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<document id="FAE9234DE102B4A042A5BD10845D0FDC" ID-CLB-Dataset="298715" ID-DOI="10.11646/zootaxa.5471.2.1" ID-GBIF-Dataset="3a469638-dc21-45a3-80f6-cb307fbf5941" ID-ISSN="1175-5326" ID-Zenodo-Dep="12189904" ID-ZooBank="BAA047F4-2D65-4E52-8798-D426E3B26DA1" IM.bibliography_approvedBy="juliana" IM.illustrations_approvedBy="juliana" IM.materialsCitations_approvedBy="juliana" IM.metadata_approvedBy="juliana" IM.tables_approvedBy="juliana" IM.taxonomicNames_approvedBy="juliana" IM.treatments_approvedBy="juliana" checkinTime="1718892848818" checkinUser="plazi" docAuthor="Kosterin, Oleg E., Onishko, Vladimir V., Ilyina, Elena V., Chepurnov, Grigory Yu. &amp; Blinov, Alexander G." docDate="2024" docId="DB0F87E19131F13DA7D3FC9079B07C7B" docLanguage="en" docName="zootaxa.5471.2.1.pdf" docOrigin="Zootaxa 5471 (2)" docSource="http://dx.doi.org/10.11646/zootaxa.5471.2.1" docStyle="DocumentStyle:647186512141C8FC8976D5BCC54AEB7D.9:Zootaxa.2013-.journal_article" docStyleId="647186512141C8FC8976D5BCC54AEB7D" docStyleName="Zootaxa.2013-.journal_article" docStyleVersion="9" docTitle="Coenagrion ponticum" docType="treatment" docVersion="4" lastPageNumber="186" masterDocId="2736FF999111F11EA744F92E7A657E46" masterDocTitle="The genus Coenagrion Kirby, 1890 (Odonata: Coenagrionidae) in the Russian part of the Caucasus" masterLastPageNumber="190" masterPageNumber="151" pageNumber="183" updateTime="1718988639371" updateUser="ExternalLinkService" zenodo-license-document="CLOSED">
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<mods:title id="A38DD892ECD4FCBC542E561D27C2D152">The genus Coenagrion Kirby, 1890 (Odonata: Coenagrionidae) in the Russian part of the Caucasus</mods:title>
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<mods:namePart id="17D94E1BFAFF2D3D96CBA3ADBF1802C1">Kosterin, Oleg E.</mods:namePart>
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<mods:namePart id="FF69BE59697DCCFBDA8BF963D80011BD">Onishko, Vladimir V.</mods:namePart>
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<mods:affiliation id="4B048264FF29B5588355013F13017832">Moscow Zoo, Department of Herpetology, Bolshaya Gruzinskaya Str. 1, Moscow 123242 Russia.</mods:affiliation>
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<mods:namePart id="7C2A3382DBCAB27030D1B6A50E3F80E1">Ilyina, Elena V.</mods:namePart>
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<mods:affiliation id="D5E9C34DBE2840EB02E414701997B379">Precaspian Institute of Biological Resources of the Daghestan Federal Research Centre of the Russian Academy of Sciences, Dagestanskiy State Nature Reserve, Gagarina Street 120, Makhachkala, Republic of Dagestan, Russia.</mods:affiliation>
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<mods:namePart id="CE9C635531C22611FE95AADBAD024D2B">Chepurnov, Grigory Yu.</mods:namePart>
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<mods:affiliation id="DBFF25FEE05EAD1B9B7C2EE5B082F4F2">Institute of Cytology &amp; Genetics SB RAS, Acad. Lavrentyev ave. 10, Novosibirsk, 630090, Russia. &amp; Institute of Molecular and Cellular Biology SB RAS, Acad. Lavrentyev ave. 8 / 2, Novosibirsk, 630090, Russia.</mods:affiliation>
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<mods:namePart id="22FDC00CA94517789954919A5E028A6D">Blinov, Alexander G.</mods:namePart>
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<mods:affiliation id="61581ED6D326141163A566A4B934EAD9">Institute of Cytology &amp; Genetics SB RAS, Acad. Lavrentyev ave. 10, Novosibirsk, 630090, Russia. &amp; Institute of Molecular and Cellular Biology SB RAS, Acad. Lavrentyev ave. 8 / 2, Novosibirsk, 630090, Russia.</mods:affiliation>
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<treatment id="DB0F87E19131F13DA7D3FC9079B07C7B" ID-DOI="http://doi.org/10.5281/zenodo.12206855" ID-GBIF-Taxon="233309477" ID-Zenodo-Dep="12206855" LSID="urn:lsid:plazi:treatment:DB0F87E19131F13DA7D3FC9079B07C7B" httpUri="http://treatment.plazi.org/id/DB0F87E19131F13DA7D3FC9079B07C7B" lastPageId="35" lastPageNumber="186" pageId="32" pageNumber="183">
<subSubSection id="1BBC657C9131F13EA7D3FC907BC57B9E" box="[151,416,1470,1496]" pageId="32" pageNumber="183" type="nomenclature">
<paragraph id="531936F79131F13EA7D3FC907BC57B9E" blockId="32.[151,416,1470,1496]" box="[151,416,1470,1496]" pageId="32" pageNumber="183">
<heading id="0851819B9131F13EA7D3FC907BC57B9E" bold="true" box="[151,416,1470,1496]" fontSize="11" level="1" pageId="32" pageNumber="183" reason="1">
<taxonomicName id="94A64D749131F13EA7D3FC907BC57B9E" ID-CoL="WTXX" authority="(Bartenef, 1929)" baseAuthorityName="Bartenef" baseAuthorityYear="1929" box="[151,416,1470,1496]" class="Insecta" family="Coenagrionidae" genus="Coenagrion" kingdom="Animalia" order="Odonata" pageId="32" pageNumber="183" phylum="Arthropoda" rank="species" species="ponticum">
<emphasis id="61D2EAE59131F13EA7D3FC907BC57B9E" bold="true" box="[151,416,1470,1496]" italics="true" pageId="32" pageNumber="183">Coenagrion ponticum</emphasis>
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<subSubSection id="1BBC657C9131F13FA7D3FF28781F7F5B" lastPageId="33" lastPageNumber="184" pageId="32" pageNumber="183" type="reference_group">
<paragraph id="531936F79131F13EA7D3FF287FE1782E" blockId="32.[151,1437,1542,2037]" pageId="32" pageNumber="183">
<taxonomicName id="94A64D749131F13EA7D3FF287B367866" ID-CoL="9WYFM" authorityName="Kosterin &amp; Onishko &amp; Ilyina &amp; Chepurnov &amp; Blinov" authorityYear="2024" box="[151,339,1542,1568]" class="Insecta" family="Coenagrionidae" genus="Agrion" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="32" pageNumber="183" phylum="Arthropoda" rank="species" species="ponticum">
<emphasis id="61D2EAE59131F13EA7D3FF287B367866" box="[151,339,1542,1568]" italics="true" pageId="32" pageNumber="183">Agrion ponticum</emphasis>
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was described by A.N.
<bibRefCitation id="37374B069131F13EA51DFF28796F7866" author="Bartenef, A. N." box="[601,778,1542,1568]" pageId="32" pageNumber="183" pagination="54 - 68" refId="ref27207" refString="Bartenef, A. N. (1929) Neue Arten und Varietaten der Odonata des West Kaukasus. Zoologischer Anzeiger, 85, 54 - 68." type="journal article" year="1929">Bartenef (1929)</bibRefCitation>
(the surname spelling as provided therein) in German, next year he reproduced the description in Russian, again using sp. n. (
<bibRefCitation id="37374B069131F13EA4CEFF047E557802" author="Bartenef, A. N." box="[906,1072,1578,1604]" pageId="32" pageNumber="183" pagination="1 - 138" refId="ref27235" refString="Bartenef, A. N. (1930) Materialy k poznaniyu Zapadnogo Kavkaza v odonatologicheskom otnoshenii [Materials for the knowledge on West Caucasus in odonatological respect]. Trudy Zapadnokavkazskoi assotsiatsii nauchno-issledovatel'skikh institutov [Proceedings of the West Caucasian Association of scientific Research Institutes], 72 (14), 1 - 138. [in Russian, with German summary but without German title]" type="journal article" year="1930">Bartenef 1930</bibRefCitation>
). The same way and in the same papers he described three other taxa of
<taxonomicName id="94A64D749131F13EA50AFF6078C8782E" box="[590,685,1614,1640]" class="Magnoliopsida" higherTaxonomySource="GBIF" kingdom="Plantae" order="Odonata" pageId="32" pageNumber="183" phylum="Tracheophyta" rank="order">Odonata</taxonomicName>
from West Caucasus, as considered in detail by
<bibRefCitation id="37374B069131F13EA38FFF607FE5782E" author="Kosterin, O. E." box="[1227,1408,1614,1640]" pageId="32" pageNumber="183" pagination="89 - 126" refId="ref28414" refString="Kosterin, O. E. (2023) Reconsideration of the three Odonata taxa described by A. N. Bartenev from the same place in West Caucasus. Odonatologica, 52 (1 / 2), 89 - 126. https: // doi. org / 10.60024 / odon. v 52 i 1 - 2. a 7" type="journal article" year="2023">Kosterin (2023)</bibRefCitation>
.
</paragraph>
<paragraph id="531936F79131F13FA783FF5C781F7F5B" blockId="32.[151,1437,1542,2037]" lastBlockId="33.[151,1437,151,2040]" lastPageId="33" lastPageNumber="184" pageId="32" pageNumber="183">
The type series consisted of numerous specimens collected in 19271929, from mid-June to early September, by Bartenev himself.
<bibRefCitation id="37374B069131F13EA6C2FFB8785278F6" author="Bartenef, A. N." box="[390,567,1686,1712]" pageId="32" pageNumber="183" pagination="54 - 68" refId="ref27207" refString="Bartenef, A. N. (1929) Neue Arten und Varietaten der Odonata des West Kaukasus. Zoologischer Anzeiger, 85, 54 - 68." type="journal article" year="1929">Bartenef (1929)</bibRefCitation>
characterised the type series origin very briefly and without further comment: “Tuapse, Sotschi, Adler, Pitzunda, Ötshimshiri, Poti, Natanebi, Kobulety, Anfang VI. bis 6.IX. (1926—1928).” The number of specimens was not mentioned. All the mentioned cities/places are at the Black Sea coast of the Caucasus. The later paper in Russian (
<bibRefCitation id="37374B069131F13EA505FE2C796B795B" author="Bartenef, A. N." box="[577,782,1794,1821]" pageId="32" pageNumber="183" pagination="1 - 138" refId="ref27235" refString="Bartenef, A. N. (1930) Materialy k poznaniyu Zapadnogo Kavkaza v odonatologicheskom otnoshenii [Materials for the knowledge on West Caucasus in odonatological respect]. Trudy Zapadnokavkazskoi assotsiatsii nauchno-issledovatel'skikh institutov [Proceedings of the West Caucasian Association of scientific Research Institutes], 72 (14), 1 - 138. [in Russian, with German summary but without German title]" type="journal article" year="1930">Bartenef 1930: 65</bibRefCitation>
) provides the provenance of the type series in more detail. It is said to originate from 15 localities (sometimes cited with some altered spelling than in 1929): seven currently in Krasnodarskiy Kray (one in Tuapse, one in Sochi and five in the Adler environs), four in
<collectingRegion id="9162F8159131F13EA32AFE647EBC7922" box="[1134,1241,1866,1892]" country="Georgia" name="Abkhazia" pageId="32" pageNumber="183">Abkhazia</collectingRegion>
(two in Pitsunda, one in Ochemchiri, one at Lake Inktit), one in
<collectingRegion id="9162F8159131F13EA5DDFE4079D079CE" box="[665,949,1902,1928]" country="Georgia" name="Samegrelo and Zemo Svaneti" pageId="32" pageNumber="183">Samegrelo-Zemo Svaneti</collectingRegion>
(Poti), one in
<collectingRegion id="9162F8159131F13EA317FE407EF679CE" box="[1107,1171,1902,1928]" country="Georgia" name="Guria" pageId="32" pageNumber="183">Guria</collectingRegion>
(Natanebi, this actually is a river) and two in
<collectingRegion id="9162F8159131F13EA6CDFEBC7BB079EA" box="[393,469,1938,1964]" country="Georgia" name="Ajaria" pageId="32" pageNumber="183">Adjara</collectingRegion>
(Kobuleti). The exact numbers of specimens are not provided again, instead there are such annotations as “many specimens”, “medium number of specimens”, “not many specimens”, “few specimens”, “several specimens”, “very few specimens”, “
<specimenCount id="45A0FD7E9131F13EA5DCFEF4788579B2" box="[664,736,2010,2036]" count="1" pageId="32" pageNumber="183" type="male">1 male</specimenCount>
(only
<specimenCount id="45A0FD7E9131F13EA46CFEF579F279B2" box="[808,919,2010,2037]" count="1" pageId="32" pageNumber="183" type="male">one male</specimenCount>
!)”, but the total number of collected specimens seems to be enormous. As for overwhelming majority of Bartenevs taxa (
<bibRefCitation id="37374B069130F13FA4A1F9B97E817EF7" author="Medvedev, A. F. &amp; Kosterin, O. E. &amp; Malikova, E. I. &amp; Schneider, W." box="[997,1252,151,177]" pageId="33" pageNumber="184" pagination="211 - 228" refId="ref28859" refString="Medvedev, A. F., Kosterin, O. E., Malikova, E. I. &amp; Schneider, W. (2013) Description date of Somatochlora exuberata Bartenev, Leucorrhinia intermedia Bartenev and Sympetrum vulgatum grandis Bartenev, the fate of A. N. Bartenev's type specimens and designation of the lectotype of L. intermedia (Anisoptera: Corduliidae, Libellulidae). Odonatologica, 42, 211 - 228." type="journal article" year="2013">
Medvedev
<emphasis id="61D2EAE59130F13FA321F9B97EFA7EF7" box="[1125,1183,151,177]" italics="true" pageId="33" pageNumber="184">et al.</emphasis>
2013
</bibRefCitation>
), the type series most probably no longer exists. Since it consisted of a long series of
<typeStatus id="8C1D88559130F13FA4FDF9957E7E7E93" box="[953,1051,187,213]" pageId="33" pageNumber="184" type="syntype">syntypes</typeStatus>
as the name bearing-type, all the localities enumerated above comprise the vast type locality of the species. It could be narrowed by a
<typeStatus id="8C1D88559130F13FA200F9F17FF87EBF" box="[1348,1437,223,249]" pageId="33" pageNumber="184" type="neotype">neotype</typeStatus>
designation but we see no necessity in this.
</paragraph>
</subSubSection>
<subSubSection id="1BBC657C9130F13DA783F80979B07C7B" lastPageId="35" lastPageNumber="186" pageId="33" pageNumber="184" type="description">
<paragraph id="531936F79130F13FA783F8097AA17CEF" blockId="33.[151,1437,151,2040]" pageId="33" pageNumber="184">
The original (that is published first) description (
<bibRefCitation id="37374B069130F13FA5A6F80979E17F07" author="Bartenef, A. N." box="[738,900,295,321]" pageId="33" pageNumber="184" pagination="54 - 68" refId="ref27207" refString="Bartenef, A. N. (1929) Neue Arten und Varietaten der Odonata des West Kaukasus. Zoologischer Anzeiger, 85, 54 - 68." type="journal article" year="1929">Bartenef 1929</bibRefCitation>
) characterised the range of the new species very briefly. Bartenev reported that “observations show that on the coast of the Black Sea from Tuapse (Gelendzhik?) to Batum are inhabited by only
<taxonomicName id="94A64D749130F13FA693F84178F67FCE" authorityName="Kosterin &amp; Onishko &amp; Ilyina &amp; Chepurnov &amp; Blinov" authorityYear="2024" box="[471,659,367,393]" class="Insecta" family="Coenagrionidae" genus="Agrion" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="33" pageNumber="184" phylum="Arthropoda" rank="species" species="ponticum">
<emphasis id="61D2EAE59130F13FA693F84178F67FCE" box="[471,659,367,393]" italics="true" pageId="33" pageNumber="184">Agrion ponticum</emphasis>
</taxonomicName>
(but neither by
<taxonomicName id="94A64D749130F13FA405F84179947FCF" baseAuthorityName="Vander Linden" baseAuthorityYear="1825" box="[833,1009,367,393]" class="Insecta" family="Coenagrionidae" genus="Coenagrion" kingdom="Animalia" order="Odonata" pageId="33" pageNumber="184" phylum="Arthropoda" rank="species" species="pulchellum">
<emphasis id="61D2EAE59130F13FA405F84179947FCF" box="[833,1009,367,393]" italics="true" pageId="33" pageNumber="184">Agr. pulchellum</emphasis>
</taxonomicName>
nor
<taxonomicName id="94A64D749130F13FA366F8417EF97FCF" baseAuthorityName="Linnaeus" baseAuthorityYear="1758" box="[1058,1180,367,393]" class="Insecta" family="Coenagrionidae" genus="Coenagrion" kingdom="Animalia" order="Odonata" pageId="33" pageNumber="184" phylum="Arthropoda" rank="species" species="puella">
<emphasis id="61D2EAE59130F13FA366F8417EF97FCF" box="[1058,1180,367,393]" italics="true" pageId="33" pageNumber="184">Agr. puella</emphasis>
</taxonomicName>
). All previous findings of the latter two species … actually belonged to the species
<taxonomicName id="94A64D749130F13FA415F8BD79957FEA" authorityName="Kosterin &amp; Onishko &amp; Ilyina &amp; Chepurnov &amp; Blinov" authorityYear="2024" box="[849,1008,403,429]" class="Insecta" family="Coenagrionidae" genus="Agrion" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="33" pageNumber="184" phylum="Arthropoda" rank="species" species="ponticum">
<emphasis id="61D2EAE59130F13FA415F8BD79957FEA" box="[849,1008,403,429]" italics="true" pageId="33" pageNumber="184">Agr. ponticum</emphasis>
</taxonomicName>
” (translated from German) (
<bibRefCitation id="37374B069130F13FA27EF8BD7A9A7F97" author="Bartenef, A. N." pageId="33" pageNumber="184" pagination="54 - 68" refId="ref27207" refString="Bartenef, A. N. (1929) Neue Arten und Varietaten der Odonata des West Kaukasus. Zoologischer Anzeiger, 85, 54 - 68." type="journal article" year="1929">Bartenef 1929: 64</bibRefCitation>
). “Widespread on the Caucasian coast of the Black Sea. Not present in the mountains.” (
<bibRefCitation id="37374B069130F13FA3B6F8997AD17FB3" author="Bartenef, A. N." pageId="33" pageNumber="184" pagination="54 - 68" refId="ref27207" refString="Bartenef, A. N. (1929) Neue Arten und Varietaten der Odonata des West Kaukasus. Zoologischer Anzeiger, 85, 54 - 68." type="journal article" year="1929">Bartenef 1929: 68</bibRefCitation>
) (Our finding of the species in Goryachiy Klyuch Ditrict, see above, disproved the latter statement).
