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      Description and etiology of paleopathological lesions in the type specimen of Parasaurolophus walkeri (Dinosauria: Hadrosauridae), with proposed reconstructions of the nuchal ligament

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          Abstract

          Paleopathology, or the study of ancient injuries and diseases, can enable the ecology and life history of extinct taxa to be deciphered. Large‐bodied ornithopods are the dinosaurs with the highest frequencies of paleopathology reported to‐date. Among these, the crested hadrosaurid Parasaurolophus walkeri is one of the most famous, largely due to its dramatic elongated and tubular nasal crest. The holotype of Parasaurolophus walkeri at the Royal Ontario Museum, Canada, displays several paleopathologies that have not been discussed in detail previously: a dental lesion in the left maxilla, perhaps related to periodontal disease; callus formation associated with fractures in three dorsal ribs; a discoidal overgrowth above dorsal neural spines six and seven; a cranially oriented spine in dorsal seven, that merges distally with spine six; a V‐shaped gap between dorsal spines seven and eight; and a ventral projection of the pubic process of the ilium which covers, and is fused with, the lateral side of the iliac process of the pubis. These lesions suggest that the animal suffered from one or more traumatic events, with the main one causing a suite of injuries to the anterior aspect of the thorax. The presence of several lesions in a single individual is a rare observation and, in comparison with a substantial database of hadrosaur paleopathological lesions, has the potential to reveal new information about the biology and behavior of these ornithopods. The precise etiology of the iliac abnormality is still unclear, although it is thought to have been an indirect consequence of the anterior trauma. The discoidal overgrowth above the two neural spines also seems to be secondary to the severe trauma inflicted on the ribs and dorsal spines, and probably represents post‐traumatic ossification of the base of the nuchal ligament. The existence of this structure has previously been considered in hadrosaurs and dinosaurs more generally through comparison of origin and insertion sites in modern diapsids ( Rhea americana, Alligator mississippiensis, Iguana iguana), but its presence, structure, and origin‐attachment sites are still debated. The V‐shaped gap is hypothesized as representing the point between the stresses of the nuchal ligament, pulling the anterior neural spines forward, and the ossified tendons pulling the posterior neural spines backward. Different reconstructions of the morphology of the structure based on the pathological conditions affecting the neural spines of ROM 768 are proposed. Finally, we review the history of reconstructions for Parasaurolophus walkeri showing how erroneous misconceptions have been perpetuated over time or have led to the development of new hypotheses, including the wide neck model supported in the current research.

          Abstract

          • The fossilized lesions in Parasaurolophus walkeri comprise periodontal disease, fractured ribs, and myositis ossificans traumatica in a number of vertebral dorsal spines and in the ilium.

          • The injuries appear to have been caused by one or more non‐threatening events.

          • The discoidal overgrowth on the dorsal spines m ight represent a pathological correlate for the origin of the nuchal ligament thereby enabling the morphology of the neck to be ascertained.

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          Most cited references101

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          The biology of fracture healing.

          The biology of fracture healing is a complex biological process that follows specific regenerative patterns and involves changes in the expression of several thousand genes. Although there is still much to be learned to fully comprehend the pathways of bone regeneration, the over-all pathways of both the anatomical and biochemical events have been thoroughly investigated. These efforts have provided a general understanding of how fracture healing occurs. Following the initial trauma, bone heals by either direct intramembranous or indirect fracture healing, which consists of both intramembranous and endochondral bone formation. The most common pathway is indirect healing, since direct bone healing requires an anatomical reduction and rigidly stable conditions, commonly only obtained by open reduction and internal fixation. However, when such conditions are achieved, the direct healing cascade allows the bone structure to immediately regenerate anatomical lamellar bone and the Haversian systems without any remodelling steps necessary. In all other non-stable conditions, bone healing follows a specific biological pathway. It involves an acute inflammatory response including the production and release of several important molecules, and the recruitment of mesenchymal stem cells in order to generate a primary cartilaginous callus. This primary callus later undergoes revascularisation and calcification, and is finally remodelled to fully restore a normal bone structure. In this article we summarise the basic biology of fracture healing. Copyright © 2011 Elsevier Ltd. All rights reserved.
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            Hadrosauridae

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              From the archives of the AFIP. Extraskeletal osseous and cartilaginous tumors of the extremities.

