New insights on the systematics, palaeoecology and palaeobiology of a plesiosaurian with soft tissue preservation from the Toarcian of Holzmaden, Germany

The Posidonienschiefer Formation (Toarcian) of Holzmaden, Baden-Wurttemberg in southwestern Germany has yielded several excellently preserved plesiosaurian specimens and received considerable research attention. The plesiosaurians found within these deposits are always significantly outnumbered by ichthyosaurs, and close examination of these rare specimens is crucial to a better understanding of the diversity and palaeoecology of Plesiosauria in this very peculiar ecosystem. The plesiosaurian specimen SMNS 51945 found in this area is a juvenile individual consisting of a partial, crushed skull and an exquisitely preserved post-cranial skeleton. Its anatomical characters seem to differ from the long-necked plesiosauroids Microcleidus brachypterygius and Seeleyosaurus guilelmiimperatoris that are the most abundant taxa within the plesiosaurian assemblage. The post-cranial skeleton preserves very likely soft tissues composed of buff-coloured and dark-coloured structures around the vertebral column and hindlimb of the animal. A network of buff-coloured fibres located posterior to the hindlimb most likely represents phosphatised collagen fibres as already found in some ichthyosaur specimens, confirming that wing area in plesiosaurians was much larger than that suggested by skeletal remains alone. The specimen also contains gastroliths (sand-sized grains mainly composed of quartz) in the stomach cavity suggesting the animal spent at least some of its time in shallow coastal waters, tens or hundreds of kilometres from the final place of burial