section_path: History of study/Soft tissue and molecular preservation
section_name: Soft tissue and molecular preservation
paragraph_text: Soft tissue and molecular preservation
  
  Dinosaur fossils are not limited to bones, but also include imprints or mineralized remains of skin coverings, organs, and other tissues. Of these, skin coverings based on keratin proteins are most easily preserved because of their cross-linked, hydrophobic molecular structure. Fossils of keratin-based skin coverings or bony skin coverings are known from most major groups of dinosaurs. Dinosaur fossils with scaly skin impressions have been found since the 19th century. Samuel Beckles discovered a sauropod forelimb with preserved skin in 1852 that was incorrectly attributed to a crocodile; it was correctly attributed by Marsh in 1888 and subject to further study by Reginald Hooley in 1917. Among ornithischians, in 1884 Jacob Wortman found skin impressions on the first known specimen of Edmontosaurus annectens, which were largely destroyed during the specimen's excavation. Owen and Hooley subsequently described skin impressions of Hypsilophodon and Iguanodon in 1885 and 1917. Since then, scale impressions have been most frequently found among hadrosaurids, where the impressions are known from nearly the entire body across multiple specimens.
  
  Starting from the 1990s, major discoveries of exceptionally preserved fossils in deposits known as conservation Lagerstätten contributed to research on dinosaur soft tissues. Chiefly among these were the rocks that produced the Jehol (Early Cretaceous) and Yanliao (Mid-to-Late Jurassic) biotas of northeastern China, from which hundreds of dinosaur specimens bearing impressions of feather-like structures (both closely related to birds and otherwise, see § Origin of birds) have been described by Xing Xu and colleagues. In living reptiles and mammals, pigment-storing cellular structures known as melanosomes are partially responsible for producing colouration. Both chemical traces of melanin and characteristically shaped melanosomes have been reported from feathers and scales of Jehol and Yanliao dinosaurs, including both theropods and ornithischians. This has enabled multiple full-body reconstructions of dinosaur colouration, such as for Sinosauropteryx and Psittacosaurus by Jakob Vinther and colleagues, and similar techniques have also been extended to dinosaur fossils from other localities. (However, some researchers have also suggested that fossilized melanosomes represent bacterial remains.) Stomach contents in some Jehol and Yanliao dinosaurs closely related to birds have also provided indirect indications of diet and digestive system anatomy (e.g., crops). More concrete evidence of internal anatomy has been reported in Scipionyx from the Pietraroja Plattenkalk of Italy. It preserves portions of the intestines, colon, liver, muscles, and windpipe.
  
  Concurrently, a line of work led by Mary Higby Schweitzer, Jack Horner, and colleagues reported various occurrences of preserved soft tissues and proteins within dinosaur bone fossils. Various mineralized structures that likely represented red blood cells and collagen fibres had been found by Schweitzer and others in tyrannosaurid bones as early as 1991. However, in 2005, Schweitzer and colleagues reported that a femur of Tyrannosaurus preserved soft, flexible tissue within, including blood vessels, bone matrix, and connective tissue (bone fibers) that had retained their microscopic structure. This discovery suggested that original soft tissues could be preserved over geological time, with multiple mechanisms having been proposed. Later, in 2009, Schweitzer and colleagues reported that a Brachylophosaurus femur preserved similar microstructures, and immunohistochemical techniques (based on antibody binding) demonstrated the presence of proteins such as collagen, elastin, and laminin. Both specimens yielded collagen protein sequences that were viable for molecular phylogenetic analyses, which grouped them with birds as would be expected. The extraction of fragmentary DNA has also been reported for both of these fossils, along with a specimen of Hypacrosaurus. In 2015, Sergio Bertazzo and colleagues reported the preservation of collagen fibres and red blood cells in eight Cretaceous dinosaur specimens that did not show any signs of exceptional preservation, indicating that soft tissue may be preserved more commonly than previously thought. Suggestions that these structures represent bacterial biofilms have been rejected, but cross-contamination remains a possibility that is difficult to detect.