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  <front>
    <journal-meta><journal-id journal-id-type="publisher">FR</journal-id><journal-title-group>
    <journal-title>Fossil Record</journal-title>
    <abbrev-journal-title abbrev-type="publisher">FR</abbrev-journal-title><abbrev-journal-title abbrev-type="nlm-ta">Foss. Rec.</abbrev-journal-title>
  </journal-title-group><issn pub-type="epub">2193-0074</issn><publisher>
    <publisher-name>Copernicus Publications</publisher-name>
    <publisher-loc>Göttingen, Germany</publisher-loc>
  </publisher></journal-meta>
    <article-meta>
      <article-id pub-id-type="doi">10.5194/fr-22-57-2019</article-id><title-group><article-title>Presence of the Jehol Biota turtle <italic>Ordosemys liaoxiensis</italic><?xmltex \hack{\break}?> in the Early Cretaceous Hengtongshan
Formation of<?xmltex \hack{\break}?> southern Jilin Province, China</article-title><alt-title>Presence of the Jehol Biota turtle <italic>Ordosemys liaoxiensis</italic></alt-title>
      </title-group><?xmltex \runningtitle{Presence of the Jehol Biota turtle \textit{Ordosemys liaoxiensis}}?><?xmltex \runningauthor{C.-F.~Zhou et al.}?>
      <contrib-group>
        <contrib contrib-type="author" corresp="yes" rid="aff1">
          <name><surname>Zhou</surname><given-names>Chang-Fu</given-names></name>
          <email>zhoucf528@163.com</email>
        <ext-link>https://orcid.org/0000-0002-3744-7573</ext-link></contrib>
        <contrib contrib-type="author" corresp="yes" rid="aff2 aff3">
          <name><surname>Wu</surname><given-names>Wen-Hao</given-names></name>
          <email>wenhao.wu@foxmail.com</email>
        <ext-link>https://orcid.org/0000-0003-0970-9713</ext-link></contrib>
        <contrib contrib-type="author" corresp="no" rid="aff4 aff5">
          <name><surname>Rabi</surname><given-names>Márton</given-names></name>
          
        </contrib>
        <aff id="aff1"><label>1</label><institution>College of Earth Science and Engineering, Shandong University of Science
and Technology,<?xmltex \hack{\break}?> Qingdao 266590, Shandong, China</institution>
        </aff>
        <aff id="aff2"><label>2</label><institution>Key Laboratory for Evolution of Past Life and Environment in Northeast
Asia, Ministry of Education,<?xmltex \hack{\break}?> Changchun 130026, Jilin, China</institution>
        </aff>
        <aff id="aff3"><label>3</label><institution>Research Center of Palaeontology and Stratigraphy, Jilin University,
Changchun 130056, Jilin, China</institution>
        </aff>
        <aff id="aff4"><label>4</label><institution>Central Natural Science Collections, Martin Luther University
Halle-Wittenberg, Halle (Saale), Germany</institution>
        </aff>
        <aff id="aff5"><label>5</label><institution>Department of Geosciences, University of Tübingen, Tübingen,
Germany</institution>
        </aff>
      </contrib-group>
      <author-notes><corresp id="corr1">Chang-Fu Zhou (zhoucf528@163.com) and Wen-Hao Wu
(wenhao.wu@foxmail.com)</corresp></author-notes><pub-date><day>12</day><month>September</month><year>2019</year></pub-date>
      
      <volume>22</volume>
      <issue>2</issue>
      <fpage>57</fpage><lpage>64</lpage>
      <history>
        <date date-type="received"><day>25</day><month>May</month><year>2019</year></date>
           <date date-type="rev-recd"><day>15</day><month>August</month><year>2019</year></date>
           <date date-type="accepted"><day>19</day><month>August</month><year>2019</year></date>
      </history>
      <permissions>
        <copyright-statement>Copyright: © 2019 </copyright-statement>
        <copyright-year>2019</copyright-year>
      <license license-type="open-access"><license-p>This work is licensed under the Creative Commons Attribution 4.0 International License. To view a copy of this licence, visit <ext-link ext-link-type="uri" xlink:href="https://creativecommons.org/licenses/by/4.0/">https://creativecommons.org/licenses/by/4.0/</ext-link></license-p></license></permissions><self-uri xlink:href="https://fr.copernicus.org/articles/.html">This article is available from https://fr.copernicus.org/articles/.html</self-uri><self-uri xlink:href="https://fr.copernicus.org/articles/.pdf">The full text article is available as a PDF file from https://fr.copernicus.org/articles/.pdf</self-uri>
      <abstract><title>Abstract</title>
    <p id="d1e137">Recently, a vertebrate assemblage of the Jehol Biota has been reported from
the Early Cretaceous Hengtongshan Formation of Xingling Town, Meihekou City,
Jilin Province, China. It is dominated by the fishes <italic>Lycoptera</italic> and <italic>Sinamia</italic> and the sinemydid
turtle <italic>Ordosemys</italic>. Here, we describe the turtle specimens and referral to <italic>Ordosemys liaoxiensis</italic>, otherwise
known from the older Yixian Formation of the Jehol Biota.  It is characterized
by a subcircular shell, wide vertebral scales, well-developed plastral
fenestrae, and a major contribution from the xiphiplastra to enclose the
hypo-xiphiplastral fenestra. As the first Mesozoic turtle of Jilin Province,
this record represents the first tetrapod to indicate the presence of the
Jehol Biota in the region. Given the geographic and temporal distance from
the Yixian Formation, future collections from the Hengtongshan Formation
have good potential for evaluating spatiotemporal patterns of the Jehol
Biota.</p>
  </abstract>
    </article-meta>
  </front>
<body>
      

