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<document ID-DOI="10.1093/zoolinnean/zlac063" ID-ISSN="0024-4082" ID-Zenodo-Dep="7690954" approvalRequired="5" approvalRequired_for_treatments="5" checkinTime="1677744395091" checkinUser="plazi" docAuthor="Macaluso, Loredana, Wencker, Lukardis C M, Castrovilli, Maria, Carnevale, Giorgio &amp; Delfino, Massimo" docDate="2023" docId="9750C307FF9C4C34FEA7F694FE08F9D6" docLanguage="en" docName="zlac063.pdf" docOrigin="Zoological Journal of the Linnean Society 197 (3)" docSource="https://academic.oup.com/zoolinnean/article/197/3/569/6693955" docStyle="DocumentStyle:36B3BD6A90C22AB4F7F465C853188CC8.7:ZoolJLinnSoc.2017-.journal_article" docStyleId="36B3BD6A90C22AB4F7F465C853188CC8" docStyleName="ZoolJLinnSoc.2017-.journal_article" docStyleVersion="7" docTitle="Salamandrella DYBOWKI 1870" docType="treatment" docVersion="2" lastPageNumber="586" masterDocId="6B69BB7FFF954C25FFBFF368FF87FFC2" masterDocTitle="A comparative atlas of selected skeletal elements of European urodeles (Amphibia: Urodela) for palaeontological investigations" masterLastPageNumber="619" masterPageNumber="569" pageNumber="578" updateTime="1677845860903" updateUser="ExternalLinkService">
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<mods:title>A comparative atlas of selected skeletal elements of European urodeles (Amphibia: Urodela) for palaeontological investigations</mods:title>
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<mods:roleTerm>Author</mods:roleTerm>
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<mods:namePart>Macaluso, Loredana</mods:namePart>
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<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Wencker, Lukardis C M</mods:namePart>
</mods:name>
<mods:name type="personal">
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<mods:roleTerm>Author</mods:roleTerm>
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<mods:namePart>Castrovilli, Maria</mods:namePart>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Carnevale, Giorgio</mods:namePart>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Delfino, Massimo</mods:namePart>
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<mods:title>Zoological Journal of the Linnean Society</mods:title>
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<mods:date>2023</mods:date>
<mods:detail type="pubDate">
<mods:number>2023-03-01</mods:number>
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<mods:detail type="volume">
<mods:number>197</mods:number>
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<mods:number>3</mods:number>
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<mods:url>https://academic.oup.com/zoolinnean/article/197/3/569/6693955</mods:url>
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<mods:classification>journal article</mods:classification>
<mods:identifier type="DOI">10.1093/zoolinnean/zlac063</mods:identifier>
<mods:identifier type="ISSN">0024-4082</mods:identifier>
<mods:identifier type="Zenodo-Dep">7690954</mods:identifier>
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<treatment ID-DOI="http://doi.org/10.5281/zenodo.7695695" ID-Zenodo-Dep="7695695" LSID="urn:lsid:plazi:treatment:9750C307FF9C4C34FEA7F694FE08F9D6" httpUri="http://treatment.plazi.org/id/9750C307FF9C4C34FEA7F694FE08F9D6" lastPageId="17" lastPageNumber="586" pageId="9" pageNumber="578">
<subSubSection box="[280,661,1532,1557]" pageId="9" pageNumber="578" type="nomenclature">
<paragraph blockId="9.[280,661,1532,1557]" box="[280,661,1532,1557]" pageId="9" pageNumber="578">
<heading box="[280,661,1532,1557]" centered="true" fontSize="9" level="2" pageId="9" pageNumber="578" reason="2">
<taxonomicName ID-CoL="7BSD" ID-ENA="288314" authorityName="DYBOWKI" authorityYear="1870" box="[280,661,1532,1557]" class="Amphibia" family="Hynobiidae" genus="Salamandrella" kingdom="Animalia" order="Caudata" pageId="9" pageNumber="578" phylum="Chordata" rank="genus">
<emphasis box="[280,476,1532,1555]" italics="true" pageId="9" pageNumber="578">SALAMANDRELLA</emphasis>
DYBOWKI, 1870
</taxonomicName>
</heading>
</paragraph>
</subSubSection>
<subSubSection box="[163,780,1572,1594]" pageId="9" pageNumber="578" type="discussion">
<paragraph blockId="9.[163,780,1572,1594]" box="[163,780,1572,1594]" pageId="9" pageNumber="578">
<emphasis box="[163,252,1573,1594]" italics="true" pageId="9" pageNumber="578">Species:</emphasis>
<taxonomicName ID-CoL="6XBLN" ID-ENA="288315" authority="* Dybowski, 1870" authorityName="* Dybowski" authorityYear="1870" box="[264,776,1572,1594]" class="Amphibia" family="Hynobiidae" genus="Salamandrella" kingdom="Animalia" order="Caudata" pageId="9" pageNumber="578" phylum="Chordata" rank="species" species="keyserlingii">
<emphasis box="[264,574,1572,1593]" italics="true" pageId="9" pageNumber="578">Salamandrella keyserlingii</emphasis>
* Dybowski, 1870
</taxonomicName>
.
