formed by highly vascularized fibrolamellar bone interrupted with cyclical growth marks. Obliquely oriented Sharpey's fibres are mostly located in the medial and lateral portions of the cortex. Secondary remodelling is evidenced by the presence of abundant secondary osteons irregularly distributed within the cortex. Both anatomical and histological evidence does not support the presence of a keratinized sheath (i.e. horn) covering the hyperelongated hemispinous processes of Amargasaurus, and either, using a parsimonious criterium, in other dicraeosaurids with similar vertebral morphology. The spatial distribution and relative orientation of the Sharpey's fibres suggest the presence of an important system of interspinous ligaments that possibly connect successive hemispinous processes in Amargasaurus. These ligaments were distributed along the entirety of the hemispinous processes. The differential distribution of secondary osteons indicates that the cervical hemispinous processes of Amargasaurus were subjected to mechanical forces that generated higher compression strain on the anterior side of the elements. Current data support the hypothesis for the presence of a 'cervical sail' in Amargasaurus and other dicraeosaurids.

Myological reconstruction of the pelvic girdle and hind limb of the giant titanosaurian sauropod dinosaur Dreadnoughtus schrani.

Osteological correlates preserve more readily than their soft tissue counterparts in the fossil record; therefore, they can more often provide insight into the soft tissue anatomy of the organism. These insights can in turn elucidate the biology of these extinct organisms. In this study, we reconstruct the pelvic girdle and hind limb musculature of the giant titanosaurian sauropod Dreadnoughtus schrani based on observations of osteological correlates and Extant Phylogenetic Bracket comparisons. Recovered fossils of Dreadnoughtus exhibit remarkably well-preserved, well-developed, and extensive muscle scars. Furthermore, this taxon is significantly larger bodied than any titanosaurian for which a myological reconstruction has previously been performed, rendering this contribution highly informative for the group. All 20 of the muscles investigated in this study are sufficiently well supported to enable reconstruction of at least one division, including reconstruction of the M. ischiocaudalis for the first time in a sauropod dinosaur. In total, 34 osteological correlates were identified on the pelvic girdle and hind limb remains of Dreadnoughtus, allowing the reconstruction of 14 muscles on the basis of Level I or Level II inferences (i.e., not Level I' or Level II' inferences). Comparisons among titanosaurians suggest widespread myological variation, yet potential phylogenetic and other paleobiologic patterns are often obscured by fragmentary preservation, infrequent myological studies, and lack of consensus on the phylogenetic placement of many taxa. However, a ventrolateral accessory process is present on the preacetabular lobe of the ilium in all of the largest titanosauriforms that preserve this skeletal element, suggesting that the presence of this process (representing the origin of the M. puboischiofemoralis internus part II) may be associated with extreme body size. By identifying such myological patterns among titanosauriforms, we can begin to address specific evolutionary and biomechanical questions related to their skeletal anatomy, how they were capable of leaving wide-gauge trackways, and resulting locomotor attributes unique to this clade.

Extinction of herbivorous dinosaurs linked to Early Jurassic global warming event.

Sauropods, the giant long-necked dinosaurs, became the dominant group of large herbivores in terrestrial ecosystems after multiple related lineages became extinct towards the end of the Early Jurassic (190-174 Ma). The causes and precise timing of this key faunal change, as well as the origin of eusauropods (true sauropods), have remained ambiguous mainly due to the scarce dinosaurian fossil record of this time. The terrestrial sedimentary successions of the Cañadón Asfalto Basin in central Patagonia (Argentina) document this critical interval of dinosaur evolution. Here, we report a new dinosaur with a nearly complete skull that is the oldest eusauropod known to date and provide high-precision U-Pb geochronology that constrains in time the rise of eusauropods in Patagonia. We show that eusauropod dominance was established after a massive magmatic event impacting southern Gondwana (180-184 Ma) and coincided with severe perturbations to the climate and a drastic decrease in the floral diversity characterized by the rise of conifers with small scaly leaves. Floral and faunal records from other regions suggest these were global changes that impacted the terrestrial ecosystems during the Toarcian warming event and formed part of a second-order mass extinction event.