<sec id="Ch1.S1" sec-type="intro">
  <label>1</label><title>Introduction</title>
      <p id="d1e161">The Jehol Biota of northeastern China (western Liaoning and conjoint areas)
is renowned for providing exceptional insights into an Early Cretaceous
ecosystem through its exquisite preservation of plant, invertebrate, and
vertebrate fossils (e.g., Zhou et al., 2003; Pan et al., 2013). In the field,
the Jehol Biota is typically recognized by the presence of the <italic>Eosestheria</italic> (crustacean)–<italic>Ephemeropsis trisetalis</italic>
(mayfly insect)–<italic>Lycoptera</italic> (fish) community (T. Shao et al., 2017). Freshwater sinemydid
turtles are among the most common tetrapod fossils of the Jehol Biota, which
makes their biostratigraphic potential of interest (e.g., Ji, 1995; Tong et
al., 2004; Zhou, 2010a, b; Zhou and Rabi, 2015; S. Shao et al., 2017, 2018; Li
et al., 2019). However, these fossil turtles have been less commonly
collected outside the typical western Liaoning area of the Jehol Biota
(e.g., Ma, 1986; Li et al., 2015; Li and Tong, 2017; Wu et al., 2018).
Recently, as the first record of the Mesozoic turtles from Jilin Province,
the sinemydid <italic>Ordosemys</italic> has been reported from the Early Cretaceous Hengtongshan
Formation of Xingling Town, Meihekou City (Wu et al., 2018). This material
has not been described yet, and its species-level taxonomy awaits
clarification. In this study, we describe this material, together with other
two specimens, and clarify their taxonomic status. We show that these
turtles represent the first tetrapod evidence for the presence of the Jehol
Biota in the Hengtongshan Formation. Previous studies correlated the
Hengtongshan Formation with the Jiufotang Formation of the Jehol Biota of
western Liaoning and conjoint areas based on radiometric dating (119–123 Myr; Y. Chen et al., 2008; Sui and Chen, 2012; J. Chen et al., 2016) and
biostratigraphic data (e.g., <italic>Lycoptera</italic>, <italic>Ephemeropsis</italic>, <italic>Eosestheria</italic>, <italic>Huashia</italic>, <italic>Diestheria</italic>, and <italic>Orthestheria</italic>) (e.g., JBGMR, 1982; Chang et
al., 2009; Wu et al., 2018).</p>
      <?pagebreak page58?><p id="d1e195">The sinemydid turtle <italic>Ordosemys</italic> has a broad paleogeographic distribution in northern
China (e.g., Danilov and Parham, 2007; Li et al., 2019). Three species are
widely accepted, including the type species <italic>Ordosemys leios</italic> from the Ordos Basin of Inner
Mongolia and Mengyin Basin of Shandong (Brinkman and Peng, 1993; Brinkman
and Wu, 1999; Li et al., 2019), <italic>O. liaoxiensis</italic> from the Yixian Formation of western
Liaoning (e.g., Ji, 1995; Tong et al., 2004), and <italic>O. brinkmania</italic> from Junggar Basin of
Xinjiang (Danilov and Parham, 2007). <italic>Ordosemys</italic> <italic>donghai</italic> (i.e., <italic>Manchurochelys donghai</italic>, Ma, 1986) as recombined by
Brinkman et al. (2008) from Heilongjiang, is possibly excluded from
<italic>Ordosemys</italic> in lacking the typical generic features (Li et al., 2019).</p>
</sec>
<sec id="Ch1.S2">
  <label>2</label><title>Geographic and geologic settings</title>
      <p id="d1e231">The fossil site is located at a small quarry near Shuanghe Village, Xingling
Town, approximately 15 km southeast of the Meihekou City, southern Jilin
Province, China (42<inline-formula><mml:math id="M1" display="inline"><mml:msup><mml:mi/><mml:mo>∘</mml:mo></mml:msup></mml:math></inline-formula>24<inline-formula><mml:math id="M2" display="inline"><mml:msup><mml:mi/><mml:mo>′</mml:mo></mml:msup></mml:math></inline-formula>54.52<inline-formula><mml:math id="M3" display="inline"><mml:msup><mml:mi/><mml:mrow><mml:mo>′</mml:mo><mml:mo>′</mml:mo></mml:mrow></mml:msup></mml:math></inline-formula> N, 125<inline-formula><mml:math id="M4" display="inline"><mml:msup><mml:mi/><mml:mo>∘</mml:mo></mml:msup></mml:math></inline-formula>51<inline-formula><mml:math id="M5" display="inline"><mml:msup><mml:mi/><mml:mo>′</mml:mo></mml:msup></mml:math></inline-formula>02.21<inline-formula><mml:math id="M6" display="inline"><mml:msup><mml:mi/><mml:mrow><mml:mo>′</mml:mo><mml:mo>′</mml:mo></mml:mrow></mml:msup></mml:math></inline-formula> E; Fig. 1). Five fossil-bearing layers can be identified in the outcrop of the
fossil site (Fig. 1b). The turtle fossils described in this study are from
the lower layer, associated with the fishes <italic>Lycoptera davidi</italic> and <italic>Sinamia</italic> sp. However, as described
by the local farmers, an upper layer is a possible second one bearing fossil
turtles. Unfortunately, the fossil turtles were lost by the local farmers.</p>

