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<article xmlns:tp="http://www.plazi.org/taxpub" xmlns:xsi="http://www.w3.org/2001/XMLSchema-instance" xmlns:xlink="http://www.w3.org/1999/xlink" xmlns:mml="http://www.w3.org/1998/Math/MathML" article-type="research-article" xml:lang="en">
  <front>
    <journal-meta>
      <journal-id journal-id-type="publisher-id">114</journal-id>
      <journal-id journal-id-type="index">urn:lsid:arphahub.com:pub:6666f368-0968-5880-b34a-cf8d3de58b35</journal-id>
      <journal-id journal-id-type="aggregator">urn:lsid:zoobank.org:pub:A96E9753-134C-4D08-A2E3-31C66B0459C6</journal-id>
      <journal-title-group>
        <journal-title xml:lang="en">Fossil Record</journal-title>
        <abbrev-journal-title xml:lang="en">foss-rec</abbrev-journal-title>
      </journal-title-group>
      <issn pub-type="ppub">2193-0066</issn>
      <issn pub-type="epub">2193-0074</issn>
      <publisher>
        <publisher-name>Pensoft Publishers</publisher-name>
      </publisher>
    </journal-meta>
    <article-meta>
      <article-id pub-id-type="doi">10.3897/fr.29.193112</article-id>
      <article-id pub-id-type="publisher-id">193112</article-id>
      <article-categories>
        <subj-group subj-group-type="heading">
          <subject>Research Article</subject>
        </subj-group>
        <subj-group subj-group-type="biological_taxon">
          <subject>Animalia</subject>
        </subj-group>
        <subj-group subj-group-type="scientific_subject">
          <subject>Vertebrate palaeontology</subject>
        </subj-group>
        <subj-group subj-group-type="geological_era">
          <subject>Mesozoic</subject>
        </subj-group>
        <subj-group subj-group-type="geographical_area">
          <subject>Americas</subject>
        </subj-group>
      </article-categories>
      <title-group>
        <article-title>A case of an expansile lesion in a caudal vertebra of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Medusaceratops">Medusaceratops</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="lokii">lokii</tp:taxon-name-part></tp:taxon-name></italic></article-title>
      </title-group>
      <contrib-group content-type="authors">
        <contrib contrib-type="author" corresp="no">
          <name name-style="western">
            <surname>Wang</surname>
            <given-names>Donghao</given-names>
          </name>
          <xref ref-type="aff" rid="A1">1</xref>
          <xref ref-type="aff" rid="A2">2</xref>
        </contrib>
        <contrib contrib-type="author" corresp="yes">
          <name name-style="western">
            <surname>Xing</surname>
            <given-names>Lida</given-names>
          </name>
          <email xlink:type="simple">xinglida@gmail.com</email>
          <uri content-type="orcid">https://orcid.org/0000-0003-3923-9206</uri>
          <xref ref-type="aff" rid="A3">3</xref>
          <xref ref-type="aff" rid="A4">4</xref>
        </contrib>
        <contrib contrib-type="author" corresp="no">
          <name name-style="western">
            <surname>Rothschild</surname>
            <given-names>Bruce M.</given-names>
          </name>
          <uri content-type="orcid">https://orcid.org/0000-0003-1327-6615</uri>
          <xref ref-type="aff" rid="A5">5</xref>
        </contrib>
        <contrib contrib-type="author" corresp="no">
          <name name-style="western">
            <surname>Du</surname>
            <given-names>Chunlei</given-names>
          </name>
          <xref ref-type="aff" rid="A4">4</xref>
        </contrib>
        <contrib contrib-type="author" corresp="no">
          <name name-style="western">
            <surname>Wen</surname>
            <given-names>Kexiang</given-names>
          </name>
          <xref ref-type="aff" rid="A4">4</xref>
        </contrib>
        <contrib contrib-type="author" corresp="no">
          <name name-style="western">
            <surname>Lin</surname>
            <given-names>Jin</given-names>
          </name>
          <xref ref-type="aff" rid="A4">4</xref>
        </contrib>
        <contrib contrib-type="author" corresp="no">
          <name name-style="western">
            <surname>Kang</surname>
            <given-names>Mengling</given-names>
          </name>
          <xref ref-type="aff" rid="A4">4</xref>
        </contrib>
        <contrib contrib-type="author" corresp="no">
          <name name-style="western">
            <surname>Su</surname>
            <given-names>Jiayin</given-names>
          </name>
          <xref ref-type="aff" rid="A6">6</xref>
        </contrib>
      </contrib-group>
      <aff id="A1">
        <label>1</label>
        <addr-line content-type="verbatim">Mineral Resources Survey Division, Harbin Center for Natural Resources Comprehensive Survey, China Geological Survey, Harbin 150039, Heilongjiang, China</addr-line>
        <institution>Mineral Resources Survey Division, Harbin Center for Natural Resources Comprehensive Survey, China Geo-logical Survey</institution>
        <addr-line content-type="city">Harbin</addr-line>
        <country>China</country>
      </aff>
      <aff id="A2">
        <label>2</label>
        <addr-line content-type="verbatim">Observation and Research Station of Earth Critical Zone in Black Soil, Ministry of Natural Resources, Harbin 150086, China</addr-line>
        <institution>Observation and Research Station of Earth Critical Zone in Black Soil, Ministry of Natural Resources</institution>
        <addr-line content-type="city">Harbin</addr-line>
        <country>China</country>
      </aff>
      <aff id="A3">
        <label>3</label>
        <addr-line content-type="verbatim">State Key Laboratory of Geomicrobiology and Environmental Changes, School of Earth Sciences and Resources, Frontiers Science Center for Deep-time Digital Earth, China University of Geosciences, Beijing, 100083, China</addr-line>
        <institution>State Key Laboratory of Geomicrobiology and Environmental Changes, School of Earth Sciences and Resources, Frontiers Science Center for Deep-time Digital Earth, China University of Geosciences</institution>
        <addr-line content-type="city">Beijin</addr-line>
        <country>China</country>
      </aff>
      <aff id="A4">
        <label>4</label>
        <addr-line content-type="verbatim">Grandview Museum of Natural Science, Guangzhou, 510620, China</addr-line>
        <institution>Grandview Museum of Natural Science</institution>
        <addr-line content-type="city">Guangzhou</addr-line>
        <country>China</country>
      </aff>
      <aff id="A5">
        <label>5</label>
        <addr-line content-type="verbatim">Department of Medicine, Indiana University Health, Muncie, IN 47303, USA</addr-line>
        <institution>Department of Medicine, Indiana University Health</institution>
        <addr-line content-type="city">Muncie</addr-line>
        <country>United States of America</country>
      </aff>
      <aff id="A6">
        <label>6</label>
        <addr-line content-type="verbatim">Department of Plastic Surgery, Fengcheng Hospital, Shanghai, 201411, China</addr-line>
        <institution>Department of Plastic Surgery, Fengcheng Hospital</institution>
        <addr-line content-type="city">Shanghai</addr-line>
        <country>China</country>
      </aff>
      <author-notes>
        <fn fn-type="corresp">
          <p>Corresponding author: Lida Xing (<email xlink:type="simple">xinglida@gmail.com</email>)</p>
        </fn>
        <fn fn-type="edited-by">
