184 lines
23 KiB
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184 lines
23 KiB
XML
<document id="C7495ABF487F5787CA6A55848B4930E1" ID-DOI="10.5852/ejt.2024.961.2681" ID-ISSN="2118-9773" ID-Zenodo-Dep="13944155" ID-ZooBank="8E064900-1289-4648-BE9A-F17461CCF25C" IM.bibliography_approvedBy="guilherme" IM.illustrations_approvedBy="felipe" IM.materialsCitations_approvedBy="guilherme" IM.metadata_approvedBy="felipe" IM.tables_approvedBy="guilherme" IM.taxonomicNames_approvedBy="guilherme" IM.treatmentCitations_approvedBy="guilherme" IM.treatments_approvedBy="guilherme" checkinTime="1729153114719" checkinUser="plazi" docAuthor="Das, Indraneil, Shankar, P. Gowri, Swamy, Priyanka, Williams, Rhiannon C., Lalremsanga, Hmar Tlawmte, Prashanth, P., Sahoo, Gunanidhi, Vijayakumar, S. P., Höglund, Jacob, Shanker, Kartik, Dutta, Sushil K., Ganesh, S. R. & Wüster, Wolfgang" docDate="2024" docId="0386D2380C4BFFBE1E84FBA7FB83F944" docLanguage="en" docName="EJT.2024.961.1-51.pdf" docOrigin="European Journal of Taxonomy 961 (1)" docSource="https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2681/12413" docStyle="DocumentStyle:EF2B578F1D15862ADE45B0C07C620911.14:EJT.2018-.journal_article.type1" docStyleId="EF2B578F1D15862ADE45B0C07C620911" docStyleName="EJT.2018-.journal_article.type1" docStyleVersion="14" docTitle="Ophiophagus Gunther 1864" docType="treatment" docVersion="1" lastPageNumber="34" masterDocId="FFBFAA400C6AFF9F1E39FFB5FF87FFF6" masterDocTitle="Taxonomic revision of the king cobra Ophiophagus hannah (Cantor, 1836) species complex (Reptilia: Serpentes: Elapidae), with the description of two new species" masterLastPageNumber="51" masterPageNumber="1" pageNumber="34" updateTime="1729778983942" updateUser="guilherme" zenodo-license-document="CC-BY-4.0" zenodo-license-figures="CC-BY-4.0" zenodo-license-treatments="UNSPECIFIED">
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<mods:title id="01525659057A3C0F3A1AC4F0C54852C6">Taxonomic revision of the king cobra Ophiophagus hannah (Cantor, 1836) species complex (Reptilia: Serpentes: Elapidae), with the description of two new species</mods:title>
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<mods:namePart id="2F5A133DA8BA371D1E672D69596320B0">Das, Indraneil</mods:namePart>
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<mods:affiliation id="F076D2690ED642E822DE34C7EE7FDDE3">Institute of Biodiversity and Environmental Conservation, Universiti Malaysia Sarawak, 94300 Kota Samarahan, Sarawak, Malaysia.</mods:affiliation>
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<mods:affiliation id="6B6C1644CF7F4B334C35A73F23AD21B4">Maharaja Shrirama Chandra Bhanja Deo University, Baripada, Takatpur, Odisha, India. & Centre for Ecological Sciences, Indian Institute of Science, Bangalore 560 012, Karnataka, India. & Department of Ecology and Genetics, Evolutionary Biology Centre, Uppsala University, SE- 751 05 Uppsala, Sweden. & Kālinga Foundation, Guddakere, Agumbe, Shivamogga 577 411, Karnataka, India.</mods:affiliation>
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<mods:affiliation id="422A28C6B5A8C5586A6286A65139E8C4">Molecular Ecology and Evolution at Bangor, School of Environmental and Natural Sciences, Bangor University, Environment Centre Wales, Bangor LL 57 2 UW, Wales, UK.</mods:affiliation>
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<mods:namePart id="A463758E7736D0C9C887979A7739877C">Lalremsanga, Hmar Tlawmte</mods:namePart>
