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<document ID-DOI="10.1016/j.ympev.2021.107115" ID-Zenodo-Dep="6643504" approvalRequired="21" approvalRequired_for_document="1" approvalRequired_for_taxonomicNames="20" checkinTime="1654106955118" checkinUser="diego" docAuthor="Bybee, Seth M., Kalkman, Vincent J., Erickson, Robert J., Frandsen, Paul B., Breinholt, Jesse W., Suvorov, Anton, Dijkstra, Klaas-Douwe B., Cordero-Rivera, Adolfo, Skevington, Jeffrey H., Abbott, John C., Herrera, Melissa Sanchez, Lemmon, Alan R., Lemmon, Emily Moriarty &amp; Ware, Jessica L." docDate="2021" docId="039687E7A86EFFD0E433AD4FFC72E922" docLanguage="en" docName="MolPhylogenetEvol.160.107115.pdf" docOrigin="Molecular Phylogenetics and Evolution 160" docTitle="Calopterygoidea Selys-Longchamps 1850" docType="treatment" docVersion="4" lastPageNumber="8" masterDocId="FFAFFF9FA869FFD7E457AC3AFFA9EC75" masterDocTitle="Phylogeny and classification of Odonata using targeted genomics" masterLastPageNumber="107115" masterPageNumber="107115" pageNumber="8" updateTime="1655236780830" updateUser="ExternalLinkService" zenodo-license-document="CC-BY-NC-ND-4.0">
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<mods:titleInfo>
<mods:title>Phylogeny and classification of Odonata using targeted genomics</mods:title>
</mods:titleInfo>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Bybee, Seth M.</mods:namePart>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Kalkman, Vincent J.</mods:namePart>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Erickson, Robert J.</mods:namePart>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Frandsen, Paul B.</mods:namePart>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Breinholt, Jesse W.</mods:namePart>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Suvorov, Anton</mods:namePart>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Dijkstra, Klaas-Douwe B.</mods:namePart>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Cordero-Rivera, Adolfo</mods:namePart>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Skevington, Jeffrey H.</mods:namePart>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Abbott, John C.</mods:namePart>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Herrera, Melissa Sanchez</mods:namePart>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Lemmon, Alan R.</mods:namePart>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Lemmon, Emily Moriarty</mods:namePart>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Ware, Jessica L.</mods:namePart>
</mods:name>
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<mods:titleInfo>
<mods:title>Molecular Phylogenetics and Evolution</mods:title>
</mods:titleInfo>
<mods:part>
<mods:date>2021</mods:date>
<mods:detail type="pubDate">
<mods:number>2021-02-18</mods:number>
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<mods:number>160</mods:number>
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<mods:classification>journal article</mods:classification>
<mods:identifier type="DOI">10.1016/j.ympev.2021.107115</mods:identifier>
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<emphasis box="[100,455,373,393]" italics="true" pageId="7" pageNumber="8">
<subSubSection box="[100,415,373,393]" pageId="7" pageNumber="8" type="nomenclature">
<paragraph blockId="7.[100,771,373,1955]" box="[100,415,373,393]" pageId="7" pageNumber="8">
<heading box="[100,415,373,393]" level="2" pageId="7" pageNumber="8">
<emphasis bold="true" box="[100,333,373,392]" italics="true" pageId="7" pageNumber="8">
4.1.5.2.
<taxonomicName authorityName="Selys-Longchamps" authorityYear="1850" box="[189,333,373,392]" kingdom="Animalia" pageId="7" pageNumber="7" rank="superFamily" superFamily="Calopterygoidea">Calopterygoidea</taxonomicName>
</emphasis>
<emphasis bold="true" box="[347,415,373,393]" italics="true" pageId="7" pageNumber="8">group 2</emphasis>
</heading>
</paragraph>
</subSubSection>
<subSubSection pageId="7" pageNumber="8" type="multiple">
<paragraph blockId="7.[100,771,373,1955]" pageId="7" pageNumber="8">
(BS =
<emphasis bold="true" box="[484,556,373,393]" italics="true" pageId="7" pageNumber="8">100, PP</emphasis>
=
<emphasis bold="true" box="[586,636,373,392]" italics="true" pageId="7" pageNumber="8">1, QS</emphasis>
=
<emphasis bold="true" italics="true" pageId="7" pageNumber="8">0.47/0.44/ 0.97).</emphasis>
</paragraph>
</subSubSection>
</emphasis>
<subSubSection pageId="7" pageNumber="8" type="discussion">
<paragraph blockId="7.[100,771,373,1955]" pageId="7" pageNumber="8">
Nine families make up Group 2. Included in these nine families, we propose two new families (
<taxonomicName authority="Bybee &amp; Kalkman &amp; Erickson &amp; Frandsen &amp; Breinholt &amp; Suvorov &amp; Dijkstra &amp; Cordero-Rivera &amp; Skevington &amp; Abbott &amp; Herrera &amp; Lemmon &amp; Lemmon &amp; Ware, 2021" authorityName="Bybee &amp; Kalkman &amp; Erickson &amp; Frandsen &amp; Breinholt &amp; Suvorov &amp; Dijkstra &amp; Cordero-Rivera &amp; Skevington &amp; Abbott &amp; Herrera &amp; Lemmon &amp; Lemmon &amp; Ware" authorityYear="2021" box="[407,549,429,448]" family="Mesagrionidae" pageId="7" pageNumber="13" rank="family" status="fam. nov.">Mesagrionidae</taxonomicName>
,
<taxonomicName authority="Bybee &amp; Kalkman &amp; Erickson &amp; Frandsen &amp; Breinholt &amp; Suvorov &amp; Dijkstra &amp; Cordero-Rivera &amp; Skevington &amp; Abbott &amp; Herrera &amp; Lemmon &amp; Lemmon &amp; Ware, 2021" authorityName="Bybee &amp; Kalkman &amp; Erickson &amp; Frandsen &amp; Breinholt &amp; Suvorov &amp; Dijkstra &amp; Cordero-Rivera &amp; Skevington &amp; Abbott &amp; Herrera &amp; Lemmon &amp; Lemmon &amp; Ware" authorityYear="2021" box="[562,687,429,448]" family="Protolestidae" pageId="7" pageNumber="13" rank="family" status="fam. nov.">Protolestidae</taxonomicName>
) and co recognize Tatocnemididae as at the family level (see below Family-level revisions to the Classification of
