421 lines
54 KiB
XML
421 lines
54 KiB
XML
<document id="D145DD876779FA1CBE10F595DBCC4369" ID-CLB-Dataset="30323" ID-DOI="http://dx.doi.org/10.3897/zookeys.748.20507" ID-GBIF-Dataset="236eed9f-0210-4fe7-8fc6-fab9de3b2354" ID-PMC="PMC5904387" ID-Pensoft-Pub="1313-2970-748-97" ID-PubMed="29674917" ID-ZooBank="7933C25326654191871DBF653A82FE06" ModsDocAuthor="" ModsDocDate="2018" ModsDocID="1313-2970-748-97" ModsDocOrigin="ZooKeys 748" ModsDocTitle="Molecular and morphological differentiation of Secret Toad-headed agama, Phrynocephalusmystaceus, with the description of a new subspecies from Iran (Reptilia, Agamidae)" checkinTime="1522997106399" checkinUser="pensoft" docAuthor="Solovyeva, Evgeniya N., Dunayev, Evgeniy N., Nazarov, Roman A., Mehdi Radjabizadeh,, Poyarkov, Nikolay A. & Jr.," docDate="2018" docId="F372CBEFEDFD55F462B34E0E07E4C47A" docLanguage="en" docName="ZooKeys 748: 97-129" docOrigin="ZooKeys 748" docSource="http://dx.doi.org/10.3897/zookeys.748.20507" docTitle="Phrynocephalus mystaceus subsp. khorasanus Solovyeva, Dunayev, Nazarov, Radjabizadeh, Poyarkov & Jr., 2018, ssp. n." docType="treatment" docUuid="6E926506-3D7A-4C99-BF64-A02C48157B5C" docUuidSource="ZooBank" docVersion="8" lastPageNumber="105" masterDocId="706EFFD94157FFB68751291CFFE0FFA3" masterDocTitle="Molecular and morphological differentiation of Secret Toad-headed agama, Phrynocephalusmystaceus, with the description of a new subspecies from Iran (Reptilia, Agamidae)" masterLastPageNumber="129" masterPageNumber="97" pageNumber="99" updateTime="1701375423160" updateUser="ExternalLinkService">
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<mods:titleInfo id="A7DBA828B9EEEF2AE8415F750E984BE0">
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<mods:title id="DD2B60C0716FB6FF6A384DCAC89E3DBB">Molecular and morphological differentiation of Secret Toad-headed agama, Phrynocephalusmystaceus, with the description of a new subspecies from Iran (Reptilia, Agamidae)</mods:title>
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<mods:namePart id="B935A61C3C258DD95419E1CC304E21BD">Solovyeva, Evgeniya N.</mods:namePart>
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<mods:roleTerm id="6E96A5B3C095E7150FFE8FDB30BA0D0D">Author</mods:roleTerm>
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<mods:namePart id="9E3F12A3BC4EA72870A575079055A83C">Dunayev, Evgeniy N.</mods:namePart>
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<mods:name id="22B34A74C98E1F096C3DFE84C59C933A" type="personal">
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<mods:roleTerm id="2763BDE6A3E3405AC21ECEFE8D882024">Author</mods:roleTerm>
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</mods:role>
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<mods:namePart id="62A84D338D870A8EB419ECCF40C1EFD7">Nazarov, Roman A.</mods:namePart>
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</mods:name>
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<mods:name id="92D24D07AD8301C870785436EC030E11" type="personal">
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<mods:roleTerm id="1AECF53BF097490F312B7F9B265424AB">Author</mods:roleTerm>
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<mods:namePart id="D457F200AA30216EF389610E79863A00">Mehdi Radjabizadeh,</mods:namePart>
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</mods:name>
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<mods:namePart id="A06E255A676ACFCA55A6D1AFCC9B777E">Poyarkov, Nikolay A.</mods:namePart>
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<mods:namePart id="FBFD53779A180218949BD907591C99E3">Jr.,</mods:namePart>
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<mods:part id="93BCE36EF211A16A5A9C7C1BF81D13DD">
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<mods:date id="70FF1354FA99E4590403339BEB5D830A">2018</mods:date>
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<mods:detail id="2AB5CB26CB1FCF7CA8B1BD6487B6FEB0" type="volume">
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<mods:number id="188B721C2F4E54D5B4E532C29E6BA382">748</mods:number>
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<mods:start id="806E4F846095596CFAAD62654F47B243">97</mods:start>
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<mods:url id="AB0BD8315B7FB556B5BF79FAA6ABC886">http://dx.doi.org/10.3897/zookeys.748.20507</mods:url>
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<mods:classification id="21728CAB3C073A223116FE12E618F4C6">journal article</mods:classification>
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<mods:identifier id="DDDA14C4C3FB3699AA0E7097F299C668" type="DOI">http://dx.doi.org/10.3897/zookeys.748.20507</mods:identifier>
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<mods:identifier id="FA63D8BCA1A0FFFCDEFC0FFC67728673" type="Pensoft-Pub">1313-2970-748-97</mods:identifier>
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<mods:identifier id="E99AF6EA606C6BDC6A6D8B970567B2FC" type="ZooBank">7933C25326654191871DBF653A82FE06</mods:identifier>
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<treatment id="F372CBEFEDFD55F462B34E0E07E4C47A" ID-GBIF-Taxon="143478655" LSID="urn:lsid:zoobank.org:act:6E926506-3D7A-4C99-BF64-A02C48157B5C" httpUri="http://treatment.plazi.org/id/F372CBEFEDFD55F462B34E0E07E4C47A" lastPageId="8" lastPageNumber="105" pageId="2" pageNumber="99">
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<subSubSection id="454BC8EC635CEECED775C56FF3F36C42" pageId="2" pageNumber="99" type="nomenclature">
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<paragraph id="E54C07A06DA29CF90BE475878BC0F71A" pageId="2" pageNumber="99">
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<taxonomicName id="959C435D031669C219B28E968CA4CDBE" ID-CoL="8KBQ6" LSID="http://zoobank.org/6E926506-3D7A-4C99-BF64-A02C48157B5C" class="Squamata" family="Agamidae" genus="Phrynocephalus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Phrynocephalus mystaceus" order="Squamata" pageId="2" pageNumber="99" phylum="Chordata" rank="subSpecies" species="mystaceus" subSpecies="khorasanus">Phrynocephalus mystaceus khorasanus</taxonomicName>
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<taxonomicNameLabel id="0CDC355EEC4138F1B8021986ECF66A71" pageId="2" pageNumber="99">ssp. n.</taxonomicNameLabel>
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Figs 6A, B; 7; 8; Table 4
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</paragraph>
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</subSubSection>
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<subSubSection id="1C6590B91F043A310FFB994B792D32FB" pageId="2" pageNumber="99" type="holotype">
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<paragraph id="A6FECAB187C8D003477A81FB13B757B2" pageId="2" pageNumber="99">Holotype.</paragraph>
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<paragraph id="058FACE40CABEE1C31DCBA8B7DC58CF3" pageId="2" pageNumber="99">ZMMU R-11913 (adult female; field number NR-1191).</paragraph>
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</subSubSection>
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<subSubSection id="F8C6501E5CEAD9B24EA21F0CF6694655" pageId="2" pageNumber="99" type="type locality">
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<paragraph id="29F3A16B8FD083EA3A3890C59E0411B2" pageId="2" pageNumber="99">Type locality.</paragraph>
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<paragraph id="DF02A26A93F79EBD205273F9E0960724" pageId="2" pageNumber="99">
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Iran, Khorasan historical area, Khorasan Razavi Province (ostan), environs of Gonabad, the right bank of the Kale-Shur River; sand dunes (see Fig. 9);
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<geoCoordinate id="9DE318614295ADB69608E4622A817E44" direction="north" orientation="latitude" precision="925" value="34.65">N34°39'</geoCoordinate>
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,
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<geoCoordinate id="65A5642FAB106ABA3CAE751F2CB47890" direction="east" orientation="longitude" precision="925" value="58.716667">E58°43'</geoCoordinate>
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; elevation 850 m a. s. l. Collected by Roman A. Nazarov and Mehdi Radjabizadeh on April 25, 2005.
