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<document ID-DOI="http://dx.doi.org/10.3897/zookeys.1036.66435" ID-GBIF-Dataset="bac822e0-f6b0-44f7-9ad8-17464faa8772" ID-PMC="PMC8128840" ID-Pensoft-Pub="1313-2970-1036-99" ID-Pensoft-UUID="87E9A246E0FD5DBA81318090D48DC9FD" ID-PubMed="34040482" ID-ZooBank="F4A9958818BD4B918DB43368AF1D7984" ModsDocID="1313-2970-1036-99" checkinTime="1620734415262" checkinUser="pensoft" docAuthor="Godunko, Roman J., Martynov, Alexander V. &amp; Staniczek, Arnold H." docDate="2021" docId="2B8AFB005E5553E7B37A3633A025E34E" docLanguage="en" docName="ZooKeys 1036: 99-120" docOrigin="ZooKeys 1036" docPubDate="2021-05-10" docSource="http://dx.doi.org/10.3897/zookeys.1036.66435" docTitle="Burmella clypeata Godunko &amp; Martynov &amp; Staniczek 2021, sp. nov." docType="treatment" docUuid="5F823D68-5C03-4138-8875-D80917D61452" docUuidSource="ZooBank" docVersion="4" id="87E9A246E0FD5DBA81318090D48DC9FD" lastPageNumber="99" masterDocId="87E9A246E0FD5DBA81318090D48DC9FD" masterDocTitle="First fossil record of the mayfly family Vietnamellidae (Insecta, Ephemeroptera) from Burmese Amber confirms its Oriental origin and gives new insights into its evolution" masterLastPageNumber="120" masterPageNumber="99" pageNumber="99" updateTime="1668150237642" updateUser="ExternalLinkService">
<mods:mods xmlns:mods="http://www.loc.gov/mods/v3">
<mods:titleInfo>
<mods:title>First fossil record of the mayfly family Vietnamellidae (Insecta, Ephemeroptera) from Burmese Amber confirms its Oriental origin and gives new insights into its evolution</mods:title>
</mods:titleInfo>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Godunko, Roman J.</mods:namePart>
<mods:nameIdentifier type="ORCID">https://orcid.org/0000-0003-2196-3327</mods:nameIdentifier>
<mods:affiliation>Biology Centre of the Czech Academy of Sciences, Institute of Entomology, Branisovska 31, 37005 Ceske Budejovice, Czech Republic &amp; Department of Invertebrate Zoology and Hydrobiology, University of Lodz, Banacha 12 / 16, 90237 Lodz, Poland &amp; State Museum of Natural History, National Academy of Sciences of Ukraine, Teatralna 18, 79008, Lviv, Ukraine</mods:affiliation>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Martynov, Alexander V.</mods:namePart>
<mods:nameIdentifier type="ORCID">https://orcid.org/0000-0002-6506-5134</mods:nameIdentifier>
<mods:affiliation>National Museum of Natural History, National Academy of Sciences of Ukraine, Bohdan Khmelnytsky 15, 01030, Kyiv, Ukraine</mods:affiliation>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Staniczek, Arnold H.</mods:namePart>
<mods:nameIdentifier type="ORCID">https://orcid.org/0000-0001-5577-137X</mods:nameIdentifier>
<mods:affiliation>Department of Entomology, State Museum of Natural History Stuttgart, Rosenstein 1, 70191, Stuttgart, Germany</mods:affiliation>
<mods:nameIdentifier type="email">arnold.staniczek@smns-bw.de</mods:nameIdentifier>
</mods:name>
<mods:typeOfResource>text</mods:typeOfResource>
<mods:relatedItem type="host">
<mods:titleInfo>
<mods:title>ZooKeys</mods:title>
</mods:titleInfo>
<mods:part>
<mods:date>2021</mods:date>
<mods:detail type="pubDate">
<mods:number>2021-05-10</mods:number>
</mods:detail>
<mods:detail type="volume">
<mods:number>1036</mods:number>
</mods:detail>
<mods:extent unit="page">
<mods:start>99</mods:start>
<mods:end>120</mods:end>
</mods:extent>
</mods:part>
</mods:relatedItem>
<mods:location>
<mods:url>http://dx.doi.org/10.3897/zookeys.1036.66435</mods:url>
</mods:location>
<mods:classification>journal article</mods:classification>
<mods:identifier type="DOI">http://dx.doi.org/10.3897/zookeys.1036.66435</mods:identifier>
<mods:identifier type="Pensoft-Pub">1313-2970-1036-99</mods:identifier>
<mods:identifier type="ZooBank">F4A9958818BD4B918DB43368AF1D7984</mods:identifier>
<mods:identifier type="Pensoft-UUID">87E9A246E0FD5DBA81318090D48DC9FD</mods:identifier>
</mods:mods>
<treatment ID-GBIF-Taxon="181974062" LSID="urn:lsid:zoobank.org:act:5F823D68-5C03-4138-8875-D80917D61452" httpUri="http://treatment.plazi.org/id/2B8AFB005E5553E7B37A3633A025E34E" lastPageNumber="99" pageId="0" pageNumber="99">
<subSubSection pageId="0" pageNumber="99" type="nomenclature">
<paragraph pageId="0" pageNumber="99">
<taxonomicName LSID="http://zoobank.org/5F823D68-5C03-4138-8875-D80917D61452" authority="Godunko &amp; Martynov &amp; Staniczek, 2021" authorityName="Godunko &amp; Martynov &amp; Staniczek" authorityYear="2021" class="Insecta" family="Vietnamellidae" genus="Burmella" higherTaxonomySource="GBIF" kingdom="Animalia" lsidName="Burmella clypeata" order="Ephemeroptera" pageId="0" pageNumber="99" phylum="Arthropoda" rank="species" species="clypeata" status="sp. nov.">Burmella clypeata</taxonomicName>
<taxonomicNameLabel pageId="0" pageNumber="99">sp. nov.</taxonomicNameLabel>