      <?xmltex \floatpos{t}?><fig id="Ch1.F1" specific-use="star"><?xmltex \currentcnt{1}?><label>Figure 1</label><caption><p id="d1e303"><bold>(a)</bold> Fossil locality (asterisk; 42<inline-formula><mml:math id="M7" display="inline"><mml:msup><mml:mi/><mml:mo>∘</mml:mo></mml:msup></mml:math></inline-formula>24<inline-formula><mml:math id="M8" display="inline"><mml:msup><mml:mi/><mml:mo>′</mml:mo></mml:msup></mml:math></inline-formula>54.52<inline-formula><mml:math id="M9" display="inline"><mml:msup><mml:mi/><mml:mrow><mml:mo>′</mml:mo><mml:mo>′</mml:mo></mml:mrow></mml:msup></mml:math></inline-formula> N,
125<inline-formula><mml:math id="M10" display="inline"><mml:msup><mml:mi/><mml:mo>∘</mml:mo></mml:msup></mml:math></inline-formula>51<inline-formula><mml:math id="M11" display="inline"><mml:msup><mml:mi/><mml:mo>′</mml:mo></mml:msup></mml:math></inline-formula>02.21<inline-formula><mml:math id="M12" display="inline"><mml:msup><mml:mi/><mml:mrow><mml:mo>′</mml:mo><mml:mo>′</mml:mo></mml:mrow></mml:msup></mml:math></inline-formula> E) of <italic>Ordosemys</italic> <italic>liaoxiensis</italic> from the Early Cretaceous Hengtongshan
Formation of Shuanghe Village, Xingling Town, Meihekou City, Jilin Province,
northeastern China. <bold>(b)</bold> Outcrop of the fossil site showing five
fossil-bearing layers; the turtle specimens described in this study are from
the lower layer.</p></caption>
        <?xmltex \igopts{width=483.69685pt}?><graphic xlink:href="https://fr.copernicus.org/articles/22/57/2019/fr-22-57-2019-f01.png"/>

      </fig>

      <p id="d1e384">The fossil-bearing beds are mainly composed of lacustrine deposits of thick
layers of greyish-green sandstones and siltstones, which belong to the
middle part of the Hengtongshan Formation (JBGMR, 1982; Wu et al., 2018).
The Hengtongshan Formation is well exposed in southern Jilin Province and
spans 119–123 Myr based on radiometric dating (e.g., Chen et al., 2008, 2016;
Sui and Chen, 2012). Based on this, the Hengtongshan
Formation is roughly coeval to the Jiufotang Formation as also suggested by
biostratigraphic data (e.g., <italic>Lycoptera, Ephemeropsis, Eosestheria, Huashia, Diestheria</italic>, and <italic>Orthestheria</italic>) (e.g., JBGMR, 1982; Chang et al., 2009;
Wu et al., 2018).</p>
</sec>
<sec id="Ch1.S3">
  <label>3</label><title>Samples and methods</title>
      <p id="d1e401">Three turtle specimens and associated fishes <italic>Lycoptera</italic> and <italic>Sinamia</italic> were collected by the
local farmers from a quarry near Shuanghe Village, Xingling Town, Meihekou
City, southern Jilin Province, China (Fig. 1). The fossils are deposited in
the Paleontological Collection of Xingling Town. Before this study, a
skillful preparation of three specimens was performed in the Research
Center of Palaeontology and Stratigraphy, Jilin University. The fossils are
deposited in greyish-green sandstones and are strongly compressed
dorsoventrally. The largest specimen (XL-009; Fig. 2) preserves much of the
plastron with several fragments and traces of the carapace; the full
ossification of the carapace indicates an adult stage for this specimen. The
other two specimens (XL-007 and XL-008; Figs. 3–4) are slightly smaller with
XL-008 exhibiting large costo-peripheral fenestrae, whereas XL-007 reduced to
closed costo-peripheral fenestrae. The three specimens are identified as the
same taxon representing different ontogenetic stages.</p>