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</paragraph>
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</subSubSection>
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<subSubSection id="408EA90E74A3B0BAB8778968361A1E8D" pageId="2" pageNumber="99" type="paratypes">
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<paragraph id="EA697C6234A4E24D6D9EE2F9BB37DD3B" pageId="2" pageNumber="99">Paratypes.</paragraph>
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<paragraph id="304B09CB6DA30C3E6346C82D13FD4211" pageId="2" pageNumber="99">
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ZMMU R-13009 (one adult male with everted hemipenial structures, field number RAN 1723; and one adult female, field number RAN 1724) was collected in Iran, Khorasan historical area, Khorasan Razavi Province, 20 km east of the town of Boshrouyeh (
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<geoCoordinate id="2C513A4F189249E1C18AEE5DF8377505" direction="north" orientation="latitude" precision="925" value="33.9">N33°54'</geoCoordinate>
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,
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<geoCoordinate id="BA866F97B4241538EB3DB0A6934C1503" direction="east" orientation="longitude" precision="925" value="57.5">E57°30'</geoCoordinate>
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; elevation 864 m a. s. l.) by Dmitriy A. Bondarenko, Roman A. Nazarov, and Mehdi Radjabizadeh on May 05, 2009. The rest of paratypes were collected in the area close to the type locality. ZMMU R-13011 (one adult male with hemipenial structures, field number RAN 1947; and one subadult female, field number RAN 1948) was collected in Iran, Khorasan Razavi Province, 60 km north of the town of Gonabad, stabilized or semi-stabilized sands (
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<geoCoordinate id="BB15AF0C06E75E4A6267AFF29F2FEF2C" direction="north" orientation="latitude" precision="925" value="34.6">N34°36'</geoCoordinate>
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,
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<geoCoordinate id="343F4B14BED8CE23CDC23AD9C46CFB03" direction="east" orientation="longitude" precision="925" value="58.233334">E58°14'</geoCoordinate>
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; elevation 867 m a. s. l.) by Roman A. Nazarov, Rustam K. Berdiev, Vlad G. Starkov, and Mehdi Radjabizadeh on June 02, 2009. ZMMU R-13169 (subadult female) was collected in Iran, Khorasan Razavi Province, 30 km north of the town of Gonabad, on sandy massif on the right bank of the Kale-Shur river (
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<geoCoordinate id="F9F6FFDB69E0C5D075DF256F0BF66785" direction="north" orientation="latitude" precision="925" value="34.583332">N34°35'</geoCoordinate>
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,
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<geoCoordinate id="86415BEF6448544B7D64ACDA022281AF" direction="east" orientation="longitude" precision="925" value="58.716667">E58°43'</geoCoordinate>
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; elevation 888 m a. s. l.) by Roman A. Nazarov, Dmitriy A. Bondarenko, and Mehdi Radjabizadeh on May 10, 2010. ZMMU R-12202 (juvenile female with slightly orange lower surface of the tail, field number N-093) was collected in Iran, Khorasan Razavi Province, 60 km north of the town of Gonabad, on sands (
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<geoCoordinate id="E4ACC40C7771C6DFFAEEA89BC0708FFC" direction="north" orientation="latitude" precision="925" value="34.6">N34°36'</geoCoordinate>
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,
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<geoCoordinate id="9DCB4723E5E95900817DDE893B22C7D6" direction="east" orientation="longitude" precision="925" value="58.733334">E58°44'</geoCoordinate>
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; elevation 881 m a.s.l.) by Dmitriy A. Bondarenko on April 20, 2006.
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</paragraph>
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<caption id="38FAA1D63AF60A08EB3F5FBA246984FE" pageId="2" pageNumber="99">
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<paragraph id="FB300E487219F40745FC15F9D9B8E1E4" pageId="2" pageNumber="99">
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Figure 6.
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<taxonomicName id="C17F56926C4B7753998DA335EB3168AF" lsidName="Ph. mystaceus" pageId="2" pageNumber="99" rank="species" species="mystaceus">Ph. mystaceus</taxonomicName>
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in life: A subadult
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<taxonomicName id="A4D2EC3DFB400905D9DEA8C126AE2864" lsidName="Ph. mystaceus" pageId="2" pageNumber="99" rank="subSpecies" species="mystaceus" subSpecies="khorasanus">Ph. mystaceus khorasanus</taxonomicName>
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ssp. n., orange lower surface of the tail is shown, Iran (photograph by R. A. Nazarov) B
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<taxonomicName id="F669EE38B8CE90404FDB94091274DC79" lsidName="Ph. mystaceus" pageId="2" pageNumber="99" rank="subSpecies" species="mystaceus" subSpecies="khorasanus">Ph. mystaceus khorasanus</taxonomicName>
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ssp. n., female, Iran (photo by R. A. Nazarov) C
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<taxonomicName id="DB0A97F6021D8D3EC8C295A5BC75CBE1" lsidName="Ph. m." pageId="2" pageNumber="99" rank="subSpecies" species="m." subSpecies="mystaceus">Ph. m. mystaceus</taxonomicName>
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, Russia, Astrakhan region, Dosang (photograph by E. A. Dunayev) D
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<taxonomicName id="52B947029D066307B2B577DF0985D902" lsidName="Ph. m." pageId="2" pageNumber="99" rank="subSpecies" species="m." subSpecies="mystaceus">Ph. m. mystaceus</taxonomicName>
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, Dagestan, Sarykum sands (photograph by E. A. Dunayev) E
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<taxonomicName id="41A6B3F641252967406E7ABC4F23B830" lsidName="Ph. m." pageId="2" pageNumber="99" rank="subSpecies" species="m." subSpecies="mystaceus">Ph. m. mystaceus</taxonomicName>
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, Uzbekistan, Qarakalpaqiston (corresponds to the previously recognized subspecies "
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<taxonomicName id="7F70B3BC5B16422D207BFD343EB5ECEA" lsidName="" pageId="2" pageNumber="99" rank="subSpecies" subSpecies="galli">galli</taxonomicName>
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"; photograph by E. A. Dunayev) F
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<taxonomicName id="E5CADE33D9E902AF1D12FCC881E1CC1A" lsidName="Ph. m." pageId="2" pageNumber="99" rank="subSpecies" species="m." subSpecies="mystaceus">Ph. m. mystaceus</taxonomicName>
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, Dagestan, Sarykum sands (corresponds to the previously recognized subspecies "
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<taxonomicName id="E26E1B2B40ECBDE610036F400342B05B" lsidName="" pageId="2" pageNumber="99" rank="subSpecies" subSpecies="dagestanica">dagestanica</taxonomicName>
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"; photograph by E. A. Dunayev) G
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<taxonomicName id="22B694717F66A3BA08754F37F063204E" lsidName="Ph. m." pageId="2" pageNumber="99" rank="subSpecies" species="m." subSpecies="aurantiacocaudatus">Ph. m. aurantiacocaudatus</taxonomicName>
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, E Kazakhstan, SE Balkash Lake (photograph by E. N. Solovyeva) H
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<taxonomicName id="01CFF7FF4945E16FC9E9EAC9A61B990D" lsidName="Ph. m." pageId="2" pageNumber="99" rank="subSpecies" species="m." subSpecies="aurantiacocaudatus">Ph. m. aurantiacocaudatus</taxonomicName>
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, E Kazakhstan, SE Balkash lake (photograph by E. N. Solovyeva) I
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<taxonomicName id="0183361E4E7545E97F3EB08783821601" lsidName="Ph. m." pageId="2" pageNumber="99" rank="subSpecies" species="m." subSpecies="mystaceus">Ph. m. mystaceus</taxonomicName>
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, Russia, Astrakhan region, Dosang (photograph by E. A. Dunayev).
