488 lines
51 KiB
XML
488 lines
51 KiB
XML
<document ID-DOI="10.5252/geodiversitas2020v42a22" ID-GBIF-Dataset="99b2432e-c149-4b82-a584-4b33d116d198" ID-ISSN="1638-9395" ID-Zenodo-Dep="4447648" ID-ZooBank="urn:lsid:zoobank.org:pub:07001ACA-EBDE-4256-BCB9-55E3159F81DC" checkinTime="1610990288905" checkinUser="felipe" docAuthor="Lapparent, France de, Bailon, Salvador, Augé, Marc Louis & Rage, Jean-Claude" docDate="2020" docId="CB2D87E12E3AFFCEFE4B4733FD4AF96A" docLanguage="en" docName="Geodiversitas.42.22.409-426.pdf" docOrigin="Geodiversitas 42 (22)" docStyle="DocumentStyle:F830B10FF475E64C1F1601E3B32DDC00.3:Geodiversitas.2018-.journal_article" docStyleId="F830B10FF475E64C1F1601E3B32DDC00" docStyleName="Geodiversitas.2018-.journal_article" docStyleVersion="3" docTitle="Varanus sp. Merrem 1820" docType="treatment" docVersion="7" lastPageNumber="420" masterDocId="3714FF992E31FFC3FFC24100FFFBFFC6" masterDocTitle="Amphibians and reptiles from the Neogene of Afghanistan" masterLastPageNumber="426" masterPageNumber="409" pageNumber="418" updateTime="1661832981842" updateUser="ExternalLinkService" zenodo-license-document="CC0-1.0" zenodo-license-figures="CC0-1.0">
|
|
<mods:mods xmlns:mods="http://www.loc.gov/mods/v3">
|
|
<mods:titleInfo>
|
|
<mods:title>Amphibians and reptiles from the Neogene of Afghanistan</mods:title>
|
|
</mods:titleInfo>
|
|
<mods:name type="personal">
|
|
<mods:role>
|
|
<mods:roleTerm>Author</mods:roleTerm>
|
|
</mods:role>
|
|
<mods:namePart>Lapparent, France de</mods:namePart>
|
|
</mods:name>
|
|
<mods:name type="personal">
|
|
<mods:role>
|
|
<mods:roleTerm>Author</mods:roleTerm>
|
|
</mods:role>
|
|
<mods:namePart>Bailon, Salvador</mods:namePart>
|
|
</mods:name>
|
|
<mods:name type="personal">
|
|
<mods:role>
|
|
<mods:roleTerm>Author</mods:roleTerm>
|
|
</mods:role>
|
|
<mods:namePart>Augé, Marc Louis</mods:namePart>
|
|
</mods:name>
|
|
<mods:name type="personal">
|
|
<mods:role>
|
|
<mods:roleTerm>Author</mods:roleTerm>
|
|
</mods:role>
|
|
<mods:namePart>Rage, Jean-Claude</mods:namePart>
|
|
</mods:name>
|
|
<mods:typeOfResource>text</mods:typeOfResource>
|
|
<mods:relatedItem type="host">
|
|
<mods:titleInfo>
|
|
<mods:title>Geodiversitas</mods:title>
|
|
</mods:titleInfo>
|
|
<mods:part>
|
|
<mods:date>2020</mods:date>
|
|
<mods:detail type="pubDate">
|
|
<mods:number>2020-09-17</mods:number>
|
|
</mods:detail>
|
|
<mods:detail type="volume">
|
|
<mods:number>42</mods:number>
|
|
</mods:detail>
|
|
<mods:detail type="issue">
|
|
<mods:number>22</mods:number>
|
|
</mods:detail>
|
|
<mods:extent unit="page">
|
|
<mods:start>409</mods:start>
|
|
<mods:end>426</mods:end>
|
|
</mods:extent>
|
|
</mods:part>
|
|
</mods:relatedItem>
|
|
<mods:classification>journal article</mods:classification>
|
|
<mods:identifier type="DOI">10.5252/geodiversitas2020v42a22</mods:identifier>
|
|
<mods:identifier type="GBIF-Dataset">99b2432e-c149-4b82-a584-4b33d116d198</mods:identifier>
|
|
<mods:identifier type="ISSN">1638-9395</mods:identifier>
|
|
<mods:identifier type="Zenodo-Dep">4447648</mods:identifier>
|
|
<mods:identifier type="ZooBank">urn:lsid:zoobank.org:pub:07001ACA-EBDE-4256-BCB9-55E3159F81DC</mods:identifier>
|
|
</mods:mods>
|
|
<treatment ID-DOI="http://doi.org/10.5281/zenodo.4447835" ID-GBIF-Taxon="177202474" ID-Zenodo-Dep="4447835" LSID="urn:lsid:plazi:treatment:CB2D87E12E3AFFCEFE4B4733FD4AF96A" httpUri="http://treatment.plazi.org/id/CB2D87E12E3AFFCEFE4B4733FD4AF96A" lastPageId="13" lastPageNumber="420" pageId="11" pageNumber="418">
|
|
<subSubSection box="[393,514,1587,1613]" pageId="11" pageNumber="418" type="nomenclature">
|
|
<paragraph blockId="11.[393,514,1587,1613]" box="[393,514,1587,1613]" pageId="11" pageNumber="418">
|
|
<heading box="[393,514,1587,1613]" centered="true" fontSize="11" level="2" pageId="11" pageNumber="418" reason="2">
|
|
<taxonomicName authorityName="Merrem" authorityYear="1820" box="[393,477,1587,1612]" class="Reptilia" family="Varanidae" genus="Varanus" kingdom="Animalia" order="Squamata" pageId="11" pageNumber="418" phylum="Chordata" rank="species" species="sp.">
|
|
<emphasis box="[393,477,1587,1612]" italics="true" pageId="11" pageNumber="418">Varanus</emphasis>
|
|
</taxonomicName>
|
|
sp.