          <p>Academic editor: Florian Witzmann</p>
        </fn>
      </author-notes>
      <pub-date pub-type="collection">
        <year>2026</year>
      </pub-date>
      <pub-date pub-type="epub">
        <day>06</day>
        <month>08</month>
        <year>2026</year>
      </pub-date>
      <volume>29</volume>
      <issue>2</issue>
      <fpage>599</fpage>
      <lpage>610</lpage>
      <uri content-type="arpha" xlink:href="http://openbiodiv.net/5687CF75-0092-569D-8DEB-7C674440773F">5687CF75-0092-569D-8DEB-7C674440773F</uri>
      <uri content-type="zoobank" xlink:href="https://zoobank.org/40D48356-BEF2-416D-ADFE-4DF6407AF284">40D48356-BEF2-416D-ADFE-4DF6407AF284</uri>
      <history>
        <date date-type="received">
          <day>27</day>
          <month>03</month>
          <year>2026</year>
        </date>
        <date date-type="accepted">
          <day>24</day>
          <month>07</month>
          <year>2026</year>
        </date>
      </history>
      <permissions>
        <license license-type="creative-commons-attribution" xlink:href="https://creativecommons.org/share-your-work/public-domain/cc0/" xlink:type="simple">
          <license-p>This is an open access article distributed under the terms of the CC0 Public Domain Dedication.</license-p>
        </license>
      </permissions>
      <self-uri content-type="zoobank" xlink:type="simple">https://zoobank.org/40D48356-BEF2-416D-ADFE-4DF6407AF284</self-uri>
      <abstract>
        <label>Abstract</label>
        <p>This study presents the first documented case of chronic osteomyelitis with a draining sinus tract in the centrosaurine ceratopsid <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Medusaceratops">Medusaceratops</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="lokii">lokii</tp:taxon-name-part></tp:taxon-name></italic>, based on a pathological seventh caudal vertebra from the Upper Cretaceous Judith River Formation of Montana. Integrated macroscopic and computed tomography analyses revealed pathognomonic features of chronic osteomyelitis secondary to trauma, including extensive hyperostosis with “cauliflower-shaped” bone overgrowths, moth-eaten osteolysis, sclerosis, a definitive sinus tract, and occult fractures associated with the destruction of the bilateral transverse processes. The development of a substantial involucrum confirms prolonged post-traumatic survival, attesting to robust immunological resilience in ceratopsid dinosaurs. A review of the 81 reported cases of dinosaurian infection indicates that osteomyelitis represents the predominant lesion type and that the great majority of cases are post-traumatic. Bone infection therefore appears to have been seeded principally through traumatic breaches rather than arising chiefly via non-traumatic routes, and the anatomical distribution of infectious lesions also differs among the major clades.</p>
      </abstract>
      <kwd-group>
        <label>Key Words</label>
        <kwd>
          <tp:taxon-name>
            <tp:taxon-name-part taxon-name-part-type="family" reg="Ceratopsidae">Ceratopsidae</tp:taxon-name-part>
          </tp:taxon-name>
        </kwd>
        <kwd>computed tomography</kwd>
        <kwd>Judith River Formation</kwd>
        <kwd>osteomyelitis</kwd>
        <kwd>paleopathology</kwd>
      </kwd-group>
      <funding-group>
        <funding-statement>This research was funded by 2025 Heilongjiang Provincial Natural Science Foundation Youth Project Type C (JJ2025QC0704), the National Natural Science Foundation of China (No. 42488201), the “Deep-time Digital Earth” Science and Technology Leading Talents Team Funds for the Central Universities for the Frontiers Science Center for Deep-time Digital Earth, China University of Geosciences (Beijing) (Fundamental Research Funds for the Central Universities; grant number: 2652023001), the 111 project (B20011) and Fundamental Research Funds for Central Universities (265QZ201903).</funding-statement>
      </funding-group>
    </article-meta>
  </front>
  <body>
    <sec sec-type="Introduction" id="sec1">
      <title>Introduction</title>
      <p>Paleopathology, an interdisciplinary field bridging paleontology and medicine, occupies an increasingly pivotal position within vertebrate paleontology. Researchers have identified evidence of antemortem pathologies in fossilized skeletal remains (e.g., <xref ref-type="bibr" rid="B58">Xing et al. 2024</xref>), eggs (e.g., <xref ref-type="bibr" rid="B23">Jackson et al. 2004</xref>), integument (e.g., <xref ref-type="bibr" rid="B40">Rothschild and DePalma 2013</xref>), and even ichnofossils (e.g., <xref ref-type="bibr" rid="B19">Herrera-Castillo et al. 2022</xref>). Non-avian dinosaurs, in particular, have been the subject of a growing number of case reports. These pathologies are diverse, encompassing fractures (e.g., <xref ref-type="bibr" rid="B54">Xing et al. 2009</xref>), infections (e.g., <xref ref-type="bibr" rid="B42">Rothschild et al. 2023</xref>), neoplasms (e.g., <xref ref-type="bibr" rid="B10">Ekhtiari et al. 2020</xref>), ankylosis secondary to arthritis (e.g., <xref ref-type="bibr" rid="B56">Xing et al. 2015</xref>), hyperostosis (e.g., <xref ref-type="bibr" rid="B13">Foth et al. 2015</xref>), dental disease (e.g., <xref ref-type="bibr" rid="B55">Xing et al. 2013</xref>), and congenital and developmental anomalies (<xref ref-type="bibr" rid="B2">Baiano et al. 2024</xref>), among others.</p>
      <p>Among these conditions, cases of infection are documented relatively frequently. Infection is defined as the invasion of a host organism by pathogenic microorganisms—such as bacteria, viruses, fungi, or parasites—and the subsequent elicitation of an immunological cascade (<xref ref-type="bibr" rid="B3">Bennett et al. 2019</xref>). This process typically results in tissue damage and induces an inflammatory response; associated clinical manifestations may include erythema, pyrexia, pain, and functional impairment (<xref ref-type="bibr" rid="B30">Murphy and Weaver 2016</xref>). Pathogens may enter the host via various routes, including the respiratory tract, gastrointestinal tract, or compromised integument (<xref ref-type="bibr" rid="B5">Cohen et al. 2016</xref>). In pathological contexts, infections in the fossil record are often discernible through osseous lesions. For instance, pathological hyperostosis (<xref ref-type="bibr" rid="B48">Tanke and Rothschild 2002</xref>) or erosive features visible in computed tomography (<abbrev xlink:title="computed tomography">CT</abbrev>) imagery (<xref ref-type="bibr" rid="B57">Xing et al. 2018</xref>) may imply past infection, typically manifesting as abnormal cortical thickening or irregular surface bone overgrowths (<xref ref-type="bibr" rid="B48">Tanke and Rothschild 2002</xref>). The analysis of such lesions facilitates inferences regarding dinosaurian behavior, environmental stressors, and the evolution of the immune system (<xref ref-type="bibr" rid="B42">Rothschild et al. 2023</xref>). As of June 2026, 81 dinosaurian cases involving infection have been scientifically reported, comprising 20 in <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="suborder" reg="Cerapoda">Cerapoda</tp:taxon-name-part></tp:taxon-name>, nine in <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="clade" reg="Thyreophora">Thyreophora</tp:taxon-name-part></tp:taxon-name>, 13 in <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="order" reg="Sauropodomorpha">Sauropodomorpha</tp:taxon-name-part></tp:taxon-name>, and 39 in <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="suborder" reg="Theropoda">Theropoda</tp:taxon-name-part></tp:taxon-name> (Suppl. material <xref ref-type="supplementary-material" rid="S1">1</xref>).</p>