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</paragraph>
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</caption>
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</subSubSection>
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<subSubSection id="5F50C404B55823C5BEEE45018E4CBD3F" pageId="2" pageNumber="99" type="diagnosis">
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<paragraph id="3B580E48F15D8DAE2BCE8160BED6DB47" pageId="2" pageNumber="99">Diagnosis.</paragraph>
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<paragraph id="025396F41C595B00399C42A54E97EEC7" pageId="2" pageNumber="99">
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A member of
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<taxonomicName id="69DE771E6776082503F8D78C85C442F8" lsidName="Ph. mystaceus" pageId="2" pageNumber="99" rank="species" species="mystaceus">Ph. mystaceus</taxonomicName>
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species complex based on the following combination of morphological attributes: (1) a large-sized
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<taxonomicName id="A12ACD373381E33B1CDA7CEFE8F4F0F4" class="Squamata" family="Agamidae" genus="Phrynocephalus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Phrynocephalus" pageId="2" pageNumber="99" phylum="Chordata" rank="genus">Phrynocephalus</taxonomicName>
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species with SVL up to 97.5 mm, tail shorter than SVL; (2) pair of cutaneous flaps present at mouth corners with numerous spiny scales along flap edges; (3) distinctly flattened body and tail; (4) toes with fringes formed by triangular scales; subdigital lamellae on toes III and IV with ridges.
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<taxonomicName id="C51D98496F8C1AF7D32885F58B2FD98C" class="Squamata" family="Agamidae" genus="Phrynocephalus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Phrynocephalus mystaceus" pageId="2" pageNumber="99" phylum="Chordata" rank="subSpecies" species="mystaceus" subSpecies="khorasanus">Phrynocephalus mystaceus khorasanus</taxonomicName>
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ssp. n. can be distinguished from the nominative subspecies of
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<taxonomicName id="8FBF812007289E1B164E10DF12C0C920" lsidName="Ph. mystaceus" pageId="2" pageNumber="99" rank="species" species="mystaceus">Ph. mystaceus</taxonomicName>
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by the following combination of two diagnostic morphological characteristics: (1) 24-27 lamellae on toe IV; (2) few supralabial scales (less than 14). In life, the new subspecies can be further distinguished from the nominative subspecies by the orange color of the lower surface of tail in young specimens (lemon to yellowish in
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<taxonomicName id="0581E12EFBE3657ED98008499D52703B" lsidName="Ph. m." pageId="2" pageNumber="99" rank="subSpecies" species="m." subSpecies="mystaceus">Ph. m. mystaceus</taxonomicName>
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except the populations from Eastern Kazakhstan and western China, formerly described as
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<taxonomicName id="66862300AD36E25BF7AF68CB6B033659" lsidName="Ph. m." pageId="2" pageNumber="99" rank="subSpecies" species="m." subSpecies="aurantiacocaudatus">Ph. m. aurantiacocaudatus</taxonomicName>
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). MtDNA sequences of
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<taxonomicName id="D100C99D9E554BF598D951E696E2B427" class="Squamata" family="Agamidae" genus="Phrynocephalus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Phrynocephalus mystaceus" pageId="2" pageNumber="99" phylum="Chordata" rank="subSpecies" species="mystaceus" subSpecies="khorasanus">Phrynocephalus mystaceus khorasanus</taxonomicName>
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ssp. n. are markedly distinct from those in all other populations of
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<taxonomicName id="7DA254B5FA9EC802D7A78ECE867CD3DF" lsidName="Ph. mystaceus" pageId="2" pageNumber="99" rank="species" species="mystaceus">Ph. mystaceus</taxonomicName>
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with sequence divergence in the range of 6.84-7.28% between them. The new subspecies is notably smaller than the representatives of southern populations of
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<taxonomicName id="EFAB21BF7CB37C7A58ABF54CF64F8A52" lsidName="Ph. m." pageId="2" pageNumber="99" rank="subSpecies" species="m." subSpecies="mystaceus">Ph. m. mystaceus</taxonomicName>
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from Uzbekistan and Turkmenistan, formerly described as
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<taxonomicName id="0EB2946C899BA7FA8632A287EC7F3656" lsidName="Ph. m." pageId="2" pageNumber="99" rank="subSpecies" species="m." subSpecies="galli">Ph. m. galli</taxonomicName>
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, which can reach SVL up to 122.7 mm (
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<bibRefCitation id="27131A855BD41E9FC84B1AE5D833C2F9" author="Anderson, SC" journalOrPublisher="Society for the Study of Amphibians and Reptiles, Ithaca, New York" pageId="8" pageNumber="105" title="The Lizards of Iran." year="1999">Anderson 1999</bibRefCitation>
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), whereas for Iranian population
|
||
<bibRefCitation id="B8F97AE3DE3BB7C9721A837A16F9A804" author="Anderson, SC" journalOrPublisher="Society for the Study of Amphibians and Reptiles, Ithaca, New York" pageId="8" pageNumber="105" title="The Lizards of Iran." year="1999">Anderson (1999)</bibRefCitation>
|
||
reported the largest specimen of
|
||
<taxonomicName id="369F39B1CC7D0A25429786214C3BCBFA" lsidName="Ph. mystaceus" pageId="2" pageNumber="99" rank="species" species="mystaceus">Ph. mystaceus</taxonomicName>
|
||
to have SVL up to 77.7 mm. SVL in the largest specimen in our sampling reached 86.0 mm, while
|
||
<bibRefCitation id="774D146AE2682090DC44647E90C7B6A7" author="Molavi, F" journalOrPublisher="Check List" pageId="9" pageNumber="106" pagination="450 - 452" title="New distribution Record of Phrynocephalusmystaceus Pallas 1776 (Sauria: Agamidae) from Iran." volume="10" year="2014">Molavi et al. (2014)</bibRefCitation>
|
||
recorded a specimen with SVL of 97.5 from Semnan Province.
|
||
</paragraph>
|
||
</subSubSection>
|
||
<subSubSection id="BF71AC541FB8E6C74037B852E66AEB44" pageId="2" pageNumber="99" type="etymology">
|
||
<paragraph id="EE70A7479FBCD0AB8ADFD77B0DE2855C" pageId="2" pageNumber="99">Etymology.</paragraph>
|
||
<paragraph id="2641A822E9FA6761D9EB03CCD02EC3C0" pageId="2" pageNumber="99">
|
||
The name of the new subspecies
|
||
<taxonomicName id="FA43EF8A923535A0B07349E7FB5176FD" lsidName="" pageId="2" pageNumber="99" rank="subSpecies" subSpecies="khorasanus">khorasanus</taxonomicName>
|
||
is a Latinized toponymic adjective, derived from Khorasan, the name of the historic area and a Khorasan Razavi Province in the northeast Iran, where the new subspecies was found. We suggest the "Khorasan Secret Toad-headed Agama" as a common name in English.