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<mods:affiliation id="1895159B2EC6317883A2C1754C631A18">Developmental Biology & Herpetology Laboratory, Department of Zoology, Mizoram University, Aizawl 796 004, Mizoram, India.</mods:affiliation>
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<mods:affiliation id="FA63DF9A12B7788E955E288E98791771">Kālinga Foundation, Guddakere, Agumbe, Shivamogga 577 411, Karnataka, India.</mods:affiliation>
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<mods:namePart id="55D3FA8CB30C2E8A322D09AFA02E7352">Sahoo, Gunanidhi</mods:namePart>
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<mods:affiliation id="9D5FACCE3FE9F9C7F8F503986FFC2364">Department of Zoology, Utkal University, Bhubaneswar 751 004, Odisha, India.</mods:affiliation>
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<mods:namePart id="00A78457D896F5967C08EF3235824AD9">Vijayakumar, S. P.</mods:namePart>
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<mods:affiliation id="C98F68340931E3DF16AE225D23202D4A">Kālinga Foundation, Guddakere, Agumbe, Shivamogga 577 411, Karnataka, India.</mods:affiliation>
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<mods:namePart id="1903AA9FA4892FC1FA172B74865DA400">Höglund, Jacob</mods:namePart>
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<mods:affiliation id="9B5E9566E716BB7278F7C2025A77E46C">Department of Ecology and Genetics, Evolutionary Biology Centre, Uppsala University, SE- 751 05 Uppsala, Sweden.</mods:affiliation>
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<mods:namePart id="91405F64922D60F1C9DA7CD6A1593FF4">Shanker, Kartik</mods:namePart>
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<mods:affiliation id="19ABBC1D14CE6FF234DCF93E23175EDC">Centre for Ecological Sciences, Indian Institute of Science, Bangalore 560 012, Karnataka, India.</mods:affiliation>
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<mods:namePart id="0416659FAE11B45FF13ADDB30CC23E10">Dutta, Sushil K.</mods:namePart>
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<mods:affiliation id="D309D8418017DB79EF2CE9F62CAF6AEF">Department of Zoology, Assam Don Bosco University, Tapesia 782 402, Assam, India.</mods:affiliation>
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<mods:namePart id="0B91C48BBB4E49D59C6265E87AAD1896">Ganesh, S. R.</mods:namePart>
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<mods:affiliation id="F8EBDD7E085E960F7D107CD54AF1327C">Kālinga Foundation, Guddakere, Agumbe, Shivamogga 577 411, Karnataka, India.</mods:affiliation>
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<mods:namePart id="735483CD07CB80F3C2FDF82672A9BE41">Wüster, Wolfgang</mods:namePart>
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<mods:affiliation id="FA82279FA6E979241E1A99AC7D484F4C">Molecular Ecology and Evolution at Bangor, School of Environmental and Natural Sciences, Bangor University, Environment Centre Wales, Bangor LL 57 2 UW, Wales, UK.</mods:affiliation>
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<mods:title id="E105AE954A1DCC295BC422DE04D3E50A">European Journal of Taxonomy</mods:title>
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<mods:part id="F489A327BEEE0C50950AA5CA78DCD06C">
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<mods:date id="8690B582EFBC3C732A70FF718DEE5B5F">2024</mods:date>