<bibRefCitation id="37374B069130F13FA278F8F57A877C5F" author="Bartenef, A. N." pageId="33" pageNumber="184" pagination="1 - 138" refId="ref27235" refString="Bartenef, A. N. (1930) Materialy k poznaniyu Zapadnogo Kavkaza v odonatologicheskom otnoshenii [Materials for the knowledge on West Caucasus in odonatological respect]. Trudy Zapadnokavkazskoi assotsiatsii nauchno-issledovatel'skikh institutov [Proceedings of the West Caucasian Association of scientific Research Institutes], 72 (14), 1 - 138. [in Russian, with German summary but without German title]" type="journal article" year="1930">Bartenef (1930)</bibRefCitation>
formulated these statements in somewhat different way—that the species is very common in the Pontic District, comprises there an overwhelming majority of
<taxonomicName id="94A64D749130F13FA443FB0D79E87C7B" authorityName="Kirby" authorityYear="1890" box="[775,909,547,573]" class="Insecta" family="Coenagrionidae" genus="Coenagrion" kingdom="Animalia" order="Odonata" pageId="33" pageNumber="184" phylum="Arthropoda" rank="genus">
<emphasis id="61D2EAE59130F13FA443FB0D79E87C7B" box="[775,909,547,573]" italics="true" pageId="33" pageNumber="184">Coenagrion</emphasis>
</taxonomicName>
and had been misidentified as either
<taxonomicName id="94A64D749130F13FA273FB0D7F187C7B" baseAuthorityName="Linnaeus" baseAuthorityYear="1758" box="[1335,1405,547,573]" class="Insecta" family="Coenagrionidae" genus="Coenagrion" kingdom="Animalia" order="Odonata" pageId="33" pageNumber="184" phylum="Arthropoda" rank="species" species="puella">
<emphasis id="61D2EAE59130F13FA273FB0D7F187C7B" box="[1335,1405,547,573]" italics="true" pageId="33" pageNumber="184">puella</emphasis>
</taxonomicName>
or
<taxonomicName id="94A64D749130F13FA7D3FB697B767C27" baseAuthorityName="Vander Linden" baseAuthorityYear="1825" box="[151,275,583,609]" class="Insecta" family="Coenagrionidae" genus="Coenagrion" kingdom="Animalia" order="Odonata" pageId="33" pageNumber="184" phylum="Arthropoda" rank="species" species="pulchellum">
<emphasis id="61D2EAE59130F13FA7D3FB697B767C27" box="[151,275,583,609]" italics="true" pageId="33" pageNumber="184">pulchellum</emphasis>
</taxonomicName>
in earlier reports. The species was registered up to the Tuapse in the north, while “in Abrau, already
<taxonomicName id="94A64D749130F13FA7D3FB457B027CC3" authorityName="Vander Linden" authorityYear="1825" box="[151,359,619,645]" class="Insecta" family="Coenagrionidae" genus="Agrion" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="33" pageNumber="184" phylum="Arthropoda" rank="species" species="pulchellum">
<emphasis id="61D2EAE59130F13FA7D3FB457B027CC3" box="[151,359,619,645]" italics="true" pageId="33" pageNumber="184">Agrion pulchellum</emphasis>
</taxonomicName>
flies, but with an inclination towards
<taxonomicName id="94A64D749130F13FA449FB4579107CC2" authorityName="Kosterin &amp; Onishko &amp; Ilyina &amp; Chepurnov &amp; Blinov" authorityYear="2024" box="[781,885,619,644]" class="Insecta" family="Coenagrionidae" genus="Agrion" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="33" pageNumber="184" phylum="Arthropoda" rank="species" species="ponticum">
<emphasis id="61D2EAE59130F13FA449FB4579107CC2" box="[781,885,619,644]" italics="true" pageId="33" pageNumber="184">ponticum</emphasis>
</taxonomicName>
” [sic!] (translated from Russian) (
<bibRefCitation id="37374B069130F13FA3B7FB457AD27CEF" author="Bartenef, A. N." pageId="33" pageNumber="184" pagination="1 - 138" refId="ref27235" refString="Bartenef, A. N. (1930) Materialy k poznaniyu Zapadnogo Kavkaza v odonatologicheskom otnoshenii [Materials for the knowledge on West Caucasus in odonatological respect]. Trudy Zapadnokavkazskoi assotsiatsii nauchno-issledovatel'skikh institutov [Proceedings of the West Caucasian Association of scientific Research Institutes], 72 (14), 1 - 138. [in Russian, with German summary but without German title]" type="journal article" year="1930">Bartenef 1930: 70</bibRefCitation>
).
</paragraph>
<paragraph id="531936F79130F13FA783FB9C7EF07DE5" blockId="33.[151,1437,151,2040]" pageId="33" pageNumber="184">
Both papers (
<bibRefCitation id="37374B069130F13FA626FB9C786F7C8B" author="Bartenef, A. N." box="[354,522,690,717]" pageId="33" pageNumber="184" pagination="54 - 68" refId="ref27207" refString="Bartenef, A. N. (1929) Neue Arten und Varietaten der Odonata des West Kaukasus. Zoologischer Anzeiger, 85, 54 - 68." type="journal article" year="1929">Bartenef 1929</bibRefCitation>
;
<bibRefCitation id="37374B069130F13FA55DFB9D78377C8B" author="Bartenef, A. N." box="[537,594,691,717]" pageId="33" pageNumber="184" pagination="1 - 138" refId="ref27235" refString="Bartenef, A. N. (1930) Materialy k poznaniyu Zapadnogo Kavkaza v odonatologicheskom otnoshenii [Materials for the knowledge on West Caucasus in odonatological respect]. Trudy Zapadnokavkazskoi assotsiatsii nauchno-issledovatel'skikh institutov [Proceedings of the West Caucasian Association of scientific Research Institutes], 72 (14), 1 - 138. [in Russian, with German summary but without German title]" type="journal article" year="1930">1930</bibRefCitation>
) summarised the characters of the new species in an identical table (see below). The former one (
<bibRefCitation id="37374B069130F13FA6EFFBF878287CB6" author="Bartenef, A. N." box="[427,589,726,752]" pageId="33" pageNumber="184" pagination="54 - 68" refId="ref27207" refString="Bartenef, A. N. (1929) Neue Arten und Varietaten der Odonata des West Kaukasus. Zoologischer Anzeiger, 85, 54 - 68." type="journal article" year="1929">Bartenef 1929</bibRefCitation>
) did not discuss the species characters beyond the table, while the latter one (
<bibRefCitation id="37374B069130F13FA7DBFBD47B247D52" author="Bartenef, A. N." box="[159,321,762,788]" pageId="33" pageNumber="184" pagination="1 - 138" refId="ref27235" refString="Bartenef, A. N. (1930) Materialy k poznaniyu Zapadnogo Kavkaza v odonatologicheskom otnoshenii [Materials for the knowledge on West Caucasus in odonatological respect]. Trudy Zapadnokavkazskoi assotsiatsii nauchno-issledovatel'skikh institutov [Proceedings of the West Caucasian Association of scientific Research Institutes], 72 (14), 1 - 138. [in Russian, with German summary but without German title]" type="journal article" year="1930">Bartenef 1930</bibRefCitation>
) claimed the species to be “intermediate in its characters between
<taxonomicName id="94A64D749130F13FA368FBD47EC27D52" baseAuthorityName="Linnaeus" baseAuthorityYear="1758" box="[1068,1191,762,788]" class="Insecta" family="Coenagrionidae" genus="Coenagrion" kingdom="Animalia" order="Odonata" pageId="33" pageNumber="184" phylum="Arthropoda" rank="species" species="puella">
<emphasis id="61D2EAE59130F13FA368FBD47EC27D52" box="[1068,1191,762,788]" italics="true" pageId="33" pageNumber="184">Agr. puella</emphasis>
</taxonomicName>
and
<taxonomicName id="94A64D749130F13FA39AFBD47FEA7D52" authority="(Bartenef 1930: 66)" authorityName="Vander Linden" authorityYear="1825" baseAuthorityName="Bartenef" baseAuthorityPageNumber="66" baseAuthorityYear="1930" box="[1246,1423,762,788]" class="Insecta" family="Coenagrionidae" genus="Agrion" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="33" pageNumber="184" phylum="Arthropoda" rank="species" species="pulchellum">
<emphasis id="61D2EAE59130F13FA39AFBD47FEA7D52" box="[1246,1423,762,788]" italics="true" pageId="33" pageNumber="184">Agr. pulchellum</emphasis>
</taxonomicName>
” (
<bibRefCitation id="37374B069130F13FA7DBFA307B087D71" author="Bartenef, A. N." box="[159,365,797,824]" pageId="33" pageNumber="184" pagination="1 - 138" refId="ref27235" refString="Bartenef, A. N. (1930) Materialy k poznaniyu Zapadnogo Kavkaza v odonatologicheskom otnoshenii [Materials for the knowledge on West Caucasus in odonatological respect]. Trudy Zapadnokavkazskoi assotsiatsii nauchno-issledovatel'skikh institutov [Proceedings of the West Caucasian Association of scientific Research Institutes], 72 (14), 1 - 138. [in Russian, with German summary but without German title]" type="journal article" year="1930">Bartenef 1930: 66</bibRefCitation>
), and that “it stands most close to
<taxonomicName id="94A64D749130F13FA5ACFA3079DC7D7E" authorityName="Vander Linden" authorityYear="1825" box="[744,953,798,824]" class="Insecta" family="Coenagrionidae" genus="Agrion" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="33" pageNumber="184" phylum="Arthropoda" rank="species" species="pulchellum">
<emphasis id="61D2EAE59130F13FA5ACFA3079DC7D7E" box="[744,953,798,824]" italics="true" pageId="33" pageNumber="184">Agrion pulchellum</emphasis>
</taxonomicName>
, but in the coloration sometimes resembles
<taxonomicName id="94A64D749130F13FA7D3FA6C7A9A7D1D" baseAuthorityName="Linnaeus" baseAuthorityYear="1758" box="[151,255,833,860]" class="Insecta" family="Coenagrionidae" genus="Coenagrion" kingdom="Animalia" order="Odonata" pageId="33" pageNumber="184" phylum="Arthropoda" rank="species" species="puella">
<emphasis id="61D2EAE59130F13FA7D3FA6C7A9A7D1D" box="[151,255,833,860]" italics="true" pageId="33" pageNumber="184">C. puella</emphasis>
</taxonomicName>
. In general, our new species is a true transition between
<taxonomicName id="94A64D749130F13FA4C8FA6C79977D1D" baseAuthorityName="Linnaeus" baseAuthorityYear="1758" box="[908,1010,833,860]" class="Insecta" family="Coenagrionidae" genus="Coenagrion" kingdom="Animalia" order="Odonata" pageId="33" pageNumber="184" phylum="Arthropoda" rank="species" species="puella">
<emphasis id="61D2EAE59130F13FA4C8FA6C79977D1D" box="[908,1010,833,860]" italics="true" pageId="33" pageNumber="184">A. puella</emphasis>
</taxonomicName>
and
<taxonomicName id="94A64D749130F13FA368FA6C7EAD7D1D" authorityName="Vander Linden" authorityYear="1825" box="[1068,1224,833,860]" class="Insecta" family="Coenagrionidae" genus="Agrion" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="33" pageNumber="184" phylum="Arthropoda" rank="species" species="pulchellum">
<emphasis id="61D2EAE59130F13FA368FA6C7EAD7D1D" box="[1068,1224,833,860]" italics="true" pageId="33" pageNumber="184">A. pulchellum</emphasis>
</taxonomicName>
. It likely is a very ancient endemic of the region, at least pre-glacial, and the contemporary
<taxonomicName id="94A64D749130F13FA48AFA4B7EFA7D39" authorityName="Vander Linden" authorityYear="1825" box="[974,1183,869,895]" class="Insecta" family="Coenagrionidae" genus="Agrion" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="33" pageNumber="184" phylum="Arthropoda" rank="species" species="pulchellum">
<emphasis id="61D2EAE59130F13FA48AFA4B7EFA7D39" box="[974,1183,869,895]" italics="true" pageId="33" pageNumber="184">Agrion pulchellum</emphasis>
</taxonomicName>
and
<taxonomicName id="94A64D749130F13FA393FA4B7F167D39" authorityName="&quot; (Skvortsov &amp; Snegovaya" authorityYear="2015" box="[1239,1395,869,895]" class="Insecta" family="Coenagrionidae" genus="Agrion" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="33" pageNumber="184" phylum="Arthropoda" rank="species" species="puella">
<emphasis id="61D2EAE59130F13FA393FA4B7F167D39" box="[1239,1395,869,895]" italics="true" pageId="33" pageNumber="184">Agrion puella</emphasis>
</taxonomicName>
are in genetic kinship with it.” (
<bibRefCitation id="37374B069130F13FA68BFAA778C57DE5" author="Bartenef, A. N." box="[463,672,905,931]" pageId="33" pageNumber="184" pagination="1 - 138" refId="ref27235" refString="Bartenef, A. N. (1930) Materialy k poznaniyu Zapadnogo Kavkaza v odonatologicheskom otnoshenii [Materials for the knowledge on West Caucasus in odonatological respect]. Trudy Zapadnokavkazskoi assotsiatsii nauchno-issledovatel'skikh institutov [Proceedings of the West Caucasian Association of scientific Research Institutes], 72 (14), 1 - 138. [in Russian, with German summary but without German title]" type="journal article" year="1930">Bartenef 1930: 70</bibRefCitation>
). The latter speculation was not argumented.
</paragraph>
<paragraph id="531936F79130F13FA783FA827BD37A74" blockId="33.[151,1437,151,2040]" pageId="33" pageNumber="184">
Both papers contain a very schematic drawing of the male appendages of
<emphasis id="61D2EAE59130F13FA37CFA837F267D80" box="[1080,1347,940,967]" italics="true" pageId="33" pageNumber="184">
<taxonomicName id="94A64D749130F13FA37CFA837EC37D80" authorityName="&quot; (Skvortsov &amp; Snegovaya" authorityYear="2015" box="[1080,1190,940,967]" class="Insecta" family="Coenagrionidae" genus="Agrion" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="33" pageNumber="184" phylum="Arthropoda" rank="species" species="puella">C. puella</taxonomicName>
,
<taxonomicName id="94A64D749130F13FA3F2FA837F267D80" authorityName="Kosterin &amp; Onishko &amp; Ilyina &amp; Chepurnov &amp; Blinov" authorityYear="2024" box="[1206,1347,941,967]" class="Insecta" family="Coenagrionidae" genus="Agrion" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="33" pageNumber="184" phylum="Arthropoda" rank="species" species="ponticum">C. ponticum</taxonomicName>
</emphasis>
and
<taxonomicName id="94A64D749130F13FA2C7FA837B767DAC" authorityName="Vander Linden" authorityYear="1825" class="Insecta" family="Coenagrionidae" genus="Agrion" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="33" pageNumber="184" phylum="Arthropoda" rank="species" species="pulchellum">
<emphasis id="61D2EAE59130F13FA2C7FA837B767DAC" italics="true" pageId="33" pageNumber="184">C. pulchellum</emphasis>
</taxonomicName>
in dorsal view and a more detailed drawing of those of
<taxonomicName id="94A64D749130F13FA438FAFF7E617DAC" authorityName="Kosterin &amp; Onishko &amp; Ilyina &amp; Chepurnov &amp; Blinov" authorityYear="2024" box="[892,1028,977,1003]" class="Insecta" family="Coenagrionidae" genus="Agrion" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="33" pageNumber="184" phylum="Arthropoda" rank="species" species="ponticum">
<emphasis id="61D2EAE59130F13FA438FAFF7E617DAC" box="[892,1028,977,1003]" italics="true" pageId="33" pageNumber="184">C. ponticum</emphasis>
</taxonomicName>
(both papers) or all the three species (
<bibRefCitation id="37374B069130F13FA7DBFADA7B247A48" author="Bartenef, A. N." box="[159,321,1012,1038]" pageId="33" pageNumber="184" pagination="1 - 138" refId="ref27235" refString="Bartenef, A. N. (1930) Materialy k poznaniyu Zapadnogo Kavkaza v odonatologicheskom otnoshenii [Materials for the knowledge on West Caucasus in odonatological respect]. Trudy Zapadnokavkazskoi assotsiatsii nauchno-issledovatel'skikh institutov [Proceedings of the West Caucasian Association of scientific Research Institutes], 72 (14), 1 - 138. [in Russian, with German summary but without German title]" type="journal article" year="1930">Bartenef 1930</bibRefCitation>
) in lateral view; that of
<taxonomicName id="94A64D749130F13FA515FADA78BF7A48" authorityName="Kosterin &amp; Onishko &amp; Ilyina &amp; Chepurnov &amp; Blinov" authorityYear="2024" box="[593,730,1012,1038]" class="Insecta" family="Coenagrionidae" genus="Agrion" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="33" pageNumber="184" phylum="Arthropoda" rank="species" species="ponticum">
<emphasis id="61D2EAE59130F13FA515FADA78BF7A48" box="[593,730,1012,1038]" italics="true" pageId="33" pageNumber="184">C. ponticum</emphasis>
</taxonomicName>
in (
<bibRefCitation id="37374B069130F13FA442FADA79CC7A48" author="Bartenef, A. N." box="[774,937,1012,1038]" pageId="33" pageNumber="184" pagination="54 - 68" refId="ref27207" refString="Bartenef, A. N. (1929) Neue Arten und Varietaten der Odonata des West Kaukasus. Zoologischer Anzeiger, 85, 54 - 68." type="journal article" year="1929">Bartenef 1929</bibRefCitation>
) is presented upside down. The drawings fit our specimens quite well.
</paragraph>
<paragraph id="531936F79130F13FA783FD157B457BD1" blockId="33.[151,1437,151,2040]" pageId="33" pageNumber="184">
The characters of
<emphasis id="61D2EAE59130F13FA6D5FD1278A07A13" box="[401,709,1083,1110]" italics="true" pageId="33" pageNumber="184">
<taxonomicName id="94A64D749130F13FA6D5FD1278547A13" authorityName="Vander Linden" authorityYear="1825" box="[401,561,1083,1110]" class="Insecta" family="Coenagrionidae" genus="Agrion" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="33" pageNumber="184" phylum="Arthropoda" rank="species" species="pulchellum">C. pulchellum</taxonomicName>
,
<taxonomicName id="94A64D749130F13FA579FD1278A07A13" authorityName="Kosterin &amp; Onishko &amp; Ilyina &amp; Chepurnov &amp; Blinov" authorityYear="2024" box="[573,709,1084,1110]" class="Insecta" family="Coenagrionidae" genus="Agrion" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="33" pageNumber="184" phylum="Arthropoda" rank="species" species="ponticum">C. ponticum</taxonomicName>
</emphasis>
and
<taxonomicName id="94A64D749130F13FA5B9FD1279067A13" authorityName="&quot; (Skvortsov &amp; Snegovaya" authorityYear="2015" box="[765,867,1083,1110]" class="Insecta" family="Coenagrionidae" genus="Agrion" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="33" pageNumber="184" phylum="Arthropoda" rank="species" species="puella">
<emphasis id="61D2EAE59130F13FA5B9FD1279067A13" box="[765,867,1083,1110]" italics="true" pageId="33" pageNumber="184">C. puella</emphasis>
</taxonomicName>
were summarised in an unnumberred comparative table which had 28 lines for males and three lines for females (
<bibRefCitation id="37374B069130F13FA40DFD7179887A3F" author="Bartenef, A. N." box="[841,1005,1119,1145]" pageId="33" pageNumber="184" pagination="54 - 68" refId="ref27207" refString="Bartenef, A. N. (1929) Neue Arten und Varietaten der Odonata des West Kaukasus. Zoologischer Anzeiger, 85, 54 - 68." type="journal article" year="1929">Bartenef 1929</bibRefCitation>
,
<bibRefCitation id="37374B069130F13FA4B3FD717E567A3F" author="Bartenef, A. N." box="[1015,1075,1119,1145]" pageId="33" pageNumber="184" pagination="1 - 138" refId="ref27235" refString="Bartenef, A. N. (1930) Materialy k poznaniyu Zapadnogo Kavkaza v odonatologicheskom otnoshenii [Materials for the knowledge on West Caucasus in odonatological respect]. Trudy Zapadnokavkazskoi assotsiatsii nauchno-issledovatel'skikh institutov [Proceedings of the West Caucasian Association of scientific Research Institutes], 72 (14), 1 - 138. [in Russian, with German summary but without German title]" type="journal article" year="1930">1930</bibRefCitation>
). Among the
<specimenCount id="45A0FD7E9130F13FA381FD717F787A3F" box="[1221,1309,1119,1145]" count="28" pageId="33" pageNumber="184" type="male">28 male</specimenCount>
characters, four concerned the pronotum, two concerned the S10 hind margin, nine concerned the appendages and one venation, other 12 characters concerned coloration. Of the
<specimenCount id="45A0FD7E9130F13FA59CFD8979027A87" box="[728,871,1191,1217]" count="3" pageId="33" pageNumber="184" type="female">three female</specimenCount>
characters, one concerned the pronotum (“as in male”) and two the coloration. These characters were not evaluated as to their importance and no truly diagnostic ones were indicated. Most of the characters, especially those concerning shape, are described in such fine detail which are no doubt individually variable, could not be reliably diagnostic and were either peculiarities of certain specimens examined or even somewhat imaginary. We find most important the more or less obvious morphological (structural) differences, which we will briefly reformulate below from (
<bibRefCitation id="37374B069130F13FA4F7FC777E327B35" author="Bartenef, A. N." box="[947,1111,1369,1395]" pageId="33" pageNumber="184" pagination="54 - 68" refId="ref27207" refString="Bartenef, A. N. (1929) Neue Arten und Varietaten der Odonata des West Kaukasus. Zoologischer Anzeiger, 85, 54 - 68." type="journal article" year="1929">Bartenef 1929</bibRefCitation>
,
<bibRefCitation id="37374B069130F13FA326FC777EFE7B35" author="Bartenef, A. N." box="[1122,1179,1369,1395]" pageId="33" pageNumber="184" pagination="1 - 138" refId="ref27235" refString="Bartenef, A. N. (1930) Materialy k poznaniyu Zapadnogo Kavkaza v odonatologicheskom otnoshenii [Materials for the knowledge on West Caucasus in odonatological respect]. Trudy Zapadnokavkazskoi assotsiatsii nauchno-issledovatel'skikh institutov [Proceedings of the West Caucasian Association of scientific Research Institutes], 72 (14), 1 - 138. [in Russian, with German summary but without German title]" type="journal article" year="1930">1930</bibRefCitation>
) in our terms, with our annotations.