      <p><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="clade" reg="Ceratopsia">Ceratopsia</tp:taxon-name-part></tp:taxon-name> represents a significant clade of Cretaceous herbivorous dinosaurs; their distinctive cranial frills and facial horns likely functioned in defense, as well as in courtship and intraspecific competition (<xref ref-type="bibr" rid="B33">Padian and Horner 2011</xref>). Existing research indicates that various ceratopsians exhibited a spectrum of pathologies. For example, a fibular fracture has been documented in <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Psittacosaurus">Psittacosaurus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="lujiatunensis">lujiatunensis</tp:taxon-name-part></tp:taxon-name></italic> (<xref ref-type="bibr" rid="B18">Hedrick et al. 2016</xref>), while <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Psittacosaurus">Psittacosaurus</tp:taxon-name-part></tp:taxon-name></italic> sp. has exhibited signs of fibular necrosis (<xref ref-type="bibr" rid="B28">Lü et al. 2007</xref>). Abnormal bone resorption was observed on the supraorbital horn of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Centrosaurus">Centrosaurus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="apertus">apertus</tp:taxon-name-part></tp:taxon-name></italic> (<xref ref-type="bibr" rid="B44">Ryan et al. 2001</xref>), alongside a rare case of osteosarcoma (<xref ref-type="bibr" rid="B10">Ekhtiari et al. 2020</xref>). Fenestrae on the squamosals of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Diabloceratops">Diabloceratops</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="eatoni">eatoni</tp:taxon-name-part></tp:taxon-name></italic> and <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Pentaceratops">Pentaceratops</tp:taxon-name-part></tp:taxon-name></italic> sp., potentially attributable to trauma, have also been identified (<xref ref-type="bibr" rid="B49">Tanke and Farke 2006</xref>; <xref ref-type="bibr" rid="B24">Kirkland and DeBlieux 2010</xref>). <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Pachyrhinosaurus">Pachyrhinosaurus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="lakustai">lakustai</tp:taxon-name-part></tp:taxon-name></italic> exhibited fractures in the metacarpals (<xref ref-type="bibr" rid="B51">Vice 2020</xref>). Furthermore, the nasal boss of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Pachyrhinosaurus">P.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="perotorum">perotorum</tp:taxon-name-part></tp:taxon-name></italic> demonstrated pathological changes consistent with fractures and osteomyelitis, while traces of trauma and infection were identified on the caudal vertebrae (<xref ref-type="bibr" rid="B12">Fiorillo and Tykoski 2023</xref>). Although <xref ref-type="bibr" rid="B50">Tanke and Rothschild (2010)</xref> documented several caudal vertebral pathologies in ceratopsian dinosaurs from Alberta, none of the reported lesions is comparable to the expansile osteomyelitic lesion described in the present study. Punctures and fractures in the horns and frills of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Triceratops">Triceratops</tp:taxon-name-part></tp:taxon-name></italic> sp. are likely associated with combat behavior (<xref ref-type="bibr" rid="B31">Nabavizadeh 2023</xref>), whereas <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Vagaceratops">Vagaceratops</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="irvinensis">irvinensis</tp:taxon-name-part></tp:taxon-name></italic> displays signs of degenerative articular changes in the cervical vertebrae and metacarpals (<xref ref-type="bibr" rid="B21">Holmes et al. 2001</xref>; <xref ref-type="bibr" rid="B46">Sampson et al. 2010</xref>).</p>
      <p><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Medusaceratops">Medusaceratops</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="lokii">lokii</tp:taxon-name-part></tp:taxon-name></italic> was originally referred to as the basal centrosaurine <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Albertaceratops">Albertaceratops</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="nesmoi">nesmoi</tp:taxon-name-part></tp:taxon-name></italic> (<xref ref-type="bibr" rid="B43">Ryan 2007</xref>). Subsequently, <xref ref-type="bibr" rid="B45">Ryan et al. (2010)</xref> reassigned the material to <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="subfamily" reg="Chasmosaurinae">Chasmosaurinae</tp:taxon-name-part></tp:taxon-name> based on the two most complete parietals, establishing the new genus and species via an analysis of specimens distinguished by the presence of only three epiparietals on either side of the midline. <xref ref-type="bibr" rid="B27">Longrich (2013)</xref> later returned the taxon to <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="subfamily" reg="Centrosaurinae">Centrosaurinae</tp:taxon-name-part></tp:taxon-name>, arguing that the number of epiparietals is variable within <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="subfamily" reg="Chasmosaurinae">Chasmosaurinae</tp:taxon-name-part></tp:taxon-name> and that the morphology of the parietosquamosal junction bears greater similarity to that observed in <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="subfamily" reg="Centrosaurinae">Centrosaurinae</tp:taxon-name-part></tp:taxon-name>. <xref ref-type="bibr" rid="B4">Chiba et al. (2018)</xref> incorporated new fossil material alongside previously described specimens into a numerical cladistic analysis, the results of which supported Longrich’s (2013) conclusion. The available evidence indicates that <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Medusaceratops">Medusaceratops</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="lokii">lokii</tp:taxon-name-part></tp:taxon-name></italic> is currently regarded as a distinct taxon from <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Albertaceratops">Albertaceratops</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="nesmoi">nesmoi</tp:taxon-name-part></tp:taxon-name></italic>, representing the prevailing scientific consensus.</p>
    </sec>
    <sec sec-type="materials|methods" id="sec2">
      <title>Materials and methods</title>
      <p>The specimen described herein was recovered from light-colored sandstones of the Upper Cretaceous Judith River Formation near Joplin (Havre), Montana, USA. It is currently housed at the Grandview Museum of Natural Science in Guangzhou, China, under the accession number HS0385 (HS = Mandarin Pinyin abbreviation for “fossil”; Fig. <xref ref-type="fig" rid="F1">1</xref>). Specimen HS0385 derives from a nearly complete skeleton, although individual elements are incomplete or damaged, most notably portions of the skull; of the caudal series, only the seventh caudal vertebra is pathological.</p>
      <fig id="F1">
        <object-id content-type="doi">10.3897/fr.29.193112.figure1</object-id>
        <object-id content-type="arpha">16F81E8C-1912-5667-AC5D-66A2617B2CBC</object-id>
        <label>Figure 1.</label>
        <caption>
          <p>Mounted skeleton (<bold>A</bold>) and line drawing of the skeleton (<bold>B</bold>) of HS0385. The purple bone represents the pathological seventh caudal vertebra.</p>
        </caption>