<taxonomicName authorityName="Selys-Longchamps" authorityYear="1854" box="[406,506,485,504]" pageId="7" pageNumber="15" rank="subOrder" subOrder="Zygoptera">Zygoptera</taxonomicName>
). Seven of these families are unique morphologically and contain either a single genus (
<taxonomicName authorityName="Novelo-Gutierrez" authorityYear="1995" class="Insecta" family="Pentaphlebiidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="7" pageNumber="8" phylum="Arthropoda" rank="family">Pentaphlebiidae</taxonomicName>
,
<taxonomicName authorityName="Tillyard &amp; Fraser" authorityYear="1938" box="[216,341,541,560]" class="Insecta" family="Hypolestidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="7" pageNumber="8" phylum="Arthropoda" rank="family">Hypolestidae</taxonomicName>
,
<taxonomicName authority="Bybee &amp; Kalkman &amp; Erickson &amp; Frandsen &amp; Breinholt &amp; Suvorov &amp; Dijkstra &amp; Cordero-Rivera &amp; Skevington &amp; Abbott &amp; Herrera &amp; Lemmon &amp; Lemmon &amp; Ware, 2021" authorityName="Bybee &amp; Kalkman &amp; Erickson &amp; Frandsen &amp; Breinholt &amp; Suvorov &amp; Dijkstra &amp; Cordero-Rivera &amp; Skevington &amp; Abbott &amp; Herrera &amp; Lemmon &amp; Lemmon &amp; Ware" authorityYear="2021" box="[355,495,541,560]" family="Mesagrionidae" pageId="7" pageNumber="13" rank="family" status="fam. nov.">Mesagrionidae</taxonomicName>
,
<taxonomicName authority="Bybee &amp; Kalkman &amp; Erickson &amp; Frandsen &amp; Breinholt &amp; Suvorov &amp; Dijkstra &amp; Cordero-Rivera &amp; Skevington &amp; Abbott &amp; Herrera &amp; Lemmon &amp; Lemmon &amp; Ware, 2021" authorityName="Bybee &amp; Kalkman &amp; Erickson &amp; Frandsen &amp; Breinholt &amp; Suvorov &amp; Dijkstra &amp; Cordero-Rivera &amp; Skevington &amp; Abbott &amp; Herrera &amp; Lemmon &amp; Lemmon &amp; Ware" authorityYear="2021" box="[509,632,541,560]" family="Protolestidae" pageId="7" pageNumber="13" rank="family" status="fam. nov.">Protolestidae</taxonomicName>
and Tatocnemididae) or two genera (
<taxonomicName authorityName="Selys" authorityYear="1853" box="[349,477,569,588]" class="Insecta" family="Dicteriadidae" kingdom="Animalia" order="Odonata" pageId="7" pageNumber="8" phylum="Arthropoda" rank="family">Dicteriadidae</taxonomicName>
and
<taxonomicName authorityName="Racenis" authorityYear="1959" box="[532,665,569,588]" class="Insecta" family="Philogeniidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="7" pageNumber="8" phylum="Arthropoda" rank="family">Philogeniidae</taxonomicName>
). Our new phylogenetic hypothesis has no taxonomic consequences regarding the existing families, except for
<taxonomicName authorityName="Racenis" authorityYear="1959" box="[385,540,624,643]" class="Insecta" family="Heteragrionidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="7" pageNumber="8" phylum="Arthropoda" rank="family">Heteragrionidae</taxonomicName>
, which is expanded to include the genera
<emphasis bold="true" box="[276,378,652,671]" italics="true" pageId="7" pageNumber="8">Dimeragion</emphasis>
and
<taxonomicName authorityName="Selys" authorityYear="1885" box="[421,568,652,671]" class="Insecta" family="Megapodagrionidae" genus="Heteropodagrion" kingdom="Animalia" order="Odonata" pageId="7" pageNumber="8" phylum="Arthropoda" rank="genus">
<emphasis bold="true" box="[421,568,652,671]" italics="true" pageId="7" pageNumber="8">Heteropodagrion</emphasis>
</taxonomicName>
. With this addition to
<taxonomicName authorityName="Racenis" authorityYear="1959" box="[100,252,680,699]" class="Insecta" family="Heteragrionidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="7" pageNumber="8" phylum="Arthropoda" rank="family">Heteragrionidae</taxonomicName>
there are now two families with more than two genera;
<taxonomicName authorityName="Racenis" authorityYear="1959" box="[100,252,708,727]" class="Insecta" family="Heteragrionidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="7" pageNumber="8" phylum="Arthropoda" rank="family">Heteragrionidae</taxonomicName>
with four genera and
<taxonomicName authorityName="Munz" authorityYear="1919" box="[465,585,708,727]" class="Insecta" family="Polythoridae" kingdom="Animalia" order="Odonata" pageId="7" pageNumber="8" phylum="Arthropoda" rank="family">Polythoridae</taxonomicName>
with seven genera. With the exception of a well-supported cluster of three Afrotropical families (Tatocnemididae,
<taxonomicName authority="Bybee &amp; Kalkman &amp; Erickson &amp; Frandsen &amp; Breinholt &amp; Suvorov &amp; Dijkstra &amp; Cordero-Rivera &amp; Skevington &amp; Abbott &amp; Herrera &amp; Lemmon &amp; Lemmon &amp; Ware, 2021" authorityName="Bybee &amp; Kalkman &amp; Erickson &amp; Frandsen &amp; Breinholt &amp; Suvorov &amp; Dijkstra &amp; Cordero-Rivera &amp; Skevington &amp; Abbott &amp; Herrera &amp; Lemmon &amp; Lemmon &amp; Ware" authorityYear="2021" box="[348,471,764,783]" family="Protolestidae" pageId="7" pageNumber="13" rank="family" status="fam. nov.">Protolestidae</taxonomicName>
and
<taxonomicName authorityName="Novelo-Gutierrez" authorityYear="1995" box="[516,670,764,783]" class="Insecta" family="Pentaphlebiidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="7" pageNumber="8" phylum="Arthropoda" rank="family">Pentaphlebiidae</taxonomicName>
; BS = 100, PP = 1, QS = 0.74/0.29/0.97) Group 2 is largely Neotropical. The relationships between the families are difficult to determine from a morphological and/or behavioural perspective as they have few characters in common. Further, although the clade is well supported overall, the relationships between many of the families are not well supported due to low QS values. This is a clade where additional taxon sampling throughout the clade itself and throughout the “calopterygoidea” in general is likely to reveal much more clarity towards the overall classification.
