Sauropod dinosaurs are a group of herbivorous dinosaurs which exceeded all other terrestrial vertebrates in mean and maximal body size. Sauropod dinosaurs were also the most successful and long-lived herbivorous tetrapod clade, but no abiological factors such as global environmental parameters conducive to their gigantism can be identified. These facts justify major efforts by evolutionary biologists and paleontologists to understand sauropods as living animals and to explain their evolutionary success and uniquely gigantic body size. Contributions to this research program have come from many fields and can be synthesized into a biological evolutionary cascade model of sauropod dinosaur gigantism (sauropod gigantism ECM). This review focuses on the sauropod gigantism ECM, providing an updated version based on the contributions to the PLoS ONE sauropod gigantism collection and on other very recent published evidence. The model consist of five separate evolutionary cascades (“Reproduction”, “Feeding”, “Head and neck”, “Avian-style lung”, and “Metabolism”). Each cascade starts with observed or inferred basal traits that either may be plesiomorphic or derived at the level of Sauropoda. Each trait confers hypothetical selective advantages which permit the evolution of the next trait. Feedback loops in the ECM consist of selective advantages originating from traits higher in the cascades but affecting lower traits. All cascades end in the trait “Very high body mass”. Each cascade is linked to at least one other cascade. Important plesiomorphic traits of sauropod dinosaurs that entered the model were ovipary as well as no mastication of food. Important evolutionary innovations (derived traits) were an avian-style respiratory system and an elevated basal metabolic rate. Comparison with other tetrapod lineages identifies factors limiting body size.
Titanosauria is an exceptionally diverse, globally-distributed clade of sauropod dinosaurs that includes the largest known land animals. Knowledge of titanosaurian pedal structure is critical to understanding the stance and locomotion of these enormous herbivores and, by extension, gigantic terrestrial vertebrates as a whole. However, completely preserved pedes are extremely rare among Titanosauria, especially as regards the truly giant members of the group. Here we describe Notocolossus gonzalezparejasi gen. et sp. nov. from the Upper Cretaceous of Mendoza Province, Argentina. With a powerfully-constructed humerus 1.76 m in length, Notocolossus is one of the largest known dinosaurs. Furthermore, the complete pes of the new taxon exhibits a strikingly compact, homogeneous metatarsus-seemingly adapted for bearing extraordinary weight-and truncated unguals, morphologies that are otherwise unknown in Sauropoda. The pes underwent a near-progressive reduction in the number of phalanges along the line to derived titanosaurs, eventually resulting in the reduced hind foot of these sauropods.
For 130 years, dinosaurs have been divided into two distinct clades-Ornithischia and Saurischia. Here we present a hypothesis for the phylogenetic relationships of the major dinosaurian groups that challenges the current consensus concerning early dinosaur evolution and highlights problematic aspects of current cladistic definitions. Our study has found a sister-group relationship between Ornithischia and Theropoda (united in the new clade Ornithoscelida), with Sauropodomorpha and Herrerasauridae (as the redefined Saurischia) forming its monophyletic outgroup. This new tree topology requires redefinition and rediagnosis of Dinosauria and the subsidiary dinosaurian clades. In addition, it forces re-evaluations of early dinosaur cladogenesis and character evolution, suggests that hypercarnivory was acquired independently in herrerasaurids and theropods, and offers an explanation for many of the anatomical features previously regarded as notable convergences between theropods and early ornithischians.
