treatments-xml/data/03/BC/E1/03BCE124681D4971FF53A15D9EBAFD66.xml

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<mods:title id="C5FBC045F090A38A3DDCE68D4992FDFB">The madicolous nymph of Heteropodagrion sanguinipes Selys (Odonata: Megapodagrionidae)</mods:title>
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<heading id="D0E2E75E681D4978FF53A15D9FB6FCD0" bold="true" box="[151,502,775,801]" fontSize="11" level="1" pageId="1" pageNumber="30" reason="1">
<taxonomicName id="4C152BB1681D4978FF53A15D9FB6FCD0" ID-CoL="4WNL" authority="Selys, 1885" authorityName="Selys" authorityYear="1885" box="[151,502,775,801]" class="Insecta" family="Megapodagrionidae" genus="Heteropodagrion" kingdom="Animalia" order="Odonata" pageId="1" pageNumber="30" phylum="Arthropoda" rank="genus">
<emphasis id="B9618C20681D4978FF53A15D9FB6FCD0" bold="true" box="[151,502,775,801]" pageId="1" pageNumber="30">
<emphasis id="B9618C20681D4978FF53A15D9F26FCD0" bold="true" box="[151,358,775,801]" italics="true" pageId="1" pageNumber="30">Heteropodagrion</emphasis>
<bibRefCitation id="EF842DC3681D4978FEA9A15D9FB6FCD0" author="Selys" box="[365,502,775,801]" pageId="1" pageNumber="38" refString="Selys Longchamps, E. de. (1885) Programme d'une Revision des Agrionines. Comptes rendus de Societe Entomologique de Belgique, 29, cxli - cxlvi." type="book chapter" year="1885">Selys, 1885</bibRefCitation>
</emphasis>
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</paragraph>
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<paragraph id="8BAA5032681D4978FF53A11D9D6CFC5D" blockId="1.[151,1437,839,2026]" pageId="1" pageNumber="30">
<emphasis id="B9618C20681D4978FF53A11D9C36FC90" bold="true" box="[151,630,839,865]" pageId="1" pageNumber="30">Description of final stadium nymph</emphasis>
(
<figureCitation id="132E4CB7681D4978FD43A1129D51FC93" box="[647,785,840,866]" captionStart-0="FIGURE 1" captionStart-1="FIGURES 2 – 11" captionStart-2="FIGURE 12" captionStartId-0="2.[151,255,1640,1664]" captionStartId-1="3.[151,269,1841,1865]" captionStartId-2="4.[151,255,1340,1364]" captionTargetBox-1="[185,1405,287,1799]" captionTargetBox-2="[166,1417,210,1309]" captionTargetId-1="figure@3.[158,1419,271,1806]" captionTargetId-2="figure@4.[151,1436,197,1316]" captionTargetPageId-1="3" captionTargetPageId-2="4" captionText-0="FIGURE 1. Heteropodagrion sanguinipes. Habitus" captionText-1="FIGURES 2 – 11. Heteropodagrion sanguinipes. 2) Right antenna; 3) Prementum, dorsal view; 4 a) ridges of prementum, dorsal view; 4 b) ridges of prementum at 45 ° angle; 5) Ligula; 6) Right palpal lobe; 7 a) Right mandible, ental view; 7 b) Left mandible, ental view; 8) Left hind femur; 9) Male cercus (c), lateral view (p = base of paraproct); 10) S 8 – 10, female, lateral view; 11) Left lateral gill, lateral view." captionText-2="FIGURE 12. Heteropodagrion sanguinipes. Posture of live nymph when disturbed." httpUri-1="https://zenodo.org/record/196542/files/figure.png" httpUri-2="https://zenodo.org/record/196543/files/figure.png" pageId="1" pageNumber="30">Figs. 1–12</figureCitation>
, 
<figureCitation id="132E4CB7681D4978FCDBA1129D02FC93" box="[799,834,840,866]" captionStart="FIGURE 14" captionStartId="6.[151,257,1737,1761]" captionTargetBox="[151,1436,197,1711]" captionTargetId="figure@6.[151,1436,197,1713]" captionTargetPageId="6" captionText="FIGURE 14. Live nymph of Heteropodagrion sanguinipes oriented head down on vertical rock face in sheet flow (photo by J. Johnson)." httpUri="https://zenodo.org/record/196545/files/figure.png" pageId="1" pageNumber="30">14</figureCitation>
). Stocky, compact body with short legs bearing numerous long hair-like setae and conspicuous patches of shorter, stout setae on dorsum of abdomen, and saccoid, setose caudal gills with filamentous tip (
<figureCitation id="132E4CB7681D4978FD12A1C89D5BFC5D" box="[726,795,914,940]" captionStart="FIGURE 1" captionStartId="2.[151,255,1640,1664]" captionText="FIGURE 1. Heteropodagrion sanguinipes. Habitus" pageId="1" pageNumber="30">Fig. 1</figureCitation>
).
</paragraph>
<paragraph id="8BAA5032681D4978FF02A1E29CF6F915" blockId="1.[151,1437,839,2026]" pageId="1" pageNumber="30">
<emphasis id="B9618C20681D4978FF02A1E29F46FC20" box="[198,262,952,977]" italics="true" pageId="1" pageNumber="30">Head</emphasis>
. 1.90–2.05 times as wide as long, 1.13–1.17 times wider than widest point of synthorax (at mesepimeral processes above the mesocoxae). Postero-lateral lobes rounded, each bearing numerous small curved setae 
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long; dorsum of epicranium appearing granular; ventral carina of occiput near occipital foramen beset with numerous long pale setae that are aligned along lateral margins of withdrawn prementum. Antenna in life and in preserved specimens curved outward, then inward, as shown in fig.1; usually 7-segmented (
<figureCitation id="132E4CB7681D4978FE5EA6289FA3FB7D" box="[410,483,1138,1164]" captionStart="FIGURES 2 – 11" captionStartId="3.[151,269,1841,1865]" captionTargetBox="[185,1405,287,1799]" captionTargetId="figure@3.[158,1419,271,1806]" captionTargetPageId="3" captionText="FIGURES 2 – 11. Heteropodagrion sanguinipes. 2) Right antenna; 3) Prementum, dorsal view; 4 a) ridges of prementum, dorsal view; 4 b) ridges of prementum at 45 ° angle; 5) Ligula; 6) Right palpal lobe; 7 a) Right mandible, ental view; 7 b) Left mandible, ental view; 8) Left hind femur; 9) Male cercus (c), lateral view (p = base of paraproct); 10) S 8 – 10, female, lateral view; 11) Left lateral gill, lateral view." httpUri="https://zenodo.org/record/196542/files/figure.png" pageId="1" pageNumber="30">Fig. 2</figureCitation>
), rarely with 6 or 8 antennomeres; Antm1 nearly as long as Antm2; Antm2 only 1.2 times longer than wide, slightly longer than remaining antennomeres. Eyes large, in life light gray with darker mottling (
<figureCitation id="132E4CB7681D4978FE9AA6E79FF5FB26" box="[350,437,1213,1239]" captionStart="FIGURE 14" captionStartId="6.[151,257,1737,1761]" captionTargetBox="[151,1436,197,1711]" captionTargetId="figure@6.[151,1436,197,1713]" captionTargetPageId="6" captionText="FIGURE 14. Live nymph of Heteropodagrion sanguinipes oriented head down on vertical rock face in sheet flow (photo by J. Johnson)." httpUri="https://zenodo.org/record/196545/files/figure.png" pageId="1" pageNumber="30">Fig. 14</figureCitation>
), in preserved specimens dark gray with slightly darker spots, ventro-lateral margins with a row of small, densely arranged, curved setae, below which are a few very fine, longer setae. Labium: articulation of pre- and post-mentum rests between bases of coxae I; prementum almost circular (
<figureCitation id="132E4CB7681D4978FA85A7529BCBFAD3" box="[1345,1419,1288,1314]" captionStart="FIGURES 2 – 11" captionStartId="3.[151,269,1841,1865]" captionTargetBox="[185,1405,287,1799]" captionTargetId="figure@3.[158,1419,271,1806]" captionTargetPageId="3" captionText="FIGURES 2 – 11. Heteropodagrion sanguinipes. 2) Right antenna; 3) Prementum, dorsal view; 4 a) ridges of prementum, dorsal view; 4 b) ridges of prementum at 45 ° angle; 5) Ligula; 6) Right palpal lobe; 7 a) Right mandible, ental view; 7 b) Left mandible, ental view; 8) Left hind femur; 9) Male cercus (c), lateral view (p = base of paraproct); 10) S 8 – 10, female, lateral view; 11) Left lateral gill, lateral view." httpUri="https://zenodo.org/record/196542/files/figure.png" pageId="1" pageNumber="30">Fig. 3</figureCitation>
), lateral margins convex, ratio of length to maximum width 0.96–1.00; a long, slightly incurved row of very small transverse ridges on each side, tightly aligned; at high magnification, about 4 ridges per 
<quantity id="4CEDFDD7681D4978FB3BA7089B1BFA9D" box="[1279,1371,1362,1388]" metricMagnitude="-4" metricUnit="m" metricValue="1.0" pageId="1" pageNumber="30" unit="mm" value="0.1">0.1 mm</quantity>
(
<figureCitation id="132E4CB7681D4978FAA8A7089EE8FA63" captionStart="FIGURES 2 – 11" captionStartId="3.[151,269,1841,1865]" captionTargetBox="[185,1405,287,1799]" captionTargetId="figure@3.[158,1419,271,1806]" captionTargetPageId="3" captionText="FIGURES 2 – 11. Heteropodagrion sanguinipes. 2) Right antenna; 3) Prementum, dorsal view; 4 a) ridges of prementum, dorsal view; 4 b) ridges of prementum at 45 ° angle; 5) Ligula; 6) Right palpal lobe; 7 a) Right mandible, ental view; 7 b) Left mandible, ental view; 8) Left hind femur; 9) Male cercus (c), lateral view (p = base of paraproct); 10) S 8 – 10, female, lateral view; 11) Left lateral gill, lateral view." httpUri="https://zenodo.org/record/196542/files/figure.png" pageId="1" pageNumber="30">Fig. 4</figureCitation>
a), each ridge a simple raised area of the integument, varying from about 10 to 15 um high (
