treatments-xml/data/03/B4/87/03B487C20008FF88FF5BFA67FD3D1194.xml

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<mods:title id="A85B7F7AB52D77ACF8A5D2401457D6F4">The Cephalozygoptera, a new, extinct suborder of Odonata with new taxa from the early Eocene Okanagan Highlands, western North America</mods:title>
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<mods:namePart id="39C3C15B0B76A31EFBD7890017DD1055">Cannings, Robert A.</mods:namePart>
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<mods:namePart id="F6BDC36F96146BDC321449E9BB9D5AB7">Erickson, Robert J.</mods:namePart>
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<mods:namePart id="36582035F86B3E7CBEAF1E00973FDA5C">Bybee, Seth M.</mods:namePart>
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<mods:namePart id="7583FD33BE53F5EA509C94EB415C45C9">Mathewes, Rolf W.</mods:namePart>
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<treatment id="03B487C20008FF88FF5BFA67FD3D1194" ID-DOI="http://doi.org/10.5281/zenodo.4672713" ID-GBIF-Taxon="180881817" ID-Zenodo-Dep="4672713" LSID="urn:lsid:plazi:treatment:03B487C20008FF88FF5BFA67FD3D1194" httpUri="http://treatment.plazi.org/id/03B487C20008FF88FF5BFA67FD3D1194" lastPageId="31" lastPageNumber="32" pageId="26" pageNumber="27">
<subSubSection id="C307655F0008FF8DFF5BFA67FC791788" box="[151,983,1410,1437]" pageId="26" pageNumber="27" type="nomenclature">
<paragraph id="8BA236D40008FF8DFF5BFA67FC791788" blockId="26.[151,983,1410,1437]" box="[151,983,1410,1437]" pageId="26" pageNumber="27">
<emphasis id="B969EAC60008FF8DFF5BFA67FC791788" bold="true" box="[151,983,1410,1437]" pageId="26" pageNumber="27">
<taxonomicName id="4C1D4D570008FF8DFF5BFA67FCB01789" authority="Archibald, Cannings &amp; Erickson" authorityName="Archibald, Cannings &amp; Erickson" box="[151,798,1410,1436]" kingdom="Animalia" pageId="26" pageNumber="27" rank="subOrder" status="subord. nov." subOrder="Cephalozygoptera">Cephalozygoptera Archibald, Cannings &amp; Erickson</taxonomicName>
, 
<taxonomicNameLabel id="A25A57BD0008FF8DFCE0FA66FC791788" box="[812,983,1411,1437]" pageId="26" pageNumber="27" rank="subOrder">new suborder</taxonomicNameLabel>
</emphasis>
</paragraph>
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<subSubSection id="C307655F0008FF8DFF5BFA2EFC9514A8" pageId="26" pageNumber="27" type="discussion">
<paragraph id="8BA236D40008FF8DFF5BFA2EFC9514A8" blockId="26.[151,1437,1483,1905]" pageId="26" pageNumber="27">
To expand the 
<taxonomicName id="4C1D4D570008FF8DFEF1FA2EFE1E17F0" box="[317,432,1483,1509]" class="Insecta" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="26" pageNumber="124" phylum="Arthropoda" rank="subOrder" subOrder="Zygoptera">Zygoptera</taxonomicName>
concept to include the head and eye morphologies of the 
<taxonomicName id="4C1D4D570008FF8DFBF5FA2EFB7217F0" box="[1081,1244,1483,1509]" class="Insecta" family="Dysagrionidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="26" pageNumber="27" phylum="Arthropoda" rank="family">Dysagrionidae</taxonomicName>
and 
<taxonomicName id="4C1D4D570008FF8DFADFFA2EFA3217F0" box="[1299,1436,1483,1509]" class="Insecta" family="Sieblosiidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="26" pageNumber="27" phylum="Arthropoda" rank="family">Sieblosiidae</taxonomicName>
(see below) would result in the loss of these few primary, unambiguous, and easily observable diagnostic character states defining the suborder. We, therefore, group these families and possibly the 
<taxonomicName id="4C1D4D570008FF8DFBFFF9F6FB531438" authority="Archibald &amp; Cannings &amp; Erickson &amp; Bybee &amp; Mathewes, 2021" authorityName="Archibald &amp; Cannings &amp; Erickson &amp; Bybee &amp; Mathewes" authorityYear="2021" box="[1075,1277,1555,1581]" family="Whetwhetaksidae" kingdom="Animalia" order="Odonata" pageId="26" pageNumber="28" rank="family" status="fam. nov.">Whetwhetaksidae</taxonomicName>
, whose wings indicate a close relationship with them, as the new suborder 
<taxonomicName id="4C1D4D570008FF8DFC9EF9D2FB8F1444" box="[850,1057,1591,1617]" kingdom="Animalia" pageId="26" pageNumber="27" rank="subOrder" subOrder="Cephalozygoptera">Cephalozygoptera</taxonomicName>
. If a future fossil (presumably in amber, as this is highly unlikely to be seen in any compression fossil) shows that the ligula differs from that of 
<taxonomicName id="4C1D4D570008FF8DFAB8F9BEFF40148C" class="Insecta" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="26" pageNumber="124" phylum="Arthropoda" rank="subOrder" subOrder="Zygoptera">Zygoptera</taxonomicName>
, this may be considered a synapomorphy of the 
<taxonomicName id="4C1D4D570008FF8DFCC1F99AFC72148C" box="[781,988,1663,1689]" kingdom="Animalia" pageId="26" pageNumber="27" rank="subOrder" subOrder="Cephalozygoptera">Cephalozygoptera</taxonomicName>
; if it is consistent with the 
<taxonomicName id="4C1D4D570008FF8DFAC7F99AFA2F148C" box="[1291,1409,1663,1689]" class="Insecta" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="26" pageNumber="124" phylum="Arthropoda" rank="subOrder" subOrder="Zygoptera">Zygoptera</taxonomicName>
, it would be a synapomorphy of 
<taxonomicName id="4C1D4D570008FF8DFE29F946FDF414A8" box="[485,602,1699,1725]" class="Insecta" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="26" pageNumber="124" phylum="Arthropoda" rank="subOrder" subOrder="Zygoptera">Zygoptera</taxonomicName>
+ 
<taxonomicName id="4C1D4D570008FF8DFDA5F946FC9614A8" box="[617,824,1699,1725]" kingdom="Animalia" pageId="26" pageNumber="27" rank="subOrder" subOrder="Cephalozygoptera">Cephalozygoptera</taxonomicName>
.
</paragraph>
</subSubSection>
<subSubSection id="C307655F0008FF8CFF0BF923FA3D1370" lastPageId="27" lastPageNumber="28" pageId="26" pageNumber="27" type="diagnosis">
<paragraph id="8BA236D40008FF8DFF0BF923FC611564" blockId="26.[151,1437,1483,1905]" pageId="26" pageNumber="27">
<emphasis id="B969EAC60008FF8DFF0BF923FEEC14F5" bold="true" box="[199,322,1734,1760]" pageId="26" pageNumber="27">Diagnosis.</emphasis>
<taxonomicName id="4C1D4D570008FF8DFE80F922FDB614F4" box="[332,536,1735,1761]" kingdom="Animalia" pageId="26" pageNumber="27" rank="subOrder" subOrder="Cephalozygoptera">Cephalozygoptera</taxonomicName>
are odonates most like 
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by a combination of: fore- and hind wings petiolate, with similar shape and venation including quadrangles that are closed and not crossed, CuA simple, and a well-developed nodus; an oblique thorax; gracile body; male clasping anal appendages comprised of a pair of dorsal cerci (without paddle-shaped extensions) and a pair of well-developed ventral paraprocts; and ocelli arranged in an equilateral triangle; but may be easily distinguished from them by any of:
</paragraph>
<paragraph id="8BA236D40008FF8DFF5BF87AFBB515C8" blockId="26.[151,1436,1951,2013]" pageId="26" pageNumber="27">
1- head width across eyes about twice the length from anterior margin of antefrons to posterior of occiput; [
<taxonomicName id="4C1D4D570008FF8DFA8BF87AFF5A15C8" class="Insecta" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="26" pageNumber="124" phylum="Arthropoda" rank="subOrder" subOrder="Zygoptera">Zygoptera</taxonomicName>
: usually about three to five times as wide]; and compound eyes that are:
</paragraph>
<paragraph id="8BA236D40009FF8CFF5BFF72FB3012C0" blockId="27.[151,1436,151,285]" pageId="27" pageNumber="28">
2- more or less adpressed to head, posterolateral corners protruding posteriorly to varying degrees, more or less rounded, sometimes acutely [
<taxonomicName id="4C1D4D570009FF8CFDC1FF5EFD2912C0" box="[525,647,187,213]" class="Insecta" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="27" pageNumber="124" phylum="Arthropoda" rank="subOrder" subOrder="Zygoptera">Zygoptera</taxonomicName>
: distinctly bulging laterally from head capsule];
</paragraph>
<paragraph id="8BA236D40009FF8CFF5BFF3AFBFA1308" blockId="27.[151,1436,151,285]" pageId="27" pageNumber="28">
3- separated by about one eye’s width dorsally (ratio of width between eyes/width eye has a range of 0.8–1.9 and a mean 1.0), n=11 (
<tableCitation id="C69F036F0009FF8CFE52FEE6FE5F1308" box="[414,497,259,285]" captionStart="TABLE 1" captionStartId="23.[151,239,485,511]" captionTargetPageId="23" captionText="TABLE 1. Head measurements of Cephalozygoptera and measurement ratios: WH, width of head; WBE, width between eyes; WE, width of one eye; LE, length of one eye; LH, length of head. See Fig. 10 for head images and text for measurement details." httpUri="http://table.plazi.org/id/DF62665C0005FF80FF5BFE00FC7F1036" pageId="27" pageNumber="28" tableUuid="DF62665C0005FF80FF5BFE00FC7F1036">Table 1</tableCitation>
) [
<taxonomicName id="4C1D4D570009FF8CFDC6FEE6FD2A1308" box="[522,644,259,285]" class="Insecta" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="27" pageNumber="124" phylum="Arthropoda" rank="subOrder" subOrder="Zygoptera">Zygoptera</taxonomicName>
: usually more than twice an eye’s width].
</paragraph>
<paragraph id="8BA236D40009FF8CFF0BFEAEFA3D1370" blockId="27.[151,1437,331,681]" box="[199,1427,331,357]" pageId="27" pageNumber="28">
<emphasis id="B969EAC60009FF8CFF0BFEAEFEDC1370" bold="true" box="[199,370,331,357]" pageId="27" pageNumber="28">Included taxa.</emphasis>
The families 
<taxonomicName id="4C1D4D570009FF8CFDC1FEAEFD1E1370" box="[525,688,331,357]" class="Insecta" family="Dysagrionidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="27" pageNumber="28" phylum="Arthropoda" rank="family">Dysagrionidae</taxonomicName>
and 
<taxonomicName id="4C1D4D570009FF8CFD2AFEAEFCDC1370" box="[742,882,331,357]" class="Insecta" family="Sieblosiidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="27" pageNumber="28" phylum="Arthropoda" rank="family">Sieblosiidae</taxonomicName>
, and possibly the 
<taxonomicName id="4C1D4D570009FF8CFBF6FEAEFAAA1370" authority="Archibald &amp; Cannings &amp; Erickson &amp; Bybee &amp; Mathewes, 2021" authorityName="Archibald &amp; Cannings &amp; Erickson &amp; Bybee &amp; Mathewes" authorityYear="2021" box="[1082,1284,331,357]" family="Whetwhetaksidae" kingdom="Animalia" order="Odonata" pageId="27" pageNumber="28" rank="family" status="fam. nov.">Whetwhetaksidae</taxonomicName>
, 
<taxonomicNameLabel id="A25A57BD0009FF8CFAC3FEAEFA211370" box="[1295,1423,331,357]" pageId="27" pageNumber="28" rank="family">new family</taxonomicNameLabel>
.
</paragraph>
</subSubSection>
<subSubSection id="C307655F0009FF8CFF0BFE8BFE9F102B" pageId="27" pageNumber="28" type="description">
<paragraph id="8BA236D40009FF8CFF0BFE8BFE9F102B" blockId="27.[151,1437,331,681]" pageId="27" pageNumber="28">
<emphasis id="B969EAC60009FF8CFF0BFE8BFEF7139D" bold="true" box="[199,345,366,392]" pageId="27" pageNumber="28">Description.</emphasis>
With character states of the diagnosis and further of the wings: Ax0 present; nodus at about a quarter wing length or more; anterior anal vein separates at a right angle from the posterior anal vein briefly before joining CuP (
<emphasis id="B969EAC60009FF8CFEAFFE52FEC513C5" box="[355,363,439,464]" italics="true" pageId="27" pageNumber="28">i</emphasis>
. 
<emphasis id="B969EAC60009FF8CFEBEFE52FED013C5" box="[370,382,439,464]" italics="true" pageId="27" pageNumber="28">e</emphasis>
., is briefly free distal to petiole); RP3-4 origin (midfork) in middle third between arculus and subnodus; antesubnodal space without crossveins; distinctive quadrangle (closed, distal side longer than proximal, posterior longer than anterior); long to very long pterostigmata (2.5 to about 10 times width if the 
<taxonomicName id="4C1D4D570009FF8CFB19FE1AFA32100C" authority="Archibald &amp; Cannings &amp; Erickson &amp; Bybee &amp; Mathewes, 2021" authorityName="Archibald &amp; Cannings &amp; Erickson &amp; Bybee &amp; Mathewes" authorityYear="2021" box="[1237,1436,511,537]" family="Whetwhetaksidae" kingdom="Animalia" order="Odonata" pageId="27" pageNumber="28" rank="family" status="fam. nov.">Whetwhetaksidae</taxonomicName>
is a member).
</paragraph>
</subSubSection>
<subSubSection id="C307655F0009FF8CFF0BFDA2FD6B10BC" pageId="27" pageNumber="28" type="etymology">
<paragraph id="8BA236D40009FF8CFF0BFDA2FD6B10BC" blockId="27.[151,1437,331,681]" pageId="27" pageNumber="28">
<emphasis id="B969EAC60009FF8CFF0BFDA2FEE01074" bold="true" box="[199,334,583,609]" pageId="27" pageNumber="28">Etymology.</emphasis>
The name 
<taxonomicName id="4C1D4D570009FF8CFE19FDA2FD0F1074" box="[469,673,583,609]" kingdom="Animalia" pageId="27" pageNumber="27" rank="subOrder" subOrder="Cephalozygoptera">Cephalozygoptera</taxonomicName>
is derived from the Greek 
<emphasis id="B969EAC60009FF8CFC29FDA3FB981074" box="[997,1078,582,609]" italics="true" pageId="27" pageNumber="28">κεφάλή</emphasis>
, 
<emphasis id="B969EAC60009FF8CFB8BFDA2FB2B1074" box="[1095,1157,583,609]" italics="true" pageId="27" pageNumber="28">kefáli</emphasis>
, “head”, and 
<taxonomicName id="4C1D4D570009FF8CFAE5FDA2FA321074" box="[1321,1436,583,609]" class="Insecta" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="27" pageNumber="124" phylum="Arthropoda" rank="subOrder" subOrder="Zygoptera">Zygoptera</taxonomicName>
(
<emphasis id="B969EAC60009FF8CFF6DFD8FFF701090" box="[161,222,618,645]" italics="true" pageId="27" pageNumber="28">ζυγός</emphasis>
, 
<emphasis id="B969EAC60009FF8CFF26FD8EFE841090" box="[234,298,619,645]" italics="true" pageId="27" pageNumber="28">zugós</emphasis>
, even + 
<emphasis id="B969EAC60009FF8CFE4AFD8FFE7F1090" box="[390,465,618,645]" italics="true" pageId="27" pageNumber="28">πτερόν</emphasis>
, 
<emphasis id="B969EAC60009FF8CFE12FD8EFD891090" box="[478,551,619,645]" italics="true" pageId="27" pageNumber="28">pterón</emphasis>
, wing, referring to the similar-shaped fore- and hind wings), indicating that they are distinguished from damselflies by their heads.
