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<taxonomicName id="DCDA4D38E020D747FF79E97EFED1FAA5" ID-CoL="62TNF" authorityName="Fraser" authorityYear="1949" box="[151,283,1374,1401]" class="Insecta" family="Chlorocyphidae" genus="Heliocypha" kingdom="Animalia" order="Odonata" pageId="21" pageNumber="230" phylum="Arthropoda" rank="genus">
<emphasis id="29AEEAA9E020D747FF79E97EFED1FAA5" bold="true" box="[151,283,1374,1401]" italics="true" pageId="21" pageNumber="230">Heliocypha</emphasis>
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The genus includes nine species ranging from Sundaland to south
<collectingCountry id="63CD762BE020D747FC91E9A3FC73FA41" box="[895,953,1411,1437]" name="India" pageId="21" pageNumber="230">India</collectingCountry>
and southern
<collectingCountry id="63CD762BE020D747FBB9E9A3FB50FA41" box="[1111,1178,1411,1437]" name="China" pageId="21" pageNumber="230">China</collectingCountry>
including
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. The habitus of the larva of
<taxonomicName id="DCDA4D38E020D747FE7AE987FD2DFA1D" authority="(Hagen)" baseAuthorityName="Hagen" box="[404,743,1447,1473]" class="Insecta" family="Chlorocyphidae" genus="Heliocypha" kingdom="Animalia" order="Odonata" pageId="21" pageNumber="230" phylum="Arthropoda" rank="species" species="bisignata">
<emphasis id="29AEEAA9E020D747FE7AE987FD4EFA1D" box="[404,644,1447,1473]" italics="true" pageId="21" pageNumber="230">Heliocypha bisignata</emphasis>
(Hagen)
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with a partial view of the mask was figured by
<bibRefCitation id="7F4B4B4AE020D747FB10E987FA5DFA1D" author="Fraser, F. C." box="[1278,1431,1447,1473]" pageId="21" pageNumber="230" pagination="683 - 691" refId="ref22898" refString="Fraser, F. C. (1928) Indian dragonflies, Part XXX. Journal of the Bombay Natural History Society, 32, 683 - 691, pls. 1 - 3, excl." type="journal article" year="1928">Fraser (1928)</bibRefCitation>
, which he later briefly described (as
<taxonomicName id="DCDA4D38E020D747FDD9E9EBFD77FA39" authorityName="Rambur" authorityYear="1842" box="[567,701,1483,1509]" class="Insecta" family="Chlorocyphidae" genus="Rhinocypha" kingdom="Animalia" order="Odonata" pageId="21" pageNumber="230" phylum="Arthropoda" rank="genus">
<emphasis id="29AEEAA9E020D747FDD9E9EBFD77FA39" box="[567,701,1483,1509]" italics="true" pageId="21" pageNumber="230">Rhinocypha</emphasis>
</taxonomicName>
), (
<bibRefCitation id="7F4B4B4AE020D747FD30E9EBFCA1FA39" author="Fraser, F. C." box="[734,875,1483,1509]" pageId="21" pageNumber="230" pagination="1 - 4" refId="ref23053" refString="Fraser, F. C. (1934) The fauna of British India, including Ceylon and Burma. Odonata. Vol. II. Taylor &amp; Francis Ltd., London, xxiii + 398 pp., pls. 1 - 4 excl." type="book chapter" year="1934">Fraser 1934</bibRefCitation>
).