<figureCitation captionStart="Figure 7" captionStartId="F7" captionText="Figure 7. Burmella clypeata sp. nov., female imago, holotype A general dorsal view B general ventral view C, D head, dorsal view E head and thorax, dorsal view. BA - basal sclerite (basalare), LPs and red line - lateroparapsidal suture, MNs and green line - mesonotal suture, MPs and yellow line - medioparapsidal suture, rc - right cercus. Scale bars: 2 mm (A, B); 0.2 mm (C, D); 0.5 mm (E)." figureDoi="10.3897/zookeys.1036.66435.figure7" httpUri="https://binary.pensoft.net/fig/541127" pageId="0" pageNumber="99">Figures 7</figureCitation>
<figureCitation captionStart="Figure 8" captionStartId="F8" captionText="Figure 8. Burmella clypeata sp. nov., female imago, holotype A tarsus of left hind leg B tarsus of right hind leg C left forewing in amber D left forewing venation and size ratio of left fore- and right hind wings E distal part of left forewing F preserved basal part of right forewing. I-V - tarsal segments, ci - basally connected intercalary vein, fi - basally free intercalary vein. Scale bars: 0.1 mm (A, B); 1 mm (C, D); 0.5 mm (E, F)." figureDoi="10.3897/zookeys.1036.66435.figure8" httpUri="https://binary.pensoft.net/fig/541128" pageId="0" pageNumber="99">, 8</figureCitation>
<figureCitation captionStart="Figure 9" captionStartId="F9" captionText="Figure 9. Burmella clypeata sp. nov., female imago, holotype A right hind wing, dorsal view B right hind wing, ventral view C right hind wing venation. Scale bars: 0.2 mm (A, B); 0.5 mm (C)." figureDoi="10.3897/zookeys.1036.66435.figure9" httpUri="https://binary.pensoft.net/fig/541129" pageId="0" pageNumber="99">, 9</figureCitation>
<figureCitation captionStart="Figure 10" captionStartId="F10" captionText="Figure 10. Burmella clypeata sp. nov., female imago, holotype A body, ventral view B, C apical part of abdomen, ventral view D basal segments of abdomen, dorsal view. I-III and V-IX - numbers of abdominal segments, gsr - remnant of gill socket, pp - paraproct plate. Scale bars: 1 mm (A); 0.5 mm (B-D)." figureDoi="10.3897/zookeys.1036.66435.figure10" httpUri="https://binary.pensoft.net/fig/541130" pageId="0" pageNumber="99">, 10</figureCitation>
,
<figureCitation captionStart="Table 1" captionStartId="T1" captionText="Table 1. Measurements of fossil representatives of the genus Burmella gen. nov." pageId="0" pageNumber="99">Table 1</figureCitation>
</paragraph>
</subSubSection>
<subSubSection pageId="0" pageNumber="99" type="materials_examined">
<paragraph pageId="0" pageNumber="99">Material examined.</paragraph>
<paragraph pageId="0" pageNumber="99">
<emphasis bold="true" pageId="0" pageNumber="99">
<emphasis italics="true" pageId="0" pageNumber="99">
<typeStatus>Holotype</typeStatus>
</emphasis>
.
</emphasis>
Female imago in Mid-Cretaceous Burmese amber, SMNS collection, inventory number: BU-321. Well preserved specimen visible in dorsal/ventral aspect. Body and left forewings preserved except of lost distal part of C and Sc; left forewing twisted, covering dorsal side of abdomen; right forewings twisted, only partly preserved, distal part missing; foretibiae damaged; right antenna, foretarsi, right middle leg and left cercus missing (Figs
<figureCitation captionStart="Figure 7" captionStartId="F7" captionText="Figure 7. Burmella clypeata sp. nov., female imago, holotype A general dorsal view B general ventral view C, D head, dorsal view E head and thorax, dorsal view. BA - basal sclerite (basalare), LPs and red line - lateroparapsidal suture, MNs and green line - mesonotal suture, MPs and yellow line - medioparapsidal suture, rc - right cercus. Scale bars: 2 mm (A, B); 0.2 mm (C, D); 0.5 mm (E)." figureDoi="10.3897/zookeys.1036.66435.figure7" httpUri="https://binary.pensoft.net/fig/541127" pageId="0" pageNumber="99">7</figureCitation>
,
<figureCitation captionStart="Figure 8" captionStartId="F8" captionText="Figure 8. Burmella clypeata sp. nov., female imago, holotype A tarsus of left hind leg B tarsus of right hind leg C left forewing in amber D left forewing venation and size ratio of left fore- and right hind wings E distal part of left forewing F preserved basal part of right forewing. I-V - tarsal segments, ci - basally connected intercalary vein, fi - basally free intercalary vein. Scale bars: 0.1 mm (A, B); 1 mm (C, D); 0.5 mm (E, F)." figureDoi="10.3897/zookeys.1036.66435.figure8" httpUri="https://binary.pensoft.net/fig/541128" pageId="0" pageNumber="99">8</figureCitation>
,
<figureCitation captionStart="Figure 10" captionStartId="F10" captionText="Figure 10. Burmella clypeata sp. nov., female imago, holotype A body, ventral view B, C apical part of abdomen, ventral view D basal segments of abdomen, dorsal view. I-III and V-IX - numbers of abdominal segments, gsr - remnant of gill socket, pp - paraproct plate. Scale bars: 1 mm (A); 0.5 mm (B-D)." figureDoi="10.3897/zookeys.1036.66435.figure10" httpUri="https://binary.pensoft.net/fig/541130" pageId="0" pageNumber="99">10</figureCitation>
). Left hind wing not visible. For measurements see Table
<tableCitation captionStart="Table 1" captionStartId="T1" captionText="Table 1. Measurements of fossil representatives of the genus Burmella gen. nov." httpUri="http://table.plazi.org/id/73C5555D08B56DEEA903CE63EDD2896A" pageId="0" pageNumber="99" tableUuid="73C5555D08B56DEEA903CE63EDD2896A">1</tableCitation>
.