</paragraph>
</subSubSection>
<caption id="DF62665C0009FF8CFF5BF8EAFBF9155E" ID-DOI="http://doi.org/10.5281/zenodo.4558824" ID-Zenodo-Dep="4558824" httpUri="https://zenodo.org/record/4558824/files/figure.png" pageId="27" pageNumber="28" startId="27.[151,250,1807,1831]" targetBox="[151,1436,705,1771]" targetPageId="27">
<paragraph id="8BA236D40009FF8CFF5BF8EAFBF9155E" blockId="27.[151,1437,1807,1868]" pageId="27" pageNumber="28">
<emphasis id="B969EAC60009FF8CFF5BF8EAFEB11532" bold="true" box="[151,287,1807,1831]" pageId="27" pageNumber="28">FIGURE 11.</emphasis>
Crossvein Ax0 indicated by arrows in: A, 
<taxonomicName id="4C1D4D570009FF8CFD1AF8EAFC361532" authorityName="Archibald &amp; Cannings &amp; Erickson &amp; Bybee &amp; Mathewes" authorityYear="2021" box="[726,920,1807,1831]" class="Insecta" family="Dysagrionidae" genus="Okanagrion" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="27" pageNumber="28" phylum="Arthropoda" rank="species" species="beardi">
<emphasis id="B969EAC60009FF8CFD1AF8EAFC361532" box="[726,920,1807,1831]" italics="true" pageId="27" pageNumber="28">Okanagrion beardi</emphasis>
</taxonomicName>
holotype, F-791; B, 
<taxonomicName id="4C1D4D570009FF8CFBA3F8F5FA981532" authorityName="Archibald &amp; Cannings &amp; Erickson &amp; Bybee &amp; Mathewes" authorityYear="2021" box="[1135,1334,1808,1831]" class="Insecta" family="Dysagrionidae" genus="Dysagrion" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="27" pageNumber="28" phylum="Arthropoda" rank="species" species="pruettae">
<emphasis id="B969EAC60009FF8CFBA3F8F5FA981532" box="[1135,1334,1808,1831]" italics="true" pageId="27" pageNumber="28">Dysagrion pruettae</emphasis>
</taxonomicName>
holotype, SR 13-005-012: C, 
<taxonomicName id="4C1D4D570009FF8CFE92F8D6FDDB155E" authorityName="Archibald &amp; Cannings &amp; Erickson &amp; Bybee &amp; Mathewes" authorityYear="2021" box="[350,629,1843,1867]" class="Insecta" family="Thaumatoneuridae" genus="Okanopteryx" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="27" pageNumber="28" phylum="Arthropoda" rank="species" species="jeppesenorum">
<emphasis id="B969EAC60009FF8CFE92F8D6FDDB155E" box="[350,629,1843,1867]" italics="true" pageId="27" pageNumber="28">Okanopteryx jeppesenorum</emphasis>
</taxonomicName>
paratype 3, SR 15-003-001. Scale bar is 1 mm.
</paragraph>
</caption>
<subSubSection id="C307655F0009FF88FF0BF89CFD3D1194" lastPageId="31" lastPageNumber="32" pageId="27" pageNumber="28" type="discussion">
<paragraph id="8BA236D40009FF8BFF0BF89CFD52139C" blockId="27.[151,1436,1913,2012]" lastBlockId="28.[151,1437,150,2013]" lastPageId="28" lastPageNumber="29" pageId="27" pageNumber="28">
<emphasis id="B969EAC60009FF8CFF0BF89CFEE21586" bold="true" box="[199,332,1913,1939]" pageId="27" pageNumber="28">Discussion.</emphasis>
<emphasis id="B969EAC60009FF8CFE99F89CFD711586" box="[341,735,1913,1939]" italics="true" pageId="27" pageNumber="28">
<taxonomicName id="4C1D4D570009FF8CFE99F89CFE701586" box="[341,478,1913,1939]" class="Insecta" family="Sieblosiidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="27" pageNumber="28" phylum="Arthropoda" rank="family">Sieblosiidae</taxonomicName>
and 
<taxonomicName id="4C1D4D570009FF8CFDD7F89CFD711586" authority="Archibald &amp; Cannings &amp; Erickson &amp; Bybee &amp; Mathewes, 2021" authorityName="Archibald &amp; Cannings &amp; Erickson &amp; Bybee &amp; Mathewes" authorityYear="2021" box="[539,735,1913,1939]" family="Whetwhetaksidae" kingdom="Animalia" order="Odonata" pageId="27" pageNumber="28" rank="family" status="fam. nov.">Whetwhetaksidae</taxonomicName>
</emphasis>
. 
<bibRefCitation id="EF8C4B250009FF8CFD23F89CFC101586" author="Handlirsch, A." box="[751,958,1913,1939]" pageId="27" pageNumber="28" pagination="675 - 1130" refId="ref58492" refString="Handlirsch, A. (1907) Die Fossilen Insekten und die Phylogenie der Rezenten Formen, IV. Tertire Insekten. Ein Handbuch fur Palaontologen und Zoologen. Leipzig: Wilhelm Engelmann, pp. 675 - 1130." type="book chapter" year="1907">Handlirsch (1907)</bibRefCitation>
excluded 
<taxonomicName id="4C1D4D570009FF8CFBF9F89CFB511586" authorityName="Belaya" baseAuthorityName="Hagen" baseAuthorityYear="1858" box="[1077,1279,1913,1939]" class="Insecta" family="Sieblosiidae" genus="Sieblosia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="27" pageNumber="28" phylum="Arthropoda" rank="species" species="jucunda">
<emphasis id="B969EAC60009FF8CFBF9F89CFB511586" box="[1077,1279,1913,1939]" italics="true" pageId="27" pageNumber="28">Sieblosia jucunda</emphasis>
</taxonomicName>
from the 
<taxonomicName id="4C1D4D570009FF8CFAB8F89FFF4015AD" class="Insecta" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="27" pageNumber="124" phylum="Arthropoda" rank="subOrder" subOrder="Zygoptera">Zygoptera</taxonomicName>
, believing that it belongs to the 
<taxonomicName id="4C1D4D570009FF8CFD87F878FD5215A2" authorityName="Handlirsch" authorityYear="1906" box="[587,764,1949,1975]" class="Insecta" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="27" pageNumber="124" phylum="Arthropoda" rank="subOrder" subOrder="Anisozygoptera">Anisozygoptera</taxonomicName>
without explanation. This view was later rejected by 
<bibRefCitation id="EF8C4B250009FF8CFA86F878FF4C15CE" author="Fischer, C." pageId="27" pageNumber="28" pagination="211 - 220" refId="ref57382" refString="Fischer, C. (1974) Systematische Stellung der Gattung Sieblosia Handlirsch, 1906 (Zygoptera, Lestinoidea: Sieblosiidae). Odonatologica, 3, 211 - 220." type="journal article" year="1974">Fischer (1974)</bibRefCitation>
and Nel 
<emphasis id="B969EAC60009FF8CFE87F827FED315CE" box="[331,381,1985,2011]" italics="true" pageId="27" pageNumber="28">et al</emphasis>
. (1993), who restored it to the 
<taxonomicName id="4C1D4D570009FF8CFD2DF827FCF615C9" box="[737,856,1986,2012]" class="Insecta" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="27" pageNumber="124" phylum="Arthropoda" rank="subOrder" subOrder="Zygoptera">Zygoptera</taxonomicName>
. The 
<typeStatus id="54A688760009FF8CFC5AF827FC6815C9" box="[918,966,1986,2012]" pageId="27" pageNumber="28">type</typeStatus>
specimen (apparently lost: 
<bibRefCitation id="EF8C4B250009FF8CFACCF824FA3A15CE" author="Fischer, C." box="[1280,1428,1985,2011]" pageId="27" pageNumber="28" pagination="211 - 220" refId="ref57382" refString="Fischer, C. (1974) Systematische Stellung der Gattung Sieblosia Handlirsch, 1906 (Zygoptera, Lestinoidea: Sieblosiidae). Odonatologica, 3, 211 - 220." type="journal article" year="1974">Fischer 1974</bibRefCitation>
) was rather complete, including the head, which Hagen noted is almost as wide as long. In his drawing (
<bibRefCitation id="EF8C4B25000EFF8BFAC7FF72FA3612A4" author="Hagen, H." box="[1291,1432,150,177]" pageId="28" pageNumber="29" pagination="121 - 124" refId="ref58469" refString="Hagen, H. (1858) Zwei Libellen aus der Braunkohle von Sieblos. Paleontographica, 5, 121 - 124." type="journal article" year="1858">Hagen 1858</bibRefCitation>
: his Plate 24, 
<figureCitation id="13262A51000EFF8BFEEBFF5EFEC912C0" box="[295,359,187,213]" captionStart="FIGURE 1" captionStartId="7.[151,250,1249,1273]" captionTargetBox="[153,1436,185,1224]" captionTargetId="figure-17@7.[151,1436,181,1225]" captionTargetPageId="7" captionText="FIGURE 1. Map of Okanagan Highlands localities with Cephalozygoptera fossils: Driftwood Canyon Provincial Park, McAbee, and Republic." figureDoi="http://doi.org/10.5281/zenodo.4558798" httpUri="https://zenodo.org/record/4558798/files/figure.png" pageId="28" pageNumber="29">Fig. 1</figureCitation>
), although lacking detail, the head closely matches those of the 
<taxonomicName id="4C1D4D57000EFF8BFBF8FF5EFB7912C0" box="[1076,1239,187,213]" class="Insecta" family="Dysagrionidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="28" pageNumber="29" phylum="Arthropoda" rank="family">Dysagrionidae</taxonomicName>
(reproduced here as our 
<figureCitation id="13262A51000EFF8BFF13FF3AFE9012EC" box="[223,318,223,249]" captionStart="FIGURE 10" captionStartId="22.[151,250,1598,1622]" captionTargetBox="[327,1259,184,1571]" captionTargetId="figure-19@22.[321,1266,181,1574]" captionTargetPageId="22" captionText="FIGURE 10. Heads of Zygoptera (A, B) and comparative Anisoptera, Anisozygoptera, Tarsophlebiidae, Stenophlebiidae (C– H); and Cephalozygoptera (I–Q).A, Zygoptera, Heteropodagrion superbum Ris redrawn from Fig. 318 of Garrison et al. (2010); B, Zygoptera, Minagrion ribeiroi (Santos) redrawn from Fig. 1842 of Garrison et al. (2010); C, Anisoptera, Gomphidae, Cyanogomphus waltheri Selys redrawn from Fig. 383 of Garrison et al. (2006); D, Anisozygoptera, Epiophlebia laidlawi Tillyard redrawn from Fig. 2 of Büsse (2016); E, Anisoptera, Aeshnidae, Limnetron antarcticum Förster redrawn from Fig. 16 of Garrison et al. (2006); F, Stenophlebia sp. redrawn from Tillyard &amp; Fraser 1940, Fig, 4C; G, Tarsophlebiidae, Tarsophlebia eximia Hagen redrawn from Fleck et al. (2004) Fig. 1C; H, photograph of Tarsophlebia eximia in MCZ collections; I, Sieblosia jucunda redrawn from Hagen (1858, Plate 24, Fig. 1); J, Cephalozygoptera incertae sedis SR 14-003-004; K, Phenacolestes parallelus UCM-4545; L, Okanopteryx macabeensis, GSC 141101; M, Okanagrion beardi paratype 1, RBCM P1546; N, Okanopteryx fraseri GSC 141104; O, Okanopteryx macabeensis F-790; P, Dysagrion fredericii redrawn from Scudder 1890, plate 6, Fig. 10; Q, Dysagrion fredericii redrawn from Scudder 1890, plate 6, Fig. 2. G and H, I, J to O to scales, all 5 mm; others, originals without scales." figureDoi="http://doi.org/10.5281/zenodo.4558820" httpUri="https://zenodo.org/record/4558820/files/figure.png" pageId="28" pageNumber="29">Fig. 10 I</figureCitation>
). He assumed, however, that it was compressed post mortem (page 121: “kopf platt gedrückt”). Fleck 
<emphasis id="B969EAC6000EFF8BFF5BFEE6FF641308" box="[151,202,259,285]" italics="true" pageId="28" pageNumber="29">et al</emphasis>
. (2004) recognised that the head of 
<taxonomicName id="4C1D4D57000EFF8BFDABFEE6FD5E1308" box="[615,752,259,285]" class="Insecta" family="Sieblosiidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="28" pageNumber="29" phylum="Arthropoda" rank="family">Sieblosiidae</taxonomicName>
is not transversely elongate with the compound eyes placed far apart. They further found that their wings do not share any unambiguous apomorphy with the 
<taxonomicName id="4C1D4D57000EFF8BFB34FEC2FAC51354" box="[1272,1387,295,321]" class="Insecta" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="28" pageNumber="124" phylum="Arthropoda" rank="subOrder" subOrder="Zygoptera">Zygoptera</taxonomicName>
and that particularly with their nodal morphology and curved CuP they may not belong to the suborder. Following this, 
<bibRefCitation id="EF8C4B25000EFF8BFF5BFE8AFE96139C" author="Bechly, G." box="[151,312,366,393]" pageId="28" pageNumber="29" pagination="423 - 464" refId="ref56191" refString="Bechly, G. (2015) Fossile Libellennachweise aus Deutschland (Odonatoptera). Libellula Supplement, 14, 423 - 464." type="journal article" year="2015">Bechly (2015)</bibRefCitation>
treated the family as “ 
<taxonomicName id="4C1D4D57000EFF8BFDF8FE8AFD40139C" authorityName="Handlirsch" authorityYear="1906" box="[564,750,367,393]" class="Insecta" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="28" pageNumber="124" phylum="Arthropoda" rank="subOrder" subOrder="Anisozygoptera">Anisozygoptera</taxonomicName>
”.