<bibRefCitation id="7F4B4B4AE020D747FC6FE9EBFBF6FA39" author="Lieftinck, M. A." box="[897,1084,1483,1509]" pageId="21" pageNumber="230" pagination="215 - 224" refId="ref24467" refString="Lieftinck, M. A. (1947) Two interesting new insular Rhinocypha from Malaysia (Odon.). Tijdschrift voor Entomologie, 88, 215 - 224." type="journal article" year="1947">Lieftinck (1947)</bibRefCitation>
provided an illustration of the habitus of
<taxonomicName id="DCDA4D38E020D747FEE3E9CFFDFEF9D5" authority="(Burmeister)" baseAuthorityName="Burmeister" box="[269,564,1519,1545]" class="Insecta" family="Chlorocyphidae" genus="Heliocypha" kingdom="Animalia" order="Odonata" pageId="21" pageNumber="230" phylum="Arthropoda" rank="species" species="fenestrata">
<emphasis id="29AEEAA9E020D747FEE3E9CFFE55F9D5" box="[269,415,1519,1545]" italics="true" pageId="21" pageNumber="230">H. fenestrata</emphasis>
(Burmeister)
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and described its habitat and later provided accurate details of the prementum and labial palp (
<bibRefCitation id="7F4B4B4AE020D747FEBDEA33FDC8F9F1" author="Lieftinck, M. A." box="[339,514,1555,1581]" pageId="21" pageNumber="230" pagination="1 - 95" refId="ref24643" refString="Lieftinck, M. A. (1962) Odonata. Insects of Micronesia, 5 (1), 1 - 95, 1 map excl." type="book chapter" year="1962">Lieftinck 1962</bibRefCitation>
).
<bibRefCitation id="7F4B4B4AE020D747FDF6EA33FCF1F9F1" author="Kumar, A. &amp; Prasad, M." box="[536,827,1555,1581]" pageId="21" pageNumber="230" pagination="547 - 554" refId="ref24362" refString="Kumar, A. &amp; Prasad, M. (1977 c) On the larvae of Rhinocypha (Odonata: Cholorocyphidae) from Garhwal hills. Oriental Insects, 11, 547 - 554. https: // doi. org / 10.1080 / 00305316.1977.11090927" type="journal article" year="1977">Kumar &amp; Prasad (1977c)</bibRefCitation>
provided for
<taxonomicName id="DCDA4D38E020D747FC33EA33FB7CF9F1" authority="(Selys)" baseAuthorityName="Selys" box="[989,1206,1555,1581]" class="Insecta" family="Chlorocyphidae" genus="Heliocypha" kingdom="Animalia" order="Odonata" pageId="21" pageNumber="230" phylum="Arthropoda" rank="species" species="biforata">
<emphasis id="29AEEAA9E020D747FC33EA33FB96F9F1" box="[989,1116,1555,1581]" italics="true" pageId="21" pageNumber="230">H. biforata</emphasis>
(Selys)
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a minutely detailed description and illustration, including the habitus. In the key “scape rough” is said to characterise this species and appears as a possible generic character. However this apparent roughness may be an illusion created by short recurved setae which in wet specimens trap fluid and give the appearance of small tubercles where none are present (Orr
<emphasis id="29AEEAA9E020D747FF23EA83FECFF961" box="[205,261,1699,1725]" italics="true" pageId="21" pageNumber="230">et al.</emphasis>
2024). The drawing of the antenna in this paper depicts the warty scape as 0.33 times the overall length of the antenna, but by the measurements given in the text and the depiction of the antennae on the habitus drawing this proportion should be 0.42, which is very close to the other species considered, hence the detailed drawing itself must be regarded as a lapsus, perhaps because it was inadvertently observed at an angle. The authors do not comment on an obviously relatively short scape. The widespread
<emphasis id="29AEEAA9E020D747FCE6EB13FC7BF891" box="[776,945,1843,1869]" italics="true" pageId="21" pageNumber="230">