<figureCitation id="132E4CB7681D4978FACFA7229B16FA63" box="[1291,1366,1400,1426]" captionStart="FIGURES 2 – 11" captionStartId="3.[151,269,1841,1865]" captionTargetBox="[185,1405,287,1799]" captionTargetId="figure@3.[158,1419,271,1806]" captionTargetPageId="3" captionText="FIGURES 2 – 11. Heteropodagrion sanguinipes. 2) Right antenna; 3) Prementum, dorsal view; 4 a) ridges of prementum, dorsal view; 4 b) ridges of prementum at 45 ° angle; 5) Ligula; 6) Right palpal lobe; 7 a) Right mandible, ental view; 7 b) Left mandible, ental view; 8) Left hind femur; 9) Male cercus (c), lateral view (p = base of paraproct); 10) S 8 – 10, female, lateral view; 11) Left lateral gill, lateral view." httpUri="https://zenodo.org/record/196542/files/figure.png" pageId="1" pageNumber="30">Fig. 4</figureCitation>
b); no conspicuous dorsal premental setae (no raptorial setae is typical of 
<taxonomicName id="4C152BB1681D4978FC7AA7C79AEEFA46" box="[958,1198,1437,1463]" class="Insecta" family="Megapodagrionidae" kingdom="Animalia" order="Odonata" pageId="1" pageNumber="30" phylum="Arthropoda" rank="family">Megapodagrionidae</taxonomicName>
) but with 8–12 fine setae visible in SEM photos; lateral margins of prementum without spinules, ventral surface scattered with long, hair-like setae slightly medial to lateral margin. Ligula convex, cleft, finely crenulated with a minuscule blade-shaped seta at the base of each notch between crenulations, and a small preapical tooth on each side of cleft (
<figureCitation id="132E4CB7681D4978FF1BA4689F6AF9BD" box="[223,298,1586,1612]" captionStart="FIGURES 2 – 11" captionStartId="3.[151,269,1841,1865]" captionTargetBox="[185,1405,287,1799]" captionTargetId="figure@3.[158,1419,271,1806]" captionTargetPageId="3" captionText="FIGURES 2 – 11. Heteropodagrion sanguinipes. 2) Right antenna; 3) Prementum, dorsal view; 4 a) ridges of prementum, dorsal view; 4 b) ridges of prementum at 45 ° angle; 5) Ligula; 6) Right palpal lobe; 7 a) Right mandible, ental view; 7 b) Left mandible, ental view; 8) Left hind femur; 9) Male cercus (c), lateral view (p = base of paraproct); 10) S 8 – 10, female, lateral view; 11) Left lateral gill, lateral view." httpUri="https://zenodo.org/record/196542/files/figure.png" pageId="1" pageNumber="30">Fig. 5</figureCitation>
). Labial palp without dorsal setae but ventral surface with a few fine hair-like setae pointing directly ventrally; distal margin below movable hook with 2 large processes and 1 smaller, rounded inner process (
<figureCitation id="132E4CB7681D4978FEC6A4279F0CF966" box="[258,332,1661,1687]" captionStart="FIGURES 2 – 11" captionStartId="3.[151,269,1841,1865]" captionTargetBox="[185,1405,287,1799]" captionTargetId="figure@3.[158,1419,271,1806]" captionTargetPageId="3" captionText="FIGURES 2 – 11. Heteropodagrion sanguinipes. 2) Right antenna; 3) Prementum, dorsal view; 4 a) ridges of prementum, dorsal view; 4 b) ridges of prementum at 45 ° angle; 5) Ligula; 6) Right palpal lobe; 7 a) Right mandible, ental view; 7 b) Left mandible, ental view; 8) Left hind femur; 9) Male cercus (c), lateral view (p = base of paraproct); 10) S 8 – 10, female, lateral view; 11) Left lateral gill, lateral view." httpUri="https://zenodo.org/record/196542/files/figure.png" pageId="1" pageNumber="30">Fig. 6</figureCitation>
); movable hook 0.77–0.88 times length of outer palpal margin. Mandibles with incisor and molar teeth (
<figureCitation id="132E4CB7681D4978FEE9A4FF9FC1F94E" box="[301,385,1701,1727]" captionStart="FIGURES 2 – 11" captionStartId="3.[151,269,1841,1865]" captionTargetBox="[185,1405,287,1799]" captionTargetId="figure@3.[158,1419,271,1806]" captionTargetPageId="3" captionText="FIGURES 2 – 11. Heteropodagrion sanguinipes. 2) Right antenna; 3) Prementum, dorsal view; 4 a) ridges of prementum, dorsal view; 4 b) ridges of prementum at 45 ° angle; 5) Ligula; 6) Right palpal lobe; 7 a) Right mandible, ental view; 7 b) Left mandible, ental view; 8) Left hind femur; 9) Male cercus (c), lateral view (p = base of paraproct); 10) S 8 – 10, female, lateral view; 11) Left lateral gill, lateral view." httpUri="https://zenodo.org/record/196542/files/figure.png" pageId="1" pageNumber="30">Figs. 7</figureCitation>
a, 7b), formula: 
<emphasis id="B9618C20681D4978FD84A4FC9BD4F94E" box="[576,1428,1698,1727]" italics="true" pageId="1" pageNumber="30">L 1+2345 Oa(m1,2,3)b b' (1&lt;3&lt;2&lt;5&lt;4) / R 1+2345 y a b (1&lt;3&lt;2&lt;4&lt;5)</emphasis>
; 6–8 long curved setae at base on ectal surface.
</paragraph>
<paragraph id="8BAA5032681D4978FF01A4AA9D5AF81B" blockId="1.[151,1437,839,2026]" pageId="1" pageNumber="30">
<emphasis id="B9618C20681D4978FF01A4AA9F5CF8F8" box="[197,284,1776,1801]" italics="true" pageId="1" pageNumber="30">Thorax</emphasis>
. Prothorax with black, triangular epimeral processes extending laterally above coxae (
<figureCitation id="132E4CB7681D4978FA85A4AA9BC9F8FB" box="[1345,1417,1776,1802]" captionStart="FIGURE 1" captionStartId="2.[151,255,1640,1664]" captionText="FIGURE 1. Heteropodagrion sanguinipes. Habitus" pageId="1" pageNumber="30">Fig. 1</figureCitation>
). Wing pads divergent, tips of hind wings reaching postero-laterally to posterior level of S6. Legs laterally depressed (width about 1.4 times thickness), mostly pale but patterned with gray blotches; femur short, apex not extending as far as tip of hind wing pad, and bearing long, hair-like setae and patches of stouter, shorter setae (
<figureCitation id="132E4CB7681D4978FF27A5DF9F6BF86E" box="[227,299,1925,1951]" captionStart="FIGURES 2 – 11" captionStartId="3.[151,269,1841,1865]" captionTargetBox="[185,1405,287,1799]" captionTargetId="figure@3.[158,1419,271,1806]" captionTargetPageId="3" captionText="FIGURES 2 – 11. Heteropodagrion sanguinipes. 2) Right antenna; 3) Prementum, dorsal view; 4 a) ridges of prementum, dorsal view; 4 b) ridges of prementum at 45 ° angle; 5) Ligula; 6) Right palpal lobe; 7 a) Right mandible, ental view; 7 b) Left mandible, ental view; 8) Left hind femur; 9) Male cercus (c), lateral view (p = base of paraproct); 10) S 8 – 10, female, lateral view; 11) Left lateral gill, lateral view." httpUri="https://zenodo.org/record/196542/files/figure.png" pageId="1" pageNumber="30">Fig. 8</figureCitation>
); tibia slightly longer than femur, patterned with gray blotches, bearing several patches of short stout setae, with numerous long hair-like setae on apical 2/3 and shorter, stouter and darker, slightly curved setae on distal 1/5; claw 
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long, 
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wide.
</paragraph>
<caption id="DF6A00BA681E497BFF53A4329CFEF971" box="[151,702,1640,1664]" pageId="2" pageNumber="31">
<paragraph id="8BAA5032681E497BFF53A4329CFEF971" blockId="2.[151,702,1640,1664]" box="[151,702,1640,1664]" pageId="2" pageNumber="31">
<emphasis id="B9618C20681E497BFF53A4329F59F971" bold="true" box="[151,281,1640,1664]" pageId="2" pageNumber="31">FIGURE 1.</emphasis>
<taxonomicName id="4C152BB1681E497BFEE4A4339C1DF971" box="[288,605,1641,1664]" class="Insecta" family="Megapodagrionidae" genus="Heteropodagrion" kingdom="Animalia" order="Odonata" pageId="2" pageNumber="31" phylum="Arthropoda" rank="species" species="sanguinipes">
<emphasis id="B9618C20681E497BFEE4A4339C1DF971" box="[288,605,1641,1664]" italics="true" pageId="2" pageNumber="31">Heteropodagrion sanguinipes</emphasis>
</taxonomicName>
. Habitus
</paragraph>
</caption>
<paragraph id="8BAA5032681E497AFF01A4F49C03FF26" blockId="2.[151,1437,1710,2035]" lastBlockId="3.[151,1436,152,215]" lastPageId="3" lastPageNumber="32" pageId="2" pageNumber="31">
<emphasis id="B9618C20681E497BFF01A4F49F74F936" box="[197,308,1710,1735]" italics="true" pageId="2" pageNumber="31">Abdomen</emphasis>
. Cylindrical dorsally but flattened ventrally; patterned in dark markings shown in fig. 1, giving the appearance of a median pale stripe; S3–10 with dorsal tufts of curved setae forming an ill-defined setal row situated on dark markings on each side of median stripe (
<figureCitation id="132E4CB7681E497BFCA6A4A39DE6F8E2" box="[866,934,1785,1811]" captionStart="FIGURE 1" captionStartId="2.[151,255,1640,1664]" captionText="FIGURE 1. Heteropodagrion sanguinipes. Habitus" pageId="2" pageNumber="31">Fig. 1</figureCitation>
). Lateral margins of S1–10 with numerous long hair-like setae; medial reddish markings on sometimes present on sternites S3–5 and S7–8. S3–8 with lateral slightly concave circular areas. Male cercus sub-rectangular with bluntly rounded apex (
<figureCitation id="132E4CB7681E497BFB2BA5199B76F8AC" box="[1263,1334,1859,1885]" captionStart="FIGURES 2 – 11" captionStartId="3.[151,269,1841,1865]" captionTargetBox="[185,1405,287,1799]" captionTargetId="figure@3.[158,1419,271,1806]" captionTargetPageId="3" captionText="FIGURES 2 – 11. Heteropodagrion sanguinipes. 2) Right antenna; 3) Prementum, dorsal view; 4 a) ridges of prementum, dorsal view; 4 b) ridges of prementum at 45 ° angle; 5) Ligula; 6) Right palpal lobe; 7 a) Right mandible, ental view; 7 b) Left mandible, ental view; 8) Left hind femur; 9) Male cercus (c), lateral view (p = base of paraproct); 10) S 8 – 10, female, lateral view; 11) Left lateral gill, lateral view." httpUri="https://zenodo.org/record/196542/files/figure.png" pageId="2" pageNumber="31">Fig. 9</figureCitation>
), female cercus triangular (