</paragraph>
<paragraph id="8BA236D4000EFF8BFF0BFE76FAC21100" blockId="28.[151,1437,150,2013]" pageId="28" pageNumber="29">
The wings of 
<taxonomicName id="4C1D4D57000EFF8BFEA8FE76FE4313B8" box="[356,493,403,429]" class="Insecta" family="Sieblosiidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="28" pageNumber="29" phylum="Arthropoda" rank="family">Sieblosiidae</taxonomicName>
and 
<taxonomicName id="4C1D4D57000EFF8BFDE9FE76FD6613B8" box="[549,712,403,429]" class="Insecta" family="Dysagrionidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="28" pageNumber="29" phylum="Arthropoda" rank="family">Dysagrionidae</taxonomicName>
have strong similarities (
<bibRefCitation id="EF8C4B25000EFF8BFC2BFE76FB3113B8" author="Nel, A. &amp; Petrulevicius, J. F. &amp; Gentilini, G. &amp; Martinez-Delclos, X." box="[999,1183,402,429]" pageId="28" pageNumber="29" pagination="219 - 233" refId="ref60383" refString="Nel, A., Petrulevicius, J. F., Gentilini, G. &amp; Martinez-Delclos, X. (2005 a) Phylogenetic analysis of the Cenozoic family Sieblo- siidae (Insecta: Odonata), with description of new taxa from Russia, Italy and France. Geobios, 38, 219 - 233. https: // doi. org / 10.1016 / j. geobios. 2003.10.007" type="journal article" year="2005">
Nel 
<emphasis id="B969EAC6000EFF8BFBD5FE76FBE513B8" box="[1049,1099,403,429]" italics="true" pageId="28" pageNumber="29">et al</emphasis>
. 2005a
</bibRefCitation>
) (
<figureCitation id="13262A51000EFF8BFB7BFE76FAA513B8" box="[1207,1291,403,429]" captionStart="FIGURE 12" captionStartId="29.[151,250,1911,1935]" captionTargetBox="[154,1432,569,1875]" captionTargetId="figure-179@29.[151,1436,561,1887]" captionTargetPageId="29" captionText="FIGURE 12. Representative Sieblosiidae, A, Stenolestes fasciata Nel et al.; B, Miostenolestes zherikhini; and Frenguelliidae, C, Treintamilun vuelvenlucha; D, Frenguellia patagonica Petrulevičius &amp; Nel. Redrawn from: A, B, Nel et al. (2005a); C, Petrulevičius (2017); D, Petrulevičius &amp; Nel (2007). An asterisk indicates the oblique vein “O” immediately to its right. Scale bars, 5 mm." figureDoi="http://doi.org/10.5281/zenodo.4558826" httpUri="https://zenodo.org/record/4558826/files/figure.png" pageId="28" pageNumber="29">Fig. 12</figureCitation>
). They share with 
<taxonomicName id="4C1D4D57000EFF8BFF03FE52FEDC13C4" box="[207,370,439,465]" class="Insecta" family="Dysagrionidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="28" pageNumber="29" phylum="Arthropoda" rank="family">Dysagrionidae</taxonomicName>
character states 2 through 7 of its diagnosis, notably the distinctive quadrangle shape, character state 3. The wings of both are broad, with a short petiole and bear a long pterostigma. Crossvein Ax0 has been found in numerous species, see 
<taxonomicName id="4C1D4D57000EFF8BFE7AFE1AFD7E100C" authorityName="Nel &amp; Fleck" authorityYear="2012" box="[438,720,511,537]" class="Insecta" family="Sieblosiidae" genus="Germanostenolestes" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="28" pageNumber="29" phylum="Arthropoda" rank="species" species="lutzi">
<emphasis id="B969EAC6000EFF8BFE7AFE1AFD7E100C" box="[438,720,511,537]" italics="true" pageId="28" pageNumber="29">Germanostenolestes lutzi</emphasis>
</taxonomicName>
(2012, their 
<figureCitation id="13262A51000EFF8BFCAFFE1AFC08100C" box="[867,934,511,537]" captionStart="FIGURE 3" captionStartId="10.[151,250,1805,1829]" captionTargetBox="[151,1436,181,1781]" captionTargetId="figure-19@10.[151,1436,181,1781]" captionTargetPageId="10" captionText="FIGURE 3. Phenacolestes parallelus. A, specimen YPMIP-220974 of the Yale Peabody Museum; B, drawing from A, photograph provided by Peabody Museum personnel; C, holotype, UCM-4503 of the University of Colorado (Boulder) paleontology collections; D, drawing of C, photograph provided by University of Colorado personnel; E, UCM-4545, University of Colorado (Boulder). A–D to scale, E to scale, both scale bars 5 mm." figureDoi="http://doi.org/10.5281/zenodo.4558803" httpUri="https://zenodo.org/record/4558803/files/figure.png" pageId="28" pageNumber="29">Fig. 3</figureCitation>
), 
<taxonomicName id="4C1D4D57000EFF8BFC71FE1AFB66100C" authority="Nel" authorityName="Nel" authorityYear="1986" box="[957,1224,511,537]" class="Insecta" family="Sieblosiidae" genus="Stenolestes" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="28" pageNumber="29" phylum="Arthropoda" rank="species" species="fischeri">
<emphasis id="B969EAC6000EFF8BFC71FE1AFB3B100C" box="[957,1173,511,537]" italics="true" pageId="28" pageNumber="29">Stenolestes fischeri</emphasis>
Nel
</taxonomicName>
, 
<typeStatus id="54A68876000EFF8BFB18FE1AFA98100C" box="[1236,1334,511,537]" pageId="28" pageNumber="29" type="holotype">holotype</typeStatus>
MNHN- LP-R.06677 (
<bibRefCitation id="EF8C4B25000EFF8BFEFEFDC6FE4A1028" author="Nel, A. &amp; Martinez-Delclos, X. &amp; Papier, F. &amp; Oudard, J." box="[306,484,547,573]" pageId="28" pageNumber="29" pagination="231 - 258" refId="ref60193" refString="Nel, A., Martinez-Delclos, X., Papier, F. &amp; Oudard, J. (1997) New Tertiary fossil Odonata from France. (Sieblosiidae, Lestidae, Coenagrioniidae, Megapodagrionidae, Libellulidae). Deutsche Entomologische Zeitschrift, 44, 231 - 258. https: // doi. org / 10.1002 / mmnd. 19970440210" type="journal article" year="1997">
Nel 
<emphasis id="B969EAC6000EFF8BFEA9FDC6FE371028" box="[357,409,547,573]" italics="true" pageId="28" pageNumber="29">et al</emphasis>
. 1997
</bibRefCitation>
, their 
<figureCitation id="13262A51000EFF8BFDE2FDC6FDDD102B" box="[558,627,547,574]" captionStart="FIGURE 6" captionStartId="13.[151,250,1943,1967]" captionTargetBox="[262,1315,184,1919]" captionTargetId="figure-17@13.[262,1326,181,1919]" captionTargetPageId="13" captionText="FIGURE 6. Dysagrion fredericii Scudder’s number 4167/4168 (now MCZ numbers 381A, B “type”): A, photograph; B, drawing from a photograph supplied by MCZ personnel. Both to scale, 5 mm." figureDoi="http://doi.org/10.5281/zenodo.4558809" httpUri="https://zenodo.org/record/4558809/files/figure.png" pageId="28" pageNumber="29">Fig. 6</figureCitation>
), and in 
<taxonomicName id="4C1D4D57000EFF8BFD10FDC6FB811028" authority="Nel et al." authorityName="Nel" box="[732,1071,547,573]" class="Insecta" family="Sieblosiidae" genus="Stenolestes" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="28" pageNumber="29" phylum="Arthropoda" rank="species" species="fasciata">
<emphasis id="B969EAC6000EFF8BFD10FDC6FC141028" box="[732,954,547,573]" italics="true" pageId="28" pageNumber="29">Stenolestes fasciata</emphasis>
Nel 
<emphasis id="B969EAC6000EFF8BFC39FDC6FB861028" box="[1013,1064,547,573]" italics="true" pageId="28" pageNumber="29">et al</emphasis>
.
</taxonomicName>
(
<bibRefCitation id="EF8C4B25000EFF8BFB8CFDC6FB501028" author="Nel, A. &amp; Petrulevicius, J. F. &amp; Gentilini, G. &amp; Martinez-Delclos, X." box="[1088,1278,546,573]" pageId="28" pageNumber="29" pagination="219 - 233" refId="ref60383" refString="Nel, A., Petrulevicius, J. F., Gentilini, G. &amp; Martinez-Delclos, X. (2005 a) Phylogenetic analysis of the Cenozoic family Sieblo- siidae (Insecta: Odonata), with description of new taxa from Russia, Italy and France. Geobios, 38, 219 - 233. https: // doi. org / 10.1016 / j. geobios. 2003.10.007" type="journal article" year="2005">
Nel 
<emphasis id="B969EAC6000EFF8BFBBFFDC6FB091028" box="[1139,1191,547,573]" italics="true" pageId="28" pageNumber="29">et al</emphasis>
. 2005a
</bibRefCitation>
, their 
<figureCitation id="13262A51000EFF8BFA84FDC6FA201028" box="[1352,1422,547,573]" captionStart="FIGURE 9" captionStartId="18.[151,250,1817,1841]" captionTargetBox="[210,1368,188,1783]" captionTargetId="figure-19@18.[206,1381,181,1793]" captionTargetPageId="18" captionText="FIGURE 9. Wing drawings of: Valerea multicellulata redrawn from Fig. 1 of Garrouste et al. (2017); Chickaloon Formation (Alaska) specimen redrawn from Fig. 3 of Garrouste &amp; Nel (2017); cf. Dysagrionidae gen. et. sp. A redrawn from Fig. 4 of Petrulevičius et al. (2008) (there as “Megapodagrionidae gen. et. sp. A”); Thanetophilosina menatensis redrawn from Fig. 21 of Nel et al. (1997); NHMUK I.9866 (Isle of Wight) redrawn from Fig. 28 of Nel &amp; Fleck (2014); Palaeodysagrion youlini redrawn from Fig. 2 of Zheng et al. (2018); Electrodysagrion neli redrawn from Fig. 2 of Zheng et al. (2019); Burmadysagrion zhangi redrawn from Fig. 3 of Zheng et al. (2016a). To associated scales, all 5 mm. Asterisks indicate quadrangles." figureDoi="http://doi.org/10.5281/zenodo.4558818" httpUri="https://zenodo.org/record/4558818/files/figure.png" pageId="28" pageNumber="29">Fig. 9</figureCitation>
), 
<taxonomicName id="4C1D4D57000EFF8BFF5BFDA2FDA81074" authority="Nel et al." authorityName="Nel" box="[151,518,583,609]" class="Insecta" family="Sieblosiidae" genus="Stenolestes" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="28" pageNumber="29" phylum="Arthropoda" rank="species" species="cerestensis">
<emphasis id="B969EAC6000EFF8BFF5BFDA2FE381075" box="[151,406,583,609]" italics="true" pageId="28" pageNumber="29">Stenolestes cerestensis</emphasis>
Nel 
<emphasis id="B969EAC6000EFF8BFE02FDA2FDAE1074" box="[462,512,583,609]" italics="true" pageId="28" pageNumber="29">et al</emphasis>
.
</taxonomicName>
(
<bibRefCitation id="EF8C4B25000EFF8BFDDAFDA2FD611074" author="Nel, A. &amp; Petrulevicius, J. F. &amp; Gentilini, G. &amp; Martinez-Delclos, X." box="[534,719,582,609]" pageId="28" pageNumber="29" pagination="219 - 233" refId="ref60383" refString="Nel, A., Petrulevicius, J. F., Gentilini, G. &amp; Martinez-Delclos, X. (2005 a) Phylogenetic analysis of the Cenozoic family Sieblo- siidae (Insecta: Odonata), with description of new taxa from Russia, Italy and France. Geobios, 38, 219 - 233. https: // doi. org / 10.1016 / j. geobios. 2003.10.007" type="journal article" year="2005">
Nel 
<emphasis id="B969EAC6000EFF8BFD84FDA2FDD41074" box="[584,634,583,609]" italics="true" pageId="28" pageNumber="29">et al</emphasis>
. 2005a
</bibRefCitation>
, their 
<figureCitation id="13262A51000EFF8BFCDBFDA2FCC41074" box="[791,874,583,609]" captionStart="FIGURE 14" captionStartId="35.[151,250,1193,1217]" captionTargetBox="[156,1427,245,1166]" captionTargetId="figure-47@35.[151,1436,237,1170]" captionTargetPageId="35" captionText="FIGURE 14. Tree resulting from parsimony-based analyses for Analysis 1: Tarsophlebiidae, the suborders of Odonata and former members of Dysagrionidae (Burmadysagrion, Electrodysagrion, Paleodysagrion). One single most-parsimonious tree (MPT) based on 23 continuous and 79 discrete characters. Symmetrical resampling values Ξ70 are labelled at the nodes." figureDoi="http://doi.org/10.5281/zenodo.4558832" httpUri="https://zenodo.org/record/4558832/files/figure.png" pageId="28" pageNumber="29">Fig. 14</figureCitation>
), and 
<taxonomicName id="4C1D4D57000EFF8BFC7CFDA2FB601074" box="[944,1230,583,609]" class="Insecta" family="Sieblosiidae" genus="Miostenolestes" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="28" pageNumber="29" phylum="Arthropoda" rank="species" species="zherikhini">
<emphasis id="B969EAC6000EFF8BFC7CFDA2FB601074" box="[944,1230,583,609]" italics="true" pageId="28" pageNumber="29">Miostenolestes zherikhini</emphasis>
</taxonomicName>
(
<bibRefCitation id="EF8C4B25000EFF8BFB12FDA2FA391074" author="Nel, A. &amp; Petrulevicius, J. F. &amp; Gentilini, G. &amp; Martinez-Delclos, X." box="[1246,1431,582,609]" pageId="28" pageNumber="29" pagination="219 - 233" refId="ref60383" refString="Nel, A., Petrulevicius, J. F., Gentilini, G. &amp; Martinez-Delclos, X. (2005 a) Phylogenetic analysis of the Cenozoic family Sieblo- siidae (Insecta: Odonata), with description of new taxa from Russia, Italy and France. Geobios, 38, 219 - 233. https: // doi. org / 10.1016 / j. geobios. 2003.10.007" type="journal article" year="2005">
Nel 
<emphasis id="B969EAC6000EFF8BFADCFDA2FAEC1074" box="[1296,1346,583,609]" italics="true" pageId="28" pageNumber="29">et al</emphasis>
. 2005a
</bibRefCitation>
, their 
<figureCitation id="13262A51000EFF8BFF1FFD8EFEB71090" box="[211,281,618,645]" captionStart="FIGURE 5" captionStartId="12.[151,250,1943,1967]" captionTargetBox="[250,1324,200,1919]" captionTargetId="figure-19@12.[250,1338,181,1919]" captionTargetPageId="12" captionText="FIGURE 5. Dysagrion lakesii Scudder type specimen MCZ 4101 (Packard number 259, not previously illustrated): A, photograph; B, drawing from a photograph supplied by MCZ personnel. Both to scale, 5 mm." figureDoi="http://doi.org/10.5281/zenodo.4558807" httpUri="https://zenodo.org/record/4558807/files/figure.png" pageId="28" pageNumber="29">Fig. 5</figureCitation>
). As in the 
<taxonomicName id="4C1D4D57000EFF8BFE6DFD8EFDE91090" box="[417,583,619,645]" class="Insecta" family="Dysagrionidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="28" pageNumber="29" phylum="Arthropoda" rank="family">Dysagrionidae</taxonomicName>
, the wing base of most published 
<taxonomicName id="4C1D4D57000EFF8BFC1CFD8EFBF71090" box="[976,1113,619,645]" class="Insecta" family="Sieblosiidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="28" pageNumber="29" phylum="Arthropoda" rank="family">Sieblosiidae</taxonomicName>
fossils where Ax0 might be found is missing or damaged, and so this represents a minimum of specimens where this may be present; judging from published drawings and photographs, none may be excluded as possessing it. As in Petrolestinae but not the 
<taxonomicName id="4C1D4D57000EFF8BFF5BFD32FE9310E4" box="[151,317,727,753]" class="Insecta" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="28" pageNumber="29" phylum="Arthropoda" rank="subFamily" subFamily="Dysagrioninae">Dysagrioninae</taxonomicName>
, the origin of IR2 is in the middle third between the arculus and nodus, close to that of RP3-4. 