|
|
</heading>
|
|
</paragraph>
|
|
</subSubSection>
|
|
<subSubSection pageId="11" pageNumber="418" type="description">
|
|
<paragraph blockId="11.[401,506,1618,1645]" box="[401,506,1618,1645]" pageId="11" pageNumber="418">
|
|
(
|
|
<figureCitation box="[408,497,1618,1645]" captionStart-0="FIG" captionStart-1="FIG" captionStart-2="FIG" captionStartId-0="11.[132,143,1323,1340]" captionStartId-1="12.[132,143,1050,1067]" captionStartId-2="14.[132,143,839,856]" captionTargetBox-0="[399,1199,213,1281]" captionTargetBox-2="[476,1109,218,800]" captionTargetId-0="figure-313@11.[378,1209,201,1291]" captionTargetId-1="figure-527@12.[416,1172,215,1028]" captionTargetId-2="figure-238@14.[472,1115,190,813]" captionTargetPageId-0="11" captionTargetPageId-1="12" captionTargetPageId-2="14" captionText-0="FIG. 4. — Varanidae, Molayan, Varanus sp., trunk vertebra MOL 4126, photos: A anterior view; B, dorsal view; C, lateral view; D, ventral view. Right humerus fragment in dorsal and ventral view, respectively. Scale bar: 5 mm." captionText-1="FIG. 5. — Varanidae, Molayan, late Miocene, Varanus sp., trunk vertebra MOL 4126, drawings: A, anterior view; B, dorsal view; C, lateral view; D, ventral view. Scale bar: 5 mm." captionText-2="FIG. 6. — Xenodermidae (Xenodermus) or Elapidae (Bungarus), Sherullah, late Miocene, photos:A, B, C, trunk vertebra AFG 1672, dorsal,ventral and right lateral views; D, trunk vertebra AFG 1673, dorsal view. Scale bar: 2 mm." figureDoi-0="http://doi.org/10.5281/zenodo.4447659" figureDoi-1="http://doi.org/10.5281/zenodo.4487961" figureDoi-2="http://doi.org/10.5281/zenodo.4447661" httpUri-0="https://zenodo.org/record/4447659/files/figure.png" httpUri-1="https://zenodo.org/record/4487961/files/figure.png" httpUri-2="https://zenodo.org/record/4447661/files/figure.png" pageId="11" pageNumber="418">Figs 4-6</figureCitation>
|
|
)
|
|
</paragraph>
|
|
<paragraph blockId="11.[132,775,1675,1726]" pageId="11" pageNumber="418">LOCALITY AND AGE. — Molayan, Khordkabul basin, late Miocene, middle Turolian in age, MN12.</paragraph>
|
|
</subSubSection>
|
|
<subSubSection pageId="11" pageNumber="418" type="materials_examined">
|
|
<paragraph blockId="11.[131,775,1749,1800]" pageId="11" pageNumber="418">
|
|
MATERIAL EXAMINED. —
|
|
<materialsCitation ID-GBIF-Occurrence="3028142301" pageId="11" pageNumber="418">Three trunk vertebrae (MOL 1853, MOL 1895, MOL 4126). 1 incomplete humerus, MOL 1854.</materialsCitation>
|
|
</paragraph>
|
|
</subSubSection>
|
|
<subSubSection pageId="11" pageNumber="418" type="description">
|
|
<paragraph blockId="11.[131,775,1842,2027]" box="[132,277,1842,1868]" pageId="11" pageNumber="418">DESCRIPTION</paragraph>
|
|
<paragraph blockId="11.[131,775,1842,2027]" box="[132,458,1873,1900]" pageId="11" pageNumber="418">
|
|
<emphasis box="[132,458,1873,1900]" italics="true" pageId="11" pageNumber="418">
|
|
Trunk vertebrae (
|
|
<figureCitation box="[311,422,1874,1900]" captionStart="FIG" captionStartId="11.[132,143,1323,1340]" captionTargetBox="[399,1199,213,1281]" captionTargetId="figure-313@11.[378,1209,201,1291]" captionTargetPageId="11" captionText="FIG. 4. — Varanidae, Molayan, Varanus sp., trunk vertebra MOL 4126, photos: A anterior view; B, dorsal view; C, lateral view; D, ventral view. Right humerus fragment in dorsal and ventral view, respectively. Scale bar: 5 mm." figureDoi="http://doi.org/10.5281/zenodo.4447659" httpUri="https://zenodo.org/record/4447659/files/figure.png" pageId="11" pageNumber="418">Figs 4A, B</figureCitation>
|
|
;
|
|
<figureCitation box="[435,449,1874,1900]" captionStart="FIG" captionStartId="12.[132,143,1050,1067]" captionTargetId="figure-527@12.[416,1172,215,1028]" captionTargetPageId="12" captionText="FIG. 5. — Varanidae, Molayan, late Miocene, Varanus sp., trunk vertebra MOL 4126, drawings: A, anterior view; B, dorsal view; C, lateral view; D, ventral view. Scale bar: 5 mm." figureDoi="http://doi.org/10.5281/zenodo.4487961" httpUri="https://zenodo.org/record/4487961/files/figure.png" pageId="11" pageNumber="418">5</figureCitation>
|
|
)
|
|
</emphasis>
|
|
</paragraph>
|
|
</subSubSection>
|
|
<subSubSection pageId="11" pageNumber="418" type="discussion">
|
|
<paragraph blockId="11.[131,775,1842,2027]" lastBlockId="11.[811,1457,1427,1804]" pageId="11" pageNumber="418">
|
|
They show the morphology that is characteristic of
|
|
<taxonomicName authorityName="Merrem" authorityYear="1820" box="[681,766,1906,1931]" class="Reptilia" family="Varanidae" genus="Varanus" kingdom="Animalia" order="Squamata" pageId="11" pageNumber="418" phylum="Chordata" rank="genus">
|
|
<emphasis box="[681,766,1906,1931]" italics="true" pageId="11" pageNumber="418">Varanus</emphasis>
|
|
</taxonomicName>
|
|
. The neural arch slopes anteriorly. In the anterior half of the vertebra it forms a marked triangular depression on either side of the neural spine. The
|
|
<emphasis box="[456,618,2001,2027]" italics="true" pageId="11" pageNumber="418">pars tectiformis</emphasis>
|
|
that roofs the anteriormost part of the neural canal is a typical feature of
|
|
<taxonomicName authorityName="Merrem" authorityYear="1820" box="[812,897,1460,1485]" class="Reptilia" family="Varanidae" genus="Varanus" kingdom="Animalia" order="Squamata" pageId="11" pageNumber="418" phylum="Chordata" rank="genus">
|
|
<emphasis box="[812,897,1460,1485]" italics="true" pageId="11" pageNumber="418">Varanus</emphasis>
|
|
</taxonomicName>
|
|
, but it is broken off in all specimens from Molayan. The condyle and cotyle are elliptical in outline. Their axis is markedly oblique: the condyle faces posterodorsally whereas the axis of the cotyle is directed anteroventrally (i.e., the cotyle is largely exposed in ventral aspect). The centrum is constricted anterior to the condyle (i.e., precondylar constriction). Apart from these features which are common to all
|
|
<taxonomicName authorityName="Merrem" authorityYear="1820" box="[932,1018,1683,1708]" class="Reptilia" family="Varanidae" genus="Varanus" kingdom="Animalia" order="Squamata" pageId="11" pageNumber="418" phylum="Chordata" rank="genus">
|
|
<emphasis box="[932,1018,1683,1708]" italics="true" pageId="11" pageNumber="418">Varanus</emphasis>
|
|
</taxonomicName>
|
|
species and it is worth mentioning that the vertebrae from Molayan are depressed, the cotyle and condyle are strongly depressed, and the ventral face of the centrum is hardly convex.
|
|
</paragraph>
|
|
</subSubSection>
|
|
<subSubSection lastPageId="12" lastPageNumber="419" pageId="11" pageNumber="418" type="description">
|
|
<paragraph blockId="11.[813,1161,1842,1868]" box="[813,1161,1842,1868]" pageId="11" pageNumber="418">
|
|
<heading box="[813,1161,1842,1868]" fontSize="11" level="2" pageId="11" pageNumber="418" reason="8">
|
|
<emphasis box="[813,1161,1842,1868]" italics="true" pageId="11" pageNumber="418">
|
|
Humerus MOL 1854 (
|
|
<figureCitation box="[1062,1148,1842,1868]" captionStart="FIG" captionStartId="11.[132,143,1323,1340]" captionTargetBox="[399,1199,213,1281]" captionTargetId="figure-313@11.[378,1209,201,1291]" captionTargetPageId="11" captionText="FIG. 4. — Varanidae, Molayan, Varanus sp., trunk vertebra MOL 4126, photos: A anterior view; B, dorsal view; C, lateral view; D, ventral view. Right humerus fragment in dorsal and ventral view, respectively. Scale bar: 5 mm." figureDoi="http://doi.org/10.5281/zenodo.4447659" httpUri="https://zenodo.org/record/4447659/files/figure.png" pageId="11" pageNumber="418">Fig. 4C</figureCitation>
|
|
).