</paragraph>
</subSubSection>
<subSubSection lastPageId="17" lastPageNumber="586" pageId="9" pageNumber="578" type="description">
<paragraph blockId="9.[163,452,1637,1661]" box="[163,452,1637,1661]" pageId="9" pageNumber="578">
<heading box="[163,452,1637,1661]" centered="true" fontSize="9" level="2" pageId="9" pageNumber="578" reason="2">
<emphasis box="[163,452,1637,1661]" italics="true" pageId="9" pageNumber="578">Oticoccipitum complex</emphasis>
</heading>
</paragraph>
<paragraph blockId="9.[163,780,1678,1761]" pageId="9" pageNumber="578">
In hynobiids, the prootic and opisthoticexoccipital remain unfused in morphologically mature specimens; therefore, there is no oticoccipitum complex (
<bibRefCitation author="Jia J &amp; Jiang JP &amp; Zhang MH &amp; Gao KQ" box="[543,697,1739,1761]" pageId="9" pageNumber="578" pagination="0211069" refId="ref36521" refString="Jia J, Jiang JP, Zhang MH, Gao KQ. 2019. Osteology of Batrachuperus yenyuanensis (Urodela, Hynobiidae), a highaltitude mountain stream salamander from western China. PLoS One 14: e 0211069." type="journal article" year="2019">
Jia
<emphasis box="[582,634,1739,1761]" italics="true" pageId="9" pageNumber="578">et al.</emphasis>
, 2019
</bibRefCitation>
).
</paragraph>
<paragraph blockId="9.[163,339,1806,1830]" box="[163,339,1806,1830]" pageId="9" pageNumber="578">
<heading box="[163,339,1806,1830]" centered="true" fontSize="9" level="2" pageId="9" pageNumber="578" reason="2">
<emphasis box="[163,339,1806,1830]" italics="true" pageId="9" pageNumber="578">
Atlas (
<figureCitation box="[243,330,1806,1830]" captionStart="Figure 6" captionStartId="8.[145,225,1761,1783]" captionTargetBox="[246,1324,199,1717]" captionTargetId="figure-24@8.[241,1328,195,1721]" captionTargetPageId="8" captionText="Figure 6. Atlantes of European urodeles. A, Salamandrella keyserlingii (BSPGM 5451), most probably not completely ossified (see the Material section). B, Proteus anguinus (BSPGM 4538). C, Speleomantes strinatii (MDHC 225), most probably not completely ossified (see the Material section). D, Salamandrina perspicillata (MDHC 407). E, Chioglossa lusitanica (MNCN 16099). F, Mertensiella caucasica (MNCN 23821). G, Salamandra salamandra (MDHC 396). From left to right: anterior, dorsal, lateral (right lateral for AC, E, F; left lateral for D, G), posterior and ventral views. Scale bars: 1 mm." figureDoi="http://doi.org/10.5281/zenodo.7690967" httpUri="https://zenodo.org/record/7690967/files/figure.png" pageId="9" pageNumber="578">Fig. 6A</figureCitation>
)
</emphasis>
</heading>
</paragraph>
<paragraph blockId="9.[163,778,1847,1900]" lastBlockId="12.[145,761,1591,1736]" lastPageId="12" lastPageNumber="581" pageId="9" pageNumber="578">
The neural canal is triangular in anterior view and is at least twice as high as each occipital joint. In posterior view, the neural canal is two or three times as wide as the circular cotyle. The occipital joints are elliptical, with the major axis horizontal (or subhorizontal). The articular surfaces of the odontoid process contact each other, such that in ventral view no groove is visible between them. In ventral view, the base of the odontoid process is wider than each occipital joint. The neural crest is absent. Based on the literature, in the completely ossified atlantes, a bulge should be present in its place (
<bibRefCitation author="Ratnikov VY &amp; Litvinchuk SN" pageId="9" pageNumber="578" pagination="57 - 68" refId="ref37221" refString="Ratnikov VY, Litvinchuk SN. 2009. Atlantal vertebrae of tailed amphibians of Russia and adjacent countries. Russian Journal of Herpetology 16: 57 - 68." type="journal article" year="2009">Ratnikov &amp; Litvinchuk, 2009</bibRefCitation>
); however, in all specimens examined in this work, the walls of the neural arch do not contact each other at the dorsal midline, and two articular surfaces are present in the middle. These are visible in dorsal view and articulated through cartilage. The secondary crests and the neural spine are absent. The lateral surface of the atlas bears only the foramen of the first spinal nerve. The incisura vertebralis cranialis is generally absent or small. In lateral view, the dorsal edge of the neural arch is sub-horizontal. The posterior margin of the lateral wall of the neural arch, ventral to the wide incisura caudalis in lateral view, is concave, inclined or sub-vertical. The maximum concavity of the incisura vertebralis caudalis is dorsal to the horizontal plane containing the maximum concavity of the incisura cranialis. The lateral crests are absent, and the inferior crests are low or also absent. In posterior view, the neural arch is dorsally convex (inverted U-shaped). In lateral view, the postzygapophyses extend posteriorly beyond the cotyle for more than half of their length, and in posterior view, these structures are sub-horizontal. In dorsal view, the neural arch is anteriorly and posteriorly concave (with V-shaped concavities). The cotyle is generally not visible in dorsal view, or only slightly visible in the middle of the incisura dorsalis. The ventral surface generally bears more than one foramen.