</paragraph>
<paragraph blockId="7.[100,771,373,1955]" pageId="7" pageNumber="8">
<taxonomicName authority="Bybee &amp; Kalkman &amp; Erickson &amp; Frandsen &amp; Breinholt &amp; Suvorov &amp; Dijkstra &amp; Cordero-Rivera &amp; Skevington &amp; Abbott &amp; Herrera &amp; Lemmon &amp; Lemmon &amp; Ware, 2021" authorityName="Bybee &amp; Kalkman &amp; Erickson &amp; Frandsen &amp; Breinholt &amp; Suvorov &amp; Dijkstra &amp; Cordero-Rivera &amp; Skevington &amp; Abbott &amp; Herrera &amp; Lemmon &amp; Lemmon &amp; Ware" authorityYear="2021" box="[132,255,1043,1062]" family="Protolestidae" pageId="7" pageNumber="13" rank="family" status="fam. nov.">Protolestidae</taxonomicName>
(BS = 100, PP = 1, QS = 1/NA/1) and Tatocnemididae are here recognised as distinct families (see below Family-level revisions to the Classification of
<taxonomicName authorityName="Selys-Longchamps" authorityYear="1854" box="[311,408,1099,1118]" pageId="7" pageNumber="15" rank="subOrder" subOrder="Zygoptera">Zygoptera</taxonomicName>
) as they have no clear relatives and are quite distinct in adult and nymphal morphology from their closest relative,
<taxonomicName authorityName="Novelo-Gutierrez" authorityYear="1995" box="[184,339,1154,1173]" class="Insecta" family="Pentaphlebiidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="7" pageNumber="8" phylum="Arthropoda" rank="family">Pentaphlebiidae</taxonomicName>
. Both
<taxonomicName authorityName="Forster" authorityYear="1899" box="[397,491,1154,1173]" class="Insecta" family="Megapodagrionidae" genus="Protolestes" kingdom="Animalia" order="Odonata" pageId="7" pageNumber="8" phylum="Arthropoda" rank="genus">
<emphasis bold="true" box="[397,491,1154,1173]" italics="true" pageId="7" pageNumber="8">Protolestes</emphasis>
</taxonomicName>
(eight species described) and
<taxonomicName authorityName="Kirby" authorityYear="1889" box="[100,200,1182,1201]" class="Insecta" family="Megapodagrionidae" genus="Tatocnemis" kingdom="Animalia" order="Odonata" pageId="7" pageNumber="8" phylum="Arthropoda" rank="genus">
<emphasis bold="true" box="[100,200,1182,1201]" italics="true" pageId="7" pageNumber="8">Tatocnemis</emphasis>
</taxonomicName>
(ten species) are restricted to rainforest streams in
<collectingCountry box="[100,214,1210,1229]" name="Madagascar" pageId="7" pageNumber="8">Madagascar</collectingCountry>
, were until recently included in
<taxonomicName authorityName="Calvert" authorityYear="1913" box="[534,723,1210,1229]" class="Insecta" family="Megapodagrionidae" kingdom="Animalia" order="Odonata" pageId="7" pageNumber="8" phylum="Arthropoda" rank="family">Megapodagrionidae</taxonomicName>
, and are poorly known and in dire need of taxonomic revision. Adults of both genera perch with wings variably closed or (half) open and the abdomen held roughly horizontal. Protolestid nymphs have fan-shaped caudal gills, a character only shared with the distantly related
<taxonomicName authority="(Kalkman et al., 2010)" baseAuthorityName="Kalkman" baseAuthorityYear="2010" class="Insecta" family="Argiolestidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="7" pageNumber="8" phylum="Arthropoda" rank="family">
Argiolestidae (
<bibRefCitation author="Kalkman, V. J. &amp; Choong, C. Y. &amp; Orr, A. G. &amp; Schutte, K." box="[108,303,1350,1369]" journalOrPublisher="Int. J. Odonatol." pageId="7" pageNumber="8" pagination="119 - 135" part="13" refId="ref18012" refString="Kalkman, V. J., Choong, C. Y., Orr, A. G., Schutte, K., 2010. Remarks on the taxonomy of Megapodagrionidae with emphasis on the larval gills (Odonata). Int. J. Odonatol. 13, 119 - 135. https: // doi. org / 10.1080 / 13887890.2010.9748366." title="Remarks on the taxonomy of Megapodagrionidae with emphasis on the larval gills (Odonata)" type="journal article" year="2010">Kalkman et al., 2010</bibRefCitation>
)
</taxonomicName>
and
<taxonomicName authority="Bybee &amp; Kalkman &amp; Erickson &amp; Frandsen &amp; Breinholt &amp; Suvorov &amp; Dijkstra &amp; Cordero-Rivera &amp; Skevington &amp; Abbott &amp; Herrera &amp; Lemmon &amp; Lemmon &amp; Ware, 2021" authorityName="Bybee &amp; Kalkman &amp; Erickson &amp; Frandsen &amp; Breinholt &amp; Suvorov &amp; Dijkstra &amp; Cordero-Rivera &amp; Skevington &amp; Abbott &amp; Herrera &amp; Lemmon &amp; Lemmon &amp; Ware" authorityYear="2021" box="[358,542,1350,1369]" family="Mesopodagrionidae" pageId="7" pageNumber="13" rank="family" status="fam. nov.">Mesopodagrionidae</taxonomicName>
(
<bibRefCitation author="Yu, X." box="[556,635,1350,1369]" journalOrPublisher="Int. J. Odonatol." pageId="7" pageNumber="8" pagination="275 - 282" part="19" refId="ref20823" refString="Yu, X., 2016. A description of the larva of Mesopodagrion tibetanum australe (Odonata: &quot; Megapodagrionidae &quot;). Int. J. Odonatol. 19, 275 - 282." title="A description of the larva of Mesopodagrion tibetanum australe (Odonata: &quot; Megapodagrionidae &quot;)" type="journal article" year="2016">Yu 2016</bibRefCitation>
), while adults have a rather wide and slender head, similar to some members of the unrelated
<taxonomicName authorityName="Jacobson &amp; Bianchi" authorityYear="1905" box="[199,359,1406,1425]" class="Insecta" family="Platycnemididae" kingdom="Animalia" order="Odonata" pageId="7" pageNumber="8" phylum="Arthropoda" rank="family">Platycnemididae</taxonomicName>
. Based on morphology, Tatocnemididae is not similar to other families: the potential apomorphy of crenulated wing tips is shared only with some genera of the unrelated
<taxonomicName authorityName="Jacobson &amp; Bianchi" authorityYear="1905" class="Insecta" family="Platycnemididae" kingdom="Animalia" order="Odonata" pageId="7" pageNumber="8" phylum="Arthropoda" rank="family">Platycnemididae</taxonomicName>
. The nymph has inflated saccoid caudal gills bearing a terminal filament as foundin several other families of