Appendicular myological reconstruction of the forelimb of the giant titanosaurian sauropod dinosaur Dreadnoughtus schrani.

Soft tissues are variably preserved in the fossil record with external tissues, such as skin and feathers, more frequently preserved than internal tissues (e.g. muscles). More commonly, soft tissues leave traces of their locations on bones and, for muscles, these clues can be used to reconstruct the musculature of extinct vertebrates, thereby enhancing our understanding of how these organisms moved and the evolution of their locomotor patterns. Herein we reconstruct the forelimb and shoulder girdle musculature of the giant titanosaurian sauropod Dreadnoughtus schrani based on observations of osteological correlates and dissections of taxa comprising the Extant Phylogenetic Bracket of non-avian dinosaurs (crocodilians and birds). Fossils of Dreadnoughtus exhibit remarkably well-preserved, well-developed, and extensive muscle scars. Furthermore, this taxon is significantly larger-bodied than any titanosaurian for which a myological reconstruction has previously been attempted, rendering this myological study highly informative for the clade. In total, 28 muscles were investigated in this study, for which 46 osteological correlates were identified; these osteological correlates allowed the reconstruction of 16 muscles on the basis of Level I or Level II inferences (i.e. not Level I' or Level II' inferences). Comparisons with other titanosaurians suggest widespread myological variation in the clade, although potential phylogenetic patterns are often obscured by fragmentary preservation, infrequent myological studies, and lack of consensus on the systematic position of many taxa. By identifying myological variations within the clade, we can begin to address specific evolutionary and biomechanical questions related to the locomotor evolution in these sauropods.

Bone histology sheds light on the nature of the "dermal armor" of the enigmatic sauropod dinosaur Agustinia ligabuei Bonaparte, 1999.

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.

A new sauropod dinosaur hindlimb from the Lower Cretaceous Wessex Formation, Isle of Wight, UK.

The Barremian-aged Wessex Formation of the Isle of Wight, UK, offers a globally significant glimpse into the sauropod dinosaur faunas of the early Cretaceous. These deposits have yielded specimens of several neosauropod lineages, such as rebbachisaurids, titanosauriforms (including some of the earliest titanosaur remains), and possible flagellicaudatans. Here, we report an undescribed sauropod partial hindlimb from the Wessex Formation (NHMUK PV R16500) and analyse its phylogenetic affinities. This hindlimb preserves the left tibia, astragalus and pes, lacking only a few phalanges. NHMUK PV R16500 can be diagnosed based on two autapomorphies: an unusually high distal end to midshaft transverse width ratio in metatarsals III and IV, and the presence of small bump-like projections located in the centre of the proximal articular surfaces of the unguals of pedal digits I and II. The phylogenetic affinities of NHMUK PV R16500 are uncertain: although our analyses recover it as an early-branching somphospondylan, a single character change moves it to close to Flagellicaudata when extended implied weighting is applied. The possibility of flagellicaudatan affinities for NHMUK PV R16500 implies a potential ghost lineage that survived the Jurassic/Cretaceous boundary; however, we present evidence that the somphospondylan position is more probable and should be preferred.

Reappraisal of sauropod dinosaur diversity in the Upper Cretaceous Winton Formation of Queensland, Australia, through 3D digitisation and description of new specimens.