H. p.
<taxonomicName id="DCDA4D38E020D747FCA9EB13FC7BF891" authorityName="Percheron" authorityYear="1835" box="[839,945,1843,1869]" class="Insecta" family="Chlorocyphidae" genus="Heliocypha" kingdom="Animalia" order="Odonata" pageId="21" pageNumber="230" phylum="Arthropoda" rank="species" species="perforata">perforata</taxonomicName>
</emphasis>
, was described with little attention to detail by
<bibRefCitation id="7F4B4B4AE020D747FF52EB77FEFFF8AD" author="Xu, Q. - H." box="[188,309,1878,1905]" pageId="21" pageNumber="230" pagination="137 - 141" refId="ref27657" refString="Xu, Q. - H. (2015) Description of the final stadium larva of Heliocypha perforata perforata (Percheron), with discussion of the taxonomic characters of the larvae of the genus Heliocypha Fraser (Odonata: Zygoptera: Chlorocyphidae). Zootaxa, 3926 (1), 137 - 141. https: // doi. org / 10.11646 / zootaxa. 3926.1.8" type="journal article" year="2015">Xu (2015)</bibRefCitation>
; it almost certain this paper overlooks the small supplementary branch on the outer process of the labial palp, as this is visible in a very young specimen of this species figured by
<bibRefCitation id="7F4B4B4AE020D747FBF9EB5BFB52F849" author="Ng, C. Y." box="[1047,1176,1915,1941]" pageId="21" pageNumber="230" refId="ref25224" refString="Ng, C. Y. (2024 c) Freshwater Macroinvertebrates in Hong Kong: Odonata: Rhinocypha perforata. Available from: https: // www. biosch. hku. hk / ecology / staffhp / dd / macroinvertebrates / Odonata / Chlorocyphidae / Rhinocypha _ perforata. html (accessed 16 May 2024)" type="url" year="2024">Ng (2024c)</bibRefCitation>
and occurs on all other members of the family examined (
<bibRefCitation id="7F4B4B4AE020D747FDFFEBBFFD6FF865" author="Fraser, F. C." box="[529,677,1951,1977]" pageId="21" pageNumber="230" pagination="197 - 198" refId="ref22755" refString="Fraser, F. C. (1919 a) The larva of Micromerus lineatus Burm. Records of the Indian Museum, 16, 197 - 198, pl. 23 excl. https: // doi. org / 10.26515 / rzsi / v 16 / i 2 / 1919 / 162807" type="journal article" year="1919">
Fraser 1919
<bibRefCitation id="7F4B4B4AE020D747FD76EBBFFD6FF865" author="Kumar, A." box="[664,677,1951,1977]" pageId="21" pageNumber="230" pagination="83 - 118" refId="ref24196" refString="Kumar, A. (1973) Descriptions of the last instar larvae of Odonata from the Dehra Dun Valley (India), with notes on biology I. (suborder Zygoptera). Oriental Insects, 7, 83 - 118. https: // doi. org / 10.1080 / 00305316.1973.10434207" type="journal article" year="1973">a</bibRefCitation>
</bibRefCitation>
, Kumar 1973,
<bibRefCitation id="7F4B4B4AE020D747FCA9EBBFFB87F865" author="Kumar, A. &amp; Prasad, M." box="[839,1101,1951,1977]" pageId="21" pageNumber="230" pagination="547 - 554" refId="ref24362" refString="Kumar, A. &amp; Prasad, M. (1977 c) On the larvae of Rhinocypha (Odonata: Cholorocyphidae) from Garhwal hills. Oriental Insects, 11, 547 - 554. https: // doi. org / 10.1080 / 00305316.1977.11090927" type="journal article" year="1977">
Kumar &amp; Prasad 1977
<bibRefCitation id="7F4B4B4AE020D747FBAFEBBFFB87F865" author="Dow, R. A. &amp; Choong, C. Y. &amp; Grinang, J. &amp; Lupiyaningdyah, P. &amp; Ngiam, R. W. J. &amp; Kalkman, V. J." box="[1089,1101,1951,1977]" pageId="21" pageNumber="230" pagination="1 - 122" refId="ref22552" refString="Dow, R. A., Choong, C. Y., Grinang, J., Lupiyaningdyah, P., Ngiam, R. W. J. &amp; Kalkman, V. J. (2024) Checklist of the Odonata (Insecta) of Sundaland and Wallacea (Malaysia, Singapore, Brunei, Indonesia and Timor Leste). Zootaxa, 5460 (1), 1 - 122. https: // doi. org / 10.11646 / zootaxa. 5460.1.1" type="journal article" year="2024">c</bibRefCitation>
</bibRefCitation>
, Orr
<emphasis id="29AEEAA9E020D747FB6AEBBFFB79F865" box="[1156,1203,1951,1977]" italics="true" pageId="21" pageNumber="230">et al</emphasis>
. 2024). However the caudal spikes in the habitus photograph conform closely with other known species. Finally, a detailed description of the Bornean endemic
<taxonomicName id="DCDA4D38E023D744FE65ECB7FDDBFF6D" authorityName="Selys" authorityYear="1859" box="[395,529,151,177]" class="Insecta" family="Chlorocyphidae" genus="Heliocypha" kingdom="Animalia" order="Odonata" pageId="22" pageNumber="231" phylum="Arthropoda" rank="species" species="biseriata">
<emphasis id="29AEEAA9E023D744FE65ECB7FDDBFF6D" box="[395,529,151,177]" italics="true" pageId="22" pageNumber="231">H. biseriata</emphasis>
</taxonomicName>
was given by Orr
<emphasis id="29AEEAA9E023D744FD31ECB7FCDAFF6D" box="[735,784,151,177]" italics="true" pageId="22" pageNumber="231">et al</emphasis>
. (2024). It was concluded that the thick but gently tapered caudal spikes and short curled setae on the scape, creating the impression of small tubercles in wet specimens, may be diagnostic at least when compared with
<taxonomicName id="DCDA4D38E023D744FD94ECFFFCE2FF25" box="[634,808,223,249]" class="Insecta" family="Chlorocyphidae" genus="Aristocypha" kingdom="Animalia" order="Odonata" pageId="22" pageNumber="231" phylum="Arthropoda" rank="species" species="undetermined">
<emphasis id="29AEEAA9E023D744FD94ECFFFCCBFF25" box="[634,769,223,249]" italics="true" pageId="22" pageNumber="231">Aristocypha</emphasis>
sp.