</paragraph>
<caption ID-DOI="http://doi.org/10.5281/zenodo.7690971" ID-Zenodo-Dep="7690971" httpUri="https://zenodo.org/record/7690971/files/figure.png" pageId="10" pageNumber="579" startId="10.[146,226,1501,1523]" targetBox="[148,1422,199,1458]" targetPageId="10">
<paragraph blockId="10.[145,1425,1501,1611]" pageId="10" pageNumber="579">
<emphasis bold="true" box="[146,253,1501,1523]" pageId="10" pageNumber="579">Figure 8.</emphasis>
Atlantes of European urodeles. A,
<taxonomicName baseAuthorityName="Linnaeus" baseAuthorityYear="1758" box="[635,847,1501,1523]" class="Amphibia" family="Salamandridae" genus="Lissotriton" kingdom="Animalia" order="Caudata" pageId="10" pageNumber="579" phylum="Chordata" rank="species" species="vulgaris">
<emphasis box="[635,847,1501,1523]" italics="true" pageId="10" pageNumber="579">Lissotriton vulgaris</emphasis>
</taxonomicName>
(MDHC 135). B,
<taxonomicName baseAuthorityName="Linnaeus" baseAuthorityYear="1758" box="[1034,1246,1501,1523]" class="Amphibia" family="Salamandridae" genus="Lissotriton" kingdom="Animalia" order="Caudata" pageId="10" pageNumber="579" phylum="Chordata" rank="species" species="vulgaris">
<emphasis box="[1034,1246,1501,1523]" italics="true" pageId="10" pageNumber="579">Lissotriton vulgaris</emphasis>
</taxonomicName>
(MDHC 133). C,
<taxonomicName baseAuthorityName="Gray" baseAuthorityYear="1835" box="[145,380,1530,1552]" class="Amphibia" family="Salamandridae" genus="Ommatotriton" kingdom="Animalia" order="Caudata" pageId="10" pageNumber="579" phylum="Chordata" rank="species" species="vittatus">
<emphasis box="[145,380,1530,1552]" italics="true" pageId="10" pageNumber="579">Ommatotriton vittatus</emphasis>
</taxonomicName>
(MNCN 13193), with right postzygapophyses missing. D,
<taxonomicName authorityName="* Michahelles" authorityYear="1830" box="[980,1159,1530,1552]" class="Amphibia" family="Salamandridae" genus="Pleurodeles" kingdom="Animalia" order="Caudata" pageId="10" pageNumber="579" phylum="Chordata" rank="species" species="aealtl">
<emphasis box="[980,1159,1530,1552]" italics="true" pageId="10" pageNumber="579">Pleurodeles waltl</emphasis>
</taxonomicName>
(MDHC 253). E,
<taxonomicName authorityName="* (Laurenti" authorityYear="1768" class="Amphibia" family="Salamandridae" genus="Triturus" kingdom="Animalia" order="Caudata" pageId="10" pageNumber="579" phylum="Chordata" rank="species" species="carnifex">
<emphasis italics="true" pageId="10" pageNumber="579">Triturus carnifex</emphasis>
</taxonomicName>
(MDHC 38). From left to right: anterior, dorsal, lateral (right lateral for A, D; left lateral for BC, E), posterior and ventral views. Scale bars: 1 mm.