</paragraph>
<caption doi="10.3897/zookeys.1036.66435.figure7" httpUri="https://binary.pensoft.net/fig/541127" pageId="0" pageNumber="99" start="Figure 7" startId="F7">
<paragraph pageId="0" pageNumber="99">
<emphasis bold="true" pageId="0" pageNumber="99">Figure 7.</emphasis>
<taxonomicName authorityName="Godunko &amp; Martynov &amp; Staniczek" authorityYear="2021" class="Insecta" family="Vietnamellidae" genus="Burmella" higherTaxonomySource="GBIF" kingdom="Animalia" lsidName="Burmella clypeata" order="Ephemeroptera" pageId="0" pageNumber="99" phylum="Arthropoda" rank="species" species="clypeata">
<emphasis italics="true" pageId="0" pageNumber="99">Burmella clypeata</emphasis>
</taxonomicName>
sp. nov., female imago,
<typeStatus>holotype</typeStatus>
<emphasis bold="true" pageId="0" pageNumber="99">A</emphasis>
general dorsal view
<emphasis bold="true" pageId="0" pageNumber="99">B</emphasis>
general ventral view
<emphasis bold="true" pageId="0" pageNumber="99">C, D</emphasis>
head, dorsal view
<emphasis bold="true" pageId="0" pageNumber="99">E</emphasis>
head and thorax, dorsal view. BA - basal sclerite (basalare), LPs and red line - lateroparapsidal suture, MNs and green line - mesonotal suture, MPs and yellow line - medioparapsidal suture, rc - right cercus. Scale bars:
<quantity metricMagnitude="-3" metricUnit="m" metricValue="2.0" unit="mm" value="2.0">2 mm</quantity>
(
<emphasis bold="true" pageId="0" pageNumber="99">A, B</emphasis>
);
<quantity metricMagnitude="-4" metricUnit="m" metricValue="2.0" unit="mm" value="0.2">0.2 mm</quantity>
(
<emphasis bold="true" pageId="0" pageNumber="99">C, D</emphasis>
);
<quantity metricMagnitude="-4" metricUnit="m" metricValue="5.0" unit="mm" value="0.5">0.5 mm</quantity>
(
<emphasis bold="true" pageId="0" pageNumber="99">E</emphasis>
).
</paragraph>
</caption>
</subSubSection>
<subSubSection pageId="0" pageNumber="99" type="derivation of name">
<paragraph pageId="0" pageNumber="99">Derivation of name.</paragraph>
<paragraph pageId="0" pageNumber="99">The species epithet refers to the laterally expanded clypeus that partly covers the eyes.</paragraph>
<caption doi="10.3897/zookeys.1036.66435.figure8" httpUri="https://binary.pensoft.net/fig/541128" pageId="0" pageNumber="99" start="Figure 8" startId="F8">
<paragraph pageId="0" pageNumber="99">
<emphasis bold="true" pageId="0" pageNumber="99">Figure 8.</emphasis>
<taxonomicName authorityName="Godunko &amp; Martynov &amp; Staniczek" authorityYear="2021" class="Insecta" family="Vietnamellidae" genus="Burmella" higherTaxonomySource="GBIF" kingdom="Animalia" lsidName="Burmella clypeata" order="Ephemeroptera" pageId="0" pageNumber="99" phylum="Arthropoda" rank="species" species="clypeata">
<emphasis italics="true" pageId="0" pageNumber="99">Burmella clypeata</emphasis>
</taxonomicName>
sp. nov., female imago, holotype
<emphasis bold="true" pageId="0" pageNumber="99">A</emphasis>
tarsus of left hind leg
<emphasis bold="true" pageId="0" pageNumber="99">B</emphasis>
tarsus of right hind leg
<emphasis bold="true" pageId="0" pageNumber="99">C</emphasis>
left forewing in amber
<emphasis bold="true" pageId="0" pageNumber="99">D</emphasis>
left forewing venation and size ratio of left fore- and right hind wings
<emphasis bold="true" pageId="0" pageNumber="99">E</emphasis>
distal part of left forewing
<emphasis bold="true" pageId="0" pageNumber="99">F</emphasis>
preserved basal part of right forewing. I-V - tarsal segments, ci - basally connected intercalary vein, fi - basally free intercalary vein. Scale bars: 0.1 mm (
<emphasis bold="true" pageId="0" pageNumber="99">A, B</emphasis>
); 1 mm (
<emphasis bold="true" pageId="0" pageNumber="99">C, D</emphasis>
); 0.5 mm (
<emphasis bold="true" pageId="0" pageNumber="99">E, F</emphasis>
).
</paragraph>
</caption>
</subSubSection>
<subSubSection pageId="0" pageNumber="99" type="diagnosis">
<paragraph pageId="0" pageNumber="99">Diagnosis.</paragraph>
<paragraph pageId="0" pageNumber="99">
<emphasis bold="true" pageId="0" pageNumber="99">
<emphasis italics="true" pageId="0" pageNumber="99">Female imago</emphasis>
:
</emphasis>
body length 7.00 mm;
<emphasis italics="true" pageId="0" pageNumber="99">forewings</emphasis>
with at least four short marginal intercalaries in MA-MP field basally attached to longitudinal veins, six free marginal intercalaries in RS field;
<emphasis italics="true" pageId="0" pageNumber="99">hind wing</emphasis>
strongly rounded, small, as long as 0.08
<normalizedToken originalValue="×">x</normalizedToken>
of forewing length, two cross veins between C-Sc, two cross veins between Sc-RA; RS forked;
<emphasis italics="true" pageId="0" pageNumber="99">subgenital plate</emphasis>
more than 2.00
<normalizedToken originalValue="×">x</normalizedToken>
as wide as long, convex and widely rounded apically;
<emphasis italics="true" pageId="0" pageNumber="99">subanal plate</emphasis>
triangular, elongated, rounded apically without cleft.