              Extraskeletal osseous and cartilaginous tumors and tumorlike conditions of the extremities can often be differentiated radiologically; for those that cannot, knowledge of the spectrum of lesions will allow a suitably ordered differential diagnosis. Of the osseous lesions--myositis ossificans, fibro-osseous pseudotumor, fibrodysplasia ossificans progressiva, soft-tissue osteoma, and extraskeletal osteosarcoma--all but myositis ossificans are relatively rare. Myositis ossificans has a distinct mineralization pattern that can be observed radiologically as a peripheral rim of lamellar bone. Fibro-osseous pseudotumor typically occurs in the digits of the hand and lacks the well-defined zoning pattern of myositis ossificans. The cartilaginous entities include the true tumors, soft-tissue chondroma and extraskeletal chondrosarcoma, and the tumorlike process, synovial osteochondromatosis. The tumors are relatively rare; synovial osteochondromatosis commonly affects middle-aged men, especially in the knee, and is associated with osteoarthritis. The differential diagnosis for these extraskeletal osseous and cartilaginous lesions includes soft-tissue sarcoma, benign mesenchymoma, malignant mesenchymoma (rare), calcified tophi in gout, melorheostosis (rare), pilomatricoma (rare), and tumoral calcinosis (rare).
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                Author and article information

                Contributors
                f.bertozzo@qub.ac.uk
                Journal
                J Anat
                J Anat
                10.1111/(ISSN)1469-7580
                JOA
                Journal of Anatomy
                John Wiley and Sons Inc. (Hoboken )
                0021-8782
                1469-7580
                01 December 2020
                May 2021
                01 December 2020
                : 238
                : 5 ( doiID: 10.1111/joa.v238.5 )
                : 1055-1069
                Affiliations
                [ 1 ] School of Natural and Built Environment Queen’s University Belfast Belfast UK
                [ 2 ] CI2Paleo Sociedade de Historia Natural Torres Vedras Portugal
                [ 3 ] Associazione Paleontologica Paleoartistica Italiana Parma Italy
                [ 4 ] Department of Musculoskeletal & Ageing Science University of Liverpool Liverpool UK
                [ 5 ] Royal Tyrrell Museum of Paleontology Drumheller AB Canada
                [ 6 ] Royal Ontario Museum Toronto ON Canada
                [ 7 ] Ecology and Evolutionary Biology University of Toronto Toronto ON Canada
                Author notes
                [*] [* ] Correspondence

                Filippo Bertozzo, School of Natural and Built Environment, Queen’s University Belfast, Belfast, Northern Ireland, UK.

                Email: f.bertozzo@ 123456qub.ac.uk

                Author information
                https://orcid.org/0000-0002-0917-9989
                Article
                JOA13363
                10.1111/joa.13363
                8053592
                33289113
                67e4d17a-dd56-439b-99a5-b6d9f05726a5
                © 2020 The Authors. Journal of Anatomy published by John Wiley & Sons Ltd on behalf of Anatomical Society

                This is an open access article under the terms of the http://creativecommons.org/licenses/by-nc/4.0/ License, which permits use, distribution and reproduction in any medium, provided the original work is properly cited and is not used for commercial purposes.

                History
                : 23 October 2020
                : 25 August 2020
                : 30 October 2020
                Page count
                Figures: 9, Tables: 0, Pages: 15, Words: 12166
                Funding
                Funded by: Dinosaur Research Institute Student Project
                Award ID: 2018
                Funded by: Horizon 2020 research and innovation programme under the MSCA grant agreement
                Award ID: 754507
                Categories
                Original Paper
                Original Papers
                Custom metadata
                2.0
                May 2021
                Converter:WILEY_ML3GV2_TO_JATSPMC version:6.0.2 mode:remove_FC converted:18.04.2021

                Anatomy & Physiology
                alberta,cretaceous,nuchal ligament,ornithopoda,trauma
                Anatomy & Physiology
                alberta, cretaceous, nuchal ligament, ornithopoda, trauma

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