<figureCitation id="132E4CB7681E497BFEB4A5339F89F872" box="[368,457,1897,1923]" captionStart="FIGURES 2 – 11" captionStartId="3.[151,269,1841,1865]" captionTargetBox="[185,1405,287,1799]" captionTargetId="figure@3.[158,1419,271,1806]" captionTargetPageId="3" captionText="FIGURES 2 – 11. Heteropodagrion sanguinipes. 2) Right antenna; 3) Prementum, dorsal view; 4 a) ridges of prementum, dorsal view; 4 b) ridges of prementum at 45 ° angle; 5) Ligula; 6) Right palpal lobe; 7 a) Right mandible, ental view; 7 b) Left mandible, ental view; 8) Left hind femur; 9) Male cercus (c), lateral view (p = base of paraproct); 10) S 8 – 10, female, lateral view; 11) Left lateral gill, lateral view." httpUri="https://zenodo.org/record/196542/files/figure.png" pageId="2" pageNumber="31">Fig. 10</figureCitation>
). Male gonapophyses small, rounded, low tubercles; tips of female gonapophyses extending slightly beyond posterior margin of S10 (
<figureCitation id="132E4CB7681E497BFCC2A5D49D1FF859" box="[774,863,1934,1960]" captionStart="FIGURES 2 – 11" captionStartId="3.[151,269,1841,1865]" captionTargetBox="[185,1405,287,1799]" captionTargetId="figure@3.[158,1419,271,1806]" captionTargetPageId="3" captionText="FIGURES 2 – 11. Heteropodagrion sanguinipes. 2) Right antenna; 3) Prementum, dorsal view; 4 a) ridges of prementum, dorsal view; 4 b) ridges of prementum at 45 ° angle; 5) Ligula; 6) Right palpal lobe; 7 a) Right mandible, ental view; 7 b) Left mandible, ental view; 8) Left hind femur; 9) Male cercus (c), lateral view (p = base of paraproct); 10) S 8 – 10, female, lateral view; 11) Left lateral gill, lateral view." httpUri="https://zenodo.org/record/196542/files/figure.png" pageId="2" pageNumber="31">Fig. 10</figureCitation>
), ventral margin of outer valves with numerous fine setae. S10 without dorso-medial excision. Caudal gills saccoid (
<figureCitation id="132E4CB7681E497BFC72A5E99A47F83C" box="[950,1031,1971,1997]" captionStart="FIGURE 1" captionStartId="2.[151,255,1640,1664]" captionText="FIGURE 1. Heteropodagrion sanguinipes. Habitus" pageId="2" pageNumber="31">Figs. 1</figureCitation>
, 
<figureCitation id="132E4CB7681E497BFBD1A5E99A72F83C" box="[1045,1074,1971,1997]" captionStart="FIGURES 2 – 11" captionStartId="3.[151,269,1841,1865]" captionTargetBox="[185,1405,287,1799]" captionTargetId="figure@3.[158,1419,271,1806]" captionTargetPageId="3" captionText="FIGURES 2 – 11. Heteropodagrion sanguinipes. 2) Right antenna; 3) Prementum, dorsal view; 4 a) ridges of prementum, dorsal view; 4 b) ridges of prementum at 45 ° angle; 5) Ligula; 6) Right palpal lobe; 7 a) Right mandible, ental view; 7 b) Left mandible, ental view; 8) Left hind femur; 9) Male cercus (c), lateral view (p = base of paraproct); 10) S 8 – 10, female, lateral view; 11) Left lateral gill, lateral view." httpUri="https://zenodo.org/record/196542/files/figure.png" pageId="2" pageNumber="31">11</figureCitation>
) but with elongate depressions on each side of dorso-medial ridge and with ventral surface concave; gills slightly more than half length of abdomen, heavily setose, and with constriction (at about ¾ length) demarcating base of narrowly tapering, flexible apical filament (
<figureCitation id="132E4CB7681F497AFE72A2E79C47FF26" box="[438,519,189,215]" captionStart="FIGURE 1" captionStartId="2.[151,255,1640,1664]" captionText="FIGURE 1. Heteropodagrion sanguinipes. Habitus" pageId="3" pageNumber="32">Figs. 1</figureCitation>
, 
<figureCitation id="132E4CB7681F497AFDD2A2E79C75FF26" box="[534,565,189,215]" captionStart="FIGURES 2 – 11" captionStartId="3.[151,269,1841,1865]" captionTargetBox="[185,1405,287,1799]" captionTargetId="figure@3.[158,1419,271,1806]" captionTargetPageId="3" captionText="FIGURES 2 – 11. Heteropodagrion sanguinipes. 2) Right antenna; 3) Prementum, dorsal view; 4 a) ridges of prementum, dorsal view; 4 b) ridges of prementum at 45 ° angle; 5) Ligula; 6) Right palpal lobe; 7 a) Right mandible, ental view; 7 b) Left mandible, ental view; 8) Left hind femur; 9) Male cercus (c), lateral view (p = base of paraproct); 10) S 8 – 10, female, lateral view; 11) Left lateral gill, lateral view." httpUri="https://zenodo.org/record/196542/files/figure.png" pageId="3" pageNumber="32">11</figureCitation>
).
</paragraph>
<caption id="DF6A00BA681F497AFF53A56B9C91F858" httpUri="https://zenodo.org/record/196542/files/figure.png" pageId="3" pageNumber="32" targetBox="[185,1405,287,1799]" targetPageId="3">
<paragraph id="8BAA5032681F497AFF53A56B9C91F858" blockId="3.[151,1437,1841,1961]" pageId="3" pageNumber="32">
<emphasis id="B9618C20681F497AFF53A56B9F0DF8B8" bold="true" box="[151,333,1841,1865]" pageId="3" pageNumber="32">FIGURES 2–11.</emphasis>
<taxonomicName id="4C152BB1681F497AFE91A5689CD3F8B8" box="[341,659,1842,1865]" class="Insecta" family="Megapodagrionidae" genus="Heteropodagrion" kingdom="Animalia" order="Odonata" pageId="3" pageNumber="32" phylum="Arthropoda" rank="species" species="sanguinipes">
<emphasis id="B9618C20681F497AFE91A5689CD3F8B8" box="[341,659,1842,1865]" italics="true" pageId="3" pageNumber="32">Heteropodagrion sanguinipes</emphasis>
</taxonomicName>
. 2) Right antenna; 3) Prementum, dorsal view; 4a) ridges of prementum, dorsal view; 4b) ridges of prementum at 45° angle; 5) Ligula; 6) Right palpal lobe; 7a) Right mandible, ental view; 7b) Left mandible, ental view; 8) Left hind femur; 9) Male cercus (c), lateral view (p = base of paraproct); 10) S8–10, female, lateral view; 11) Left lateral gill, lateral view.
</paragraph>
</caption>
<caption id="DF6A00BA6818497DFF53A7669A57FAA2" box="[151,1047,1340,1364]" httpUri="https://zenodo.org/record/196543/files/figure.png" pageId="4" pageNumber="33" targetBox="[166,1417,210,1309]" targetPageId="4">
<paragraph id="8BAA50326818497DFF53A7669A57FAA2" blockId="4.[151,1047,1340,1364]" box="[151,1047,1340,1364]" pageId="4" pageNumber="33">
<emphasis id="B9618C206818497DFF53A7669F66FAA5" bold="true" box="[151,294,1340,1364]" pageId="4" pageNumber="33">FIGURE 12.</emphasis>
<taxonomicName id="4C152BB16818497DFEE9A7669C2BFAA2" box="[301,619,1340,1363]" class="Insecta" family="Megapodagrionidae" genus="Heteropodagrion" kingdom="Animalia" order="Odonata" pageId="4" pageNumber="33" phylum="Arthropoda" rank="species" species="sanguinipes">
<emphasis id="B9618C206818497DFEE9A7669C2BFAA2" box="[301,619,1340,1363]" italics="true" pageId="4" pageNumber="33">Heteropodagrion sanguinipes</emphasis>
</taxonomicName>
. Posture of live nymph when disturbed.
</paragraph>
</caption>
<paragraph id="8BAA50326818497DFF01A7DB9A71F9FD" blockId="4.[151,1437,1409,2033]" pageId="4" pageNumber="33">
<emphasis id="B9618C206818497DFF01A7DB9C10FA6A" bold="true" box="[197,592,1409,1436]" pageId="4" pageNumber="33">
Measurements, 
<taxonomicName id="4C152BB16818497DFE49A7D89C06FA6D" box="[397,582,1410,1436]" class="Insecta" family="Megapodagrionidae" genus="Heteropodagrion" kingdom="Animalia" order="Odonata" pageId="4" pageNumber="33" phylum="Arthropoda" rank="species" species="sanguinipes">
<emphasis id="B9618C206818497DFE49A7D89C06FA6D" bold="true" box="[397,582,1410,1436]" italics="true" pageId="4" pageNumber="33">H. sanguinipes</emphasis>
</taxonomicName>
:
</emphasis>
(males N=7, females N=4). Total length 8.75–11.5. Head length 1.67– 1.85, maximum width 3.40–3.60. Prementum length 1.90–1.94, maximum width 1.92–2.00. 
<collectingCountry id="F30210A26818497DFACAA7FD9B0BFA30" box="[1294,1355,1447,1473]" name="India" pageId="4" pageNumber="33">Hind</collectingCountry>
femur length 2.55–2.70; hind tibia length 2.75–3.02. Abdomen length 3.8–6.5; cercus length 3 0.44–0.48, Ƥ 0.42– 0.43. Median caudal gill length 3.05–3.40; lateral caudal gill length 2.60–2.95.
</paragraph>
<paragraph id="8BAA50326818497DFF01A44D9DFBF9A7" blockId="4.[151,1437,1409,2033]" pageId="4" pageNumber="33">
F-1 measurements: Total length 7.15 
<emphasis id="B9618C206818497DFD61A44D9CF4F9C1" box="[677,692,1559,1584]" italics="true" pageId="4" pageNumber="33">–</emphasis>
7.40, head width 2.68–2.72, prementum length 1.52–1.54, prementum width 1.52–1.54, median gill 2.45, lateral gill 2.10–2.15.
</paragraph>
<paragraph id="8BAA50326818497DFF01A43B9DB6F91D" blockId="4.[151,1437,1409,2033]" pageId="4" pageNumber="33">
<emphasis id="B9618C206818497DFF01A43B9CD0F98A" bold="true" box="[197,656,1633,1660]" pageId="4" pageNumber="33">
Measurements, 
<taxonomicName id="4C152BB16818497DFE4BA4389C22F98D" box="[399,610,1634,1660]" class="Insecta" family="Megapodagrionidae" genus="Heteropodagrion" kingdom="Animalia" order="Odonata" pageId="4" pageNumber="33" phylum="Arthropoda" rank="genus">
<emphasis id="B9618C206818497DFE4BA4389C22F98D" bold="true" box="[399,610,1634,1660]" italics="true" pageId="4" pageNumber="33">Heteropodagrion</emphasis>
</taxonomicName>
sp:
</emphasis>
(males N=1, females N=2). Total length 9.60–9.90. Head length 1.75–1.83, maximum width 3.50–3.65. Prementum length 1.94–2.01, maximum width 2.01–2.09. 
<collectingCountry id="F30210A26818497DFAD0A4DD9B0FF950" box="[1300,1359,1671,1697]" name="India" pageId="4" pageNumber="33">Hind</collectingCountry>
femur length 2.70–2.85; hind tibia length 2.94–3.09. Abdomen length 4.4–4.9; cercus length 3 0.42–0.43, Ƥ 0.24– 0.32. Median caudal gill length 3.9–4.0; lateral caudal gill length 3.3–3.4.