</paragraph>
</subSubSection>
<subSubSection pageId="0" pageNumber="99" type="description">
<paragraph pageId="0" pageNumber="99">Description.</paragraph>
<paragraph pageId="0" pageNumber="99">
General colouration of body relatively pale, light brown to dark brown. Ventral side of body slightly darker than dorsal side. Body covered by blackish maculation (Figs
<figureCitation captionStart="Figure 7" captionStartId="F7" captionText="Figure 7. Burmella clypeata sp. nov., female imago, holotype A general dorsal view B general ventral view C, D head, dorsal view E head and thorax, dorsal view. BA - basal sclerite (basalare), LPs and red line - lateroparapsidal suture, MNs and green line - mesonotal suture, MPs and yellow line - medioparapsidal suture, rc - right cercus. Scale bars: 2 mm (A, B); 0.2 mm (C, D); 0.5 mm (E)." figureDoi="10.3897/zookeys.1036.66435.figure7" httpUri="https://binary.pensoft.net/fig/541127" pageId="0" pageNumber="99">7A, B</figureCitation>
,
<figureCitation captionStart="Figure 10" captionStartId="F10" captionText="Figure 10. Burmella clypeata sp. nov., female imago, holotype A body, ventral view B, C apical part of abdomen, ventral view D basal segments of abdomen, dorsal view. I-III and V-IX - numbers of abdominal segments, gsr - remnant of gill socket, pp - paraproct plate. Scale bars: 1 mm (A); 0.5 mm (B-D)." figureDoi="10.3897/zookeys.1036.66435.figure10" httpUri="https://binary.pensoft.net/fig/541130" pageId="0" pageNumber="99">10</figureCitation>
).
</paragraph>
<paragraph pageId="0" pageNumber="99">
<emphasis bold="true" pageId="0" pageNumber="99">
<emphasis italics="true" pageId="0" pageNumber="99">Head</emphasis>
.
</emphasis>
Clypeus expanded anterolaterally, partly covering anterior part of eyes. Eyes brown, elongated, relatively large, widely separated medially; facets of eyes hexagonal. Distance between eyes 0.73
<normalizedToken originalValue="×">x</normalizedToken>
of head width. Ocelli well preserved, large, without conspicuous colouration. Facial keel small. Antenna brown, approximately as long as head; segmentation hardly distinguishable, therefore not depicted (see Fig.
<figureCitation captionStart="Figure 7" captionStartId="F7" captionText="Figure 7. Burmella clypeata sp. nov., female imago, holotype A general dorsal view B general ventral view C, D head, dorsal view E head and thorax, dorsal view. BA - basal sclerite (basalare), LPs and red line - lateroparapsidal suture, MNs and green line - mesonotal suture, MPs and yellow line - medioparapsidal suture, rc - right cercus. Scale bars: 2 mm (A, B); 0.2 mm (C, D); 0.5 mm (E)." figureDoi="10.3897/zookeys.1036.66435.figure7" httpUri="https://binary.pensoft.net/fig/541127" pageId="0" pageNumber="99">7C-D</figureCitation>
).
</paragraph>
<paragraph pageId="0" pageNumber="99">
<emphasis bold="true" pageId="0" pageNumber="99">
<emphasis italics="true" pageId="0" pageNumber="99">Thorax</emphasis>
.
</emphasis>
General colouration brown to dark brown. Lateral aspect of thorax not visible. Prothorax narrow, brown. Mesonotal suture transverse, expressed; medioparapsidal suture poorly visible, straight; lateroparapsidal suture distinctly curved laterally; no preserved natural colouration of pigmented area of mesonotum. Ventral side of mesothorax poorly visible; basisternum relatively short and wide distally, furcasternal protuberances distinctly separated. Metathorax brown to dark brown, blackish maculation dorsally (Fig.
<figureCitation captionStart="Figure 7" captionStartId="F7" captionText="Figure 7. Burmella clypeata sp. nov., female imago, holotype A general dorsal view B general ventral view C, D head, dorsal view E head and thorax, dorsal view. BA - basal sclerite (basalare), LPs and red line - lateroparapsidal suture, MNs and green line - mesonotal suture, MPs and yellow line - medioparapsidal suture, rc - right cercus. Scale bars: 2 mm (A, B); 0.2 mm (C, D); 0.5 mm (E)." figureDoi="10.3897/zookeys.1036.66435.figure7" httpUri="https://binary.pensoft.net/fig/541127" pageId="0" pageNumber="99">7E</figureCitation>
).
</paragraph>
<paragraph pageId="0" pageNumber="99">
<emphasis bold="true" pageId="0" pageNumber="99">
<emphasis italics="true" pageId="0" pageNumber="99">Wings</emphasis>
.
</emphasis>
<emphasis italics="true" pageId="0" pageNumber="99">Forewings</emphasis>
hyaline, translucent, relatively narrow; venation poorly recognizable due to wing deformation, pollution on surface and resin influxes [left wing], and damage of distal part [right wing]; venation well visible from dorsal, and partly from lateral side. Veins light brown to brown; relatively small number of cross veins; no jagged edge along of ventral margin (Fig.
<figureCitation captionStart="Figure 8" captionStartId="F8" captionText="Figure 8. Burmella clypeata sp. nov., female imago, holotype A tarsus of left hind leg B tarsus of right hind leg C left forewing in amber D left forewing venation and size ratio of left fore- and right hind wings E distal part of left forewing F preserved basal part of right forewing. I-V - tarsal segments, ci - basally connected intercalary vein, fi - basally free intercalary vein. Scale bars: 0.1 mm (A, B); 1 mm (C, D); 0.5 mm (E, F)." figureDoi="10.3897/zookeys.1036.66435.figure8" httpUri="https://binary.pensoft.net/fig/541128" pageId="0" pageNumber="99">8C-F</figureCitation>
).
</paragraph>
<paragraph pageId="0" pageNumber="99">
General pattern of forewing venation similar to those of male imago of
<taxonomicName authorityName="Godunko &amp; Martynov &amp; Staniczek" authorityYear="2021" class="Insecta" family="Vietnamellidae" genus="Burmella" higherTaxonomySource="GBIF" kingdom="Animalia" lsidName="Burmella paucivenosa" order="Ephemeroptera" pageId="0" pageNumber="99" phylum="Arthropoda" rank="species" species="paucivenosa">
<emphasis italics="true" pageId="0" pageNumber="99">Burmella paucivenosa</emphasis>
</taxonomicName>
sp. nov., except for the following features: six free intercalary veins at least in RS field and CuA-CuP; at least four intercalaries in MA-MP field basally attached to longitudinal veins (Fig.