</paragraph>
<paragraph id="531936F79130F13FA783FC8F7F317BFD" blockId="33.[151,1437,151,2040]" box="[199,1364,1441,1467]" pageId="33" pageNumber="184">
The structural characters of
<taxonomicName id="94A64D749130F13FA545FC8F78EF7BFD" authorityName="Kosterin &amp; Onishko &amp; Ilyina &amp; Chepurnov &amp; Blinov" authorityYear="2024" box="[513,650,1441,1467]" class="Insecta" family="Coenagrionidae" genus="Agrion" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="33" pageNumber="184" phylum="Arthropoda" rank="species" species="ponticum">
<emphasis id="61D2EAE59130F13FA545FC8F78EF7BFD" box="[513,650,1441,1467]" italics="true" pageId="33" pageNumber="184">C. ponticum</emphasis>
</taxonomicName>
intermediate between those of
<taxonomicName id="94A64D749130F13FA4ADFC8F7EE07BFD" authorityName="Vander Linden" authorityYear="1825" box="[1001,1157,1441,1467]" class="Insecta" family="Coenagrionidae" genus="Agrion" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="33" pageNumber="184" phylum="Arthropoda" rank="species" species="pulchellum">
<emphasis id="61D2EAE59130F13FA4ADFC8F7EE07BFD" box="[1001,1157,1441,1467]" italics="true" pageId="33" pageNumber="184">C. pulchellum</emphasis>
</taxonomicName>
and
<taxonomicName id="94A64D749130F13FA3F8FC8F7F467BFD" authorityName="&quot; (Skvortsov &amp; Snegovaya" authorityYear="2015" box="[1212,1315,1441,1467]" class="Insecta" family="Coenagrionidae" genus="Agrion" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="33" pageNumber="184" phylum="Arthropoda" rank="species" species="puella">
<emphasis id="61D2EAE59130F13FA3F8FC8F7F467BFD" box="[1212,1315,1441,1467]" italics="true" pageId="33" pageNumber="184">C. puella</emphasis>
</taxonomicName>
are:
</paragraph>
<paragraph id="531936F79130F13FA783FCEB7F51782B" blockId="33.[151,1437,151,2040]" pageId="33" pageNumber="184">
—Pronotum lateral lobes in both sexes less expressed in
<taxonomicName id="94A64D749130F13FA402FCEB79AA7B98" authorityName="Kosterin &amp; Onishko &amp; Ilyina &amp; Chepurnov &amp; Blinov" authorityYear="2024" box="[838,975,1477,1503]" class="Insecta" family="Coenagrionidae" genus="Agrion" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="33" pageNumber="184" phylum="Arthropoda" rank="species" species="ponticum">
<emphasis id="61D2EAE59130F13FA402FCEB79AA7B98" box="[838,975,1477,1503]" italics="true" pageId="33" pageNumber="184">C. ponticum</emphasis>
</taxonomicName>
than in
<taxonomicName id="94A64D749130F13FA36FFCEB7EAD7B98" authorityName="Vander Linden" authorityYear="1825" box="[1067,1224,1476,1503]" class="Insecta" family="Coenagrionidae" genus="Agrion" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="33" pageNumber="184" phylum="Arthropoda" rank="species" species="pulchellum">
<emphasis id="61D2EAE59130F13FA36FFCEB7EAD7B98" box="[1067,1224,1476,1503]" italics="true" pageId="33" pageNumber="184">C. pulchellum</emphasis>
</taxonomicName>
, but not as weakly as in
<taxonomicName id="94A64D749130F13FA795FCC77B527844" authorityName="&quot; (Skvortsov &amp; Snegovaya" authorityYear="2015" box="[209,311,1512,1539]" class="Insecta" family="Coenagrionidae" genus="Agrion" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="33" pageNumber="184" phylum="Arthropoda" rank="species" species="puella">
<emphasis id="61D2EAE59130F13FA795FCC77B527844" box="[209,311,1512,1539]" italics="true" pageId="33" pageNumber="184">C. puella</emphasis>
</taxonomicName>
, the central lobe being relatively more expressed as compared to the lateral lobes in
<taxonomicName id="94A64D749130F13FA39AFCC77F037844" authorityName="Kosterin &amp; Onishko &amp; Ilyina &amp; Chepurnov &amp; Blinov" authorityYear="2024" box="[1246,1382,1513,1539]" class="Insecta" family="Coenagrionidae" genus="Agrion" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="33" pageNumber="184" phylum="Arthropoda" rank="species" species="ponticum">
<emphasis id="61D2EAE59130F13FA39AFCC77F037844" box="[1246,1382,1513,1539]" italics="true" pageId="33" pageNumber="184">C. ponticum</emphasis>
</taxonomicName>
than in the two other species. In females, the depressions between the lobes are deep and symmetrical in
<taxonomicName id="94A64D749130F13FA3BBFF227FF97860" authorityName="Vander Linden" authorityYear="1825" box="[1279,1436,1548,1574]" class="Insecta" family="Coenagrionidae" genus="Agrion" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="33" pageNumber="184" phylum="Arthropoda" rank="species" species="pulchellum">
<emphasis id="61D2EAE59130F13FA3BBFF227FF97860" box="[1279,1436,1548,1574]" italics="true" pageId="33" pageNumber="184">C. pulchellum</emphasis>
</taxonomicName>
(
<figureCitation id="CB9D2A729130F13FA7DBFF1E7A8B780C" box="[159,238,1584,1610]" captionStart="FIGURE 7" captionStartId="15.[151,250,1565,1589]" captionTargetBox="[447,1140,184,1541]" captionTargetId="figure-19@15.[447,1140,181,1541]" captionTargetPageId="15" captionText="FIGURE 7. Female pronotum of Coenagrion spp. of the puella-group in the broad sense, ah—in dorsal, with respect to the body main axis (above), and anteriodorsal, perpendicular to the posterior lobe surface (below), views; ij—in dorsal view only; a—C. australocaspicum; bg—C. puella; h—C. ponticum; i—C. persicum; j—C. pulchellum; a—Russia, Dagestan, Magaramkent District, Samur Delta Forest, 2 km NW of Primorskiy village, a swampy estuary of a forest rivulet, 24.06.2021; b—Georgia, Dzau District, 4 km NNE Kvaisa, Lake Koz (Kvedi), 2830.06.2021; c—Sergokala District, 3 km SW of Myurego village, the Inchkheozen River upper valley, 19.06.2021; df—Russia, Dagestan, Rutul District; the Samur River left arm between Kufa and Kala villages, 26.06.2021; g—Russia, Karachay-Cherkes Republic, Zelenchuk District, 10 km WSW of Arkhyz, Lunnaya Polyana terrain, pools in the Dukka River valley left side, 2.07.2019; h—Russia, Chelyabinsk Province, Nagaybakskiy District, small oxbow lakes at the Gumbeyka River left bank at Ostrolenskiy Settlement SE margin, 6.07.2021; i—Russia, Krasnodarskiy Kray, Goryachiy Klyuch Municipality, Fanagoriyskoe village NW environs, ponds, 27.04.2019; j—Iran, Markazi Ostan, Shazand Shakhrestan, the Garehchay River at Gavigodar village, 34.1078-1111° N 49.3627-3657° E, 18031805 m a.s.l., 19.05.2017; k—Russia, Dagestan, Makhachkala N env., Muraveynik Protected Area at the Sulak River mouth, pools among coastal dunes, 1.06.2021. Scale bar 0.5 mm." figureDoi="http://doi.org/10.5281/zenodo.12189924" httpUri="https://zenodo.org/record/12189924/files/figure.png" pageId="33" pageNumber="184">Fig. 7j</figureCitation>
), less expressed and asymmetrical in
<taxonomicName id="94A64D749130F13FA5E4FF1E794E780C" authorityName="Kosterin &amp; Onishko &amp; Ilyina &amp; Chepurnov &amp; Blinov" authorityYear="2024" box="[672,811,1584,1610]" class="Insecta" family="Coenagrionidae" genus="Agrion" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="33" pageNumber="184" phylum="Arthropoda" rank="species" species="ponticum">
<emphasis id="61D2EAE59130F13FA5E4FF1E794E780C" box="[672,811,1584,1610]" italics="true" pageId="33" pageNumber="184">C. ponticum</emphasis>
</taxonomicName>
(
<figureCitation id="CB9D2A729130F13FA478FF1E79F4780C" box="[828,913,1584,1610]" captionStart="FIGURE 7" captionStartId="15.[151,250,1565,1589]" captionTargetBox="[447,1140,184,1541]" captionTargetId="figure-19@15.[447,1140,181,1541]" captionTargetPageId="15" captionText="FIGURE 7. Female pronotum of Coenagrion spp. of the puella-group in the broad sense, ah—in dorsal, with respect to the body main axis (above), and anteriodorsal, perpendicular to the posterior lobe surface (below), views; ij—in dorsal view only; a—C. australocaspicum; bg—C. puella; h—C. ponticum; i—C. persicum; j—C. pulchellum; a—Russia, Dagestan, Magaramkent District, Samur Delta Forest, 2 km NW of Primorskiy village, a swampy estuary of a forest rivulet, 24.06.2021; b—Georgia, Dzau District, 4 km NNE Kvaisa, Lake Koz (Kvedi), 2830.06.2021; c—Sergokala District, 3 km SW of Myurego village, the Inchkheozen River upper valley, 19.06.2021; df—Russia, Dagestan, Rutul District; the Samur River left arm between Kufa and Kala villages, 26.06.2021; g—Russia, Karachay-Cherkes Republic, Zelenchuk District, 10 km WSW of Arkhyz, Lunnaya Polyana terrain, pools in the Dukka River valley left side, 2.07.2019; h—Russia, Chelyabinsk Province, Nagaybakskiy District, small oxbow lakes at the Gumbeyka River left bank at Ostrolenskiy Settlement SE margin, 6.07.2021; i—Russia, Krasnodarskiy Kray, Goryachiy Klyuch Municipality, Fanagoriyskoe village NW environs, ponds, 27.04.2019; j—Iran, Markazi Ostan, Shazand Shakhrestan, the Garehchay River at Gavigodar village, 34.1078-1111° N 49.3627-3657° E, 18031805 m a.s.l., 19.05.2017; k—Russia, Dagestan, Makhachkala N env., Muraveynik Protected Area at the Sulak River mouth, pools among coastal dunes, 1.06.2021. Scale bar 0.5 mm." figureDoi="http://doi.org/10.5281/zenodo.12189924" httpUri="https://zenodo.org/record/12189924/files/figure.png" pageId="33" pageNumber="184">Fig. 7h</figureCitation>
), scarcely expressed in
<taxonomicName id="94A64D749130F13FA3E7FF1E7F69780C" authorityName="&quot; (Skvortsov &amp; Snegovaya" authorityYear="2015" box="[1187,1292,1584,1610]" class="Insecta" family="Coenagrionidae" genus="Agrion" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="33" pageNumber="184" phylum="Arthropoda" rank="species" species="puella">
<emphasis id="61D2EAE59130F13FA3E7FF1E7F69780C" box="[1187,1292,1584,1610]" italics="true" pageId="33" pageNumber="184">C. puella</emphasis>
</taxonomicName>
(
<figureCitation id="CB9D2A729130F13FA259FF1E7FF4780C" box="[1309,1425,1584,1610]" captionStart="FIGURE 7" captionStartId="15.[151,250,1565,1589]" captionTargetBox="[447,1140,184,1541]" captionTargetId="figure-19@15.[447,1140,181,1541]" captionTargetPageId="15" captionText="FIGURE 7. Female pronotum of Coenagrion spp. of the puella-group in the broad sense, ah—in dorsal, with respect to the body main axis (above), and anteriodorsal, perpendicular to the posterior lobe surface (below), views; ij—in dorsal view only; a—C. australocaspicum; bg—C. puella; h—C. ponticum; i—C. persicum; j—C. pulchellum; a—Russia, Dagestan, Magaramkent District, Samur Delta Forest, 2 km NW of Primorskiy village, a swampy estuary of a forest rivulet, 24.06.2021; b—Georgia, Dzau District, 4 km NNE Kvaisa, Lake Koz (Kvedi), 2830.06.2021; c—Sergokala District, 3 km SW of Myurego village, the Inchkheozen River upper valley, 19.06.2021; df—Russia, Dagestan, Rutul District; the Samur River left arm between Kufa and Kala villages, 26.06.2021; g—Russia, Karachay-Cherkes Republic, Zelenchuk District, 10 km WSW of Arkhyz, Lunnaya Polyana terrain, pools in the Dukka River valley left side, 2.07.2019; h—Russia, Chelyabinsk Province, Nagaybakskiy District, small oxbow lakes at the Gumbeyka River left bank at Ostrolenskiy Settlement SE margin, 6.07.2021; i—Russia, Krasnodarskiy Kray, Goryachiy Klyuch Municipality, Fanagoriyskoe village NW environs, ponds, 27.04.2019; j—Iran, Markazi Ostan, Shazand Shakhrestan, the Garehchay River at Gavigodar village, 34.1078-1111° N 49.3627-3657° E, 18031805 m a.s.l., 19.05.2017; k—Russia, Dagestan, Makhachkala N env., Muraveynik Protected Area at the Sulak River mouth, pools among coastal dunes, 1.06.2021. Scale bar 0.5 mm." figureDoi="http://doi.org/10.5281/zenodo.12189924" httpUri="https://zenodo.org/record/12189924/files/figure.png" pageId="33" pageNumber="184">Fig. 7bg</figureCitation>
). These pronotum characters are well seen in our female specimens but is not evident in males (see above).
</paragraph>
<paragraph id="531936F79130F13FA783FF597A97789F" blockId="33.[151,1437,151,2040]" pageId="33" pageNumber="184">
—S10 hind margin dorsal incision in males is deeper than half of its length in
<taxonomicName id="94A64D749130F13FA37DFF567EB378D7" authorityName="Vander Linden" authorityYear="1825" box="[1081,1238,1655,1682]" class="Insecta" family="Coenagrionidae" genus="Agrion" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="33" pageNumber="184" phylum="Arthropoda" rank="species" species="pulchellum">
<emphasis id="61D2EAE59130F13FA37DFF567EB378D7" box="[1081,1238,1655,1682]" italics="true" pageId="33" pageNumber="184">C. pulchellum</emphasis>
</taxonomicName>
, about the half in
<taxonomicName id="94A64D749130F13FA7D3FFB57B4478F3" authorityName="Kosterin &amp; Onishko &amp; Ilyina &amp; Chepurnov &amp; Blinov" authorityYear="2024" box="[151,289,1691,1717]" class="Insecta" family="Coenagrionidae" genus="Agrion" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="33" pageNumber="184" phylum="Arthropoda" rank="species" species="ponticum">
<emphasis id="61D2EAE59130F13FA7D3FFB57B4478F3" box="[151,289,1691,1717]" italics="true" pageId="33" pageNumber="184">C. ponticum</emphasis>
</taxonomicName>
and less in
<taxonomicName id="94A64D749130F13FA6EEFFB5787778F3" authorityName="&quot; (Skvortsov &amp; Snegovaya" authorityYear="2015" box="[426,530,1691,1717]" class="Insecta" family="Coenagrionidae" genus="Agrion" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="33" pageNumber="184" phylum="Arthropoda" rank="species" species="puella">
<emphasis id="61D2EAE59130F13FA6EEFFB5787778F3" box="[426,530,1691,1717]" italics="true" pageId="33" pageNumber="184">C. puella</emphasis>
</taxonomicName>
. This is correct but the character is weak as quantitative and without any hiatus (
<figureCitation id="CB9D2A729130F13FA7DBFF917A80789F" box="[159,229,1727,1753]" captionStart="FIGURE 2" captionStartId="9.[151,250,1421,1445]" captionTargetBox="[381,1200,187,1391]" captionTargetId="figure-19@9.[381,1206,181,1397]" captionTargetPageId="9" captionText="FIGURE 2. Male appendages of Coenagrion spp. of the puella-group in the broad sense, from the Caucasus and elsewhere, in dorsal view: ac—C. australocaspicum (b—with the appendages expanded); d-k—C. puella; lm—C. ponticum (males with a more (c) and less (d) expanded black abdominal maculation); n—C. persicum; op—C. pulchellum; ab—Russia, Dagestan, Magaramkent District, Samur Delta Forest, 2 km NW of Primorskiy village, a swampy estuary of a forest rivulet, 24.06.2021; c—Russia, Dagestan, Kumtorkala District, the Shura-Ozen River at Kumtorkala Station, next to Sarykum Barkhan, 3.05.2021; d—Georgia, Dzau District, 4 km NNE Kvaisa, Lake Koz (Kvedi), 2830.06.2021; ef—Russia, Dagestan, Rutul District; the Samur River left arm between Kufa and Kala villages, 26.06.2021; gh—Russia, Dagestan, Magaramkent District, Samur Delta Forest, Bil-Bil Kazmalyar village, 25.06.2021; i—Russia, Dagestan, Buynaksk District, Buynaksk S environs, Nizhnee Kazanishche village environs, a water reservoir, 2020; j—Russia, Karachay-Cherkes Republic, Zelenchuk District, 10 km WSW of Arkhyz, Lunnaya Polyana terrain, pools in the Dukka River valley left side, 2.07.2019; k—Russia, Republic of Bashkortostan, Zinachurinskiy District, a small roadside oxbow lake 0.7 km E of Verkhniy Sarabil village, 5.07.2021; lm— Russia, Krasnodarskiy Kray, Goryachiy Klyuch Municipality, Fanagoriyskoe village NW environs, ponds, 27.04.2019; n—Iran, Markazi Ostan, Shazand Shakhrestan, Emarat village env., a spring and swampy reservoir, 20.05.2017; o—Russia, Dagestan, Makhachkala N suburbs, pools between dunes at the Krivaya Balka River left bank in its lowermost reaches, 23.05.2023; p—Russia, N Caucasus, Karachay-Cherkes Republic, Zelenchuk District, Dausuz village W environs, S banks of a reservoir at the Bolzhoy Zelenchuk River, 1012 m a.s.l.; q—Russia, Dagestan, Makhachkala N env., Muraveynik Protected Area at the Sulak River mouth, pools among coastal dunes, 1.06.2021. Scale bar 0.5 mm." figureDoi="http://doi.org/10.5281/zenodo.12189910" httpUri="https://zenodo.org/record/12189910/files/figure.png" pageId="33" pageNumber="184">Fig. 2</figureCitation>
).