<bibRefCitation id="EF8C4B25000EFF8BFA9BFD32FEFF1100" author="Nel, A. &amp; Paicheler, J. - C." pageId="28" pageNumber="29" pagination="1 - 59" refId="ref60306" refString="Nel, A. &amp; Paicheler, J. - C. (1994) Les Lestoidea (Odonata, Zygoptera) fossiles: Un inventaire critique. Annales de Paleontologie, 80, 1 - 59." type="journal article" year="1994">Nel &amp; Paicheler (1994)</bibRefCitation>
suggested that 
<taxonomicName id="4C1D4D57000EFF8BFE32FD1EFD291100" box="[510,647,763,789]" class="Insecta" family="Sieblosiidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="28" pageNumber="29" phylum="Arthropoda" rank="family">Sieblosiidae</taxonomicName>
and 
<taxonomicName id="4C1D4D57000EFF8BFD72FD1EFCCF1100" box="[702,865,763,789]" class="Insecta" family="Dysagrionidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="28" pageNumber="29" phylum="Arthropoda" rank="family">Dysagrionidae</taxonomicName>
might be an example of convergent evolution.
</paragraph>
<paragraph id="8BA236D4000EFF8BFF0BFCFAFADD11DC" blockId="28.[151,1437,150,2013]" pageId="28" pageNumber="29">
<taxonomicName id="4C1D4D57000EFF8BFF0BFCFAFEFE112C" box="[199,336,799,825]" class="Insecta" family="Sieblosiidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="28" pageNumber="29" phylum="Arthropoda" rank="family">Sieblosiidae</taxonomicName>
differ from the above diagnosis of the 
<taxonomicName id="4C1D4D57000EFF8BFCDAFCFAFC17112C" box="[790,953,799,825]" class="Insecta" family="Dysagrionidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="28" pageNumber="29" phylum="Arthropoda" rank="family">Dysagrionidae</taxonomicName>
by character state 8, width of the CuA–A space, which is usually two or more cells wide as in 
<taxonomicName id="4C1D4D57000EFF8BFD22FCA6FC3A1148" box="[750,916,835,861]" class="Insecta" family="Dysagrionidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="28" pageNumber="29" phylum="Arthropoda" rank="family">Dysagrionidae</taxonomicName>
, but may be one cell wide (
<emphasis id="B969EAC6000EFF8BFB03FCA6FB591149" box="[1231,1271,835,860]" italics="true" pageId="28" pageNumber="29">e.g.</emphasis>
, see 
<figureCitation id="13262A51000EFF8BFAFEFCA6FA361148" box="[1330,1432,835,861]" captionStart="FIGURE 12" captionStartId="29.[151,250,1911,1935]" captionTargetBox="[154,1432,569,1875]" captionTargetId="figure-179@29.[151,1436,561,1887]" captionTargetPageId="29" captionText="FIGURE 12. Representative Sieblosiidae, A, Stenolestes fasciata Nel et al.; B, Miostenolestes zherikhini; and Frenguelliidae, C, Treintamilun vuelvenlucha; D, Frenguellia patagonica Petrulevičius &amp; Nel. Redrawn from: A, B, Nel et al. (2005a); C, Petrulevičius (2017); D, Petrulevičius &amp; Nel (2007). An asterisk indicates the oblique vein “O” immediately to its right. Scale bars, 5 mm." figureDoi="http://doi.org/10.5281/zenodo.4558826" httpUri="https://zenodo.org/record/4558826/files/figure.png" pageId="28" pageNumber="29">Fig. 12B</figureCitation>
, 
<taxonomicName id="4C1D4D57000EFF8BFF5BFC82FE191194" box="[151,439,871,897]" class="Insecta" family="Sieblosiidae" genus="Miostenolestes" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="28" pageNumber="29" phylum="Arthropoda" rank="species" species="zherikhini">
<emphasis id="B969EAC6000EFF8BFF5BFC82FE191194" box="[151,439,871,897]" italics="true" pageId="28" pageNumber="29">Miostenolestes zherikhini</emphasis>
</taxonomicName>
); by 9, CuA terminates on the margin proximal to the mid-wing; and 1, crossvein O is present. The nodus of 
<taxonomicName id="4C1D4D57000EFF8BFE5DFC6EFDB411B0" box="[401,538,907,933]" class="Insecta" family="Sieblosiidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="28" pageNumber="29" phylum="Arthropoda" rank="family">Sieblosiidae</taxonomicName>
differs as well by ScP passing through it and terminating on the anterior margin at an oblique angle. Although the pterostigmata of 
<taxonomicName id="4C1D4D57000EFF8BFD1CFC4AFCDD11DC" box="[720,883,943,969]" class="Insecta" family="Dysagrionidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="28" pageNumber="29" phylum="Arthropoda" rank="family">Dysagrionidae</taxonomicName>
are long, those of the 
<taxonomicName id="4C1D4D57000EFF8BFBA1FC4AFB5811DC" box="[1133,1270,943,969]" class="Insecta" family="Sieblosiidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="28" pageNumber="29" phylum="Arthropoda" rank="family">Sieblosiidae</taxonomicName>
are longer.
</paragraph>
<paragraph id="8BA236D4000EFF8BFF0BFC36FDCE164C" blockId="28.[151,1437,150,2013]" pageId="28" pageNumber="29">
The new family 
<taxonomicName id="4C1D4D57000EFF8BFE4EFC36FDE711F8" authorityName="Archibald &amp; Cannings &amp; Erickson &amp; Bybee &amp; Mathewes" authorityYear="2021" box="[386,585,979,1005]" family="Whetwhetaksidae" kingdom="Animalia" order="Odonata" pageId="28" pageNumber="29" rank="family">Whetwhetaksidae</taxonomicName>
is only known by wings, which share character states 1 and 3 through 9 of the 
<taxonomicName id="4C1D4D57000EFF8BFF0DFC12FECA1604" box="[193,356,1015,1041]" class="Insecta" family="Dysagrionidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="28" pageNumber="29" phylum="Arthropoda" rank="family">Dysagrionidae</taxonomicName>
diagnosis and their general shape is like that of the 
<taxonomicName id="4C1D4D57000EFF8BFC67FC12FBE01604" box="[939,1102,1015,1041]" class="Insecta" family="Dysagrionidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="28" pageNumber="29" phylum="Arthropoda" rank="family">Dysagrionidae</taxonomicName>
and 
<taxonomicName id="4C1D4D57000EFF8BFB49FC12FABC1604" box="[1157,1298,1015,1041]" class="Insecta" family="Sieblosiidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="28" pageNumber="29" phylum="Arthropoda" rank="family">Sieblosiidae</taxonomicName>
: broad, with a short petiole. The origin of IR2 is in the middle third between the arculus and nodus as in 
<taxonomicName id="4C1D4D57000EFF8BFB74FBFEFAEF1620" box="[1208,1345,1051,1077]" class="Insecta" family="Sieblosiidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="28" pageNumber="29" phylum="Arthropoda" rank="family">Sieblosiidae</taxonomicName>
and the Petrolestinae, but not the 
<taxonomicName id="4C1D4D57000EFF8BFE79FBDAFDF2164C" box="[437,604,1087,1113]" class="Insecta" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="28" pageNumber="29" phylum="Arthropoda" rank="subFamily" subFamily="Dysagrioninae">Dysagrioninae</taxonomicName>
.
</paragraph>
<paragraph id="8BA236D4000EFF8BFF0BFB86FAB31740" blockId="28.[151,1437,150,2013]" pageId="28" pageNumber="29">
These wings are notably distinct, however, as the arculus is in a proximal position near Ax1 as is found in some odonates outside of the 
<taxonomicName id="4C1D4D57000EFF8BFE52FB62FDBB16B4" box="[414,533,1159,1185]" class="Insecta" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="28" pageNumber="124" phylum="Arthropoda" rank="subOrder" subOrder="Zygoptera">Zygoptera</taxonomicName>
. This feature is unknown in extant 
<taxonomicName id="4C1D4D57000EFF8BFC55FB62FBA216B4" box="[921,1036,1159,1185]" class="Insecta" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="28" pageNumber="124" phylum="Arthropoda" rank="subOrder" subOrder="Zygoptera">Zygoptera</taxonomicName>
except in the Amazonian 
<taxonomicName id="4C1D4D57000EFF8BFAE0FB62FEC616D0" authority="Selys" authorityName="Selys" class="Insecta" family="Dicteriadidae" genus="Heliocharis" kingdom="Animalia" order="Odonata" pageId="28" pageNumber="29" phylum="Arthropoda" rank="species" species="amazona">
<emphasis id="B969EAC6000EFF8BFAE0FB62FE8C16D1" italics="true" pageId="28" pageNumber="29">Heliocharis amazona</emphasis>
Selys
</taxonomicName>
(Calopterygoidea, 
<taxonomicName id="4C1D4D57000EFF8BFD8EFB4EFD7016D3" authorityName="Selys" authorityYear="1853" box="[578,734,1195,1222]" class="Insecta" family="Dicteriadidae" kingdom="Animalia" order="Odonata" pageId="28" pageNumber="29" phylum="Arthropoda" rank="family">Dicteriadidae</taxonomicName>
), and is present in two Paleogene 
<taxonomicName id="4C1D4D57000EFF8BFBA8FB4EFB4916D0" baseAuthorityName="Jacobson &amp; Bianchi" baseAuthorityYear="1905" box="[1124,1255,1195,1221]" class="Insecta" family="Euphaeidae" kingdom="Animalia" order="Odonata" pageId="28" pageNumber="29" phylum="Arthropoda" rank="family">Euphaeidae</taxonomicName>
(the Priabonian 
<taxonomicName id="4C1D4D57000EFF8BFF5BFB2AFE7416FC" authority="Bechly" authorityName="Bechly" authorityYear="1998" box="[151,474,1231,1257]" class="Insecta" family="Epallagidae" genus="Litheuphaea" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="28" pageNumber="29" phylum="Arthropoda" rank="species" species="ludwigi">
<emphasis id="B969EAC6000EFF8BFF5BFB2AFE2F16FC" box="[151,385,1231,1257]" italics="true" pageId="28" pageNumber="29">Litheuphaea ludwigi</emphasis>
Bechly
</taxonomicName>
and the Chattian 
<taxonomicName id="4C1D4D57000EFF8BFD60FB2AFB8C16FC" authority="Nel" authorityName="Nel" authorityYear="1988" box="[684,1058,1231,1257]" class="Insecta" family="Epallagidae" genus="Parazacallites" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="28" pageNumber="29" phylum="Arthropoda" rank="species" species="aquisextanea">
<emphasis id="B969EAC6000EFF8BFD60FB2AFC4516FD" box="[684,1003,1231,1257]" italics="true" pageId="28" pageNumber="29">Parazacallites aquisextanea</emphasis>
Nel
</taxonomicName>
: Bechly 1998; 
<bibRefCitation id="EF8C4B25000EFF8BFB1EFB2AFAEF16FC" author="Nel, A." box="[1234,1345,1231,1257]" pageId="28" pageNumber="29" pagination="175 - 179" refId="ref59539" refString="Nel, A. (1988) Parazacallitinae, nouvelle sous-famille et premier Epallagidae' de l'Oligocene europeen (Odonata, Zygoptera). Bulletin du Museum National d'Histoire Naturelle, Paris, 10, 175 - 179." type="journal article" year="1988">Nel 1988</bibRefCitation>
). 
<taxonomicName id="4C1D4D57000EFF8BFA9AFB2AFE8F1718" authority="Archibald &amp; Cannings &amp; Erickson &amp; Bybee &amp; Mathewes, 2021" authorityName="Archibald &amp; Cannings &amp; Erickson &amp; Bybee &amp; Mathewes" authorityYear="2021" family="Whetwhetaksidae" kingdom="Animalia" order="Odonata" pageId="28" pageNumber="28" rank="family" status="fam. nov.">Whetwhetaksidae</taxonomicName>
possess an extremely long pterostigma, unknown in 
<taxonomicName id="4C1D4D57000EFF8BFCBCFB16FC481718" box="[880,998,1267,1293]" class="Insecta" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="28" pageNumber="124" phylum="Arthropoda" rank="subOrder" subOrder="Zygoptera">Zygoptera</taxonomicName>
, but like those found in the anisopteran 
<taxonomicName id="4C1D4D57000EFF8BFF5BFAF2FEB71724" box="[151,281,1303,1329]" class="Insecta" family="Petaluridae" kingdom="Animalia" order="Odonata" pageId="28" pageNumber="29" phylum="Arthropoda" rank="family">Petaluridae</taxonomicName>
. It is up to about ten times longer than wide, and is about a quarter to almost a third the length of the nodus to the wing apex. Further, the nodus is distant from the wing base, at about 38–40% wing length.