      <?xmltex \floatpos{t}?><fig id="Ch1.F2" specific-use="star"><?xmltex \currentcnt{2}?><label>Figure 2</label><caption><p id="d1e412"><italic>Ordosemys</italic> <italic>liaoxiensis</italic> (XL-009; most in visceral view) from the Early Cretaceous
Hengtongshan Formation of Shuanghe Village, Xingling Town, Meihekou City,
Jilin Province, northeastern China. <bold>(a, b)</bold>   Carapace before
preparation; <bold>(c, d)</bold> plastron and appendicular elements after
preparation. Abbreviations: 1–2, neural plates 1–2; c1–c6, costal plates
1–6; cf, central plastral fenestra; co, coracoid; d2–d6, dorsal vertebrae
2–6; fe, femur; fi, fibula; hyo, hyoplastron; hyp, hypoplastron; is,
ischium; lf, lateral plastral fenestra; pe, peripheral plate; pf, posterior
medial plastral fenestra; pu, pubis; ti, tibia; v1–v2, vertebral scales 1–2;
xi, xiphiplastron.</p></caption>
        <?xmltex \igopts{width=412.564961pt}?><graphic xlink:href="https://fr.copernicus.org/articles/22/57/2019/fr-22-57-2019-f02.jpg"/>

      </fig>

      <?xmltex \floatpos{p}?><fig id="Ch1.F3" specific-use="star"><?xmltex \currentcnt{3}?><label>Figure 3</label><caption><p id="d1e434"><italic>Ordosemys</italic> <italic>liaoxiensis</italic> (XL-007; in dorsal view) from the Early Cretaceous Hengtongshan
Formation of Shuanghe Village, Xingling Town, Meihekou City, Jilin Province,
northeastern China. Abbreviations: 3–4, neural plates 3–4; c2–c8, costal
plates 2–8; ib, inguinal buttress of the hypoplastron; p4–p9, peripheral
plates 4–9; v3, vertebral scale 3.</p></caption>
        <?xmltex \igopts{width=455.244094pt}?><graphic xlink:href="https://fr.copernicus.org/articles/22/57/2019/fr-22-57-2019-f03.jpg"/>

      </fig>

      <?xmltex \floatpos{p}?><fig id="Ch1.F4" specific-use="star"><?xmltex \currentcnt{4}?><label>Figure 4</label><caption><p id="d1e451"><italic>Ordosemys</italic> <italic>liaoxiensis</italic> (XL-008; in dorsal view) from the Early Cretaceous Hengtongshan
Formation of Shuanghe Village, Xingling Town, Meihekou City, Jilin Province,
northeastern China. Abbreviations: 6, neural plate 6; c1–c8, costal plates
1–8; cf, central plastral fenestra; cs, cervical scale; cv, cervical
vertebrae; hu, humerus; hyo, hyoplastron; hyp, hypoplastron; lf, lateral
plastral fenestra; pe, peripheral plates; ul, ulna.</p></caption>
        <?xmltex \igopts{width=455.244094pt}?><graphic xlink:href="https://fr.copernicus.org/articles/22/57/2019/fr-22-57-2019-f04.jpg"/>

      </fig>

<sec id="Ch1.S3.SSx1" specific-use="unnumbered">
  <title>Institution abbreviations</title>
      <p id="d1e470">IVPG, Institute of Vertebrate Paleontology, Gansu
Agricultural University, Lanzhou, China; GSGM, Gansu Geological Museum,
Lanzhou, China; PMOL, Paleontological Museum of Liaoning, Shenyang Normal
University, Shenyang, China; XL, Paleontological Collection of Xingling
Town, Meihekou, China.</p>
</sec>
</sec>
<sec id="Ch1.S4">
  <label>4</label><title>Systematic paleontology</title>
      <p id="d1e482"><list list-type="custom">
          <list-item><label> </label>

      <p id="d1e487"><bold>Testudinata</bold> Klein, 1760</p>
          </list-item>
          <list-item><label> </label>

      <p id="d1e495"><bold>Testudines</bold> Batsch, 1788</p>
          </list-item>
          <list-item><label> </label>

      <p id="d1e503"><bold>Pan-Cryptodira</bold> Joyce, Parham &amp; Gauthier, 2004</p>
          </list-item>
          <list-item><label> </label>

      <p id="d1e511"><bold>Sinemydidae</bold> sensu Rabi, Sukhanov, Egorova, Danilov &amp; Joyce, 2014</p>
          </list-item>
          <list-item><label> </label>