Skeletal remains of sauropod dinosaurs have been known from Australia for over 100 years. Unfortunately, the classification of the majority of these specimens to species level has historically been impeded by their incompleteness. This has begun to change in the last 15 years, primarily through the discovery and description of several partial skeletons from the Cenomanian-lower Turonian (lower Upper Cretaceous) Winton Formation in central Queensland, with four species erected to date: Australotitan cooperensis, Diamantinasaurus matildae, Savannasaurus elliottorum, and Wintonotitan wattsi. The first three of these appear to form a clade (Diamantinasauria) of early diverging titanosaurs (or close relatives of titanosaurs), whereas Wintonotitan wattsi is typically recovered as a distantly related non-titanosaurian somphospondylan. Through the use of 3D scanning, we digitised numerous specimens of Winton Formation sauropods, facilitating enhanced comparison between type and referred specimens, and heretofore undescribed specimens. We present new anatomical information on the holotype specimen of Diamantinasaurus matildae, and describe new remains pertaining to twelve sauropod individuals. Firsthand observations and digital analysis enabled previously proposed autapomorphic features of all four named Winton Formation sauropod species to be identified in the newly described specimens, with some specimens exhibiting putative autapomorphies of more than one species, prompting a reassessment of their taxonomic validity. Supported by a specimen-level phylogenetic analysis, we suggest that Australotitan cooperensis is probably a junior synonym of Diamantinasaurus matildae, but conservatively regard it herein as an indeterminate diamantinasaurian, meaning that the Winton Formation sauropod fauna now comprises three (rather than four) valid diamantinasaurian species: Diamantinasaurus matildae, Savannasaurus elliottorum, and Wintonotitan wattsi, with the latter robustly supported as a member of the clade for the first time. We refer some of the newly described specimens to these three species and provide revised diagnoses, with some previously proposed autapomorphies now regarded as diamantinasaurian synapomorphies. Our newly presented anatomical data and critical reappraisal of the Winton Formation sauropods facilitates a more comprehensive understanding of the mid-Cretaceous sauropod palaeobiota of central Queensland.

The dentition of the Late Jurassic dwarf sauropod Europasaurus holgeri from northern Germany: ontogeny, function, and implications for a rhamphotheca-like structure in Sauropoda.

The basal macronarian sauropod Europasaurus holgeri is known only from the Late Jurassic of the Langenberg Quarry near Goslar, Lower Saxony, Germany. Europasaurus has been identified as an insular dwarf and shows a clear resemblance to Camarasaurus and Giraffatitan. This study provides a detailed description of the dentition of Europasaurus based on an array of fossils outstanding in their abundance, variety of preservation, and ontogenetic range. Dental morphology for the replacement and functional dentitions, the tooth replacement pattern, and implications for food intake are described for the Europasaurus dentition, which is characterized by broad-crowned teeth. Characteristic features for Europasaurus are the presence of denticles on replacement teeth, the wrinkled enamel, and large wear facets both on the apex and on the carinae of the tooth crowns. The partially articulated skull SNHM-2207-R and isolated tooth rows DfmMh/FV 580.1 and DfmMh/FV 896.7 suggest the presence of strong connective tissue partially covering the teeth. This connective tissue would have provided stability and protection for the teeth. Evidence for this connective tissue include exposed tooth necks, in-situ teeth with strongly resorbed roots which no longer would have been connected to the jaw bone, and wrinkled enamel and its surface pattern. The same features can be observed in other sauropod taxa as well. We therefore suggest that eusauropods in general possessed this connective tissue structure, which may be an autapomorphy of the group. Possibly, this hypothetical structure is homologous to the rhamphotheca in birds and some non-avian theropods, which, however rarely, show such a close integration of keratinous tissue and teeth that we hypothesize here.

A new juvenile sauropod specimen from the Middle Jurassic Dongdaqiao Formation of East Tibet.

Jurassic strata are widely distributed in the eastern part of Tibet Autonomous Region, and have yielded many dinosaur bones. However, none of these specimens has been studied extensively, and some remain unprepared. Here we provide a detailed description of some new sauropod material, including several cervical vertebrae and a nearly complete scapula, recovered from the Middle Jurassic of Chaya County, East Tibet. The cervical vertebrae have short centra that bear ventral midline keels, as in many non-neosauropod sauropods such as Shunosaurus. Moreover, the cervical centra display deep lateral excavations, partitioned by a septum. The scapula has proximal and distal ends that are both expanded as in mamenchisaurids and neosauropods. However, relatively small body size and lack of fusion of neurocentral sutures in the cervical vertebrae suggest that the available material is from a juvenile, and the length of the cervical centra may have increased relative to the size of the rest of the skeleton in later ontogenetic stages. Phylogenetic analysis provides limited evidence that the new Tibetan sauropod specimen belongs to Eusauropoda, being more derived than Shunosaurus, but is basal to Mamenchisauridae. The new material provides important information on the morphological transition between Shunosaurus and mamenchisaurids, and extends the known biogeographic range of early-diverging sauropods in the Middle Jurassic of East Asia.