|
|
</emphasis>
|
|
</heading>
|
|
</paragraph>
|
|
<paragraph blockId="11.[811,1457,1873,2028]" pageId="11" pageNumber="418">
|
|
Distal end of a right humerus, the ulnar and radial condyles are lacking. It is tentatively referred to
|
|
<taxonomicName authorityName="Merrem" authorityYear="1820" box="[1260,1346,1906,1931]" class="Reptilia" family="Varanidae" genus="Varanus" kingdom="Animalia" order="Squamata" pageId="11" pageNumber="418" phylum="Chordata" rank="genus">
|
|
<emphasis box="[1260,1346,1906,1931]" italics="true" pageId="11" pageNumber="418">Varanus</emphasis>
|
|
</taxonomicName>
|
|
sp. based on size and varanid features. The diaphysis appears mediodistally enlarged. Laterally, the ectepicondyle is broken but it seems prolonged proximally by a small ascending crest
|
|
</paragraph>
|
|
<paragraph blockId="12.[434,451,240,264]" box="[434,451,240,264]" pageId="12" pageNumber="419">A</paragraph>
|
|
<paragraph pageId="12" pageNumber="419">B D C</paragraph>
|
|
<caption ID-DOI="http://doi.org/10.5281/zenodo.4487961" ID-Zenodo-Dep="4487961" httpUri="https://zenodo.org/record/4487961/files/figure.png" pageId="12" pageNumber="419" targetBox="[416,1172,215,1028]" targetPageId="12">
|
|
<paragraph blockId="12.[132,1457,1050,1089]" pageId="12" pageNumber="419">
|
|
FIG. 5. —
|
|
<taxonomicName authority=", Molayan" authorityName="Molayan" box="[213,381,1050,1067]" class="Reptilia" family="Varanidae" kingdom="Animalia" order="Squamata" pageId="12" pageNumber="419" phylum="Chordata" rank="family">Varanidae, Molayan</taxonomicName>
|
|
, late Miocene,
|
|
<taxonomicName authorityName="Merrem" authorityYear="1820" box="[508,575,1050,1067]" class="Reptilia" family="Varanidae" genus="Varanus" kingdom="Animalia" order="Squamata" pageId="12" pageNumber="419" phylum="Chordata" rank="genus">
|
|
<emphasis box="[508,575,1050,1067]" italics="true" pageId="12" pageNumber="419">Varanus</emphasis>
|
|
</taxonomicName>
|
|
sp., trunk vertebra MOL 4126, drawings:
|
|
<emphasis bold="true" box="[925,938,1050,1067]" pageId="12" pageNumber="419">A</emphasis>
|
|
, anterior view;
|
|
<emphasis bold="true" box="[1067,1080,1050,1067]" pageId="12" pageNumber="419">B</emphasis>
|
|
, dorsal view;
|
|
<emphasis bold="true" box="[1196,1210,1050,1067]" pageId="12" pageNumber="419">C</emphasis>
|
|
, lateral view;
|
|
<emphasis bold="true" box="[1326,1340,1050,1067]" pageId="12" pageNumber="419">D</emphasis>
|
|
, ventral view. Scale bar: 5 mm.
|
|
</paragraph>
|
|
</caption>
|
|
<paragraph blockId="12.[131,776,1140,1772]" pageId="12" pageNumber="419">(ectepicondylar crest). The entepicondyle is projected medially. In ventral view, a large triangular, shallow depression is present, just proximal to the condyles and a small foramen opens on the proximal rim of the depression.</paragraph>
|
|
</subSubSection>
|
|
<subSubSection pageId="12" pageNumber="419" type="discussion">
|
|
<paragraph blockId="12.[131,776,1140,1772]" pageId="12" pageNumber="419">
|
|
The possible presence of an ectepicondylar crest is consistent with referral of this humerus to varanids, as well as the enlarged diaphysis (
|
|
<bibRefCitation author="LECURU S." box="[356,500,1331,1357]" pageId="12" pageNumber="419" pagination="515 - 558" refId="ref15915" refString="LECURU S. 1969. - Etude morphologique de l'humerus des Lacertiliens. Annales des Sciences naturelles, Zoologie, Paris, 12 e serie 11: 515 - 558." type="journal article" year="1969">Lécuru 1969</bibRefCitation>
|
|
). In addition, the large, shallow, triangular depression above the condyles, pierced by a small foramen that opens on the proximal rim of the depression are characteristic of varanid lizards. According to
|
|
<bibRefCitation author="LECURU S." box="[131,293,1459,1485]" pageId="12" pageNumber="419" pagination="515 - 558" refId="ref15915" refString="LECURU S. 1969. - Etude morphologique de l'humerus des Lacertiliens. Annales des Sciences naturelles, Zoologie, Paris, 12 e serie 11: 515 - 558." type="journal article" year="1969">Lécuru (1969)</bibRefCitation>
|
|
and
|
|
<bibRefCitation author="SMITH K. T." box="[349,498,1459,1485]" pageId="12" pageNumber="419" pagination="299 - 358" refId="ref17431" refString="SMITH K. T. 2009. - A new lizard assemblage from the earliest Eocene (Zone WAO) of the Bighorn Basin, Wyoming, USA: Biogeography during the warmest interval of the Cenozoic. Journal of Systematic Palaeontology 7 (3): 299 - 358. https: // doi. org / 10.1017 / S 1477201909002752" type="journal article" year="2009">Smith (2009)</bibRefCitation>
|
|
this foramen is observed in nearly all limbed squamates, but it is quite reduced or even absent in
|
|
<taxonomicName authorityName="Merrem" authorityYear="1820" box="[291,376,1523,1548]" class="Reptilia" family="Varanidae" genus="Varanus" kingdom="Animalia" order="Squamata" pageId="12" pageNumber="419" phylum="Chordata" rank="genus">
|
|
<emphasis box="[291,376,1523,1548]" italics="true" pageId="12" pageNumber="419">Varanus</emphasis>
|
|
</taxonomicName>
|
|
.
|
|
<bibRefCitation author="GEORGALIS G. L. & RAGE J. - C. & BONIS L. DE & KOUFOS G." box="[391,633,1523,1549]" pageId="12" pageNumber="419" pagination="169 - 181" refId="ref14568" refString="GEORGALIS G. L., RAGE J. - C., BONIS L. DE & KOUFOS G. 2018. - Lizards and snakes from the late Miocene hominoid locality of Ravin de la Pluie (Axios Valley, Greece). Swiss Journal of Geosciences 111: 169 - 181. https: // doi. org / 10.1007 / s 00015 - 017 - 0291 - 2" type="journal article" year="2018">
|
|
Georgalis
|
|
<emphasis box="[501,552,1523,1549]" italics="true" pageId="12" pageNumber="419">et al.</emphasis>
|
|
(2018)
|
|
</bibRefCitation>
|
|
described an humerus from the late Miocene of Ravin de la Pluie (
|
|
<collectingCountry box="[684,764,1555,1581]" name="Greece" pageId="12" pageNumber="419">Greece</collectingCountry>
|
|
). The authors state that Miocene European
|
|
<taxonomicName authorityName="Merrem" authorityYear="1820" box="[581,666,1587,1612]" class="Reptilia" family="Varanidae" genus="Varanus" kingdom="Animalia" order="Squamata" pageId="12" pageNumber="419" phylum="Chordata" rank="genus">
|
|
<emphasis box="[581,666,1587,1612]" italics="true" pageId="12" pageNumber="419">Varanus</emphasis>
|
|
</taxonomicName>
|
|
had short and robust limbs. Unfortunately, the humerus studied here is too fragmentry to allow thorough comparisons with the material from
|
|
<collectingCountry box="[287,366,1682,1708]" name="Greece" pageId="12" pageNumber="419">Greece</collectingCountry>
|
|
. However, the diaphysis, as preserved, seems robustly built like the humerus described by
|
|
<bibRefCitation author="GEORGALIS G. L. & RAGE J. - C. & BONIS L. DE & KOUFOS G." pageId="12" pageNumber="419" pagination="169 - 181" refId="ref14568" refString="GEORGALIS G. L., RAGE J. - C., BONIS L. DE & KOUFOS G. 2018. - Lizards and snakes from the late Miocene hominoid locality of Ravin de la Pluie (Axios Valley, Greece). Swiss Journal of Geosciences 111: 169 - 181. https: // doi. org / 10.1007 / s 00015 - 017 - 0291 - 2" type="journal article" year="2018">
|
|
Georgalis
|
|
<emphasis box="[188,241,1746,1772]" italics="true" pageId="12" pageNumber="419">et al.</emphasis>
|
|
(2018)
|
|
</bibRefCitation>
|
|
.
|
|
</paragraph>
|
|
<paragraph blockId="12.[131,776,1810,2028]" box="[132,262,1810,1836]" pageId="12" pageNumber="419">COMMENTS</paragraph>
|
|
<paragraph blockId="12.[131,776,1810,2028]" pageId="12" pageNumber="419">
|
|
The referral of these specimens to the genus
|
|
<taxonomicName authorityName="Merrem" authorityYear="1820" box="[601,686,1842,1867]" class="Reptilia" family="Varanidae" genus="Varanus" kingdom="Animalia" order="Squamata" pageId="12" pageNumber="419" phylum="Chordata" rank="genus">
|
|
<emphasis box="[601,686,1842,1867]" italics="true" pageId="12" pageNumber="419">Varanus</emphasis>
|
|
</taxonomicName>
|
|
appears to be unquestionable (at least for the vertebrae) but assignment at species level would require comparisons with all living species, which is not possible. Here, preliminary comparisons are made only with extinct species the vertebrae of which are known.