</paragraph>
<paragraph id="8BA236D4000EFF8BFF0BFABAFE2F1460" blockId="28.[151,1437,150,2013]" pageId="28" pageNumber="29">
<emphasis id="B969EAC6000EFF8BFF0BFABAFD10176C" box="[199,702,1375,1401]" italics="true" pageId="28" pageNumber="29">Groups with similar wings that are excluded</emphasis>
. The 
<taxonomicName id="4C1D4D57000EFF8BFD33FABAFC0F176C" authorityName="Petrulevicius &amp; Nel" authorityYear="2003" box="[767,929,1375,1401]" class="Insecta" family="Frenguelliidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="28" pageNumber="29" phylum="Arthropoda" rank="family">Frenguelliidae</taxonomicName>
are odonates from the Ypresian and Lutetian of Patagonia, 
<collectingCountry id="F30A7644000EFF8BFEFCFA66FE0C1788" box="[304,418,1411,1437]" name="Argentina" pageId="28" pageNumber="29">Argentina</collectingCountry>
, known only from wings (
<figureCitation id="13262A51000EFF8BFD0BFA66FCFE1788" box="[711,848,1411,1437]" captionStart="FIGURE 12" captionStartId="29.[151,250,1911,1935]" captionTargetBox="[154,1432,569,1875]" captionTargetId="figure-179@29.[151,1436,561,1887]" captionTargetPageId="29" captionText="FIGURE 12. Representative Sieblosiidae, A, Stenolestes fasciata Nel et al.; B, Miostenolestes zherikhini; and Frenguelliidae, C, Treintamilun vuelvenlucha; D, Frenguellia patagonica Petrulevičius &amp; Nel. Redrawn from: A, B, Nel et al. (2005a); C, Petrulevičius (2017); D, Petrulevičius &amp; Nel (2007). An asterisk indicates the oblique vein “O” immediately to its right. Scale bars, 5 mm." figureDoi="http://doi.org/10.5281/zenodo.4558826" httpUri="https://zenodo.org/record/4558826/files/figure.png" pageId="28" pageNumber="29">Fig. 12C, D</figureCitation>
). They are considered by some authors not to belong to the 
<taxonomicName id="4C1D4D57000EFF8BFF12FA42FEFF17D4" box="[222,337,1447,1473]" class="Insecta" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="28" pageNumber="124" phylum="Arthropoda" rank="subOrder" subOrder="Zygoptera">Zygoptera</taxonomicName>
by their curved CuP (
<bibRefCitation id="EF8C4B25000EFF8BFD8BFA42FCC417D4" author="Petrulevicius, J. F. &amp; Nel, A." box="[583,874,1447,1473]" pageId="28" pageNumber="29" pagination="2909 - 2917" refId="ref60790" refString="Petrulevicius, J. F. &amp; Nel, A. (2003) Frenguelliidae, a new family of dragonflies from the earliest Eocene of Argentina (Insecta: Odonata): phylogenetic relationships within Odonata. Journal of Natural History, 37, 2909 - 2917. https: // doi. org / 10.1080 / 0022293021000007543" type="journal article" year="2003">Petrulevičius &amp; Nel 2003</bibRefCitation>
, 2007, 2013; 
<bibRefCitation id="EF8C4B25000EFF8BFBCFFA42FB7917D4" author="Petrulevicius, J. F." box="[1027,1239,1447,1473]" pageId="28" pageNumber="29" pagination="367 - 374" refId="ref60663" refString="Petrulevicius, J. F. (2017) First Frenguelliidae (Insecta: Odonata) from the middle Eocene of Rio Pichileufu, Patagonia, Argen- tina. Arquivos Entomoloxicos, 18, 367 - 374." type="journal article" year="2017">Petrulevičius 2017</bibRefCitation>
, 
<bibRefCitation id="EF8C4B25000EFF8BFB2EFA42FAB117D4" author="Petrulevicius, J. F." box="[1250,1311,1447,1473]" pageId="28" pageNumber="29" pagination="591 - 595" refId="ref60701" refString="Petrulevicius, J. F. (2019) A new Frenguelliidae (Insecta: Odonata) from the Eocene of Arroyo Chacay, Patagonia, Argentina. Palaeoentomology, 002, 591 - 595. https: // doi. org / 10.11646 / palaeoentomology. 2.6.9" type="book chapter" year="2019">2019</bibRefCitation>
). The wing of 
<emphasis id="B969EAC6000EFF8BFF7BFA2EFE4A17F0" box="[183,484,1483,1509]" italics="true" pageId="28" pageNumber="29">Treintamilun vuelvenlucha</emphasis>
Petrulevičius resembles those of the 
<taxonomicName id="4C1D4D57000EFF8BFC5CFA2EFB9D17F0" box="[912,1075,1483,1509]" class="Insecta" family="Dysagrionidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="28" pageNumber="29" phylum="Arthropoda" rank="family">Dysagrionidae</taxonomicName>
in many aspects, but differs by its vertical subnodus and—importantly—its curved CuP. Those of 
<emphasis id="B969EAC6000EFF8BFCB1FA0AFBA7141C" box="[893,1033,1519,1545]" italics="true" pageId="28" pageNumber="29">Nelala chori</emphasis>
Petrulevičius lack the characteristic 
<taxonomicName id="4C1D4D57000EFF8BFF5BF9F6FECD1438" box="[151,355,1555,1581]" kingdom="Animalia" pageId="28" pageNumber="27" rank="subOrder" subOrder="Cephalozygoptera">Cephalozygoptera</taxonomicName>
quadrangle shape. The suborder affinity of 
<taxonomicName id="4C1D4D57000EFF8BFC8BF9F6FC471438" authorityName="Petrulevicius &amp; Nel" authorityYear="2003" box="[839,1001,1555,1581]" class="Insecta" family="Frenguelliidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="28" pageNumber="29" phylum="Arthropoda" rank="family">Frenguelliidae</taxonomicName>
is unclear, and we follow 
<bibRefCitation id="EF8C4B25000EFF8BFAC6F9F6FF751444" author="Petrulevicius, J. F." pageId="28" pageNumber="29" pagination="367 - 374" refId="ref60663" refString="Petrulevicius, J. F. (2017) First Frenguelliidae (Insecta: Odonata) from the middle Eocene of Rio Pichileufu, Patagonia, Argen- tina. Arquivos Entomoloxicos, 18, 367 - 374." type="journal article" year="2017">Petrulevičius (2017</bibRefCitation>
, 
<bibRefCitation id="EF8C4B25000EFF8BFF24F9D2FE8C1444" author="Petrulevicius, J. F." box="[232,290,1591,1617]" pageId="28" pageNumber="29" pagination="591 - 595" refId="ref60701" refString="Petrulevicius, J. F. (2019) A new Frenguelliidae (Insecta: Odonata) from the Eocene of Arroyo Chacay, Patagonia, Argentina. Palaeoentomology, 002, 591 - 595. https: // doi. org / 10.11646 / palaeoentomology. 2.6.9" type="book chapter" year="2019">2019</bibRefCitation>
) and 
<bibRefCitation id="EF8C4B25000EFF8BFEAEF9D2FD3E1444" author="Petrulevicius, J. F. &amp; Nel, A." box="[354,656,1591,1617]" pageId="28" pageNumber="29" pagination="2909 - 2917" refId="ref60790" refString="Petrulevicius, J. F. &amp; Nel, A. (2003) Frenguelliidae, a new family of dragonflies from the earliest Eocene of Argentina (Insecta: Odonata): phylogenetic relationships within Odonata. Journal of Natural History, 37, 2909 - 2917. https: // doi. org / 10.1080 / 0022293021000007543" type="journal article" year="2003">Petrulevičius &amp; Nel (2003</bibRefCitation>
, 2007, 2013) in treating it as 
<taxonomicName id="4C1D4D57000EFF8BFC2CF9D2FB911444" box="[992,1087,1591,1617]" class="Insecta" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="28" pageNumber="29" phylum="Arthropoda" rank="order">Odonata</taxonomicName>
suborder indet., pending more complete specimens.
</paragraph>
<paragraph id="8BA236D4000EFF8BFF0BF99AFD891564" blockId="28.[151,1437,150,2013]" pageId="28" pageNumber="29">
The wings of 
<taxonomicName id="4C1D4D57000EFF8BFEA4F99AFD36148C" authority="(Statz)" baseAuthorityName="Statz" baseAuthorityYear="1930" box="[360,664,1663,1689]" class="Insecta" family="Sieblosiidae" genus="Oligolestes" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="28" pageNumber="29" phylum="Arthropoda" rank="species" species="grandis">
<emphasis id="B969EAC6000EFF8BFEA4F99AFDEB148C" box="[360,581,1663,1689]" italics="true" pageId="28" pageNumber="29">Oligolestes grandis</emphasis>
(Statz)
</taxonomicName>
from the Chattian of Rott, 
<collectingCountry id="F30A7644000EFF8BFC1FF99AFB95148C" box="[979,1083,1663,1689]" name="Germany" pageId="28" pageNumber="29">Germany</collectingCountry>
and 
<taxonomicName id="4C1D4D57000EFF8BFBB9F99AFF1A14A8" authority="Nel et al." authorityName="Nel" class="Insecta" genus="Italolestes" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="28" pageNumber="29" phylum="Arthropoda" rank="species" species="stroppai">
<emphasis id="B969EAC6000EFF8BFBB9F99AFAE3148D" box="[1141,1357,1663,1689]" italics="true" pageId="28" pageNumber="29">Italolestes stroppai</emphasis>
Nel 
<emphasis id="B969EAC6000EFF8BFA44F99AFF1A14A8" italics="true" pageId="28" pageNumber="29">et al.</emphasis>
</taxonomicName>
from the Messinian (late Miocene) of 
<collectingCountry id="F30A7644000EFF8BFDA6F946FD3314A8" box="[618,669,1699,1725]" name="Italy" pageId="28" pageNumber="29">Italy</collectingCountry>
(
<bibRefCitation id="EF8C4B25000EFF8BFD60F946FC1914A8" author="Gentilini, G. &amp; Bagli, L." box="[684,951,1699,1725]" pageId="28" pageNumber="29" pagination="17 - 44" refId="ref57986" refString="Gentilini, G. &amp; Bagli, L. (2004) Fossil Zygoptera and Anisoptera from the Upper Miocene of Monte Castellaro (Pesaro, Marches, Central Ital). Quaderno di Studi e Notizie di Storio Naturale dello Romagna, 19, 17 - 44." type="journal article" year="2004">Gentilini &amp; Bagli 2004</bibRefCitation>
; 
<bibRefCitation id="EF8C4B25000EFF8BFC0FF946FBD414A8" author="Nel, A. &amp; Petrulevicius, J. F. &amp; Gentilini, G. &amp; Martinez-Delclos, X." box="[963,1146,1698,1725]" pageId="28" pageNumber="29" pagination="219 - 233" refId="ref60383" refString="Nel, A., Petrulevicius, J. F., Gentilini, G. &amp; Martinez-Delclos, X. (2005 a) Phylogenetic analysis of the Cenozoic family Sieblo- siidae (Insecta: Odonata), with description of new taxa from Russia, Italy and France. Geobios, 38, 219 - 233. https: // doi. org / 10.1016 / j. geobios. 2003.10.007" type="journal article" year="2005">
Nel 
<emphasis id="B969EAC6000EFF8BFC3FF946FB8B14A8" box="[1011,1061,1699,1725]" italics="true" pageId="28" pageNumber="29">et al</emphasis>
. 2005 
<bibRefCitation id="EF8C4B25000EFF8BFBA2F947FBD414A8" author="Nel, A. &amp; Petrulevicius, J. F. &amp; Gentilini, G. &amp; Martinez-Delclos, X." box="[1134,1146,1698,1725]" pageId="28" pageNumber="29" pagination="188" refId="ref60098" refString="Nel, A., Petrulevicius, J. F., Gentilini, G. &amp; Martinez-Delclos, X. (2005 c) Un nouvel Odonate du Miocene d'Italie (Odon.). Bulletin de la Societe entomologique de France, 110, 188." type="journal article" year="2005">a</bibRefCitation>
</bibRefCitation>
, 2005c) share many traits with the 
<taxonomicName id="4C1D4D57000EFF8BFF30F922FE2B14F4" box="[252,389,1735,1761]" class="Insecta" family="Sieblosiidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="28" pageNumber="29" phylum="Arthropoda" rank="family">Sieblosiidae</taxonomicName>
(
<figureCitation id="13262A51000EFF8BFE5AF922FE4614F4" box="[406,488,1735,1761]" captionStart="FIGURE 13" captionStartId="30.[151,250,1454,1478]" captionTargetBox="[166,1415,565,1417]" captionTargetId="figure-181@30.[151,1436,561,1430]" captionTargetPageId="30" captionText="FIGURE 13. Drawings of wing of Oligolestes and wings of Italolestes stroppai. Oligolestes is redrawn from Schmidt (1985, Fig. 2), which appears accurate, cf. his photograph (his Fig. 1). Carpenter’s drawing (1992, Fig. 52.4) correctly reproduced this, but Statz’s drawing (1930, unnumbered figure) appears inaccurate in part and includes some presumptive morphology not on the fossil. These had no scale bar. Drawings of Italolestes stroppai wings are redrawn from Nel et al. (2005a, Figs. 2, 3, there called genus and species A), bottom wing portion reproduced to show nodal region morphology contrary to Sieblosiidae that is damaged in the more complete wing reproduced above it, scale bar is 5 mm. An asterisk indicates the oblique vein “O” immediately to its right; “s”, subnodus." figureDoi="http://doi.org/10.5281/zenodo.4558828" httpUri="https://zenodo.org/record/4558828/files/figure.png" pageId="28" pageNumber="29">Fig. 13</figureCitation>
) but differ in important ways, 
<emphasis id="B969EAC6000EFF8BFC85F922FCC414F5" box="[841,874,1735,1760]" italics="true" pageId="28" pageNumber="29">e.g</emphasis>
., the quadrangle of 
<taxonomicName id="4C1D4D57000EFF8BFB9CF922FA9A14F4" authority="Schmidt" authorityName="Schmidt" authorityYear="1958" box="[1104,1332,1735,1761]" class="Insecta" family="Sieblosiidae" genus="Oligolestes" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="28" pageNumber="29" phylum="Arthropoda" rank="genus">
<emphasis id="B969EAC6000EFF8BFB9CF922FB6014F4" box="[1104,1230,1735,1761]" italics="true" pageId="28" pageNumber="29">Oligolestes</emphasis>
Schmidt
</taxonomicName>
is somewhat narrower and the bases of RP3-4 and IR2 are in a distinctly different position. Both genera differ significantly from the 
<taxonomicName id="4C1D4D57000EFF8BFF31F8EAFE28153C" box="[253,390,1807,1833]" class="Insecta" family="Sieblosiidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="28" pageNumber="29" phylum="Arthropoda" rank="family">Sieblosiidae</taxonomicName>
in possessing a subnodus of normal obliquity and ScP not passing through the nodus (see 
<bibRefCitation id="EF8C4B25000EFF8BFAB8F8EAFE8F1558" author="Nel, A. &amp; Petrulevicius, J. F. &amp; Gentilini, G. &amp; Martinez-Delclos, X." pageId="28" pageNumber="29" pagination="219 - 233" refId="ref60383" refString="Nel, A., Petrulevicius, J. F., Gentilini, G. &amp; Martinez-Delclos, X. (2005 a) Phylogenetic analysis of the Cenozoic family Sieblo- siidae (Insecta: Odonata), with description of new taxa from Russia, Italy and France. Geobios, 38, 219 - 233. https: // doi. org / 10.1016 / j. geobios. 2003.10.007" type="journal article" year="2005">
Nel 
<emphasis id="B969EAC6000EFF8BFF5BF8D6FF671558" box="[151,201,1843,1869]" italics="true" pageId="28" pageNumber="29">et al</emphasis>
. 2005a
</bibRefCitation>
). We agree with 
<bibRefCitation id="EF8C4B25000EFF8BFE11F8D6FD0A1558" author="Nel, A. &amp; Petrulevicius, J. F. &amp; Gentilini, G. &amp; Martinez-Delclos, X." box="[477,676,1842,1869]" pageId="28" pageNumber="29" pagination="219 - 233" refId="ref60383" refString="Nel, A., Petrulevicius, J. F., Gentilini, G. &amp; Martinez-Delclos, X. (2005 a) Phylogenetic analysis of the Cenozoic family Sieblo- siidae (Insecta: Odonata), with description of new taxa from Russia, Italy and France. Geobios, 38, 219 - 233. https: // doi. org / 10.1016 / j. geobios. 2003.10.007" type="journal article" year="2005">
Nel 
<emphasis id="B969EAC6000EFF8BFDC1F8D6FD911558" box="[525,575,1843,1869]" italics="true" pageId="28" pageNumber="29">et al</emphasis>
. (2005a)
</bibRefCitation>
that they might be best treated as of uncertain position pending the discovery of more complete fossils.
</paragraph>
<paragraph id="8BA236D4000EFF8AFF0BF89EFE1B12EC" blockId="28.[151,1437,150,2013]" lastBlockId="29.[151,1437,151,537]" lastPageId="29" lastPageNumber="30" pageId="28" pageNumber="29">
<emphasis id="B969EAC6000EFF8BFF0BF89EFD271580" box="[199,649,1915,1941]" italics="true" pageId="28" pageNumber="29">
<taxonomicName id="4C1D4D57000EFF8BFF0BF89EFE3A1580" box="[199,404,1915,1941]" kingdom="Animalia" pageId="28" pageNumber="27" rank="subOrder" subOrder="Cephalozygoptera">Cephalozygoptera</taxonomicName>
terminal appendages
</emphasis>
. Only the male terminal appendages in 
<taxonomicName id="4C1D4D57000EFF8BFB80F89EFAB61580" box="[1100,1304,1915,1941]" kingdom="Animalia" pageId="28" pageNumber="27" rank="subOrder" subOrder="Cephalozygoptera">Cephalozygoptera</taxonomicName>
are known, unless those of the female are present on some 
<taxonomicName id="4C1D4D57000EFF8BFD64F87AFC9F15AC" box="[680,817,1951,1977]" class="Insecta" family="Sieblosiidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="28" pageNumber="29" phylum="Arthropoda" rank="family">Sieblosiidae</taxonomicName>
fossils of which we are not aware. The well-developed cerci and paraprocts at the apex of the abdomen are used in extant 
<taxonomicName id="4C1D4D57000EFF8BFCB2F826FC5F15C8" box="[894,1009,1987,2013]" class="Insecta" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="28" pageNumber="124" phylum="Arthropoda" rank="subOrder" subOrder="Zygoptera">Zygoptera</taxonomicName>
to grasp the female during mating. As the terminal appendages of male 
<taxonomicName id="4C1D4D57000FFF8AFDC9FF72FD7F12A4" box="[517,721,151,177]" kingdom="Animalia" pageId="29" pageNumber="27" rank="subOrder" subOrder="Cephalozygoptera">Cephalozygoptera</taxonomicName>
are so like those of 
<taxonomicName id="4C1D4D57000FFF8AFC61FF72FB8D12A4" box="[941,1059,151,177]" class="Insecta" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="29" pageNumber="124" phylum="Arthropoda" rank="subOrder" subOrder="Zygoptera">Zygoptera</taxonomicName>
, we predict that their female genitalia will be found to be correspondingly similar. We are not aware of any 
<taxonomicName id="4C1D4D57000FFF8AFC24FF5EFB1A12C0" box="[1000,1204,187,213]" kingdom="Animalia" pageId="29" pageNumber="27" rank="subOrder" subOrder="Cephalozygoptera">Cephalozygoptera</taxonomicName>
fossil that preserves secondary male genitalia.