</paragraph>
</caption>
<caption ID-DOI="http://doi.org/10.5281/zenodo.7690973" ID-Zenodo-Dep="7690973" httpUri="https://zenodo.org/record/7690973/files/figure.png" pageId="11" pageNumber="580" startId="11.[163,242,1750,1772]" targetBox="[206,1399,196,1706]" targetPageId="11">
<paragraph blockId="11.[163,1443,1750,1889]" pageId="11" pageNumber="580">
<emphasis bold="true" box="[163,268,1750,1772]" pageId="11" pageNumber="580">Figure 9.</emphasis>
Precaudal vertebrae of European urodeles. A,
<taxonomicName authorityName="* Dybowski" authorityYear="1870" box="[765,1053,1750,1772]" class="Amphibia" family="Hynobiidae" genus="Salamandrella" kingdom="Animalia" order="Caudata" pageId="11" pageNumber="580" phylum="Chordata" rank="species" species="keyserlingii">
<emphasis box="[765,1053,1750,1772]" italics="true" pageId="11" pageNumber="580">Salamandrella keyserlingii</emphasis>
</taxonomicName>
(BSPGM 5451). B,
<taxonomicName authorityName="* Laurenti" authorityYear="1768" box="[1258,1443,1750,1772]" class="Amphibia" family="Proteidae" genus="Proteus" kingdom="Animalia" order="Caudata" pageId="11" pageNumber="580" phylum="Chordata" rank="species" species="anguinus">
<emphasis box="[1258,1443,1750,1772]" italics="true" pageId="11" pageNumber="580">Proteus anguinus</emphasis>
</taxonomicName>
(BSPGM 4539). C, first precaudal vertebra of
<taxonomicName baseAuthorityName="Aellen" baseAuthorityYear="1958" box="[646,882,1779,1801]" class="Amphibia" family="Plethodontidae" genus="Speleomantes" kingdom="Animalia" order="Caudata" pageId="11" pageNumber="580" phylum="Chordata" rank="species" species="strinatii">
<emphasis box="[646,882,1779,1801]" italics="true" pageId="11" pageNumber="580">Speleomantes strinatii</emphasis>
</taxonomicName>
(MDHC 225). D, precaudal vertebra of
<taxonomicName baseAuthorityName="Aellen" baseAuthorityYear="1958" class="Amphibia" family="Plethodontidae" genus="Speleomantes" kingdom="Animalia" order="Caudata" pageId="11" pageNumber="580" phylum="Chordata" rank="species" species="strinatii">
<emphasis italics="true" pageId="11" pageNumber="580">Speleomantes strinatii</emphasis>
</taxonomicName>
(MDHC 225) close to the pelvis. E,
<taxonomicName authorityName="* (Savi" authorityYear="1821" box="[638,937,1809,1831]" class="Amphibia" family="Salamandridae" genus="Salamandrina" kingdom="Animalia" order="Caudata" pageId="11" pageNumber="580" phylum="Chordata" rank="species" species="perspicillata">
<emphasis box="[638,937,1809,1831]" italics="true" pageId="11" pageNumber="580">Salamandrina perspicillata</emphasis>
</taxonomicName>
(MDHC 228). F,
<taxonomicName authorityName="Bocage" authorityYear="1864" box="[1121,1353,1809,1831]" class="Amphibia" family="Salamandridae" genus="Chioglossa" kingdom="Animalia" order="Caudata" pageId="11" pageNumber="580" phylum="Chordata" rank="species" species="lusitanica">
<emphasis box="[1121,1353,1809,1831]" italics="true" pageId="11" pageNumber="580">Chioglossa lusitanica</emphasis>
</taxonomicName>
(MNCN 16099). From left to right: anterior, dorsal, lateral (right lateral for A, CF; left lateral for B), posterior and ventral views. Scale bars: 1 mm.
</paragraph>
</caption>
<caption ID-DOI="http://doi.org/10.5281/zenodo.7690975" ID-Zenodo-Dep="7690975" httpUri="https://zenodo.org/record/7690975/files/figure.png" pageId="12" pageNumber="581" startId="12.[145,226,1443,1465]" targetBox="[148,1426,199,1395]" targetPageId="12">
<paragraph blockId="12.[145,1426,1443,1553]" pageId="12" pageNumber="581">
<emphasis bold="true" box="[145,268,1443,1465]" pageId="12" pageNumber="581">Figure 10.</emphasis>
Precaudal vertebrae of European urodeles. A,
<taxonomicName baseAuthorityName="Waga" baseAuthorityYear="1876" box="[789,1038,1443,1465]" class="Amphibia" family="Salamandridae" genus="Mertensiella" kingdom="Animalia" order="Caudata" pageId="12" pageNumber="581" phylum="Chordata" rank="species" species="caucasica">
<emphasis box="[789,1038,1443,1465]" italics="true" pageId="12" pageNumber="581">Mertensiella caucasica</emphasis>
</taxonomicName>
(MNCN 23821). BD,
<taxonomicName authorityName="* (Linnaeus" authorityYear="1758" class="Amphibia" family="Salamandridae" genus="Salamandra" kingdom="Animalia" order="Caudata" pageId="12" pageNumber="581" phylum="Chordata" rank="species" species="salamandra">
<emphasis italics="true" pageId="12" pageNumber="581">Salamandra salamandra</emphasis>
</taxonomicName>
(MDHC 396): B, one of the first trunk vertebrae; C, trunk vertebra close to the pelvis; and D, sacral vertebra. E,
<taxonomicName baseAuthorityName="Duges" baseAuthorityYear="1852" box="[173,342,1502,1524]" class="Amphibia" family="Salamandridae" genus="Calotriton" kingdom="Animalia" order="Caudata" pageId="12" pageNumber="581" phylum="Chordata" rank="species" species="asper">
<emphasis box="[173,342,1502,1524]" italics="true" pageId="12" pageNumber="581">Calotriton asper</emphasis>
</taxonomicName>
(MNCN 16122). From left to right: anterior, dorsal, lateral (right lateral for B, D; left lateral for A, C, E), posterior and ventral views. Scale bars: 1 mm.