Massospondylus carinatus is a basal sauropodomorph dinosaur from the early Jurassic Elliot Formation of South Africa. It is one of the best-represented fossil dinosaur taxa, known from hundreds of specimens including at least 13 complete or nearly complete skulls. Surprisingly, the internal cranial anatomy of M. carinatus has never been described using computed tomography (CT) methods. Using CT scans and 3D digital representations, we digitally reconstruct the bones of the facial skeleton, braincase, and palate of a complete, undistorted cranium of M. carinatus (BP/1/5241). We describe the anatomical features of the cranial bones, and compare them to other closely related sauropodomorph taxa such as Plateosaurus erlenbergiensis, Lufengosaurus huenei, Sarahsaurus aurifontanalis and Efraasia minor. We identify a suite of character states of the skull and braincase for M. carinatus that sets it apart from other taxa, but these remain tentative due to the lack of comparative sauropodomorph braincase descriptions in the literature. Furthermore, we hypothesize 27 new cranial characters useful for determining relationships in non-sauropodan Sauropodomorpha, delete five pre-existing characters and revise the scores of several existing cranial characters to make more explicit homology statements. All the characters that we hypothesized or revised are illustrated. Using parsimony as an optimality criterion, we then test the relationships of M. carinatus (using BP/1/5241 as a specimen-level exemplar) in our revised phylogenetic data matrix.
Dinosauromorpha includes dinosaurs and other much less diverse dinosaur precursors of Triassic age, such as lagerpetids . Joint occurrences of these taxa with dinosaurs are rare but more common during the latest part of that period (Norian-Rhaetian, 228-201 million years ago [mya]) [2, 3]. In contrast, the new lagerpetid and saurischian dinosaur described here were unearthed from one of the oldest rock units with dinosaur fossils worldwide, the Carnian (237-228 mya) Santa Maria Formation of south Brazil , a record only matched in age by much more fragmentary remains from Argentina . This is the first time nearly complete dinosaur and non-dinosaur dinosauromorph remains are found together in the same excavation, clearly showing that these animals were contemporaries since the first stages of dinosaur evolution. The new lagerpetid preserves the first skull, scapular and forelimb elements, plus associated vertebrae, known for the group, revealing how dinosaurs acquired several of their typical anatomical traits. Furthermore, a novel phylogenetic analysis shows the new dinosaur as the most basal Sauropodomorpha. Its plesiomorphic teeth, strictly adapted to faunivory, provide crucial data to infer the feeding behavior of the first dinosaurs.
Osteocytes harbour much potential for paleobiological studies. Synchrotron radiation and spectroscopic analyses are providing fascinating data on osteocyte density, size and orientation in fossil taxa. However, such studies may be costly and time consuming. Here we describe an uncomplicated and inexpensive method to measure osteocyte lacunar densities in bone thin sections. We report on cell lacunar densities in the long bones of various extant and extinct tetrapods, with a focus on sauropodomorph dinosaurs, and how lacunar densities can help us understand bone formation rates in the iconic sauropod dinosaurs. Ordinary least square and phylogenetic generalized least square regressions suggest that sauropodomorphs have lacunar densities higher than scaled up or comparably sized mammals. We also found normal mammalian-like osteocyte densities for the extinct bovid Myotragus, questioning its crocodilian-like physiology. When accounting for body mass effects and phylogeny, growth rates are a main factor determining the density of the lacunocanalicular network. However, functional aspects most likely play an important role as well. Observed differences in cell strategies between mammals and dinosaurs likely illustrate the convergent nature of fast growing bone tissues in these groups.
The evolutionary history of dinosaurs might date back to the first stages of the Triassic (c. 250-240 Ma), but the oldest unequivocal records of the group come from Late Triassic (Carnian - c. 230 Ma) rocks of South America. Here, we present the first braincase endocast of a Carnian dinosaur, the sauropodomorph Saturnalia tupiniquim, and provide new data regarding the evolution of the floccular and parafloccular lobe of the cerebellum (FFL), which has been extensively discussed in the field of palaeoneurology. Previous studies proposed that the development of a permanent quadrupedal stance was one of the factors leading to the volume reduction of the FFL of sauropods. However, based on the new data for S. tupiniquim we identified a first moment of FFL volume reduction in non-sauropodan Sauropodomorpha, preceding the acquisition of a fully quadrupedal stance. Analysing variations in FFL volume alongside other morphological changes in the group, we suggest that this reduction is potentially related to the adoption of a more restricted herbivore diet. In this context, the FFL of sauropods might represent a vestigial trait, retained in a reduced version from the bipedal and predatory early sauropodomorphs.