        <graphic xlink:href="fr-29-599-g001.jpg" id="oo_1739212.jpg">
          <uri content-type="original_file">https://binary.pensoft.net/fig/1739212</uri>
        </graphic>
      </fig>
      <p>The morphological characteristics of HS0385 conform to the diagnosis of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Medusaceratops">Medusaceratops</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="lokii">lokii</tp:taxon-name-part></tp:taxon-name></italic> established by <xref ref-type="bibr" rid="B4">Chiba et al. (2018)</xref>. Specifically, it is identified as a centrosaurine ceratopsid possessing five epiparietals on the posterolateral parietal ramus, arranged medially to laterally as follows: epiparietal 1 (<abbrev xlink:title="epiparietal 1">ep 1</abbrev>) is small and slightly procurved; epiparietal 2 (<abbrev xlink:title="epiparietal 2">ep 2</abbrev>) is a comparatively large, broad-based pachyostotic projection that curves strongly anterolaterally; epiparietal 3 (<abbrev xlink:title="epiparietal 3">ep 3</abbrev>) forms a smaller pachyostotic process with an anterolateral curvature similar to that of <abbrev xlink:title="epiparietal 2">ep 2</abbrev>; and epiparietals 4 and 5 (<abbrev xlink:title="epiparietals 4">ep 4</abbrev> and <abbrev xlink:title="epiparietals 5">ep 5</abbrev>) are unmodified, manifesting as small, triangular ossifications. The pathology of HS0385 is present in the seventh caudal vertebra.</p>
      <p><abbrev xlink:title="computed tomography">CT</abbrev> imagery was acquired using an ANATOM P419 scanner (120 kV, 350 mA, 280 mAs), designed and manufactured by Shenzhen Anke High-tech Co., Ltd., Shenzhen, China. The external surface of the specimen was scanned using an Artec Spider structured-light scanner, and the acquired data were processed and reconstructed in Artec Studio.</p>
      <p>A dataset comprising 81 cases of dinosaurian infection was compiled (Suppl. material <xref ref-type="supplementary-material" rid="S1">1</xref>), and statistical analyses were performed in PAST v5.3 (<xref ref-type="bibr" rid="B16">Hammer et al. 2001</xref>) to test for associations between taxon or ecological niche and both trauma type and the anatomical location of infection. For 2 × 2 contingency tables, Fisher’s exact test was used, whereas larger contingency tables were analyzed using the chi-square test in combination with Monte Carlo permutation tests (999 replicates). A similar approach was employed by <xref ref-type="bibr" rid="B2">Baiano et al. (2024)</xref> to investigate the relationship between taxonomic affiliation and ecological niche. Effect sizes are reported as Cramér’s <italic>V</italic>, and the significance threshold was set at <italic>α</italic> = 0.05.</p>
    </sec>
    <sec sec-type="Results" id="sec3">
      <title>Results</title>
      <sec sec-type="Description" id="sec4">
        <title>Description</title>
        <p>The seventh caudal vertebra exhibits gross expansile pathological deformation. The lateral surfaces and posterior articular surface of the centrum are covered by reactive bone exhibiting a honeycomb-like texture. The anterior and posterior articular facets of the centrum remain discernible; the anterior facet measures 50.1 mm in width and 68.8 mm in height, while the posterior facet measures 55.8 mm in width and 61.7 mm in height (Fig. <xref ref-type="fig" rid="F2">2</xref>). The remaining caudal vertebrae are slightly smaller than the corresponding elements in <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Styracosaurus">Styracosaurus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="albertensis">albertensis</tp:taxon-name-part></tp:taxon-name></italic> (<xref ref-type="bibr" rid="B20">Holmes and Ryan 2013</xref>), whereas the seventh caudal vertebra is distinctly smaller than both its counterpart in <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Styracosaurus">S.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="albertensis">albertensis</tp:taxon-name-part></tp:taxon-name></italic> and the immediately adjacent sixth and eighth caudal vertebrae.</p>
        <fig id="F2">
          <object-id content-type="doi">10.3897/fr.29.193112.figure2</object-id>
          <object-id content-type="arpha">6452893D-A49D-588C-82D9-A80DD97A5BD0</object-id>
          <label>Figure 2.</label>
          <caption>
            <p>The seventh caudal vertebra of HS0385: <bold>A<sub>1</sub></bold>. Right lateral view of the 3D reconstruction; <bold>B<sub>1</sub></bold>. Cranial view of the 3D reconstruction; <bold>C<sub>1</sub></bold>. Caudal view of the 3D reconstruction; <bold>D<sub>1</sub></bold>. Left lateral view of the 3D reconstruction; <bold>A<sub>2</sub></bold>. Right lateral view; <bold>B<sub>2</sub></bold>. Cranial view; <bold>C<sub>2</sub></bold>. Caudal view; <bold>D<sub>2</sub></bold>. Left lateral view of the actual fossil. Light pink represents bone overgrowth No. 1, and teal represents bone overgrowth No. 2; red arrows indicate the honeycomb-like texture; black arrows indicate the remnant of the left transverse process. The sixth caudal vertebra of HS0385: <bold>E<sub>1</sub></bold>. Right lateral view of the 3D reconstruction; <bold>E<sub>2</sub></bold>. Left lateral view of the 3D reconstruction; and <bold>E<sub>3</sub></bold>. Caudal view of the 3D reconstruction. The 3D reconstructions (<bold>A<sub>1</sub>–D<sub>1</sub>, E<sub>1–3</sub></bold>) are surface models generated from data acquired using the Artec Spider scanner; <bold>A<sub>2</sub>–D<sub>2</sub></bold> are photographs of the corresponding views.</p>
          </caption>
          <graphic xlink:href="fr-29-599-g002.jpg" id="oo_1739213.jpg">
            <uri content-type="original_file">https://binary.pensoft.net/fig/1739213</uri>
          </graphic>
        </fig>
        <p>The vertebra is characterized by the conspicuous absence of the bilateral transverse processes (Fig. <xref ref-type="fig" rid="F2">2</xref>). A partial remnant of the left transverse process is preserved (Fig. <xref ref-type="fig" rid="F2">2D</xref><sub>1</sub>), measuring 27.6 mm in maximum width, 40.8 mm in height, and 35.4 mm in length. Its cranial cortical surface remains relatively smooth and continuous, whereas the remaining surfaces exhibit a honeycomb texture. This structure represents the vestige of a fractured transverse process, with evident hyperostosis at its terminus. The right side lacks the distinct fracture trace observed on the left (Fig. <xref ref-type="fig" rid="F2">2A</xref><sub>1</sub>); however, slightly dorsal to the relative position of the left process, there is an elliptical hyperostotic feature with raised margins and a depressed center, characterized by rough cortical bone. The dimensions and relative position of this elliptical trace are consistent with those of a transverse process; consequently, this area is interpreted as the vestige of the right transverse process.</p>
        <p>The centrum exhibits multiple exophytic bone overgrowths, two of which exceed 20 mm in width. Bone overgrowth No. 1 connects to the ventral surface of the centrum via a broad craniocaudal pedicle, extending cranially to form a plane roughly parallel to the anterior articular facet. This surface is honeycombed, and the bone overgrowth displays an irregular, “cauliflower-shaped” morphology. The lateral cortical margin of bone overgrowth No. 1 is relatively smooth, whereas other areas lack a continuous, smooth cortex. It measures 28.6 mm in maximum width and 41.5 mm in length, with the pedicle constituting the majority of its length. Bone overgrowth No. 2 is “beak-shaped,” measuring 38.1 mm in maximum width, 57.3 mm in height, and 44.1 mm in length. The ventral expansion of bone overgrowth No. 2 appears to overlie bone overgrowth No. 1. Additionally, smaller bone overgrowths with surface textures similar to those of bone overgrowths No. 1 and No. 2 are present on the right lateral aspect of the centrum.</p>