</paragraph>
<paragraph id="8BAA50326818497DFF01A4AD9CEAF8C7" blockId="4.[151,1437,1409,2033]" pageId="4" pageNumber="33">F-1 measurements: Total length 7.2–8.3, head width 3.1–3.8, prementum length 1.62, prementum width 1.65, median gill 3.2–3.4, lateral gill 2.6–2.8.</paragraph>
</subSubSection>
<subSubSection id="C30F03B968184971FF02A51B9EBAFD66" lastPageId="8" lastPageNumber="37" pageId="4" pageNumber="33" type="discussion">
<paragraph id="8BAA50326818497CFF02A51B9EBAFE46" blockId="4.[151,1437,1409,2033]" lastBlockId="5.[151,1437,152,924]" lastPageId="5" lastPageNumber="34" pageId="4" pageNumber="33">
<emphasis id="B9618C206818497DFF02A51B9C4FF8AA" bold="true" box="[198,527,1857,1883]" pageId="4" pageNumber="33">Discussion and diagnosis.</emphasis>
<taxonomicName id="4C152BB16818497DFDD9A5189CA7F8AA" box="[541,743,1858,1883]" class="Insecta" family="Megapodagrionidae" genus="Heteropodagrion" kingdom="Animalia" order="Odonata" pageId="4" pageNumber="33" phylum="Arthropoda" rank="genus">
<emphasis id="B9618C206818497DFDD9A5189CA7F8AA" box="[541,743,1858,1883]" italics="true" pageId="4" pageNumber="33">Heteropodagrion</emphasis>
</taxonomicName>
is in need of critical taxonomic study, as adults of 
<taxonomicName id="4C152BB16818497DFABBA5189F63F871" class="Insecta" family="Megapodagrionidae" genus="Heteropodagrion" kingdom="Animalia" order="Odonata" pageId="4" pageNumber="33" phylum="Arthropoda" rank="species" species="sanguinipes">
<emphasis id="B9618C206818497DFABBA5189F63F871" italics="true" pageId="4" pageNumber="33">H. sanguinipes</emphasis>
</taxonomicName>
appear to be variable in color pattern and morphology of the cerci (Garrison, pers. comm.). From the two described species, it has been suggested that 
<taxonomicName id="4C152BB16818497DFCC9A5D79AC4F857" authority="Ris" authorityName="Ris" box="[781,1156,1932,1958]" class="Insecta" family="Megapodagrionidae" genus="Heteropodagrion" kingdom="Animalia" order="Odonata" pageId="4" pageNumber="33" phylum="Arthropoda" rank="species" species="superbum">
<emphasis id="B9618C206818497DFCC9A5D79A15F857" box="[781,1109,1933,1958]" italics="true" pageId="4" pageNumber="33">Heteropodagrion superbum</emphasis>
Ris
</taxonomicName>
could be a synonym of 
<taxonomicName id="4C152BB16818497DFF53A5E89C11F83D" authority="Garrison et al. 2010" authorityName="Garrison et al." authorityYear="2010" box="[151,593,1970,1996]" class="Insecta" family="Megapodagrionidae" genus="Heteropodagrion" kingdom="Animalia" order="Odonata" pageId="4" pageNumber="33" phylum="Arthropoda" rank="species" species="sanguinipes">
<emphasis id="B9618C206818497DFF53A5E89F07F83A" box="[151,327,1970,1995]" italics="true" pageId="4" pageNumber="33">H. sanguinipes</emphasis>
(Garrison 
<emphasis id="B9618C206818497DFE02A5E89C42F83A" box="[454,514,1970,1995]" italics="true" pageId="4" pageNumber="33">et al.</emphasis>
2010)
</taxonomicName>
. It is possible that an additional taxon is involved in the samples I took, as adult males taken in Bolivar Province appear to be distinct from 
<taxonomicName id="4C152BB16818497DFC27A58D9ADCF801" box="[995,1180,2007,2032]" class="Insecta" family="Megapodagrionidae" genus="Heteropodagrion" kingdom="Animalia" order="Odonata" pageId="4" pageNumber="33" phylum="Arthropoda" rank="species" species="sanguinipes">
<emphasis id="B9618C206818497DFC27A58D9ADCF801" box="[995,1180,2007,2032]" italics="true" pageId="4" pageNumber="33">H. sanguinipes</emphasis>
</taxonomicName>
. Nymphs of the two populations are basically identical in general appearance and morphological details; however, I observed several differences: The molar crest of the right mandible of 
<taxonomicName id="4C152BB16819497CFCB3A2E79A6BFF27" box="[887,1067,189,214]" class="Insecta" family="Megapodagrionidae" genus="Heteropodagrion" kingdom="Animalia" order="Odonata" pageId="5" pageNumber="34" phylum="Arthropoda" rank="species" species="sanguinipes">
<emphasis id="B9618C206819497CFCB3A2E79A6BFF27" box="[887,1067,189,214]" italics="true" pageId="5" pageNumber="34">H. sanguinipes</emphasis>
</taxonomicName>
has teeth “a” and “b” equal in length, whereas in 
<emphasis id="B9618C206819497CFEB7A2B89FD0FF0A" box="[371,400,226,251]" italics="true" pageId="5" pageNumber="34">H.</emphasis>
sp., tooth “a” is longer than tooth “b”. The movable hook in 
<taxonomicName id="4C152BB16819497CFB92A2B89B45FF0A" box="[1110,1285,226,251]" class="Insecta" family="Megapodagrionidae" genus="Heteropodagrion" kingdom="Animalia" order="Odonata" pageId="5" pageNumber="34" phylum="Arthropoda" rank="species" species="sanguinipes">
<emphasis id="B9618C206819497CFB92A2B89B45FF0A" box="[1110,1285,226,251]" italics="true" pageId="5" pageNumber="34">H. sanguinipes</emphasis>
</taxonomicName>
is 0.77–0.80 the length of the palpal lobe vs. 
<quantity id="4CEDFDD76819497CFDD3A3529CECFED3" box="[535,684,264,290]" metricMagnitude="-2" metricUnit="m" metricValue="2.2098" metricValueMax="2.2352" metricValueMin="2.1844" pageId="5" pageNumber="34" unit="in" value="0.87" valueMax="0.88" valueMin="0.86">0.86–0.88 in</quantity>
<emphasis id="B9618C206819497CFD70A3529C91FED0" box="[692,721,264,289]" italics="true" pageId="5" pageNumber="34">H.</emphasis>
sp. The hind femora of 
<taxonomicName id="4C152BB16819497CFC37A3529AE5FED0" box="[1011,1189,264,289]" class="Insecta" family="Megapodagrionidae" genus="Heteropodagrion" kingdom="Animalia" order="Odonata" pageId="5" pageNumber="34" phylum="Arthropoda" rank="species" species="sanguinipes">
<emphasis id="B9618C206819497CFC37A3529AE5FED0" box="[1011,1189,264,289]" italics="true" pageId="5" pageNumber="34">H. sanguinipes</emphasis>
</taxonomicName>
have only one distal patch of conspicuous stout setae on the dorsal carina vs. three or four in 
<emphasis id="B9618C206819497CFC3EA3779A57FEB7" box="[1018,1047,301,326]" italics="true" pageId="5" pageNumber="34">H.</emphasis>
sp. Lateral gills varied in length from 
<quantity id="4CEDFDD76819497CFF1CA3089F02FE9D" box="[216,322,338,364]" metricMagnitude="-3" metricUnit="m" metricValue="2.58" pageId="5" pageNumber="34" unit="mm" value="2.58">2.58 mm</quantity>
to 
<quantity id="4CEDFDD76819497CFEADA3089F92FE9D" box="[361,466,338,364]" metricMagnitude="-3" metricUnit="m" metricValue="2.95" pageId="5" pageNumber="34" unit="mm" value="2.95">2.95 mm</quantity>
in 
<taxonomicName id="4C152BB16819497CFE3DA3089CEBFE9A" box="[505,683,338,363]" class="Insecta" family="Megapodagrionidae" genus="Heteropodagrion" kingdom="Animalia" order="Odonata" pageId="5" pageNumber="34" phylum="Arthropoda" rank="species" species="sanguinipes">
<emphasis id="B9618C206819497CFE3DA3089CEBFE9A" box="[505,683,338,363]" italics="true" pageId="5" pageNumber="34">H. sanguinipes</emphasis>
</taxonomicName>
vs. 
<quantity id="4CEDFDD76819497CFD19A3089DE1FE9D" box="[733,929,338,364]" metricMagnitude="-3" metricUnit="m" metricValue="3.3499999999999996" metricValueMax="3.4" metricValueMin="3.3" pageId="5" pageNumber="34" unit="mm" value="3.3499999999999996" valueMax="3.4" valueMin="3.3">3.30 to 3.40 mm</quantity>
in 
<emphasis id="B9618C206819497CFC0CA3089DA5FE9A" box="[968,997,338,363]" italics="true" pageId="5" pageNumber="34">H.</emphasis>
sp. In 
<taxonomicName id="4C152BB16819497CFBF3A3089B7CFE9D" authority="Antm" authorityName="Antm" box="[1079,1340,338,364]" class="Insecta" family="Megapodagrionidae" genus="Heteropodagrion" kingdom="Animalia" order="Odonata" pageId="5" pageNumber="34" phylum="Arthropoda" rank="species" species="sanguinipes">
<emphasis id="B9618C206819497CFBF3A3089AA9FE9A" box="[1079,1257,338,363]" italics="true" pageId="5" pageNumber="34">H. sanguinipes</emphasis>
, Antm
</taxonomicName>
2 length was equal to or slightly greater than Antm1 length (ratio 1.00–1.04) vs. shorter than Antm1 (ratio 0.86–0.93) in 
<emphasis id="B9618C206819497CFF71A3C79E92FE47" box="[181,210,413,438]" italics="true" pageId="5" pageNumber="34">H.</emphasis>
sp.