</paragraph>
</caption>
<paragraph blockId="12.[145,502,1795,1819]" box="[145,502,1795,1819]" pageId="12" pageNumber="581">
<heading box="[145,502,1795,1819]" centered="true" fontSize="9" level="2" pageId="12" pageNumber="581" reason="2">
<emphasis box="[145,502,1795,1819]" italics="true" pageId="12" pageNumber="581">
Precaudal vertebrae (
<figureCitation box="[406,493,1795,1819]" captionStart="Figure 9" captionStartId="11.[163,242,1750,1772]" captionTargetBox="[206,1399,196,1706]" captionTargetId="figure-24@11.[203,1402,195,1710]" captionTargetPageId="11" captionText="Figure 9. Precaudal vertebrae of European urodeles. A, Salamandrella keyserlingii (BSPGM 5451). B, Proteus anguinus (BSPGM 4539). C, first precaudal vertebra of Speleomantes strinatii (MDHC 225). D, precaudal vertebra of Speleomantes strinatii (MDHC 225) close to the pelvis. E, Salamandrina perspicillata (MDHC 228). F, Chioglossa lusitanica (MNCN 16099). From left to right: anterior, dorsal, lateral (right lateral for A, CF; left lateral for B), posterior and ventral views. Scale bars: 1 mm." figureDoi="http://doi.org/10.5281/zenodo.7690973" httpUri="https://zenodo.org/record/7690973/files/figure.png" pageId="12" pageNumber="581">Fig. 9A</figureCitation>
)
</emphasis>
</heading>
</paragraph>
<paragraph blockId="12.[145,761,1835,1888]" lastBlockId="17.[163,779,197,955]" lastPageId="17" lastPageNumber="586" pageId="12" pageNumber="581">The precaudal vertebrae are amphicoelous. The neural canal is pentagonal or circular in anterior view, slightly higher or lower than the cotyle. The cotyles are circular or slightly elliptical, with the major axis horizontal. In lateral view, the anterior edge of the neural arch between the cotyle and prezygapophyses is concave or vertical. Diapophyses and parapophyses are usually fused and not recognizable (unicipital transverse processes). The transverse processes are rectangular in anterior view and slightly pointed posteroventrally. The transverse processes do not cover (or cover only in part) the posterior edge of the neural arch in lateral view. In lateral view, the vertebrae are dorsoventrally compressed, and most of the height of the vertebrae is formed by the centrum and the neural canal (only one-fifth of the height of the vertebra is formed by the neural arch dorsal to the postzygapophyses). In lateral view, the neural arch dorsal to the prezygapophyses is not visible or only slightly visible. The neural crest is absent. The neural spine is present in the form of a low and thick bulge that is posteriorly convex and extends posteriorly as far as the postzygapophyses. The anterior and posterior zygapophyseal and ventral crests are absent. The lateral surface of the vertebrae is smooth. The only foramen present is visible in anterior and lateral views, in the ventral half of the proximal edge of the transverse processes. In lateral view, the dorsal edge of the neural arch is anteriorly sub-horizontal. The incisura vertebralis caudalis is not deep and only present as a small posterior concavity (wider in the first precaudal vertebrae), and the neural arch between the centrum and postzygapophyses is slightly concave or convex. In posterior view, the neural arch is dorsally flat or convex (inverted U-shaped). In lateral view, the postzygapophyses extend posteriorly beyond the cotyle for more than half of their length, and in posterior view, postzygapophyses are sub-horizontal. In dorsal view, the neural arch is anteriorly concave (U-shaped), and posteriorly, the incisura dorsalis is not visible. The edge of the anterior cotyle is visible in dorsal view, whereas the posterior cotyle is not visible. In lateral view, the ventral profile of the centrum is strongly concave. The ventral surface is generally smooth.</paragraph>
<caption ID-DOI="http://doi.org/10.5281/zenodo.7690977" ID-Zenodo-Dep="7690977" httpUri="https://zenodo.org/record/7690977/files/figure.png" pageId="13" pageNumber="582" startId="13.[163,241,1803,1825]" targetBox="[262,1342,198,1760]" targetPageId="13">
<paragraph blockId="13.[163,1444,1803,1913]" pageId="13" pageNumber="582">
<emphasis bold="true" box="[163,280,1803,1825]" pageId="13" pageNumber="582">Figure 11.</emphasis>
Precaudal vertebrae of European urodeles. A,