        <p>The prezygapophyseal articular facets exhibit pathological abnormalities. The left facet displays expansile changes with a rough, honeycombed surface. The postzygapophyseal facets show similar anomalies with varying degrees of expansion; the changes are more pronounced on the right than on the left. The left caudoventral margin is honeycombed, while the entire surface of the right facet exhibits this texture. The neural spine adjacent to the postzygapophyses displays comparable expansile changes over a vertical range of 24.6 mm, featuring discontinuous cortical bone and a honeycombed surface.</p>
      </sec>
      <sec sec-type="Imaging manifestations" id="sec5">
        <title>Imaging manifestations</title>
        <p><abbrev xlink:title="computed tomography">CT</abbrev> imaging reveals “moth-eaten” osteolytic destruction within both the bone overgrowths and the vertebral body (Fig. <xref ref-type="fig" rid="F3">3</xref>). All bone overgrowths demonstrate continuity with the cortical and cancellous bone of the centrum. The two primary bone overgrowths are particularly distinct in the <abbrev xlink:title="computed tomography">CT</abbrev> imagery. High-density regions are evident at the lateral margins of the bone overgrowths and the centrum; density decreases heterogeneously toward the interior, forming focal osteolytic lesions. The periosteum of the non-pathological portions of the centrum is difficult to distinguish accurately. Localized regions of homogeneous density are present within the centrum. Distinct cavities and fissures are observable within the bone overgrowths and the internal structure of the normal bone, surrounded by marked, homogeneous high-density zones (sclerosis).</p>
        <fig id="F3">
          <object-id content-type="doi">10.3897/fr.29.193112.figure3</object-id>
          <object-id content-type="arpha">7628B7FC-0D91-5F81-A061-F2DDD4738A94</object-id>
          <label>Figure 3.</label>
          <caption>
            <p><abbrev xlink:title="computed tomography">CT</abbrev> imaging of the seventh caudal vertebra of HS0385: <bold>A</bold>. Axial section; <bold>B</bold>. Coronal section; <bold>C</bold>. Sagittal section; and <bold>D</bold>. 3D right lateral view. Red arrows indicate the sinus tract; white arrows indicate sclerosis.</p>
          </caption>
          <graphic xlink:href="fr-29-599-g003.jpg" id="oo_1739214.jpg">
            <uri content-type="original_file">https://binary.pensoft.net/fig/1739214</uri>
          </graphic>
        </fig>
        <p>A sinus tract is identified at the margin of the right-sided region of the centrum; in coronal sections, it constitutes a clearly visible channel draining to the exterior of the bone (Fig. <xref ref-type="fig" rid="F3">3</xref>). The prezygapophyses, postzygapophyses, and adjacent neural spine form heterogeneous regions of mixed high and low density. Occult fracture lines are identifiable in the axial and bilateral sagittal sections (Fig. <xref ref-type="fig" rid="F4">4</xref>). The fracture line is penetrating in the cranial and right sections but does not extend deeply into the centrum; in the left sagittal section, only a minimal portion of the fracture line is visible.</p>
        <fig id="F4">
          <object-id content-type="doi">10.3897/fr.29.193112.figure4</object-id>
          <object-id content-type="arpha">5DA93F6D-185F-5C8C-9FD0-ABE3EFD175B6</object-id>
          <label>Figure 4.</label>
          <caption>
            <p>Location of the occult fracture line in the seventh caudal vertebra of HS0385 in axial (<bold>A</bold>) and right sagittal (<bold>B</bold>) <abbrev xlink:title="computed tomography">CT</abbrev> sections and in 3D right lateral (<bold>C</bold>), cranial (<bold>D</bold>), and left lateral (<bold>E</bold>) views.</p>
          </caption>
          <graphic xlink:href="fr-29-599-g004.jpg" id="oo_1739215.jpg">
            <uri content-type="original_file">https://binary.pensoft.net/fig/1739215</uri>
          </graphic>
        </fig>
      </sec>
    </sec>
    <sec sec-type="Discussion" id="sec6">
      <title>Discussion</title>
      <p>Synthesizing the morphological alterations and <abbrev xlink:title="computed tomography">CT</abbrev> manifestations of the seventh caudal vertebra, the pathological characteristics may be summarized as follows: (1) Extensive expansile hyperostosis and honeycomb-like surface changes: The vertebra exhibits gross expansile abnormalities. In the affected regions, the cortical bone is discontinuous, and a rough, honeycomb-like texture is widely distributed across the surface of the element, indicating an active periosteal reaction or bone remodeling secondary to chronic inflammation. (2) Prominent heteromorphic bone overgrowths: Significant bone overgrowths have developed on the ventral aspect of the centrum, displaying “cauliflower-shaped” or “beak-like” morphologies with substantial volume (both maximum length and height exceed 20 mm). <abbrev xlink:title="computed tomography">CT</abbrev> imaging confirms that these bone overgrowths are continuous with the cortical and cancellous bone of the centrum and contain internal cavities and fissures surrounded by sclerotic high-density zones. (3) “Moth-eaten” osteolytic destruction and mixed-density manifestations: The lesion area displays highly heterogeneous density, characterized by a “high-density margin, low-density center” pattern accompanied by focal osteolytic lesions. (4) Traces of old trauma and occult fractures: The right transverse process is absent and replaced by a hyperostotic stump or elliptical osseous trace, consistent with healed, historic trauma following fracture of the process. Furthermore, <abbrev xlink:title="computed tomography">CT</abbrev> scans reveal occult fracture lines in the transverse and right sagittal sections; these lines partially penetrate but do not extend deeply into the vertebral interior, further confirming that the vertebra suffered external trauma that resulted in incomplete healing or malunion. (5) Sinus tract identification: A definitive sinus tract is identified at the right margin of the centrum. In the coronal section, this manifests as a channel penetrating the cortical bone and connecting the internal pathological focus with the external environment. (6) Given that the remaining caudal vertebrae are comparable in size to those of adult individuals, the relatively small centrum of the seventh caudal vertebra may indicate that the lesion developed during ontogeny and subsequently influenced the growth of this element.</p>
      <sec sec-type="Diagnosis and behavioral implications" id="sec7">
        <title>Diagnosis and behavioral implications</title>