</paragraph>
<paragraph id="8BAA50326819497CFF01A3989CA7FDF3" blockId="5.[151,1437,152,924]" pageId="5" pageNumber="34">
Based on nymphs known to date, the genera of Neotropical 
<taxonomicName id="4C152BB16819497CFC54A3989A3CFE2D" box="[912,1148,450,476]" class="Insecta" family="Megapodagrionidae" kingdom="Animalia" order="Odonata" pageId="5" pageNumber="34" phylum="Arthropoda" rank="family">Megapodagrionidae</taxonomicName>
can be divided into two groups with the following character combinations:
</paragraph>
<paragraph id="8BAA50326819497CFF01A0579C7EFD13" blockId="5.[151,1437,152,924]" pageId="5" pageNumber="34">
Group 1). Antm1 relatively short (length less than half the distance between antennal bases), femora laterally compressed, S6–9 without dorsal hooks, and gills saccoid with a jointed, terminal filament [includes 
<taxonomicName id="4C152BB16819497CFF53A0029F71FD80" box="[151,305,600,625]" class="Insecta" family="Megapodagrionidae" genus="Dimeragrion" kingdom="Animalia" order="Odonata" pageId="5" pageNumber="34" phylum="Arthropoda" rank="genus">
<emphasis id="B9618C206819497CFF53A0029F71FD80" box="[151,305,600,625]" italics="true" pageId="5" pageNumber="34">Dimeragrion</emphasis>
</taxonomicName>
(
<bibRefCitation id="EF842DC36819497CFE84A0029C5BFD83" author="De" box="[320,539,600,626]" pageId="5" pageNumber="37" refString="De Marmels, J. (1999) A new species of Dimeragrion Calvert 1913 from Pantepui, Venezuela (Odonata: Megapodagrionidae). Boletin de Entomologia Venezolana, 14 (1), 27 - 36." type="journal article" year="1999">De Marmels 1999</bibRefCitation>
; pers. comm.), 
<taxonomicName id="4C152BB16819497CFD17A0029D25FD80" box="[723,869,600,625]" class="Insecta" family="Megapodagrionidae" genus="Heteragrion" kingdom="Animalia" order="Odonata" pageId="5" pageNumber="34" phylum="Arthropoda" rank="genus">
<emphasis id="B9618C206819497CFD17A0029D25FD80" box="[723,869,600,625]" italics="true" pageId="5" pageNumber="34">Heteragrion</emphasis>
</taxonomicName>
(
<bibRefCitation id="EF842DC36819497CFCB0A0029A49FD83" author="Santos" box="[884,1033,600,626]" pageId="5" pageNumber="38" refString="Santos, N. D. (1968) Contribuicao ao conhecimento da fauna do Estado da Guanabara. 65. Descricao da ninfa de Heteragrion aurantiacum Selys, 1862 e notas sobre o imago (Odonata: Megapodagrionidae). Atas da Sociedade de Biologia do Rio de Janeiro, 12 (1), 13 - 15." type="journal article" year="1968">Santos 1968</bibRefCitation>
, 
<bibRefCitation id="EF842DC36819497CFBD2A0029B6CFD83" author="Novelo-Gutierrez" box="[1046,1324,600,626]" pageId="5" pageNumber="37" refString="Novelo-Gutierrez, R. (1987) Las nayades de Heteragrion albifrons, H. alienum y H. tricellulare (Odonata: Megapodagrionidae); su descripcion y habitos. Folia Entomologica Mexicana, 73,11 - 22." type="book" year="1987">Novelo-Gutiérrez 1987</bibRefCitation>
, 
<bibRefCitation id="EF842DC36819497CFAFCA0029E97FD66" author="Ramirez" pageId="5" pageNumber="38" refString="Ramirez, A. (1992) Description and natural history of Costa Rican dragonfly larvae. I: Heteragrion erythrogastrum Selys, 1886 (Zygoptera: Megapodagrionidae). Odonatologica, 21, 361 - 365." type="journal article" year="1992">Ramírez 1992</bibRefCitation>
), 
<taxonomicName id="4C152BB16819497CFF28A0279FFBFD67" box="[236,443,637,662]" class="Insecta" family="Megapodagrionidae" genus="Heteropodagrion" kingdom="Animalia" order="Odonata" pageId="5" pageNumber="34" phylum="Arthropoda" rank="genus">
<emphasis id="B9618C206819497CFF28A0279FFBFD67" box="[236,443,637,662]" italics="true" pageId="5" pageNumber="34">Heteropodagrion</emphasis>
</taxonomicName>
, 
<taxonomicName id="4C152BB16819497CFE0FA0279C0EFD67" box="[459,590,637,662]" class="Insecta" family="Megapodagrionidae" genus="Hypolestes" kingdom="Animalia" order="Odonata" pageId="5" pageNumber="34" phylum="Arthropoda" rank="genus">
<emphasis id="B9618C206819497CFE0FA0279C0EFD67" box="[459,590,637,662]" italics="true" pageId="5" pageNumber="34">Hypolestes</emphasis>
</taxonomicName>
(Westfall &amp; 
<date id="FFAB76F26819497CFD28A0279D2BFD66" box="[748,875,637,663]" pageId="5" pageNumber="34" value="2000-05">May 2000</date>
), 
<taxonomicName id="4C152BB16819497CFC45A0279AB9FD66" authority="De Marmels 1987" authorityName="De Marmels" authorityYear="1987" box="[897,1273,637,663]" class="Insecta" family="Megapodagrionidae" genus="Oxystigma" kingdom="Animalia" order="Odonata" pageId="5" pageNumber="34" phylum="Arthropoda" rank="genus">
<emphasis id="B9618C206819497CFC45A0279A41FD67" box="[897,1025,637,662]" italics="true" pageId="5" pageNumber="34">Oxystigma</emphasis>
(
<bibRefCitation id="EF842DC36819497CFBD5A0279AB1FD66" author="De" box="[1041,1265,637,663]" pageId="5" pageNumber="37" refString="De Marmels, J. (1987) On the type specimens of some Neotropical Megapodagrionidae, with a description of Heteragrion pemon spec. nov. and Oxystigma caerulans spec. nov. from Venezuela (Zygoptera). Odonatologica, 16 (3), 225 - 238." type="journal article" year="1987">De Marmels 1987</bibRefCitation>
)
</taxonomicName>
, 
<taxonomicName id="4C152BB16819497CFAC2A0279F83FD4D" authority="Novelo-Gutierrez 2008" authorityName="Novelo-Gutierrez" authorityYear="2008" class="Insecta" family="Megapodagrionidae" genus="Paraphlebia" kingdom="Animalia" order="Odonata" pageId="5" pageNumber="34" phylum="Arthropoda" rank="genus">
<emphasis id="B9618C206819497CFAC2A0279BDCFD67" box="[1286,1436,637,662]" italics="true" pageId="5" pageNumber="34">Paraphlebia</emphasis>
(
<bibRefCitation id="EF842DC36819497CFF5BA0F89FFAFD4D" author="Novelo-Gutierrez" box="[159,442,674,700]" pageId="5" pageNumber="37" refString="Novelo-Gutierrez, R. (2008) Description of the larva of Paraphlebia zoe Selys in Hagen, 1861 (Odonata: Megapodagrionidae). Zootaxa, 1876, 29 - 34." type="journal article" year="2008">Novelo-Gutiérrez 2008</bibRefCitation>
)
</taxonomicName>
, 
<taxonomicName id="4C152BB16819497CFE0BA0F89C11FD4A" box="[463,593,674,699]" class="Insecta" family="Megapodagrionidae" genus="Philogenia" kingdom="Animalia" order="Odonata" pageId="5" pageNumber="34" phylum="Arthropoda" rank="genus">
<emphasis id="B9618C206819497CFE0BA0F89C11FD4A" box="[463,593,674,699]" italics="true" pageId="5" pageNumber="34">Philogenia</emphasis>
</taxonomicName>
(
<bibRefCitation id="EF842DC36819497CFDA6A0F89A13FD4D" author="Ramirez-Ulate" box="[610,1107,674,700]" pageId="5" pageNumber="37" refString="Ramirez-Ulate, A. &amp; Novelo-Gutierrez, R. (1994) Megapodagrionidae (Odonata, Zygoptera) de Mexico y Centro America I: Las nayades de Philogenia carrillica, P. peacocki y P. t e r r a b a. Acta Zoologica Mexicana, 63, 61 - 73." type="journal article" year="1994">Ramírez-Ulate &amp; Novelo-Gutiérrez 1994</bibRefCitation>
; 
<bibRefCitation id="EF842DC36819497CFBA4A0F89BD4FD4D" author="Bybee" box="[1120,1428,674,700]" pageId="5" pageNumber="37" refString="Bybee, S. M. &amp; Tennessen, K. J. (2008) Description of the female and nymph of Philogenia mangosisa from southern Ecuador (Odonata: Megapodagrionidae). Zootaxa, 1787, 63 - 68." type="journal article" year="2008">Bybee &amp; Tennessen 2008</bibRefCitation>
) and 
<taxonomicName id="4C152BB16819497CFF0CA0929C73FD13" authority="De Marmels 2004" authorityName="De Marmels" authorityYear="2004" box="[200,563,712,738]" class="Insecta" family="Megapodagrionidae" genus="Sciotropis" kingdom="Animalia" order="Odonata" pageId="5" pageNumber="34" phylum="Arthropoda" rank="genus">
<emphasis id="B9618C206819497CFF0CA0929F7EFD10" box="[200,318,712,737]" italics="true" pageId="5" pageNumber="34">Sciotropis</emphasis>
(
<bibRefCitation id="EF842DC36819497CFE8AA0929C6AFD13" author="De" box="[334,554,712,738]" pageId="5" pageNumber="37" refString="De Marmels, J. (2004) Heteragrion makiritare sp. nov., with descriptions of hitherto unknown females and larvae of other species from Venezuela (Odonata: Megapodagrionidae, Lestidae). International Journal of Odonatology, 7 (3), 439 - 458." type="journal article" year="2004">De Marmels 2004</bibRefCitation>
)
</taxonomicName>
].
</paragraph>
<paragraph id="8BAA50326819497CFF01A0B79D36FC6D" blockId="5.[151,1437,152,924]" pageId="5" pageNumber="34">
Group 2). Antm1 relatively long (length more than half the distance between antennal bases), femora not laterally compressed, S6–9 with dorsal hooks, and gills elongate, lacking a terminal filament [includes 
<taxonomicName id="4C152BB16819497CFF53A1629F12FCA0" box="[151,338,824,849]" class="Insecta" family="Megapodagrionidae" genus="Teinopodagrion" kingdom="Animalia" order="Odonata" pageId="5" pageNumber="34" phylum="Arthropoda" rank="genus">
<emphasis id="B9618C206819497CFF53A1629F12FCA0" box="[151,338,824,849]" italics="true" pageId="5" pageNumber="34">Teinopodagrion</emphasis>
</taxonomicName>
(
<bibRefCitation id="EF842DC36819497CFEA5A1629C7CFCA3" author="De" box="[353,572,824,850]" pageId="5" pageNumber="37" refString="De Marmels, J. (2001) Revision of Megapodagrion Selys, 1886 (Insecta, Odonata: Megapodagrionidae). Dissertation, University of Zurich, 220 pp." type="book" year="2001">De Marmels 2001</bibRefCitation>
, von 
<bibRefCitation id="EF842DC36819497CFD43A1629D10FCA3" author="Ellenrieder" box="[647,848,824,850]" pageId="5" pageNumber="38" refString="von Ellenrieder, N. (2006) The larvae of Teinopodagrion decipiens De Marmels and T. meridionale De Marmels (Zygoptera: Megapodagrionidae). Odonatologica, 35 (3), 281 - 287." type="journal article" year="2006">Ellenrieder 2006</bibRefCitation>
, 
<bibRefCitation id="EF842DC36819497CFC98A1629A20FCA3" author="Perez-Gutierrez" box="[860,1120,824,850]" pageId="5" pageNumber="37" refString="Perez-Gutierrez, L. A. (2007) The larvae of Teinopodagrion caquetanum De Marmels and T. vallenatum De Marmels (Zygoptera: Megapodagrionidae). Odonatologica, 36 (3), 307 - 313." type="journal article" year="2007">Pérez-Gutiérrez 2007</bibRefCitation>
)]. Although the nymphs of 
<taxonomicName id="4C152BB16819497CFF53A1079F09FC87" box="[151,329,861,886]" class="Insecta" family="Megapodagrionidae" genus="Allopodagrion" kingdom="Animalia" order="Odonata" pageId="5" pageNumber="34" phylum="Arthropoda" rank="genus">
<emphasis id="B9618C206819497CFF53A1079F09FC87" box="[151,329,861,886]" italics="true" pageId="5" pageNumber="34">Allopodagrion</emphasis>
</taxonomicName>
and 
<taxonomicName id="4C152BB16819497CFE41A1079C08FC87" box="[389,584,861,886]" class="Insecta" family="Megapodagrionidae" genus="Megapodagrion" kingdom="Animalia" order="Odonata" pageId="5" pageNumber="34" phylum="Arthropoda" rank="genus">
<emphasis id="B9618C206819497CFE41A1079C08FC87" box="[389,584,861,886]" italics="true" pageId="5" pageNumber="34">Megapodagrion</emphasis>
</taxonomicName>
are unknown, they probably belong in Group 2 as they are related to 
<taxonomicName id="4C152BB16819497CFF53A1D89F11FC6A" box="[151,337,898,923]" class="Insecta" family="Megapodagrionidae" genus="Teinopodagrion" kingdom="Animalia" order="Odonata" pageId="5" pageNumber="34" phylum="Arthropoda" rank="genus">
<emphasis id="B9618C206819497CFF53A1D89F11FC6A" box="[151,337,898,923]" italics="true" pageId="5" pageNumber="34">Teinopodagrion</emphasis>
</taxonomicName>
based on adult characters (
<bibRefCitation id="EF842DC36819497CFD4BA1D89D29FC6D" author="De" box="[655,873,898,924]" pageId="5" pageNumber="37" refString="De Marmels, J. (2001) Revision of Megapodagrion Selys, 1886 (Insecta, Odonata: Megapodagrionidae). Dissertation, University of Zurich, 220 pp." type="book" year="2001">De Marmels 2001</bibRefCitation>
).