</paragraph>
<paragraph id="8BA236D4000FFF8AFF0BFEE7FD88100C" blockId="29.[151,1437,151,537]" pageId="29" pageNumber="30">
<emphasis id="B969EAC6000FFF8AFF0BFEE7FE0D1308" bold="true" box="[199,419,258,285]" pageId="29" pageNumber="30">Cladistic analyses.</emphasis>
Searches performed using morphological data (Appendix C) recovered a single most-parsimonious tree (MPT). Analysis 1, of the relationships of the suborders of 
<taxonomicName id="4C1D4D57000FFF8AFC34FEC2FB651354" authority=", Burmese" authorityName="Burmese" box="[1016,1227,295,321]" class="Insecta" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="29" pageNumber="30" phylum="Arthropoda" rank="order">Odonata, Burmese</taxonomicName>
amber genera not assigned to suborder, and 
<taxonomicName id="4C1D4D57000FFF8AFE71FEAEFDDC1370" box="[445,626,331,357]" class="Insecta" family="Tarsophlebiidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="29" pageNumber="30" phylum="Arthropoda" rank="family">Tarsophlebiidae</taxonomicName>
, produced one MPT (
<figureCitation id="13262A51000FFF8AFCA5FEAEFC131370" box="[873,957,331,357]" captionStart="FIGURE 14" captionStartId="35.[151,250,1193,1217]" captionTargetBox="[156,1427,245,1166]" captionTargetId="figure-47@35.[151,1436,237,1170]" captionTargetPageId="35" captionText="FIGURE 14. Tree resulting from parsimony-based analyses for Analysis 1: Tarsophlebiidae, the suborders of Odonata and former members of Dysagrionidae (Burmadysagrion, Electrodysagrion, Paleodysagrion). One single most-parsimonious tree (MPT) based on 23 continuous and 79 discrete characters. Symmetrical resampling values Ξ70 are labelled at the nodes." figureDoi="http://doi.org/10.5281/zenodo.4558832" httpUri="https://zenodo.org/record/4558832/files/figure.png" pageId="29" pageNumber="30">Fig. 14</figureCitation>
). Constraining 
<taxonomicName id="4C1D4D57000FFF8AFBA7FEAEFAB31370" box="[1131,1309,331,357]" class="Insecta" family="Tarsophlebiidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="29" pageNumber="30" phylum="Arthropoda" rank="family">Tarsophlebiidae</taxonomicName>
as the outgroup recovers 
<taxonomicName id="4C1D4D57000FFF8AFE8DFE8AFE12139C" box="[321,444,367,393]" class="Insecta" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="29" pageNumber="4" phylum="Arthropoda" rank="subOrder" subOrder="Anisoptera">Anisoptera</taxonomicName>
and 
<taxonomicName id="4C1D4D57000FFF8AFE22FE8AFD31139C" authorityName="Handlirsch" authorityYear="1906" box="[494,671,367,393]" class="Insecta" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="29" pageNumber="124" phylum="Arthropoda" rank="subOrder" subOrder="Anisozygoptera">Anisozygoptera</taxonomicName>
as sister taxa with high support, consistent with other recent analyses (
<bibRefCitation id="EF8C4B25000FFF8AFF52FE76FED613B8" author="Westfall, M. J. &amp; May, M. L. &amp; Needham, J. G." box="[158,376,403,429]" pageId="29" pageNumber="30" refId="ref62218" refString="Westfall, M. J., May, M. L. &amp; Needham, J. G. (2014) Dragonflies of North America: The Odonata (Anisoptera) Fauna of Canada, the Continental United States, Northern Mexico and the Greater Antilles. Scientific Publishers, Gainesville, Florida, xiv + 648 pp." type="book" year="2014">
Westfall 
<emphasis id="B969EAC6000FFF8AFECFFE76FE9D13B8" box="[259,307,403,429]" italics="true" pageId="29" pageNumber="30">et al</emphasis>
. 2014
</bibRefCitation>
and references therein). 
<taxonomicName id="4C1D4D57000FFF8AFD46FE76FCF813B8" box="[650,854,403,429]" kingdom="Animalia" pageId="29" pageNumber="27" rank="subOrder" subOrder="Cephalozygoptera">Cephalozygoptera</taxonomicName>
and 
<taxonomicName id="4C1D4D57000FFF8AFC46FE76FC5313B8" box="[906,1021,403,429]" class="Insecta" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="29" pageNumber="124" phylum="Arthropoda" rank="subOrder" subOrder="Zygoptera">Zygoptera</taxonomicName>
are recovered as sister taxa with high support, while 
<taxonomicName id="4C1D4D57000FFF8AFEF2FE52FE5413C4" box="[318,506,439,465]" class="Insecta" family="Dysagrionidae" genus="Burmadysagrion" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="29" pageNumber="30" phylum="Arthropoda" rank="genus">
<emphasis id="B969EAC6000FFF8AFEF2FE52FE5413C4" box="[318,506,439,465]" italics="true" pageId="29" pageNumber="30">Burmadysagrion</emphasis>
</taxonomicName>
, 
<taxonomicName id="4C1D4D57000FFF8AFDC5FE52FD6413C4" box="[521,714,439,465]" class="Insecta" family="Dysagrionidae" genus="Electrodysagrion" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="29" pageNumber="30" phylum="Arthropoda" rank="genus">
<emphasis id="B969EAC6000FFF8AFDC5FE52FD6413C4" box="[521,714,439,465]" italics="true" pageId="29" pageNumber="30">Electrodysagrion</emphasis>
</taxonomicName>
, and 
<emphasis id="B969EAC6000FFF8AFCCBFE52FC1613C4" box="[775,952,439,465]" italics="true" pageId="29" pageNumber="30">Paleodysagrion</emphasis>
, formerly considered to belong to the 
<taxonomicName id="4C1D4D57000FFF8AFAAAFE52FEBE13E0" class="Insecta" family="Dysagrionidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="29" pageNumber="30" phylum="Arthropoda" rank="family">Dysagrionidae</taxonomicName>
, are recovered as members of 
<taxonomicName id="4C1D4D57000FFF8AFDAAFE3EFD7713E0" box="[614,729,475,501]" class="Insecta" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="29" pageNumber="124" phylum="Arthropoda" rank="subOrder" subOrder="Zygoptera">Zygoptera</taxonomicName>
with moderate to high support. 
<taxonomicName id="4C1D4D57000FFF8AFB8DFE3EFB1913E0" box="[1089,1207,475,501]" class="Insecta" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="29" pageNumber="124" phylum="Arthropoda" rank="subOrder" subOrder="Zygoptera">Zygoptera</taxonomicName>
, therefore, was used as an outgroup in Analyses 2 and 3.
</paragraph>
<caption id="DF62665C000FFF8AFF5BF892FEBE15EF" ID-DOI="http://doi.org/10.5281/zenodo.4558826" ID-Zenodo-Dep="4558826" httpUri="https://zenodo.org/record/4558826/files/figure.png" pageId="29" pageNumber="30" startId="29.[151,250,1911,1935]" targetBox="[154,1432,569,1875]" targetPageId="29">
<paragraph id="8BA236D4000FFF8AFF5BF892FEBE15EF" blockId="29.[151,1437,1911,2043]" pageId="29" pageNumber="30">
<emphasis id="B969EAC6000FFF8AFF5BF892FE8E159A" bold="true" box="[151,288,1911,1935]" pageId="29" pageNumber="30">FIGURE 12.</emphasis>
Representative 
<taxonomicName id="4C1D4D57000FFF8AFE09F892FDED159A" box="[453,579,1911,1935]" class="Insecta" family="Sieblosiidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="29" pageNumber="30" phylum="Arthropoda" rank="family">Sieblosiidae</taxonomicName>
, A, 
<taxonomicName id="4C1D4D57000FFF8AFDA1F892FC34159B" authority="Nel et al." authorityName="Nel" box="[621,922,1911,1935]" class="Insecta" family="Sieblosiidae" genus="Stenolestes" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="29" pageNumber="30" phylum="Arthropoda" rank="species" species="fasciata">
<emphasis id="B969EAC6000FFF8AFDA1F892FC9A159A" box="[621,820,1911,1935]" italics="true" pageId="29" pageNumber="30">Stenolestes fasciata</emphasis>
Nel 
<emphasis id="B969EAC6000FFF8AFCABF89DFC3A159A" box="[871,916,1911,1935]" italics="true" pageId="29" pageNumber="30">et al</emphasis>
.
</taxonomicName>
; B, 
<taxonomicName id="4C1D4D57000FFF8AFC0BF892FB64159A" box="[967,1226,1911,1935]" class="Insecta" family="Sieblosiidae" genus="Miostenolestes" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="29" pageNumber="30" phylum="Arthropoda" rank="species" species="zherikhini">
<emphasis id="B969EAC6000FFF8AFC0BF892FB64159A" box="[967,1226,1911,1935]" italics="true" pageId="29" pageNumber="30">Miostenolestes zherikhini</emphasis>
</taxonomicName>
; and 
<taxonomicName id="4C1D4D57000FFF8AFAC8F892FA37159A" authorityName="Petrulevicius &amp; Nel" authorityYear="2003" box="[1284,1433,1911,1935]" class="Insecta" family="Frenguelliidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="29" pageNumber="30" phylum="Arthropoda" rank="family">Frenguelliidae</taxonomicName>
, C, 
<emphasis id="B969EAC6000FFF8AFF74F87EFE6715A6" box="[184,457,1947,1971]" italics="true" pageId="29" pageNumber="30">Treintamilun vuelvenlucha</emphasis>
; D, 
<taxonomicName id="4C1D4D57000FFF8AFE36F87EFB8015A6" authority="Petrulevicius &amp; Nel. Redrawn" authorityName="Petrulevicius &amp; Nel. Redrawn" authorityYear="2003" box="[506,1070,1947,1971]" class="Insecta" family="Frenguelliidae" genus="Frenguellia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="29" pageNumber="30" phylum="Arthropoda" rank="species" species="patagonica">
<emphasis id="B969EAC6000FFF8AFE36F87EFD4215A6" box="[506,748,1947,1971]" italics="true" pageId="29" pageNumber="30">Frenguellia patagonica</emphasis>
Petrulevičius &amp; Nel. Redrawn
</taxonomicName>
from: A, B, 
<bibRefCitation id="EF8C4B25000FFF8AFB76F87EFAD915A6" author="Nel, A. &amp; Petrulevicius, J. F. &amp; Gentilini, G. &amp; Martinez-Delclos, X." box="[1210,1399,1947,1971]" pageId="29" pageNumber="30" pagination="219 - 233" refId="ref60383" refString="Nel, A., Petrulevicius, J. F., Gentilini, G. &amp; Martinez-Delclos, X. (2005 a) Phylogenetic analysis of the Cenozoic family Sieblo- siidae (Insecta: Odonata), with description of new taxa from Russia, Italy and France. Geobios, 38, 219 - 233. https: // doi. org / 10.1016 / j. geobios. 2003.10.007" type="journal article" year="2005">
Nel 
<emphasis id="B969EAC6000FFF8AFB24F879FAB915A6" box="[1256,1303,1947,1971]" italics="true" pageId="29" pageNumber="30">et al</emphasis>
. (2005a)
</bibRefCitation>
; C, 
<bibRefCitation id="EF8C4B25000FFF8AFF5BF85AFEC615C2" author="Petrulevicius, J. F." box="[151,360,1983,2007]" pageId="29" pageNumber="30" pagination="367 - 374" refId="ref60663" refString="Petrulevicius, J. F. (2017) First Frenguelliidae (Insecta: Odonata) from the middle Eocene of Rio Pichileufu, Patagonia, Argen- tina. Arquivos Entomoloxicos, 18, 367 - 374." type="journal article" year="2017">Petrulevičius (2017)</bibRefCitation>
; D, Petrulevičius &amp; Nel (2007). An asterisk indicates the oblique vein “O” immediately to its right. Scale bars, 5 mm.
</paragraph>
</caption>
<paragraph id="8BA236D4000CFF89FF0BFF72FB7512C0" blockId="30.[151,1437,151,538]" pageId="30" pageNumber="31">
In Analysis 2, the relationship of 
<taxonomicName id="4C1D4D57000CFF89FD87FF72FCBC12A4" authority="Archibald &amp; Cannings &amp; Erickson &amp; Bybee &amp; Mathewes, 2021" authorityName="Archibald &amp; Cannings &amp; Erickson &amp; Bybee &amp; Mathewes" authorityYear="2021" box="[587,786,151,177]" family="Whetwhetaksidae" kingdom="Animalia" order="Odonata" pageId="30" pageNumber="28" rank="family" status="fam. nov.">Whetwhetaksidae</taxonomicName>
to 
<taxonomicName id="4C1D4D57000CFF89FCF7FF72FBA912A4" box="[827,1031,151,177]" kingdom="Animalia" pageId="30" pageNumber="27" rank="subOrder" subOrder="Cephalozygoptera">Cephalozygoptera</taxonomicName>
and its families, produced a single MPT (
<figureCitation id="13262A51000CFF89FF2CFF5EFE9F12C0" box="[224,305,186,213]" captionStart="FIGURE 15" captionStartId="35.[151,250,1929,1953]" captionTargetBox="[161,1428,1323,1898]" captionTargetId="figure-102@35.[151,1436,1314,1906]" captionTargetPageId="35" captionText="FIGURE 15. Tree resulting from parsimony-based analyses for Analysis 2 (relationship Whetwhetaksidae to confirmed families of Cephalozygoptera). One single most-parsimonious tree (MPT) based on 23 continuous and 79 discrete characters. Symmetrical resampling values Ξ70 are labeled at the nodes." figureDoi="http://doi.org/10.5281/zenodo.4558834" httpUri="https://zenodo.org/record/4558834/files/figure.png" pageId="30" pageNumber="31">Fig. 15</figureCitation>
) that provides weak support for 
<taxonomicName id="4C1D4D57000CFF89FD51FF5EFCEE12C0" box="[669,832,187,213]" class="Insecta" family="Dysagrionidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="30" pageNumber="31" phylum="Arthropoda" rank="family">Dysagrionidae</taxonomicName>
and 
<taxonomicName id="4C1D4D57000CFF89FCBBFF5EFB9012C0" authority="Archibald &amp; Cannings &amp; Erickson &amp; Bybee &amp; Mathewes, 2021" authorityName="Archibald &amp; Cannings &amp; Erickson &amp; Bybee &amp; Mathewes" authorityYear="2021" box="[887,1086,187,213]" family="Whetwhetaksidae" kingdom="Animalia" order="Odonata" pageId="30" pageNumber="28" rank="family" status="fam. nov.">Whetwhetaksidae</taxonomicName>
as sister taxa.