        <p>Expansile osseous lesions are typically differentially diagnosed as follows: (1) infection-induced expansile lesions, such as osteomyelitis (<xref ref-type="bibr" rid="B29">Meyer et al. 2011</xref>), tuberculosis (<xref ref-type="bibr" rid="B15">Greenspan and Remagen 1998</xref>), and parasitic infection (<xref ref-type="bibr" rid="B35">Resnick 2002</xref>; <xref ref-type="bibr" rid="B32">Nielsen and Rosenberg 2017</xref>); (2) metabolic diseases, such as Paget’s disease of bone (<xref ref-type="bibr" rid="B53">Witzmann et al. 2011</xref>); (3) benign lesions, including osteomas (<xref ref-type="bibr" rid="B11">Fechner and Mills 1993</xref>), ossifying fibromas (<xref ref-type="bibr" rid="B8">Dunfee et al. 2005</xref>), bone cysts (<xref ref-type="bibr" rid="B52">Wang et al. 2003</xref>), enchondromas (<xref ref-type="bibr" rid="B29">Meyer et al. 2011</xref>), chondroblastoma (<xref ref-type="bibr" rid="B25">Kong and Cai 2021</xref>), and giant cell tumors (<xref ref-type="bibr" rid="B29">Meyer et al. 2011</xref>); (4) malignant lesions, such as osteosarcoma (<xref ref-type="bibr" rid="B25">Kong and Cai 2021</xref>), chondrosarcoma (<xref ref-type="bibr" rid="B25">Kong and Cai 2021</xref>), fibrosarcoma (<xref ref-type="bibr" rid="B25">Kong and Cai 2021</xref>), chordoma (<xref ref-type="bibr" rid="B25">Kong and Cai 2021</xref>), and myeloma (<xref ref-type="bibr" rid="B25">Kong and Cai 2021</xref>); and (5) arthropathies, such as spondyloarthropathy (<xref ref-type="bibr" rid="B7">de Souza Barbosa et al. 2018</xref>).</p>
        <p>Given the irregular morphology of the bone overgrowths and the rough, discontinuous cortex observed in this specimen, bone cysts, fibrous dysplasia, enchondromas, chondroblastoma, giant cell tumors, and the majority of benign tumors can be excluded (<xref ref-type="bibr" rid="B25">Kong and Cai 2021</xref>). In cases of malignant neoplasms and infectious lesions, persistent pathological stimulation of the periosteum induces abnormal proliferation, resulting in the formation of irregular bone overgrowths (<xref ref-type="bibr" rid="B25">Kong and Cai 2021</xref>). The rapid formation of these bone overgrowths often outpaces cortical formation or produces a cortex lacking normal structural integrity, leading to a discontinuous surface. <abbrev xlink:title="computed tomography">CT</abbrev> imagery reveals clear bone overgrowth boundaries with high-density marginal layering, indicative of hyperostosis and a periosteal reaction typical of chronic inflammation; this permits the exclusion of metastatic tumors, osteosarcoma, and myeloma (<xref ref-type="bibr" rid="B25">Kong and Cai 2021</xref>). Under the prolonged influence of chronic inflammation or stress fractures, the periosteum is continually stimulated to generate new bone, which subsequently calcifies and hardens, forming layered high-density structures. This stratification typically results from repeated periosteal or osseous proliferation. In this specimen, such features are observed not only in the bone overgrowths and centrum but also in the neural arch. The “moth-eaten” osteolytic erosion observed in <abbrev xlink:title="computed tomography">CT</abbrev> scans is predominantly associated with malignant tumors and chronic infections, allowing for the exclusion of parasites and Paget’s disease of bone (<xref ref-type="bibr" rid="B25">Kong and Cai 2021</xref>). Arthropathies such as spondyloarthropathy (<xref ref-type="bibr" rid="B7">de Souza Barbosa et al. 2018</xref>) typically involve the intervertebral articulations and produce enthesiophytes (syndesmophytes) on multiple adjacent vertebrae. By contrast, in HS0385, bony overgrowth is confined to the seventh caudal vertebra, making spondyloarthropathy an unlikely diagnosis. Malignant cells or infectious pathogens destroy the bone matrix via the release of proteases and lytic enzymes, causing local osteolysis. These agents invade trabecular bone, forming small, irregular lytic defects. Chronic inflammatory lesions, such as chronic osteomyelitis, are accompanied by the release of local immune cells; the inflammatory cytokines produced by these cells also stimulate osteolysis, resulting in irregular bone defects. The neural arch of this specimen exhibits identical manifestations (<xref ref-type="bibr" rid="B25">Kong and Cai 2021</xref>). The sinus tract is a specific pathological feature of chronic osteomyelitis. Its formation mechanism involves intramedullary hypertension caused by pyogenic infection within the medullary cavity, forcing fibriscess material and necrotic tissue outward as they erode the cortical bone and form a drainage channel to relieve pressure (<xref ref-type="bibr" rid="B22">Huchzermeyer and Cooper 2000</xref>). In paleopathological and radiographic diagnosis, the presence of a sinus tract not only confirms the connection between a deep infectious focus and the external environment but also serves as a critical functional criterion for distinguishing deep osteomyelitis from simple superficial periostitis (<xref ref-type="bibr" rid="B35">Resnick 2002</xref>). In conclusion, based on the fractured bilateral transverse processes and the evidence of historic trauma and occult fractures, the diagnosis is chronic osteomyelitis secondary to trauma.</p>
        <p>The severe post-traumatic chronic osteomyelitis present in this specimen provides key clues to the behavior of cerapodans. The seventh caudal vertebra is located at the tail base, a primary target zone for theropod predation and a common point of impact in cerapodan intraspecific combat (e.g., flank-butting or tail-swinging) (<xref ref-type="bibr" rid="B47">Siviero et al. 2020</xref>; <xref ref-type="bibr" rid="B6">Cruzado-Caballero et al. 2021</xref>). The destruction of the bilateral transverse processes and traces of open trauma imply that this individual survived an episode of extreme physical confrontation. Notably, despite suffering a deep infection accompanied by a draining sinus, the presence of a massive involucrum and extensive internal sclerosis confirms that the individual survived for a considerable period following the severe injury. <xref ref-type="bibr" rid="B2">Baiano et al. (2024)</xref> conducted a statistical survey of theropod paleopathologies and analyzed the resulting dataset. Their results indicated a non-random anatomical distribution of lesions, with the axial skeleton showing the highest frequency of pathological involvement across all pathology types combined rather than for infectious cases specifically. This may reflect the recurrent exposure of the axial skeleton to mechanical stress, intraspecific interactions, or predation-related injuries during life. Although HS0385 is not a theropod, the occurrence of chronic osteomyelitis in a caudal vertebra is broadly consistent with the pattern reported by <xref ref-type="bibr" rid="B2">Baiano et al. (2024)</xref>.</p>
        <p>The seventh caudal vertebra of HS0385 is situated in the anterior portion of the tail, a region of functional importance for normal activity (<xref ref-type="bibr" rid="B34">Persons and Currie 2011</xref>). Following the initial trauma, prolonged tail movement may have generated additional stress fractures, thereby increasing the likelihood of wound opening and facilitating progression to severe infection. The seventh caudal vertebra is situated within the origin of the <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Medusaceratops">M.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="caudofemoralis">caudofemoralis</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="subspecies" reg="longus">longus</tp:taxon-name-part></tp:taxon-name></italic>, the principal femoral retractor muscle responsible for hindlimb retraction during locomotion (<xref ref-type="bibr" rid="B34">Persons and Currie 2011</xref>). Consequently, this vertebra would have experienced sustained mechanical loading throughout locomotion, thereby increasing the likelihood and severity of mechanical damage. The inability of the animal to reduce the mechanical load on the injured caudal vertebra may have perpetuated callus disruption and promoted microbial invasion of cancellous bone, a recognized pathway for contiguous-spread osteomyelitis (<xref ref-type="bibr" rid="B35">Resnick 2002</xref>; <xref ref-type="bibr" rid="B26">Lew and Waldvogel 2004</xref>). The large-scale gregarious behavior documented in centrosaurines (<xref ref-type="bibr" rid="B44">Ryan et al. 2001</xref>; <xref ref-type="bibr" rid="B9">Eberth and Getty 2005</xref>) may also have increased the exposure of injured individuals to environmental pathogens through substrate contamination associated with herd living.</p>