</paragraph>
<caption id="DF6A00BA6819497CFF53A5EA9B56F839" box="[151,1302,1968,1992]" httpUri="https://zenodo.org/record/196544/files/figure.png" pageId="5" pageNumber="34" targetBox="[151,1435,981,1943]" targetPageId="5">
<paragraph id="8BAA50326819497CFF53A5EA9B56F839" blockId="5.[151,1302,1968,1992]" box="[151,1302,1968,1992]" pageId="5" pageNumber="34">
<emphasis id="B9618C206819497CFF53A5EA9F66F839" bold="true" box="[151,294,1968,1992]" pageId="5" pageNumber="34">FIGURE 13.</emphasis>
<taxonomicName id="4C152BB16819497CFEE9A5EB9C2BF839" box="[301,619,1969,1992]" class="Insecta" family="Megapodagrionidae" genus="Heteropodagrion" kingdom="Animalia" order="Odonata" pageId="5" pageNumber="34" phylum="Arthropoda" rank="species" species="sanguinipes">
<emphasis id="B9618C206819497CFEE9A5EB9C2BF839" box="[301,619,1969,1992]" italics="true" pageId="5" pageNumber="34">Heteropodagrion sanguinipes</emphasis>
</taxonomicName>
. habitat of 
<taxonomicName id="4C152BB16819497CFD20A5EB9A62F839" box="[740,1058,1969,1992]" class="Insecta" family="Megapodagrionidae" genus="Heteropodagrion" kingdom="Animalia" order="Odonata" pageId="5" pageNumber="34" phylum="Arthropoda" rank="species" species="sanguinipes">
<emphasis id="B9618C206819497CFD20A5EB9A62F839" box="[740,1058,1969,1992]" italics="true" pageId="5" pageNumber="34">Heteropodagrion sanguinipes</emphasis>
</taxonomicName>
(photo by J. Johnson).
</paragraph>
</caption>
<caption id="DF6A00BA681A497FFF53A4939FC3F8F1" httpUri="https://zenodo.org/record/196545/files/figure.png" pageId="6" pageNumber="35" targetBox="[151,1436,197,1711]" targetPageId="6">
<paragraph id="8BAA5032681A497FFF53A4939FC3F8F1" blockId="6.[151,1436,1737,1792]" pageId="6" pageNumber="35">
<emphasis id="B9618C20681A497FFF53A4939F6BF910" bold="true" box="[151,299,1737,1761]" pageId="6" pageNumber="35">FIGURE 14.</emphasis>
Live nymph of 
<taxonomicName id="4C152BB1681A497FFE2CA4939D73F911" box="[488,819,1737,1760]" class="Insecta" family="Megapodagrionidae" genus="Heteropodagrion" kingdom="Animalia" order="Odonata" pageId="6" pageNumber="35" phylum="Arthropoda" rank="species" species="sanguinipes">
<emphasis id="B9618C20681A497FFE2CA4939D73F911" box="[488,819,1737,1760]" italics="true" pageId="6" pageNumber="35">Heteropodagrion sanguinipes</emphasis>
</taxonomicName>
oriented head down on vertical rock face in sheet flow (photo by J. Johnson).
</paragraph>
</caption>
<paragraph id="8BAA5032681A497EFF01A5759A61FF43" blockId="6.[151,1436,1839,2014]" lastBlockId="7.[151,1437,152,2007]" lastPageId="7" lastPageNumber="36" pageId="6" pageNumber="35">
<bibRefCitation id="EF842DC3681A497FFF01A5759F8FF8B8" box="[197,463,1839,1865]" pageId="6" pageNumber="37" refString="Kalkman, V. J., Choong, C. Y., Orr, A. G. &amp; Schutte, K. (2010) Remarks on the taxonomy of Megapodagrionidae with emphasis on the larval gills (Odonata). International Journal of Odonatology 13 (1), 119 - 135." type="journal article">
Kalkman 
<emphasis id="B9618C20681A497FFEFFA5759F38F8B9" box="[315,376,1839,1864]" italics="true" pageId="6" pageNumber="35">et al.</emphasis>
(2010)
</bibRefCitation>
termed 
<taxonomicName id="4C152BB1681A497FFDF7A5759D62F8B8" box="[563,802,1839,1865]" class="Insecta" family="Megapodagrionidae" kingdom="Animalia" order="Odonata" pageId="6" pageNumber="35" phylum="Arthropoda" rank="family">Megapodagrionidae</taxonomicName>
nymphs with saccoid gills “Balloon megapods” and listed all of the genera in Group 1 except two, 
<taxonomicName id="4C152BB1681A497FFD07A50E9D1DF89C" box="[707,861,1876,1901]" class="Insecta" family="Megapodagrionidae" genus="Dimeragrion" kingdom="Animalia" order="Odonata" pageId="6" pageNumber="35" phylum="Arthropoda" rank="genus">
<emphasis id="B9618C20681A497FFD07A50E9D1DF89C" box="[707,861,1876,1901]" italics="true" pageId="6" pageNumber="35">Dimeragrion</emphasis>
</taxonomicName>
and 
<taxonomicName id="4C152BB1681A497FFC5CA50E9A24F89C" box="[920,1124,1876,1901]" class="Insecta" family="Megapodagrionidae" genus="Heteropodagrion" kingdom="Animalia" order="Odonata" pageId="6" pageNumber="35" phylum="Arthropoda" rank="genus">
<emphasis id="B9618C20681A497FFC5CA50E9A24F89C" box="[920,1124,1876,1901]" italics="true" pageId="6" pageNumber="35">Heteropodagrion</emphasis>
</taxonomicName>
, the latter unknown at the time of their publication. 
<taxonomicName id="4C152BB1681A497FFE00A5239C1DF863" box="[452,605,1913,1938]" class="Insecta" family="Megapodagrionidae" genus="Dimeragrion" kingdom="Animalia" order="Odonata" pageId="6" pageNumber="35" phylum="Arthropoda" rank="genus">
<emphasis id="B9618C20681A497FFE00A5239C1DF863" box="[452,605,1913,1938]" italics="true" pageId="6" pageNumber="35">Dimeragrion</emphasis>
</taxonomicName>
was not included because the gills are nearly flat (
<bibRefCitation id="EF842DC3681A497FFB70A5239BCEF862" author="De" box="[1204,1422,1913,1939]" pageId="6" pageNumber="37" refString="De Marmels, J. (1999) A new species of Dimeragrion Calvert 1913 from Pantepui, Venezuela (Odonata: Megapodagrionidae). Boletin de Entomologia Venezolana, 14 (1), 27 - 36." type="journal article" year="1999">De Marmels 1999</bibRefCitation>
); the genus was therefore grouped with the “Fan megapods”. However, I believe that 
<taxonomicName id="4C152BB1681A497FFBBBA5C59B59F849" box="[1151,1305,1951,1976]" class="Insecta" family="Megapodagrionidae" genus="Dimeragrion" kingdom="Animalia" order="Odonata" pageId="6" pageNumber="35" phylum="Arthropoda" rank="genus">
<emphasis id="B9618C20681A497FFBBBA5C59B59F849" box="[1151,1305,1951,1976]" italics="true" pageId="6" pageNumber="35">Dimeragrion</emphasis>
</taxonomicName>
belongs to the “Balloon megapods” because the gills have a terminal filament (not present in “Fan megapods”) and are slightly inflated with a thickened dorsal keel making them three dimensional.
</paragraph>
<paragraph id="8BAA5032681B497EFF01A2E79F12FD13" blockId="7.[151,1437,152,2007]" pageId="7" pageNumber="36">
Several characters separate nymphs of 
<taxonomicName id="4C152BB1681B497EFD78A2E79DD8FF27" box="[700,920,189,214]" class="Insecta" family="Megapodagrionidae" genus="Heteropodagrion" kingdom="Animalia" order="Odonata" pageId="7" pageNumber="36" phylum="Arthropoda" rank="genus">
<emphasis id="B9618C20681B497EFD78A2E79DD8FF27" box="[700,920,189,214]" italics="true" pageId="7" pageNumber="36">Heteropodagrion</emphasis>
</taxonomicName>
from the other genera in the first group (characters for Group 
<quantity id="4CEDFDD7681B497EFE60A2B89F92FF0D" box="[420,466,226,252]" metricMagnitude="-2" metricUnit="m" metricValue="2.54" pageId="7" pageNumber="36" unit="in" value="1.0">1 in</quantity>
parentheses): 1) prementum with a long, slightly curved row of small ridges on each side (without ridges); 2) ventral epicranial carinae near foramen (paralleling sides of prementum) with many long, hair-like setae (without setae or only one or two long setae); 3) Antm1–3 roughly equal in length (with either Antm2 and/or 3 at least twice as long as Antm1; exception 
<taxonomicName id="4C152BB1681B497EFC1BA3089A32FE9A" box="[991,1138,338,363]" class="Insecta" family="Megapodagrionidae" genus="Paraphlebia" kingdom="Animalia" order="Odonata" pageId="7" pageNumber="36" phylum="Arthropoda" rank="genus">
<emphasis id="B9618C20681B497EFC1BA3089A32FE9A" box="[991,1138,338,363]" italics="true" pageId="7" pageNumber="36">Paraphlebia</emphasis>
</taxonomicName>
in which Antm2 is about 1.6 times as long as Antm1 and Antm3 is shorter than Antm1); 4) femora with small, isolated dorsal patches of two or more stout setae (without patches of stout setae); 5) hind femur short, when fully extended posteriorly apex is proximal to tip of hind wing pad of final stadium (extending beyond tip of hind wing pad). 