|
|
</paragraph>
|
|
</subSubSection>
|
|
<subSubSection pageId="12" pageNumber="419" type="reference_group">
|
|
<paragraph blockId="12.[811,1460,1140,2028]" pageId="12" pageNumber="419">
|
|
The oldest fossils attributed to
|
|
<taxonomicName authorityName="Merrem" authorityYear="1820" box="[1159,1243,1141,1166]" class="Reptilia" family="Varanidae" genus="Varanus" kingdom="Animalia" order="Squamata" pageId="12" pageNumber="419" phylum="Chordata" rank="genus">
|
|
<emphasis box="[1159,1243,1141,1166]" italics="true" pageId="12" pageNumber="419">Varanus</emphasis>
|
|
</taxonomicName>
|
|
consist of vertebrae from the late Eocene (Priabonian) and the early Oligocene of
|
|
<collectingCountry box="[842,907,1204,1230]" name="Egypt" pageId="12" pageNumber="419">Egypt</collectingCountry>
|
|
(
|
|
<bibRefCitation author="HOLMES R. B. & MURRAY A. M. & YOUSRY S. & ATTIA Y. S. & SIMONS E. L. & CHATRATH P." box="[926,1141,1204,1230]" pageId="12" pageNumber="419" pagination="1099 - 1110" refId="ref15229" refString="HOLMES R. B., MURRAY A. M., YOUSRY S., ATTIA Y. S., SIMONS E. L. & CHATRATH P. 2010. - Oldest known Varanus (Squamata: Varanidae) from the upper Eocene and lower Oligocene of Egypt: Support for an African origin of the genus. Palaeontology 53 (5): 1099 - 1110. https: // doi. org / 10.1111 / j. 1475 - 4983.2010.00994. x" type="journal article" year="2010">
|
|
Holmes
|
|
<emphasis box="[1022,1076,1204,1230]" italics="true" pageId="12" pageNumber="419">et al.</emphasis>
|
|
2010
|
|
</bibRefCitation>
|
|
;
|
|
<bibRefCitation author="SMITH K. T. & BHULLAR B. A. & HOLROYD P. A." box="[1157,1352,1204,1230]" pageId="12" pageNumber="419" pagination="909 - 913" refId="ref17493" refString="SMITH K. T., BHULLAR B. A. & HOLROYD P. A. 2008. - Earliest African Record of the Varanus stem clade (Squamata: Varanidae) from the Early Oligocene of Egypt. Journal of Vertebrate Paleontology 28 (3): 909 - 913. https: // doi. org / dhs 5 g 4" type="journal article" year="2008">
|
|
Smith
|
|
<emphasis box="[1232,1287,1204,1230]" italics="true" pageId="12" pageNumber="419">et al.</emphasis>
|
|
2008
|
|
</bibRefCitation>
|
|
;
|
|
<bibRefCitation author="AUGE M. L. & GUEVEL B." pageId="12" pageNumber="419" pagination="1410483" refId="ref12689" refString="AUGE M. L. & GUEVEL B. 2018. - New varanid remains from the Miocene (MN 4 - MN 5) of France: inferring fossil lizard phylogeny from subsets of large morphological data sets. Journal of Vertebrate Paleontology 38 (1): e 1410483. https: // doi. org / 10.10 80 / 02724634.2017.1410483" type="journal article" year="2018">Augé & Guével 2018</bibRefCitation>
|
|
).
|
|
</paragraph>
|
|
<paragraph blockId="12.[811,1460,1140,2028]" pageId="12" pageNumber="419">
|
|
Earliest Miocene
|
|
<taxonomicName authorityName="Merrem" authorityYear="1820" box="[1034,1120,1268,1293]" class="Reptilia" family="Varanidae" genus="Varanus" kingdom="Animalia" order="Squamata" pageId="12" pageNumber="419" phylum="Chordata" rank="genus">
|
|
<emphasis box="[1034,1120,1268,1293]" italics="true" pageId="12" pageNumber="419">Varanus</emphasis>
|
|
</taxonomicName>
|
|
material come from the early Miocene (Orleanian, i.e. Burdigalian) of Africa and Europe (
|
|
<bibRefCitation author="CLOS L. M." box="[822,937,1331,1358]" pageId="12" pageNumber="419" pagination="254 - 267" refId="ref13554" refString="CLOS L. M. 1995. - A new species of Varanus (Reptilia: Sauria) from the Miocene of Kenya. Journal of Vertebrate Paleontology 15: 254 - 267. https: // doi. org / 10.1080 / 02724634.1995.10011228" type="journal article" year="1995">Clos 1995</bibRefCitation>
|
|
; Hoffstetter 1969,
|
|
<bibRefCitation author="IVANOV M. & RUTA M. & KLEMBARA J. & BOHME M." box="[1154,1354,1331,1358]" pageId="12" pageNumber="419" pagination="767 - 797" refId="ref15316" refString="IVANOV M., RUTA M., KLEMBARA J. & BOHME M. 2018. - A new species of Varanus (Anguimorpha: Varanidae) from the early Miocene of the Czech Republic, and its relationships and palaeoecology. Journal of Systematic Palaeontology 16: 767 - 797. https: // doi. org / 1 0.1080 / 14772019.2017.1355338" type="journal article" year="2018">
|
|
Ivanov
|
|
<emphasis box="[1236,1290,1331,1357]" italics="true" pageId="12" pageNumber="419">et al.</emphasis>
|
|
2018
|
|
</bibRefCitation>
|
|
).
|
|
</paragraph>
|
|
</subSubSection>
|
|
<subSubSection lastPageId="13" lastPageNumber="420" pageId="12" pageNumber="419" type="discussion">
|
|
<paragraph blockId="12.[811,1460,1140,2028]" lastBlockId="13.[130,777,215,1709]" lastPageId="13" lastPageNumber="420" pageId="12" pageNumber="419">
|
|
Many varanid vertebrae have been described and several species have also been named but now serious doubts are raised about the possibility of alpha taxonomy based on isolated vertebrae.