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<mods:number id="4122DDF34C5C8AA5A1A03FEE8688D809">2024-10-16</mods:number>
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<mods:identifier id="A324E61397EBA868288A62657B7BCF3B" type="DOI">10.5852/ejt.2024.961.2681</mods:identifier>
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<treatment id="0386D2380C4BFFBE1E84FBA7FB83F944" LSID="urn:lsid:zoobank.org:act:1D2264EB-DFE1-4A71-AAA3-0EC155480486" httpUri="http://treatment.plazi.org/id/0386D2380C4BFFBE1E84FBA7FB83F944" lastPageNumber="34" pageId="33" pageNumber="34">
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<subSubSection id="C33530A50C4BFFBE1E84FBA7FEECFBD8" box="[189,363,1042,1070]" pageId="33" pageNumber="34" type="nomenclature">
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<paragraph id="8B90632E0C4BFFBE1E84FBA7FEECFBD8" blockId="33.[189,363,1042,1070]" box="[189,363,1042,1070]" pageId="33" pageNumber="34">
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<taxonomicName id="4C2F18AD0C4BFFBE1E84FBA7FEECFBD8" ID-CoL="CDQ8Q" ID-ENA="8664" authority="Gunther, 1864" authorityName="Gunther" authorityYear="1864" box="[189,363,1042,1070]" class="Reptilia" family="Elapidae" genus="Ophiophagus" kingdom="Animalia" order="Squamata" pageId="33" pageNumber="34" phylum="Chordata" rank="genus">
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<emphasis id="B95BBF3C0C4BFFBE1E84FBA7FEECFBD8" bold="true" box="[189,363,1042,1070]" italics="true" pageId="33" pageNumber="34">Comparisons</emphasis>
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<subSubSection id="C33530A50C4BFFBE1E84FBF3FB83F944" pageId="33" pageNumber="34" type="discussion">
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<paragraph id="8B90632E0C4BFFBE1E84FBF3FE97F9F1" blockId="33.[187,1401,1094,1543]" pageId="33" pageNumber="34">
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Early attempts to understand variation in the complex suffered from a variety of shortcomings, due to the variation within (
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<figureCitation id="13147FAB0C4BFFBE1FBAFBDCFD82FB72" box="[387,517,1129,1156]" captionStart-0="Fig" captionStart-1="Fig" captionStartId-0="27.[189,232,1892,1918]" captionStartId-1="34.[189,231,1892,1918]" captionTargetBox-0="[208,1374,1254,1820]" captionTargetBox-1="[283,1300,766,1842]" captionTargetId-0="figure-341@27.[189,1398,1233,1852]" captionTargetId-1="figure-148@34.[189,1398,756,1853]" captionTargetPageId-0="27" captionTargetPageId-1="34" captionText-0="Fig. 11. Variation in dorsal bands in adults of the four species of Ophiophagus Günther, 1864 recognised in this study. A. O. kaalinga Gowri Shankar, Das & Ganesh sp. nov. B. O. hannah (Cantor, 1836). C. O. bungarus (Schlegel, 1837) comb. nov. D. O. salvatana Gowri Shankar, Das & Wüster sp. nov." captionText-1="Fig. 12. Variation in dorsal body bands in juveniles of the four species of Ophiophagus Günther, 1864 recognised in this study.A. O. hannah (Cantor, 1836) (photo: H.T. Lalremsanga).B. O. bungarus (Schlegel, 1837) comb. nov. (photo: T. Charlton). C. O. kaalinga Gowri Shankar, Das & Ganesh sp. nov. (photo: P. Gowri Shankar). D. O. salvatana Gowri Shankar, Das & Wüster sp. nov. (photo: Bernard Tarun)." figureDoi-0="http://doi.org/10.5281/zenodo.13944181" figureDoi-1="http://doi.org/10.5281/zenodo.13944183" httpUri-0="https://zenodo.org/record/13944181/files/figure.png" httpUri-1="https://zenodo.org/record/13944183/files/figure.png" pageId="33" pageNumber="34">Figs 11–12</figureCitation>