</paragraph>
<paragraph id="8BA236D4000CFF89FF0BFF3AFC5B12EC" blockId="30.[151,1437,151,538]" box="[199,1013,223,249]" pageId="30" pageNumber="31">
Analysis 3, the internal phylogeny of 
<taxonomicName id="4C1D4D57000CFF89FDA1FF3AFCBD12EC" box="[621,787,223,249]" class="Insecta" family="Dysagrionidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="30" pageNumber="31" phylum="Arthropoda" rank="family">Dysagrionidae</taxonomicName>
, is discussed below.
</paragraph>
<paragraph id="8BA236D4000CFF89FF0BFEE7FD30100C" blockId="30.[151,1437,151,538]" pageId="30" pageNumber="31">
<emphasis id="B969EAC6000CFF89FF0BFEE7FCEA1309" bold="true" box="[199,836,258,285]" pageId="30" pageNumber="31">
History and biogeography of the 
<taxonomicName id="4C1D4D57000CFF89FDA8FEE7FCEE1309" box="[612,832,258,284]" kingdom="Animalia" pageId="30" pageNumber="27" rank="subOrder" subOrder="Cephalozygoptera">Cephalozygoptera</taxonomicName>
.
</emphasis>
The 
<taxonomicName id="4C1D4D57000CFF89FC49FEE6FBFF1308" box="[901,1105,259,285]" kingdom="Animalia" pageId="30" pageNumber="27" rank="subOrder" subOrder="Cephalozygoptera">Cephalozygoptera</taxonomicName>
has been found in localities across the Northern Hemisphere (
<tableCitation id="C69F036F000CFF89FDD4FEC2FDDE1354" box="[536,624,295,321]" captionStart="TABLE 3" captionStartId="31.[151,239,944,968]" captionTargetBox="[166,1139,985,1975]" captionTargetPageId="31" captionText="TABLE 3. Cephalozygoptera and tentative Cephalozygoptera occurrences." httpUri="http://table.plazi.org/id/DF62665C000DFF88FF5BFC55FC1711DD" pageId="30" pageNumber="31" tableUuid="DF62665C000DFF88FF5BFC55FC1711DD">Table 3</tableCitation>
) from the Aptian of 
<collectingCountry id="F30A7644000CFF89FCAEFEC2FC061354" box="[866,936,295,321]" name="China" pageId="30" pageNumber="31">China</collectingCountry>
; possibly the Thanetian of 
<collectingCountry id="F30A7644000CFF89FB2EFEC2FA801354" box="[1250,1326,295,321]" name="France" pageId="30" pageNumber="31">France</collectingCountry>
(
<taxonomicName id="4C1D4D57000CFF89FA8DFEC2FA3B1354" box="[1345,1429,295,321]" class="Insecta" genus="Valerea" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="30" pageNumber="31" phylum="Arthropoda" rank="genus">
<emphasis id="B969EAC6000CFF89FA8DFEC2FA3B1354" box="[1345,1429,295,321]" italics="true" pageId="30" pageNumber="31">Valerea</emphasis>
</taxonomicName>
, 
<taxonomicName id="4C1D4D57000CFF89FF5BFEAEFEF51370" box="[151,347,331,357]" class="Insecta" genus="Thanetophilosina" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="30" pageNumber="31" phylum="Arthropoda" rank="genus">
<emphasis id="B969EAC6000CFF89FF5BFEAEFEF51370" box="[151,347,331,357]" italics="true" pageId="30" pageNumber="31">Thanetophilosina</emphasis>
</taxonomicName>
) and the Thanetian or Ypresian of 
<collectingRegion id="49D9F836000CFF89FD3BFEAEFCE61370" box="[759,840,331,357]" country="United States of America" name="Alaska" pageId="30" pageNumber="31">Alaska</collectingRegion>
, 
<collectingCountry id="F30A7644000CFF89FC9BFEAEFBD21370" box="[855,1148,331,357]" name="United States of America" pageId="30" pageNumber="31">United States of America</collectingCountry>
(
<bibRefCitation id="EF8C4B25000CFF89FB42FEAEFA361370" author="Garrouste, R. &amp; Nel, A." box="[1166,1432,331,357]" pageId="30" pageNumber="31" pagination="1939 - 1946" refId="ref57867" refString="Garrouste, R. &amp; Nel, A. (2019) Alaskan Palaeogene insects: a challenge for a better knowledge of the Beringian ' route' (Odonata: Aeshnidae, Dysagrionidae). Journal of Systematic Palaeontology, 17, 1939 - 1946. https: // doi. org / 10.1080 / 14772019.2019.1572235" type="journal article" year="2019">Garrouste &amp; Nel 2019</bibRefCitation>
: unnamed); the Ypresian of 
<collectingCountry id="F30A7644000CFF89FE0BFE8AFD9C139C" box="[455,562,367,393]" name="Denmark" pageId="30" pageNumber="31">Denmark</collectingCountry>
, 
<collectingCountry id="F30A7644000CFF89FDF2FE8AFD3B139C" box="[574,661,367,393]" name="Canada" pageId="30" pageNumber="31">Canada</collectingCountry>
, and the 
<collectingCountry id="F30A7644000CFF89FD36FE8AFBB7139C" box="[762,1049,367,393]" name="United States of America" pageId="30" pageNumber="31">United States of America</collectingCountry>
; the Lutetian of 
<collectingCountry id="F30A7644000CFF89FB1DFE8AFA93139C" box="[1233,1341,367,393]" name="Germany" pageId="30" pageNumber="31">Germany</collectingCountry>
; the Priabonian of 
<collectingRegion id="49D9F836000CFF89FED8FE76FED113B8" box="[276,383,403,429]" country="United States of America" name="Colorado" pageId="30" pageNumber="31">Colorado</collectingRegion>
, 
<collectingCountry id="F30A7644000CFF89FE45FE76FD0A13B8" box="[393,676,403,429]" name="United States of America" pageId="30" pageNumber="31">United States of America</collectingCountry>
, the 
<collectingCountry id="F30A7644000CFF89FD1BFE76FC3913B8" box="[727,919,403,429]" name="United Kingdom" pageId="30" pageNumber="31">United Kingdom</collectingCountry>
, and 
<collectingCountry id="F30A7644000CFF89FC1CFE76FBB513B8" box="[976,1051,403,429]" name="Russia" pageId="30" pageNumber="31">Russia</collectingCountry>
(European and Pacific); the Rupelian (early Oligocene) of 
<collectingCountry id="F30A7644000CFF89FE7DFE52FE5213C4" box="[433,508,439,465]" name="Russia" pageId="30" pageNumber="31">Russia</collectingCountry>
(Caucasus) and 
<collectingCountry id="F30A7644000CFF89FD78FE52FC8E13C4" box="[692,800,439,465]" name="Germany" pageId="30" pageNumber="31">Germany</collectingCountry>
; Chattian (late Oligocene) of 
<collectingCountry id="F30A7644000CFF89FBA6FE52FB1413C4" box="[1130,1210,439,465]" name="France" pageId="30" pageNumber="31">France</collectingCountry>
; Langhian/Serravallian (middle Miocene) of 
<collectingCountry id="F30A7644000CFF89FE7AFE3EFDAF13E0" box="[438,513,475,501]" name="Russia" pageId="30" pageNumber="31">Russia</collectingCountry>
(Caucasus) and 
<collectingCountry id="F30A7644000CFF89FD75FE3EFCB313E0" box="[697,797,475,501]" name="Bulgaria" pageId="30" pageNumber="31">Bulgaria</collectingCountry>
; Serravallian (middle Miocene) of 
<collectingCountry id="F30A7644000CFF89FB68FE3EFB5813E0" box="[1188,1270,475,501]" name="Croatia" pageId="30" pageNumber="31">Croatia</collectingCountry>
and 
<collectingCountry id="F30A7644000CFF89FAE0FE3EFA3613E0" box="[1324,1432,475,501]" name="Germany" pageId="30" pageNumber="31">Germany</collectingCountry>
; Tortonian (late Miocene) of 
<collectingCountry id="F30A7644000CFF89FE18FE1AFDBA100C" box="[468,532,511,537]" name="Spain" pageId="30" pageNumber="31">Spain</collectingCountry>
and 
<collectingCountry id="F30A7644000CFF89FD86FE1AFD34100C" box="[586,666,511,537]" name="France" pageId="30" pageNumber="31">France</collectingCountry>
.
</paragraph>
<caption id="DF62665C000CFF89FF5BFA4BFDA5148B" ID-DOI="http://doi.org/10.5281/zenodo.4558828" ID-Zenodo-Dep="4558828" httpUri="https://zenodo.org/record/4558828/files/figure.png" pageId="30" pageNumber="31" startId="30.[151,250,1454,1478]" targetBox="[166,1415,565,1417]" targetPageId="30">
<paragraph id="8BA236D4000CFF89FF5BFA4BFDA5148B" blockId="30.[151,1437,1454,1694]" pageId="30" pageNumber="31">
<emphasis id="B969EAC6000CFF89FF5BFA4BFE8F17D3" bold="true" box="[151,289,1454,1478]" pageId="30" pageNumber="31">FIGURE 13.</emphasis>
Drawings of wing of 
<taxonomicName id="4C1D4D57000CFF89FDC9FA4BFDD917D3" authorityName="Schmidt" authorityYear="1958" box="[517,631,1454,1478]" class="Insecta" family="Sieblosiidae" genus="Oligolestes" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="30" pageNumber="31" phylum="Arthropoda" rank="genus">
<emphasis id="B969EAC6000CFF89FDC9FA4BFDD917D3" box="[517,631,1454,1478]" italics="true" pageId="30" pageNumber="31">Oligolestes</emphasis>
</taxonomicName>
and wings of 
<taxonomicName id="4C1D4D57000CFF89FCC6FA4BFC6317D3" box="[778,973,1454,1478]" class="Insecta" genus="Italolestes" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="30" pageNumber="31" phylum="Arthropoda" rank="species" species="stroppai">
<emphasis id="B969EAC6000CFF89FCC6FA4BFC6317D3" box="[778,973,1454,1478]" italics="true" pageId="30" pageNumber="31">Italolestes stroppai</emphasis>
</taxonomicName>
. 
<taxonomicName id="4C1D4D57000CFF89FC17FA4BFBE317D3" authorityName="Schmidt" authorityYear="1958" box="[987,1101,1454,1478]" class="Insecta" family="Sieblosiidae" genus="Oligolestes" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="30" pageNumber="31" phylum="Arthropoda" rank="genus">
<emphasis id="B969EAC6000CFF89FC17FA4BFBE317D3" box="[987,1101,1454,1478]" italics="true" pageId="30" pageNumber="31">Oligolestes</emphasis>
</taxonomicName>
is redrawn from Schmidt (1985, Fig. 2), which appears accurate, 
<emphasis id="B969EAC6000CFF89FE12FA37FE5E17FF" box="[478,496,1490,1514]" italics="true" pageId="30" pageNumber="31">cf</emphasis>
. his photograph (his Fig. 1). Carpenter’s drawing (1992, Fig. 52.4) correctly reproduced this, but Statz’s drawing (1930, unnumbered figure) appears inaccurate in part and includes some presumptive morphology not on the fossil. These had no scale bar. Drawings of 
<taxonomicName id="4C1D4D57000CFF89FDB1F9FFFCEE1427" box="[637,832,1562,1586]" class="Insecta" genus="Italolestes" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="30" pageNumber="31" phylum="Arthropoda" rank="species" species="stroppai">
<emphasis id="B969EAC6000CFF89FDB1F9FFFCEE1427" box="[637,832,1562,1586]" italics="true" pageId="30" pageNumber="31">Italolestes stroppai</emphasis>
</taxonomicName>
wings are redrawn from 
<bibRefCitation id="EF8C4B25000CFF89FB8EF9FFFB5F1427" author="Nel, A. &amp; Petrulevicius, J. F. &amp; Gentilini, G. &amp; Martinez-Delclos, X." box="[1090,1265,1562,1586]" pageId="30" pageNumber="31" pagination="219 - 233" refId="ref60383" refString="Nel, A., Petrulevicius, J. F., Gentilini, G. &amp; Martinez-Delclos, X. (2005 a) Phylogenetic analysis of the Cenozoic family Sieblo- siidae (Insecta: Odonata), with description of new taxa from Russia, Italy and France. Geobios, 38, 219 - 233. https: // doi. org / 10.1016 / j. geobios. 2003.10.007" type="journal article" year="2005">
Nel 
<emphasis id="B969EAC6000CFF89FBA2F9FEFB351427" box="[1134,1179,1562,1586]" italics="true" pageId="30" pageNumber="31">et al</emphasis>
. (2005a
</bibRefCitation>
, Figs. 2, 3, there called genus and species A), bottom wing portion reproduced to show nodal region morphology contrary to 
<taxonomicName id="4C1D4D57000CFF89FB3FF9DBFAC11443" box="[1267,1391,1598,1622]" class="Insecta" family="Sieblosiidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="30" pageNumber="31" phylum="Arthropoda" rank="family">Sieblosiidae</taxonomicName>
that is damaged in the more complete wing reproduced above it, scale bar is 5 mm. An asterisk indicates the oblique vein “O” immediately to its right; “s”, subnodus.
</paragraph>
</caption>
<paragraph id="8BA236D4000CFF89FF0BF929FBB9153B" blockId="30.[151,1437,1740,2019]" pageId="30" pageNumber="31">
Bechly (pers. comm.) reports a specimen from the Ypresian Green River Formation that bears the distinctive 
<taxonomicName id="4C1D4D57000CFF89FF5BF915FECD151F" box="[151,355,1776,1802]" kingdom="Animalia" pageId="30" pageNumber="27" rank="subOrder" subOrder="Cephalozygoptera">Cephalozygoptera</taxonomicName>
head morphology, but is in ways quite different from the families that we include, suggesting that the suborder may have been larger and more diverse than we understand it here.