<figureCitation captionStart="Figure 8" captionStartId="F8" captionText="Figure 8. Burmella clypeata sp. nov., female imago, holotype A tarsus of left hind leg B tarsus of right hind leg C left forewing in amber D left forewing venation and size ratio of left fore- and right hind wings E distal part of left forewing F preserved basal part of right forewing. I-V - tarsal segments, ci - basally connected intercalary vein, fi - basally free intercalary vein. Scale bars: 0.1 mm (A, B); 1 mm (C, D); 0.5 mm (E, F)." figureDoi="10.3897/zookeys.1036.66435.figure8" httpUri="https://binary.pensoft.net/fig/541128" pageId="0" pageNumber="99">8C-F</figureCitation>
).
</paragraph>
<paragraph pageId="0" pageNumber="99">
<emphasis bold="true" italics="true" pageId="0" pageNumber="99">Hind wings</emphasis>
hyaline, translucent, small, as long as 0.08 of forewing length; preserved wing is deformed due to embedding, but most probable was naturally strongly rounded, with shallow costal process; venation brown, significantly simplifies; strong reduction of number of longitudinal and cross veins; no jagged edge along of ventral margin. General structure and pattern of hind wing venation similar to those in male imago of
<taxonomicName authorityName="Godunko &amp; Martynov &amp; Staniczek" authorityYear="2021" class="Insecta" family="Vietnamellidae" genus="Burmella" higherTaxonomySource="GBIF" kingdom="Animalia" lsidName="Burmella paucivenosa" order="Ephemeroptera" pageId="0" pageNumber="99" phylum="Arthropoda" rank="species" species="paucivenosa">
<emphasis italics="true" pageId="0" pageNumber="99">Burmella paucivenosa</emphasis>
</taxonomicName>
sp. nov., except for the following features: a few cross veins between C-Sc (2 veins), and Sc-RA (2 veins); fork RS present, iRS short, no cross veins in RS field; costal process not prominent (Fig.
<figureCitation captionStart="Figure 9" captionStartId="F9" captionText="Figure 9. Burmella clypeata sp. nov., female imago, holotype A right hind wing, dorsal view B right hind wing, ventral view C right hind wing venation. Scale bars: 0.2 mm (A, B); 0.5 mm (C)." figureDoi="10.3897/zookeys.1036.66435.figure9" httpUri="https://binary.pensoft.net/fig/541129" pageId="0" pageNumber="99">9A-C</figureCitation>
).
</paragraph>
<caption doi="10.3897/zookeys.1036.66435.figure9" httpUri="https://binary.pensoft.net/fig/541129" pageId="0" pageNumber="99" start="Figure 9" startId="F9">
<paragraph pageId="0" pageNumber="99">
<emphasis bold="true" pageId="0" pageNumber="99">Figure 9.</emphasis>
<taxonomicName authorityName="Godunko &amp; Martynov &amp; Staniczek" authorityYear="2021" class="Insecta" family="Vietnamellidae" genus="Burmella" higherTaxonomySource="GBIF" kingdom="Animalia" lsidName="Burmella clypeata" order="Ephemeroptera" pageId="0" pageNumber="99" phylum="Arthropoda" rank="species" species="clypeata">
<emphasis italics="true" pageId="0" pageNumber="99">Burmella clypeata</emphasis>
</taxonomicName>
sp. nov., female imago, holotype
<emphasis bold="true" pageId="0" pageNumber="99">A</emphasis>
right hind wing, dorsal view
<emphasis bold="true" pageId="0" pageNumber="99">B</emphasis>
right hind wing, ventral view
<emphasis bold="true" pageId="0" pageNumber="99">C</emphasis>
right hind wing venation. Scale bars: 0.2 mm (
<emphasis bold="true" pageId="0" pageNumber="99">A, B</emphasis>
); 0.5 mm (
<emphasis bold="true" pageId="0" pageNumber="99">C</emphasis>
).
</paragraph>
</caption>
<paragraph pageId="0" pageNumber="99">
<emphasis bold="true" italics="true" pageId="0" pageNumber="99">Legs</emphasis>
well preserved, except for both forelegs with partly missing tibiae and tarsi; no visible strong spines or setae on margins of leg segments. Preserved part of forelegs darker than middle and hind legs, brown to intensively brown (Fig.
<figureCitation captionStart="Figure 8" captionStartId="F8" captionText="Figure 8. Burmella clypeata sp. nov., female imago, holotype A tarsus of left hind leg B tarsus of right hind leg C left forewing in amber D left forewing venation and size ratio of left fore- and right hind wings E distal part of left forewing F preserved basal part of right forewing. I-V - tarsal segments, ci - basally connected intercalary vein, fi - basally free intercalary vein. Scale bars: 0.1 mm (A, B); 1 mm (C, D); 0.5 mm (E, F)." figureDoi="10.3897/zookeys.1036.66435.figure8" httpUri="https://binary.pensoft.net/fig/541128" pageId="0" pageNumber="99">8A, B</figureCitation>
). For measurements of leg segments see Table
<tableCitation captionStart="Table 1" captionStartId="T1" captionText="Table 1. Measurements of fossil representatives of the genus Burmella gen. nov." httpUri="http://table.plazi.org/id/73C5555D08B56DEEA903CE63EDD2896A" pageId="0" pageNumber="99" tableUuid="73C5555D08B56DEEA903CE63EDD2896A">1</tableCitation>
.