</paragraph>
<paragraph id="531936F79130F13FA783FFCC7FFC7902" blockId="33.[151,1437,151,2040]" pageId="33" pageNumber="184">
—Male cercus length is somewhat less than half of S10 length in
<emphasis id="61D2EAE59130F13FA4DBFFCD7E4B78BA" box="[927,1070,1763,1789]" italics="true" pageId="33" pageNumber="184">
<taxonomicName id="94A64D749130F13FA4DBFFCD7E4F78BA" authorityName="Kosterin &amp; Onishko &amp; Ilyina &amp; Chepurnov &amp; Blinov" authorityYear="2024" box="[927,1066,1763,1789]" class="Insecta" family="Coenagrionidae" genus="Agrion" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="33" pageNumber="184" phylum="Arthropoda" rank="species" species="ponticum">C. ponticum</taxonomicName>
,
</emphasis>
versus somewhat more than half in
<taxonomicName id="94A64D749130F13FA7F1FE297B797966" authorityName="&quot; (Skvortsov &amp; Snegovaya" authorityYear="2015" box="[181,284,1798,1825]" class="Insecta" family="Coenagrionidae" genus="Agrion" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="33" pageNumber="184" phylum="Arthropoda" rank="species" species="puella">
<emphasis id="61D2EAE59130F13FA7F1FE297B797966" box="[181,284,1798,1825]" italics="true" pageId="33" pageNumber="184">C. puella</emphasis>
</taxonomicName>
and four times as short in
<taxonomicName id="94A64D749130F13FA508FE29788C7966" authorityName="Vander Linden" authorityYear="1825" box="[588,745,1798,1825]" class="Insecta" family="Coenagrionidae" genus="Agrion" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="33" pageNumber="184" phylum="Arthropoda" rank="species" species="pulchellum">
<emphasis id="61D2EAE59130F13FA508FE29788C7966" box="[588,745,1798,1825]" italics="true" pageId="33" pageNumber="184">C. pulchellum</emphasis>
</taxonomicName>
. We find these values incorrect, since in
<taxonomicName id="94A64D749130F13FA3FFFE297F467966" authorityName="&quot; (Skvortsov &amp; Snegovaya" authorityYear="2015" box="[1211,1315,1798,1825]" class="Insecta" family="Coenagrionidae" genus="Agrion" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="33" pageNumber="184" phylum="Arthropoda" rank="species" species="puella">
<emphasis id="61D2EAE59130F13FA3FFFE297F467966" box="[1211,1315,1798,1825]" italics="true" pageId="33" pageNumber="184">C. puella</emphasis>
</taxonomicName>
the cercus may look shorter than half of S10, depending on its position, while reaches 1/3 of the S10 length in
<taxonomicName id="94A64D749130F13FA3B1FE047FF47902" authorityName="Vander Linden" authorityYear="1825" box="[1269,1425,1834,1860]" class="Insecta" family="Coenagrionidae" genus="Agrion" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="33" pageNumber="184" phylum="Arthropoda" rank="species" species="pulchellum">
<emphasis id="61D2EAE59130F13FA3B1FE047FF47902" box="[1269,1425,1834,1860]" italics="true" pageId="33" pageNumber="184">C. pulchellum</emphasis>
</taxonomicName>
.
</paragraph>
<paragraph id="531936F79130F13FA783FE60798E79BE" blockId="33.[151,1437,151,2040]" pageId="33" pageNumber="184">
—Male cerci contact each other in dorsal view at or just behind S10 hind margin, while only at their bases in
<taxonomicName id="94A64D749130F13FA7D3FE5C7A9B79CA" authorityName="&quot; (Skvortsov &amp; Snegovaya" authorityYear="2015" box="[151,254,1906,1932]" class="Insecta" family="Coenagrionidae" genus="Agrion" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="33" pageNumber="184" phylum="Arthropoda" rank="species" species="puella">
<emphasis id="61D2EAE59130F13FA7D3FE5C7A9B79CA" box="[151,254,1906,1932]" italics="true" pageId="33" pageNumber="184">C. puella</emphasis>
</taxonomicName>
and at the middle of their trunks with their bulges in
<emphasis id="61D2EAE59130F13FA40BFE5C799679CA" box="[847,1011,1906,1932]" italics="true" pageId="33" pageNumber="184">
<taxonomicName id="94A64D749130F13FA40BFE5C798A79CA" authorityName="Vander Linden" authorityYear="1825" box="[847,1007,1906,1932]" class="Insecta" family="Coenagrionidae" genus="Agrion" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="33" pageNumber="184" phylum="Arthropoda" rank="species" species="pulchellum">C. pulchellum</taxonomicName>
.
</emphasis>
We find this is true for
<taxonomicName id="94A64D749130F13FA3BEFE5C7F0579CA" authorityName="&quot; (Skvortsov &amp; Snegovaya" authorityYear="2015" box="[1274,1376,1906,1932]" class="Insecta" family="Coenagrionidae" genus="Agrion" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="33" pageNumber="184" phylum="Arthropoda" rank="species" species="puella">
<emphasis id="61D2EAE59130F13FA3BEFE5C7F0579CA" box="[1274,1376,1906,1932]" italics="true" pageId="33" pageNumber="184">C. puella</emphasis>
</taxonomicName>
(
<figureCitation id="CB9D2A729130F13FA22BFE5C7AB479F6" captionStart="FIGURE 2" captionStartId="9.[151,250,1421,1445]" captionTargetBox="[381,1200,187,1391]" captionTargetId="figure-19@9.[381,1206,181,1397]" captionTargetPageId="9" captionText="FIGURE 2. Male appendages of Coenagrion spp. of the puella-group in the broad sense, from the Caucasus and elsewhere, in dorsal view: ac—C. australocaspicum (b—with the appendages expanded); d-k—C. puella; lm—C. ponticum (males with a more (c) and less (d) expanded black abdominal maculation); n—C. persicum; op—C. pulchellum; ab—Russia, Dagestan, Magaramkent District, Samur Delta Forest, 2 km NW of Primorskiy village, a swampy estuary of a forest rivulet, 24.06.2021; c—Russia, Dagestan, Kumtorkala District, the Shura-Ozen River at Kumtorkala Station, next to Sarykum Barkhan, 3.05.2021; d—Georgia, Dzau District, 4 km NNE Kvaisa, Lake Koz (Kvedi), 2830.06.2021; ef—Russia, Dagestan, Rutul District; the Samur River left arm between Kufa and Kala villages, 26.06.2021; gh—Russia, Dagestan, Magaramkent District, Samur Delta Forest, Bil-Bil Kazmalyar village, 25.06.2021; i—Russia, Dagestan, Buynaksk District, Buynaksk S environs, Nizhnee Kazanishche village environs, a water reservoir, 2020; j—Russia, Karachay-Cherkes Republic, Zelenchuk District, 10 km WSW of Arkhyz, Lunnaya Polyana terrain, pools in the Dukka River valley left side, 2.07.2019; k—Russia, Republic of Bashkortostan, Zinachurinskiy District, a small roadside oxbow lake 0.7 km E of Verkhniy Sarabil village, 5.07.2021; lm— Russia, Krasnodarskiy Kray, Goryachiy Klyuch Municipality, Fanagoriyskoe village NW environs, ponds, 27.04.2019; n—Iran, Markazi Ostan, Shazand Shakhrestan, Emarat village env., a spring and swampy reservoir, 20.05.2017; o—Russia, Dagestan, Makhachkala N suburbs, pools between dunes at the Krivaya Balka River left bank in its lowermost reaches, 23.05.2023; p—Russia, N Caucasus, Karachay-Cherkes Republic, Zelenchuk District, Dausuz village W environs, S banks of a reservoir at the Bolzhoy Zelenchuk River, 1012 m a.s.l.; q—Russia, Dagestan, Makhachkala N env., Muraveynik Protected Area at the Sulak River mouth, pools among coastal dunes, 1.06.2021. Scale bar 0.5 mm." figureDoi="http://doi.org/10.5281/zenodo.12189910" httpUri="https://zenodo.org/record/12189910/files/figure.png" pageId="33" pageNumber="184">Fig. 2dk</figureCitation>
) while the point of contact may be similar in
<taxonomicName id="94A64D749130F13FA5A5FEB8791A79F6" authorityName="Vander Linden" authorityYear="1825" box="[737,895,1942,1968]" class="Insecta" family="Coenagrionidae" genus="Agrion" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="33" pageNumber="184" phylum="Arthropoda" rank="species" species="pulchellum">
<emphasis id="61D2EAE59130F13FA5A5FEB8791A79F6" box="[737,895,1942,1968]" italics="true" pageId="33" pageNumber="184">C. pulchellum</emphasis>
</taxonomicName>
(
<figureCitation id="CB9D2A729130F13FA4D4FEB87E6579F6" box="[912,1024,1942,1968]" captionStart="FIGURE 2" captionStartId="9.[151,250,1421,1445]" captionTargetBox="[381,1200,187,1391]" captionTargetId="figure-19@9.[381,1206,181,1397]" captionTargetPageId="9" captionText="FIGURE 2. Male appendages of Coenagrion spp. of the puella-group in the broad sense, from the Caucasus and elsewhere, in dorsal view: ac—C. australocaspicum (b—with the appendages expanded); d-k—C. puella; lm—C. ponticum (males with a more (c) and less (d) expanded black abdominal maculation); n—C. persicum; op—C. pulchellum; ab—Russia, Dagestan, Magaramkent District, Samur Delta Forest, 2 km NW of Primorskiy village, a swampy estuary of a forest rivulet, 24.06.2021; c—Russia, Dagestan, Kumtorkala District, the Shura-Ozen River at Kumtorkala Station, next to Sarykum Barkhan, 3.05.2021; d—Georgia, Dzau District, 4 km NNE Kvaisa, Lake Koz (Kvedi), 2830.06.2021; ef—Russia, Dagestan, Rutul District; the Samur River left arm between Kufa and Kala villages, 26.06.2021; gh—Russia, Dagestan, Magaramkent District, Samur Delta Forest, Bil-Bil Kazmalyar village, 25.06.2021; i—Russia, Dagestan, Buynaksk District, Buynaksk S environs, Nizhnee Kazanishche village environs, a water reservoir, 2020; j—Russia, Karachay-Cherkes Republic, Zelenchuk District, 10 km WSW of Arkhyz, Lunnaya Polyana terrain, pools in the Dukka River valley left side, 2.07.2019; k—Russia, Republic of Bashkortostan, Zinachurinskiy District, a small roadside oxbow lake 0.7 km E of Verkhniy Sarabil village, 5.07.2021; lm— Russia, Krasnodarskiy Kray, Goryachiy Klyuch Municipality, Fanagoriyskoe village NW environs, ponds, 27.04.2019; n—Iran, Markazi Ostan, Shazand Shakhrestan, Emarat village env., a spring and swampy reservoir, 20.05.2017; o—Russia, Dagestan, Makhachkala N suburbs, pools between dunes at the Krivaya Balka River left bank in its lowermost reaches, 23.05.2023; p—Russia, N Caucasus, Karachay-Cherkes Republic, Zelenchuk District, Dausuz village W environs, S banks of a reservoir at the Bolzhoy Zelenchuk River, 1012 m a.s.l.; q—Russia, Dagestan, Makhachkala N env., Muraveynik Protected Area at the Sulak River mouth, pools among coastal dunes, 1.06.2021. Scale bar 0.5 mm." figureDoi="http://doi.org/10.5281/zenodo.12189910" httpUri="https://zenodo.org/record/12189910/files/figure.png" pageId="33" pageNumber="184">Fig. 2oq</figureCitation>
) and
<taxonomicName id="94A64D749130F13FA306FEB87EA979F6" authorityName="Kosterin &amp; Onishko &amp; Ilyina &amp; Chepurnov &amp; Blinov" authorityYear="2024" box="[1090,1228,1942,1968]" class="Insecta" family="Coenagrionidae" genus="Agrion" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="33" pageNumber="184" phylum="Arthropoda" rank="species" species="ponticum">
<emphasis id="61D2EAE59130F13FA306FEB87EA979F6" box="[1090,1228,1942,1968]" italics="true" pageId="33" pageNumber="184">C. ponticum</emphasis>
</taxonomicName>
(
<figureCitation id="CB9D2A729130F13FA39AFEB87F3479F6" box="[1246,1361,1942,1968]" captionStart="FIGURE 2" captionStartId="9.[151,250,1421,1445]" captionTargetBox="[381,1200,187,1391]" captionTargetId="figure-19@9.[381,1206,181,1397]" captionTargetPageId="9" captionText="FIGURE 2. Male appendages of Coenagrion spp. of the puella-group in the broad sense, from the Caucasus and elsewhere, in dorsal view: ac—C. australocaspicum (b—with the appendages expanded); d-k—C. puella; lm—C. ponticum (males with a more (c) and less (d) expanded black abdominal maculation); n—C. persicum; op—C. pulchellum; ab—Russia, Dagestan, Magaramkent District, Samur Delta Forest, 2 km NW of Primorskiy village, a swampy estuary of a forest rivulet, 24.06.2021; c—Russia, Dagestan, Kumtorkala District, the Shura-Ozen River at Kumtorkala Station, next to Sarykum Barkhan, 3.05.2021; d—Georgia, Dzau District, 4 km NNE Kvaisa, Lake Koz (Kvedi), 2830.06.2021; ef—Russia, Dagestan, Rutul District; the Samur River left arm between Kufa and Kala villages, 26.06.2021; gh—Russia, Dagestan, Magaramkent District, Samur Delta Forest, Bil-Bil Kazmalyar village, 25.06.2021; i—Russia, Dagestan, Buynaksk District, Buynaksk S environs, Nizhnee Kazanishche village environs, a water reservoir, 2020; j—Russia, Karachay-Cherkes Republic, Zelenchuk District, 10 km WSW of Arkhyz, Lunnaya Polyana terrain, pools in the Dukka River valley left side, 2.07.2019; k—Russia, Republic of Bashkortostan, Zinachurinskiy District, a small roadside oxbow lake 0.7 km E of Verkhniy Sarabil village, 5.07.2021; lm— Russia, Krasnodarskiy Kray, Goryachiy Klyuch Municipality, Fanagoriyskoe village NW environs, ponds, 27.04.2019; n—Iran, Markazi Ostan, Shazand Shakhrestan, Emarat village env., a spring and swampy reservoir, 20.05.2017; o—Russia, Dagestan, Makhachkala N suburbs, pools between dunes at the Krivaya Balka River left bank in its lowermost reaches, 23.05.2023; p—Russia, N Caucasus, Karachay-Cherkes Republic, Zelenchuk District, Dausuz village W environs, S banks of a reservoir at the Bolzhoy Zelenchuk River, 1012 m a.s.l.; q—Russia, Dagestan, Makhachkala N env., Muraveynik Protected Area at the Sulak River mouth, pools among coastal dunes, 1.06.2021. Scale bar 0.5 mm." figureDoi="http://doi.org/10.5281/zenodo.12189910" httpUri="https://zenodo.org/record/12189910/files/figure.png" pageId="33" pageNumber="184">Fig. 2lm</figureCitation>
), at ca proximal ¼ of the trunk, while the actual difference is the shape of the bulge which is a well expressed blunt angle in
<taxonomicName id="94A64D749130F13FA7F0FEF07B3479BE" authorityName="Vander Linden" authorityYear="1825" box="[180,337,2014,2040]" class="Insecta" family="Coenagrionidae" genus="Agrion" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="33" pageNumber="184" phylum="Arthropoda" rank="species" species="pulchellum">
<emphasis id="61D2EAE59130F13FA7F0FEF07B3479BE" box="[180,337,2014,2040]" italics="true" pageId="33" pageNumber="184">C. pulchellum</emphasis>
</taxonomicName>
(
<figureCitation id="CB9D2A729130F13FA624FEF07BAB79BE" box="[352,462,2014,2040]" captionStart="FIGURE 2" captionStartId="9.[151,250,1421,1445]" captionTargetBox="[381,1200,187,1391]" captionTargetId="figure-19@9.[381,1206,181,1397]" captionTargetPageId="9" captionText="FIGURE 2. Male appendages of Coenagrion spp. of the puella-group in the broad sense, from the Caucasus and elsewhere, in dorsal view: ac—C. australocaspicum (b—with the appendages expanded); d-k—C. puella; lm—C. ponticum (males with a more (c) and less (d) expanded black abdominal maculation); n—C. persicum; op—C. pulchellum; ab—Russia, Dagestan, Magaramkent District, Samur Delta Forest, 2 km NW of Primorskiy village, a swampy estuary of a forest rivulet, 24.06.2021; c—Russia, Dagestan, Kumtorkala District, the Shura-Ozen River at Kumtorkala Station, next to Sarykum Barkhan, 3.05.2021; d—Georgia, Dzau District, 4 km NNE Kvaisa, Lake Koz (Kvedi), 2830.06.2021; ef—Russia, Dagestan, Rutul District; the Samur River left arm between Kufa and Kala villages, 26.06.2021; gh—Russia, Dagestan, Magaramkent District, Samur Delta Forest, Bil-Bil Kazmalyar village, 25.06.2021; i—Russia, Dagestan, Buynaksk District, Buynaksk S environs, Nizhnee Kazanishche village environs, a water reservoir, 2020; j—Russia, Karachay-Cherkes Republic, Zelenchuk District, 10 km WSW of Arkhyz, Lunnaya Polyana terrain, pools in the Dukka River valley left side, 2.07.2019; k—Russia, Republic of Bashkortostan, Zinachurinskiy District, a small roadside oxbow lake 0.7 km E of Verkhniy Sarabil village, 5.07.2021; lm— Russia, Krasnodarskiy Kray, Goryachiy Klyuch Municipality, Fanagoriyskoe village NW environs, ponds, 27.04.2019; n—Iran, Markazi Ostan, Shazand Shakhrestan, Emarat village env., a spring and swampy reservoir, 20.05.2017; o—Russia, Dagestan, Makhachkala N suburbs, pools between dunes at the Krivaya Balka River left bank in its lowermost reaches, 23.05.2023; p—Russia, N Caucasus, Karachay-Cherkes Republic, Zelenchuk District, Dausuz village W environs, S banks of a reservoir at the Bolzhoy Zelenchuk River, 1012 m a.s.l.; q—Russia, Dagestan, Makhachkala N env., Muraveynik Protected Area at the Sulak River mouth, pools among coastal dunes, 1.06.2021. Scale bar 0.5 mm." figureDoi="http://doi.org/10.5281/zenodo.12189910" httpUri="https://zenodo.org/record/12189910/files/figure.png" pageId="33" pageNumber="184">Fig. 2oq</figureCitation>
) but gently rounded in
<taxonomicName id="94A64D749130F13FA590FEF0793879BE" authorityName="Kosterin &amp; Onishko &amp; Ilyina &amp; Chepurnov &amp; Blinov" authorityYear="2024" box="[724,861,2014,2040]" class="Insecta" family="Coenagrionidae" genus="Agrion" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="33" pageNumber="184" phylum="Arthropoda" rank="species" species="ponticum">
<emphasis id="61D2EAE59130F13FA590FEF0793879BE" box="[724,861,2014,2040]" italics="true" pageId="33" pageNumber="184">C. ponticum</emphasis>
</taxonomicName>
(
<figureCitation id="CB9D2A729130F13FA429FEF079BA79BE" box="[877,991,2014,2040]" captionStart="FIGURE 2" captionStartId="9.[151,250,1421,1445]" captionTargetBox="[381,1200,187,1391]" captionTargetId="figure-19@9.[381,1206,181,1397]" captionTargetPageId="9" captionText="FIGURE 2. Male appendages of Coenagrion spp. of the puella-group in the broad sense, from the Caucasus and elsewhere, in dorsal view: ac—C. australocaspicum (b—with the appendages expanded); d-k—C. puella; lm—C. ponticum (males with a more (c) and less (d) expanded black abdominal maculation); n—C. persicum; op—C. pulchellum; ab—Russia, Dagestan, Magaramkent District, Samur Delta Forest, 2 km NW of Primorskiy village, a swampy estuary of a forest rivulet, 24.06.2021; c—Russia, Dagestan, Kumtorkala District, the Shura-Ozen River at Kumtorkala Station, next to Sarykum Barkhan, 3.05.2021; d—Georgia, Dzau District, 4 km NNE Kvaisa, Lake Koz (Kvedi), 2830.06.2021; ef—Russia, Dagestan, Rutul District; the Samur River left arm between Kufa and Kala villages, 26.06.2021; gh—Russia, Dagestan, Magaramkent District, Samur Delta Forest, Bil-Bil Kazmalyar village, 25.06.2021; i—Russia, Dagestan, Buynaksk District, Buynaksk S environs, Nizhnee Kazanishche village environs, a water reservoir, 2020; j—Russia, Karachay-Cherkes Republic, Zelenchuk District, 10 km WSW of Arkhyz, Lunnaya Polyana terrain, pools in the Dukka River valley left side, 2.07.2019; k—Russia, Republic of Bashkortostan, Zinachurinskiy District, a small roadside oxbow lake 0.7 km E of Verkhniy Sarabil village, 5.07.2021; lm— Russia, Krasnodarskiy Kray, Goryachiy Klyuch Municipality, Fanagoriyskoe village NW environs, ponds, 27.04.2019; n—Iran, Markazi Ostan, Shazand Shakhrestan, Emarat village env., a spring and swampy reservoir, 20.05.2017; o—Russia, Dagestan, Makhachkala N suburbs, pools between dunes at the Krivaya Balka River left bank in its lowermost reaches, 23.05.2023; p—Russia, N Caucasus, Karachay-Cherkes Republic, Zelenchuk District, Dausuz village W environs, S banks of a reservoir at the Bolzhoy Zelenchuk River, 1012 m a.s.l.; q—Russia, Dagestan, Makhachkala N env., Muraveynik Protected Area at the Sulak River mouth, pools among coastal dunes, 1.06.2021. Scale bar 0.5 mm." figureDoi="http://doi.org/10.5281/zenodo.12189910" httpUri="https://zenodo.org/record/12189910/files/figure.png" pageId="33" pageNumber="184">Fig. 2lm</figureCitation>
).