Bite and tooth marks on sauropod dinosaurs from the Morrison Formation.

Tooth-marked bones provide important evidence for feeding choices made by extinct carnivorous animals. In the case of the dinosaurs, most bite traces are attributed to the large and robust osteophagous tyrannosaurs, but those of other large carnivores remain underreported. Here we report on an extensive survey of the literature and some fossil collections cataloging a large number of sauropod bones (68) from the Upper Jurassic Morrison Formation of the USA that bear bite traces that can be attributed to theropods. We find that such bites on large sauropods, although less common than in tyrannosaur-dominated faunas, are known in large numbers from the Morrison Formation, and that none of the observed traces showed evidence of healing. The presence of tooth wear in non-tyrannosaur theropods further shows that they were biting into bone, but it remains difficult to assign individual bite traces to theropod taxa in the presence of multiple credible candidate biters. The widespread occurrence of bite traces without evidence of perimortem bites or healed bite traces, and of theropod tooth wear in Morrison Formation taxa suggests preferential feeding by theropods on juvenile sauropods, and likely scavenging of large-sized sauropod carcasses.

A nearly complete skull of the sauropod dinosaur Diamantinasaurus matildae from the Upper Cretaceous Winton Formation of Australia and implications for the early evolution of titanosaurs.

Titanosaurian sauropod dinosaurs were diverse and abundant throughout the Cretaceous, with a global distribution. However, few titanosaurian taxa are represented by multiple skeletons, let alone skulls. Diamantinasaurus matildae, from the lower Upper Cretaceous Winton Formation of Queensland, Australia, was heretofore represented by three specimens, including one that preserves a braincase and several other cranial elements. Herein, we describe a fourth specimen of Diamantinasaurus matildae that preserves a more complete skull-including numerous cranial elements not previously known for this taxon-as well as a partial postcranial skeleton. The skull of Diamantinasaurus matildae shows many similarities to that of the coeval Sarmientosaurus musacchioi from Argentina (e.g. quadratojugal with posterior tongue-like process; braincase with more than one ossified exit for cranial nerve V; compressed-cone-chisel-like teeth), providing further support for the inclusion of both taxa within the clade Diamantinasauria. The replacement teeth within the premaxilla of the new specimen are morphologically congruent with teeth previously attributed to Diamantinasaurus matildae, and Diamantinasauria more broadly, corroborating those referrals. Plesiomorphic characters of the new specimen include a sacrum comprising five vertebrae (also newly demonstrated in the holotype of Diamantinasaurus matildae), rather than the six or more that typify other titanosaurs. However, we demonstrate that there have been a number of independent acquisitions of a six-vertebrae sacrum among Somphospondyli and/or that there have been numerous reversals to a five-vertebrae sacrum, suggesting that sacral count is relatively plastic. Other newly identified plesiomorphic features include: the overall skull shape, which is more similar to brachiosaurids than 'derived' titanosaurs; anterior caudal centra that are amphicoelous, rather than procoelous; and a pedal phalangeal formula estimated as 2-2-3-2-0. These features are consistent with either an early-branching position within Titanosauria, or a position just outside the titanosaurian radiation, for Diamantinasauria, as indicated by alternative character weighting approaches applied in our phylogenetic analyses, and help to shed light on the early assembly of titanosaurian anatomy that has until now been obscured by a poor fossil record.