</paragraph>
<paragraph pageId="0" pageNumber="99">
Forelegs partly preserved [due to damage of foretibiae the ratio of femur/tibia is not calculated]. Left middle leg completely preserved: length ratio of femur/tibia/tarsus = 1/2.23/0.70; length ratio of tarsomeres: 1/0.88/0.88/1.00/1.50 (5&gt; 4&gt; 3 = 2 &lt;1). Right hind leg much shorter than left one, probably re-grown after previous injury, therefore with changed proportions of tarsomeres: length ratio of femur/tibia/tarsus =1/0.79/0.62; length ratio of tarsomeres: 1/0.89/0.78/0.89/1.33 (5&gt; 4&gt; 3 &lt;2 &lt;1). Left hind leg: length ratio of femur/tibia/tarsus = 1/1.02/0.57; length ratio of tarsomeres: 1/1/0.91/0.91/1.27 (5&gt; 4 = 3 &lt;2 = 1) (Figs
<figureCitation captionStart="Figure 8" captionStartId="F8" captionText="Figure 8. Burmella clypeata sp. nov., female imago, holotype A tarsus of left hind leg B tarsus of right hind leg C left forewing in amber D left forewing venation and size ratio of left fore- and right hind wings E distal part of left forewing F preserved basal part of right forewing. I-V - tarsal segments, ci - basally connected intercalary vein, fi - basally free intercalary vein. Scale bars: 0.1 mm (A, B); 1 mm (C, D); 0.5 mm (E, F)." figureDoi="10.3897/zookeys.1036.66435.figure8" httpUri="https://binary.pensoft.net/fig/541128" pageId="0" pageNumber="99">8A, B</figureCitation>
,
<figureCitation captionStart="Figure 10" captionStartId="F10" captionText="Figure 10. Burmella clypeata sp. nov., female imago, holotype A body, ventral view B, C apical part of abdomen, ventral view D basal segments of abdomen, dorsal view. I-III and V-IX - numbers of abdominal segments, gsr - remnant of gill socket, pp - paraproct plate. Scale bars: 1 mm (A); 0.5 mm (B-D)." figureDoi="10.3897/zookeys.1036.66435.figure10" httpUri="https://binary.pensoft.net/fig/541130" pageId="0" pageNumber="99">10A</figureCitation>
). Other leg characters similar to those in male imago of
<taxonomicName authorityName="Godunko &amp; Martynov &amp; Staniczek" authorityYear="2021" class="Insecta" family="Vietnamellidae" genus="Burmella" higherTaxonomySource="GBIF" kingdom="Animalia" lsidName="Burmella paucivenosa" order="Ephemeroptera" pageId="0" pageNumber="99" phylum="Arthropoda" rank="species" species="paucivenosa">
<emphasis italics="true" pageId="0" pageNumber="99">Burmella paucivenosa</emphasis>
</taxonomicName>
sp. nov.
</paragraph>
<paragraph pageId="0" pageNumber="99">
<emphasis bold="true" italics="true" pageId="0" pageNumber="99">Abdominal segments</emphasis>
completely preserved, light brown to brown, with blackish maculation on terga and sterna; ventral side of abdomen paler than dorsal side. Vestigial gill sockets, not finger-like, well recognizable on segments II, V, and IV; on other segments gill sockets not distinguishable due to body position in amber. Abdominal segments without large and prominent posterolateral projections; no conspicuous elongation of distal segments compared to proximal ones. Subgenital plate relatively broad, more than 2.00
<normalizedToken originalValue="×">x</normalizedToken>
as wide as long, convex and widely rounded apically. Subanal plate triangular, elongated, moderately narrow and rounded apically without apical cleft. Right cercus completely preserved, brown, darker proximally, approximately as long as body (Fig.
<figureCitation captionStart="Figure 10" captionStartId="F10" captionText="Figure 10. Burmella clypeata sp. nov., female imago, holotype A body, ventral view B, C apical part of abdomen, ventral view D basal segments of abdomen, dorsal view. I-III and V-IX - numbers of abdominal segments, gsr - remnant of gill socket, pp - paraproct plate. Scale bars: 1 mm (A); 0.5 mm (B-D)." figureDoi="10.3897/zookeys.1036.66435.figure10" httpUri="https://binary.pensoft.net/fig/541130" pageId="0" pageNumber="99">10A-D</figureCitation>
).
</paragraph>
<caption doi="10.3897/zookeys.1036.66435.figure10" httpUri="https://binary.pensoft.net/fig/541130" pageId="0" pageNumber="99" start="Figure 10" startId="F10">
<paragraph pageId="0" pageNumber="99">
<emphasis bold="true" pageId="0" pageNumber="99">Figure 10.</emphasis>
<taxonomicName authorityName="Godunko &amp; Martynov &amp; Staniczek" authorityYear="2021" class="Insecta" family="Vietnamellidae" genus="Burmella" higherTaxonomySource="GBIF" kingdom="Animalia" lsidName="Burmella clypeata" order="Ephemeroptera" pageId="0" pageNumber="99" phylum="Arthropoda" rank="species" species="clypeata">
<emphasis italics="true" pageId="0" pageNumber="99">Burmella clypeata</emphasis>
</taxonomicName>
sp. nov., female imago, holotype
<emphasis bold="true" pageId="0" pageNumber="99">A</emphasis>
body, ventral view
<emphasis bold="true" pageId="0" pageNumber="99">B, C</emphasis>
apical part of abdomen, ventral view
<emphasis bold="true" pageId="0" pageNumber="99">D</emphasis>
basal segments of abdomen, dorsal view. I-III and V-IX - numbers of abdominal segments, gsr - remnant of gill socket, pp - paraproct plate. Scale bars: 1 mm (
<emphasis bold="true" pageId="0" pageNumber="99">A</emphasis>
); 0.5 mm (
<emphasis bold="true" pageId="0" pageNumber="99">B-D</emphasis>
).
</paragraph>
</caption>
</subSubSection>
<subSubSection pageId="0" pageNumber="99" type="affinities">
<paragraph pageId="0" pageNumber="99">Affinities.</paragraph>
<paragraph pageId="0" pageNumber="99">
Attribution of
<taxonomicName authorityName="Godunko &amp; Martynov &amp; Staniczek" authorityYear="2021" class="Insecta" family="Vietnamellidae" genus="Burmella" higherTaxonomySource="GBIF" kingdom="Animalia" lsidName="Burmella clypeata" order="Ephemeroptera" pageId="0" pageNumber="99" phylum="Arthropoda" rank="species" species="clypeata">
<emphasis italics="true" pageId="0" pageNumber="99">Burmella clypeata</emphasis>
</taxonomicName>
sp. nov. to the newly described genus is confirmed based on the shape of hind wings, and specific venation.