</paragraph>
<paragraph id="531936F79133F13CA783F9B97A857F5B" blockId="34.[151,1437,151,2013]" pageId="34" pageNumber="185">
—Apex of male cercus trunk moderately convex, while very convex and roundish in
<taxonomicName id="94A64D749133F13CA335F9B97F697EF7" authorityName="Vander Linden" authorityYear="1825" box="[1137,1292,151,177]" class="Insecta" family="Coenagrionidae" genus="Agrion" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="34" pageNumber="185" phylum="Arthropoda" rank="species" species="pulchellum">
<emphasis id="61D2EAE59133F13CA335F9B97F697EF7" box="[1137,1292,151,177]" italics="true" pageId="34" pageNumber="185">C. pulchellum</emphasis>
</taxonomicName>
and scarcely convex or straight in
<taxonomicName id="94A64D749133F13CA6C7F9957B987E93" baseAuthorityName="Linnaeus" baseAuthorityYear="1758" box="[387,509,187,213]" class="Insecta" family="Coenagrionidae" genus="Coenagrion" kingdom="Animalia" order="Odonata" pageId="34" pageNumber="185" phylum="Arthropoda" rank="species" species="puellum">
<emphasis id="61D2EAE59133F13CA6C7F9957B987E93" box="[387,509,187,213]" italics="true" pageId="34" pageNumber="185">C. puellum</emphasis>
</taxonomicName>
. We would reformulate that in lateral view, it is bluntly angled in
<taxonomicName id="94A64D749133F13CA39CF9957F057E92" authorityName="Kosterin &amp; Onishko &amp; Ilyina &amp; Chepurnov &amp; Blinov" authorityYear="2024" box="[1240,1376,187,213]" class="Insecta" family="Coenagrionidae" genus="Agrion" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="34" pageNumber="185" phylum="Arthropoda" rank="species" species="ponticum">
<emphasis id="61D2EAE59133F13CA39CF9957F057E92" box="[1240,1376,187,213]" italics="true" pageId="34" pageNumber="185">C. ponticum</emphasis>
</taxonomicName>
(
<figureCitation id="CB9D2A729133F13CA22BF9957AB07EBF" captionStart="FIGURE3" captionStartId="10.[151,250,1432,1456]" captionTargetBox="[393,1180,188,1398]" captionTargetId="figure-24@10.[393,1194,181,1408]" captionTargetPageId="10" captionText="FIGURE3. Maleappendages of Coenagrion spp.ofthe puella-groupin thebroadsense inlateral view:ac—C.australocaspicum (b—with the appendages expanded); dk—C. puella; lm—C. ponticum (males with a more (c) and less (d) expanded black abdominal maculation); n—C. persicum; oq—C. pulchellum; ab—Russia, Dagestan, Magaramkent District, Samur Delta Forest, 2 km NW of Primorskiy village, a swampy estuary of a forest rivulet, 24.06.2021; c—Russia, Dagestan, Kumtorkala District, the Shura-Ozen River at Kumtorkala Station, next to Sarykum Barkhan, 3.05.2021; d —Georgia, Dzau District, 4 km NNE Kvaisa, Lake Koz (Kvedi), 2830.06.2021; ef—Russia, Dagestan, Rutul District; the Samur River left arm between Kufa and Kala villages, 26.06.2021; gh—Russia, Dagestan, Magaramkent District, Samur Delta Forest, Bil-Bil Kazmalyar village, 25.06.2021; i—Russia, Dagestan, Buynaksk District, Buynaksk S environs, Nizhnee Kazanishche village environs, a water reservoir, 2020; k—Russia, Karachay-Cherkes Republic, Zelenchuk District, 10 km WSW of Arkhyz, Lunnaya Polyana terrain, pools in the Dukka River valley left side, 2.07.2019; k—Russia, Republic of Bashkortostan, Zinachurinskiy District, a small roadside oxbow lake 0.7 km E of Verkhniy Sarabil village, 5.07.2021; lm—Russia, Krasnodarskiy Kray, Goryachiy Klyuch Municipality, Fanagoriyskoe village NW environs, ponds, 27.04.2019; n—Iran, Markazi Ostan, Shazand Shakhrestan, Emarat village env., a spring and swampy reservoir, 20.05.2017; o—Russia, Dagestan, Makhachkala N suburbs, pools between dunes at the Krivaya Balka River left bank in its lowermost reaches, 23.05.2023; p—Russia, N Caucasus, Karachay-Cherkes Republic, Zelenchuk District, Dausuz village W environs, S banks of a reservoir at the Bolzhoy Zelenchuk River, 1012 m a.s.l.; q—Russia, Dagestan, Makhachkala N env., Muraveynik Protected Area at the Sulak River mouth, pools among coastal dunes, 1.06.2021. Scale bar 0.5 mm." figureDoi="http://doi.org/10.5281/zenodo.12189912" httpUri="https://zenodo.org/record/12189912/files/figure.png" pageId="34" pageNumber="185">Fig. 3lm</figureCitation>
), narrowly processed in
<taxonomicName id="94A64D749133F13CA6A3F9F178E17EBF" authorityName="Vander Linden" authorityYear="1825" box="[487,644,223,249]" class="Insecta" family="Coenagrionidae" genus="Agrion" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="34" pageNumber="185" phylum="Arthropoda" rank="species" species="pulchellum">
<emphasis id="61D2EAE59133F13CA6A3F9F178E17EBF" box="[487,644,223,249]" italics="true" pageId="34" pageNumber="185">C. pulchellum</emphasis>
</taxonomicName>
(
<figureCitation id="CB9D2A729133F13CA5D0F9F179677EBF" box="[660,770,223,249]" captionStart="FIGURE3" captionStartId="10.[151,250,1432,1456]" captionTargetBox="[393,1180,188,1398]" captionTargetId="figure-24@10.[393,1194,181,1408]" captionTargetPageId="10" captionText="FIGURE3. Maleappendages of Coenagrion spp.ofthe puella-groupin thebroadsense inlateral view:ac—C.australocaspicum (b—with the appendages expanded); dk—C. puella; lm—C. ponticum (males with a more (c) and less (d) expanded black abdominal maculation); n—C. persicum; oq—C. pulchellum; ab—Russia, Dagestan, Magaramkent District, Samur Delta Forest, 2 km NW of Primorskiy village, a swampy estuary of a forest rivulet, 24.06.2021; c—Russia, Dagestan, Kumtorkala District, the Shura-Ozen River at Kumtorkala Station, next to Sarykum Barkhan, 3.05.2021; d —Georgia, Dzau District, 4 km NNE Kvaisa, Lake Koz (Kvedi), 2830.06.2021; ef—Russia, Dagestan, Rutul District; the Samur River left arm between Kufa and Kala villages, 26.06.2021; gh—Russia, Dagestan, Magaramkent District, Samur Delta Forest, Bil-Bil Kazmalyar village, 25.06.2021; i—Russia, Dagestan, Buynaksk District, Buynaksk S environs, Nizhnee Kazanishche village environs, a water reservoir, 2020; k—Russia, Karachay-Cherkes Republic, Zelenchuk District, 10 km WSW of Arkhyz, Lunnaya Polyana terrain, pools in the Dukka River valley left side, 2.07.2019; k—Russia, Republic of Bashkortostan, Zinachurinskiy District, a small roadside oxbow lake 0.7 km E of Verkhniy Sarabil village, 5.07.2021; lm—Russia, Krasnodarskiy Kray, Goryachiy Klyuch Municipality, Fanagoriyskoe village NW environs, ponds, 27.04.2019; n—Iran, Markazi Ostan, Shazand Shakhrestan, Emarat village env., a spring and swampy reservoir, 20.05.2017; o—Russia, Dagestan, Makhachkala N suburbs, pools between dunes at the Krivaya Balka River left bank in its lowermost reaches, 23.05.2023; p—Russia, N Caucasus, Karachay-Cherkes Republic, Zelenchuk District, Dausuz village W environs, S banks of a reservoir at the Bolzhoy Zelenchuk River, 1012 m a.s.l.; q—Russia, Dagestan, Makhachkala N env., Muraveynik Protected Area at the Sulak River mouth, pools among coastal dunes, 1.06.2021. Scale bar 0.5 mm." figureDoi="http://doi.org/10.5281/zenodo.12189912" httpUri="https://zenodo.org/record/12189912/files/figure.png" pageId="34" pageNumber="185">Fig. 3oq</figureCitation>
) and scarcely convex and gently rounded in
<taxonomicName id="94A64D749133F13CA3BCF9F17F3A7EBF" authorityName="&quot; (Skvortsov &amp; Snegovaya" authorityYear="2015" box="[1272,1375,223,249]" class="Insecta" family="Coenagrionidae" genus="Agrion" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="34" pageNumber="185" phylum="Arthropoda" rank="species" species="puella">
<emphasis id="61D2EAE59133F13CA3BCF9F17F3A7EBF" box="[1272,1375,223,249]" italics="true" pageId="34" pageNumber="185">C. puella</emphasis>
</taxonomicName>
(
<figureCitation id="CB9D2A729133F13CA22BF9F17AB67F5B" captionStart="FIGURE3" captionStartId="10.[151,250,1432,1456]" captionTargetBox="[393,1180,188,1398]" captionTargetId="figure-24@10.[393,1194,181,1408]" captionTargetPageId="10" captionText="FIGURE3. Maleappendages of Coenagrion spp.ofthe puella-groupin thebroadsense inlateral view:ac—C.australocaspicum (b—with the appendages expanded); dk—C. puella; lm—C. ponticum (males with a more (c) and less (d) expanded black abdominal maculation); n—C. persicum; oq—C. pulchellum; ab—Russia, Dagestan, Magaramkent District, Samur Delta Forest, 2 km NW of Primorskiy village, a swampy estuary of a forest rivulet, 24.06.2021; c—Russia, Dagestan, Kumtorkala District, the Shura-Ozen River at Kumtorkala Station, next to Sarykum Barkhan, 3.05.2021; d —Georgia, Dzau District, 4 km NNE Kvaisa, Lake Koz (Kvedi), 2830.06.2021; ef—Russia, Dagestan, Rutul District; the Samur River left arm between Kufa and Kala villages, 26.06.2021; gh—Russia, Dagestan, Magaramkent District, Samur Delta Forest, Bil-Bil Kazmalyar village, 25.06.2021; i—Russia, Dagestan, Buynaksk District, Buynaksk S environs, Nizhnee Kazanishche village environs, a water reservoir, 2020; k—Russia, Karachay-Cherkes Republic, Zelenchuk District, 10 km WSW of Arkhyz, Lunnaya Polyana terrain, pools in the Dukka River valley left side, 2.07.2019; k—Russia, Republic of Bashkortostan, Zinachurinskiy District, a small roadside oxbow lake 0.7 km E of Verkhniy Sarabil village, 5.07.2021; lm—Russia, Krasnodarskiy Kray, Goryachiy Klyuch Municipality, Fanagoriyskoe village NW environs, ponds, 27.04.2019; n—Iran, Markazi Ostan, Shazand Shakhrestan, Emarat village env., a spring and swampy reservoir, 20.05.2017; o—Russia, Dagestan, Makhachkala N suburbs, pools between dunes at the Krivaya Balka River left bank in its lowermost reaches, 23.05.2023; p—Russia, N Caucasus, Karachay-Cherkes Republic, Zelenchuk District, Dausuz village W environs, S banks of a reservoir at the Bolzhoy Zelenchuk River, 1012 m a.s.l.; q—Russia, Dagestan, Makhachkala N env., Muraveynik Protected Area at the Sulak River mouth, pools among coastal dunes, 1.06.2021. Scale bar 0.5 mm." figureDoi="http://doi.org/10.5281/zenodo.12189912" httpUri="https://zenodo.org/record/12189912/files/figure.png" pageId="34" pageNumber="185">Fig. 3dk</figureCitation>
).
</paragraph>
<paragraph id="531936F79133F13CA783F8097F267FCF" blockId="34.[151,1437,151,2013]" pageId="34" pageNumber="185">
—Male paraprocts scarcely slanting, directed more behind than up, while strongly slanting, directed more up than behind in
<taxonomicName id="94A64D749133F13CA604F8657BB87F23" authorityName="Vander Linden" authorityYear="1825" box="[320,477,331,357]" class="Insecta" family="Coenagrionidae" genus="Agrion" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="34" pageNumber="185" phylum="Arthropoda" rank="species" species="pulchellum">
<emphasis id="61D2EAE59133F13CA604F8657BB87F23" box="[320,477,331,357]" italics="true" pageId="34" pageNumber="185">C. pulchellum</emphasis>
</taxonomicName>
and almost horizontal in
<taxonomicName id="94A64D749133F13CA5B9F86579017F23" authorityName="&quot; (Skvortsov &amp; Snegovaya" authorityYear="2015" box="[765,868,331,357]" class="Insecta" family="Coenagrionidae" genus="Agrion" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="34" pageNumber="185" phylum="Arthropoda" rank="species" species="puella">
<emphasis id="61D2EAE59133F13CA5B9F86579017F23" box="[765,868,331,357]" italics="true" pageId="34" pageNumber="185">C. puella</emphasis>
</taxonomicName>
. We find that
<taxonomicName id="94A64D749133F13CA340F8657E0E7F23" authorityName="&quot; (Skvortsov &amp; Snegovaya" authorityYear="2015" box="[1028,1131,331,357]" class="Insecta" family="Coenagrionidae" genus="Agrion" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="34" pageNumber="185" phylum="Arthropoda" rank="species" species="puella">
<emphasis id="61D2EAE59133F13CA340F8657E0E7F23" box="[1028,1131,331,357]" italics="true" pageId="34" pageNumber="185">C. puella</emphasis>
</taxonomicName>
(
<figureCitation id="CB9D2A729133F13CA33FF8657E8C7F23" box="[1147,1257,331,357]" captionStart="FIGURE3" captionStartId="10.[151,250,1432,1456]" captionTargetBox="[393,1180,188,1398]" captionTargetId="figure-24@10.[393,1194,181,1408]" captionTargetPageId="10" captionText="FIGURE3. Maleappendages of Coenagrion spp.ofthe puella-groupin thebroadsense inlateral view:ac—C.australocaspicum (b—with the appendages expanded); dk—C. puella; lm—C. ponticum (males with a more (c) and less (d) expanded black abdominal maculation); n—C. persicum; oq—C. pulchellum; ab—Russia, Dagestan, Magaramkent District, Samur Delta Forest, 2 km NW of Primorskiy village, a swampy estuary of a forest rivulet, 24.06.2021; c—Russia, Dagestan, Kumtorkala District, the Shura-Ozen River at Kumtorkala Station, next to Sarykum Barkhan, 3.05.2021; d —Georgia, Dzau District, 4 km NNE Kvaisa, Lake Koz (Kvedi), 2830.06.2021; ef—Russia, Dagestan, Rutul District; the Samur River left arm between Kufa and Kala villages, 26.06.2021; gh—Russia, Dagestan, Magaramkent District, Samur Delta Forest, Bil-Bil Kazmalyar village, 25.06.2021; i—Russia, Dagestan, Buynaksk District, Buynaksk S environs, Nizhnee Kazanishche village environs, a water reservoir, 2020; k—Russia, Karachay-Cherkes Republic, Zelenchuk District, 10 km WSW of Arkhyz, Lunnaya Polyana terrain, pools in the Dukka River valley left side, 2.07.2019; k—Russia, Republic of Bashkortostan, Zinachurinskiy District, a small roadside oxbow lake 0.7 km E of Verkhniy Sarabil village, 5.07.2021; lm—Russia, Krasnodarskiy Kray, Goryachiy Klyuch Municipality, Fanagoriyskoe village NW environs, ponds, 27.04.2019; n—Iran, Markazi Ostan, Shazand Shakhrestan, Emarat village env., a spring and swampy reservoir, 20.05.2017; o—Russia, Dagestan, Makhachkala N suburbs, pools between dunes at the Krivaya Balka River left bank in its lowermost reaches, 23.05.2023; p—Russia, N Caucasus, Karachay-Cherkes Republic, Zelenchuk District, Dausuz village W environs, S banks of a reservoir at the Bolzhoy Zelenchuk River, 1012 m a.s.l.; q—Russia, Dagestan, Makhachkala N env., Muraveynik Protected Area at the Sulak River mouth, pools among coastal dunes, 1.06.2021. Scale bar 0.5 mm." figureDoi="http://doi.org/10.5281/zenodo.12189912" httpUri="https://zenodo.org/record/12189912/files/figure.png" pageId="34" pageNumber="185">Fig. 3dk</figureCitation>
) does not differ much from
<taxonomicName id="94A64D749133F13CA65EF8417BC77FCE" authorityName="Kosterin &amp; Onishko &amp; Ilyina &amp; Chepurnov &amp; Blinov" authorityYear="2024" box="[282,418,367,393]" class="Insecta" family="Coenagrionidae" genus="Agrion" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="34" pageNumber="185" phylum="Arthropoda" rank="species" species="ponticum">
<emphasis id="61D2EAE59133F13CA65EF8417BC77FCE" box="[282,418,367,393]" italics="true" pageId="34" pageNumber="185">C. ponticum</emphasis>
</taxonomicName>
(
<figureCitation id="CB9D2A729133F13CA6F6F84178477FCF" box="[434,546,367,393]" captionStart="FIGURE3" captionStartId="10.[151,250,1432,1456]" captionTargetBox="[393,1180,188,1398]" captionTargetId="figure-24@10.[393,1194,181,1408]" captionTargetPageId="10" captionText="FIGURE3. Maleappendages of Coenagrion spp.ofthe puella-groupin thebroadsense inlateral view:ac—C.australocaspicum (b—with the appendages expanded); dk—C. puella; lm—C. ponticum (males with a more (c) and less (d) expanded black abdominal maculation); n—C. persicum; oq—C. pulchellum; ab—Russia, Dagestan, Magaramkent District, Samur Delta Forest, 2 km NW of Primorskiy village, a swampy estuary of a forest rivulet, 24.06.2021; c—Russia, Dagestan, Kumtorkala District, the Shura-Ozen River at Kumtorkala Station, next to Sarykum Barkhan, 3.05.2021; d —Georgia, Dzau District, 4 km NNE Kvaisa, Lake Koz (Kvedi), 2830.06.2021; ef—Russia, Dagestan, Rutul District; the Samur River left arm between Kufa and Kala villages, 26.06.2021; gh—Russia, Dagestan, Magaramkent District, Samur Delta Forest, Bil-Bil Kazmalyar village, 25.06.2021; i—Russia, Dagestan, Buynaksk District, Buynaksk S environs, Nizhnee Kazanishche village environs, a water reservoir, 2020; k—Russia, Karachay-Cherkes Republic, Zelenchuk District, 10 km WSW of Arkhyz, Lunnaya Polyana terrain, pools in the Dukka River valley left side, 2.07.2019; k—Russia, Republic of Bashkortostan, Zinachurinskiy District, a small roadside oxbow lake 0.7 km E of Verkhniy Sarabil village, 5.07.2021; lm—Russia, Krasnodarskiy Kray, Goryachiy Klyuch Municipality, Fanagoriyskoe village NW environs, ponds, 27.04.2019; n—Iran, Markazi Ostan, Shazand Shakhrestan, Emarat village env., a spring and swampy reservoir, 20.05.2017; o—Russia, Dagestan, Makhachkala N suburbs, pools between dunes at the Krivaya Balka River left bank in its lowermost reaches, 23.05.2023; p—Russia, N Caucasus, Karachay-Cherkes Republic, Zelenchuk District, Dausuz village W environs, S banks of a reservoir at the Bolzhoy Zelenchuk River, 1012 m a.s.l.; q—Russia, Dagestan, Makhachkala N env., Muraveynik Protected Area at the Sulak River mouth, pools among coastal dunes, 1.06.2021. Scale bar 0.5 mm." figureDoi="http://doi.org/10.5281/zenodo.12189912" httpUri="https://zenodo.org/record/12189912/files/figure.png" pageId="34" pageNumber="185">Fig. 3lm</figureCitation>
) in this respect, since the former also may have the paraprocts slanting.