      <p id="d1e519"><italic>Ordosemys liaoxiensis</italic> (Ji 1995)</p>

      <p id="d1e524">(Figs. 2–4)</p>
          </list-item>
        </list></p>
<sec id="Ch1.S4.SSx1" specific-use="unnumbered">
  <title>Specimens</title>
      <p id="d1e534">XL-007 (Fig. 3), a subadult specimen with most of the shell; XL-008 (Fig. 4), a juvenile skeleton including a partial skull, cervicals, shell and
forelimbs; XL-009 (Fig. 2), an adult skeleton including a partial carapace,
much of the plastron, and some appendicular bones. These fossils are
deposited in the Paleontological Collection of Xingling Town (XL).</p>
</sec>
<sec id="Ch1.S4.SSx2" specific-use="unnumbered">
  <title>Locality and horizon</title>
      <p id="d1e543">A quarry (GPS: 42<inline-formula><mml:math id="M13" display="inline"><mml:msup><mml:mi/><mml:mo>∘</mml:mo></mml:msup></mml:math></inline-formula>24<inline-formula><mml:math id="M14" display="inline"><mml:msup><mml:mi/><mml:mo>′</mml:mo></mml:msup></mml:math></inline-formula>54.52<inline-formula><mml:math id="M15" display="inline"><mml:msup><mml:mi/><mml:mrow><mml:mo>′</mml:mo><mml:mo>′</mml:mo></mml:mrow></mml:msup></mml:math></inline-formula> N, 125<inline-formula><mml:math id="M16" display="inline"><mml:msup><mml:mi/><mml:mo>∘</mml:mo></mml:msup></mml:math></inline-formula>51<inline-formula><mml:math id="M17" display="inline"><mml:msup><mml:mi/><mml:mo>′</mml:mo></mml:msup></mml:math></inline-formula>02.21<inline-formula><mml:math id="M18" display="inline"><mml:msup><mml:mi/><mml:mrow><mml:mo>′</mml:mo><mml:mo>′</mml:mo></mml:mrow></mml:msup></mml:math></inline-formula> E) near
Shuanghe Village, Xingling Town, Meihekou City, southern Jilin Province,
China (Fig. 1); the Early Cretaceous Hengtongshan Formation (119–123 Myr).</p>
</sec>
</sec>
<sec id="Ch1.S5">
  <label>5</label><title>Description</title>
<sec id="Ch1.S5.SS1">
  <label>5.1</label><title>Osteology of XL-009</title>
      <p id="d1e624">XL-009 is the largest specimen with an estimated plastral width of 144 cm.
The carapace is mostly missing, but the imprints reveal that the
costo-peripheral fenestrae were fully ossified (Fig. 2a, b). Neurals 1 and 2 are partially preserved together with the sulcus of vertebrals 1–2 across
neural 1 and<?pagebreak page59?> costal 1. The sulcus is slightly protruding anteriorly, which
is also known for <italic>Ordosemys leios</italic> and <italic>O. brinkmania</italic> (Danilov and Parham, 2007; Li et al., 2019). The
preneural plate is typical for <italic>Ordosemys</italic> spp., and its suture with neural 1 is
generally positioned behind the sulcus of vertebrals 1–2 in <italic>Ordosemys</italic> spp. (e.g.,
Brinkman and Peng, 1993; Tong et al., 2004; Danilov and Parham, 2007; Li et
al., 2019), or confluent with the sulcus of the vertebrals 1–2 (a juvenile
of <italic>O. leios</italic>; IVPG-T001-2; Li et al., 2019). However, the suture is uncertain in the
corresponding region of XL-009.</p>
      <p id="d1e642">The plastron is well exposed with the hyoplastron, hypoplastron, and
xiphiplastron in visceral view (Fig. 2c, d). The plastral fenestrae are
open, including the two medial and two lateral fenestrae, as in <italic>Ordosemys leios, O. liaoxiensis</italic>, and
<italic>Liaochelys jianchangensis</italic> (Brinkman and Peng, 1993; Tong et al., 2004; Zhou, 2010a). The central and
lateral plastral fenestrae are enlarged in size, whereas the
hypo-xiphiplastral fenestra is much smaller. The xiphiplastra contributes to
the hypo-xiphiplastral fenestra to a greater extent than the hypoplastra. In
contrast, the fenestra is enclosed by the xiphiplastra and hypoplastra
subequally in <italic>O. leios, O. liaoxiensis</italic>, and <italic>L. jianchangensis</italic> (Brinkman and Peng, 1993; Tong et al., 2004; Zhou,
2010a).</p>
      <p id="d1e657">The right hyoplastron is 72 mm in width. The hyoplastron and hypoplastron
are firmly articulated and form the plastral bridge. The bridge is 45 mm in
length. Laterally, the hyoplastron and hypoplastron are characteristic of
the peg-like processes which articulated with the carapace through
ligaments. The hypoplastra enclosed the posterior medial fenestra with the
xiphiplastra. The xiphiplastra is largely covered by the pelvic girdle. The
xiphiplastron seems to be elongated as in <italic>Ordosemys</italic> spp. but different from the
condition seen in <italic>Liaochelys jianchangensis</italic>.</p>
      <p id="d1e666">The pectoral girdle is only preserved with the two strap-like coracoids. The
right pelvic girdle is exposed with the pubis and ischium. The pubis is
triradiate with the lateral pubic process, the medial pectineal process, and
the dorsal acetabular process. The lateral pubic process is comparable with
the pectineal process in size. The acetabular process is stronger and firmly
articulates with ischium. The right femur is preserved in articulation with
the acetabulum and is about 40 mm long. The tibia and fibula are only
preserved with their proximal ends.</p>
</sec>
<sec id="Ch1.S5.SS2">
  <label>5.2</label><title>Osteology of XL-007</title>
      <p id="d1e677">The carapace appears to be subcircular with an estimated width of 130 mm
across costal 4 (Fig. 3). The carapace is smooth and sculptured by the scale
sulci. Costals 2–8 are preserved and the costo-peripheral fenestrae seem to
be open between costals 2–4 and peripherals 4–7, but the posterior ones are
fully closed. In contrast, the similar region is completely ossified in the
larger specimen XL-009, based on the imprints of the carapace (Fig. 2a, b).
Therefore, the open condition implies that XL-007 represents a subadult
individual.</p>
      <p id="d1e680">Distally, the rib ends are exposed similarly to the condition in other
sinemydids. Costal 3 is not expanded distally, different from the condition
of <italic>Liaochelys jianchangensis</italic> (Zhou, 2010a).</p>
      <p id="d1e686">The sulci of vertebral and pleural scales are deep and identifiable. The
lateral termination of vertebral 3 can be traced. Its width is about 70 mm,
more than half the width of the carapace indicating that the pleural is
narrower than<?pagebreak page60?> the vertebral as also seen in <italic>Ordosemys</italic> spp., <italic>Liaochelys jianchangensis</italic>, and <italic>Xiaochelys ningchengensis</italic> (e.g., Brinkman and
Peng, 1993; Tong et al., 2004; Danilov and Parham, 2007; Zhou, 2010a; Zhou
and Rabi, 2015).</p>
      <p id="d1e698">The plastron has a limited exposure with the inguinal buttress of the
hypoplastron. The inguinal buttress terminates at peripheral 8.</p>
</sec>
<sec id="Ch1.S5.SS3">
  <label>5.3</label><title>Osteology of XL-008</title>
      <p id="d1e709">XL-008 is a juvenile skeleton and is exposed dorsally (Fig. 4). The partial
skull is poorly preserved in articulation with the damaged cervical series.
The forelimbs are partially preserved with the distal end of the humeri, the