<taxonomicName authorityName="Selys-Longchamps" authorityYear="1854" box="[514,612,1518,1537]" pageId="7" pageNumber="15" rank="subOrder" subOrder="Zygoptera">Zygoptera</taxonomicName>
. Tatocnemididae were originally described by
<bibRefCitation author="R'acenis, J." box="[384,528,1544,1564]" journalOrPublisher="Acta Biologica Venezolana" pageId="7" pageNumber="8" pagination="335 - 367" part="2" refId="ref19766" refString="R'acenis, J., 1959. Notas taxonomicas sobre la familia Megapodagrionidae (Odonata: Zygoptera) con la sinopsis de las especies venezolanas. Acta Biologica Venezolana 2 (30), 335 - 367." title="Notas taxonomicas sobre la familia Megapodagrionidae (Odonata: Zygoptera) con la sinopsis de las especies venezolanas" type="journal article" year="1959">R´acenis (1959)</bibRefCitation>
as a subfamily of
<taxonomicName authorityName="Calvert" authorityYear="1913" class="Insecta" family="Megapodagrionidae" kingdom="Animalia" order="Odonata" pageId="7" pageNumber="8" phylum="Arthropoda" rank="family">Megapodagrionidae</taxonomicName>
to include
<taxonomicName authorityName="Kirby" authorityYear="1889" box="[339,439,1573,1592]" class="Insecta" family="Megapodagrionidae" genus="Tatocnemis" kingdom="Animalia" order="Odonata" pageId="7" pageNumber="8" phylum="Arthropoda" rank="genus">
<emphasis bold="true" box="[339,439,1573,1592]" italics="true" pageId="7" pageNumber="8">Tatocnemis</emphasis>
</taxonomicName>
and
<taxonomicName authorityName="Kennedy" authorityYear="1939" box="[485,647,1573,1592]" class="Insecta" family="Megapodagrionidae" genus="Archaeopodagrion" kingdom="Animalia" order="Odonata" pageId="7" pageNumber="8" phylum="Arthropoda" rank="genus">
<emphasis bold="true" box="[485,647,1573,1592]" italics="true" pageId="7" pageNumber="8">Archaeopodagrion</emphasis>
</taxonomicName>
, but are now restricted to the genus
<taxonomicName authorityName="Kirby" authorityYear="1889" box="[317,417,1601,1620]" class="Insecta" family="Megapodagrionidae" genus="Tatocnemis" kingdom="Animalia" order="Odonata" pageId="7" pageNumber="8" phylum="Arthropoda" rank="genus">
<emphasis bold="true" box="[317,417,1601,1620]" italics="true" pageId="7" pageNumber="8">Tatocnemis</emphasis>
</taxonomicName>
.
</paragraph>
<paragraph blockId="7.[100,771,373,1955]" lastBlockId="7.[818,1488,148,1367]" pageId="7" pageNumber="8">
The endemic species
<emphasis bold="true" box="[347,559,1628,1648]" italics="true" pageId="7" pageNumber="8">
<taxonomicName authorityName="Selys" authorityYear="1885" box="[347,438,1629,1648]" class="Insecta" family="Megapodagrionidae" genus="Mesagrion" kingdom="Animalia" order="Odonata" pageId="7" pageNumber="8" phylum="Arthropoda" rank="genus">Mesagrion</taxonomicName>
leucorhinum
</emphasis>
from the Colombian Andes is found as sister to two small families,
<taxonomicName authorityName="Selys" authorityYear="1853" box="[527,653,1657,1676]" class="Insecta" family="Dicteriadidae" kingdom="Animalia" order="Odonata" pageId="7" pageNumber="8" phylum="Arthropoda" rank="family">Dicteriadidae</taxonomicName>
(two genera each with a single species from the Amazonian region) and
<taxonomicName authorityName="Tillyard &amp; Fraser" authorityYear="1938" box="[647,770,1685,1704]" class="Insecta" family="Hypolestidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="7" pageNumber="8" phylum="Arthropoda" rank="family">Hypolestidae</taxonomicName>
(one genus, three species from the Greater Antilles), but with low QS values (BS = 100, PP = 1, QS = -0.45/0.28/0.99). Both of these families have well defined apomorphies, are fairly distinctive and do not seem particularly close to
<emphasis bold="true" box="[292,431,1796,1815]" italics="true" pageId="7" pageNumber="8">M. leucorhinum</emphasis>
from a morphological or behavioral perspective.
<emphasis bold="true" box="[222,432,1824,1843]" italics="true" pageId="7" pageNumber="8">
<taxonomicName authorityName="Selys" authorityYear="1885" box="[222,313,1824,1843]" class="Insecta" family="Megapodagrionidae" genus="Mesagrion" kingdom="Animalia" order="Odonata" pageId="7" pageNumber="8" phylum="Arthropoda" rank="genus">Mesagrion</taxonomicName>
leucorhinum
</emphasis>
was not included in previous molecular analyses (
<bibRefCitation author="Bybee, S. M. &amp; Ogden, T. H. &amp; Branham, M. A. &amp; Whiting, M. F." box="[266,442,1852,1871]" journalOrPublisher="Cladistics" pageId="7" pageNumber="8" pagination="477 - 514" part="24" refId="ref16610" refString="Bybee, S. M., Ogden, T. H., Branham, M. A., Whiting, M. F., 2008. Molecules, morphology and fossils: a comprehensive approach to odonate phylogeny and the evolution of the odonate wing. Cladistics 24, 477 - 514." title="Molecules, morphology and fossils: a comprehensive approach to odonate phylogeny and the evolution of the odonate wing" type="journal article" year="2008">Bybee et al., 2008</bibRefCitation>
;
<bibRefCitation author="Davis, R. B. &amp; Nicholson, D. B. &amp; Saunders, E. L. R. &amp; Mayhew, P. J." box="[453,623,1852,1871]" journalOrPublisher="BMC Evol. Biol." pageId="7" pageNumber="8" pagination="252" part="11" refId="ref17013" refString="Davis, R. B., Nicholson, D. B., Saunders, E. L. R., Mayhew, P. J., 2011. Fossil gaps inferred from phylogenies alter the apparent nature of diversification in dragonflies and their relatives. BMC Evol. Biol. 11, 252." title="Fossil gaps inferred from phylogenies alter the apparent nature of diversification in dragonflies and their relatives" type="journal article" year="2011">Davis et al., 2011</bibRefCitation>
;
<bibRefCitation author="Dijkstra, K. - D. - B. &amp; Kalkman, V. J. &amp; Dow, R. A. &amp; Stokvis, F. R. &amp; Van Tol, J. A. N." journalOrPublisher="Syst. Entomol." pageId="7" pageNumber="8" pagination="68 - 96" part="39" refId="ref17175" refString="Dijkstra, K. - D. - B., Kalkman, V. J., Dow, R. A., Stokvis, F. R., Van Tol, J. A. N., 2014. Redefining the damselfly families: a comprehensive molecular phylogeny of Z ygoptera (O donata). Syst. Entomol. 39, 68 - 96." title="Redefining the damselfly families: a comprehensive molecular phylogeny of Z ygoptera (O donata)" type="journal article" year="2014">Dijkstra et al., 2014</bibRefCitation>