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) and among populations (
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<figureCitation id="13147FAB0C4BFFBE1D08FBDCFC32FB75" box="[817,949,1129,1155]" captionStart-0="Fig" captionStart-1="Fig" captionStart-2="Fig" captionStart-3="Fig" captionStartId-0="35.[189,232,1928,1954]" captionStartId-1="36.[189,232,1892,1918]" captionStartId-2="37.[189,232,1892,1918]" captionStartId-3="38.[188,230,1965,1991]" captionTargetBox-0="[230,1332,860,1851]" captionTargetBox-1="[363,1252,924,1810]" captionTargetBox-2="[234,1367,878,1844]" captionTargetBox-3="[189,1398,1477,1910]" captionTargetId-0="figure-192@35.[189,1398,830,1889]" captionTargetId-1="figure-237@36.[279,1309,905,1846]" captionTargetId-2="figure-221@37.[189,1398,844,1855]" captionTargetId-3="figure-494@38.[189,1398,1469,1920]" captionTargetPageId-0="35" captionTargetPageId-1="36" captionTargetPageId-2="37" captionTargetPageId-3="38" captionText-0="Fig. 13. Live unvouchered adults of Ophiophagus hannah (Cantor, 1836) showing variations in colour pattern. A. Mizoram, India (photo: H.T. Lalremsanga). B. China (Adam Francis). C. Thailand (photo: P. Gowri Shankar). D. Andaman Islands, India (photo: Aaron Fernandes)." captionText-1="Fig. 14. Live unvouchered adults of Ophiophagus bungarus (Schlegel, 1837) comb. nov. showing variations in colour pattern. A. Peninsular Malaysia (photo: Ahmad Khaldun Ismail). B. Java, Indonesia (photo: Nathan Rusli). C. Bali, Indonesia (photo: Shinta Sukum). D. East Malaysia (photo: P. Gowri Shankar)." captionText-2="Fig. 15. Live unvouchered adults of Ophiophagus kaalinga Gowri Shankar, Das & Ganesh sp. nov. showing variation in colour pattern. A. Kalakkad-Mundanthurai Tiger Reserve, India (photo: Naveen Joseph). B–C. Agumbe, Karnataka, India (photo: P. Gowri Shankar). D. Goa, India (photo: P. Gowri Shankar)." captionText-3="Fig. 16. Live unvouchered adults of Ophiophagus salvatana Gowri Shankar, Das & Wüster sp. nov. showing colouration. A. Entire. B. Head close-up profiles, Luzon, Philippines (both photos Emerson Sy)." figureDoi-0="http://doi.org/10.5281/zenodo.13944185" figureDoi-1="http://doi.org/10.5281/zenodo.13944187" figureDoi-2="http://doi.org/10.5281/zenodo.13944191" figureDoi-3="http://doi.org/10.5281/zenodo.13944193" httpUri-0="https://zenodo.org/record/13944185/files/figure.png" httpUri-1="https://zenodo.org/record/13944187/files/figure.png" httpUri-2="https://zenodo.org/record/13944191/files/figure.png" httpUri-3="https://zenodo.org/record/13944193/files/figure.png" pageId="33" pageNumber="34">Figs 13–16</figureCitation>
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) and scarcity of vouchered specimens, especially entire specimens of adults. For instance, all four species recognised here demonstrate dramatic ontogenetic colour and pattern changes, with the banded juveniles losing the pattern in adults consistently in one species (
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<taxonomicName id="4C2F18AD0C4BFFBE1F4DFB66FD8CFB1B" authorityName="Das & Shankar & Swamy & Williams & Lalremsanga & Prashanth & Sahoo & Vijayakumar & Höglund & Shanker & Dutta & Ganesh & Wüster" authorityYear="2024" box="[372,523,1235,1261]" class="Reptilia" family="Elapidae" genus="Ophiophagus" kingdom="Animalia" order="Squamata" pageId="33" pageNumber="34" phylum="Chordata" rank="species" species="salvatana" status="sp. nov.">
