Biology of the sauropod dinosaurs: the evolution of gigantism.

The herbivorous sauropod dinosaurs of the Jurassic and Cretaceous periods were the largest terrestrial animals ever, surpassing the largest herbivorous mammals by an order of magnitude in body mass. Several evolutionary lineages among Sauropoda produced giants with body masses in excess of 50 metric tonnes by conservative estimates. With body mass increase driven by the selective advantages of large body size, animal lineages will increase in body size until they reach the limit determined by the interplay of bauplan, biology, and resource availability. There is no evidence, however, that resource availability and global physicochemical parameters were different enough in the Mesozoic to have led to sauropod gigantism. We review the biology of sauropod dinosaurs in detail and posit that sauropod gigantism was made possible by a specific combination of plesiomorphic characters (phylogenetic heritage) and evolutionary innovations at different levels which triggered a remarkable evolutionary cascade. Of these key innovations, the most important probably was the very long neck, the most conspicuous feature of the sauropod bauplan. Compared to other herbivores, the long neck allowed more efficient food uptake than in other large herbivores by covering a much larger feeding envelope and making food accessible that was out of the reach of other herbivores. Sauropods thus must have been able to take up more energy from their environment than other herbivores. The long neck, in turn, could only evolve because of the small head and the extensive pneumatization of the sauropod axial skeleton, lightening the neck. The small head was possible because food was ingested without mastication. Both mastication and a gastric mill would have limited food uptake rate. Scaling relationships between gastrointestinal tract size and basal metabolic rate (BMR) suggest that sauropods compensated for the lack of particle reduction with long retention times, even at high uptake rates. The extensive pneumatization of the axial skeleton resulted from the evolution of an avian-style respiratory system, presumably at the base of Saurischia. An avian-style respiratory system would also have lowered the cost of breathing, reduced specific gravity, and may have been important in removing excess body heat. Another crucial innovation inherited from basal dinosaurs was a high BMR. This is required for fueling the high growth rate necessary for a multi-tonne animal to survive to reproductive maturity. The retention of the plesiomorphic oviparous mode of reproduction appears to have been critical as well, allowing much faster population recovery than in megaherbivore mammals. Sauropods produced numerous but small offspring each season while land mammals show a negative correlation of reproductive output to body size. This permitted lower population densities in sauropods than in megaherbivore mammals but larger individuals. Our work on sauropod dinosaurs thus informs us about evolutionary limits to body size in other groups of herbivorous terrestrial tetrapods. Ectothermic reptiles are strongly limited by their low BMR, remaining small. Mammals are limited by their extensive mastication and their vivipary, while ornithsichian dinosaurs were only limited by their extensive mastication, having greater average body sizes than mammals.

A new basal sauropod dinosaur from the middle Jurassic of Niger and the early evolution of sauropoda.

BACKGROUND: The early evolution of sauropod dinosaurs is poorly understood because of a highly incomplete fossil record. New discoveries of Early and Middle Jurassic sauropods have a great potential to lead to a better understanding of early sauropod evolution and to reevaluate the patterns of sauropod diversification. PRINCIPAL FINDINGS: A new sauropod from the Middle Jurassic of Niger, Spinophorosaurus nigerensis n. gen. et sp., is the most complete basal sauropod currently known. The taxon shares many anatomical characters with Middle Jurassic East Asian sauropods, while it is strongly dissimilar to Lower and Middle Jurassic South American and Indian forms. A possible explanation for this pattern is a separation of Laurasian and South Gondwanan Middle Jurassic sauropod faunas by geographic barriers. Integration of phylogenetic analyses and paleogeographic data reveals congruence between early sauropod evolution and hypotheses about Jurassic paleoclimate and phytogeography. CONCLUSIONS: Spinophorosaurus demonstrates that many putatively derived characters of Middle Jurassic East Asian sauropods are plesiomorphic for eusauropods, while South Gondwanan eusauropods may represent a specialized line. The anatomy of Spinophorosaurus indicates that key innovations in Jurassic sauropod evolution might have taken place in North Africa, an area close to the equator with summer-wet climate at that time. Jurassic climatic zones and phytogeography possibly controlled early sauropod diversification.

Vertebral pathology in an ornithopod dinosaur: a hemivertebra in Dysalotosaurus lettowvorbecki from the Jurassic of Tanzania.

A vertebral fragment of the Late Jurassic ornithopod dinosaur Dysalotosaurus lettowvorbecki from Tanzania is described. It consists of a hemivertebra that is co-ossified with a complete vertebral centrum. The hemivertebra causes a hyperkyphotic posture of the vertebral column with an angle of approximately 35 degrees between the end plates of the vertebra, that is, a dorsal bending of the vertebral column. Associated with this is a 15 degrees lateral bending, which suggests a scoliosis. Micro-CT scans reveal thickening of the cortical bone in the hemivertebra and the complete vertebral centrum as compared to a "normal" vertebra. This can be interpreted as a reaction of the bone to the abnormal direction of forces arising from the defective configuration of the vertebral column. No signs of vertebral fracture are present. The arrangement of the Foramina venosa and the trapezoidal outline of the complete centrum that is co-ossified with the hemivertebra indicate that the hemivertebra in Dysalotosaurus developed early in embryogenesis probably by "hemimetameric segmental shift", that is, a defect of the fusion of the paired vertebral anlagen. This finding illustrates that hemivertebrae represent a fundamental defect of the vertebrate ontogenetic program.

Discovery of a short-necked sauropod dinosaur from the Late Jurassic period of Patagonia.

Sauropod dinosaurs are one of the most conspicuous groups of Mesozoic terrestrial vertebrates. They show general trends towards an overall increase in size and elongation of the neck, by means of considerable elongation of the length of individual vertebrae and a cervical vertebra count that, in some cases, increases to 19 (ref. 1). The long neck is a particular hallmark of sauropod dinosaurs and is usually regarded as a key feeding adaptation. Here we describe a new dicraeosaurid sauropod, from the latest Jurassic period of Patagonia, that has a particularly short neck. With a neck that is about 40% shorter than in other known dicraeosaurs, this taxon demonstrates a trend opposite to that seen in most sauropods and indicates that the ecology of dicraeosaurids might have differed considerably from that of other sauropods. The new taxon indicates that there was a rapid radiation and dispersal of dicraeosaurids in the Late Jurassic of the Southern Hemisphere, after the separation of Gondwana from the northern continents by the late Middle Jurassic.