New specimens of Baurutitan britoi and a taxonomic reassessment of the titanosaur dinosaur fauna (Sauropoda) from the Serra da Galga Formation (Late Cretaceous) of Brazil.

The description of new titanosaur specimens unearthed from deposits of the Serra da Galga Formation (Bauru Group, Late Cretaceous) at the BR-262 site, near Peirópolis (Uberaba, Minas Gerais State, Brazil), sheds light on the taxonomy of two taxa previously known from the same area and geological unit: Baurutitan britoi and Trigonosaurus pricei. A comparative revision indicates that T. pricei represents a junior synonym of Ba. britoi, and that the BR-262 specimens belong to that latter species. The information provided by the new specimens also revealed that the paratype of T. pricei (MCT 1719-R), a caudal vertebral series, actually represents a new taxon, named here as Caieiria allocaudata gen. et sp. nov.

New macronarian from the Middle Jurassic of Chongqing, China: phylogenetic and biogeographic implications for neosauropod dinosaur evolution.

Macronaria is a clade of gigantic body-sized sauropod dinosaurs widely distributed from the Late Jurassic to the Late Cretaceous globally. However, its origin, early diversification, and dispersal are still controversial. Here, we report a new macronarian Yuzhoulong qurenensis gen. et sp. nov. excavated from the Middle Jurassic (Bathonian) Lower Shaximiao Formation. Yuzhoulong qurenensis bears a unique combination of features, such as two accessory fossae that exist on the posterior surface of dorsal diapophyses of anterior dorsal vertebrae. Results of phylogenetic analyses demonstrate it is one of the earliest-diverging macronarians. This new material represents a Middle Jurassic fossil record of macronarian sauropod worldwide and improves the understanding of the early diversity and dispersal of the Neosauropoda. This discovery further supports that sauropods achieved a more rapid and varied morphological diversity and palaeogeographical dispersal in the Middle Jurassic.

Sauropod dinosaur teeth from the lower Upper Cretaceous Winton Formation of Queensland, Australia and the global record of early titanosauriforms.

The Upper Cretaceous Winton Formation of Queensland, Australia, has produced several partial sauropod skeletons, but cranial remains-including teeth-remain rare. Herein, we present the first description of sauropod teeth from this formation, based on specimens from three separate sites. An isolated tooth and a dentary fragment from the Diamantinasaurus matildae type locality are considered to be referable to that titanosaurian taxon. A single tooth from the D. matildae referred specimen site is similarly regarded as being part of that individual. Seventeen teeth from a new site that are morphologically uniform, and similar to the teeth from the two Diamantinasaurus sites, are assigned to Diamantinasauria. All sauropod teeth recovered from the Winton Formation to date are compressed-cone-chisel-shaped, have low slenderness index values (2.00-2.88), are lingually curved at their apices, mesiodistally convex on their lingual surfaces, and lack prominent carinae and denticles. They are markedly different from the chisel-like teeth of derived titanosaurs, more closely resembling the teeth of early branching members of the titanosauriform radiation. This provides further support for a 'basal' titanosaurian position for Diamantinasauria. Scanning electron microscope microwear analysis of the wear facets of several teeth reveals more scratches than pits, implying that diamantinasaurians were mid-height (1-10 m) feeders. With a view to assessing the spatio-temporal distribution of sauropod tooth morphotypes before and after deposition of the Winton Formation, we provide a comprehensive continent-by-continent review of the early titanosauriform global record (Early to early Late Cretaceous). This indicates that throughout the Early-early Late Cretaceous, sauropod faunas transitioned from being quite diverse at higher phylogenetic levels and encompassing a range of tooth morphologies at the start of the Berriasian, to faunas comprising solely titanosaurs with limited dental variability by the end-Turonian. Furthermore, this review highlights the different ways in which this transition unfolded on each continent, including the earliest records of titanosaurs with narrow-crowned teeth on each continent.

Climatic constraints on the biogeographic history of Mesozoic dinosaurs.