|
|
<taxonomicName authority="Clos, 1995" authorityName="Clos" authorityYear="1995" box="[1018,1353,1459,1486]" class="Reptilia" family="Varanidae" genus="Varanus" kingdom="Animalia" order="Squamata" pageId="12" pageNumber="419" phylum="Chordata" rank="species" species="Rusingensis">
|
|
<emphasis box="[1018,1225,1460,1485]" italics="true" pageId="12" pageNumber="419">Varanus rusingensis</emphasis>
|
|
<bibRefCitation author="CLOS L. M." box="[1232,1353,1459,1486]" pageId="12" pageNumber="419" pagination="254 - 267" refId="ref13554" refString="CLOS L. M. 1995. - A new species of Varanus (Reptilia: Sauria) from the Miocene of Kenya. Journal of Vertebrate Paleontology 15: 254 - 267. https: // doi. org / 10.1080 / 02724634.1995.10011228" type="journal article" year="1995">Clos, 1995</bibRefCitation>
|
|
</taxonomicName>
|
|
from the Orleanian of
|
|
<collectingCountry box="[957,1026,1491,1517]" name="Kenya" pageId="12" pageNumber="419">Kenya</collectingCountry>
|
|
(
|
|
<bibRefCitation author="CLOS L. M." box="[1042,1155,1491,1518]" pageId="12" pageNumber="419" pagination="254 - 267" refId="ref13554" refString="CLOS L. M. 1995. - A new species of Varanus (Reptilia: Sauria) from the Miocene of Kenya. Journal of Vertebrate Paleontology 15: 254 - 267. https: // doi. org / 10.1080 / 02724634.1995.10011228" type="journal article" year="1995">Clos 1995</bibRefCitation>
|
|
),
|
|
<taxonomicName authority="Roger, 1898" authorityName="Roger" authorityYear="1898" box="[1178,1450,1491,1517]" class="Reptilia" family="Varanidae" genus="Varanus" kingdom="Animalia" order="Squamata" pageId="12" pageNumber="419" phylum="Chordata" rank="species" species="Hofmanni">
|
|
<emphasis box="[1178,1308,1491,1517]" italics="true" pageId="12" pageNumber="419">V. hofmanni</emphasis>
|
|
Roger, 1898
|
|
</taxonomicName>
|
|
, from the early (Orleanian), middle (Astaracian) and perhaps late (Vallesian) of western Europe (
|
|
<bibRefCitation author="ESTES R." box="[1188,1305,1555,1581]" pageId="12" pageNumber="419" refId="ref13909" refString="ESTES R. 1983. - Sauriaterrestria, Amphisbaenia, in WELLNHOFER P. (ed.), Handbuch der Palaoherpetologie. Volume 10 A. Gustav Fischer, Stuttgart, xxii + 249 p." type="book" year="1983">Estes 1983</bibRefCitation>
|
|
),
|
|
<taxonomicName authority="Ivanov, Ivanov, Ruta, Klembara & Bohme, 2018" authorityName="Ivanov, Ivanov, Ruta, Klembara & Bohme" authorityYear="2018" class="Reptilia" family="Varanidae" genus="Varanus" kingdom="Animalia" order="Squamata" pageId="12" pageNumber="419" phylum="Chordata" rank="species" species="Mokrensis">
|
|
<emphasis box="[1327,1455,1554,1580]" italics="true" pageId="12" pageNumber="419">V. mokrensis</emphasis>
|
|
<bibRefCitation author="IVANOV M. & RUTA M. & KLEMBARA J. & BOHME M." box="[813,1348,1586,1613]" pageId="12" pageNumber="419" pagination="767 - 797" refId="ref15316" refString="IVANOV M., RUTA M., KLEMBARA J. & BOHME M. 2018. - A new species of Varanus (Anguimorpha: Varanidae) from the early Miocene of the Czech Republic, and its relationships and palaeoecology. Journal of Systematic Palaeontology 16: 767 - 797. https: // doi. org / 1 0.1080 / 14772019.2017.1355338" type="journal article" year="2018">Ivanov, Ivanov, Ruta, Klembara & Böhme, 2018</bibRefCitation>
|
|
</taxonomicName>
|
|
, from the early Miocene (Orleanian, MN4) of
|
|
<collectingCountry box="[1236,1418,1618,1644]" name="Czech Republic" pageId="12" pageNumber="419">Czech Republic</collectingCountry>
|
|
,
|
|
<taxonomicName authority="Zerova & Ckhikvadze, 1986" authorityName="Zerova & Ckhikvadze" authorityYear="1986" class="Reptilia" family="Varanidae" genus="Varanus" kingdom="Animalia" order="Squamata" pageId="12" pageNumber="419" phylum="Chordata" rank="species" species="Pronini">
|
|
<emphasis italics="true" pageId="12" pageNumber="419">V. pronini</emphasis>
|
|
Zerova & Čkhikvadze, 1986
|
|
</taxonomicName>
|
|
(from middle Miocene of
|
|
<collectingCountry box="[839,969,1682,1709]" name="Kazakhstan" pageId="12" pageNumber="419">Kazakhstan</collectingCountry>
|
|
),
|
|
<taxonomicName authority="Zerova & Ckhikvadze, 1983 in Lungu, Zerova & Ckhikvadze, 1983" authorityName="Zerova & Ckhikvadze, 1983 in Lungu, Zerova & Ckhikvadze" authorityYear="1983" class="Reptilia" family="Varanidae" genus="Varanus" kingdom="Animalia" order="Squamata" pageId="12" pageNumber="419" phylum="Chordata" rank="species" species="Tyrasiensis">
|
|
<emphasis box="[985,1114,1683,1708]" italics="true" pageId="12" pageNumber="419">V. tyrasiensis</emphasis>
|
|
Zerova & Čkhikvadze,
|
|
<quantity box="[1371,1454,1682,1708]" metricMagnitude="1" metricUnit="m" metricValue="5.03682" pageId="12" pageNumber="419" unit="in" value="1983.0">
|
|
1983
|
|
<emphasis box="[1432,1454,1683,1708]" italics="true" pageId="12" pageNumber="419">in</emphasis>
|
|
</quantity>
|
|
Lungu, Zerova & Čkhikvadze, 1983
|
|
</taxonomicName>
|
|
, and
|
|
<taxonomicName authority="Zerova & Ckhikvadze, 1983" authorityName="Zerova & Ckhikvadze" authorityYear="1983" class="Reptilia" family="Varanidae" genus="Varanus" kingdom="Animalia" order="Squamata" pageId="12" pageNumber="419" phylum="Chordata" rank="species" species="Lungui">
|
|
<emphasis box="[1253,1346,1714,1740]" italics="true" pageId="12" pageNumber="419">V. lungui</emphasis>
|
|
Zerova & Čkhikvadze, 1983
|
|
</taxonomicName>
|
|
from the late Miocene (Sarmatian) of
|
|
<collectingCountry box="[815,913,1778,1804]" name="Moldova" pageId="12" pageNumber="419">Moldova</collectingCountry>
|
|
(Zerova & Čkhikvadze 1983, 1986),
|
|
<taxonomicName authority="Zerova & Ckhikvadze, 1986" authorityName="Zerova & Ckhikvadze" authorityYear="1986" class="Reptilia" family="Varanidae" genus="Varanus" kingdom="Animalia" order="Squamata" pageId="12" pageNumber="419" phylum="Chordata" rank="species" species="Semjonovi">
|
|
<emphasis box="[1324,1455,1779,1804]" italics="true" pageId="12" pageNumber="419">V. semjonovi</emphasis>
|
|
Zerova & Čkhikvadze, 1986
|
|
</taxonomicName>
|
|
from the late Miocene (Meotian) of
|
|
<collectingCountry box="[841,934,1841,1867]" name="Ukraine" pageId="12" pageNumber="419">Ukraine</collectingCountry>
|
|
,
|
|
<taxonomicName authority="Conrad, Balcarcel & Mehling, 2012" authorityName="Conrad, Balcarcel & Mehling" authorityYear="2012" class="Reptilia" family="Varanidae" genus="Varanus" kingdom="Animalia" order="Squamata" pageId="12" pageNumber="419" phylum="Chordata" rank="species" species="Amnhophilis">
|
|
<emphasis box="[949,1109,1841,1867]" italics="true" pageId="12" pageNumber="419">V. amnhophilis</emphasis>
|
|
<bibRefCitation author="CONRAD J. L. & BALCARCEL A. M. & MEHLING C. M." pageId="12" pageNumber="419" pagination="41767" refId="ref13598" refString="CONRAD J. L., BALCARCEL A. M. & MEHLING C. M. 2012. - Earliest example of a Giant Monitor Lizard (Varanus, Varanidae, Squamata). PLOS One 7 (8): e 41767. https: // doi. org / 10.1371 / journal. pone. 0041767" type="journal article" year="2012">Conrad, Balcarcel & Mehling, 2012</bibRefCitation>
|
|
</taxonomicName>
|
|
from the late Miocene (Turolian) of Samos,
|
|
<collectingCountry box="[1372,1451,1874,1900]" name="Greece" pageId="12" pageNumber="419">Greece</collectingCountry>
|
|
,
|
|
<taxonomicName authority="Weithofer, 1888" authorityName="Weithofer" authorityYear="1888" box="[813,1174,1905,1931]" class="Reptilia" family="Varanidae" genus="Varanus" kingdom="Animalia" order="Squamata" pageId="12" pageNumber="419" phylum="Chordata" rank="species" species="Marathonensis">
|
|
<emphasis box="[813,990,1905,1931]" italics="true" pageId="12" pageNumber="419">V. marathonensis</emphasis>