|
||
</paragraph>
|
||
</subSubSection>
|
||
<subSubSection id="62FA77B7EE967968720357CCC5861C1C" lastPageId="4" lastPageNumber="101" pageId="2" pageNumber="99" type="description">
|
||
<paragraph id="6F1BEAD17C99BD5B513027EF818D586D" pageId="2" pageNumber="99">Description of holotype.</paragraph>
|
||
<paragraph id="DE7EE42E9C650D242BB0D31674FD0EA4" pageId="2" pageNumber="99">Medium-sized agamid lizard, adult female, specimen in good state of preservation; body dissected on ventral side along the midline of belly (dissection ca. 20 cm in length). Measurements and counts of the holotype are presented in Table 4.</paragraph>
|
||
<paragraph id="37D790B3CE459CE86F305087A6BA932F" lastPageId="4" lastPageNumber="101" pageId="2" pageNumber="99">
|
||
Head large, rounded, distinctly wider than neck region (see Fig. 7A); body and tail notably flattened. Snout abruptly blunt, head almost vertical in profile view (see Fig. 7E), nostrils invisible dorsally (see Fig. 7C). Nasals separated from each other by single scale (see Fig. 7D). Dorsal surface of head with distinct pileus consisting of small slightly keeled scales; ca. 30 scales across the pileus. Pineal scale separated from nasals by 13 smaller scales; scales covering orbital area somewhat smaller than those on frontal surface of head; occipital scales not enlarged. Five scales contacting subnasal ventrally (see Fig. 7D). Subnasal scale not in contact with inner (medial) side of supranasal. Supralabials separated from subnasal scale by 6 rows of small granular scales (see Fig. 7D). Pair of skin-folds form characteristic ear-shaped flaps in mouth corners, edges of each flap with enlarged conical scales, two groups of similarly enlarged conical scales on each side of head posterior to the mouth angle at tympanal area (see Fig. 7E). Supralabial scales anterior to cutaneous fold at mouth angle 11/12 (hereafter data for symmetrical characteristics is given in Right/Left order); 9/9 of anterior supralabials notably flattened, 2/3 posterior supralabials conical-shaped; supralabials separated from small granular scales of lower eyelid by 3/4 rows of scales, ventral row of these
|
||
<pageBreakToken id="63EAF43506FD70C7F96F1BE5C89C44C4" pageId="3" pageNumber="100" start="start">scales</pageBreakToken>
|
||
almost the same size as supralabials (see Fig. 7E). Single small scale between the posteriormost supralabial and insertion of cutaneous fold at mouth angle. Infralabial scales anterior to cutaneous fold - 6/6, 3/3 of anterior infralabials notably flattened, posterior infralabials cone-shaped. Posterior corner of eye and insertion of cutaneous fold at mouth angle separated by row of three enlarged flat scales (see Fig. 7E). Vertebral scales not enlarged. Scales at middle of dorsum slightly bigger than scales on dorsolateral and lateral surfaces of body. Dorsal scales with weak keels, becoming cone-shaped laterally, forming almost triangular spines on the flanks. Notably enlarged spiny scale (about four times the size of adjacent scales) on each side of thorax behind maxilla, two groups of enlarged spiny scales on each lateral surface of neck region (see Fig. 7E). Tail notably flattened along its whole length. Scales on dorsal surface of tail
|
||
<pageBreakToken id="93ED39BE087190B49B06D0EA49230951" pageId="4" pageNumber="101" start="start">and</pageBreakToken>
|
||
on ventral surface of tail posterior half notably keeled; scales on lateral sides of tail with well-pronounced spines. Limbs comparatively long: hindlimb length greater than distance from cloaca to gular fold. Toe IV bearing a single row of subdigital lamellae, each with a well-pronounced ridge on its volar surface; lateral sides of toe IV with two rows of enlarged triangular scales that form distinct serrated fringe (see Fig. 7F). Similar crests present on lateral surfaces of toe III, triangular scales on toe III notably smaller compared to those on toe IV (see Fig. 7F). Number of lamellae on toe IV 24/24, on toe III 16/16; number of enlarged triangular scales on toe IV 20/20, on toe III 9/9.
|
||
</paragraph>
|
||
<caption id="C3FFD00083AE17BC24C965D2850920D9" pageId="4" pageNumber="101">
|
||
<paragraph id="06B3C6F5013FE3BCCCB9197BB5A0B34D" pageId="4" pageNumber="101">
|
||
Figure 7. Holotype of
|
||
<taxonomicName id="06A723C0CCD6EA96F78E6A3B7907BE55" lsidName="Ph. mystaceus" pageId="4" pageNumber="101" rank="subSpecies" species="mystaceus" subSpecies="khorasanus">Ph. mystaceus khorasanus</taxonomicName>
|
||
ssp. n. in preservative: A dorsal view B ventral view C head in dorsal view D head in frontal view E head in lateral view; F right foot in thenar view (photographs by E. N. Solovyeva).
|
||
</paragraph>
|
||
</caption>
|
||
</subSubSection>
|
||
<subSubSection id="FAD6B4F91CDB54F96F6C4D888264C14B" pageId="4" pageNumber="101" type="color of holotype in life">
|
||
<paragraph id="6FA7F5742A9D86C0386EC49FEE8B550C" pageId="4" pageNumber="101">Color of holotype in life.</paragraph>
|
||
<paragraph id="DF29080E0B9D2C5B9827769774FB055D" pageId="4" pageNumber="101">In life dorsum sandy-beige; with numerous small black and white dots and reticulations; row of three pairs of irregular-shaped larger dark blotches on each side of vertebral line; ventral surfaces of body, limbs and proximal part of tail white; ventral surface of tail tip black, chin and throat with gray reticulations, chest with blackish longitudinal blotch. Ten brownish transverse bars (wider than interspaces) on dorsal surface of tail, faint at tail basis, get more distinct towards tail tip. Internal surfaces of mouth angle cutaneous flaps in life are pinkish, and may become red when animal displays a threatening posture.</paragraph>
|
||
</subSubSection>
|
||
<subSubSection id="F23E94E5C9E518FD6FB6B24F9EC7FFF6" pageId="4" pageNumber="101" type="color of holotype in preservative">
|
||
<paragraph id="2998CF611FD3FDA7D8004E0E55470CDD" pageId="4" pageNumber="101">Color of holotype in preservative.</paragraph>
|
||
<paragraph id="AF0882E07AAFB06AFD681DADEBBF1CC8" pageId="4" pageNumber="101">
|
||
In preservative, numerous dark spots and mottling are distinct on dull sandy-gray background color of dorsum. They form vermiculate patterns ca. 1-2 scales wide. On lateral parts of dorsum these lines form 6-7 indistinct dark transverse bands. Ten dark transverse bars on dorsal side of tail are well-distinct (Fig. 7A). Three posterior dark bars have a distinct light-beige longitudinal line between them along midline of tail. Tail ventral surface light yellowish-white. Ventral surface of head with distinctive dark greyish marbling (Fig. 7A). Distinct triangular longitudinal black spot in the middle of chest area resembling a
|
||
<normalizedToken id="2574D72346C199385EC25EC388283288" originalValue="“necktie”">"necktie"</normalizedToken>
|
||
, ca. 8.8 mm in length. Black coloration of distal part of ventral surface of tail 24 mm in length.
|
||
</paragraph>
|
||
</subSubSection>
|
||
<subSubSection id="5EE4E79A22A30B20A223837524DC696B" lastPageId="5" lastPageNumber="102" pageId="4" pageNumber="101" type="paratype variation">
|
||
<paragraph id="3685B4B685002695141930AAD1058CC0" pageId="4" pageNumber="101">Paratype variation.</paragraph>
|
||
<paragraph id="94CD2496DDDAF729A3606AD6E282343D" lastPageId="5" lastPageNumber="102" pageId="4" pageNumber="101">
|
||
Variations of morphological characteristics in the type series are shown in Table 4 and in Fig. 8. In general, morphology of paratypes corresponds well to morphology of the holotype. SVL of new subspecies varies in range of 85.0-86.0 mm in two males, and in range of 54.0-70.0 in five females; tail length 76.0 mm in males, 51.0-67.0 mm in females; tail comparatively shorter in male specimens (SVL/TL ratio 1.12-1.13) than in females (SVL/TL ratio 1.00-1.06); however, the sample size is too small to detect significant differences. Length of dark distal part of ventral surface of tail varies from 20 to 27 mm. Number of subdigital lamellae on toe III varies from 17 to 20, from 25 to 28 on toe IV. Number of enlarged triangular scales of lateral fringes on toe III from 7 to 11, on toe IV from 18 to 21. Number of flattened anterior supralabials 6-11, total number of supralabials (to insertion of cutaneous fold at mouth angle) varies from 10 to 15. Number of small scales ventrally in contact with subnasal scale 3-6. Subnasal scale in all paratypes (except one specimen ZMMU R-13009) touches supranasal along medial edge of latter. In nearly all paratypes supralabials are separated from subnasal by five rows of small scales (only in ZMMU R-13009 by 4/5 rows of small scales). In most specimens, there is one small scale between last supralabial and insertion of cutaneous fold at mouth angle (specimen ZMMU R-13011 has two scales, ZMMU R-13169 lacks such scales). Number
|
||
<pageBreakToken id="F9D2E053BDF58FE1E6456ABD14705504" pageId="5" pageNumber="102" start="start">of</pageBreakToken>
|
||
flat anterior infralabials varies from 2 to 4, total number of infralabials to insertion of cutaneous fold at mouth angle varies from 5 to 7 (only ZMMU R-13009 has 3/3 infralabials). Number of black irregularly shaped spots on dorsum also may vary: from 4 to 6 pairs of black spots on each side of vertebral line (see Fig. 8A).