        <p>Stress fractures are relatively common across <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="superorder" reg="Dinosauria">Dinosauria</tp:taxon-name-part></tp:taxon-name> (e.g., <xref ref-type="bibr" rid="B37">Rothschild and Tanke 1992</xref>; <xref ref-type="bibr" rid="B38">Rothschild and Molnar 2005</xref>). Among ceratopsians, they are most frequently documented in <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Pachyrhinosaurus">Pachyrhinosaurus</tp:taxon-name-part></tp:taxon-name></italic>, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Styracosaurus">Styracosaurus</tp:taxon-name-part></tp:taxon-name></italic>, and <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Triceratops">Triceratops</tp:taxon-name-part></tp:taxon-name></italic>, predominantly affecting the proximal phalanges of pedal digits III and IV (<xref ref-type="bibr" rid="B36">Rothschild 1988</xref>; <xref ref-type="bibr" rid="B37">Rothschild and Tanke 1992</xref>). In sauropods, both the manus and pes represent common sites of stress-related injury (<xref ref-type="bibr" rid="B38">Rothschild and Molnar 2005</xref>; <xref ref-type="bibr" rid="B42">Rothschild et al. 2023</xref>). Among theropods, stress fractures have been reported in <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Allosaurus">Allosaurus</tp:taxon-name-part></tp:taxon-name></italic>, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Ceratosaurus">Ceratosaurus</tp:taxon-name-part></tp:taxon-name></italic>, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Tarbosaurus">Tarbosaurus</tp:taxon-name-part></tp:taxon-name></italic>, and dromaeosaurids (<xref ref-type="bibr" rid="B41">Rothschild et al. 2001</xref>; <xref ref-type="bibr" rid="B39">Rothschild and Tanke 2005</xref>; <xref ref-type="bibr" rid="B1">Anné et al. 2015</xref>), with additional occurrences noted in other tyrannosaurids (<xref ref-type="bibr" rid="B42">Rothschild et al. 2023</xref>). Notably, stress fractures in the <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Allosaurus">Allosaurus</tp:taxon-name-part></tp:taxon-name></italic> pes occur at approximately equal frequencies across digits II–IV, which does not support a running-induced etiology; if running were the primary cause, a higher incidence would be expected in digit III. <xref ref-type="bibr" rid="B41">Rothschild et al. (2001)</xref> instead proposed that these injuries may have been associated with predatory behavior.</p>
        <p>Theropods may have mitigated loading on an injured hind limb through compensatory use of the contralateral pes. The trackway described by <xref ref-type="bibr" rid="B19">Herrera-Castillo et al. (2022)</xref>, in which the left pes shows digital abnormalities, whereas the right footprints indicate greater loading, is consistent with such compensatory behavior. <xref ref-type="bibr" rid="B6">Cruzado-Caballero et al. (2021)</xref> likewise argued that pathological conditions do not necessarily result directly in severe locomotor impairment. Several <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Allosaurus">Allosaurus</tp:taxon-name-part></tp:taxon-name></italic> specimens further show that pedal infections could be effectively contained as localized involucra rather than progressing to diffuse osteomyelitis (<xref ref-type="bibr" rid="B17">Hanna 2002</xref>; <xref ref-type="bibr" rid="B13">Foth et al. 2015</xref>). Together with the comparatively high metabolic rates inferred for theropods (<xref ref-type="bibr" rid="B14">Grady et al. 2014</xref>), this has been interpreted as potentially reflecting a more vigorous innate immune response.</p>
      </sec>
      <sec sec-type="Infection statistics" id="sec8">
        <title>Infection statistics</title>
        <p>Infectious cases are relatively common in dinosaurian paleopathology; as of June 2026, 81 such cases have been compiled (Suppl. material <xref ref-type="supplementary-material" rid="S1">1</xref>). Osteomyelitis represents the predominant category in the infection dataset, accounting for 84.0% of cases (68/81), whereas the remaining cases include septic arthritis, parasitism, respiratory infection, and other less frequent conditions. This predominance is consistent with osteomyelitis being the infectious lesion most readily recognized in fossil bone or, alternatively, the type most likely to be preserved in the fossil record. The great majority of cases are post-traumatic infections that preserve evidence of antecedent trauma, accounting for 59.0% of the dataset (48/81), whereas non-traumatic cases comprise only 21.0% (17/81). Among the post-traumatic cases, the recorded specific injury mechanisms include fractures other than stress fractures (28 cases), open or puncture wounds (14 cases), cranial impact injuries (four cases), and limb amputation (one case). The marked predominance of post-traumatic cases indicates that, within the diagnosable dinosaurian fossil record, bone infection was seeded principally through traumatic breaches of the integument or skeleton, consistent with contiguous-spread osteomyelitis (<xref ref-type="bibr" rid="B35">Resnick 2002</xref>; <xref ref-type="bibr" rid="B26">Lew and Waldvogel 2004</xref>), rather than arising chiefly from hematogenous or other non-traumatic routes. These data describe the composition of reported cases and may therefore be subject to overrepresentation. Because the sample includes only infected individuals, post-traumatic osteomyelitis with associated fractures or wounds is more readily recognized than primary hematogenous osteomyelitis, and approximately one-quarter of cases have an indeterminate trauma mechanism, these proportions are likely to be influenced by diagnostic and taphonomic biases.</p>
        <p>The anatomical distribution of infectious lesions also differs among the major clades (Fig. <xref ref-type="fig" rid="F5">5</xref>). Axial-skeletal involvement predominates in thyreophorans and sauropodomorphs; theropod lesions are more evenly distributed across the cranial, axial, and appendicular regions and girdle; and cerapodan lesions occur mainly in the cranial and axial skeleton. The association between clade and anatomical region is statistically detectable (<italic>χ</italic><sup>2</sup> = 25, df = 9, <italic>P</italic> = 0.004; Cramér’s <italic>V</italic> = 0.32). Caudal lesions are significantly overrepresented in <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="order" reg="Sauropodomorpha">Sauropodomorpha</tp:taxon-name-part></tp:taxon-name> relative to other clades (Fisher’s exact test, <italic>P</italic> = 0.004; Cramér’s <italic>V</italic> = 0.35), a pattern that may be related to the functional and behavioral importance of the sauropod tail. However, these associations should be regarded as descriptive and hypothesis-generating rather than as confirmatory evidence of causal relationships.</p>
