Comparative bone histology of two thalattosaurians (Diapsida: Thalattosauria): Askeptosaurus italicus from the Alpine Triassic (Middle Triassic) and a Thalattosauroidea indet. from the Carnian of Oregon (Late Triassic).
N Klein, P M Sander, J Liu, P Druckenmiller, E T Metz, N P Kelley, T M Scheyer
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Abstract
Here, we present the first bone histological and microanatomical study of thalattosaurians, an enigmatic group among Triassic marine reptiles. Two taxa of thalattosaurians, the askeptosauroid Askeptosaurus italicus and one as yet undescribed thalattosauroid, are examined. Both taxa have a rather different microanatomy, tissue type, and growth pattern. Askeptosaurus italicus from the late Anisian middle Besano Formation of the southern Alpine Triassic shows very compact tissue in vertebrae, rib, a gastralium, and femora, and all bones are without medullary cavities. The tissue shows moderate to low vascularization, dominated by highly organized and very coarse parallel-fibred bone, resembling interwoven tissue. Vascularization is dominated by simple longitudinal vascular canals, except for the larger femur of Askeptosaurus, where simple vascular canals dominate in a radial arrangement. Growth marks stratify the cortex of femora. The vertebrae and humeri from the undescribed thalattosauroid from the late Carnian of Oregon have primary and secondary cancellous bone, resulting in an overall low bone compactness. Two dorsal vertebral centra show dominantly secondary trabeculae, whereas a caudal vertebral centrum shows much primary trabecular bone, globuli ossei, and cartilage, indicating an earlier ontogenetic stage of the specimens or paedomorphosis. The humeri of the thalattosauroid show large, simple vascular canals that are dominantly radially oriented in a scaffold of woven and loosely organized parallel-fibred tissue. Few of the simple vascular canals are thinly but only incompletely lined by parallel-fibered tissue. In the Oregon material, changes in growth rate are only indicated by changes in vascular organization but no distinct growth marks were identified. The compact bone of Askeptosaurus is best comparable to some pachypleurosaurs, whereas its combination of tissue and vascularity is similar to eosauropterygians in general, except for the coarse nature of its parallel-fibred tissue. The cancellous bone of the Oregon thalattosauroid resembles what is documented in ichthyosaurs and plesiosaurs. However, in contrast to these its tissue does not consist of fibro-lamellar bone type. Tissue types of both thalattosaurian taxa indicate rather different growth rates and growth patterns, associated with different life history strategies. The microanatomy reflects different life styles that fit to the different environments in which they had been found (intraplatform basin vs. open marine). Both thalattosaurian taxa differ from each other but in sum also from all other marine reptile taxa studied so far. Thalattosaurian bone histology documents once more that bone histology provides for certain groups (i.e., Triassic Diapsida) only a poor phylogenetic signal and is more influenced by exogenous factors. Differences in lifestyle, life history traits, and growth rate and pattern enabled all these Triassic marine reptiles to live contemporaneously in the same habitat managing to avoid substantial competition.
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