|
||
</paragraph>
|
||
<caption id="328D07722A0890F157EBAA0881940604" pageId="5" pageNumber="102">
|
||
<paragraph id="1DA5C8C8505786B31E8234AAC7F7650C" pageId="5" pageNumber="102">
|
||
Figure 8. Paratypes of
|
||
<taxonomicName id="76514196119B3E1B060D52F3965B2BD9" lsidName="Ph. mystaceus" pageId="5" pageNumber="102" rank="subSpecies" species="mystaceus" subSpecies="khorasanus">Ph. mystaceus khorasanus</taxonomicName>
|
||
ssp. n. in preservative: A in dorsal view B in ventral view (photographs by E. N. Solovyeva).
|
||
</paragraph>
|
||
</caption>
|
||
<paragraph id="9F3C1FFF3B931E6EFE29DEBA93D4169B" pageId="5" pageNumber="102">
|
||
We were unable to detect sexual dimorphism in morphometric and meristic characteristics of
|
||
<taxonomicName id="9BA5065F9202F4A7C804C08A1F744C01" lsidName="Ph. mystaceus" pageId="5" pageNumber="102" rank="subSpecies" species="mystaceus" subSpecies="khorasanus">Ph. mystaceus khorasanus</taxonomicName>
|
||
ssp. n., however our sample size (N = 7) was too small.
|
||
<bibRefCitation id="8991FD152791AC07E3F95A570373EC00" author="Molavi, F" journalOrPublisher="Check List" pageId="9" pageNumber="106" pagination="450 - 452" title="New distribution Record of Phrynocephalusmystaceus Pallas 1776 (Sauria: Agamidae) from Iran." volume="10" year="2014">Molavi et al. (2014)</bibRefCitation>
|
||
, who also examined seven specimens of both sexes from Semnan Province, was also unable to detect sexual dimorphism in morphological features in their sample.
|
||
</paragraph>
|
||
</subSubSection>
|
||
<subSubSection id="16609FBC429C97DB2DBE2C467829C9F1" lastPageId="6" lastPageNumber="103" pageId="5" pageNumber="102" type="distribution">
|
||
<paragraph id="FF5FAB017B9478E75DFF2E6E8CC445CC" pageId="5" pageNumber="102">Distribution.</paragraph>
|
||
<paragraph id="FB624E1E84B1871882F7A88EACCAC919" lastPageId="6" lastPageNumber="103" pageId="5" pageNumber="102">
|
||
To date, the new subspecies is known from two major localities in southwestern part of Khorasan Razavi Province (environs of the towns of Gonabad and Boshrouyeh, this study) and from a single locality in the easternmost part of Semnan
|
||
<pageBreakToken id="CF2BA28E4063A7D6D8775CC2ECC9B97D" pageId="6" pageNumber="103" start="start">Province</pageBreakToken>
|
||
of Iran (Ahmad Abad village, Molavi 2014). The record from the environs of the town of Boshrouyeh appears to be the southernmost known locality for
|
||
<taxonomicName id="225C9EF9DDE03BCCF7C4E875065C5A23" lsidName="Ph. mystaceus" pageId="6" pageNumber="103" rank="species" species="mystaceus">Ph. mystaceus</taxonomicName>
|
||
complex known to date. The three records of
|
||
<taxonomicName id="8E669410084E0095308D9B2E5502CF73" lsidName="Ph. mystaceus" pageId="6" pageNumber="103" rank="species" species="mystaceus">Ph. mystaceus</taxonomicName>
|
||
by
|
||
<bibRefCitation id="E65547D0CCF48AA6B7518423176B3C20" author="Anderson, SC" journalOrPublisher="Society for the Study of Amphibians and Reptiles, Ithaca, New York" pageId="8" pageNumber="105" title="The Lizards of Iran." year="1999">Anderson (1999)</bibRefCitation>
|
||
from the northern part of Khorasan Razavi Province, North Khorasan and Golestan provinces are all located along the border with Turkmenistan. These populations most likely correspond to
|
||
<taxonomicName id="074246CD94CBEE66D1BE97933A3701A9" lsidName="Ph. m." pageId="6" pageNumber="103" rank="subSpecies" species="m." subSpecies="mystaceus">Ph. m. mystaceus</taxonomicName>
|
||
rather than to
|
||
<taxonomicName id="870BA92B88333E5293440BC1EA46B9F0" lsidName="Ph. mystaceus" pageId="6" pageNumber="103" rank="subSpecies" species="mystaceus" subSpecies="khorasanus">Ph. mystaceus khorasanus</taxonomicName>
|
||
ssp. n. as they are close to the range of the nominative form and there are no biogeographic barriers that separate these populations. On the contrary, localities in Khorasan Razavi and Semnan provinces are situated on different elevations and sand massifs are isolated from the range of
|
||
<taxonomicName id="979D03FBF5000784154EB2A00E1A1567" lsidName="Ph. m." pageId="6" pageNumber="103" rank="subSpecies" species="m." subSpecies="mystaceus">Ph. m. mystaceus</taxonomicName>
|
||
by at least 200 km of habitats unsuitable for
|
||
<taxonomicName id="6E9E1BC1810FA087D83E516DF4E541B5" lsidName="Ph. mystaceus" pageId="6" pageNumber="103" rank="species" species="mystaceus">Ph. mystaceus</taxonomicName>
|
||
. We anticipate new records of the new subspecies in sandy areas of Khorasan Razavi, Semnan and, possibly, northern part of Yazd and South Khorasan provinces.
|
||
</paragraph>
|
||
</subSubSection>
|
||
<subSubSection id="9813AB24F34ECA7E4062FE141BA5E3F2" pageId="6" pageNumber="103" type="habitat">
|
||
<paragraph id="9CC391ECC8DDD48E93EE00FF1F621A4F" pageId="6" pageNumber="103">Habitat.</paragraph>
|
||
<paragraph id="04F3BE25742E962A83A49A99B5F5AA26" pageId="6" pageNumber="103">
|
||
<taxonomicName id="8BF232E359206AE10C54E4666C3FD4B4" lsidName="Ph. mystaceus" pageId="6" pageNumber="103" rank="subSpecies" species="mystaceus" subSpecies="khorasanus">Ph. mystaceus khorasanus</taxonomicName>
|
||
ssp. n. inhabits sandy areas with sparse vegetation in northeast Iran at comparatively higher altitudes, than other
|
||
<taxonomicName id="8CD5DC3B95F64CDA3222C50EF4F41EA0" lsidName="Ph. mystaceus" pageId="6" pageNumber="103" rank="species" species="mystaceus">Ph. mystaceus</taxonomicName>
|
||
subspecies. The usual habitat is represented by dunes of loose sands and semi-stabilized dunes with rare grass, occasional bushes of
|
||
<taxonomicName id="4EEC2184D6A7484988F039984DE9E1F8" class="Magnoliopsida" family="Amaranthaceae" genus="Haloxylon" higherTaxonomySource="CoL" kingdom="Plantae" lsidName="Haloxylon" order="Caryophyllales" pageId="6" pageNumber="103" phylum="Tracheophyta" rank="genus">Haloxylon</taxonomicName>
|
||
sp. and
|
||
<taxonomicName id="16C9FDF25DD3056F9CB67DFA11214A6D" class="Magnoliopsida" family="Tamaricaceae" genus="Tamarix" higherTaxonomySource="CoL" kingdom="Plantae" lsidName="Tamarix" order="Caryophyllales" pageId="6" pageNumber="103" phylum="Tracheophyta" rank="genus">Tamarix</taxonomicName>
|
||
sp. and large open sandy areas (Fig. 9). The areas inhabited by the new subspecies receive almost no rainfall during the year. In the town of Gonabad the average annual temperature is 17.3 °C, the average temperature in July reaches 29.2 °C, the average temperature in January is 4.8 °C; In Boshrouyeh the average annual temperature is 19.7 °C, the average temperature in July is 31.9 °C, the average temperature in January is 6.6 °C. (http://www.climate-data.org).