<taxonomicName id="4C152BB1681B497EFF53A3B29F28FDF0" box="[151,360,488,513]" class="Insecta" family="Megapodagrionidae" genus="Heteropodagrion" kingdom="Animalia" order="Odonata" pageId="7" pageNumber="36" phylum="Arthropoda" rank="genus">
<emphasis id="B9618C20681B497EFF53A3B29F28FDF0" box="[151,360,488,513]" italics="true" pageId="7" pageNumber="36">Heteropodagrion</emphasis>
</taxonomicName>
nymphs are most similar to 
<taxonomicName id="4C152BB1681B497EFD03A3B29D1FFDF0" box="[711,863,488,513]" class="Insecta" family="Megapodagrionidae" genus="Paraphlebia" kingdom="Animalia" order="Odonata" pageId="7" pageNumber="36" phylum="Arthropoda" rank="genus">
<emphasis id="B9618C20681B497EFD03A3B29D1FFDF0" box="[711,863,488,513]" italics="true" pageId="7" pageNumber="36">Paraphlebia</emphasis>
</taxonomicName>
and 
<taxonomicName id="4C152BB1681B497EFC58A3B29A56FDF0" box="[924,1046,488,513]" class="Insecta" family="Megapodagrionidae" genus="Sciotropis" kingdom="Animalia" order="Odonata" pageId="7" pageNumber="36" phylum="Arthropoda" rank="genus">
<emphasis id="B9618C20681B497EFC58A3B29A56FDF0" box="[924,1046,488,513]" italics="true" pageId="7" pageNumber="36">Sciotropis</emphasis>
</taxonomicName>
, sharing the following traits: 1) antennomeres short and thick; 2) gills with many long, hair-like setae from base to apex; and 3) terminal filament relatively short and thick. In 
<taxonomicName id="4C152BB1681B497EFD9DA0689CACFDBA" box="[601,748,562,587]" class="Insecta" family="Megapodagrionidae" genus="Paraphlebia" kingdom="Animalia" order="Odonata" pageId="7" pageNumber="36" phylum="Arthropoda" rank="genus">
<emphasis id="B9618C20681B497EFD9DA0689CACFDBA" box="[601,748,562,587]" italics="true" pageId="7" pageNumber="36">Paraphlebia</emphasis>
</taxonomicName>
, the prementum lacks the rows of small ridges, the dorsal carina of all femora have a singular row of short, stout setae, and the hind femora in the final instar extend beyond the tips of the hind wing pads (
<bibRefCitation id="EF842DC3681B497EFDAEA0279DC7FD66" author="Novelo-Gutierrez" box="[618,903,637,663]" pageId="7" pageNumber="37" refString="Novelo-Gutierrez, R. (2008) Description of the larva of Paraphlebia zoe Selys in Hagen, 1861 (Odonata: Megapodagrionidae). Zootaxa, 1876, 29 - 34." type="journal article" year="2008">Novelo-Gutiérrez 2008</bibRefCitation>
). 
<taxonomicName id="4C152BB1681B497EFC5EA0279A51FD67" box="[922,1041,637,662]" class="Insecta" family="Megapodagrionidae" genus="Sciotropis" kingdom="Animalia" order="Odonata" pageId="7" pageNumber="36" phylum="Arthropoda" rank="genus">
<emphasis id="B9618C20681B497EFC5EA0279A51FD67" box="[922,1041,637,662]" italics="true" pageId="7" pageNumber="36">Sciotropis</emphasis>
</taxonomicName>
is similar to 
<taxonomicName id="4C152BB1681B497EFB74A0279B3CFD67" box="[1200,1404,637,662]" class="Insecta" family="Megapodagrionidae" genus="Heteropodagrion" kingdom="Animalia" order="Odonata" pageId="7" pageNumber="36" phylum="Arthropoda" rank="genus">
<emphasis id="B9618C20681B497EFB74A0279B3CFD67" box="[1200,1404,637,662]" italics="true" pageId="7" pageNumber="36">Heteropodagrion</emphasis>
</taxonomicName>
in most characters, but the prementum is slightly longer than wide and lacks the rows of small ridges (
<bibRefCitation id="EF842DC3681B497EFAB3A0F89F06FD13" author="De" pageId="7" pageNumber="37" refString="De Marmels, J. (2004) Heteragrion makiritare sp. nov., with descriptions of hitherto unknown females and larvae of other species from Venezuela (Odonata: Megapodagrionidae, Lestidae). International Journal of Odonatology, 7 (3), 439 - 458." type="journal article" year="2004">De Marmels 2004</bibRefCitation>
).
</paragraph>
<paragraph id="8BAA5032681B497EFF01A0B79FE0FBC3" blockId="7.[151,1437,152,2007]" pageId="7" pageNumber="36">
The tiny ridges on the prementum do not appear as highly effective aids for holding onto struggling prey. However, the ridges could increase friction and drag on prey when engaging small, segmented organisms, and because of the parallel, longitudinal orientation, could help direct the prey toward the mouth. Several small larvae of 
<taxonomicName id="4C152BB1681B497EFECDA1079FF4FC86" box="[265,436,861,887]" class="Insecta" family="Chironomidae" kingdom="Animalia" order="Diptera" pageId="7" pageNumber="36" phylum="Arthropoda" rank="family">Chironomidae</taxonomicName>
were found in the madicolous microhabitat. Because the nymphs must cling to the rocky surface, I would expect tarsal claws of madicolous nymphs to be short and stout, more so than in nymphs that occupy detritus or sediments in streams. This difference appears to be the case, as length by width ratios of 
<taxonomicName id="4C152BB1681B497EFE89A1979C59FC17" box="[333,537,973,998]" class="Insecta" family="Megapodagrionidae" genus="Heteropodagrion" kingdom="Animalia" order="Odonata" pageId="7" pageNumber="36" phylum="Arthropoda" rank="genus">
<emphasis id="B9618C20681B497EFE89A1979C59FC17" box="[333,537,973,998]" italics="true" pageId="7" pageNumber="36">Heteropodagrion</emphasis>
</taxonomicName>
and 
<taxonomicName id="4C152BB1681B497EFD90A1979D56FC17" box="[596,790,973,998]" class="Insecta" family="Megapodagrionidae" genus="Paraphlebia" kingdom="Animalia" order="Odonata" pageId="7" pageNumber="36" phylum="Arthropoda" rank="species" species="zoe">
<emphasis id="B9618C20681B497EFD90A1979D56FC17" box="[596,790,973,998]" italics="true" pageId="7" pageNumber="36">Paraphlebia zoe</emphasis>
</taxonomicName>
(also an inhabitant of vertical sheet flow, see Novelo- 
<bibRefCitation id="EF842DC3681B497EFF53A1A89F14FBFD" author="Gutierrez" box="[151,340,1010,1036]" pageId="7" pageNumber="37" refString="Novelo-Gutierrez, R. (2008) Description of the larva of Paraphlebia zoe Selys in Hagen, 1861 (Odonata: Megapodagrionidae). Zootaxa, 1876, 29 - 34." type="journal article" year="2008">Gutiérrez 2008</bibRefCitation>
.) varied from 2.0 to 2.4, compared to ratios of 2.8 and 
<quantity id="4CEDFDD7681B497EFC32A1A89A7CFBFD" box="[1014,1084,1010,1036]" metricMagnitude="-2" metricUnit="m" metricValue="8.382" pageId="7" pageNumber="36" unit="in" value="3.3">3.3 in</quantity>
<taxonomicName id="4C152BB1681B497EFB81A1A89A8AFBFA" box="[1093,1226,1010,1035]" class="Insecta" family="Megapodagrionidae" genus="Philogenia" kingdom="Animalia" order="Odonata" pageId="7" pageNumber="36" phylum="Arthropoda" rank="genus">
<emphasis id="B9618C20681B497EFB81A1A89A8AFBFA" box="[1093,1226,1010,1035]" italics="true" pageId="7" pageNumber="36">Philogenia</emphasis>
</taxonomicName>
and 
<taxonomicName id="4C152BB1681B497EFAC2A1A89BDCFBFA" box="[1286,1436,1010,1035]" class="Insecta" family="Megapodagrionidae" genus="Heteragrion" kingdom="Animalia" order="Odonata" pageId="7" pageNumber="36" phylum="Arthropoda" rank="genus">
<emphasis id="B9618C20681B497EFAC2A1A89BDCFBFA" box="[1286,1436,1010,1035]" italics="true" pageId="7" pageNumber="36">Heteragrion</emphasis>
</taxonomicName>
which inhabit streams.
</paragraph>
<paragraph id="8BAA5032681B497EFF01A6679C0AFB36" blockId="7.[151,1437,152,2007]" pageId="7" pageNumber="36">
The generic description of the nymph of 
<taxonomicName id="4C152BB1681B497EFD06A6679DD6FBA7" box="[706,918,1085,1110]" class="Insecta" family="Megapodagrionidae" genus="Heteropodagrion" kingdom="Animalia" order="Odonata" pageId="7" pageNumber="36" phylum="Arthropoda" rank="genus">
<emphasis id="B9618C20681B497EFD06A6679DD6FBA7" box="[706,918,1085,1110]" italics="true" pageId="7" pageNumber="36">Heteropodagrion</emphasis>
</taxonomicName>
is based on two possibly distinct species. Adults of two possible new species are known to exist in 
<collectingCountry id="F30210A2681B497EFC9CA6389DFDFB8D" box="[856,957,1122,1148]" name="Ecuador" pageId="7" pageNumber="36">Ecuador</collectingCountry>
on the eastern side of the Andes (to be studied by J. J. Daigle, pers. comm.); discovery of the nymphs of these species is necessary to confirm or alter the generic characters proposed here.