right ulna, and radius. The ulna is about 20 mm long.</p>
      <?pagebreak page62?><p id="d1e712"><?xmltex \hack{\newpage}?>The carapace is also strongly damaged, resulting in the extensive exposure
of the plastron in dorsal view. Large costo-peripheral fenestrae are
present, implying a juvenile stage for this specimen. The carapace shows a
subcircular profile. The preserved length is about 120 mm, but the estimated
length is about 130 mm. The maximum width is about 130 mm and comparable
with that of XL-007. The cervical scale is wide. Other sulci are obscured
due to the broken surface of the carapace. The nuchal emargination is
moderately developed.</p>
      <p id="d1e716">The hyoplastron and hyoplastron are enclosing the large central fenestra and
lateral fenestrae. The bridge is about 34 mm long.</p>
</sec>
</sec>
<sec id="Ch1.S6" sec-type="conclusions">
  <label>6</label><title>Discussion</title>
<sec id="Ch1.S6.SS1">
  <label>6.1</label><title>Taxonomy</title>
      <p id="d1e735">The three specimens represent an ontogenetic series from juvenile (XL-008),
to subadult (XL-007), and to adult (XL-009) stages. As they show no distinct
morphological differences besides the ontogenetic variance and were
collected from the same locality and horizon, we refer them to the same
taxon. The specimens are similar to <italic>Ordosemys</italic> spp. and differ from other sinemydids in
having a subcircular shell, wide vertebral scales, and large central and lateral
fenestrae of plastron (e.g., Brinkman et al., 2008; Li and Tong, 2017; Li et
al., 2019). The presence of a preneural plate is also diagnostic for
<italic>Ordosemys</italic> spp. (e.g., Brinkman et al., 2008; Li et al., 2019), but it is uncertain in
all three specimens due to the poor preservation.</p>
      <p id="d1e744">Three species are widely accepted in the genus <italic>Ordosemys</italic>: <italic>O. leios</italic>, <italic>O. liaoxiensis</italic>, and <italic>O. brinkmania</italic> (Danilov and
Parham, 2007; Brinkman et al., 2008; Li and Tong, 2017; Li et al., 2019).
Two further species, <italic>O. perforata</italic> (i.e., <italic>Asiachelys perforata</italic> sensu Sukhanov and Narmandakh, 2006; Danilov
and Parham, 2007)   and <italic>O. donghai</italic> (i.e., <italic>Machurochelys donghai</italic> sensu Ma, 1986; Brinkman et al., 2008; Li and
Tong, 2017), have been questioned in their affinities to <italic>Ordosemys</italic> in lacking the
preneural plate and wide vertebral scales (e.g., Li et al., 2019).</p>
      <p id="d1e775">The Jilin specimens differ from <italic>O. brinkmania</italic> in the presence of a hypo-xiphiplastral
fenestra. The fenestra is also present in <italic>O. leios</italic> and <italic>O. liaoxiensis</italic>, but <italic>O. leios</italic> lacks lateral
plastral fenestrae. In the former two species, the hypo-xiphiplastral
fenestra is enclosed by the hypoplastra and xiphiplastra subequally. In the
adult XL-009, however, the xiphiplastra have a major contribution relative
to the hypoplastra in enclosing the fenestra (Fig. 2). The
hypo-xiphiplastral fenestra is also known in other sinemydids including
<italic>Liaochelys jianchangensis</italic> (the adult specimen PMOL-AR00140; Zhou, 2010a), <italic>Changmachelys bohlini</italic> (the subadult specimen GSGM
05-CM-016, but uncertain in adult specimen GSGM 05 IG-02-0902; Brinkman et
al., 2013), and <italic>Jeholochelys lingyuanensis</italic> (the juvenile specimen PMOL-AR00213, but absent in adult
specimens; Shao et al., 2018). The fenestra is therefore subject to
ontogenetic variation although the hypo-xiphiplastral fenestra is always
enclosed subequally by the hypoplastra and xiphiplastra in these sinemydids.
The condition in the adult XL-009 is therefore unusual, but there is still
much left to be learned about the ontogeny and individual morphological
variation of sinemydids (Shao et al., 2018). Given that the Jilin material
only differs from <italic>O. liaoxiensis</italic> in this single character, we refer all specimens to this
species.</p><?xmltex \hack{\newpage}?>
</sec>
<?pagebreak page63?><sec id="Ch1.S6.SS2">
  <label>6.2</label><?xmltex \opttitle{Paleobiogeography of \textit{Ordosemys}}?><title>Paleobiogeography of <italic>Ordosemys</italic></title>
      <p id="d1e815">Among freshwater sinemydid turtles, <italic>Ordosemys</italic> spp. is considered to have the broadest
paleobiogeographic distribution in northern China (e.g., Danilov and Parham,
2007; Li et al., 2019). The type species <italic>Ordosemys leios</italic> is from the Ordos Basin of Inner
Mongolia (Brinkman and Peng, 1993; see also Brinkman and Wu, 1999) and has
been recently reported from the Mengyin Basin of Shandong (Li et al., 2019).
The second species, <italic>O. liaoxiensis</italic>, is known from the lacustrine deposits of Yixian
Formation of Beipiao, western Liaoning (e.g., Ji, 1995; Tong et al., 2004).
<italic>Ordosemys</italic> <italic>brinkmania</italic> comes from the fluvial deposits of the Tugulu Formation of Wuerho, eastern
Xinjiang (Danilov and Parham, 2007). <italic>Ordosemys</italic> <italic>donghai</italic> (i.e., <italic>Manchurochelys donghai</italic> sensu Ma, 1986; Brinkman et al.,
2008; Li and Tong, 2017) from Helongjiang was previously considered the
easternmost record of <italic>Ordosemys</italic> in northeastern China (Danilov and Parham, 2007), but
this taxon may not belong to <italic>Ordosemys</italic> because of the lack of a preneural and wide
vertebral scales   (e.g., Li et al., 2019). <italic>Ordosemys</italic> <italic>liaoxiensis</italic> is so far restricted to
northeastern China and the Yixian Formation of the Jehol Biota. The new
record expands the distribution of this species from western Liaoning to
southern Jilin. This occurrence thus furthermore supports that <italic>Ordosemys</italic> spp. may have
had a broader paleogeographic distribution compared to other sinemydid
turtles. As the first Mesozoic turtle of Jilin Province, <italic>Ordosemys liaoxiensis</italic> from the Early
Cretaceous Hengtongshan Formation represents the first tetrapod to indicate
the presence of the Jehol Biota in the region. It also extends the
stratigraphic range of <italic>O. liaoxiensis</italic>, which was previously only known from the lower
Yixian Formation and has not been reported from the overlying Jiufotang
Formation of Liaoning, the equivalent of the Hengtongshan Formation of Jilin
Province (e.g., Wu et al., 2018). This is likely due to a sampling bias
because we are aware of unpublished specimens from the Jiufotang Formation
that resemble <italic>Ordosemys</italic> spp. Given the morphological difference from the typical <italic>O. liaoxiensis</italic> from
the Yixian Formation and given the geographic and age difference, future
collections in the Hengtongshan Formation have good potential for evaluating
spatial patterns of the Jehol Biota.</p>
</sec>
</sec>