|
||
</paragraph>
|
||
<caption id="34A08AE2F9D480266119E056475E7AC0" pageId="6" pageNumber="103">
|
||
<paragraph id="AC995E5DD356CE422402E2E9D7D176B9" pageId="6" pageNumber="103">
|
||
Figure 9. Typical habitat of
|
||
<taxonomicName id="39A9E7F12231E2106418D41FE83ABB67" lsidName="Ph. mystaceus" pageId="6" pageNumber="103" rank="subSpecies" species="mystaceus" subSpecies="khorasanus">Ph. mystaceus khorasanus</taxonomicName>
|
||
ssp. n. at the type locality in the vicinity of Gonabad, Khorasan Razavi Province, Iran (photo by R. A. Nazarov).
|
||
</paragraph>
|
||
</caption>
|
||
<paragraph id="4FC105B4EB3E5A9B3935F77912A137EA" pageId="6" pageNumber="103">Lizards burrow in sand, digging short tunnels and chambers; they can quickly dig into sand by rapid lateral movements of the body (Anderson, 1999).</paragraph>
|
||
</subSubSection>
|
||
<subSubSection id="0E7242285EC54FB68FCE386E1A03D9A1" lastPageId="7" lastPageNumber="104" pageId="6" pageNumber="103" type="comparisons">
|
||
<paragraph id="0E409487526EFB16FBD2C785F3D70CCA" pageId="6" pageNumber="103">Comparisons with other subspecies.</paragraph>
|
||
<paragraph id="B12FC431F3BC414BFA31E92306B1BEAC" pageId="6" pageNumber="103">
|
||
Comparisons of the new subspecies from Khorasan Razavi and Semnan provinces of Iran with the nominative subspecies
|
||
<taxonomicName id="80C4D4CFF02A98CC1A7CBD86FEB89CDD" lsidName="Ph. m." pageId="6" pageNumber="103" rank="subSpecies" species="m." subSpecies="mystaceus">Ph. m. mystaceus</taxonomicName>
|
||
sensu lato from Middle Asia, Caspian basin, and westernmost Xinjiang (China) are summarized below. In preservative, the new subspecies can be differentiated from specimens of
|
||
<taxonomicName id="2063C2324EB404763CBD3FCB4A5DFC62" lsidName="Ph. m." pageId="6" pageNumber="103" rank="subSpecies" species="m." subSpecies="mystaceus">Ph. m. mystaceus</taxonomicName>
|
||
by the following combination of morphological attributes: lower number of subdigital lamellae on the IVth toe (SLIV 25.7 (24-27; N = 7) in vs. 30.2 (25-35; N = 70) in
|
||
<taxonomicName id="0D8ABB701E6CA8BB12705514FE246B15" lsidName="Ph. m." pageId="6" pageNumber="103" rank="subSpecies" species="m." subSpecies="mystaceus">Ph. m. mystaceus</taxonomicName>
|
||
sensu lato); comparatively lower number of supralabials (SL 12.1 (10-14; N = 7) vs. 14.9 (10-19; N = 70) in
|
||
<taxonomicName id="51A5FFF9C3F6D95392F44AB3089D0D4F" lsidName="Ph. m." pageId="6" pageNumber="103" rank="subSpecies" species="m." subSpecies="mystaceus">Ph. m. mystaceus</taxonomicName>
|
||
sensu lato) and by the comparatively shorter black distal part on the tail ventral surface (TL-black/TL 0.38 (0.36-0.40; N = 7) vs. 0.42 (0.32-0.48; N = 70) in
|
||
<taxonomicName id="FFDA325E844417C9C1C8DDC3F98F6464" lsidName="Ph. m." pageId="6" pageNumber="103" rank="subSpecies" species="m." subSpecies="mystaceus">Ph. m. mystaceus</taxonomicName>
|
||
sensu lato). In life, juvenile and young specimens of the new subspecies can be further distinguished from Middle Asian / Caspian Basin populations of
|
||
<taxonomicName id="5450A98E57DB38BB75D57467FD3BDBE2" lsidName="Ph. mystaceus" pageId="6" pageNumber="103" rank="species" species="mystaceus">Ph. mystaceus</taxonomicName>
|
||
by is rusty orange color of the proximal part of tail ventral surface (vs. lemon-yellow in
|
||
<taxonomicName id="C4DBBA1B0A781013CE622B30DDD38DEA" lsidName="Ph. m." pageId="6" pageNumber="103" rank="subSpecies" species="m." subSpecies="mystaceus">Ph. m. mystaceus</taxonomicName>
|
||
sensu stricto), but is similar to orange tail coloration in juveniles of East Kazakhstan - western China populations described as
|
||
<taxonomicName id="698D8E330A7CB29FC19DB0F3EDBB4093" lsidName="Ph. m." pageId="6" pageNumber="103" rank="subSpecies" species="m." subSpecies="aurantiacocaudatus">Ph. m. aurantiacocaudatus</taxonomicName>
|
||
.
|
||
</paragraph>
|
||
<paragraph id="57D835BEEABC768820377D4FB813EB14" lastPageId="7" lastPageNumber="104" pageId="6" pageNumber="103">
|
||
We do not recognize
|
||
<taxonomicName id="87BF4D0AD5462501D18AE4BDFA89881D" lsidName="Ph. m." pageId="6" pageNumber="103" rank="subSpecies" species="m." subSpecies="galli">Ph. m. galli</taxonomicName>
|
||
as a separate subspecies due to the absence of stable genetic and morphological differences of this subspecies from
|
||
<taxonomicName id="48346FEE9448B34CC12D8F4D96E65134" lsidName="Ph. m." pageId="6" pageNumber="103" rank="subSpecies" species="m." subSpecies="mystaceus">Ph. m. mystaceus</taxonomicName>
|
||
(see above). The
|
||
<taxonomicName id="0839B79E286DC109893C7744281FDDB6" class="Squamata" family="Agamidae" genus="Phrynocephalus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Phrynocephalus mystaceus" pageId="6" pageNumber="103" phylum="Chordata" rank="subSpecies" species="mystaceus" subSpecies="dagestanica">Phrynocephalus mystaceus dagestanica</taxonomicName>
|
||
form from Daghestan (
|
||
<bibRefCitation id="8A245F8313E023884D3EAB30218DACD5" pageId="6" pageNumber="103">
|
||
Ananjeva
|
||
<normalizedToken id="255DB39F879EDD1C40122A5F84F636E9" originalValue="“1986”">"1986"</normalizedToken>
|
||
1987
|
||
</bibRefCitation>
|
||
) is very close to the populations from the Volga River basin and was
|
||
<pageBreakToken id="49FFC3E6386D7FEEC96879CC2F5316BE" pageId="7" pageNumber="104" start="start">considered</pageBreakToken>
|
||
a synonym of
|
||
<taxonomicName id="74E5316A2BE19BE96ABF421AC266445C" lsidName="Ph. m." pageId="7" pageNumber="104" rank="subSpecies" species="m." subSpecies="mystaceus">Ph. m. mystaceus</taxonomicName>
|
||
by several authors (
|
||
<bibRefCitation id="C201B135AEF35F1366770A89DFEA690B" author="Semenov, DV" journalOrPublisher="Description of a new subspecies with remarks on the taxonomic status of Phrynocephalusmystaceus (Reptilia, Agamidae) [In Russian, English summary) Zoologicheskiy Zhurnal (USSR)" pageId="9" pageNumber="106" pagination="76 - 81" title="Species of the genus Phrynocephalus of the USSR fauna." volume="69" year="1990">Semenov and Shenbrot 1990</bibRefCitation>
|
||
;
|
||
<bibRefCitation id="7117978FAF4B2B7AFAFCED171A69EB8B" author="Barabanov, AV" journalOrPublisher="Zootaxa" pageId="8" pageNumber="105" pagination="1 - 56" title="Catalogue of the available scientific species-group names for lizards of the genus Phrynocephalus Kaup, 1825 (Reptilia, Sauria, Agamidae)." url="https://doi.org/10.11646/zootaxa.1399.1.1" volume="1399" year="2007">Barabanov and Ananjeva 2007</bibRefCitation>
|
||
). Our molecular and morphometric data do not support monophyly or significant differentiation of
|
||
<taxonomicName id="0EF0B07A303B18757D308B3F7DC87FDE" lsidName="Ph. m." pageId="7" pageNumber="104" rank="subSpecies" species="m." subSpecies="aurantiacocaudatus">Ph. m. aurantiacocaudatus</taxonomicName>
|
||
from Eastern Kazakhstan and western China. The only stable difference between this population and
|
||
<taxonomicName id="662CF6FB5611E1A5EBE89A615C64DA15" lsidName="Ph. m." pageId="7" pageNumber="104" rank="subSpecies" species="m." subSpecies="mystaceus">Ph. m. mystaceus</taxonomicName>
|
||
sensu stricto is the tail coloration in juveniles. We consider that additional genetic and morphological data is needed to clarify taxonomic status of East Kazakhstan
|
||
<taxonomicName id="26863B172393779DAAA3EDF55155FE02" lsidName="Ph. mystaceus" pageId="7" pageNumber="104" rank="species" species="mystaceus">Ph. mystaceus</taxonomicName>
|
||
populations.