</paragraph>
<paragraph id="531936F79133F13CA783F8BD79FE7FEB" blockId="34.[151,1437,151,2013]" box="[199,923,403,429]" pageId="34" pageNumber="185">
The structural characters of
<taxonomicName id="94A64D749133F13CA545F8BD78EF7FEA" authorityName="Kosterin &amp; Onishko &amp; Ilyina &amp; Chepurnov &amp; Blinov" authorityYear="2024" box="[513,650,403,429]" class="Insecta" family="Coenagrionidae" genus="Agrion" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="34" pageNumber="185" phylum="Arthropoda" rank="species" species="ponticum">
<emphasis id="61D2EAE59133F13CA545F8BD78EF7FEA" box="[513,650,403,429]" italics="true" pageId="34" pageNumber="185">C. ponticum</emphasis>
</taxonomicName>
as in
<taxonomicName id="94A64D749133F13CA588F8BD790C7FEB" authorityName="Vander Linden" authorityYear="1825" box="[716,873,403,429]" class="Insecta" family="Coenagrionidae" genus="Agrion" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="34" pageNumber="185" phylum="Arthropoda" rank="species" species="pulchellum">
<emphasis id="61D2EAE59133F13CA588F8BD790C7FEB" box="[716,873,403,429]" italics="true" pageId="34" pageNumber="185">C. pulchellum</emphasis>
</taxonomicName>
are:
</paragraph>
<paragraph id="531936F79133F13CA783F89978A57FB3" blockId="34.[151,1437,151,2013]" pageId="34" pageNumber="185">
—Male S10 hind margin in lateral view with two bends (one in
<taxonomicName id="94A64D749133F13CA4E5F8997E6C7F97" authorityName="&quot; (Skvortsov &amp; Snegovaya" authorityYear="2015" box="[929,1033,439,465]" class="Insecta" family="Coenagrionidae" genus="Agrion" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="34" pageNumber="185" phylum="Arthropoda" rank="species" species="puella">
<emphasis id="61D2EAE59133F13CA4E5F8997E6C7F97" box="[929,1033,439,465]" italics="true" pageId="34" pageNumber="185">C. puella</emphasis>
</taxonomicName>
). We find those bends scarcely and variably expressed and do not see this difference.
</paragraph>
<paragraph id="531936F79133F13CA783F8D179007C7B" blockId="34.[151,1437,151,2013]" pageId="34" pageNumber="185">
—Cercus process long and straight, reaching paraproct projection (
<figureCitation id="CB9D2A729133F13CA4A6F8D17E327C5F" box="[994,1111,511,537]" captionStart="FIGURE 4" captionStartId="11.[151,250,1054,1078]" captionTargetBox="[189,1398,181,1021]" captionTargetId="figure-19@11.[189,1398,181,1030]" captionTargetPageId="11" captionText="FIGURE 4. Male appendages of Coenagrion spp. of the puella-group in the broad sense in posterior view: ab—C. australocaspicum (b—with the appendages expanded); cd—C. ponticum (males with a more (c) and less (d) expanded black abdominal maculation); e-f—C. pulchellum; gk—C. puella; l—C. persicum; ab—Russia, Dagestan, Magaramkent District, Samur Delta Forest, 2 km NW of Primorskiy village, a swampy estuary of a forest rivulet, 24.06.2021; cd—Russia, Krasnodarskiy Kray, Goryachiy Klyuch Municipality, Fanagoriyskoe village NW environs, ponds, 27.04.2019; e—Russia, Dagestan, Makhachkala N suburbs, pools between dunes at the Krivaya Balka River left bank in its lowermost reaches, 23.05.2023; f—Russia, Dagestan, Makhachkala N env., Muraveynik Protected Area at the Sulak River mouth, pools among coastal dunes, 1.06.2021; gh—Russia, Dagestan, Rutul District; the Samur River left arm between Kufa and Kala villages, 26.06.2021; i—Russia, Dagestan, Magaramkent District, Samur Delta Forest, Bil-Bil Kazmalyar village, a brook and pond, 25.06.2021; j—Karachay-Cherkes Republic, Zelenchuk District, 10 km WSW of Arkhyz, Lunnaya Polyana terrain, pools in the Dukka River valley left side, 2.07.2019; k—Russia, Republic of Bashkortostan, Zinachurinskiy District, a small roadside oxbow lake 0.7 km E of Verkhniy Sarabil village, 5.07.2021; l—Iran, Markazi Ostan, Shazand Shakhrestan, Emarat village env., a spring and swampy reservoir, 20.05.2017. Scale bar 0.5 mm." figureDoi="http://doi.org/10.5281/zenodo.12189914" httpUri="https://zenodo.org/record/12189914/files/figure.png" pageId="34" pageNumber="185">Fig. 4cd</figureCitation>
), said to be short (scarcely expanding beyond tubercle) and curved in
<taxonomicName id="94A64D749133F13CA532FB0D78B87C7B" authorityName="&quot; (Skvortsov &amp; Snegovaya" authorityYear="2015" box="[630,733,547,573]" class="Insecta" family="Coenagrionidae" genus="Agrion" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="34" pageNumber="185" phylum="Arthropoda" rank="species" species="puella">
<emphasis id="61D2EAE59133F13CA532FB0D78B87C7B" box="[630,733,547,573]" italics="true" pageId="34" pageNumber="185">C. puella</emphasis>
</taxonomicName>
(
<figureCitation id="CB9D2A729133F13CA5A8FB0D793D7C7B" box="[748,856,547,573]" captionStart="FIGURE 4" captionStartId="11.[151,250,1054,1078]" captionTargetBox="[189,1398,181,1021]" captionTargetId="figure-19@11.[189,1398,181,1030]" captionTargetPageId="11" captionText="FIGURE 4. Male appendages of Coenagrion spp. of the puella-group in the broad sense in posterior view: ab—C. australocaspicum (b—with the appendages expanded); cd—C. ponticum (males with a more (c) and less (d) expanded black abdominal maculation); e-f—C. pulchellum; gk—C. puella; l—C. persicum; ab—Russia, Dagestan, Magaramkent District, Samur Delta Forest, 2 km NW of Primorskiy village, a swampy estuary of a forest rivulet, 24.06.2021; cd—Russia, Krasnodarskiy Kray, Goryachiy Klyuch Municipality, Fanagoriyskoe village NW environs, ponds, 27.04.2019; e—Russia, Dagestan, Makhachkala N suburbs, pools between dunes at the Krivaya Balka River left bank in its lowermost reaches, 23.05.2023; f—Russia, Dagestan, Makhachkala N env., Muraveynik Protected Area at the Sulak River mouth, pools among coastal dunes, 1.06.2021; gh—Russia, Dagestan, Rutul District; the Samur River left arm between Kufa and Kala villages, 26.06.2021; i—Russia, Dagestan, Magaramkent District, Samur Delta Forest, Bil-Bil Kazmalyar village, a brook and pond, 25.06.2021; j—Karachay-Cherkes Republic, Zelenchuk District, 10 km WSW of Arkhyz, Lunnaya Polyana terrain, pools in the Dukka River valley left side, 2.07.2019; k—Russia, Republic of Bashkortostan, Zinachurinskiy District, a small roadside oxbow lake 0.7 km E of Verkhniy Sarabil village, 5.07.2021; l—Iran, Markazi Ostan, Shazand Shakhrestan, Emarat village env., a spring and swampy reservoir, 20.05.2017. Scale bar 0.5 mm." figureDoi="http://doi.org/10.5281/zenodo.12189914" httpUri="https://zenodo.org/record/12189914/files/figure.png" pageId="34" pageNumber="185">Fig. 4fk</figureCitation>
).
</paragraph>
<paragraph id="531936F79133F13CA783FB6979017C27" blockId="34.[151,1437,151,2013]" box="[199,868,583,609]" pageId="34" pageNumber="185">
The structural characters of
<taxonomicName id="94A64D749133F13CA545FB6978EF7C26" authorityName="Kosterin &amp; Onishko &amp; Ilyina &amp; Chepurnov &amp; Blinov" authorityYear="2024" box="[513,650,583,609]" class="Insecta" family="Coenagrionidae" genus="Agrion" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="34" pageNumber="185" phylum="Arthropoda" rank="species" species="ponticum">
<emphasis id="61D2EAE59133F13CA545FB6978EF7C26" box="[513,650,583,609]" italics="true" pageId="34" pageNumber="185">C. ponticum</emphasis>
</taxonomicName>
as in
<taxonomicName id="94A64D749133F13CA588FB6979567C27" authorityName="&quot; (Skvortsov &amp; Snegovaya" authorityYear="2015" box="[716,819,583,609]" class="Insecta" family="Coenagrionidae" genus="Agrion" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="34" pageNumber="185" phylum="Arthropoda" rank="species" species="puella">
<emphasis id="61D2EAE59133F13CA588FB6979567C27" box="[716,819,583,609]" italics="true" pageId="34" pageNumber="185">C. puella</emphasis>
</taxonomicName>
are:
</paragraph>
<paragraph id="531936F79133F13CA783FB457F1D7CEF" blockId="34.[151,1437,151,2013]" pageId="34" pageNumber="185">
—Tubercle present, and said to absent from
<taxonomicName id="94A64D749133F13CA58CFB45790D7CC3" authorityName="Vander Linden" authorityYear="1825" box="[712,872,619,645]" class="Insecta" family="Coenagrionidae" genus="Agrion" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="34" pageNumber="185" phylum="Arthropoda" rank="species" species="pulchellum">
<emphasis id="61D2EAE59133F13CA58CFB45790D7CC3" box="[712,872,619,645]" italics="true" pageId="34" pageNumber="185">C. pulchellum</emphasis>
</taxonomicName>
, which is not correct: it is well expressed in this species but somewhat obscured by the trunk in dorsal view (
<figureCitation id="CB9D2A729133F13CA47FFBA179CE7CEF" box="[827,939,655,681]" captionStart="FIGURE 2" captionStartId="9.[151,250,1421,1445]" captionTargetBox="[381,1200,187,1391]" captionTargetId="figure-19@9.[381,1206,181,1397]" captionTargetPageId="9" captionText="FIGURE 2. Male appendages of Coenagrion spp. of the puella-group in the broad sense, from the Caucasus and elsewhere, in dorsal view: ac—C. australocaspicum (b—with the appendages expanded); d-k—C. puella; lm—C. ponticum (males with a more (c) and less (d) expanded black abdominal maculation); n—C. persicum; op—C. pulchellum; ab—Russia, Dagestan, Magaramkent District, Samur Delta Forest, 2 km NW of Primorskiy village, a swampy estuary of a forest rivulet, 24.06.2021; c—Russia, Dagestan, Kumtorkala District, the Shura-Ozen River at Kumtorkala Station, next to Sarykum Barkhan, 3.05.2021; d—Georgia, Dzau District, 4 km NNE Kvaisa, Lake Koz (Kvedi), 2830.06.2021; ef—Russia, Dagestan, Rutul District; the Samur River left arm between Kufa and Kala villages, 26.06.2021; gh—Russia, Dagestan, Magaramkent District, Samur Delta Forest, Bil-Bil Kazmalyar village, 25.06.2021; i—Russia, Dagestan, Buynaksk District, Buynaksk S environs, Nizhnee Kazanishche village environs, a water reservoir, 2020; j—Russia, Karachay-Cherkes Republic, Zelenchuk District, 10 km WSW of Arkhyz, Lunnaya Polyana terrain, pools in the Dukka River valley left side, 2.07.2019; k—Russia, Republic of Bashkortostan, Zinachurinskiy District, a small roadside oxbow lake 0.7 km E of Verkhniy Sarabil village, 5.07.2021; lm— Russia, Krasnodarskiy Kray, Goryachiy Klyuch Municipality, Fanagoriyskoe village NW environs, ponds, 27.04.2019; n—Iran, Markazi Ostan, Shazand Shakhrestan, Emarat village env., a spring and swampy reservoir, 20.05.2017; o—Russia, Dagestan, Makhachkala N suburbs, pools between dunes at the Krivaya Balka River left bank in its lowermost reaches, 23.05.2023; p—Russia, N Caucasus, Karachay-Cherkes Republic, Zelenchuk District, Dausuz village W environs, S banks of a reservoir at the Bolzhoy Zelenchuk River, 1012 m a.s.l.; q—Russia, Dagestan, Makhachkala N env., Muraveynik Protected Area at the Sulak River mouth, pools among coastal dunes, 1.06.2021. Scale bar 0.5 mm." figureDoi="http://doi.org/10.5281/zenodo.12189910" httpUri="https://zenodo.org/record/12189910/files/figure.png" pageId="34" pageNumber="185">Fig. 2op</figureCitation>
). So this character by Bartenev is wrong.
</paragraph>
<paragraph id="531936F79133F13CA783FB9D79617CB7" blockId="34.[151,1437,151,2013]" pageId="34" pageNumber="185">
—Male paraproct 1.52 times as long as S
<quantity id="945E9B129133F13CA5DDFB9D78B77C8B" box="[665,722,691,717]" metricMagnitude="-1" metricUnit="m" metricValue="2.54" pageId="34" pageNumber="185" unit="in" value="10.0">10 in</quantity>
lateral and dorsal views, versus as long as in
<emphasis id="61D2EAE59133F13CA380FB9D7F1C7C8B" box="[1220,1401,691,717]" italics="true" pageId="34" pageNumber="185">
<taxonomicName id="94A64D749133F13CA380FB9D7F397C8B" authorityName="Vander Linden" authorityYear="1825" box="[1220,1372,691,717]" class="Insecta" family="Coenagrionidae" genus="Agrion" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="34" pageNumber="185" phylum="Arthropoda" rank="species" species="pulchellum">C. pulchellum</taxonomicName>
</emphasis>
but this is a matter of its direction rather than actual length.
</paragraph>
<paragraph id="531936F79133F13CA783FBD57E187D7F" blockId="34.[151,1437,151,2013]" pageId="34" pageNumber="185">
—Male paraproct prominence shorter than cercus (as long in
<taxonomicName id="94A64D749133F13CA4D1FBD57E507D53" authorityName="Vander Linden" authorityYear="1825" box="[917,1077,763,789]" class="Insecta" family="Coenagrionidae" genus="Agrion" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="34" pageNumber="185" phylum="Arthropoda" rank="species" species="pulchellum">
<emphasis id="61D2EAE59133F13CA4D1FBD57E507D53" box="[917,1077,763,789]" italics="true" pageId="34" pageNumber="185">C. pulchellum</emphasis>
</taxonomicName>
). This character works but the prominence is yet somewhat shorter than the cercus in
<taxonomicName id="94A64D749133F13CA5BAFA3179FE7D7F" authorityName="Vander Linden" authorityYear="1825" box="[766,923,799,825]" class="Insecta" family="Coenagrionidae" genus="Agrion" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="34" pageNumber="185" phylum="Arthropoda" rank="species" species="pulchellum">
<emphasis id="61D2EAE59133F13CA5BAFA3179FE7D7F" box="[766,923,799,825]" italics="true" pageId="34" pageNumber="185">C. pulchellum</emphasis>
</taxonomicName>
as well (
<figureCitation id="CB9D2A729133F13CA344FA317E147D7F" box="[1024,1137,799,825]" captionStart="FIGURE3" captionStartId="10.[151,250,1432,1456]" captionTargetBox="[393,1180,188,1398]" captionTargetId="figure-24@10.[393,1194,181,1408]" captionTargetPageId="10" captionText="FIGURE3. Maleappendages of Coenagrion spp.ofthe puella-groupin thebroadsense inlateral view:ac—C.australocaspicum (b—with the appendages expanded); dk—C. puella; lm—C. ponticum (males with a more (c) and less (d) expanded black abdominal maculation); n—C. persicum; oq—C. pulchellum; ab—Russia, Dagestan, Magaramkent District, Samur Delta Forest, 2 km NW of Primorskiy village, a swampy estuary of a forest rivulet, 24.06.2021; c—Russia, Dagestan, Kumtorkala District, the Shura-Ozen River at Kumtorkala Station, next to Sarykum Barkhan, 3.05.2021; d —Georgia, Dzau District, 4 km NNE Kvaisa, Lake Koz (Kvedi), 2830.06.2021; ef—Russia, Dagestan, Rutul District; the Samur River left arm between Kufa and Kala villages, 26.06.2021; gh—Russia, Dagestan, Magaramkent District, Samur Delta Forest, Bil-Bil Kazmalyar village, 25.06.2021; i—Russia, Dagestan, Buynaksk District, Buynaksk S environs, Nizhnee Kazanishche village environs, a water reservoir, 2020; k—Russia, Karachay-Cherkes Republic, Zelenchuk District, 10 km WSW of Arkhyz, Lunnaya Polyana terrain, pools in the Dukka River valley left side, 2.07.2019; k—Russia, Republic of Bashkortostan, Zinachurinskiy District, a small roadside oxbow lake 0.7 km E of Verkhniy Sarabil village, 5.07.2021; lm—Russia, Krasnodarskiy Kray, Goryachiy Klyuch Municipality, Fanagoriyskoe village NW environs, ponds, 27.04.2019; n—Iran, Markazi Ostan, Shazand Shakhrestan, Emarat village env., a spring and swampy reservoir, 20.05.2017; o—Russia, Dagestan, Makhachkala N suburbs, pools between dunes at the Krivaya Balka River left bank in its lowermost reaches, 23.05.2023; p—Russia, N Caucasus, Karachay-Cherkes Republic, Zelenchuk District, Dausuz village W environs, S banks of a reservoir at the Bolzhoy Zelenchuk River, 1012 m a.s.l.; q—Russia, Dagestan, Makhachkala N env., Muraveynik Protected Area at the Sulak River mouth, pools among coastal dunes, 1.06.2021. Scale bar 0.5 mm." figureDoi="http://doi.org/10.5281/zenodo.12189912" httpUri="https://zenodo.org/record/12189912/files/figure.png" pageId="34" pageNumber="185">Fig. 3oq</figureCitation>
).