<taxonomicName authorityName="* (Gravenhorst" authorityYear="1829" box="[773,1030,1803,1825]" class="Amphibia" family="Salamandridae" genus="Euproctus" kingdom="Animalia" order="Caudata" pageId="13" pageNumber="582" phylum="Chordata" rank="species" species="platycephalus">
<emphasis box="[773,1030,1803,1825]" italics="true" pageId="13" pageNumber="582">Euproctus platycephalus</emphasis>
</taxonomicName>
(MDHC 405). B,
<taxonomicName authorityName="* (Laurenti" authorityYear="1768" box="[1210,1443,1803,1825]" class="Amphibia" family="Salamandridae" genus="Ichthyosaura" kingdom="Animalia" order="Caudata" pageId="13" pageNumber="582" phylum="Chordata" rank="species" species="alpestris">
<emphasis box="[1210,1443,1803,1825]" italics="true" pageId="13" pageNumber="582">Ichthyosaura alpestris</emphasis>
</taxonomicName>
(MDHC 352). C,
<taxonomicName baseAuthorityName="Linnaeus" baseAuthorityYear="1758" box="[329,528,1832,1854]" class="Amphibia" family="Salamandridae" genus="Lissotriton" kingdom="Animalia" order="Caudata" pageId="13" pageNumber="582" phylum="Chordata" rank="species" species="vulgaris">
<emphasis box="[329,528,1832,1854]" italics="true" pageId="13" pageNumber="582">Lissotriton vulgaris</emphasis>
</taxonomicName>
(MDHC 135). D,
<taxonomicName baseAuthorityName="Gray" baseAuthorityYear="1835" box="[702,930,1832,1854]" class="Amphibia" family="Salamandridae" genus="Ommatotriton" kingdom="Animalia" order="Caudata" pageId="13" pageNumber="582" phylum="Chordata" rank="species" species="vittatus">
<emphasis box="[702,930,1832,1854]" italics="true" pageId="13" pageNumber="582">Ommatotriton vittatus</emphasis>
</taxonomicName>
(MNCN 13193). E,
<taxonomicName authorityName="* Michahelles" authorityYear="1830" box="[1128,1302,1832,1854]" class="Amphibia" family="Salamandridae" genus="Pleurodeles" kingdom="Animalia" order="Caudata" pageId="13" pageNumber="582" phylum="Chordata" rank="species" species="aealtl">
<emphasis box="[1128,1302,1832,1854]" italics="true" pageId="13" pageNumber="582">Pleurodeles waltl</emphasis>
</taxonomicName>
(MDHC 253). F, G,
<taxonomicName authorityName="* (Laurenti" authorityYear="1768" box="[211,382,1862,1884]" class="Amphibia" family="Salamandridae" genus="Triturus" kingdom="Animalia" order="Caudata" pageId="13" pageNumber="582" phylum="Chordata" rank="species" species="carnifex">
<emphasis box="[211,382,1862,1884]" italics="true" pageId="13" pageNumber="582">Triturus carnifex</emphasis>
</taxonomicName>
(MDHC 38): F, one of the first trunk vertebrae; and G, trunk vertebra close to the pelvis. From left to right: anterior, dorsal, lateral (left lateral for AC, E, F; right lateral for D, G), posterior and ventral views. Scale bars: 1 mm.
</paragraph>
</caption>
<caption ID-DOI="http://doi.org/10.5281/zenodo.7690979" ID-Zenodo-Dep="7690979" httpUri="https://zenodo.org/record/7690979/files/figure.png" pageId="14" pageNumber="583" startId="14.[145,225,1786,1808]" targetBox="[379,1190,197,1744]" targetPageId="14">
<paragraph blockId="14.[145,1426,1786,1896]" pageId="14" pageNumber="583">
<emphasis bold="true" box="[145,265,1786,1808]" pageId="14" pageNumber="583">Figure 12.</emphasis>
Caudal vertebrae of European urodeles. A,
<taxonomicName authorityName="* Dybowski" authorityYear="1870" box="[739,1030,1786,1808]" class="Amphibia" family="Hynobiidae" genus="Salamandrella" kingdom="Animalia" order="Caudata" pageId="14" pageNumber="583" phylum="Chordata" rank="species" species="keyserlingii">
<emphasis box="[739,1030,1786,1808]" italics="true" pageId="14" pageNumber="583">Salamandrella keyserlingii</emphasis>
</taxonomicName>
(BSPGM 5451). B,
<taxonomicName authorityName="* Laurenti" authorityYear="1768" box="[1238,1425,1786,1808]" class="Amphibia" family="Proteidae" genus="Proteus" kingdom="Animalia" order="Caudata" pageId="14" pageNumber="583" phylum="Chordata" rank="species" species="anguinus">
<emphasis box="[1238,1425,1786,1808]" italics="true" pageId="14" pageNumber="583">Proteus anguinus</emphasis>
</taxonomicName>
(BSPGM 4538). C,
<taxonomicName baseAuthorityName="Aellen" baseAuthorityYear="1958" box="[352,599,1815,1837]" class="Amphibia" family="Plethodontidae" genus="Speleomantes" kingdom="Animalia" order="Caudata" pageId="14" pageNumber="583" phylum="Chordata" rank="species" species="strinatii">