|
||
</paragraph>
|
||
</subSubSection>
|
||
<subSubSection id="4F57C9FAB7FA815F934233FC361051E0" lastPageId="8" lastPageNumber="105" pageId="7" pageNumber="104" type="discussion">
|
||
<paragraph id="DD8306543F035B50842271ED88915241" pageId="7" pageNumber="104">Discussion.</paragraph>
|
||
<paragraph id="8D66E39CD2DAC5C72607E4367BF447DA" lastPageId="8" lastPageNumber="105" pageId="7" pageNumber="104">
|
||
Our study indicates deep genetic divergence between Iranian populations of
|
||
<taxonomicName id="140D6B3C4AD6E1778FDCAB3AB88D0029" lsidName="Ph. m." pageId="7" pageNumber="104" rank="subSpecies" species="m." subSpecies="khorasanus">Ph. m. khorasanus</taxonomicName>
|
||
ssp. n. and the rest of the populations within the range of the species. However, morphological differentiation within
|
||
<taxonomicName id="65A24D5F9F8945FF89A48522F6B6FAD5" lsidName="Ph. mystaceus" pageId="7" pageNumber="104" rank="species" species="mystaceus">Ph. mystaceus</taxonomicName>
|
||
complex is less clear with only a few morphological characteristics that reliably separate these two lineages. Differentiation pattern for the mtDNA COI gene within the Middle Asian and Caspian populations of
|
||
<taxonomicName id="5299E8033394172399D30449C56FC6BE" lsidName="Ph. mystaceus" pageId="7" pageNumber="104" rank="species" species="mystaceus">Ph. mystaceus</taxonomicName>
|
||
complex suggests that East Kazakhstan was populated by
|
||
<taxonomicName id="ACECCC29258310D4042F17816B4A0CE9" lsidName="Ph. mystaceus" pageId="7" pageNumber="104" rank="species" species="mystaceus">Ph. mystaceus</taxonomicName>
|
||
earlier than the rest of Middle Asia. After that, a dispersal process from the east to the west likely took place. Morphologically different populations of
|
||
<taxonomicName id="F26CC67AE63F8D015B47E265F2401D19" lsidName="Ph. mystaceus" pageId="7" pageNumber="104" rank="species" species="mystaceus">Ph. mystaceus</taxonomicName>
|
||
across Middle Asia present considerable amount of variation both in body size and in such morphological features as the relative size of cutaneous flaps in the mouth angles, relative tail length, etc. This high morphological plasticity may be connected with psam
|
||
<pageBreakToken id="F232585245F48DA4451F8DED747CF02E" pageId="8" pageNumber="105" start="start">mophilous</pageBreakToken>
|
||
biology of this species, as it was suggested by previous researchers (
|
||
<bibRefCitation id="4984FBF55C0FC980CE8A390CD6C87FB8" author="Vel'dre, SR" journalOrPublisher="Primeneniye matematicheskikh metodov v biologii [Application of mathematical methods in biology], Leningrad" pageId="9" pageNumber="106" pagination="75 - 85" title="On correlational structure of external morphological characters in secret toad-headed agama Phrynocephalusmystaceus." volume="3" year="1964 a">
|
||
<normalizedToken id="5966ACBF6C73F00EC302F6090FFF1216" originalValue="Vel’dre">Vel'dre</normalizedToken>
|
||
1964a
|
||
</bibRefCitation>
|
||
,
|
||
<bibRefCitation id="D2BE68EF8DF313FE3B2BB58FAB143F96" author="Vel'dre, SR" journalOrPublisher="Vestnik Leningardskogo Universiteta [Vestnik of Leningrad University]" pageId="9" pageNumber="106" pagination="34 - 40" title="On reality of subspecies in secret toad-headed agama." volume="3" year="1964 b">1964b</bibRefCitation>
|
||
;
|
||
<bibRefCitation id="C9FF3F0AD4C568BBBDAB55053611A04E" author="Semenov, DV" journalOrPublisher="Description of a new subspecies with remarks on the taxonomic status of Phrynocephalusmystaceus (Reptilia, Agamidae) [In Russian, English summary) Zoologicheskiy Zhurnal (USSR)" pageId="9" pageNumber="106" pagination="76 - 81" title="Species of the genus Phrynocephalus of the USSR fauna." volume="69" year="1990">Semenov and Shenbrot 1990</bibRefCitation>
|
||
;
|
||
<bibRefCitation id="E6D0D338C82414E0D0562A2D7763CA9E" author="Golubev, ML" journalOrPublisher="Vestnik Zoologii, Kiev" pageId="8" pageNumber="105" pagination="26 - 32" title="On Intraspecific Structure and Intraspecific Relations of the Ear-Folded Toad Agama Phrynocephalusmystaceus (Reptilia, Agamidae)." volume="3" year="1992">Golubev and Sattorov 1992</bibRefCitation>
|
||
).
|
||
</paragraph>
|
||
<paragraph id="CA7FF4890652F33346E1F860C1D6F3A0" pageId="8" pageNumber="105">
|
||
The data of phylogenetic analyses in the present paper clearly indicates that the whole territory of Middle Asia, including westernmost China and Caspian region, is inhabited by a single poorly differentiated mtDNA lineage.
|
||
<bibRefCitation id="F8891B5933B1019498BE1FBE015E44E1" author="Golubev, ML" journalOrPublisher="Vestnik Zoologii, Kiev" pageId="8" pageNumber="105" pagination="26 - 32" title="On Intraspecific Structure and Intraspecific Relations of the Ear-Folded Toad Agama Phrynocephalusmystaceus (Reptilia, Agamidae)." volume="3" year="1992">Golubev and Sattorov (1992)</bibRefCitation>
|
||
argued that coloration of the ventral tail surface in juveniles of
|
||
<taxonomicName id="061EF7AB0E8D94F0B10D1AFDCBF43D73" lsidName="Ph. mystaceus" pageId="8" pageNumber="105" rank="species" species="mystaceus">Ph. mystaceus</taxonomicName>
|
||
is also subject to high variation, and
|
||
<normalizedToken id="9E3F6DFCC06C945F5AB6D02358C2BA23" originalValue="“orange-”">"orange-"</normalizedToken>
|
||
and
|
||
<normalizedToken id="A62DA88881998B040F7E44DCCB8EABB3" originalValue="“yellow-tailed”">"yellow-tailed"</normalizedToken>
|
||
specimens can be occasionally recorded within the same population, thus suggesting that subspecies within
|
||
<taxonomicName id="C95BBC005110639A556FD4B424F6B131" lsidName="Ph. mystaceus" pageId="8" pageNumber="105" rank="species" species="mystaceus">Ph. mystaceus</taxonomicName>
|
||
should not be recognized. Our mtDNA genealogy indicates that both
|
||
<taxonomicName id="28A396FF7557AF139C97DC1687D7B623" lsidName="Ph. m." pageId="8" pageNumber="105" rank="species" species="m.">Ph. m.</taxonomicName>
|
||
"
|
||
<taxonomicName id="416B2F76A827093FE1C56C3A30B730F9" lsidName="" pageId="8" pageNumber="105" rank="subSpecies" subSpecies="galli">galli</taxonomicName>
|
||
" and
|
||
<taxonomicName id="67D57368FE78D338743E7DAF9C93BFF0" lsidName="Ph. m." pageId="8" pageNumber="105" rank="species" species="m.">Ph. m.</taxonomicName>
|
||
"
|
||
<taxonomicName id="D6AA6772F2221CF338DE37C1168F508E" lsidName="" pageId="8" pageNumber="105" rank="subSpecies" subSpecies="aurantiacocaudatus">aurantiacocaudatus</taxonomicName>
|
||
" do not form a respective monophyletic units and are genetically indistinguishable or very close the nominative subspecies
|
||
<taxonomicName id="12BF905D76EDF8A8F043C33FEE50CE78" lsidName="P. m." pageId="8" pageNumber="105" rank="subSpecies" species="m." subSpecies="mystaceus">P. m. mystaceus</taxonomicName>
|
||
sensu stricto (p-distance 1.65-1.87% in case of East Kazakhstan populations).