</taxonomicName>
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<emphasis id="29AEEAA9E020D747FF79E81EFEEBFB85" bold="true" box="[151,289,1086,1113]" italics="true" pageId="21" pageNumber="230">Aristocypha</emphasis>
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The genus includes 12 species ranging from the Himalayas to southern
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and
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,
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,
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, Indochina and West
<collectingCountry id="63CD762BE020D747FE90E8A7FE22FB7D" box="[382,488,1159,1185]" name="Malaysia" pageId="21" pageNumber="230">Malaysia</collectingCountry>
. Larvae of
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<emphasis id="29AEEAA9E020D747FD86E8A7FC61FB7D" box="[616,939,1159,1185]" italics="true" pageId="21" pageNumber="230">Aristocypha quadrimaculata</emphasis>
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and
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<emphasis id="29AEEAA9E020D747FBD4E8A7FB07FB7D" box="[1082,1229,1159,1185]" italics="true" pageId="21" pageNumber="230">A. trifasciata</emphasis>
(Selys)
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were both described and illustrated with habitus and numerous structural details by
<bibRefCitation id="7F4B4B4AE020D747FC01E88BFAD2FB19" author="Kumar, A. &amp; Prasad, M." box="[1007,1304,1195,1221]" pageId="21" pageNumber="230" pagination="547 - 554" refId="ref24362" refString="Kumar, A. &amp; Prasad, M. (1977 c) On the larvae of Rhinocypha (Odonata: Cholorocyphidae) from Garhwal hills. Oriental Insects, 11, 547 - 554. https: // doi. org / 10.1080 / 00305316.1977.11090927" type="journal article" year="1977">Kumar &amp; Prasad (1977c)</bibRefCitation>
. The larval habitus of
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<emphasis id="29AEEAA9E020D747FEE2E8EFFE6AFB35" box="[268,416,1231,1257]" italics="true" pageId="21" pageNumber="230">A. fenestrella</emphasis>
(Rambur)
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was illustrated by
<bibRefCitation id="7F4B4B4AE020D747FD0BE8EFFC97FB35" author="Orr, A. G." box="[741,861,1230,1257]" pageId="21" pageNumber="230" refId="ref25754" refString="Orr, A. G. (2005) Dragonflies of Peninsular Malaysia and Singapore. Natural History Publications (Borneo), Kota Kinabalu, vi + 127 pp." type="book" year="2005">Orr (2005)</bibRefCitation>
and a male larva of the same species was described in detail by Orr
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. (2024). Comparisons between these descriptions suggests that the form of the caudal spikes and the presence of longer setae on the scape may distinguish this genus from
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<emphasis id="29AEEAA9E020D747FBEAE937FB4FFAED" box="[1028,1157,1303,1329]" italics="true" pageId="21" pageNumber="230">Heliocypha</emphasis>
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in known species.