</paragraph>
<paragraph id="8BA236D4000CFF88FF0BF8DDFD44130B" blockId="30.[151,1437,1740,2019]" lastBlockId="31.[151,1437,151,897]" lastPageId="31" lastPageNumber="32" pageId="30" pageNumber="31">
The 
<taxonomicName id="4C1D4D57000CFF89FF37F8DDFE301547" box="[251,414,1848,1874]" class="Insecta" family="Dysagrionidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="30" pageNumber="31" phylum="Arthropoda" rank="family">Dysagrionidae</taxonomicName>
dominanted the Eocene 
<taxonomicName id="4C1D4D57000CFF89FD74F8DDFC2A1547" box="[696,900,1848,1874]" kingdom="Animalia" pageId="30" pageNumber="27" rank="subOrder" subOrder="Cephalozygoptera">Cephalozygoptera</taxonomicName>
across the Holarctic (see 
<tableCitation id="C69F036F000CFF89FB66F8DDFB501547" box="[1194,1278,1848,1874]" captionStart="TABLE 3" captionStartId="31.[151,239,944,968]" captionTargetBox="[166,1139,985,1975]" captionTargetPageId="31" captionText="TABLE 3. Cephalozygoptera and tentative Cephalozygoptera occurrences." httpUri="http://table.plazi.org/id/DF62665C000DFF88FF5BFC55FC1711DD" pageId="30" pageNumber="31" tableUuid="DF62665C000DFF88FF5BFC55FC1711DD">Table 3</tableCitation>
and references therein). They were by far the dominant 
<taxonomicName id="4C1D4D57000CFF89FD41F8B9FD421563" box="[653,748,1884,1910]" class="Insecta" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="30" pageNumber="31" phylum="Arthropoda" rank="order">Odonata</taxonomicName>
in more temperate far-western montane North America, the Okanagan Highlands (in upper microthermal mean annual temperatures, 
<emphasis id="B969EAC6000CFF89FC0BF864FC44158F" box="[967,1002,1921,1946]" italics="true" pageId="30" pageNumber="31">i.e.</emphasis>
13°C and below: 
<bibRefCitation id="EF8C4B25000CFF89FB7FF865FA39158F" author="Wolfe, J. A. &amp; Wehr, W. C." box="[1203,1431,1920,1946]" pageId="30" pageNumber="31" pagination="1 - 25" refId="ref62471" refString="Wolfe, J. A. &amp; Wehr, W. C. (1987) Middle Eocene Dicotyledonous plants from Republic, northeast Washington. United States Geological Survey Bulletin, 1597, 1 - 25, 16 pls." type="journal article" year="1987">Wolfe &amp; Wehr 1987</bibRefCitation>
; 
<bibRefCitation id="EF8C4B25000CFF89FF5BF841FE1415AB" author="Archibald, S. B. &amp; Farrell, B. D." box="[151,442,1956,1982]" pageId="30" pageNumber="31" pagination="17 - 23" refId="ref55625" refString="Archibald, S. B. &amp; Farrell, B. D. (2003) Wheeler's dilemma. Proceedings of the Second Paleoentomological Congress, Acta Zoologica Crakoviensia, 46 (Supplement), Fossil Insects, 17 - 23." type="book chapter" year="2003">Archibald &amp; Farrell 2003</bibRefCitation>
; 
<bibRefCitation id="EF8C4B25000CFF89FE0BF841FD6315AA" author="Greenwood, D. R. &amp; Archibald, S. B. &amp; Mathewes, R. W. &amp; Moss, P. T." box="[455,717,1956,1983]" pageId="30" pageNumber="31" pagination="167 - 185" refId="ref58279" refString="Greenwood, D. R., Archibald, S. B., Mathewes, R. W. &amp; Moss, P. T. (2005) Fossil biotas from the Okanagan Highlands, southern British Columbia and northern Washington State: climates and ecosystems across an Eocene landscape. Canadian Journal of Earth Sciences, 42, 167 - 185. https: // doi. org / 10.1139 / e 04 - 100" type="journal article" year="2005">
Greenwood 
<emphasis id="B969EAC6000CFF89FD9EF840FD2A15AB" box="[594,644,1956,1982]" italics="true" pageId="30" pageNumber="31">et al</emphasis>
. 2005
</bibRefCitation>
; 
<bibRefCitation id="EF8C4B25000CFF89FD15F841FC6315AB" author="Archibald, S. B. &amp; Morse, G. E. &amp; Greenwood, D. R. &amp; Mathewes, R. W." box="[729,973,1956,1983]" pageId="30" pageNumber="31" pagination="8095 - 8100" refId="ref55931" refString="Archibald, S. B., Morse, G. E., Greenwood, D. R. &amp; Mathewes, R. W. (2014) Fossil palm beetles refine upland winter tempera- tures in the Early Eocene Climatic Optimum. Proceedings of the National Academy of Sciences of the United States of America, 111, 8095 - 8100. https: // doi. org / 10.1073 / pnas. 1323269111" type="journal article" year="2014">
Archibald 
<emphasis id="B969EAC6000CFF89FC9DF840FC2D15AB" box="[849,899,1956,1982]" italics="true" pageId="30" pageNumber="31">et al</emphasis>
. 2014
</bibRefCitation>
), with 70 fossils in seventeen species assigned to four genera and one genus-level parataxon genus, contrasted with seven specimens of the single species of 
<taxonomicName id="4C1D4D57000DFF88FF5BFF72FECC12A4" authority="Archibald &amp; Cannings &amp; Erickson &amp; Bybee &amp; Mathewes, 2021" authorityName="Archibald &amp; Cannings &amp; Erickson &amp; Bybee &amp; Mathewes" authorityYear="2021" box="[151,354,151,177]" family="Whetwhetaksidae" kingdom="Animalia" order="Odonata" pageId="31" pageNumber="28" rank="family" status="fam. nov.">Whetwhetaksidae</taxonomicName>
; nine fossil dragonflies there in two families, seven genera and eight species; and one undescribed fossil damselfly wing. It is unclear if the larger wings of dragonflies depress their fossil abundance relative to their community abundance by taphonomic bias through increased floating time before deposition on the substrate (
<bibRefCitation id="EF8C4B25000DFF88FAAFFF3AFE541308" author="Martinez-Delclos, X. &amp; Martinell, J." pageId="31" pageNumber="32" pagination="133 - 144" refId="ref59251" refString="Martinez-Delclos, X. &amp; Martinell, J. (1993) Insect taphonomy experiments. Their application to the Cretaceous outcrops of lithographic limestones from Spain. Kaupia. Darmstadter Beitrage zur Naturgeschichte, 2, 133 - 144." type="journal article" year="1993">Martínez-Delclòs &amp; Martinell 1993</bibRefCitation>
, 
<bibRefCitation id="EF8C4B25000DFF88FDC9FEE6FD73130B" author="Wagner, T. &amp; Neinhuis, C. &amp; Barthlott, W." box="[517,733,259,286]" pageId="31" pageNumber="32" pagination="213 - 225" refId="ref62057" refString="Wagner, T., Neinhuis, C. &amp; Barthlott, W. (1996) Wettability and contaminability of insect wings as a function of their surface sculptures. Acta Zoologica, 77, 213 - 225. https: // doi. org / 10.1111 / j. 1463 - 6395.1996. tb 01265. x" type="journal article" year="1996">
Wagner 
<emphasis id="B969EAC6000DFF88FDAEFEE6FD3D1308" box="[610,659,259,285]" italics="true" pageId="31" pageNumber="32">et al</emphasis>
. 1996
</bibRefCitation>
).
</paragraph>
<paragraph id="8BA236D4000DFF88FF0BFEC2FB8613C4" blockId="31.[151,1437,151,897]" pageId="31" pageNumber="32">
They were less common, but still well-represented in the hot lowlands of mid-continental Ypresian 
<collectingRegion id="49D9F836000DFF88FAF8FEC2FA321354" box="[1332,1436,295,321]" country="United States of America" name="Colorado" pageId="31" pageNumber="32">Colorado</collectingRegion>
and 
<collectingRegion id="49D9F836000DFF88FF06FEAEFE961370" box="[202,312,331,357]" country="United States of America" name="Wyoming" pageId="31" pageNumber="32">Wyoming</collectingRegion>
(mean annual temperatures upper mesothermal to megathermal, megathermal = 20°C and over: 
<bibRefCitation id="EF8C4B25000DFF88FF5BFE8AFE34139F" author="Archibald, S. B. &amp; Johnson, K. R. &amp; Mathewes, R. W. &amp; Greenwood, D. R." box="[151,410,367,394]" pageId="31" pageNumber="32" pagination="3679 - 3686" refId="ref55797" refString="Archibald, S. B., Johnson, K. R., Mathewes, R. W. &amp; Greenwood, D. R. (2011 b) Intercontinental dispersal of giant thermophilic ants across the Arctic during early Eocene hyperthermals. Proceedings of the Royal Society B, 278, 3679 - 3686. https: // doi. org / 10.1098 / rspb. 2011.0729" type="journal article" year="2011">
Archibald 
<emphasis id="B969EAC6000DFF88FEC3FE8AFEEC139C" box="[271,322,367,393]" italics="true" pageId="31" pageNumber="32">et al</emphasis>
. 2011b
</bibRefCitation>
), and are fewer yet in the upland Priabonian of 
<collectingRegion id="49D9F836000DFF88FC76FE8AFB8C139C" box="[954,1058,367,393]" country="United States of America" name="Colorado" pageId="31" pageNumber="32">Colorado</collectingRegion>
(likely similar mean annual temperature as the Okanagan Highlands: 
<bibRefCitation id="EF8C4B25000DFF88FDF1FE76FCA513B8" author="Allena, S. E. &amp; Lowe, A. J. &amp; Peppe, D. J. &amp; Meyer, H. W." box="[573,779,403,429]" pageId="31" pageNumber="32" refId="ref55393" refString="Allena, S. E., Lowe, A. J., Peppe, D. J. &amp; Meyer, H. W. (2020) Paleoclimate and paleoecology of the latest Eocene Florissant flora of central Colorado, U. S. A. Palaeogeography, Palaeoclimatology, Palaeoecology, 551. [in press] https: // doi. org / 10.1016 / j. palaeo. 2020.109678" type="book" year="2020">
Allena 
<emphasis id="B969EAC6000DFF88FD43FE76FD6913B8" box="[655,711,403,429]" italics="true" pageId="31" pageNumber="32">et al.</emphasis>
2020
</bibRefCitation>
), where they were mixed with a variety of dragonflies and damselflies, and were present to lesser degrees in the Eocene of Europe and Asia.
</paragraph>
<paragraph id="8BA236D4000DFF88FF0BFE3EFBF41074" blockId="31.[151,1437,151,897]" pageId="31" pageNumber="32">
After the Eocene, the 
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become rare, with one (
<taxonomicName id="4C1D4D57000DFF88FC4BFE3EFB0C13E0" box="[903,1186,475,501]" class="Insecta" family="Thaumatoneuridae" genus="Primorilestes" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="31" pageNumber="32" phylum="Arthropoda" rank="species" species="magnificus">
<emphasis id="B969EAC6000DFF88FC4BFE3EFB0C13E0" box="[903,1186,475,501]" italics="true" pageId="31" pageNumber="32">Primorilestes magnificus</emphasis>
</taxonomicName>
), perhaps two (
<taxonomicName id="4C1D4D57000DFF88FA95FE3EFF50100C" authorityName="Cockerell" authorityYear="1908" class="Insecta" family="Dysagrionidae" genus="Phenacolestes" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="31" pageNumber="32" phylum="Arthropoda" rank="genus">
<emphasis id="B969EAC6000DFF88FA95FE3EFF50100C" italics="true" pageId="31" pageNumber="32">Phenacolestes</emphasis>
</taxonomicName>
? 
<emphasis id="B969EAC6000DFF88FEDFFE1AFED3100C" box="[275,381,511,537]" italics="true" pageId="31" pageNumber="32">coloratus</emphasis>
) occurrences in the Miocene. 
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dominate the 
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in the Oligocene and Miocene, the globally cooler post-Eocene icehouse world climatic regime (
<bibRefCitation id="EF8C4B25000DFF88FBC6FDC6FB731028" author="Zachos, J. &amp; Pagani, M. &amp; Sloan, L. &amp; Thomas, E. &amp; Billups, K." box="[1034,1245,547,573]" pageId="31" pageNumber="32" pagination="686 - 693" refId="ref62565" refString="Zachos, J., Pagani, M., Sloan, L., Thomas, E. &amp; Billups, K. (2001) Trends, rhythms, and aberrations in global climate 65 Ma to present. Science, 292, 686 - 693. https: // doi. org / 10.1126 / science. 1059412" type="journal article" year="2001">
Zachos 
<emphasis id="B969EAC6000DFF88FBAFFDC6FB3D1028" box="[1123,1171,547,573]" italics="true" pageId="31" pageNumber="32">et al</emphasis>
. 2001
</bibRefCitation>
), restricted to the Palearctic Realm. Their last record, and so that of the suborder, is in the late Miocene.
</paragraph>
<paragraph id="8BA236D4000DFF88FF0BFD8EFD3D1194" blockId="31.[151,1437,151,897]" pageId="31" pageNumber="32">
The cool, higher elevations of the Okanagan Highlands is where we see many of the earliest occurrences of the floral taxa and community associations of modern temperate Northern Hemisphere forests that later descended to lower elevations and spread throughout and predominate in much of the Holarctic as temperature seasonality increased outside of the tropics bringing colder winters, the latitudinal gradient of mean annual temperature steepened, and cooler global temperatures were established following the close of the Eocene (
<emphasis id="B969EAC6000DFF88FB5FFD1EFB1B1101" box="[1171,1205,763,788]" italics="true" pageId="31" pageNumber="32">e.g</emphasis>
., 
<bibRefCitation id="EF8C4B25000DFF88FB07FD1EFAC61100" author="Graham, A." box="[1227,1384,763,789]" pageId="31" pageNumber="32" refId="ref58235" refString="Graham, A. (1999) Late Cretaceous and Cenozoic history of North American vegetation north of Mexico. Oxford University Press, New York, xviii + 350 pp. https: // doi. org / 10.1093 / oso / 9780195113426.001.0001" type="book" year="1999">Graham 1999</bibRefCitation>
; 
<bibRefCitation id="EF8C4B25000DFF88FAB9FD1EFEE4112C" author="Zachos, J. &amp; Pagani, M. &amp; Sloan, L. &amp; Thomas, E. &amp; Billups, K." pageId="31" pageNumber="32" pagination="686 - 693" refId="ref62565" refString="Zachos, J., Pagani, M., Sloan, L., Thomas, E. &amp; Billups, K. (2001) Trends, rhythms, and aberrations in global climate 65 Ma to present. Science, 292, 686 - 693. https: // doi. org / 10.1126 / science. 1059412" type="journal article" year="2001">
Zachos 
<emphasis id="B969EAC6000DFF88FF1DFCFAFEAC112C" box="[209,258,799,825]" italics="true" pageId="31" pageNumber="32">et al</emphasis>
. 2001
</bibRefCitation>
, 
<bibRefCitation id="EF8C4B25000DFF88FE99FCFAFE3C112C" author="Zachos, J. C. &amp; Dickens, G. R. &amp; Zeebe, R. E." box="[341,402,799,825]" pageId="31" pageNumber="32" pagination="279 - 283" refId="ref62512" refString="Zachos, J. C., Dickens, G. R. &amp; Zeebe, R. E. (2008) An early Cenozoic perspective on greenhouse warming and carbon-cycle dynamics. Nature, 451, 279 - 283. https: // doi. org / 10.1038 / nature 06588" type="journal article" year="2008">2008</bibRefCitation>
). As the 
<taxonomicName id="4C1D4D57000DFF88FE3CFCFAFD12112C" box="[496,700,799,825]" kingdom="Animalia" pageId="31" pageNumber="27" rank="subOrder" subOrder="Cephalozygoptera">Cephalozygoptera</taxonomicName>
genera 
<taxonomicName id="4C1D4D57000DFF88FCD8FCFAFC35112C" authorityName="Archibald &amp; Cannings &amp; Erickson &amp; Bybee &amp; Mathewes" authorityYear="2021" box="[788,923,799,825]" class="Insecta" family="Dysagrionidae" genus="Okanagrion" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="31" pageNumber="32" phylum="Arthropoda" rank="genus">
<emphasis id="B969EAC6000DFF88FCD8FCFAFC35112C" box="[788,923,799,825]" italics="true" pageId="31" pageNumber="32">Okanagrion</emphasis>
</taxonomicName>
and 
<taxonomicName id="4C1D4D57000DFF88FC1CFCFAFBCF112C" authorityName="Archibald &amp; Cannings &amp; Erickson &amp; Bybee &amp; Mathewes" authorityYear="2021" box="[976,1121,799,825]" class="Insecta" family="Thaumatoneuridae" genus="Okanopteryx" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="31" pageNumber="32" phylum="Arthropoda" rank="genus">
<emphasis id="B969EAC6000DFF88FC1CFCFAFBCF112C" box="[976,1121,799,825]" italics="true" pageId="31" pageNumber="32">Okanopteryx</emphasis>
</taxonomicName>
were the dominant 
<taxonomicName id="4C1D4D57000DFF88FAF1FCFAFA32112C" box="[1341,1436,799,825]" class="Insecta" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="31" pageNumber="32" phylum="Arthropoda" rank="order">Odonata</taxonomicName>
of the montane Okanagan Highlands forests, why did they not spread with them and flourish across the Holarctic to the present? There is no obvious explanation.
</paragraph>
</subSubSection>
</treatment>
</document>