</paragraph>
<paragraph id="531936F79133F13CA783FA6D79617DAB" blockId="34.[151,1437,151,2013]" pageId="34" pageNumber="185">
It is noteworthy that Bartenevs drawing (
<bibRefCitation id="37374B069133F13CA5D0FA6D795D7D1B" author="Bartenef, A. N." box="[660,824,835,861]" pageId="34" pageNumber="185" pagination="54 - 68" refId="ref27207" refString="Bartenef, A. N. (1929) Neue Arten und Varietaten der Odonata des West Kaukasus. Zoologischer Anzeiger, 85, 54 - 68." type="journal article" year="1929">Bartenef 1929</bibRefCitation>
: abb. 13; 1930: fig. 28) show the paraproct prominence rounded (more like in
<taxonomicName id="94A64D749133F13CA6D7FA497B9F7DC7" authorityName="&quot; (Skvortsov &amp; Snegovaya" authorityYear="2015" box="[403,506,871,897]" class="Insecta" family="Coenagrionidae" genus="Agrion" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="34" pageNumber="185" phylum="Arthropoda" rank="species" species="puella">
<emphasis id="61D2EAE59133F13CA6D7FA497B9F7DC7" box="[403,506,871,897]" italics="true" pageId="34" pageNumber="185">C. puella</emphasis>
</taxonomicName>
but better expressed) rather than processed as in
<taxonomicName id="94A64D749133F13CA367FA497EA57DC7" authorityName="Vander Linden" authorityYear="1825" box="[1059,1216,871,897]" class="Insecta" family="Coenagrionidae" genus="Agrion" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="34" pageNumber="185" phylum="Arthropoda" rank="species" species="pulchellum">
<emphasis id="61D2EAE59133F13CA367FA497EA57DC7" box="[1059,1216,871,897]" italics="true" pageId="34" pageNumber="185">C. pulchellum</emphasis>
</taxonomicName>
, as observed in our specimens of
<taxonomicName id="94A64D749133F13CA671FAA57BA57DE2" authorityName="Kosterin &amp; Onishko &amp; Ilyina &amp; Chepurnov &amp; Blinov" authorityYear="2024" box="[309,448,907,933]" class="Insecta" family="Coenagrionidae" genus="Agrion" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="34" pageNumber="185" phylum="Arthropoda" rank="species" species="ponticum">
<emphasis id="61D2EAE59133F13CA671FAA57BA57DE2" box="[309,448,907,933]" italics="true" pageId="34" pageNumber="185">C. ponticum</emphasis>
</taxonomicName>
(
<figureCitation id="CB9D2A729133F13CA696FAA578237DE3" box="[466,582,907,933]" captionStart="FIGURE3" captionStartId="10.[151,250,1432,1456]" captionTargetBox="[393,1180,188,1398]" captionTargetId="figure-24@10.[393,1194,181,1408]" captionTargetPageId="10" captionText="FIGURE3. Maleappendages of Coenagrion spp.ofthe puella-groupin thebroadsense inlateral view:ac—C.australocaspicum (b—with the appendages expanded); dk—C. puella; lm—C. ponticum (males with a more (c) and less (d) expanded black abdominal maculation); n—C. persicum; oq—C. pulchellum; ab—Russia, Dagestan, Magaramkent District, Samur Delta Forest, 2 km NW of Primorskiy village, a swampy estuary of a forest rivulet, 24.06.2021; c—Russia, Dagestan, Kumtorkala District, the Shura-Ozen River at Kumtorkala Station, next to Sarykum Barkhan, 3.05.2021; d —Georgia, Dzau District, 4 km NNE Kvaisa, Lake Koz (Kvedi), 2830.06.2021; ef—Russia, Dagestan, Rutul District; the Samur River left arm between Kufa and Kala villages, 26.06.2021; gh—Russia, Dagestan, Magaramkent District, Samur Delta Forest, Bil-Bil Kazmalyar village, 25.06.2021; i—Russia, Dagestan, Buynaksk District, Buynaksk S environs, Nizhnee Kazanishche village environs, a water reservoir, 2020; k—Russia, Karachay-Cherkes Republic, Zelenchuk District, 10 km WSW of Arkhyz, Lunnaya Polyana terrain, pools in the Dukka River valley left side, 2.07.2019; k—Russia, Republic of Bashkortostan, Zinachurinskiy District, a small roadside oxbow lake 0.7 km E of Verkhniy Sarabil village, 5.07.2021; lm—Russia, Krasnodarskiy Kray, Goryachiy Klyuch Municipality, Fanagoriyskoe village NW environs, ponds, 27.04.2019; n—Iran, Markazi Ostan, Shazand Shakhrestan, Emarat village env., a spring and swampy reservoir, 20.05.2017; o—Russia, Dagestan, Makhachkala N suburbs, pools between dunes at the Krivaya Balka River left bank in its lowermost reaches, 23.05.2023; p—Russia, N Caucasus, Karachay-Cherkes Republic, Zelenchuk District, Dausuz village W environs, S banks of a reservoir at the Bolzhoy Zelenchuk River, 1012 m a.s.l.; q—Russia, Dagestan, Makhachkala N env., Muraveynik Protected Area at the Sulak River mouth, pools among coastal dunes, 1.06.2021. Scale bar 0.5 mm." figureDoi="http://doi.org/10.5281/zenodo.12189912" httpUri="https://zenodo.org/record/12189912/files/figure.png" pageId="34" pageNumber="185">Fig. 3lm</figureCitation>
). A similar shape is shown for “Sp. 5” from
<collectingCountry id="2BB176679133F13CA30CFAA57EA67DE3" box="[1096,1219,907,933]" name="Azerbaijan" pageId="34" pageNumber="185">Azerbaijan</collectingCountry>
associated with
<taxonomicName id="94A64D749133F13CA2C7FAA57A9A7D8E" authorityName="Kosterin &amp; Onishko &amp; Ilyina &amp; Chepurnov &amp; Blinov" authorityYear="2024" class="Insecta" family="Coenagrionidae" genus="Agrion" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="34" pageNumber="185" phylum="Arthropoda" rank="species" species="ponticum">
<emphasis id="61D2EAE59133F13CA2C7FAA57A9A7D8E" italics="true" pageId="34" pageNumber="185">C. ponticum</emphasis>
</taxonomicName>
by
<bibRefCitation id="37374B069133F13CA66CFA8178E77D8F" author="Skvortsov, V. E. &amp; Snegovaya, N. Y." box="[296,642,942,969]" pageId="34" pageNumber="185" pagination="1 - 38" refId="ref29550" refString="Skvortsov, V. E. &amp; Snegovaya, N. Y. (2015) A second addition to the Odonata fauna of Azerbaijan. International Dragonfly Fund Report, 87, 1 - 38." type="journal article" year="2015">Skvortsov &amp; Snegovaya (2015</bibRefCitation>
: fig. 28). This may indicate that populations of
<taxonomicName id="94A64D749133F13CA3D4FA817F7D7D8E" authorityName="Kosterin &amp; Onishko &amp; Ilyina &amp; Chepurnov &amp; Blinov" authorityYear="2024" box="[1168,1304,943,969]" class="Insecta" family="Coenagrionidae" genus="Agrion" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="34" pageNumber="185" phylum="Arthropoda" rank="species" species="ponticum">
<emphasis id="61D2EAE59133F13CA3D4FA817F7D7D8E" box="[1168,1304,943,969]" italics="true" pageId="34" pageNumber="185">C. ponticum</emphasis>
</taxonomicName>
still vary in expression of the puella-like and pulchellum-like traits.
</paragraph>
<paragraph id="531936F79133F13CA783FAD97E067AE7" blockId="34.[151,1437,151,2013]" pageId="34" pageNumber="185">
Much attention was paid in the original description to fine peculiarities of the black maculation in males, just reflecting its overall expression. We find it best characterised by the extent of the black spot (its main part, not lateral processes) in S3, which, according to
<bibRefCitation id="37374B069133F13CA57EFD1178807A1F" author="Bartenef, A. N." box="[570,741,1087,1113]" pageId="34" pageNumber="185" pagination="54 - 68" refId="ref27207" refString="Bartenef, A. N. (1929) Neue Arten und Varietaten der Odonata des West Kaukasus. Zoologischer Anzeiger, 85, 54 - 68." type="journal article" year="1929">Bartenef (1929</bibRefCitation>
;
<bibRefCitation id="37374B069133F13CA5B4FD1179497A1F" author="Bartenef, A. N." box="[752,812,1087,1113]" pageId="34" pageNumber="185" pagination="1 - 138" refId="ref27235" refString="Bartenef, A. N. (1930) Materialy k poznaniyu Zapadnogo Kavkaza v odonatologicheskom otnoshenii [Materials for the knowledge on West Caucasus in odonatological respect]. Trudy Zapadnokavkazskoi assotsiatsii nauchno-issledovatel'skikh institutov [Proceedings of the West Caucasian Association of scientific Research Institutes], 72 (14), 1 - 138. [in Russian, with German summary but without German title]" type="journal article" year="1930">1930</bibRefCitation>
), occupies from 1/2 to 5/6 of its length in
<taxonomicName id="94A64D749133F13CA3BDFD117FF07A1F" authorityName="Vander Linden" authorityYear="1825" box="[1273,1429,1087,1113]" class="Insecta" family="Coenagrionidae" genus="Agrion" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="34" pageNumber="185" phylum="Arthropoda" rank="species" species="pulchellum">
<emphasis id="61D2EAE59133F13CA3BDFD117FF07A1F" box="[1273,1429,1087,1113]" italics="true" pageId="34" pageNumber="185">C. pulchellum</emphasis>
</taxonomicName>
, from 1/4 to 4/
<quantity id="945E9B129133F13CA674FD4C7B3D7A3B" box="[304,344,1122,1149]" metricMagnitude="-1" metricUnit="m" metricValue="1.27" pageId="34" pageNumber="185" unit="in" value="5.0">5 in</quantity>
<taxonomicName id="94A64D749133F13CA619FD4D7B807A3A" authorityName="Kosterin &amp; Onishko &amp; Ilyina &amp; Chepurnov &amp; Blinov" authorityYear="2024" box="[349,485,1123,1149]" class="Insecta" family="Coenagrionidae" genus="Agrion" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="34" pageNumber="185" phylum="Arthropoda" rank="species" species="ponticum">
<emphasis id="61D2EAE59133F13CA619FD4D7B807A3A" box="[349,485,1123,1149]" italics="true" pageId="34" pageNumber="185">C. ponticum</emphasis>
</taxonomicName>
and 1/
<quantity id="945E9B129133F13CA568FD4D783D7A3B" box="[556,600,1123,1149]" metricMagnitude="-1" metricUnit="m" metricValue="1.016" pageId="34" pageNumber="185" unit="in" value="4.0">4 in</quantity>
<taxonomicName id="94A64D749133F13CA519FD4D78A67A3B" authorityName="&quot; (Skvortsov &amp; Snegovaya" authorityYear="2015" box="[605,707,1123,1149]" class="Insecta" family="Coenagrionidae" genus="Agrion" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="34" pageNumber="185" phylum="Arthropoda" rank="species" species="puella">
<emphasis id="61D2EAE59133F13CA519FD4D78A67A3B" box="[605,707,1123,1149]" italics="true" pageId="34" pageNumber="185">C. puella</emphasis>
</taxonomicName>
. Hence its range in
<taxonomicName id="94A64D749133F13CA4D8FD4D7E467A3A" authorityName="Kosterin &amp; Onishko &amp; Ilyina &amp; Chepurnov &amp; Blinov" authorityYear="2024" box="[924,1059,1123,1149]" class="Insecta" family="Coenagrionidae" genus="Agrion" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="34" pageNumber="185" phylum="Arthropoda" rank="species" species="ponticum">
<emphasis id="61D2EAE59133F13CA4D8FD4D7E467A3A" box="[924,1059,1123,1149]" italics="true" pageId="34" pageNumber="185">C. ponticum</emphasis>
</taxonomicName>
, although intermediate with those of
<taxonomicName id="94A64D749133F13CA7F2FDA97B377AE7" authorityName="Vander Linden" authorityYear="1825" box="[182,338,1159,1185]" class="Insecta" family="Coenagrionidae" genus="Agrion" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="34" pageNumber="185" phylum="Arthropoda" rank="species" species="pulchellum">
<emphasis id="61D2EAE59133F13CA7F2FDA97B377AE7" box="[182,338,1159,1185]" italics="true" pageId="34" pageNumber="185">C. pulchellum</emphasis>
</taxonomicName>
and
<taxonomicName id="94A64D749133F13CA6CDFDA97B957AE7" authorityName="&quot; (Skvortsov &amp; Snegovaya" authorityYear="2015" box="[393,496,1159,1185]" class="Insecta" family="Coenagrionidae" genus="Agrion" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="34" pageNumber="185" phylum="Arthropoda" rank="species" species="puella">
<emphasis id="61D2EAE59133F13CA6CDFDA97B957AE7" box="[393,496,1159,1185]" italics="true" pageId="34" pageNumber="185">C. puella</emphasis>
</taxonomicName>
, overlaps with both and by no means can be diagnostic.
</paragraph>
<paragraph id="531936F79133F13CA783FD8579A07AAF" blockId="34.[151,1437,151,2013]" pageId="34" pageNumber="185">
Importantly, females of
<taxonomicName id="94A64D749133F13CA692FD85783A7A82" authorityName="Kosterin &amp; Onishko &amp; Ilyina &amp; Chepurnov &amp; Blinov" authorityYear="2024" box="[470,607,1195,1221]" class="Insecta" family="Coenagrionidae" genus="Agrion" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="34" pageNumber="185" phylum="Arthropoda" rank="species" species="ponticum">
<emphasis id="61D2EAE59133F13CA692FD85783A7A82" box="[470,607,1195,1221]" italics="true" pageId="34" pageNumber="185">C. ponticum</emphasis>
</taxonomicName>
have S8 black with only very small blue markings, as in
<taxonomicName id="94A64D749133F13CA3ACFD857F2A7A83" authorityName="&quot; (Skvortsov &amp; Snegovaya" authorityYear="2015" box="[1256,1359,1195,1221]" class="Insecta" family="Coenagrionidae" genus="Agrion" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="34" pageNumber="185" phylum="Arthropoda" rank="species" species="puella">
<emphasis id="61D2EAE59133F13CA3ACFD857F2A7A83" box="[1256,1359,1195,1221]" italics="true" pageId="34" pageNumber="185">C. puella</emphasis>
</taxonomicName>
, while blue with some black spots in all Caucasian specimens of
<taxonomicName id="94A64D749133F13CA465FDE179DB7AAF" authorityName="Vander Linden" authorityYear="1825" box="[801,958,1231,1257]" class="Insecta" family="Coenagrionidae" genus="Agrion" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="34" pageNumber="185" phylum="Arthropoda" rank="species" species="pulchellum">
<emphasis id="61D2EAE59133F13CA465FDE179DB7AAF" box="[801,958,1231,1257]" italics="true" pageId="34" pageNumber="185">C. pulchellum</emphasis>
</taxonomicName>
.
</paragraph>
<paragraph id="531936F79133F13CA783FDDD7EB47B13" blockId="34.[151,1437,151,2013]" pageId="34" pageNumber="185">
So, our specimens generally fit the original description of
<taxonomicName id="94A64D749133F13CA42AFDDD799F7B4A" authorityName="Kosterin &amp; Onishko &amp; Ilyina &amp; Chepurnov &amp; Blinov" authorityYear="2024" box="[878,1018,1267,1293]" class="Insecta" family="Coenagrionidae" genus="Agrion" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="34" pageNumber="185" phylum="Arthropoda" rank="species" species="ponticum">
<emphasis id="61D2EAE59133F13CA42AFDDD799F7B4A" box="[878,1018,1267,1293]" italics="true" pageId="34" pageNumber="185">C. ponticum</emphasis>
</taxonomicName>
except for some characters, and for identification of this species we recommend to use the short character annotation provided above in the annotated list and the key below rather than the big table by Bartenev overloaded with unnecessary details.
</paragraph>
<paragraph id="531936F79133F13CA783FC71796478DF" blockId="34.[151,1437,151,2013]" pageId="34" pageNumber="185">
While
<bibRefCitation id="37374B069133F13CA655FC717BA77B3F" author="Bartenef, A. N." box="[273,450,1375,1401]" pageId="34" pageNumber="185" pagination="1 - 138" refId="ref27235" refString="Bartenef, A. N. (1930) Materialy k poznaniyu Zapadnogo Kavkaza v odonatologicheskom otnoshenii [Materials for the knowledge on West Caucasus in odonatological respect]. Trudy Zapadnokavkazskoi assotsiatsii nauchno-issledovatel'skikh institutov [Proceedings of the West Caucasian Association of scientific Research Institutes], 72 (14), 1 - 138. [in Russian, with German summary but without German title]" type="journal article" year="1930">Bartenef (1930)</bibRefCitation>
considered
<taxonomicName id="94A64D749133F13CA503FC7178AA7B3E" authorityName="Kosterin &amp; Onishko &amp; Ilyina &amp; Chepurnov &amp; Blinov" authorityYear="2024" box="[583,719,1375,1401]" class="Insecta" family="Coenagrionidae" genus="Agrion" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="34" pageNumber="185" phylum="Arthropoda" rank="species" species="ponticum">
<emphasis id="61D2EAE59133F13CA503FC7178AA7B3E" box="[583,719,1375,1401]" italics="true" pageId="34" pageNumber="185">C. ponticum</emphasis>
</taxonomicName>
a very ancient endemic of the region, we find it more logical to alternatively suppose that this taxon originated from some Caucasian population which was a product of occasional hybridisation of
<taxonomicName id="94A64D749133F13CA612FC897B907B87" authorityName="Vander Linden" authorityYear="1825" box="[342,501,1447,1473]" class="Insecta" family="Coenagrionidae" genus="Agrion" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="34" pageNumber="185" phylum="Arthropoda" rank="species" species="pulchellum">
<emphasis id="61D2EAE59133F13CA612FC897B907B87" box="[342,501,1447,1473]" italics="true" pageId="34" pageNumber="185">C. pulchellum</emphasis>
</taxonomicName>
and
<taxonomicName id="94A64D749133F13CA576FC8978FE7B87" authorityName="&quot; (Skvortsov &amp; Snegovaya" authorityYear="2015" box="[562,667,1447,1473]" class="Insecta" family="Coenagrionidae" genus="Agrion" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="34" pageNumber="185" phylum="Arthropoda" rank="species" species="puella">
<emphasis id="61D2EAE59133F13CA576FC8978FE7B87" box="[562,667,1447,1473]" italics="true" pageId="34" pageNumber="185">C. puella</emphasis>
</taxonomicName>
. If under some condition of the past, unfavourable for
<emphasis id="61D2EAE59133F13CA259FC897FF97B87" box="[1309,1436,1447,1473]" italics="true" pageId="34" pageNumber="185">Coenarion,</emphasis>
both parental species had temporarily vanished from the Black Sea coast while that hybrid population by chance remained the only one in the region, it could later spread as all over the region. The enormous, even intra-population continuous variation of the male black pattern over the whole span between that of both presumed parental species, as well as difference in the paraproct prominence expression between our specimens and those by
<bibRefCitation id="37374B069133F13CA3AFFF197FF27817" author="Bartenef, A. N." box="[1259,1431,1591,1617]" pageId="34" pageNumber="185" pagination="54 - 68" refId="ref27207" refString="Bartenef, A. N. (1929) Neue Arten und Varietaten der Odonata des West Kaukasus. Zoologischer Anzeiger, 85, 54 - 68." type="journal article" year="1929">Bartenef (1929</bibRefCitation>
;
<bibRefCitation id="37374B069133F13CA7D3FF757AB57833" author="Bartenef, A. N." box="[151,208,1627,1653]" pageId="34" pageNumber="185" pagination="1 - 138" refId="ref27235" refString="Bartenef, A. N. (1930) Materialy k poznaniyu Zapadnogo Kavkaza v odonatologicheskom otnoshenii [Materials for the knowledge on West Caucasus in odonatological respect]. Trudy Zapadnokavkazskoi assotsiatsii nauchno-issledovatel'skikh institutov [Proceedings of the West Caucasian Association of scientific Research Institutes], 72 (14), 1 - 138. [in Russian, with German summary but without German title]" type="journal article" year="1930">1930</bibRefCitation>
) and
<bibRefCitation id="37374B069133F13CA64AFF7578157833" author="Skvortsov, V. E. &amp; Snegovaya, N. Y." box="[270,624,1626,1653]" pageId="34" pageNumber="185" pagination="1 - 38" refId="ref29550" refString="Skvortsov, V. E. &amp; Snegovaya, N. Y. (2015) A second addition to the Odonata fauna of Azerbaijan. International Dragonfly Fund Report, 87, 1 - 38." type="journal article" year="2015">Skvortsov &amp; Snegovaya (2015)</bibRefCitation>
may suggest that the
<taxonomicName id="94A64D749133F13CA426FF7579CD7833" authorityName="&quot; (Skvortsov &amp; Snegovaya" authorityYear="2015" box="[866,936,1627,1653]" class="Insecta" family="Coenagrionidae" genus="Agrion" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="34" pageNumber="185" phylum="Arthropoda" rank="species" species="puella">
<emphasis id="61D2EAE59133F13CA426FF7579CD7833" box="[866,936,1627,1653]" italics="true" pageId="34" pageNumber="185">puella</emphasis>
</taxonomicName>
- and
<taxonomicName id="94A64D749133F13CA4A3FF757E067833" authorityName="Vander Linden" authorityYear="1825" box="[999,1123,1627,1653]" class="Insecta" family="Coenagrionidae" genus="Agrion" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="34" pageNumber="185" phylum="Arthropoda" rank="species" species="pulchellum">
<emphasis id="61D2EAE59133F13CA4A3FF757E067833" box="[999,1123,1627,1653]" italics="true" pageId="34" pageNumber="185">pulchellum</emphasis>
</taxonomicName>
-alleles are still co-exist and maybe not yet well mixed in this hybridogenic species.