      
      </body>
    <back><notes notes-type="dataavailability"><title>Data availability</title>

      <p id="d1e876">Three specimens of <italic>Ordosemys</italic> described in this study are stored in Paleontological Collection of Xingling Town, Meihekou City, Jilin Province, China (accession numbers: XL-007, XL-008, and XL-009).</p>
  </notes><?xmltex \hack{\newpage}?><notes notes-type="authorcontribution"><title>Author contributions</title>

      <p id="d1e886">CFZ and WHW conceived the project and created figures. CFZ, MR and WHW
wrote and approved the paper.</p>
  </notes><notes notes-type="competinginterests"><title>Competing interests</title>

      <p id="d1e892">The authors declare that they have no conflict of interest.</p>
  </notes><ack><title>Acknowledgements</title><p id="d1e898">We would like to thank the director of the Research Center of Palaeontology
and Stratigraphy, Jilin University,  Chun-Li Sun, for his
encouragement.  Juliana Sterli and an anonymous reviewer are thanked for
their helpful comments that improved the quality of this paper.</p></ack><notes notes-type="financialsupport"><title>Financial support</title>

      <p id="d1e903">This research has been supported by the Taishan Scholar Program of Shandong Province (grant no. tsqn201812070), the Shandong Provincial Natural Science Foundation (grant no. ZR2017MD031), and the Research Fund of Shandong University of Science and Technology (grant no. 2015TDJH101) to Chang-Fu Zhou. Maton Rabi received funding from the Volkswagen Foundation
(grant no. 90 978).</p>
  </notes><notes notes-type="reviewstatement"><title>Review statement</title>