</paragraph>
<paragraph pageId="0" pageNumber="99">
On the other hand, some aspects of the venation of fore- and hind wings differ between
<taxonomicName authorityName="Godunko &amp; Martynov &amp; Staniczek" authorityYear="2021" class="Insecta" family="Vietnamellidae" genus="Burmella" higherTaxonomySource="GBIF" kingdom="Animalia" lsidName="Burmella clypeata" order="Ephemeroptera" pageId="0" pageNumber="99" phylum="Arthropoda" rank="species" species="clypeata">
<emphasis italics="true" pageId="0" pageNumber="99">Burmella clypeata</emphasis>
</taxonomicName>
sp. nov. and
<taxonomicName authorityName="Godunko &amp; Martynov &amp; Staniczek" authorityYear="2021" class="Insecta" family="Vietnamellidae" genus="Burmella" higherTaxonomySource="GBIF" kingdom="Animalia" lsidName="Burmella paucivenosa" order="Ephemeroptera" pageId="0" pageNumber="99" phylum="Arthropoda" rank="species" species="paucivenosa">
<emphasis italics="true" pageId="0" pageNumber="99">Burmella paucivenosa</emphasis>
</taxonomicName>
sp. nov. The forewings of
<taxonomicName authorityName="Godunko &amp; Martynov &amp; Staniczek" authorityYear="2021" class="Insecta" family="Vietnamellidae" genus="Burmella" higherTaxonomySource="GBIF" kingdom="Animalia" lsidName="Burmella clypeata" order="Ephemeroptera" pageId="0" pageNumber="99" phylum="Arthropoda" rank="species" species="clypeata">
<emphasis italics="true" pageId="0" pageNumber="99">Burmella clypeata</emphasis>
</taxonomicName>
sp. nov. differ by the presence of numerous free marginal intercalaries between iRS and CuP, as well as the presence of at least one cross vein between A1 and A2. In the hind wings differences between the extinct species described here refer to the number of cross veins between C-Sc and Sc-RA. The presence of RS furcation and blunt costal process in
<taxonomicName authorityName="Godunko &amp; Martynov &amp; Staniczek" authorityYear="2021" class="Insecta" family="Vietnamellidae" genus="Burmella" higherTaxonomySource="GBIF" kingdom="Animalia" lsidName="Burmella clypeata" order="Ephemeroptera" pageId="0" pageNumber="99" phylum="Arthropoda" rank="species" species="clypeata">
<emphasis italics="true" pageId="0" pageNumber="99">Burmella clypeata</emphasis>
</taxonomicName>
sp. nov. are also suitable for the separation of both species. In contrast to all other representatives of
<taxonomicName class="Insecta" family="Vietnamellidae" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" lsidName="" order="Ephemeroptera" pageId="0" pageNumber="99" phylum="Arthropoda" rank="family">Vietnamellidae</taxonomicName>
, the clypeus in the female of
<taxonomicName authorityName="Godunko &amp; Martynov &amp; Staniczek" authorityYear="2021" class="Insecta" family="Vietnamellidae" genus="Burmella" higherTaxonomySource="GBIF" kingdom="Animalia" lsidName="Burmella clypeata" order="Ephemeroptera" pageId="0" pageNumber="99" phylum="Arthropoda" rank="species" species="clypeata">
<emphasis italics="true" pageId="0" pageNumber="99">Burmella clypeata</emphasis>
</taxonomicName>
sp. nov. is anterolaterally expanded, as a result the anterior portion of eyes is partly covered by this clypeal shield (Fig.
<figureCitation captionStart="Figure 7" captionStartId="F7" captionText="Figure 7. Burmella clypeata sp. nov., female imago, holotype A general dorsal view B general ventral view C, D head, dorsal view E head and thorax, dorsal view. BA - basal sclerite (basalare), LPs and red line - lateroparapsidal suture, MNs and green line - mesonotal suture, MPs and yellow line - medioparapsidal suture, rc - right cercus. Scale bars: 2 mm (A, B); 0.2 mm (C, D); 0.5 mm (E)." figureDoi="10.3897/zookeys.1036.66435.figure7" httpUri="https://binary.pensoft.net/fig/541127" pageId="0" pageNumber="99">7C-D</figureCitation>
; compare with e.g.,
<bibRefCitation DOI="https://doi.org/10.3897/zookeys.902.46844" author="Auychinda, Ch" journalOrPublisher="ZooKeys" pageId="0" pageNumber="99" pagination="17 - 36" refId="B1" refString="Auychinda, Ch, Sartori, M, Boonsoong, B, 2020a. Vietnamellidae (Insecta, Ephemeroptera) of Thailand. ZooKeys 902: 17 - 36, DOI: https://doi.org/10.3897/zookeys.902.46844" title="Vietnamellidae (Insecta, Ephemeroptera) of Thailand." url="https://doi.org/10.3897/zookeys.902.46844" volume="902" year="2020 a">Auychinda et al. 2020a</bibRefCitation>
: 9, figs A-E; 2020b: 30, fig. 9A).