;
<bibRefCitation author="Dumont, H. J. &amp; Vierstraete, A. &amp; Vanfleteren, J. R." box="[162,357,1880,1899]" journalOrPublisher="Syst. Entomol." pageId="7" pageNumber="8" pagination="6 - 18" part="35" refId="ref17305" refString="Dumont, H. J., Vierstraete, A., Vanfleteren, J. R., 2010. A molecular phylogeny of the Odonata (Insecta). Syst. Entomol. 35, 6 - 18." title="A molecular phylogeny of the Odonata (Insecta)" type="journal article" year="2010">Dumont et al., 2010</bibRefCitation>
;
<bibRefCitation author="van Tol, J. &amp; Reijnen, B. T." box="[369,625,1880,1899]" journalOrPublisher="Leiden University, Leiden" pageId="7" pageNumber="8" refId="ref20410" refString="van Tol, J., Reijnen, B. T., 2009. Phylogeny and biogeography of the Platystictidae (Odonata). Leiden University, Leiden, p. PhD thesis." title="Phylogeny and biogeography of the Platystictidae (Odonata)" type="book" year="2009">van Tol and Reijnen, 2009</bibRefCitation>
), but based on morphology it was tentatively placed in an
<emphasis bold="true" box="[504,626,1907,1927]" italics="true" pageId="7" pageNumber="8">Incertae Sedis</emphasis>
group together with
<taxonomicName authorityName="Calvert" authorityYear="1913" box="[150,260,1936,1955]" class="Insecta" family="Megapodagrionidae" genus="Dimeragrion" kingdom="Animalia" order="Odonata" pageId="7" pageNumber="8" phylum="Arthropoda" rank="genus">
<emphasis bold="true" box="[150,260,1936,1955]" italics="true" pageId="7" pageNumber="8">Dimeragrion</emphasis>
</taxonomicName>
and
<taxonomicName authorityName="Selys" authorityYear="1885" box="[308,455,1935,1954]" class="Insecta" family="Megapodagrionidae" genus="Heteropodagrion" kingdom="Animalia" order="Odonata" pageId="7" pageNumber="8" phylum="Arthropoda" rank="genus">
<emphasis bold="true" box="[308,455,1935,1954]" italics="true" pageId="7" pageNumber="8">Heteropodagrion</emphasis>
</taxonomicName>
. In our analyses
<taxonomicName authorityName="Selys" authorityYear="1885" box="[619,710,1936,1955]" class="Insecta" family="Megapodagrionidae" genus="Mesagrion" kingdom="Animalia" order="Odonata" pageId="7" pageNumber="8" phylum="Arthropoda" rank="genus">
<emphasis bold="true" box="[619,710,1936,1955]" italics="true" pageId="7" pageNumber="8">Mesagrion</emphasis>
</taxonomicName>
is not found to be close to these genera. As there are no other likely candidates to be the closest relative of
<taxonomicName authorityName="Selys" authorityYear="1885" box="[1068,1159,176,195]" class="Insecta" family="Megapodagrionidae" genus="Mesagrion" kingdom="Animalia" order="Odonata" pageId="7" pageNumber="8" phylum="Arthropoda" rank="genus">
<emphasis bold="true" box="[1068,1159,176,195]" italics="true" pageId="7" pageNumber="8">Mesagrion</emphasis>
</taxonomicName>
, we propose to regard it as a family in its own right Family-level revisions to the Classification of
<taxonomicName authorityName="Selys-Longchamps" authorityYear="1854" box="[1386,1484,204,223]" pageId="7" pageNumber="15" rank="subOrder" subOrder="Zygoptera">Zygoptera</taxonomicName>
. Apomorphies for this family (although not unique within
<taxonomicName authorityName="Selys-Longchamps" authorityYear="1854" box="[1350,1447,232,251]" pageId="7" pageNumber="15" rank="subOrder" subOrder="Zygoptera">Zygoptera</taxonomicName>
) are the scarcely sclerotized dorsum of segment eight in the female and the long paraprocts which are serrated at the distal fourth of the dorsal margin (
<bibRefCitation author="Garrison, R. W. &amp; von Ellenrieder, N. &amp; Louton, J. A." box="[902,1109,315,334]" journalOrPublisher="Johns Hopkins University Press" pageId="7" pageNumber="8" refId="ref17591" refString="Garrison, R. W., von Ellenrieder, N., Louton, J. A., 2010. Damselfly Genera of the New World: An Illustrated and Annotated Key to the Zygoptera. Johns Hopkins University Press." title="Damselfly Genera of the New World: An Illustrated and Annotated Key to the Zygoptera" type="book" year="2010">Garrison et al., 2010</bibRefCitation>
;
<bibRefCitation author="P'erez-Guti'errez, L. A &amp; Montes-Fontalvo, J. M." journalOrPublisher="Int. J. Odonatol." pageId="7" pageNumber="8" pagination="91 - 100" part="14" refId="ref19524" refString="P'erez-Guti'errez, L. A., Montes-Fontalvo, J. M., 2011. Rediscovery of Mesagrion leucorrhinum (Zygoptera: Megapodagrionidae): a &quot; formal &quot; description of female and ultimate stadium of larva with notes on habits. Int. J. Odonatol. 14, 91 - 100." title="Rediscovery of Mesagrion leucorrhinum (Zygoptera: Megapodagrionidae): a &quot; formal &quot; description of female and ultimate stadium of larva with notes on habits" type="journal article" year="2011">P´erez-Guti´errez and Montes-Fontalvo, 2011</bibRefCitation>
). The species rests with its wings closed, which was regarded as an additional indication that
<taxonomicName authorityName="Selys" authorityYear="1885" box="[1087,1178,371,390]" class="Insecta" family="Megapodagrionidae" genus="Mesagrion" kingdom="Animalia" order="Odonata" pageId="7" pageNumber="8" phylum="Arthropoda" rank="genus">
<emphasis bold="true" box="[1087,1178,371,390]" italics="true" pageId="7" pageNumber="8">Mesagrion</emphasis>
</taxonomicName>
was close to
<emphasis bold="true" box="[1298,1451,370,389]" italics="true" pageId="7" pageNumber="8">
<taxonomicName authorityName="Selys" authorityYear="1885" box="[1298,1447,370,389]" class="Insecta" family="Megapodagrionidae" genus="Heteropodagrion" kingdom="Animalia" order="Odonata" pageId="7" pageNumber="8" phylum="Arthropoda" rank="genus">Heteropodagrion</taxonomicName>
,
</emphasis>
but it is now clear that this habit evolved several times within the families previously grouped into Calopterygoidea.