Dinosaurs dominated Mesozoic terrestrial ecosystems globally. However, whereas a pole-to-pole geographic distribution characterized ornithischians and theropods, sauropods were restricted to lower latitudes. Here, we evaluate the role of climate in shaping these biogeographic patterns through the Jurassic-Cretaceous (201-66 mya), combining dinosaur fossil occurrences, past climate data from Earth System models, and habitat suitability modeling. Results show that, uniquely among dinosaurs, sauropods occupied climatic niches characterized by high temperatures and strongly bounded by minimum cold temperatures. This constrained the distribution and dispersal pathways of sauropods to tropical areas, excluding them from latitudinal extremes, especially in the Northern Hemisphere. The greater availability of suitable habitat in the southern continents, particularly in the Late Cretaceous, might be key to explaining the high diversity of sauropods there, relative to northern landmasses. Given that ornithischians and theropods show a flattened or bimodal latitudinal biodiversity gradient, with peaks at higher latitudes, the closer correspondence of sauropods to a subtropical concentration could hint at fundamental thermophysiological differences to the other two clades.

Ecomorphospace occupation of large herbivorous dinosaurs from Late Jurassic through to Late Cretaceous time in North America.

Following the Late Jurassic, megaherbivore communities in North America undergo a dramatic turnover in faunal composition: sauropods decline to the point of becoming relatively minor components of ecosystems, stegosaurs become extinct, and hadrosaurids, ceratopsids and ankylosaurs rise in diversity and abundance. Although a variety of causes have been proposed to account for the dramatic decrease in sauropod diversity following the Late Jurassic and could have also been applicable to the disappearance of stegosaurs, the potential for competitive replacement of sauropods by hadrosauroids as an explanation has been previously dismissed due to morphological differences without further investigation. Using twelve ecomorphological correlates of the skull, this study provides a preliminary investigation into ecomorphospace occupation of major megaherbivore clades from the Late Jurassic through to the Late Cretaceous of North America and assess if morphological differences were enough to have potentially facilitated dietary niche partitioning between sauropods and iguanodontians and stegosaurs and ankylosaurs. Overlap in reconstructed ecomorphospace was observed between sauropods (particularly non-diplodocid sauropods) and iguanodontians, as would be expected if morphological differences were not enough to facilitate niche partitioning, contrary to original claims used to dismiss the competitive replacement hypothesis. Overlap was also observed between stegosaurs and ankylosaurs, particularly between Late Cretaceous ankylosaurs. Whether this overlap is reflective competitive replacement or opportunistic occupation of recently vacated niches will require further assessment as sampling of some clades prior to the Late Cretaceous is too poor to make a reliable assessment and several underlying assumptions necessary for competition to occur (e.g., resource limitation) still need investigation. Teasing out the cause(s) of the 'sauropod decline' and extinction of stegosaurs in North America following the Late Jurassic will require future research not only into the competitive exclusion hypothesis, but other hypotheses as well with better sampling from Early Cretaceous and Late Jurassic intervals.

Rapid growth preceded gigantism in sauropodomorph evolution.

Sauropod dinosaurs include the largest land animals to have walked the earth, mostly weighing 10-70 tons (e.g., Sander et al.1 and Carballido et al.2). Osteohistology suggests that derived physiological traits evolved near the origin of sauropod gigantism, including both rapid and uninterrupted growth from juvenile to adult with little developmental plasticity.1,3,4 This differs from the slower, seasonally interrupted growth of their direct ancestors, as evident in most non-sauropodan sauropodomorphs, which also show developmental plasticity in some groups. Accelerated but seasonally interrupted growth is present in Lessemsauridae, the sister clade to Sauropoda, which also attained giant adult body sizes (>10 tons).5 These observations suggest a correlation between giant size and accelerated growth. However, testing this evolutionary connection has been limited by the incomplete understanding of the growth patterns in some of the closest non-giant relatives of sauropods. We present the osteohistology of two such taxa, Aardonyx celestae and Sefapanosaurus zatronensis. Both exhibit highly vascularized woven-parallel complexes, with fibrolamellar complexes during early to mid-ontogeny, containing regular growth marks. These observations provide strong evidence for rapid but seasonally interrupted growth with limited developmental plasticity (indicated by the regular spacing of growth marks). Combined with our review of early branching sauropodomorph osteohistology, these results show that highly accelerated growth rates originated among smaller, non-sauropodan sauropodomorphs weighing 1 to 2 tons but preceded the origins of giant size (>10 tons). Therefore, the capacity for rapid bone tissue formation, a derived aspect of rapid growth seen in sauropods, did not evolve specifically to enable giant body sizes but may have been a prerequisite for them.