|
||
<emphasis id="B95BBF3C0C4BFFBE1F4DFB66FD8CFB1B" box="[372,523,1235,1261]" italics="true" pageId="33" pageNumber="34">O. salvatana</emphasis>
|
||
</taxonomicName>
|
||
<taxonomicNameLabel id="A26802470C4BFFBE1C2DFB61FDF7FB1B" box="[532,624,1235,1262]" pageId="33" pageNumber="34" rank="species">sp. nov.</taxonomicNameLabel>
|
||
) and sometimes in another (
|
||
<taxonomicName id="4C2F18AD0C4BFFBE1DF9FB66FBD4FB1B" authorityName="Das & Shankar & Swamy & Williams & Lalremsanga & Prashanth & Sahoo & Vijayakumar & Höglund & Shanker & Dutta & Ganesh & Wüster" authorityYear="2024" baseAuthorityName="Schlegel" baseAuthorityYear="1837" box="[960,1107,1235,1261]" class="Reptilia" family="Elapidae" genus="Ophiophagus" kingdom="Animalia" order="Squamata" pageId="33" pageNumber="34" phylum="Chordata" rank="species" species="bungarus" status="comb. nov.">
|
||
<emphasis id="B95BBF3C0C4BFFBE1DF9FB66FBD4FB1B" box="[960,1107,1235,1261]" italics="true" pageId="33" pageNumber="34">O. bungarus</emphasis>
|
||
</taxonomicName>
|
||
<taxonomicNameLabel id="A26802470C4BFFBE1A65FB66FB5AFB18" box="[1116,1245,1235,1262]" pageId="33" pageNumber="34" rank="species">comb. nov.</taxonomicNameLabel>
|
||
), retaining it into adulthood in a third population (
|
||
<taxonomicName id="4C2F18AD0C4BFFBE1C52FB42FD71FAE6" authorityName="Das & Shankar & Swamy & Williams & Lalremsanga & Prashanth & Sahoo & Vijayakumar & Höglund & Shanker & Dutta & Ganesh & Wüster" authorityYear="2024" box="[619,758,1270,1297]" class="Reptilia" family="Elapidae" genus="Ophiophagus" kingdom="Animalia" order="Squamata" pageId="33" pageNumber="34" phylum="Chordata" rank="species" species="kaalinga" status="sp. nov.">
|
||
<emphasis id="B95BBF3C0C4BFFBE1C52FB42FD71FAE6" box="[619,758,1270,1297]" italics="true" pageId="33" pageNumber="34">O. kaalinga</emphasis>
|
||
</taxonomicName>
|
||
<taxonomicNameLabel id="A26802470C4BFFBE1CC5FB42FCDDFAE6" box="[764,858,1270,1297]" pageId="33" pageNumber="34" rank="species">sp. nov.</taxonomicNameLabel>
|
||
), and developing dark edges to the band in the fourth (
|
||
<taxonomicName id="4C2F18AD0C4BFFBE1F21FAAFFE12FAC5" baseAuthorityName="Cantor" baseAuthorityYear="1836" box="[280,405,1305,1332]" class="Reptilia" family="Elapidae" genus="Ophiophagus" kingdom="Animalia" order="Squamata" pageId="33" pageNumber="34" phylum="Chordata" rank="species" species="hannah">
|
||
<emphasis id="B95BBF3C0C4BFFBE1F21FAAFFE12FAC5" box="[280,405,1305,1332]" italics="true" pageId="33" pageNumber="34">O. hannah</emphasis>
|
||
</taxonomicName>
|
||
). Further, the allopatric populations of the four species recognized in this paper do not show categorical differences in the counts of the major scales that have been traditionally employed for taxonomic differentiation of species within snakes. However, a number of characters are useful to unequivocally diagnose the four species recognized here. Character states useful for identification and diagnosis of the members of the complex include dorsal ground colour and pattern, nature of body bands in adults (unbanded and banded, the bands with or without a dark edge) and pterygoid tooth/tooth socket counts.