</paragraph>
<paragraph blockId="7.[818,1488,148,1367]" pageId="7" pageNumber="8">
Recent hypotheses proposed
<taxonomicName authorityName="Racenis" authorityYear="1959" box="[1119,1271,455,474]" class="Insecta" family="Heteragrionidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="7" pageNumber="8" phylum="Arthropoda" rank="family">Heteragrionidae</taxonomicName>
to be composed of two South American genera:
<taxonomicName authorityName="Selys" authorityYear="1862" box="[1051,1155,482,501]" class="Insecta" family="Megapodagrionidae" genus="Heteragrion" kingdom="Animalia" order="Odonata" pageId="7" pageNumber="8" phylum="Arthropoda" rank="genus">
<emphasis bold="true" box="[1051,1155,482,501]" italics="true" pageId="7" pageNumber="8">Heteragrion</emphasis>
</taxonomicName>
and
<taxonomicName authorityName="Selys" authorityYear="1862" box="[1202,1296,482,501]" class="Magnoliopsida" family="Fabaceae" genus="Oxystigma" kingdom="Plantae" order="Fabales" pageId="7" pageNumber="8" phylum="Tracheophyta" rank="genus">
<emphasis bold="true" box="[1202,1296,482,501]" italics="true" pageId="7" pageNumber="8">Oxystigma</emphasis>
</taxonomicName>
(
<bibRefCitation author="Bybee, S. M. &amp; Ogden, T. H. &amp; Branham, M. A. &amp; Whiting, M. F." box="[1311,1483,482,502]" journalOrPublisher="Cladistics" pageId="7" pageNumber="8" pagination="477 - 514" part="24" refId="ref16610" refString="Bybee, S. M., Ogden, T. H., Branham, M. A., Whiting, M. F., 2008. Molecules, morphology and fossils: a comprehensive approach to odonate phylogeny and the evolution of the odonate wing. Cladistics 24, 477 - 514." title="Molecules, morphology and fossils: a comprehensive approach to odonate phylogeny and the evolution of the odonate wing" type="journal article" year="2008">Bybee et al., 2008</bibRefCitation>
;
<bibRefCitation author="Davis, R. B. &amp; Nicholson, D. B. &amp; Saunders, E. L. R. &amp; Mayhew, P. J." box="[818,981,510,530]" journalOrPublisher="BMC Evol. Biol." pageId="7" pageNumber="8" pagination="252" part="11" refId="ref17013" refString="Davis, R. B., Nicholson, D. B., Saunders, E. L. R., Mayhew, P. J., 2011. Fossil gaps inferred from phylogenies alter the apparent nature of diversification in dragonflies and their relatives. BMC Evol. Biol. 11, 252." title="Fossil gaps inferred from phylogenies alter the apparent nature of diversification in dragonflies and their relatives" type="journal article" year="2011">Davis et al., 2011</bibRefCitation>
;
<bibRefCitation author="Dijkstra, K. - D. - B. &amp; Kalkman, V. J. &amp; Dow, R. A. &amp; Stokvis, F. R. &amp; Van Tol, J. A. N." box="[989,1174,510,530]" journalOrPublisher="Syst. Entomol." pageId="7" pageNumber="8" pagination="68 - 96" part="39" refId="ref17175" refString="Dijkstra, K. - D. - B., Kalkman, V. J., Dow, R. A., Stokvis, F. R., Van Tol, J. A. N., 2014. Redefining the damselfly families: a comprehensive molecular phylogeny of Z ygoptera (O donata). Syst. Entomol. 39, 68 - 96." title="Redefining the damselfly families: a comprehensive molecular phylogeny of Z ygoptera (O donata)" type="journal article" year="2014">Dijkstra et al., 2014</bibRefCitation>
;
<bibRefCitation author="Dumont, H. J. &amp; Vierstraete, A. &amp; Vanfleteren, J. R." box="[1182,1369,510,530]" journalOrPublisher="Syst. Entomol." pageId="7" pageNumber="8" pagination="6 - 18" part="35" refId="ref17305" refString="Dumont, H. J., Vierstraete, A., Vanfleteren, J. R., 2010. A molecular phylogeny of the Odonata (Insecta). Syst. Entomol. 35, 6 - 18." title="A molecular phylogeny of the Odonata (Insecta)" type="journal article" year="2010">Dumont et al., 2010</bibRefCitation>
;
<bibRefCitation author="van Tol, J. &amp; Reijnen, B. T." journalOrPublisher="Leiden University, Leiden" pageId="7" pageNumber="8" refId="ref20410" refString="van Tol, J., Reijnen, B. T., 2009. Phylogeny and biogeography of the Platystictidae (Odonata). Leiden University, Leiden, p. PhD thesis." title="Phylogeny and biogeography of the Platystictidae (Odonata)" type="book" year="2009">van Tol and Reijnen, 2009</bibRefCitation>
). In their Bayesian analyses (
<bibRefCitation author="Dijkstra, K. - D. - B. &amp; Kalkman, V. J. &amp; Dow, R. A. &amp; Stokvis, F. R. &amp; Van Tol, J. A. N." box="[1241,1435,538,558]" journalOrPublisher="Syst. Entomol." pageId="7" pageNumber="8" pagination="68 - 96" part="39" refId="ref17175" refString="Dijkstra, K. - D. - B., Kalkman, V. J., Dow, R. A., Stokvis, F. R., Van Tol, J. A. N., 2014. Redefining the damselfly families: a comprehensive molecular phylogeny of Z ygoptera (O donata). Syst. Entomol. 39, 68 - 96." title="Redefining the damselfly families: a comprehensive molecular phylogeny of Z ygoptera (O donata)" type="journal article" year="2014">Dijkstra et al., 2014</bibRefCitation>
), the South American genera
<taxonomicName authorityName="Calvert" authorityYear="1913" box="[1044,1154,566,585]" class="Insecta" family="Megapodagrionidae" genus="Dimeragrion" kingdom="Animalia" order="Odonata" pageId="7" pageNumber="8" phylum="Arthropoda" rank="genus">
<emphasis bold="true" box="[1044,1154,566,585]" italics="true" pageId="7" pageNumber="8">Dimeragrion</emphasis>
</taxonomicName>
and
<taxonomicName authorityName="Selys" authorityYear="1885" box="[1201,1348,566,585]" class="Insecta" family="Megapodagrionidae" genus="Heteropodagrion" kingdom="Animalia" order="Odonata" pageId="7" pageNumber="8" phylum="Arthropoda" rank="genus">