        <fig id="F5">
          <object-id content-type="doi">10.3897/fr.29.193112.figure5</object-id>
          <object-id content-type="arpha">D430E948-24E2-5C67-93F7-9F26E6365636</object-id>
          <label>Figure 5.</label>
          <caption>
            <p>Heatmap showing the anatomical distribution of infectious lesions by clade across 81 reported cases of dinosaurian infection. Cell values indicate lesion counts, and shading represents within-clade proportions. Cranial includes the skull and mandible; axial includes vertebrae, ribs, and osteoderms. One non-skeletal lesion was excluded.</p>
          </caption>
          <graphic xlink:href="fr-29-599-g005.jpg" id="oo_1739216.jpg">
            <uri content-type="original_file">https://binary.pensoft.net/fig/1739216</uri>
          </graphic>
        </fig>
      </sec>
    </sec>
    <sec sec-type="Conclusion" id="sec9">
      <title>Conclusion</title>
      <p>This study presents the first documented case of chronic osteomyelitis with a draining sinus tract in <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Medusaceratops">Medusaceratops</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="lokii">lokii</tp:taxon-name-part></tp:taxon-name></italic>, diagnosed from a pathological seventh caudal vertebra (HS0385) recovered from the Upper Cretaceous Judith River Formation of Montana. Integrated macroscopic and <abbrev xlink:title="computed tomography">CT</abbrev> analyses reveal pathognomonic features of chronic osteomyelitis secondary to trauma, including moth-eaten osteolysis, extensive involucrum formation, and occult fractures associated with the destruction of the bilateral transverse processes. The presence of a definitive sinus tract—a hallmark of deep osteomyelitis—together with the massive involucrum and internal sclerosis, confirms that the individual survived for a prolonged period following the initial traumatic insult, attesting to robust immunological resilience in ceratopsid dinosaurs.</p>
      <p>A review of the 81 reported cases of dinosaurian infection further indicates that osteomyelitis is the predominant lesion type and that the great majority of cases are post-traumatic. The anatomical distribution of infectious lesions also differs among clades: in theropods, lesions are concentrated mainly in the cranium and the axial and appendicular skeletons; in thyreophorans and sauropodomorphs, they are concentrated in the axial skeleton; and in cerapodans, lesions occur predominantly in the cranial and axial skeleton.</p>
    </sec>
  </body>
  <back>
    <ack>
      <title>Acknowledgments</title>
      <p>The authors thank two anonymous reviewers for their constructive comments and helpful suggestions.</p>
    </ack>
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    <sec sec-type="Additional information" id="sec10">
      <title>Additional information</title>
      <sec sec-type="Conflict of interest" id="sec11">
        <title>Conflict of interest</title>
        <p>The authors have declared that no competing interests exist.</p>
      </sec>
      <sec sec-type="Ethical statement" id="sec12">
        <title>Ethical statement</title>
        <p>No ethical statement was reported.</p>
      </sec>
      <sec sec-type="Artificial Intelligence (AI) use" id="sec13">
        <title>Artificial Intelligence (AI) use</title>
        <p>The authors accept full responsibility for the content of the manuscript, including the disclosure of any use of AI.</p>
        <p>Regarding the use of AI in the preparation of this manuscript, the authors declare the following: Claude.</p>
        <p><bold>Used for</bold>: Other.</p>
        <p>The authors used AI for references checking and manuscript proofreading.</p>
      </sec>
      <sec sec-type="Funding" id="sec14">
        <title>Funding</title>
        <p>This research was funded by the 2025 Heilongjiang Provincial Natural Science Foundation Youth Project Type C (JJ2025QC0704), the National Natural Science Foundation of China (No. 42488201), the “Deep-time Digital Earth” Science and Technology Leading Talents Team Funds for the Central Universities for the Frontiers Science Center for Deep-time Digital Earth, China University of Geosciences (Beijing) (Fundamental Research Funds for the Central Universities; grant number: 2652023001), the 111 Project (B20011), and the Fundamental Research Funds for the Central Universities (265QZ201903).</p>
      </sec>
      <sec sec-type="Author contributions" id="sec15">
        <title>Author contributions</title>
        <p>Donghao Wang conducted the specimen description and diagnosis, collected and analysed the data, and drafted the manuscript. Lida Xing contributed to the specimen description, diagnosis, and manuscript preparation. Bruce M. Rothschild contributed to the diagnosis, data collection, and manuscript preparation. Chunlei Du, Kexiang Wen, Jin Lin, and Mengling Kang carried out specimen photography, <abbrev xlink:title="computed tomography">CT</abbrev> scanning, and structured-light scanning. Jiayin Su contributed to the diagnosis. All authors reviewed and approved the final version of the manuscript.</p>
      </sec>
      <sec sec-type="Author ORCIDs" id="sec16">
        <title>Author ORCIDs</title>
        <p>L. Xing <ext-link xlink:href="https://orcid.org/0000-0003-3923-9206" ext-link-type="uri">https://orcid.org/0000-0003-3923-9206</ext-link></p>
        <p>B.M. Rothschild <ext-link xlink:href="https://orcid.org/0000-0003-1327-6615" ext-link-type="uri">https://orcid.org/0000-0003-1327-6615</ext-link></p>
      </sec>
      <sec sec-type="Data availability" id="sec17">
        <title>Data availability</title>
        <p>The Supplementary material is available at <ext-link xlink:href="10.6084/m9.figshare.31850866" ext-link-type="doi">https://doi.org/10.6084/m9.figshare.31850866</ext-link>.</p>
      </sec>
    </sec>
    <sec sec-type="supplementary-material">
      <title>Supplementary materials</title>
      <supplementary-material id="S1" position="float" orientation="portrait" xlink:type="simple">
        <object-id content-type="arpha">6DC49EED-9CE2-5023-85EF-C5219176196D</object-id>
        <label>Supplementary material 1</label>
        <caption>
          <p>Supplementary information</p>
        </caption>
        <statement content-type="dataType">
          <label>Data type</label>
          <p>xlsx</p>
        </statement>
        <media xlink:href="fr-29-599-s001.xlsx" mimetype="application" mime-subtype="vnd.openxmlformats-officedocument.spreadsheetml.sheet" position="float" orientation="portrait" id="oo_1739217.xlsx">
          <uri content-type="original_file">https://binary.pensoft.net/file/1739217</uri>
        </media>
        <permissions>
          <license>
            <license-p>This dataset is made available under the Open Database License (<ext-link ext-link-type="uri" xlink:href="http://opendatacommons.org/licenses/odbl/1.0">http://opendatacommons.org/licenses/odbl/1.0</ext-link>). The Open Database License (ODbL) is a license agreement intended to allow users to freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited.</license-p>
          </license>
        </permissions>
        <attrib specific-use="authors"> Donghao Wang, Lida Xing, Bruce M. Rothschild, Chunlei Du, Kexiang Wen, Jin Lin, Mengling Kang, Jiayin Su</attrib>
      </supplementary-material>
    </sec>
  </back>
</article>