|
||
</paragraph>
|
||
<paragraph id="8B90632E0C4BFFBE1E84F99AFB83F944" blockId="33.[189,1401,1582,1714]" pageId="33" pageNumber="34">
|
||
Means, ranges and sample size of characters differentiating the four lineages identified in this study and supported by our mitochondrial phylogeny (see
|
||
<bibRefCitation id="EFBE1EDF0C4BFFBE1CC6F9E4FBD8F99A" author="Gowri Shankar P. & Swamy P. & Williams R. C. & Ganesh S. R. & Moss M. & Hoglund J. & Das I. & Sahoo G. & Vijayakumar S. P. & Shankar K. & Wuster W. & Dutta S. K." box="[767,1119,1617,1644]" pageId="33" pageNumber="34" refId="ref27441" refString="Gowri Shankar P., Swamy P., Williams R. C., Ganesh S. R., Moss M., Hoglund J., Das I., Sahoo G., Vijayakumar S. P., Shankar K., Wuster W. & Dutta S. K. 2021. King or royal family? Testing for species boundaries in the king cobra, Ophiophagus hanah (Cantor, 1836), using morphology and multilocus DNA analyses. Molecular Phylogenetics and Evolution 164: e 107300. https: // doi. org / 10.1016 / j. ympev. 2021.107300" type="journal volume" year="2021">
|
||
Gowri Shankar
|
||
<emphasis id="B95BBF3C0C4BFFBE1D86F9E7FC74F99D" box="[959,1011,1617,1643]" italics="true" pageId="33" pageNumber="34">et al</emphasis>
|
||
. 2021: 5
|
||
</bibRefCitation>
|
||
) are shown in
|
||
<tableCitation id="C6AD56950C4BFFBE1B2FF9E4FAF7F99D" box="[1302,1392,1617,1643]" captionStart="Table 1" captionStartId="31.[188,258,265,291]" captionText="Table 1. Summary of morphological characters associated with species of Ophiophagus Günther, 1864 recognised in this study. See ‘Terminology for character analysis’ for details. Note: pale dorsal bands are sometimes absent in adult O. bungarus (Schlegel, 1837) comb. nov. (in images of live specimens examined), and consistently absent in adult O. salvatana Gowri Shankar, Das & Wüster sp. nov. Further, juveniles of the latter show pale bands (in images of three live specimens examined; no voucher specimens of juvenile O. salvatana were available for study)." pageId="33" pageNumber="34">Table 1</tableCitation>
|
||
. Measurement data are in
|
||
<tableCitation id="C6AD56950C4BFFBE1FDDF9C1FDBAF979" box="[484,573,1652,1679]" captionStart="Table 2" captionStartId="32.[189,259,265,291]" captionText="Table 2. Summary of measurements (range [n], in mm) of species of Ophiophagus Günther, 1864 recognised in this study. See ‘Terminology for character analysis’ for details. Abbreviations: BW = Midbody diameter; ED = Eye width; EYEN = Eye to nostril distance; EYES = Eye to snout distance; FL = Fang length from base; FW = Maximum fang width; HD = Head depth; HL = Head length; HW = Head width; IORB = Interorbital width; NOD = Nostril diameter; OCCL = Occipital length; ROSL = Rostral length; ROSW = Rostral width; SVL = Snout-vent length; TL = Tail length (if entire)." pageId="33" pageNumber="34">Table 2</tableCitation>
|
||
. Furthermore, we provide here a dichotomous identification key for the identification of the species of the genus
|
||
<taxonomicName id="4C2F18AD0C4BFFBE1CA4F92DFCBCF944" authorityName="Gunther" authorityYear="1864" box="[669,827,1688,1714]" class="Reptilia" family="Elapidae" genus="Ophiophagus" kingdom="Animalia" order="Squamata" pageId="33" pageNumber="34" phylum="Chordata" rank="genus">
|
||
<emphasis id="B95BBF3C0C4BFFBE1CA4F92DFCBCF944" box="[669,827,1688,1714]" italics="true" pageId="33" pageNumber="34">Ophiophagus</emphasis>
|
||
</taxonomicName>
|
||
recognized here.
|
||
</paragraph>
|
||
</subSubSection>
|
||
</treatment>
|
||
</document> |