<emphasis bold="true" box="[1201,1348,566,585]" italics="true" pageId="7" pageNumber="8">Heteropodagrion</emphasis>
</taxonomicName>
were found to be sister to
<taxonomicName authorityName="Racenis" authorityYear="1959" box="[928,1080,594,613]" class="Insecta" family="Heteragrionidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="7" pageNumber="8" phylum="Arthropoda" rank="family">Heteragrionidae</taxonomicName>
and would have been included too were it not for the ML analyses which showed these two genera to be close to
<taxonomicName authorityName="Racenis" authorityYear="1959" box="[818,970,650,669]" class="Insecta" family="Heteragrionidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="7" pageNumber="8" phylum="Arthropoda" rank="family">Heteragrionidae</taxonomicName>
but with
<taxonomicName authorityName="Needham" authorityYear="1934" box="[1083,1173,649,668]" class="Insecta" family="Amphipterygidae" genus="Rimanella" kingdom="Animalia" order="Odonata" pageId="7" pageNumber="8" phylum="Arthropoda" rank="genus">
<emphasis bold="true" box="[1083,1173,649,668]" italics="true" pageId="7" pageNumber="8">Rimanella</emphasis>
</taxonomicName>
(
<taxonomicName box="[1192,1319,650,669]" class="Odonata" family="Rimanellidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Insecta" pageId="7" pageNumber="8" phylum="Arthropoda" rank="family">Rimanellidae</taxonomicName>
) and
<taxonomicName box="[1388,1488,649,668]" class="Insecta" family="Dicteriadidae" genus="Heliocharis" kingdom="Animalia" order="Odonata" pageId="7" pageNumber="8" phylum="Arthropoda" rank="genus">
<emphasis bold="true" box="[1388,1488,649,668]" italics="true" pageId="7" pageNumber="8">Heliocharis</emphasis>
</taxonomicName>
(
<taxonomicName authorityName="Selys" authorityYear="1853" box="[824,954,678,697]" class="Insecta" family="Dicteriadidae" kingdom="Animalia" order="Odonata" pageId="7" pageNumber="8" phylum="Arthropoda" rank="family">Dicteriadidae</taxonomicName>
) intermingled. In our analyses
<taxonomicName authorityName="Calvert" authorityYear="1913" box="[1253,1363,678,697]" class="Insecta" family="Megapodagrionidae" genus="Dimeragrion" kingdom="Animalia" order="Odonata" pageId="7" pageNumber="8" phylum="Arthropoda" rank="genus">
<emphasis bold="true" box="[1253,1363,678,697]" italics="true" pageId="7" pageNumber="8">Dimeragrion</emphasis>
</taxonomicName>
and
<taxonomicName authorityName="Selys" authorityYear="1862" class="Insecta" family="Megapodagrionidae" genus="Heteragrion" kingdom="Animalia" order="Odonata" pageId="7" pageNumber="8" phylum="Arthropoda" rank="genus">
<emphasis bold="true" italics="true" pageId="7" pageNumber="8">Heteragrion</emphasis>
</taxonomicName>
form a monophyletic group (BS = 100, PP = 1, QS = 1/NA/1) suggesting that
<emphasis bold="true" box="[983,1085,733,752]" italics="true" pageId="7" pageNumber="8">Dimeragion</emphasis>
and
<taxonomicName authorityName="Selys" authorityYear="1885" box="[1148,1295,733,752]" class="Insecta" family="Megapodagrionidae" genus="Heteropodagrion" kingdom="Animalia" order="Odonata" pageId="7" pageNumber="8" phylum="Arthropoda" rank="genus">
<emphasis bold="true" box="[1148,1295,733,752]" italics="true" pageId="7" pageNumber="8">Heteropodagrion</emphasis>
</taxonomicName>
indeed should be included as members of
<taxonomicName authorityName="Racenis" authorityYear="1959" box="[1061,1216,761,780]" class="Insecta" family="Heteragrionidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="7" pageNumber="8" phylum="Arthropoda" rank="family">Heteragrionidae</taxonomicName>
. In this new definition, the
<taxonomicName authorityName="Racenis" authorityYear="1959" box="[818,970,789,808]" class="Insecta" family="Heteragrionidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="7" pageNumber="8" phylum="Arthropoda" rank="family">Heteragrionidae</taxonomicName>
include four genera from tropical South-American:
<taxonomicName authorityName="Calvert" authorityYear="1913" box="[818,928,817,836]" class="Insecta" family="Megapodagrionidae" genus="Dimeragrion" kingdom="Animalia" order="Odonata" pageId="7" pageNumber="8" phylum="Arthropoda" rank="genus">
<emphasis bold="true" box="[818,928,817,836]" italics="true" pageId="7" pageNumber="8">Dimeragrion</emphasis>
</taxonomicName>
(5 species),
<taxonomicName authorityName="Selys" authorityYear="1862" box="[1060,1164,817,836]" class="Insecta" family="Megapodagrionidae" genus="Heteragrion" kingdom="Animalia" order="Odonata" pageId="7" pageNumber="8" phylum="Arthropoda" rank="genus">
<emphasis bold="true" box="[1060,1164,817,836]" italics="true" pageId="7" pageNumber="8">Heteragrion</emphasis>
</taxonomicName>
(56 species),
<taxonomicName authorityName="Selys" authorityYear="1885" box="[1309,1456,817,836]" class="Insecta" family="Megapodagrionidae" genus="Heteropodagrion" kingdom="Animalia" order="Odonata" pageId="7" pageNumber="8" phylum="Arthropoda" rank="genus">
<emphasis bold="true" box="[1309,1456,817,836]" italics="true" pageId="7" pageNumber="8">Heteropodagrion</emphasis>
</taxonomicName>
(5 species) and
<taxonomicName authorityName="Selys" authorityYear="1862" box="[952,1046,845,864]" class="Magnoliopsida" family="Fabaceae" genus="Oxystigma" kingdom="Plantae" order="Fabales" pageId="7" pageNumber="8" phylum="Tracheophyta" rank="genus">
<emphasis bold="true" box="[952,1046,845,864]" italics="true" pageId="7" pageNumber="8">Oxystigma</emphasis>
