Long bones (femora, humeri) are the most abundant remains of sauropod dinosaurs. Their length is a good proxy for body length and body mass, and their histology is informative about ontogenetic age. Here we provide a comparative assessment of histologic changes in growth series of several sauropod taxa, including diplodocids (Apatosaurus, Diplodocus, indeterminate Diplodocinae from the Tendaguru Beds and from the Morrison Formation), basal macronarians (Camarasaurus, Brachiosaurus, Europasaurus), and titanosaurs (Phuwiangosaurus, Ampelosaurus). A total of 167 long bones, mainly humeri and femora, and 18 limb girdle bones were sampled. Sampling was performed by core drilling at prescribed locations at midshaft, and 13 histologic ontogenetic stages (HOS stages) were recognized. Because growth of all sauropod long bones is quite uniform, with laminar fibrolamellar bone being the dominant tissue, HOS stages could be recognized across taxa, although with minor differences. Histologic ontogenetic stages generally correlate closely with body size and thus provide a means to resolve important issue like the ontogenetic status of questionable specimens. We hypothesize that sexual maturity was attained at HOS-8, well before maximum size was attained, but we did not find sexually differentiated growth trajectories subsequent to HOS-8. On the basis of HOS stages, we detected two morphotypes in the Camarasaurus sample, a small one (type 1) and a larger one (type 2), presumably representing different species or sexual dimorphism.
BackgroundSauropterygia is an abundant and successful group of Triassic marine reptiles. Phylogenetic relationships of Triassic Sauropterygia have always been unstable and recently questioned. Although specimens occur in high numbers, the main problems are rareness of diagnostic material from the Germanic Basin and uniformity of postcranial morphology of eosauropterygians. In the current paper, morphotypes of humeri along with their corresponding bone histologies for Lower to Middle Muschelkalk sauropterygians are described and interpreted for the first time in a phylogenetic context.Methodology/Principal Findings
Nothosaurus shows a typical plesiomorphic lamellar-zonal bone type, but varying growth patterns and the occurrence of a new humerus morphotype point to a higher taxonomic diversity than was known. In contrast to the enormous morphological variability of eosauropterygian humeri not assigned to Nothosaurus, their long bone histology is relatively uniform and can be divided into two histotypes. Unexpectedly, both of these histotypes reveal abundant fibrolamellar bone throughout the cortex. This pushes the origin of fibrolamellar bone in Sauropterygia back from the Cretaceous to the early Middle Triassic (early Anisian). Histotype A is assigned to Cymatosaurus, a basal member of the Pistosauroidea, which includes the plesiosaurs as derived members. Histotype B is related to the pachypleurosaur Anarosaurus. Contrary to these new finds, the stratigraphically younger pachypleurosaur Neusticosaurus shows the plesiomorphic lamellar-zonal bone type and an incomplete endochondral ossification, like Nothosaurus.Conclusions/SignificanceHistological results hypothetically discussed in a phylogenetical context have a large impact on the current phylogenetic hypothesis of Sauropterygia, leaving the pachypleurosaurs polyphyletic. On the basis of histological data, Neusticosaurus would be related to Nothosaurus, whereas Anarosaurus would follow the pistosaur clade. Furthermore, the presence of fibrolamellar bone, which is accompanied with increased growth rates and presumably even with increased metabolic rates, already in Anarosaurus and Cymatosaurus can explain the success of the Pistosauroidea, the only sauropterygian group to survive into the Jurassic and give rise to the pelagic plesiosaur radiation.
Long-bone histology indicates that the most common early dinosaur, the prosauropod Plateosaurus engelhardti from the Upper Triassic of Central Europe, had variable life histories. Although Plateosaurus grew at the fast rates typical for dinosaurs, as indicated by fibrolamellar bone, qualitative (growth stop) and quantitative (growth-mark counts) features of its histology are poorly correlated with body size. Individual life histories of P. engelhardti were influenced by environmental factors, as in modern ectothermic reptiles, but not in mammals, birds, or other dinosaurs.
Sauropterygia was the most successful marine reptile radiation in history, spanning almost the entire Mesozoic and exploiting a wide range of habitats and ecological niches. Here we report a new, exceptionally preserved skull of a juvenile stem placodont from the early Middle Triassic of the Netherlands, thus indicating a western Tethyan (European) origin for Placodontia, the most basal group of sauropterygians. A single row of teeth on an enlarged palatine supports this close relationship, although these are small and pointed instead of broad and flat, as is the case in placodonts, which demonstrate the strongest adaptation to a durophagous diet known in any reptile. Peg-like, slightly procumbent premaxillary teeth and an 'L-shaped' jugal also confirm a close relationship to basal placodonts. The new taxon provides insight into the evolution of placodont dentition, representing a transitional morphology between the plesiomorphic diapsid condition of palatal denticles and the specialized crushing teeth of placodonts.
Numerous amniote groups adapted to an aquatic life. This change of habitat naturally led to numerous convergences. The various adaptive traits vary depending on the degree of adaptation to an aquatic life, notably between shallow water taxa still able to occasionally locomote on land and open-marine forms totally independent from the terrestrial environment, but also between surface swimmers and deep divers. As a consequence, despite convergences, there is a high diversity within aquatic amniotes in e.g., shape, size, physiology, swimming mode. Bone microanatomy is considered to be strongly associated with bone biomechanics and is thus a powerful tool to understand bone adaptation to functional constraints and to make functional inferences on extinct taxa. Two opposing major microanatomical specializations have been described in aquatic amniotes, referred to as bone mass increase and a spongious organization, respectively. They are assumed to be essentially linked with the hydrostatic or hydrodynamic control of buoyancy and body trim and with swimming abilities and velocity. However, between extremes in these specializations, a wide range of intermediary patterns occurs. The present study provides a state-of-the-art review of these inner bone adaptations in semi-aquatic and aquatic amniotes. The analysis of the various microanatomical patterns observed in long bones, vertebrae, and ribs of a large sample of (semi-)aquatic extant and extinct amniotes reveals the wide diversity in microanatomical patterns and the variation in combination of these different patterns within a single skeleton. This enables us to discuss the link between microanatomical features and habitat, swimming abilities, and thus functional requirements in the context of amniote adaptation to an aquatic lifestyle.
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