</paragraph>
<paragraph id="8BAA5032681B497EFF01A6889D4CF87D" blockId="7.[151,1437,152,2007]" pageId="7" pageNumber="36">
Adult 
<taxonomicName id="4C152BB1681B497EFED7A6899FA6FB1D" box="[275,486,1235,1260]" class="Insecta" family="Megapodagrionidae" genus="Heteropodagrion" kingdom="Animalia" order="Odonata" pageId="7" pageNumber="36" phylum="Arthropoda" rank="genus">
<emphasis id="B9618C20681B497EFED7A6899FA6FB1D" box="[275,486,1235,1260]" italics="true" pageId="7" pageNumber="36">Heteropodagrion</emphasis>
</taxonomicName>
perch with wings closed in shade along small montane streams, usually near waterfalls. I am convinced nymphs do not inhabit the actual streams, as I have sampled these habitats for at least 100 man hours without finding 
<taxonomicName id="4C152BB1681B497EFDAFA7479D03FAC7" box="[619,835,1309,1334]" class="Insecta" family="Megapodagrionidae" genus="Heteropodagrion" kingdom="Animalia" order="Odonata" pageId="7" pageNumber="36" phylum="Arthropoda" rank="genus">
<emphasis id="B9618C20681B497EFDAFA7479D03FAC7" box="[619,835,1309,1334]" italics="true" pageId="7" pageNumber="36">Heteropodagrion</emphasis>
</taxonomicName>
nymphs. I finally found the nymphs by closely inspecting near-vertical boulders covered in sheet flow (
<figureCitation id="132E4CB7681B497EFCF5A7189DC8FAAD" box="[817,904,1346,1372]" captionStart="FIGURE 13" captionStartId="5.[151,255,1968,1992]" captionTargetBox="[151,1435,981,1943]" captionTargetId="figure@5.[151,1436,981,1945]" captionTargetPageId="5" captionText="FIGURE 13. Heteropodagrion sanguinipes. habitat of Heteropodagrion sanguinipes (photo by J. Johnson)." httpUri="https://zenodo.org/record/196544/files/figure.png" pageId="7" pageNumber="36">Fig. 13</figureCitation>
). Motionless nymphs were difficult to detect because their color pattern of yellowish orange and reddish brown blended with the background (
<figureCitation id="132E4CB7681B497EFAFEA7329BD3FA73" box="[1338,1427,1384,1410]" captionStart="FIGURE 14" captionStartId="6.[151,257,1737,1761]" captionTargetBox="[151,1436,197,1711]" captionTargetId="figure@6.[151,1436,197,1713]" captionTargetPageId="6" captionText="FIGURE 14. Live nymph of Heteropodagrion sanguinipes oriented head down on vertical rock face in sheet flow (photo by J. Johnson)." httpUri="https://zenodo.org/record/196545/files/figure.png" pageId="7" pageNumber="36">Fig. 14</figureCitation>
) resulting in remarkable protective coloration. Possible predators I saw in this habitat included spiders and beetle larvae, although there are undoubtedly others. The slightly concave and flattened underside of 
<taxonomicName id="4C152BB1681B497EFF53A7829F25FA00" box="[151,357,1496,1521]" class="Insecta" family="Megapodagrionidae" genus="Heteropodagrion" kingdom="Animalia" order="Odonata" pageId="7" pageNumber="36" phylum="Arthropoda" rank="genus">
<emphasis id="B9618C20681B497EFF53A7829F25FA00" box="[151,357,1496,1521]" italics="true" pageId="7" pageNumber="36">Heteropodagrion</emphasis>
</taxonomicName>
nymphs is an adaptation to these living conditions, designed for occupying sheet flow on high-gradient rock surfaces. The venter of the thorax is slightly concave, the abdominal venter is nearly flat, and the ventral surfaces of the lateral caudal gills are concave, all which undoubtedly aid the nymph in adhering to and maneuvering on the rock in the swift flow. I estimated the velocity of the water to be roughly 
<quantity id="4CEDFDD7681B497EFF53A4379EA4F976" box="[151,228,1645,1671]" metricMagnitude="-1" metricUnit="m" metricValue="2.0" pageId="7" pageNumber="36" unit="cm" value="20.0">20 cm</quantity>
/sec. Despite the velocity, there is probably little drag on the nymphs due to frictional forces, as the water sheet itself tends to “cling” to the rock surface. I suspect that the contour and shape of the nymphs offers little in the way of resistance. The short, thick legs appear particularly strong. Disturbed nymphs ran or made herky-jerky movements in all directions on the vertical rock surfaces without losing their hold. When held in the palm of the hand and disturbed by a finger tip, nymphs would run and curl the tip of the abdomen up and forward over their dorsum (fig. 12); they appeared to poke the gills at the intruding finger tip. This behavior could be directed at an approaching predator; a detached hairy gill in the mouth of a predator might buy enough time for a 
<taxonomicName id="4C152BB1681B497EFEA9A5299C77F87D" box="[365,567,1907,1932]" class="Insecta" family="Megapodagrionidae" genus="Heteropodagrion" kingdom="Animalia" order="Odonata" pageId="7" pageNumber="36" phylum="Arthropoda" rank="genus">
<emphasis id="B9618C20681B497EFEA9A5299C77F87D" box="[365,567,1907,1932]" italics="true" pageId="7" pageNumber="36">Heteropodagrion</emphasis>
</taxonomicName>
nymph to escape.
</paragraph>
<paragraph id="8BAA5032681B497EFF01A5C29BDCF826" blockId="7.[151,1437,152,2007]" pageId="7" pageNumber="36">
At the locality where I found 
<taxonomicName id="4C152BB1681B497EFDFEA5C29D4FF840" box="[570,783,1944,1969]" class="Insecta" family="Megapodagrionidae" genus="Heteropodagrion" kingdom="Animalia" order="Odonata" pageId="7" pageNumber="36" phylum="Arthropoda" rank="genus">
<emphasis id="B9618C20681B497EFDFEA5C29D4FF840" box="[570,783,1944,1969]" italics="true" pageId="7" pageNumber="36">Heteropodagrion</emphasis>
</taxonomicName>
sp., I collected one nymph of 
<taxonomicName id="4C152BB1681B497EFB4AA5C29B6CF843" box="[1166,1324,1944,1970]" class="Insecta" family="Polythoridae" kingdom="Animalia" order="Odonata" pageId="7" pageNumber="36" phylum="Arthropoda" rank="family">Polythoridae</taxonomicName>
that was clinging to the rock surface in open sheet flow within a few centimeters of several 
<taxonomicName id="4C152BB1681B497EFBA2A5E79B70F827" box="[1126,1328,1981,2006]" class="Insecta" family="Megapodagrionidae" genus="Heteropodagrion" kingdom="Animalia" order="Odonata" pageId="7" pageNumber="36" phylum="Arthropoda" rank="genus">
<emphasis id="B9618C20681B497EFBA2A5E79B70F827" box="[1126,1328,1981,2006]" italics="true" pageId="7" pageNumber="36">Heteropodagrion</emphasis>
</taxonomicName>
nymphs.
</paragraph>
<paragraph id="8BAA503268144971FF53A2C29EBAFD66" blockId="8.[151,1437,152,663]" pageId="8" pageNumber="37">
This nymph appears to be a species of 
<taxonomicName id="4C152BB168144971FDACA2C29C9CFF40" box="[616,732,152,177]" class="Insecta" family="Polythoridae" genus="Polythore" kingdom="Animalia" order="Odonata" pageId="8" pageNumber="37" phylum="Arthropoda" rank="genus">
<emphasis id="B9618C2068144971FDACA2C29C9CFF40" box="[616,732,152,177]" italics="true" pageId="8" pageNumber="37">Polythore</emphasis>
</taxonomicName>
, although I was unable to rear it and no adult 
<taxonomicName id="4C152BB168144971FAC1A2C29BDCFF43" box="[1285,1436,152,178]" class="Insecta" family="Polythoridae" kingdom="Animalia" order="Odonata" pageId="8" pageNumber="37" phylum="Arthropoda" rank="family">Polythoridae</taxonomicName>
were seen at this locality. The nymph is amazingly similar in overall appearance to 
<taxonomicName id="4C152BB168144971FB4DA2E79B17FF27" box="[1161,1367,189,214]" class="Insecta" family="Megapodagrionidae" genus="Heteropodagrion" kingdom="Animalia" order="Odonata" pageId="8" pageNumber="37" phylum="Arthropoda" rank="genus">
<emphasis id="B9618C2068144971FB4DA2E79B17FF27" box="[1161,1367,189,214]" italics="true" pageId="8" pageNumber="37">Heteropodagrion</emphasis>
</taxonomicName>
, with compact tapering body, relatively short prementum (L/W = 1.12), angular projections on thoracic dorsum, setal patches on legs, and gills heavily setose with ventral surfaces concave. Despite its larger size and dorsal hooks, in dorsal view the resemblance of the polythorid nymph to that of 
<taxonomicName id="4C152BB168144971FBCDA3779A96FEB7" box="[1033,1238,301,326]" class="Insecta" family="Megapodagrionidae" genus="Heteropodagrion" kingdom="Animalia" order="Odonata" pageId="8" pageNumber="37" phylum="Arthropoda" rank="genus">
<emphasis id="B9618C2068144971FBCDA3779A96FEB7" box="[1033,1238,301,326]" italics="true" pageId="8" pageNumber="37">Heteropodagrion</emphasis>
</taxonomicName>
is obvious. This might be a case of convergent evolution, in which a similar body style/shape evolved in distinctly separate lineages as a result of selective pressures imposed by a unique habitat. 
<bibRefCitation id="EF842DC368144971FBD1A3229B0CFE63" author="Novelo-Gutierrez" box="[1045,1356,376,402]" pageId="8" pageNumber="37" refString="Novelo-Gutierrez, R. (2008) Description of the larva of Paraphlebia zoe Selys in Hagen, 1861 (Odonata: Megapodagrionidae). Zootaxa, 1876, 29 - 34." type="journal article" year="2008">Novelo-Gutiérrez (2008)</bibRefCitation>
found similarities between nymphs of the megapodagrionid 
<taxonomicName id="4C152BB168144971FCDCA3C79A64FE46" authority="Selys" authorityName="Selys" box="[792,1060,413,439]" class="Insecta" family="Megapodagrionidae" genus="Paraphlebia" kingdom="Animalia" order="Odonata" pageId="8" pageNumber="37" phylum="Arthropoda" rank="species" species="zoe">
<emphasis id="B9618C2068144971FCDCA3C79D9CFE47" box="[792,988,413,438]" italics="true" pageId="8" pageNumber="37">Paraphlebia zoe</emphasis>
Selys
</taxonomicName>
and coenagrionids in the 
<emphasis id="B9618C2068144971FA9EA3C79F10FE2A" italics="true" pageId="8" pageNumber="37">
<taxonomicName id="4C152BB168144971FA9EA3C79BDCFE47" box="[1370,1436,413,438]" class="Insecta" family="Coenagrionidae" genus="Argia" kingdom="Animalia" order="Odonata" pageId="8" pageNumber="37" phylum="Arthropoda" rank="genus">Argia</taxonomicName>
extranea-vivida
</emphasis>
group, both inhabiting seepage habitats on rocky walls or nearby on the ground. I examined 
<taxonomicName id="4C152BB168144971FF53A3B29E9AFDF0" box="[151,218,488,513]" class="Insecta" family="Coenagrionidae" genus="Argia" kingdom="Animalia" order="Odonata" pageId="8" pageNumber="37" phylum="Arthropoda" rank="genus">
<emphasis id="B9618C2068144971FF53A3B29E9AFDF0" box="[151,218,488,513]" italics="true" pageId="8" pageNumber="37">Argia</emphasis>
</taxonomicName>
nymphs from a vertical rocky seep near a waterfall in Cañar Province, 
<collectingCountry id="F30210A268144971FBFBA3B29AE5FDF3" box="[1087,1189,488,514]" name="Ecuador" pageId="8" pageNumber="37">Ecuador</collectingCountry>
, which are probably 
<taxonomicName id="4C152BB168144971FF53A0579F80FDD6" authority="Calvert" authorityName="Calvert" box="[151,448,525,551]" class="Insecta" family="Coenagrionidae" genus="Argia" kingdom="Animalia" order="Odonata" pageId="8" pageNumber="37" phylum="Arthropoda" rank="species" species="talamanca">
<emphasis id="B9618C2068144971FF53A0579F1DFDD7" box="[151,349,525,550]" italics="true" pageId="8" pageNumber="37">Argia talamanca</emphasis>
Calvert
</taxonomicName>
; they have an overall appearance similar to 
<taxonomicName id="4C152BB168144971FC0FA0579AD7FDD7" box="[971,1175,525,550]" class="Insecta" family="Megapodagrionidae" genus="Heteropodagrion" kingdom="Animalia" order="Odonata" pageId="8" pageNumber="37" phylum="Arthropoda" rank="genus">
<emphasis id="B9618C2068144971FC0FA0579AD7FDD7" box="[971,1175,525,550]" italics="true" pageId="8" pageNumber="37">Heteropodagrion</emphasis>
</taxonomicName>
, although they do not possess the angular projections on the dorsum of the thorax. Future research should be directed at the comparative morphology and behavior of various damselflies that occupy sheet flow and normal stream habitats.
</paragraph>
</subSubSection>
</treatment>
</document>