|
|
Weithofer, 1888
|
|
</taxonomicName>
|
|
from the middle (Gaudry 1862-1867;
|
|
<bibRefCitation author="VILLA A. & ABELLA J. & ALBA D. M. & ALMECIJA S. & BOLET A. & KOUFOS G. D. & KNOLL F. & LUJAN A. H. & MORALES J. & ROBLES J. M. & SANCHEZ I. M. & DELFINO M." box="[952,1132,1937,1963]" pageId="12" pageNumber="419" pagination="0207719" refId="ref17825" refString="VILLA A., ABELLA J., ALBA D. M., ALMECIJA S., BOLET A., KOUFOS G. D., KNOLL F., LUJAN A. H., MORALES J., ROBLES J. M., SANCHEZ I. M. & DELFINO M. 2018. - Revision of Varanus marathonensis (Squamata, Varanidae) based on historogical and new material: morphology, systematics, and paleobiogeography of the European monitor lizards. PLoS ONE 13 (12): e 0207719. https: // doi. org / 10.1371 / journal. pone. 0207719" type="journal article" year="2018">
|
|
Villa
|
|
<emphasis box="[1013,1067,1937,1963]" italics="true" pageId="12" pageNumber="419">et al.</emphasis>
|
|
2018
|
|
</bibRefCitation>
|
|
) to late Miocene, early Pliocene (Ruscinian), and from the early Pliocene (Ruscinian) of
|
|
<collectingCountry box="[844,938,2001,2027]" name="Turkey" pageId="12" pageNumber="419">Anatolia</collectingCountry>
|
|
(
|
|
<bibRefCitation author="RAGE J. - C. & SEN S." box="[956,1157,2001,2028]" pageId="12" pageNumber="419" pagination="127 - 134" refId="ref16649" refString="RAGE J. - C. & SEN S. 1976. - Les Amphibiens et les Reptiles du Pliocene superieur de Calta (Turquie). Geologie mediterraneenne 3: 127 - 134. https: // doi. org / 10.3406 / geolm. 1976.969" type="journal article" year="1976">Rage & Sen 1976</bibRefCitation>
|
|
), and
|
|
<taxonomicName authority="Fejervary, 1935" authorityName="Fejervary" authorityYear="1935" class="Reptilia" family="Varanidae" genus="Varanus" kingdom="Animalia" lastPageId="13" lastPageNumber="420" order="Squamata" pageId="12" pageNumber="419" phylum="Chordata" rank="species" species="Bolkayi">
|
|
<emphasis box="[1234,1342,2001,2027]" italics="true" pageId="12" pageNumber="419">V. bolkayi</emphasis>
|
|
<bibRefCitation author="FEJERVARY G." lastPageId="13" lastPageNumber="420" pageId="12" pageNumber="419" pagination="1 - 130" refId="ref14027" refString="FEJERVARY G. 1935. - Further contributions to a monograph of the Megalanidae and fossil Varanidae - with notes on recent varanians. Annales Musei Historico-Naturalis Hungarici 29: 1 - 130." type="journal article" year="1935">Fejérváry, 1935</bibRefCitation>
|
|
</taxonomicName>
|
|
from the Pleistocene of
|
|
<collectingCountry box="[471,583,215,241]" name="Indonesia" pageId="13" pageNumber="420">Indonesia</collectingCountry>
|
|
. Fossils from the middle Pleistocene of
|
|
<collectingCountry box="[383,459,247,273]" name="Greece" pageId="13" pageNumber="420">Greece</collectingCountry>
|
|
represent the last occurence of varanid lizards in Europe (
|
|
<bibRefCitation author="GEORGALIS G. L. & VILLA A. & DELFINO M." box="[458,688,279,305]" pageId="13" pageNumber="420" pagination="1301946" refId="ref14640" refString="GEORGALIS G. L., VILLA A. & DELFINO M. 2017. - The last European varanid: demise and extinction of monitor lizards (Squamata, Varanidae) from Europe. Journal of Vertebrate Paleontology 37 (2): e 1301946. https: // doi. org / 10.1080 / 0272463 4.2017. 1301946" type="journal article" year="2017">
|
|
Georgalis
|
|
<emphasis box="[571,624,279,305]" italics="true" pageId="13" pageNumber="420">et al.</emphasis>
|
|
2017
|
|
</bibRefCitation>
|
|
).
|
|
</paragraph>
|
|
<paragraph blockId="13.[130,777,215,1709]" pageId="13" pageNumber="420">
|
|
In addition, in Asia, rare vertebrae belonging to
|
|
<taxonomicName authorityName="Merrem" authorityYear="1820" box="[693,777,312,337]" class="Reptilia" family="Varanidae" genus="Varanus" kingdom="Animalia" order="Squamata" pageId="13" pageNumber="420" phylum="Chordata" rank="genus">
|
|
<emphasis box="[693,777,312,337]" italics="true" pageId="13" pageNumber="420">Varanus</emphasis>
|
|
</taxonomicName>
|
|
have been recovered from the Siwalik beds; one of these vertebrae, of presumed Pliocene age, was referred to the extinct species
|
|
<taxonomicName authority="Falconer, 1868" authorityName="Falconer" authorityYear="1868" box="[209,485,406,432]" class="Reptilia" family="Varanidae" genus="Varanus" kingdom="Animalia" order="Squamata" pageId="13" pageNumber="420" phylum="Chordata" rank="species" species="Sivalensis">
|
|
<emphasis box="[209,326,406,432]" italics="true" pageId="13" pageNumber="420">V. sivalensis</emphasis>
|
|
<bibRefCitation author="FALCONER H." box="[329,485,406,432]" pageId="13" pageNumber="420" refId="ref13986" refString="FALCONER H. 1868. - Palaeontological Memoirs and Notes of the late Hugh Falconer. 2 Vols. AM, MD, London, R. Hardwicke. https: // www. biodiversitylibrary. org / bibliography / 49217" type="url" year="1868">Falconer, 1868</bibRefCitation>
|
|
</taxonomicName>
|
|
that is based on a humerus. The assignment of this vertebra to
|
|
<taxonomicName authorityName="Falconer" authorityYear="1868" box="[493,611,438,464]" class="Reptilia" family="Varanidae" genus="Varanus" kingdom="Animalia" order="Squamata" pageId="13" pageNumber="420" phylum="Chordata" rank="species" species="Sivalensis">
|
|
<emphasis box="[493,611,438,464]" italics="true" pageId="13" pageNumber="420">V. sivalensis</emphasis>
|
|
</taxonomicName>
|
|
is not justified. Another vertebra, from the late Miocene of Siwalik deposits has been referred to as
|
|
<taxonomicName authorityName="Merrem" authorityYear="1820" box="[326,409,503,528]" class="Reptilia" family="Varanidae" genus="Varanus" kingdom="Animalia" order="Squamata" pageId="13" pageNumber="420" phylum="Chordata" rank="genus">
|
|
<emphasis box="[326,409,503,528]" italics="true" pageId="13" pageNumber="420">Varanus</emphasis>
|
|
</taxonomicName>
|
|
sp. (
|
|
<bibRefCitation author="RAGE J. C. & GUPTA S. S. & PRASAD G. V. R." box="[460,628,502,529]" pageId="13" pageNumber="420" pagination="197 - 205" refId="ref16860" refString="RAGE J. C., GUPTA S. S. & PRASAD G. V. R. 2001. - Amphibians and Squamates from the Neogene Siwalik beds of Jammu and Kashmir, India. Palaontologische Zeitschrift 75 (2): 197 - 205. https: // doi. org / 10.1007 / BF 02988013" type="journal article" year="2001">
|
|
Rage
|
|
<emphasis box="[518,567,502,528]" italics="true" pageId="13" pageNumber="420">et al.</emphasis>
|
|
2001
|
|
</bibRefCitation>
|
|
). In addition,
|
|
<bibRefCitation author="CERNANSKY A. & SYROMYATNIKOVA E. V. & JABLONSKI D." box="[130,394,534,564]" pageId="13" pageNumber="420" pagination="869 - 879" refId="ref13304" refString="CERNANSKY A., SYROMYATNIKOVA E. V. & JABLONSKI D. 2018. - The first record of amphisbaenian and anguimorph lizards (Reptilia, Squamata) from the upper Miocene Solnechnodolsk locality in Russia. Historical Biology 32 (7): 869 - 879. https: // doi. org / 10.1 080 / 08912963.2018.1539973" type="journal article" year="2018">
|
|
Čerňanský
|
|
<emphasis box="[257,311,534,560]" italics="true" pageId="13" pageNumber="420">et al.</emphasis>
|
|
(2018)
|
|
</bibRefCitation>
|
|
described vertebrae of
|
|
<taxonomicName authorityName="Merrem" authorityYear="1820" box="[653,737,535,560]" class="Reptilia" family="Varanidae" genus="Varanus" kingdom="Animalia" order="Squamata" pageId="13" pageNumber="420" phylum="Chordata" rank="genus">
|
|
<emphasis box="[653,737,535,560]" italics="true" pageId="13" pageNumber="420">Varanus</emphasis>
|
|
</taxonomicName>
|
|
sp. from the late Miocene of Solnechnodolsk (
|
|
<collectingCountry box="[592,667,566,593]" name="Russia" pageId="13" pageNumber="420">Russia</collectingCountry>
|
|
).