Skeletal pneumaticity is found in the presacral vertebrae of most sauropod dinosaurs, but pneumaticity is much less common in the vertebrae of the tail. We describe previously unrecognized pneumatic fossae in the mid-caudal vertebrae of specimens of Giraffatitan and Apatosaurus. In both taxa, the most distal pneumatic vertebrae are separated from other pneumatic vertebrae by sequences of three to seven apneumatic vertebrae. Caudal pneumaticity is not prominent in most individuals of either of these taxa, and its unpredictable development means that it may be more widespread than previously recognised within Sauropoda and elsewhere in Saurischia. The erratic patterns of caudal pneumatization in Giraffatitan and Apatosaurus, including the pneumatic hiatuses, show that pneumatic diverticula were more broadly distributed in the bodies of the living animals than are their traces in the skeleton. Together with recently published evidence of cryptic diverticula-those that leave few or no skeletal traces-in basal sauropodomorphs and in pterosaurs, this is further evidence that pneumatic diverticula were widespread in ornithodirans, both across phylogeny and throughout anatomy.
The early evolution of sauropod dinosaurs remains poorly understood, with a paucity of unequivocal sauropod taxa known from the first twenty million years of the Jurassic. Recently, the Early Jurassic of South Africa has yielded an assemblage of dental and post-cranial remains displaying a more apomorphic character suite than any other similarly aged sauropodomorph. These remains are interpreted as a new species of basal sauropod and recovered cladistically as the sister taxon to Vulcanodon +more derived Sauropoda, underscoring its importance for our understanding of this pivotal period of sauropod evolution. Key changes in the dentition, axial skeleton and forelimb of this new species suggest a genuine functional distinction occurring at the sauropodiform-sauropod boundary. With reference to these changes, we propose a scenario in which interdependent refinements of the locomotory and feeding apparatus occurred in tandem with, or were effected by, restrictions in the amount of vertical forage initially available to the earliest sauropods. The hypothesized instance of niche-partitioning between basal sauropodan taxa and higher-browsing non-sauropodan sauropodomorphs may partially explain the rarity of true sauropods in the basal rocks of the Jurassic, while having the added corollary of couching the origins of Sauropoda in terms of an ecologically delimited ‘event’.
To estimate the body temperature (BT) of seven dinosaurs Gillooly, Alleen, and Charnov (2006) used an equation that predicts BT from the body mass and maximum growth rate (MGR) with the latter preserved in ontogenetic growth trajectories (BT-equation). The results of these authors evidence inertial homeothermy in Dinosauria and suggest that, due to overheating, the maximum body size in Dinosauria was ultimately limited by BT. In this paper, I revisit this hypothesis of Gillooly, Alleen, and Charnov (2006). I first studied whether BTs derived from the BT-equation of today’s crocodiles, birds and mammals are consistent with core temperatures of animals. Second, I applied the BT-equation to a larger number of dinosaurs than Gillooly, Alleen, and Charnov (2006) did. In particular, I estimated BT of Archaeopteryx (from two MGRs), ornithischians (two), theropods (three), prosauropods (three), and sauropods (nine). For extant species, the BT value estimated from the BT-equation was a poor estimate of an animal’s core temperature. For birds, BT was always strongly overestimated and for crocodiles underestimated; for mammals the accuracy of BT was moderate. I argue that taxon-specific differences in the scaling of MGR (intercept and exponent of the regression line, log-log-transformed) and in the parameterization of the Arrhenius model both used in the BT-equation as well as ecological and evolutionary adaptations of species cause these inaccuracies. Irrespective of the found inaccuracy of BTs estimated from the BT-equation and contrary to the results of Gillooly, Alleen, and Charnov (2006) I found no increase in BT with increasing body mass across all dinosaurs (Sauropodomorpha, Sauropoda) studied. This observation questions that, due to overheating, the maximum size in Dinosauria was ultimately limited by BT. However, the general high inaccuracy of dinosaurian BTs derived from the BT-equation makes a reliable test of whether body size in dinosaurs was ultimately limited by overheating impossible.