</paragraph>
<paragraph id="531936F79133F13DA783FF8D7BC07F5B" blockId="34.[151,1437,151,2013]" lastBlockId="35.[151,1437,151,573]" lastPageId="35" lastPageNumber="186" pageId="34" pageNumber="185">
In view of this supposition it is interesting to note that Bliek (1963) reported and illustrated a presumed male hybrid between
<taxonomicName id="94A64D749133F13CA60FFFE97BD778A7" authorityName="&quot; (Skvortsov &amp; Snegovaya" authorityYear="2015" box="[331,434,1735,1761]" class="Insecta" family="Coenagrionidae" genus="Agrion" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="34" pageNumber="185" phylum="Arthropoda" rank="species" species="puella">
<emphasis id="61D2EAE59133F13CA60FFFE97BD778A7" box="[331,434,1735,1761]" italics="true" pageId="34" pageNumber="185">C. puella</emphasis>
</taxonomicName>
and
<taxonomicName id="94A64D749133F13CA6ACFFE978E178A7" authorityName="Vander Linden" authorityYear="1825" box="[488,644,1735,1761]" class="Insecta" family="Coenagrionidae" genus="Agrion" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="34" pageNumber="185" phylum="Arthropoda" rank="species" species="pulchellum">
<emphasis id="61D2EAE59133F13CA6ACFFE978E178A7" box="[488,644,1735,1761]" italics="true" pageId="34" pageNumber="185">C. pulchellum</emphasis>
</taxonomicName>
found near Tutzing (
<collectingRegion id="9162F8159133F13CA42BFFE979A178A7" box="[879,964,1735,1761]" country="Germany" name="Bayern" pageId="34" pageNumber="185">Bayern</collectingRegion>
,
<collectingCountry id="2BB176679133F13CA48BFFE97E5878A7" box="[975,1085,1735,1761]" name="Germany" pageId="34" pageNumber="185">Germany</collectingCountry>
). The illustrated S2 mark (Abb. 2c in loc. cit.) resembles that of
<taxonomicName id="94A64D749133F13CA552FFC578E57943" authorityName="&quot; (Skvortsov &amp; Snegovaya" authorityYear="2015" box="[534,640,1771,1797]" class="Insecta" family="Coenagrionidae" genus="Agrion" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="34" pageNumber="185" phylum="Arthropoda" rank="species" species="puella">
<emphasis id="61D2EAE59133F13CA552FFC578E57943" box="[534,640,1771,1797]" italics="true" pageId="34" pageNumber="185">C. puella</emphasis>
</taxonomicName>
but is thicker, that is inclining towards the
<taxonomicName id="94A64D749133F13CA3C5FFC57F447943" authorityName="Vander Linden" authorityYear="1825" box="[1153,1313,1771,1797]" class="Insecta" family="Coenagrionidae" genus="Agrion" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="34" pageNumber="185" phylum="Arthropoda" rank="species" species="pulchellum">
<emphasis id="61D2EAE59133F13CA3C5FFC57F447943" box="[1153,1313,1771,1797]" italics="true" pageId="34" pageNumber="185">C. pulchellum</emphasis>
</taxonomicName>
condition. The pronotum hind lobe shape (Abb. 1c) is intermediate between that in the two parental species. Unfortunately, the caudal appendages of the hybrid were illustrated only in the posterior view (Abb. 3c), and the tips of the cercal processes are shown hidden behind the paraproct bases, that is very strange. Anyway, the cerci were shown robust, contacting with their bulges and with scarcely curved processes, that is resembling those of
<taxonomicName id="94A64D749133F13CA3BCFE557FF079D3" authorityName="Vander Linden" authorityYear="1825" box="[1272,1429,1915,1941]" class="Insecta" family="Coenagrionidae" genus="Agrion" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="34" pageNumber="185" phylum="Arthropoda" rank="species" species="pulchellum">
<emphasis id="61D2EAE59133F13CA3BCFE557FF079D3" box="[1272,1429,1915,1941]" italics="true" pageId="34" pageNumber="185">C. pulchellum</emphasis>
</taxonomicName>
. The paraprocts were shown as in
<taxonomicName id="94A64D749133F13CA565FEB178A479FF" authorityName="Vander Linden" authorityYear="1825" box="[545,705,1951,1977]" class="Insecta" family="Coenagrionidae" genus="Agrion" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="34" pageNumber="185" phylum="Arthropoda" rank="species" species="pulchellum">
<emphasis id="61D2EAE59133F13CA565FEB178A479FF" box="[545,705,1951,1977]" italics="true" pageId="34" pageNumber="185">C. pulchellum</emphasis>
</taxonomicName>
but less divergent. In sum, the pronotum lobe of the hybrid is intermediate between the parental species, the S2 markings close to that of
<taxonomicName id="94A64D749133F13CA49AFEED7E21799B" authorityName="&quot; (Skvortsov &amp; Snegovaya" authorityYear="2015" box="[990,1092,1987,2013]" class="Insecta" family="Coenagrionidae" genus="Agrion" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="34" pageNumber="185" phylum="Arthropoda" rank="species" species="puella">
<emphasis id="61D2EAE59133F13CA49AFEED7E21799B" box="[990,1092,1987,2013]" italics="true" pageId="34" pageNumber="185">C. puella</emphasis>
</taxonomicName>
and the appendages resembles those of
<taxonomicName id="94A64D749132F13DA7BCF9B97BF17EF7" authorityName="Vander Linden" authorityYear="1825" box="[248,404,151,177]" class="Insecta" family="Coenagrionidae" genus="Agrion" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="35" pageNumber="186" phylum="Arthropoda" rank="species" species="pulchellum">
<emphasis id="61D2EAE59132F13DA7BCF9B97BF17EF7" box="[248,404,151,177]" italics="true" pageId="35" pageNumber="186">C. pulchellum</emphasis>
</taxonomicName>
. These characteristics are remarkably shared by
<taxonomicName id="94A64D749132F13DA4F1F9B97E5B7EF6" authorityName="Kosterin &amp; Onishko &amp; Ilyina &amp; Chepurnov &amp; Blinov" authorityYear="2024" box="[949,1086,151,177]" class="Insecta" family="Coenagrionidae" genus="Agrion" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="35" pageNumber="186" phylum="Arthropoda" rank="species" species="ponticum">
<emphasis id="61D2EAE59132F13DA4F1F9B97E5B7EF6" box="[949,1086,151,177]" italics="true" pageId="35" pageNumber="186">C. ponticum</emphasis>
</taxonomicName>
. The S2 and appendages of the hybrid as depicted by Bliek (1963: Abb. 2c, 3c) could be identifiable as
<taxonomicName id="94A64D749132F13DA4F8F9957E217E92" authorityName="Kosterin &amp; Onishko &amp; Ilyina &amp; Chepurnov &amp; Blinov" authorityYear="2024" box="[956,1092,187,213]" class="Insecta" family="Coenagrionidae" genus="Agrion" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="35" pageNumber="186" phylum="Arthropoda" rank="species" species="ponticum">
<emphasis id="61D2EAE59132F13DA4F8F9957E217E92" box="[956,1092,187,213]" italics="true" pageId="35" pageNumber="186">C. ponticum</emphasis>
</taxonomicName>
, but the pronotum hind lobe is too sinuous, strictly intermediate between the
<taxonomicName id="94A64D749132F13DA5D1F9F1789E7EBF" authorityName="&quot; (Skvortsov &amp; Snegovaya" authorityYear="2015" box="[661,763,223,249]" class="Insecta" family="Coenagrionidae" genus="Agrion" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="35" pageNumber="186" phylum="Arthropoda" rank="species" species="puella">
<emphasis id="61D2EAE59132F13DA5D1F9F1789E7EBF" box="[661,763,223,249]" italics="true" pageId="35" pageNumber="186">C. puella</emphasis>
</taxonomicName>
and
<taxonomicName id="94A64D749132F13DA46BF9F179AE7EBF" authorityName="Vander Linden" authorityYear="1825" box="[815,971,223,249]" class="Insecta" family="Coenagrionidae" genus="Agrion" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="35" pageNumber="186" phylum="Arthropoda" rank="species" species="pulchellum">
<emphasis id="61D2EAE59132F13DA46BF9F179AE7EBF" box="[815,971,223,249]" italics="true" pageId="35" pageNumber="186">C. pulchellum</emphasis>
</taxonomicName>
, while in males of
<taxonomicName id="94A64D749132F13DA3D9F9F17F407EBE" authorityName="Kosterin &amp; Onishko &amp; Ilyina &amp; Chepurnov &amp; Blinov" authorityYear="2024" box="[1181,1317,223,249]" class="Insecta" family="Coenagrionidae" genus="Agrion" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="35" pageNumber="186" phylum="Arthropoda" rank="species" species="ponticum">
<emphasis id="61D2EAE59132F13DA3D9F9F17F407EBE" box="[1181,1317,223,249]" italics="true" pageId="35" pageNumber="186">C. ponticum</emphasis>
</taxonomicName>
it is closer to
<taxonomicName id="94A64D749132F13DA7F0F82D7B7E7F5B" authorityName="&quot; (Skvortsov &amp; Snegovaya" authorityYear="2015" box="[180,283,259,285]" class="Insecta" family="Coenagrionidae" genus="Agrion" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="35" pageNumber="186" phylum="Arthropoda" rank="species" species="puella">
<emphasis id="61D2EAE59132F13DA7F0F82D7B7E7F5B" box="[180,283,259,285]" italics="true" pageId="35" pageNumber="186">C. puella</emphasis>
</taxonomicName>
(
<figureCitation id="CB9D2A729132F13DA66EF82D7BF27F5B" box="[298,407,258,285]" captionStart="FIGURE 5" captionStartId="13.[151,250,1570,1594]" captionTargetBox="[151,1436,181,1541]" captionTargetId="figure-19@13.[151,1436,181,1546]" captionTargetPageId="13" captionText="FIGURE 5. Male pronotum of Coenagrion spp. of the puella-group in the broad sense in dorsal view: a—C. persicum; bc—C. pulchellum; de—C. ponticum (males with a more (d) and less (e) expanded black abdominal maculation); f—C. australocaspicum; gl—C. puella; a—Iran, Markazi Ostan, Shazand Shakhrestan, Emarat village env., a spring and swampy reservoir, 20.05.2017; b—Russia, Dagestan, Makhachkala N env., Muraveynik Protected Area at the Sulak River mouth, pools among coastal dunes, 1.06.2021; c—Russia, Dagestan, Makhachkala N suburbs, pools between dunes at the Krivaya Balka River left bank in its lowermost reaches, 23.05.2023; de—Russia, Krasnodarskiy Kray, Goryachiy Klyuch Municipality, Fanagoriyskoe village NW environs, ponds, 27.04.2019; f—Russia, Dagestan, Magaramkent District, Samur Delta Forest, 2 km NW of Primorskiy village, a swampy estuary of a forest rivulet, 24.06.2021; g —Georgia, Dzau District, 4 km NNE Kvaisa, Lake Koz (Kvedi), 2830.06.2021; hi—Rutul District, the Samur River left arm between Kufa and Kala villages, 26.06.2021; j—Russia, Dagestan, Magaramkent District, Samur Delta Forest, Bil-Bil Kazmalyar village, 25.06.2021; k—Russia, Dagestan, Buynaksk District, Buynaksk S environs, Nizhnee Kazanishche village environs, a water reservoir, 2020; l—Russia, Karachay- Cherkes Republic, Zelenchuk District, 10 km WSW of Arkhyz, Lunnaya Polyana terrain, pools in the Dukka River valley left side, 2.07.2019. Scale bar 0.5 mm." figureDoi="http://doi.org/10.5281/zenodo.12189916" httpUri="https://zenodo.org/record/12189916/files/figure.png" pageId="35" pageNumber="186">Fig. 5d, e</figureCitation>
).
</paragraph>
<paragraph id="531936F79132F13DA783F80978E67FEB" blockId="35.[151,1437,151,573]" pageId="35" pageNumber="186">
On the other hand, so far there was just a single report of such a hybrid while
<bibRefCitation id="37374B069132F13DA378F8097F6A7F07" author="Lowe, C. D. &amp; Harvey, I. F. &amp; Thompson, D. J. &amp; Watts, P. C." box="[1084,1295,295,321]" pageId="35" pageNumber="186" pagination="55 - 63" refId="ref28710" refString="Lowe, C. D., Harvey, I. F., Thompson, D. J. &amp; Watts, P. C. (2008) Strong genetic divergence indicates that congeneric damselflies Coenagrion puella and C. pulchellum (Odonata: Zygoptera: Coenagrionidae) do not hybridise. Hydrobiologia, 605, 55 - 63. https: // doi. org / 10.1007 / s 10750 - 008 - 9300 - 9" type="journal article" year="2008">
Lowe
<emphasis id="61D2EAE59132F13DA3C7F8097ED87F07" box="[1155,1213,295,321]" italics="true" pageId="35" pageNumber="186">et al.</emphasis>
(2008)
</bibRefCitation>
analysed 12 microsatellite loci in
<taxonomicName id="94A64D749132F13DA6C2F8657B887F23" authorityName="&quot; (Skvortsov &amp; Snegovaya" authorityYear="2015" box="[390,493,331,357]" class="Insecta" family="Coenagrionidae" genus="Agrion" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="35" pageNumber="186" phylum="Arthropoda" rank="species" species="puella">
<emphasis id="61D2EAE59132F13DA6C2F8657B887F23" box="[390,493,331,357]" italics="true" pageId="35" pageNumber="186">C. puella</emphasis>
</taxonomicName>
and
<taxonomicName id="94A64D749132F13DA560F86578A47F23" authorityName="Vander Linden" authorityYear="1825" box="[548,705,331,357]" class="Insecta" family="Coenagrionidae" genus="Agrion" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="35" pageNumber="186" phylum="Arthropoda" rank="species" species="pulchellum">
<emphasis id="61D2EAE59132F13DA560F86578A47F23" box="[548,705,331,357]" italics="true" pageId="35" pageNumber="186">C. pulchellum</emphasis>
</taxonomicName>
from British Isles and revealed no sign of hybridisation between these species. This suggests that hybridisation between the two species is not easy, yet it could take place in the complicated history of the Caucasian fauna.
</paragraph>
<paragraph id="531936F79132F13DA783F89979B07C7B" blockId="35.[151,1437,151,573]" pageId="35" pageNumber="186">
It should be noted that
<collectingCountry id="2BB176679132F13DA68DF89978717F97" box="[457,532,439,465]" name="Russia" pageId="35" pageNumber="186">Russia</collectingCountry>
was missed from the range of
<taxonomicName id="94A64D749132F13DA429F89979937F96" authorityName="Kosterin &amp; Onishko &amp; Ilyina &amp; Chepurnov &amp; Blinov" authorityYear="2024" box="[877,1014,439,465]" class="Insecta" family="Coenagrionidae" genus="Agrion" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="35" pageNumber="186" phylum="Arthropoda" rank="species" species="ponticum">
<emphasis id="61D2EAE59132F13DA429F89979937F96" box="[877,1014,439,465]" italics="true" pageId="35" pageNumber="186">C. ponticum</emphasis>
</taxonomicName>
in the the IUCN Red List assessment of this species (
<bibRefCitation id="37374B069132F13DA603F8F57B857FB3" author="Boudot, J. - P." box="[327,480,475,501]" pageId="35" pageNumber="186" pagination="12324645" refId="ref27457" refString="Boudot, J. - P. (2009) Coenagrion ponticum. The IUCN Red List of Threatened Species 2009: e. T 60288 A 12324645. https: // doi. org / 10.2305 / IUCN. UK. 2009 - 2. RLTS. T 60288 A 12324645. en" type="journal article" year="2009">Boudot 2009</bibRefCitation>
), although it comprises a part of its broad
<typeStatus id="8C1D88559132F13DA484F8F579957FB3" box="[960,1008,475,501]" pageId="35" pageNumber="186">type</typeStatus>
locality, while
<collectingCountry id="2BB176679132F13DA3DAF8F57F657FB3" box="[1182,1280,475,501]" name="Armenia" pageId="35" pageNumber="186">Armenia</collectingCountry>
was included without a reference. The range is adequately shown in
<bibRefCitation id="37374B069132F13DA5B7F8D179A97C5F" author="Boudot, J. - P. &amp; Borisov, S. &amp; De Knijf, G. &amp; Van Grunsven, R. &amp; Schroter, A. &amp; Kalkman, V. J." box="[755,972,511,537]" pageId="35" pageNumber="186" pagination="3 - 248" refId="ref27573" refString="Boudot, J. - P., Borisov, S., De Knijf, G., Van Grunsven, R., Schroter, A. &amp; Kalkman, V. J. (2021) Atlas of the dragonflies and damselflies of West and Central Asia. Brachytron, 22, Supplement, 3 - 248." type="journal article" year="2021">Boudot et al (2021)</bibRefCitation>
, without
<collectingCountry id="2BB176679132F13DA375F8D17EF67C5F" box="[1073,1171,511,537]" name="Armenia" pageId="35" pageNumber="186">Armenia</collectingCountry>
but including the Black Sea coast of
<collectingCountry id="2BB176679132F13DA661FB0D7B107C7B" box="[293,373,547,573]" name="Turkey" pageId="35" pageNumber="186">Turkey</collectingCountry>
and a locality in North
<collectingCountry id="2BB176679132F13DA53BFB0D78C97C7B" box="[639,684,547,573]" name="Iran" pageId="35" pageNumber="186">Iran</collectingCountry>
by
<bibRefCitation id="37374B069132F13DA592FB0D79B47C7B" author="Schneider, T. &amp; Ikemeyer, D. &amp; Ferreira, S. &amp; Muller, O." box="[726,977,547,573]" pageId="35" pageNumber="186" pagination="1 - 40" refId="ref29254" refString="Schneider, T., Ikemeyer, D., Ferreira, S. &amp; Muller, O. (2018) Checklist of the dragonflies (Odonata) of Iran with new records and notes on distribution and taxonomy. Zootaxa, 4394 (1), 1 - 40. https: // doi. org / 10.11646 / zootaxa. 4394.1.1" type="journal article" year="2018">Schneider et al (2018)</bibRefCitation>
.
</paragraph>
</subSubSection>
</treatment>
</document>