<emphasis box="[352,599,1815,1837]" italics="true" pageId="14" pageNumber="583">Speleomantes strinatii</emphasis>
</taxonomicName>
(MDHC 225). D,
<taxonomicName authorityName="* (Savi" authorityYear="1821" box="[792,1097,1815,1837]" class="Amphibia" family="Salamandridae" genus="Salamandrina" kingdom="Animalia" order="Caudata" pageId="14" pageNumber="583" phylum="Chordata" rank="species" species="perspicillata">
<emphasis box="[792,1097,1815,1837]" italics="true" pageId="14" pageNumber="583">Salamandrina perspicillata</emphasis>
</taxonomicName>
(MDHC 407). E,
<taxonomicName authority="(BSPGM 2730)" baseAuthorityName="BSPGM" baseAuthorityYear="2730" class="Amphibia" family="Salamandridae" genus="Mertensiella" kingdom="Animalia" order="Caudata" pageId="14" pageNumber="583" phylum="Chordata" rank="species" species="caucasica">
<emphasis italics="true" pageId="14" pageNumber="583">Mertensiella caucasica</emphasis>
(BSPGM 2730)
</taxonomicName>
. F,
<taxonomicName authorityName="* (Linnaeus" authorityYear="1758" box="[442,708,1845,1867]" class="Amphibia" family="Salamandridae" genus="Salamandra" kingdom="Animalia" order="Caudata" pageId="14" pageNumber="583" phylum="Chordata" rank="species" species="salamandra">
<emphasis box="[442,708,1845,1867]" italics="true" pageId="14" pageNumber="583">Salamandra salamandra</emphasis>
</taxonomicName>
(MDHC 234). G,
<taxonomicName authorityName="* (Linnaeus" authorityYear="1758" box="[886,1152,1845,1867]" class="Amphibia" family="Salamandridae" genus="Salamandra" kingdom="Animalia" order="Caudata" pageId="14" pageNumber="583" phylum="Chordata" rank="species" species="salamandra">
<emphasis box="[886,1152,1845,1867]" italics="true" pageId="14" pageNumber="583">Salamandra salamandra</emphasis>
</taxonomicName>
(MDHC 396). From left to right: anterior, dorsal, lateral (right lateral for A, B, DF; left lateral for C, G), posterior and ventral views. Scale bars: 1 mm.
</paragraph>
</caption>
<caption ID-DOI="http://doi.org/10.5281/zenodo.7690981" ID-Zenodo-Dep="7690981" httpUri="https://zenodo.org/record/7690981/files/figure.png" pageId="15" pageNumber="584" startId="15.[163,241,1815,1837]" targetBox="[351,1254,215,1772]" targetPageId="15">
<paragraph blockId="15.[163,1443,1815,1895]" pageId="15" pageNumber="584">
<emphasis bold="true" box="[163,279,1815,1837]" pageId="15" pageNumber="584">Figure 13.</emphasis>
Caudal vertebrae of European urodeles. A,
<taxonomicName baseAuthorityName="Duges" baseAuthorityYear="1852" box="[732,900,1815,1837]" class="Amphibia" family="Salamandridae" genus="Calotriton" kingdom="Animalia" order="Caudata" pageId="15" pageNumber="584" phylum="Chordata" rank="species" species="asper">
<emphasis box="[732,900,1815,1837]" italics="true" pageId="15" pageNumber="584">Calotriton asper</emphasis>
</taxonomicName>
(MNCN 16122). B,
<taxonomicName authorityName="* (Gravenhorst" authorityYear="1829" box="[1100,1355,1815,1837]" class="Amphibia" family="Salamandridae" genus="Euproctus" kingdom="Animalia" order="Caudata" pageId="15" pageNumber="584" phylum="Chordata" rank="species" species="platycephalus">
<emphasis box="[1100,1355,1815,1837]" italics="true" pageId="15" pageNumber="584">Euproctus platycephalus</emphasis>
</taxonomicName>
(MDHC 405). C, D,
<taxonomicName authorityName="* (Laurenti" authorityYear="1768" box="[280,520,1844,1866]" class="Amphibia" family="Salamandridae" genus="Ichthyosaura" kingdom="Animalia" order="Caudata" pageId="15" pageNumber="584" phylum="Chordata" rank="species" species="alpestris">
<emphasis box="[280,520,1844,1866]" italics="true" pageId="15" pageNumber="584">Ichthyosaura alpestris</emphasis>
</taxonomicName>
(MDHC 391). E,
<taxonomicName baseAuthorityName="Linnaeus" baseAuthorityYear="1758" box="[705,916,1844,1866]" class="Amphibia" family="Salamandridae" genus="Lissotriton" kingdom="Animalia" order="Caudata" pageId="15" pageNumber="584" phylum="Chordata" rank="species" species="vulgaris">
<emphasis box="[705,916,1844,1866]" italics="true" pageId="15" pageNumber="584">Lissotriton vulgaris</emphasis>
</taxonomicName>
(MDHC 135). From left to right: anterior, dorsal, lateral (left lateral for A, E; right lateral for BD), posterior and ventral views. Scale bars: 1 mm.