Anatomy and systematics of the diplodocoid Amphicoelias altus supports high sauropod dinosaur diversity in the Upper Jurassic Morrison Formation of the USA.

Sauropod dinosaurs were an abundant and diverse component of the Upper Jurassic Morrison Formation of the USA, with 24 currently recognized species. However, some authors consider this high diversity to have been ecologically unviable and the validity of some species has been questioned, with suggestions that they represent growth series (ontogimorphs) of other species. Under this scenario, high sauropod diversity in the Late Jurassic of North America is greatly overestimated. One putative ontogimorph is the enigmatic diplodocoid Amphicoelias altus, which has been suggested to be synonymous with Diplodocus. Given that Amphicoelias was named first, it has priority and thus Diplodocus would become its junior synonym. Here, we provide a detailed re-description of A. altus in which we restrict it to the holotype individual and support its validity, based on three autapomorphies. Constraint analyses demonstrate that its phylogenetic position within Diplodocoidea is labile, but it seems unlikely that Amphicoelias is synonymous with Diplodocus. As such, our re-evaluation also leads us to retain Diplodocus as a distinct genus. There is no evidence to support the view that any of the currently recognized Morrison sauropod species are ontogimorphs. Available data indicate that sauropod anatomy did not dramatically alter once individuals approached maturity. Furthermore, subadult sauropod individuals are not prone to stemward slippage in phylogenetic analyses, casting doubt on the possibility that their taxonomic affinities are substantially misinterpreted. An anatomical feature can have both an ontogenetic and phylogenetic signature, but the former does not outweigh the latter when other characters overwhelmingly support the affinities of a taxon. Many Morrison Formation sauropods were spatio-temporally and/or ecologically separated from one another. Combined with the biases that cloud our reading of the fossil record, we contend that the number of sauropod dinosaur species in the Morrison Formation is currently likely to be underestimated, not overestimated.

A diverse Late Cretaceous vertebrate tracksite from the Winton Formation of Queensland, Australia.

The Upper Cretaceous 'upper' Winton Formation of Queensland, Australia is world famous for hosting Dinosaur Stampede National Monument at Lark Quarry Conservation Park, a somewhat controversial tracksite that preserves thousands of tridactyl dinosaur tracks attributed to ornithopods and theropods. Herein, we describe the Snake Creek Tracksite, a new vertebrate ichnoassemblage from the 'upper' Winton Formation, originally situated on Karoola Station but now relocated to the Australian Age of Dinosaurs Museum of Natural History. This site preserves the first sauropod tracks reported from eastern Australia, a small number of theropod and ornithopod tracks, the first fossilised crocodyliform and ?turtle tracks reported from Australia, and possible lungfish and actinopterygian feeding traces. The sauropod trackways are wide-gauge, with manus tracks bearing an ungual impression on digit I, and anteriorly tapered pes tracks with straight or concave forward posterior margins. These tracks support the hypothesis that at least one sauropod taxon from the 'upper' Winton Formation retained a pollex claw (previously hypothesised for Diamantinasaurus matildae based on body fossils). Many of the crocodyliform trackways indicate underwater walking. The Snake Creek Tracksite reconciles the sauropod-, crocodyliform-, turtle-, and lungfish-dominated body fossil record of the 'upper' Winton Formation with its heretofore ornithopod- and theropod-dominated ichnofossil record.