|
|
</paragraph>
|
|
<paragraph blockId="13.[130,777,215,1709]" pageId="13" pageNumber="420">
|
|
<taxonomicName authority="(Hoffstetter, 1969)" baseAuthorityName="Hoffstetter" baseAuthorityYear="1969" box="[155,641,598,625]" class="Reptilia" family="Varanidae" genus="Iberovaranus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Squamata" pageId="13" pageNumber="420" phylum="Chordata" rank="species" species="Catalaunicus">
|
|
<emphasis box="[155,428,598,624]" italics="true" pageId="13" pageNumber="420">Iberovaranus catalaunicus</emphasis>
|
|
(Hoffstetter, 1969)
|
|
</taxonomicName>
|
|
comes from the early Miocene (MN3/MN4) of
|
|
<collectingCountry box="[526,591,630,656]" name="Spain" pageId="13" pageNumber="420">Spain</collectingCountry>
|
|
; it has been synonymized with
|
|
<taxonomicName authorityName="Merrem" authorityYear="1820" box="[312,398,662,687]" class="Reptilia" family="Varanidae" genus="Varanus" kingdom="Animalia" order="Squamata" pageId="13" pageNumber="420" phylum="Chordata" rank="genus">
|
|
<emphasis box="[312,398,662,687]" italics="true" pageId="13" pageNumber="420">Varanus</emphasis>
|
|
</taxonomicName>
|
|
and the species
|
|
<taxonomicName baseAuthorityName="Hoffstetter" baseAuthorityYear="1969" box="[587,750,662,688]" class="Reptilia" family="Varanidae" genus="Iberovaranus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Squamata" pageId="13" pageNumber="420" phylum="Chordata" rank="species" species="Catalaunicus">
|
|
<emphasis box="[587,750,662,688]" italics="true" pageId="13" pageNumber="420">V. catalaunicus</emphasis>
|
|
</taxonomicName>
|
|
is now regarded as a nomen dubium (
|
|
<bibRefCitation author="DELFINO M. & RAGE J. - C. & BOLET A. & ALBA D. M." box="[541,758,693,720]" pageId="13" pageNumber="420" pagination="731 - 735" refId="ref13845" refString="DELFINO M., RAGE J. - C., BOLET A. & ALBA D. M. 2013. - Early Miocene dispersal of the lizard Varanus into Europe: Reassessment of vertebral material from Spain. Acta Palaeontologica Polonica 58 (4): 731 - 735. https: // doi. org / 10.4202 / app. 2012.0025" type="journal article" year="2013">
|
|
Delfino
|
|
<emphasis box="[636,691,693,719]" italics="true" pageId="13" pageNumber="420">et al.</emphasis>
|
|
2013
|
|
</bibRefCitation>
|
|
). (See a revision of the fossil record of the genus
|
|
<taxonomicName authority="in Molnar (2004))" authorityName="in Molnar" authorityYear="2004" class="Reptilia" family="Varanidae" genus="Varanus" kingdom="Animalia" order="Squamata" pageId="13" pageNumber="420" phylum="Chordata" rank="genus">
|
|
<emphasis box="[659,745,726,751]" italics="true" pageId="13" pageNumber="420">Varanus</emphasis>
|
|
in
|
|
<bibRefCitation author="MOLNAR R. E." box="[133,312,757,784]" pageId="13" pageNumber="420" pagination="10 - 67" refId="ref16191" refString="MOLNAR R. E. 2004. - The long and honorable History of Monitors and their Kins, in PIANKA E. R. & KING D. R. (eds), Varanoid Lizards of theWorld. Indiana University Press, Bloomington: 10 - 67." type="book chapter" year="2004">Molnar (2004))</bibRefCitation>
|
|
</taxonomicName>
|
|
.
|
|
</paragraph>
|
|
<paragraph blockId="13.[130,777,215,1709]" pageId="13" pageNumber="420">
|
|
<bibRefCitation author="IVANOV M. & RUTA M. & KLEMBARA J. & BOHME M." box="[155,391,789,816]" pageId="13" pageNumber="420" pagination="767 - 797" refId="ref15316" refString="IVANOV M., RUTA M., KLEMBARA J. & BOHME M. 2018. - A new species of Varanus (Anguimorpha: Varanidae) from the early Miocene of the Czech Republic, and its relationships and palaeoecology. Journal of Systematic Palaeontology 16: 767 - 797. https: // doi. org / 1 0.1080 / 14772019.2017.1355338" type="journal article" year="2018">
|
|
Ivanov
|
|
<emphasis box="[243,302,789,815]" italics="true" pageId="13" pageNumber="420">et al.</emphasis>
|
|
(2018)
|
|
</bibRefCitation>
|
|
retained only four valid species (
|
|
<taxonomicName authorityName="Clos" authorityYear="1995" box="[142,282,822,847]" class="Reptilia" family="Varanidae" genus="Varanus" kingdom="Animalia" order="Squamata" pageId="13" pageNumber="420" phylum="Chordata" rank="species" species="Rusingensis">
|
|
<emphasis box="[142,282,822,847]" italics="true" pageId="13" pageNumber="420">V. rusingensis</emphasis>
|
|
</taxonomicName>
|
|
,
|
|
<emphasis box="[294,597,821,847]" italics="true" pageId="13" pageNumber="420">
|
|
<taxonomicName authorityName="Roger" authorityYear="1898" box="[294,429,821,847]" class="Reptilia" family="Varanidae" genus="Varanus" kingdom="Animalia" order="Squamata" pageId="13" pageNumber="420" phylum="Chordata" rank="species" species="Hofmanni">V. hofmanni</taxonomicName>
|
|
,
|
|
<taxonomicName authorityName="Conrad, Balcarcel & Mehling" authorityYear="2012" box="[438,597,821,847]" class="Reptilia" family="Varanidae" genus="Varanus" kingdom="Animalia" order="Squamata" pageId="13" pageNumber="420" phylum="Chordata" rank="species" species="Amnhophilis">V. amnhophilis</taxonomicName>
|
|
</emphasis>
|
|
and
|
|
<taxonomicName authorityName="Weithofer" authorityYear="1888" class="Reptilia" family="Varanidae" genus="Varanus" kingdom="Animalia" order="Squamata" pageId="13" pageNumber="420" phylum="Chordata" rank="species" species="Marathonensis">
|
|
<emphasis italics="true" pageId="13" pageNumber="420">V. marathonensis</emphasis>
|
|
</taxonomicName>
|
|
) in addition to the new species
|
|
<taxonomicName authorityName="Ivanov, Ivanov, Ruta, Klembara & Bohme" authorityYear="2018" box="[538,669,853,879]" class="Reptilia" family="Varanidae" genus="Varanus" kingdom="Animalia" order="Squamata" pageId="13" pageNumber="420" phylum="Chordata" rank="species" species="Mokrensis">
|
|
<emphasis box="[538,669,853,879]" italics="true" pageId="13" pageNumber="420">V. mokrensis</emphasis>
|
|
</taxonomicName>
|
|
and
|
|
<bibRefCitation author="VILLA A. & ABELLA J. & ALBA D. M. & ALMECIJA S. & BOLET A. & KOUFOS G. D. & KNOLL F. & LUJAN A. H. & MORALES J. & ROBLES J. M. & SANCHEZ I. M. & DELFINO M." pageId="13" pageNumber="420" pagination="0207719" refId="ref17825" refString="VILLA A., ABELLA J., ALBA D. M., ALMECIJA S., BOLET A., KOUFOS G. D., KNOLL F., LUJAN A. H., MORALES J., ROBLES J. M., SANCHEZ I. M. & DELFINO M. 2018. - Revision of Varanus marathonensis (Squamata, Varanidae) based on historogical and new material: morphology, systematics, and paleobiogeography of the European monitor lizards. PLoS ONE 13 (12): e 0207719. https: // doi. org / 10.1371 / journal. pone. 0207719" type="journal article" year="2018">
|
|
Villa
|
|
<emphasis box="[132,187,885,911]" italics="true" pageId="13" pageNumber="420">et al.</emphasis>
|
|
(2018)
|
|
</bibRefCitation>
|
|
suggest that
|
|
<taxonomicName authorityName="Conrad, Balcarcel & Mehling" authorityYear="2012" box="[426,588,885,911]" class="Reptilia" family="Varanidae" genus="Varanus" kingdom="Animalia" order="Squamata" pageId="13" pageNumber="420" phylum="Chordata" rank="species" species="Amnhophilis">
|
|
<emphasis box="[426,588,885,911]" italics="true" pageId="13" pageNumber="420">V. amnhophilis</emphasis>
|
|
</taxonomicName>
|
|
is a synonym of
|
|
<taxonomicName authorityName="Weithofer" authorityYear="1888" box="[134,314,917,943]" class="Reptilia" family="Varanidae" genus="Varanus" kingdom="Animalia" order="Squamata" pageId="13" pageNumber="420" phylum="Chordata" rank="species" species="Marathonensis">
|
|
<emphasis box="[134,314,917,943]" italics="true" pageId="13" pageNumber="420">V. marathonensis</emphasis>
|
|
</taxonomicName>
|
|
. Many nominal fossil varanid species are based on insufficient material (isolated vertebrae) and are regarded as
|
|
<emphasis box="[261,412,980,1006]" italics="true" pageId="13" pageNumber="420">nomina dubia</emphasis>
|
|
by several authors and the status of these taxa awaits a thorough revision.