</paragraph>
</caption>
<caption ID-DOI="http://doi.org/10.5281/zenodo.7690983" ID-Zenodo-Dep="7690983" httpUri="https://zenodo.org/record/7690983/files/figure.png" pageId="16" pageNumber="585" startId="16.[145,225,1678,1700]" targetBox="[240,1323,199,1634]" targetPageId="16">
<paragraph blockId="16.[145,1426,1678,1759]" pageId="16" pageNumber="585">
<emphasis bold="true" box="[145,264,1678,1700]" pageId="16" pageNumber="585">Figure 14.</emphasis>
Caudal vertebrae of European urodeles. A,
<taxonomicName authorityName="* (Litvinchuk, Zuiderwijk, Borkin &amp; Rosanov" authorityYear="2005" box="[735,1004,1678,1700]" class="Amphibia" family="Salamandridae" genus="Ommatotriton" kingdom="Animalia" order="Caudata" pageId="16" pageNumber="585" phylum="Chordata" rank="species" species="ophryticus">
<emphasis box="[735,1004,1678,1700]" italics="true" pageId="16" pageNumber="585">Ommatotriton ophryticus</emphasis>
</taxonomicName>
(MNCN 40462). B, C,
<taxonomicName authorityName="* Michahelles" authorityYear="1830" box="[1242,1426,1678,1700]" class="Amphibia" family="Salamandridae" genus="Pleurodeles" kingdom="Animalia" order="Caudata" pageId="16" pageNumber="585" phylum="Chordata" rank="species" species="aealtl">
<emphasis box="[1242,1426,1678,1700]" italics="true" pageId="16" pageNumber="585">Pleurodeles waltl</emphasis>
</taxonomicName>
(MDHC 253). D,
<taxonomicName authorityName="* (Laurenti" authorityYear="1768" box="[316,492,1708,1730]" class="Amphibia" family="Salamandridae" genus="Triturus" kingdom="Animalia" order="Caudata" pageId="16" pageNumber="585" phylum="Chordata" rank="species" species="carnifex">
<emphasis box="[316,492,1708,1730]" italics="true" pageId="16" pageNumber="585">Triturus carnifex</emphasis>
</taxonomicName>
(MDHC 299). E,
<taxonomicName authorityName="* (Laurenti" authorityYear="1768" box="[669,845,1708,1730]" class="Amphibia" family="Salamandridae" genus="Triturus" kingdom="Animalia" order="Caudata" pageId="16" pageNumber="585" phylum="Chordata" rank="species" species="carnifex">
<emphasis box="[669,845,1708,1730]" italics="true" pageId="16" pageNumber="585">Triturus carnifex</emphasis>
</taxonomicName>
(MDHC 145). From left to right: anterior, dorsal, lateral (right lateral for A, C; left lateral for B, D, E), posterior and ventral views. Scale bars: 1 mm.
</paragraph>
</caption>
<paragraph blockId="17.[163,501,1003,1027]" box="[163,501,1003,1027]" pageId="17" pageNumber="586">
<heading box="[163,501,1003,1027]" centered="true" fontSize="9" level="2" pageId="17" pageNumber="586" reason="2">
<emphasis box="[163,501,1003,1027]" italics="true" pageId="17" pageNumber="586">
Caudal vertebrae (
<figureCitation box="[390,492,1003,1027]" captionStart="Figure 12" captionStartId="14.[145,225,1786,1808]" captionTargetBox="[379,1190,197,1744]" captionTargetId="figure-25@14.[377,1192,195,1746]" captionTargetPageId="14" captionText="Figure 12. Caudal vertebrae of European urodeles. A, Salamandrella keyserlingii (BSPGM 5451). B, Proteus anguinus (BSPGM 4538). C, Speleomantes strinatii (MDHC 225). D, Salamandrina perspicillata (MDHC 407). E, Mertensiella caucasica (BSPGM 2730). F, Salamandra salamandra (MDHC 234). G, Salamandra salamandra (MDHC 396). From left to right: anterior, dorsal, lateral (right lateral for A, B, DF; left lateral for C, G), posterior and ventral views. Scale bars: 1 mm." figureDoi="http://doi.org/10.5281/zenodo.7690979" httpUri="https://zenodo.org/record/7690979/files/figure.png" pageId="17" pageNumber="586">Fig. 12A</figureCitation>
)
</emphasis>
</heading>
</paragraph>
<paragraph blockId="17.[163,779,1044,1556]" pageId="17" pageNumber="586">The caudal vertebrae are not particularly high (height/length ratio &lt;1.25). The neural canal is pentagonal or circular, whereas the haemal canal is elliptical or U-shaped. The neural canal is wider and lower than the haemal canal, and the haemal arch is generally slender. The transverse processes are long and slender, sub-cylindrical and pointing ventrally. The neural crest is absent or low; a posterodorsal bulge is present. Zygapophyseal and ventral crests are absent. The lateral surface is smooth, with a single foramen visible at the base of the haemal arch. In lateral view, the anterior edge of the haemal arch is variably inclined, convex or concave, whereas the posteroventral edge of the haemal arch forms a sharp tip in lateral view. In ventral view, the posterior edge of the haemal arch is not forked. The haemal crest is either low or absent.</paragraph>
</subSubSection>
</treatment>
</document>