</taxonomicName>
(3 species). With the exception of
<taxonomicName authorityName="Selys" authorityYear="1885" class="Insecta" family="Megapodagrionidae" genus="Heteropodagrion" kingdom="Animalia" order="Odonata" pageId="7" pageNumber="8" phylum="Arthropoda" rank="genus">
<emphasis bold="true" italics="true" pageId="7" pageNumber="8">Heteropodagrion</emphasis>
</taxonomicName>
all these genera have their wings open at rest. The caudal gills of the nymphs of
<taxonomicName authorityName="Selys" authorityYear="1862" box="[978,1082,901,920]" class="Insecta" family="Megapodagrionidae" genus="Heteragrion" kingdom="Animalia" order="Odonata" pageId="7" pageNumber="8" phylum="Arthropoda" rank="genus">
<emphasis bold="true" box="[978,1082,901,920]" italics="true" pageId="7" pageNumber="8">Heteragrion</emphasis>
</taxonomicName>
,
<taxonomicName authorityName="Selys" authorityYear="1885" box="[1093,1240,900,919]" class="Insecta" family="Megapodagrionidae" genus="Heteropodagrion" kingdom="Animalia" order="Odonata" pageId="7" pageNumber="8" phylum="Arthropoda" rank="genus">
<emphasis bold="true" box="[1093,1240,900,919]" italics="true" pageId="7" pageNumber="8">Heteropodagrion</emphasis>
</taxonomicName>
and
<taxonomicName authorityName="Selys" authorityYear="1862" box="[1285,1379,901,920]" class="Magnoliopsida" family="Fabaceae" genus="Oxystigma" kingdom="Plantae" order="Fabales" pageId="7" pageNumber="8" phylum="Tracheophyta" rank="genus">
<emphasis bold="true" box="[1285,1379,901,920]" italics="true" pageId="7" pageNumber="8">Oxystigma</emphasis>
</taxonomicName>
are saccoid witha constriction at about ¾ length witha slender apical filament. The caudal gills of the nymph of
<taxonomicName authorityName="Calvert" authorityYear="1913" box="[1098,1208,957,976]" class="Insecta" family="Megapodagrionidae" genus="Dimeragrion" kingdom="Animalia" order="Odonata" pageId="7" pageNumber="8" phylum="Arthropoda" rank="genus">
<emphasis bold="true" box="[1098,1208,957,976]" italics="true" pageId="7" pageNumber="8">Dimeragrion</emphasis>
</taxonomicName>
are nearly flat (
<bibRefCitation author="De Marmels, J." journalOrPublisher="Boletin de Entomologia Venezolana" pageId="7" pageNumber="8" pagination="27 - 36" part="14" refId="ref17071" refString="De Marmels, J., 1999. A new species of Dimeragrion Calvert 1913 from Pantepui, Venezuela (Odonata: Megapodagrionidae). Boletin de Entomologia Venezolana 14, 27 - 36." title="A new species of Dimeragrion Calvert 1913 from Pantepui, Venezuela (Odonata: Megapodagrionidae)" type="journal article" year="1999">De Marmels, 1999</bibRefCitation>
), but do have a terminal filament and are slightly inflated with a thickened dorsal keel making them three-dimensional (
<bibRefCitation author="Tennessen, K. J." pageId="7" pageNumber="8" refId="ref20211" refString="Tennessen, K. J., 2010. The Madicolous Nymph Of Heteropodagrion Sanguinipes Selys (Odonata: Megapodagrionidae). https: // doi. org / 10.5281 / ZENODO. 196541." title="The Madicolous Nymph Of Heteropodagrion Sanguinipes Selys (Odonata: Megapodagrionidae)" type="book" year="2010">Tennessen, 2010</bibRefCitation>
).
</paragraph>
<paragraph blockId="7.[818,1488,148,1367]" pageId="7" pageNumber="8">
The last family included in this section of the Calopterygoidea is
<taxonomicName authorityName="Munz" authorityYear="1919" box="[818,938,1096,1115]" class="Insecta" family="Polythoridae" kingdom="Animalia" order="Odonata" pageId="7" pageNumber="8" phylum="Arthropoda" rank="family">Polythoridae</taxonomicName>
(BS = 100, PP = 1, QS = 1/NA/1) which has several apomorphies in the nymphal stage such as lateral abdominal gills on the second to seventh segment, dorsal abdominal knobs and swollen caudal gills with angular or finger-like projections. The molecular revision of the family by
<bibRefCitation author="Sanchez Herrera, M. &amp; Beatty, C. &amp; Nunes, R. &amp; Realpe, E. &amp; Salazar, C. &amp; Ware, J. L." box="[953,1237,1208,1227]" journalOrPublisher="Syst. Entomol." pageId="7" pageNumber="8" pagination="56 - 67" part="43" refId="ref19918" refString="Sanchez Herrera, M., Beatty, C., Nunes, R., Realpe, E., Salazar, C., Ware, J. L., 2018. A molecular systematic analysis of the Neotropical banner winged damselflies (Polythoridae: Odonata): Phylogenetic relationships of Polythoridae. Syst. Entomol. 43, 56 - 67." title="A molecular systematic analysis of the Neotropical banner winged damselflies (Polythoridae: Odonata): Phylogenetic relationships of Polythoridae." type="book" year="2018">Sanchez Herrera et al. (2018)</bibRefCitation>
showed that the family is monophyletic. The lateral abdominal gills of the nymphs are reminiscent of those of
<taxonomicName baseAuthorityName="Jacobson &amp; Bianchi" baseAuthorityYear="1905" box="[936,1048,1263,1282]" class="Insecta" family="Euphaeidae" kingdom="Animalia" order="Odonata" pageId="7" pageNumber="8" phylum="Arthropoda" rank="family">Euphaeidae</taxonomicName>
, which has led to the suggestion that these families might be related. Our phylogeny shows clearly that these two are not close and that lateral abdominal gills evolved at least twice within
<taxonomicName authorityName="Selys-Longchamps" authorityYear="1854" box="[885,983,1348,1367]" pageId="7" pageNumber="15" rank="subOrder" subOrder="Zygoptera">Zygoptera</taxonomicName>
.
</paragraph>
</subSubSection>
</treatment>
</document>