|
|
</paragraph>
|
|
</subSubSection>
|
|
<subSubSection pageId="13" pageNumber="420" type="description">
|
|
<paragraph blockId="13.[130,777,215,1709]" pageId="13" pageNumber="420">
|
|
As recognized by previous authors, isolated vertebrae are poorly diagnostic, at least at the species rank, their identification is hindered by poor knowledge of their variability (intra or inter specific). (See discussion in
|
|
<bibRefCitation author="GEORGALIS G. L. & RAGE J. - C. & BONIS L. DE & KOUFOS G." box="[543,775,1140,1166]" pageId="13" pageNumber="420" pagination="169 - 181" refId="ref14568" refString="GEORGALIS G. L., RAGE J. - C., BONIS L. DE & KOUFOS G. 2018. - Lizards and snakes from the late Miocene hominoid locality of Ravin de la Pluie (Axios Valley, Greece). Swiss Journal of Geosciences 111: 169 - 181. https: // doi. org / 10.1007 / s 00015 - 017 - 0291 - 2" type="journal article" year="2018">
|
|
Georgalis
|
|
<emphasis box="[654,709,1140,1166]" italics="true" pageId="13" pageNumber="420">et al.</emphasis>
|
|
2018
|
|
</bibRefCitation>
|
|
about the problematic status of vertebral based varanid taxa).
|
|
</paragraph>
|
|
</subSubSection>
|
|
<subSubSection pageId="13" pageNumber="420" type="discussion">
|
|
<paragraph blockId="13.[130,777,215,1709]" pageId="13" pageNumber="420">
|
|
The strongly depressed condyle and cotyle represent the main distinctive feature of the vertebrae from Molayan. Cursory comparison shows that the Afghan fossil is clearly distinguished from the purported
|
|
<taxonomicName authorityName="in Molnar" authorityYear="2004" box="[532,620,1300,1325]" class="Reptilia" family="Varanidae" genus="Varanus" kingdom="Animalia" order="Squamata" pageId="13" pageNumber="420" phylum="Chordata" rank="genus">
|
|
<emphasis box="[532,620,1300,1325]" italics="true" pageId="13" pageNumber="420">Varanus</emphasis>
|
|
</taxonomicName>
|
|
species listed above, except
|
|
<taxonomicName authorityName="Zerova & Ckhikvadze, 1983 in Lungu, Zerova & Ckhikvadze" authorityYear="1983" box="[299,440,1332,1357]" class="Reptilia" family="Varanidae" genus="Varanus" kingdom="Animalia" order="Squamata" pageId="13" pageNumber="420" phylum="Chordata" rank="species" species="Tyrasiensis">
|
|
<emphasis box="[299,440,1332,1357]" italics="true" pageId="13" pageNumber="420">V. tyrasiensis</emphasis>
|
|
</taxonomicName>
|
|
and
|
|
<taxonomicName authorityName="Zerova & Ckhikvadze" authorityYear="1986" box="[503,644,1332,1357]" class="Reptilia" family="Varanidae" genus="Varanus" kingdom="Animalia" order="Squamata" pageId="13" pageNumber="420" phylum="Chordata" rank="species" species="Semjonovi">
|
|
<emphasis box="[503,644,1332,1357]" italics="true" pageId="13" pageNumber="420">V. semjonovi</emphasis>
|
|
</taxonomicName>
|
|
. Comparisons between the cotyle/condyle of the vertebrae from Molayan and those of
|
|
<taxonomicName authorityName="Zerova & Ckhikvadze, 1983 in Lungu, Zerova & Ckhikvadze" authorityYear="1983" box="[380,513,1396,1421]" class="Reptilia" family="Varanidae" genus="Varanus" kingdom="Animalia" order="Squamata" pageId="13" pageNumber="420" phylum="Chordata" rank="species" species="Tyrasiensis">
|
|
<emphasis box="[380,513,1396,1421]" italics="true" pageId="13" pageNumber="420">V. tyrasiensis</emphasis>
|
|
</taxonomicName>
|
|
cannot be made. However, the vertebrae of the latter species appear to be less depressed than the specimens from Molayan. The cotyle and condyle of
|
|
<taxonomicName authorityName="Zerova & Ckhikvadze" authorityYear="1986" box="[318,458,1492,1517]" class="Reptilia" family="Varanidae" genus="Varanus" kingdom="Animalia" order="Squamata" pageId="13" pageNumber="420" phylum="Chordata" rank="species" species="Semjonovi">
|
|
<emphasis box="[318,458,1492,1517]" italics="true" pageId="13" pageNumber="420">V. semjonovi</emphasis>
|
|
</taxonomicName>
|
|
are similar to those of the Molayan specimens; nevertheless, the latter vertebrae differ from those of
|
|
<taxonomicName authorityName="Zerova & Ckhikvadze" authorityYear="1986" box="[341,480,1555,1580]" class="Reptilia" family="Varanidae" genus="Varanus" kingdom="Animalia" order="Squamata" pageId="13" pageNumber="420" phylum="Chordata" rank="species" species="Semjonovi">
|
|
<emphasis box="[341,480,1555,1580]" italics="true" pageId="13" pageNumber="420">V. semjonovi</emphasis>
|
|
</taxonomicName>
|
|
in being more depressed and in having a deeper interzygapophyseal constriction. Finally, the fossil from Molayan differs from all extinct species. Nevertheless, the possibility that this
|
|
<taxonomicName authorityName="in Molnar" authorityYear="2004" box="[593,679,1651,1676]" class="Reptilia" family="Varanidae" genus="Varanus" kingdom="Animalia" order="Squamata" pageId="13" pageNumber="420" phylum="Chordata" rank="genus">
|
|
<emphasis box="[593,679,1651,1676]" italics="true" pageId="13" pageNumber="420">Varanus</emphasis>
|
|
</taxonomicName>
|
|
pertains to an extant species cannot be definitely excluded.
|
|
</paragraph>
|
|
</subSubSection>
|
|
</treatment>
|
|
</document> |