      <p id="d1e909">This paper was edited by Florian Witzmann and reviewed by Juliana Sterli and one anonymous referee.</p>
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  </ref-list></back>
    <!--<article-title-html>Presence of the Jehol Biota turtle <i>Ordosemys liaoxiensis</i> in the Early Cretaceous Hengtongshan Formation of southern Jilin Province, China</article-title-html>
<abstract-html><p>Recently, a vertebrate assemblage of the Jehol Biota has been reported from
the Early Cretaceous Hengtongshan Formation of Xingling Town, Meihekou City,
Jilin Province, China. It is dominated by the fishes <i>Lycoptera</i> and <i>Sinamia</i> and the sinemydid
turtle <i>Ordosemys</i>. Here, we describe the turtle specimens and referral to <i>Ordosemys liaoxiensis</i>, otherwise
known from the older Yixian Formation of the Jehol Biota.  It is characterized
by a subcircular shell, wide vertebral scales, well-developed plastral
fenestrae, and a major contribution from the xiphiplastra to enclose the
hypo-xiphiplastral fenestra. As the first Mesozoic turtle of Jilin Province,
this record represents the first tetrapod to indicate the presence of the
Jehol Biota in the region. Given the geographic and temporal distance from
the Yixian Formation, future collections from the Hengtongshan Formation
have good potential for evaluating spatiotemporal patterns of the Jehol
Biota.</p></abstract-html>
<ref-html id="bib1.bib1"><label>1</label><mixed-citation>
Batsch, A. J. G. C.: Versuch einer Anleitung, zur Kenntniÿund Geschichte
der Thiere und Mineralien, Akademische Buchhandlung, Jena, 53–59, 1788.
</mixed-citation></ref-html>
<ref-html id="bib1.bib2"><label>2</label><mixed-citation>
Brinkman, D. B. and Peng, J.-H.: <i>Ordosemys leios</i>, n. gen., n. sp., a new turtle from the
Early Cretaceous of the Ordos Basin, Inner Mongolia, Can. J. Earth Sci., 30,
2128–2138, 1993.
</mixed-citation></ref-html>
<ref-html id="bib1.bib3"><label>3</label><mixed-citation>
Brinkman, D. B. and Wu, X.-C.: The skull of <i>Ordosemys</i>, an Early Cretaceous turtle
from Inner Mongolia, People's Republic of China, and the interrelationships
of Eucryptodira (Chelonia, Cryptodira), Paludicola, 2, 134–147, 1999.
</mixed-citation></ref-html>
<ref-html id="bib1.bib4"><label>4</label><mixed-citation>
Brinkman, D. B., Li, J.-L., and Ye, X.-K.: Order Testudines, in: The Chinese
fossil reptiles and their kin, edited by: Li, J. L., Wu, X. C., and Zhang, F.
C., Science Press, Beijing, 35–102, 2008.
</mixed-citation></ref-html>
<ref-html id="bib1.bib5"><label>5</label><mixed-citation>
Brinkman, D. B., Yuan, C.-X., Ji, Q., Li, D.-Q., and You, H.-L.: A new
turtle from the Xiagou Formation (Early Cretaceous) of Changma Basin, Gansu
Province, P. R. China. Palaeobiol. Palaeoenv., 93, 367–382, 2013.
</mixed-citation></ref-html>
<ref-html id="bib1.bib6"><label>6</label><mixed-citation>
Chang, S.-C., Zhang, H., Renne, P. R., and Fang, Y.: High-precision
<sup>40</sup>Ar∕<sup>39</sup>Ar age for the Jehol Biota, Palaeogeogr. Palaeocl., 280, 94–104, 2009.
</mixed-citation></ref-html>
<ref-html id="bib1.bib7"><label>7</label><mixed-citation>
Chen, J., Li, B., Yao, Y., Liu, M., Yang, F., Xing D., Li, W., and Wang, Y.:
Comparison between Mesozoic volcanic rock strata in northeast of
Liaoning-south of Jilin and Yixian Formation in west of Liaoning, Acta Geol.
Sin.-Engl., 90, 2733–2746, 2016 (in Chinese with English abstract).
</mixed-citation></ref-html>
<ref-html id="bib1.bib8"><label>8</label><mixed-citation>
Chen, Y., Sun, C., Sun, Y., and Sun, W.: Cretaceous volcanic events in
southeastern Jilin Province, China: evidence from single zircon U-Pb ages,
Acta Geol. Sin.-Engl., 82, 1194–1200, 2008.
</mixed-citation></ref-html>
<ref-html id="bib1.bib9"><label>9</label><mixed-citation>
Danilov, I. G. and Parham, J. F.: The type series of “<i>Sinemys</i>” <i>wuerhoensis</i>, a problematic
turtle from the Lower Cretaceous of China, includes at least three taxa,
Palaeontology, 50, 431–444, 2007.
</mixed-citation></ref-html>
<ref-html id="bib1.bib10"><label>10</label><mixed-citation>
Ji, S.-A.: Reptiles, in: Fauna and Stratigraphy of Jurassic-Cretaceous in
Beijing and the Adjacent Areas, edited by: Ren, D., Lu, L.-W., Guo, Z.-G.,
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