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 tetrapo...

Sr/ 86 Sr reference maps (isoscapes) are a key tool for investigating past human and animal migrations. However, there is little understanding of which biosphere samples are best proxies for local bioavailable Sr when dealing with movements of past populations. In this study, biological and geological samples (ground vegetation, tree leaves, rock leachates, water, soil extracts, as well as modern and archeological animal teeth and snail shells) were collected in the vicinity of two early medieval cemeteries ("Thuringians", 5-6th century AD) in central Germany, in order to characterize 87 Sr/ 86 Sr of the local biosphere. Animal tooth enamel is not appropriate in this specific context to provide a reliable 87 Sr/ 86 Sr baseline for investigating past human migration. Archeological faunal teeth data (pig, sheep/goat, and cattle) indicates a different feeding area compared to that of the human population and modern deer teeth 87 Sr/ 86 Sr such as freshwater bivalve shells and tree cores were examined in order to track potential historic anthropogenic contamination of the water and the vegetation. The data obtained from the archeological bivalve shells show that the modern rivers yield 87 Sr/ 86 Sr ratios which are similar to those of the past. However, the tree cores registered decreasing 87 Sr/ 86 Sr values over time towards present day likely mirroring anthropogenic activities such as forest liming, coal mining and/or soil acidification. The comparison of 87 Sr/ 86 Sr of the Thuringian skeletons excavated in the same area also shows that the vegetation samples are very likely anthropogenically influenced to some extent, affecting especially 87 Sr/ 86 Sr of the shallow rooted plants.

The stable isotope compositions of biologically precipitated apatite in bone, teeth, and scales are widely used to obtain information on the diet, behavior, and physiology of extinct organisms and to reconstruct past climate. Here we report the application of a new type of geochemical measurement to bioapatite, a "clumpedisotope" paleothermometer, based on the thermodynamically driven preference for 13 C and 18 O to bond with each other within carbonate ions in the bioapatite crystal lattice. This effect is dependent on temperature but, unlike conventional stable isotope paleothermometers, is independent from the isotopic composition of water from which the mineral formed. We show that the abundance of 13 C-18 O bonds in the carbonate component of tooth bioapatite from modern specimens decreases with increasing body temperature of the animal, following a relationship between isotope "clumping" and temperature that is statistically indistinguishable from inorganic calcite. This result is in agreement with a theoretical model of isotopic ordering in carbonate ion groups in apatite and calcite. This thermometer constrains body temperatures of bioapatite-producing organisms with an accuracy of 1-2°C. Analyses of fossilized tooth enamel of both Pleistocene and Miocene age yielded temperatures within error of those derived from similar modern taxa. Clumped-isotope analysis of bioapatite represents a new approach in the study of the thermophysiology of extinct species, allowing the first direct measurement of their body temperatures. It will also open new avenues in the study of paleoclimate, as the measurement of clumped isotopes in phosphorites and fossils has the potential to reconstruct environmental temperatures.apatite | isotope | paleoclimate | thermophysiology | paleothermometry T he mechanisms by which animals regulate their body temperatures are among the most fundamental aspects of their biology. The acquisition of endothermy, the ability to maintain high and stable body temperatures through internal heat production, is a major physiological change that occurred at an unknown stage during the evolutionary transition to mammals and birds from their ancestors among the nonmammalian therapsids and nonavian dinosaurs, respectively (1). Approaches to understanding the physiology of extinct animals and the evolution of endothermy have largely focused on biophysical modeling, anatomical observations, growth rate analysis from bone histology, and behavioral studies such as estimating predator/prey ratios (1-7). The validity of each of these approaches is uncertain (for contrasting viewpoints on approaches to dinosaur thermoregulation see refs. 4 and 5).Modern endothermic mammals and ectotherms, such as alligators and crocodiles, generally have significant differences in average body temperatures. With rare exceptions, mammals have high and stable body temperatures around 36-38°C regardless of their environment, whereas the body temperatures of ectotherms are generally lower on average and often fluctuate depending o...

The nature of the physiology and thermal regulation of the nonavian dinosaurs is the subject of debate. Previously, arguments have been made for both endothermic and ectothermic metabolisms on the basis of differing methodologies. We used clumped isotope thermometry to determine body temperatures from the fossilized teeth of large Jurassic sauropods. Our data indicate body temperatures of 36° to 38°C, which are similar to those of most modern mammals. This temperature range is 4° to 7°C lower than predicted by a model that showed scaling of dinosaur body temperature with mass, which could indicate that sauropods had mechanisms to prevent excessively high body temperatures being reached because of their gigantic size.

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Our understanding of the evolutionary transitions leading to the modern endothermic state of birds and mammals is incomplete, partly because tools available to study the thermophysiology of extinct vertebrates are limited. Here we show that clumped isotope analysis of eggshells can be used to determine body temperatures of females during periods of ovulation. Late Cretaceous titanosaurid eggshells yield temperatures similar to large modern endotherms. In contrast, oviraptorid eggshells yield temperatures lower than most modern endotherms but ∼ 6 °C higher than co-occurring abiogenic carbonates, implying that this taxon did not have thermoregulation comparable to modern birds, but was able to elevate its body temperature above environmental temperatures. Therefore, we observe no strong evidence for end-member ectothermy or endothermy in the species examined. Body temperatures for these two species indicate that variable thermoregulation likely existed among the non-avian dinosaurs and that not all dinosaurs had body temperatures in the range of that seen in modern birds.

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