|
||
</paragraph>
|
||
<paragraph id="539EBAFEC27C6333C0DB0DA2BDB62A25" pageId="8" pageNumber="105">
|
||
On the contrary, the Khorasan population described herein as
|
||
<taxonomicName id="2C57E5D574B1F98FE19E62CAB1666C34" lsidName="Ph. m." pageId="8" pageNumber="105" rank="subSpecies" species="m." subSpecies="khorasanus">Ph. m. khorasanus</taxonomicName>
|
||
shows very deep genetic divergence in mtDNA which is comparable to the species-level divergence in
|
||
<taxonomicName id="C3F1E6E9EA86ED2C0692E6FB6381D5A6" class="Squamata" family="Agamidae" genus="Phrynocephalus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Phrynocephalus" pageId="8" pageNumber="105" phylum="Chordata" rank="genus">Phrynocephalus</taxonomicName>
|
||
, but is only moderately differentiated morphologically. Indeed, previous research on four mtDNA genes also showed significant differentiation between
|
||
<taxonomicName id="FFD11D292C02CEBA8C2F5A603170D83C" lsidName="Ph. mystaceus" pageId="8" pageNumber="105" rank="species" species="mystaceus">Ph. mystaceus</taxonomicName>
|
||
from Khorasan and
|
||
<taxonomicName id="43CE796E08C74907518DD51EF8322248" lsidName="Ph. m." pageId="8" pageNumber="105" rank="subSpecies" species="m." subSpecies="mystaceus">Ph. m. mystaceus</taxonomicName>
|
||
(p-distances: COI - 7.18%; ND4 - 6.6%; ND2 - 8.0%; and cyt b - 6.6%) (see
|
||
<bibRefCitation id="8775692BC2D998A7856584CCBFB8C92D" author="Solovyeva, EN" journalOrPublisher="Doklady Biological Sciences" pageId="9" pageNumber="106" pagination="119 - 124" title="Phylogenetic relationships and subgeneric taxonomy of toad-headed agamas Phrynocephalus (Reptilia, Squamata, Agamidae) based on mitochondrial DNA sequence data." volume="455" year="2014">Solovyeva et al. 2014</bibRefCitation>
|
||
). According to our unpublished data on molecular dating of 4 mtDNA genes these two forms diverged during Pliocene about 3.7 Ma (Solovyeva et al., 2018). Further studies are required to verify the taxonomic status of
|
||
<taxonomicName id="669A395DD8C467C2C77CE1D7ABDA40BA" lsidName="Ph. m." pageId="8" pageNumber="105" rank="subSpecies" species="m." subSpecies="khorasanus">Ph. m. khorasanus</taxonomicName>
|
||
ssp. n., including morphological examination of larger samples and molecular analysis of the nuclear DNA markers in order to check the presence of possible isolation between the Iranian and Middle Asian forms of
|
||
<taxonomicName id="37BC76F9EE5885C7CF87AC5EAE82E9FE" lsidName="Ph. mystaceus" pageId="8" pageNumber="105" rank="species" species="mystaceus">Ph. mystaceus</taxonomicName>
|
||
. The new subspecies inhabits sand dunes in the northeastern Iran; this desert area is separated from the range of
|
||
<taxonomicName id="25715C88C53E1577F78C40DF94E5F70E" lsidName="Ph. m." pageId="8" pageNumber="105" rank="subSpecies" species="m." subSpecies="mystaceus">Ph. m. mystaceus</taxonomicName>
|
||
by Kopet-Dagh Mountain Ridge making the possibility of gene flow between these populations quite low. However, the taxonomic status of
|
||
<taxonomicName id="F9092A40D27DAE188F15A7552CA48E30" lsidName="Ph. mystaceus" pageId="8" pageNumber="105" rank="species" species="mystaceus">Ph. mystaceus</taxonomicName>
|
||
populations reported by
|
||
<bibRefCitation id="344AE9FACAFE08BB742A2269334FB9D7" author="Anderson, SC" journalOrPublisher="Society for the Study of Amphibians and Reptiles, Ithaca, New York" pageId="8" pageNumber="105" title="The Lizards of Iran." year="1999">Anderson (1999)</bibRefCitation>
|
||
from northern Iran (northern parts of Golestan, North Khorasan and Khorasan Razavi provinces) is unclear and require verification. Additional fieldwork in northern Iran, western Afghanistan, and southern Middle Asia is required to recover new populations of
|
||
<taxonomicName id="2B7F27D2CCA2BBC24CB817A548FF2769" lsidName="Ph. mystaceus" pageId="8" pageNumber="105" rank="species" species="mystaceus">Ph. mystaceus</taxonomicName>
|
||
complex. Further progress in understanding of the phylogenetic relationships within
|
||
<taxonomicName id="EECF01AA39380A2BDDA95C72936C85EF" lsidName="Ph. mystaceus" pageId="8" pageNumber="105" rank="species" species="mystaceus">Ph. mystaceus</taxonomicName>
|
||
complex might lead to reconsideration of the taxonomic status of the Khorasan population as a full species.
|
||
</paragraph>
|
||
<caption id="C4CAB56E4CCF673F47C78CCFEBB852EE" pageId="8" pageNumber="105">
|
||
<paragraph id="BE140BF62A6711CCC5F7B97EC4091F65" pageId="8" pageNumber="105">
|
||
Figure 10. ZMMU R-6413, lectotype of
|
||
<taxonomicName id="A8012EDED74837F869AA27572D813C06" genus="Phrynocepahlus" lsidName="Phrynocepahlus mystaceus" pageId="8" pageNumber="105" rank="subSpecies" species="mystaceus" subSpecies="galli">Phrynocepahlus mystaceus galli</taxonomicName>
|
||
Krassowsky, 1932 in preservative: A dorsal view B ventral view C head in dorsal view D head in frontal view E head in lateral view F left foot in thenar view (photographs by E. N. Solovyeva).
|
||
</paragraph>
|
||
</caption>
|
||
<caption id="CAB948EA58BF060C44DFFC83E30A64E8" pageId="8" pageNumber="105">
|
||
<paragraph id="1A67096C09CAB32A46C6FAA07B2013DF" pageId="8" pageNumber="105">
|
||
Figure 11. ZMMU R-6412, holotype of
|
||
<taxonomicName id="E0CB6921321E61B50FB1EEC930B3770F" genus="Phrynocepahlus" lsidName="Phrynocepahlus mystaceus" pageId="8" pageNumber="105" rank="subSpecies" species="mystaceus" subSpecies="aurantiacocaudatus">Phrynocepahlus mystaceus aurantiacocaudatus</taxonomicName>
|
||
Semenov & Shenbrot, 1990 in preservative: A dorsal view B ventral view C head in dorsal view D head in frontal view E head in lateral view F right foot in thenar view (photographs by E. N. Solovyeva).
|
||
</paragraph>
|
||
</caption>
|
||
</subSubSection>
|
||
</treatment>
|
||
</document> |