</paragraph>
<paragraph pageId="0" pageNumber="99">
We do however not per se exclude a possible conspecifity of both fossil specimens. This may be supported by a similar, small body size of both specimens, with similar proportions of male/female body length as in extant
<taxonomicName class="Insecta" family="Vietnamellidae" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" lsidName="" order="Ephemeroptera" pageId="0" pageNumber="99" phylum="Arthropoda" rank="family">Vietnamellidae</taxonomicName>
(for
<taxonomicName authorityName="Godunko &amp; Martynov &amp; Staniczek" authorityYear="2021" class="Insecta" family="Vietnamellidae" genus="Burmella" higherTaxonomySource="GBIF" kingdom="Animalia" lsidName="Burmella" order="Ephemeroptera" pageId="0" pageNumber="99" phylum="Arthropoda" rank="genus">
<emphasis italics="true" pageId="0" pageNumber="99">Burmella</emphasis>
</taxonomicName>
gen. nov. the ratio is 0.82; for
<taxonomicName class="Insecta" family="Vietnamellidae" genus="Vietnamella" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Vietnamella" order="Ephemeroptera" pageId="0" pageNumber="99" phylum="Arthropoda" rank="genus">
<emphasis italics="true" pageId="0" pageNumber="99">Vietnamella</emphasis>
</taxonomicName>
between 0.92 and 0.96). Also, the anterolaterally expanded clypeus in
<taxonomicName lsidName="B. clypeata" pageId="0" pageNumber="99" rank="species" species="clypeata">
<emphasis italics="true" pageId="0" pageNumber="99">B. clypeata</emphasis>
</taxonomicName>
may not exclude their conspecifity. Similar clypeal expansions present in one sex only have been reported in several extant and fossil species of
<taxonomicName authorityName="Needham" authorityYear="1901" class="Insecta" family="Heptageniidae" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" lsidName="" order="Ephemeroptera" pageId="0" pageNumber="99" phylum="Arthropoda" rank="family">Heptageniidae</taxonomicName>
(e.g. in the subgenus
<taxonomicName authority="" class="Insecta" family="Heptageniidae" genus="Ecdyonurus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Ecdyonurus (Nestormeus)" order="Ephemeroptera" pageId="0" pageNumber="99" phylum="Arthropoda" rank="subGenus" subGenus="Nestormeus">Ecdyonurus (Nestormeus)</taxonomicName>
Godunko, 2004), representing a morphological trait independently occurring in several unrelated taxa within the family (see
<bibRefCitation DOI="https://doi.org/10.5281/zenodo.179949" author="Godunko, RJ" journalOrPublisher="Zootaxa" pageId="0" pageNumber="99" pagination="63 - 68" refId="B4" refString="Godunko, RJ, 2007. Contribution to the knowledge of the fossil subgenus Nestormeus Godunko, 2004 (Ephemeroptera: Heptageniidae: Ecdyonurus) from the Baltic amber (Eocene). Zootaxa 1661: 63 - 68, DOI: https://doi.org/10.5281/zenodo.179949" title="Contribution to the knowledge of the fossil subgenus Nestormeus Godunko, 2004 (Ephemeroptera: Heptageniidae: Ecdyonurus) from the Baltic amber (Eocene)." url="https://doi.org/10.5281/zenodo.179949" volume="1661" year="2007">Godunko 2007</bibRefCitation>
: 66, figs 1, 2;
<bibRefCitation DOI="https://doi.org/10.11646/zootaxa.4500.2.3" author="Hrivniak, L" journalOrPublisher="Zootaxa" pageId="0" pageNumber="99" pagination="195 - 221" refId="B9" refString="Hrivniak, L, Sroka, P, Godunko, RJ, Palatov, D, Polasek, M, Manko, P, Obona, J, 2018. Diversity of Armenian mayflies (Ephemeroptera) with the description of a new species of the genus Ecdyonurus (Heptageniidae). Zootaxa 4500 (2): 195 - 221, DOI: https://doi.org/10.11646/zootaxa.4500.2.3" title="Diversity of Armenian mayflies (Ephemeroptera) with the description of a new species of the genus Ecdyonurus (Heptageniidae)." url="https://doi.org/10.11646/zootaxa.4500.2.3" volume="4500" year="2018">Hrivniak et al. 2018</bibRefCitation>
: 199, 204-205, figs 2-5). However, a clear difference in the venation of fore- and hind wings between
<taxonomicName lsidName="B. paucivenosa" pageId="0" pageNumber="99" rank="species" species="paucivenosa">
<emphasis italics="true" pageId="0" pageNumber="99">B. paucivenosa</emphasis>
</taxonomicName>
sp. nov. and
<taxonomicName lsidName="B. clypeata" pageId="0" pageNumber="99" rank="species" species="clypeata">
<emphasis italics="true" pageId="0" pageNumber="99">B. clypeata</emphasis>
</taxonomicName>
sp. nov. rather points to the presence of two different fossil species.
</paragraph>
<paragraph pageId="0" pageNumber="99">
In any case, unless specimens of different sex are syninclusions and fossilized in mating position, we tend to describe males and females of the same genus as different species also to maintain nomenclatural stability (see e.g.
<bibRefCitation DOI="https://doi.org/10.11646/zootaxa.4103.1.1" author="Staniczek, AH" journalOrPublisher="Zootaxa" pageId="0" pageNumber="99" pagination="1 - 24" refId="B20" refString="Staniczek, AH, Godunko, RJ, 2016. Revision of fossil Metretopodidae (Insecta: Ephemeroptera) in Baltic amber - Part 3: Description of two new species of Siphloplecton Clemens, 1915, with notes on the re-discovered lectotype of Siphloplecton macrops (Pictet-Baraban &amp; Hagen, 1856). Zootaxa 4103 (1): 1 - 24, DOI: https://doi.org/10.11646/zootaxa.4103.1.1" title="Revision of fossil Metretopodidae (Insecta: Ephemeroptera) in Baltic amber - Part 3: Description of two new species of Siphloplecton Clemens, 1915, with notes on the re-discovered lectotype of Siphloplecton macrops (Pictet-Baraban &amp; Hagen, 1856)." url="https://doi.org/10.11646/zootaxa.4103.1.1" volume="4103" year="2016">Staniczek and Godunko 2016</bibRefCitation>
;
<bibRefCitation DOI="https://doi.org/10.3897/zookeys.898.47118" author="Godunko, RJ" journalOrPublisher="Zootaxa" pageId="0" pageNumber="99" refId="B6" refString="Godunko, RJ, Neumann, Ch, Staniczek, AH, 2019. Revision of fossil Metretopodidae (Insecta, Ephemeroptera) in Baltic amber - Part 4: Description of two new species of Siphloplecton Clemens, 1915, with notes on the new S. jaegeri species group and with key to fossil male adults of Siphloplecton. ZooKeys 898: 1-26. https://doi.org/10.3897/zookeys.898.47118" title="Revision of fossil Metretopodidae (Insecta, Ephemeroptera) in Baltic amber - Part 4: Description of two new species of Siphloplecton Clemens, 1915, with notes on the new S. jaegeri species group and with key to fossil male adults of Siphloplecton. ZooKeys 898: 1 - 26." url="https://doi.org/10.3897/zookeys.898.47118" year="2019">Godunko et al. 2019</bibRefCitation>
).
</paragraph>
</subSubSection>
</treatment>
</document>