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Just two species of this genus occur in parts of Sundaland. The larva of
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. (2024) and differs clearly from known chlorocyphid larvae in a number of characters, including possessing a very long prementum, proportionally very long antennae, reduced anterior tubercles on the pronotum, long, thin evenly tapered caudal spikes, numerous small spines on the venter of the terminal abdominal segments and distinctive outer female gonapophyses. The other species of the genus,
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Orr
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, is confined to
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and adjoining areas of
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and its larva is expected to be very like that of
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, with the caveat that closely related species can show surprising differences in some characters as in the Japanese
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<emphasis id="29AEEAA9E022D745FF79EDFAFE5BFE28" bold="true" box="[151,401,474,500]" pageId="23" pageNumber="232">
Family
<taxonomicName id="DCDA4D38E022D745FF1CEDFAFE5BFE28" ID-CoL="9WZ26" authorityName="Dijkstra, Kalkman, Dow, Stokvis &amp; van Tol" authorityYear="2014" box="[242,401,474,500]" class="Insecta" family="Devadattidae" higherTaxonomySource="CoL,GBIF" kingdom="Animalia" order="Odonata" pageId="23" pageNumber="232" phylum="Arthropoda" rank="family">Devadattidae</taxonomicName>
</emphasis>
</heading>
</paragraph>
</subSubSection>
<subSubSection id="53C06530E022D745FF79EDDFFD80FC5D" pageId="23" pageNumber="232" type="discussion">
<paragraph id="1B6536BBE022D745FF79EDDFFD80FC5D" blockId="23.[151,1437,474,897]" pageId="23" pageNumber="232">
The family includes just a single genus
<taxonomicName id="DCDA4D38E022D745FD86EDDFFD17FDC5" authorityName="Kirby" authorityYear="1890" box="[616,733,511,537]" class="Insecta" family="Amphipterygidae" genus="Devadatta" kingdom="Animalia" order="Odonata" pageId="23" pageNumber="232" phylum="Arthropoda" rank="genus">
<emphasis id="29AEEAA9E022D745FD86EDDFFD17FDC5" box="[616,733,511,537]" italics="true" pageId="23" pageNumber="232">Devadatta</emphasis>
</taxonomicName>
represented by 13 species which range from southern
<collectingCountry id="63CD762BE022D745FAB7EDDFFA56FDC5" box="[1369,1436,511,537]" name="China" pageId="23" pageNumber="232">China</collectingCountry>
(
<collectingRegion id="D91EF859E022D745FF71EE03FF33FDE1" box="[159,249,547,573]" country="China" name="Yunnan" pageId="23" pageNumber="232">Yunnan</collectingRegion>
,
<collectingRegion id="D91EF859E022D745FEE6EE03FEA7FDE1" box="[264,365,547,573]" country="China" name="Guangxi" pageId="23" pageNumber="232">Guangxi</collectingRegion>
), Indochina and
<collectingCountry id="63CD762BE022D745FDDEEE03FD5CFDE1" box="[560,662,547,573]" name="Thailand" pageId="23" pageNumber="232">Thailand</collectingCountry>
, to peninsular
<collectingCountry id="63CD762BE022D745FCAAEE03FC67FDE1" box="[836,941,547,573]" name="Malaysia" pageId="23" pageNumber="232">Malaysia</collectingCountry>
, Borneo and Sumatra. The larva is known only for
<taxonomicName id="DCDA4D38E022D745FF19EE67FE16FDBD" authority="(Selys)" baseAuthorityName="Selys" box="[247,476,583,609]" class="Insecta" family="Amphipterygidae" genus="Devadatta" kingdom="Animalia" order="Odonata" pageId="23" pageNumber="232" phylum="Arthropoda" rank="species" species="argyoides">
<emphasis id="29AEEAA9E022D745FF19EE67FE4CFDBD" box="[247,390,583,609]" italics="true" pageId="23" pageNumber="232">D. argyoides</emphasis>
(Selys)
</taxonomicName>
(
<figureCitation id="83E12A3EE022D745FE05EE67FDFEFDBD" box="[491,564,583,609]" captionStart="FIGURES 8–11. 8" captionStartId="4.[151,264,1914,1938]" captionTargetBox="[200,1382,177,1900]" captionTargetId="figure-21@4.[193,1394,169,1908]" captionTargetPageId="4" captionText="FIGURES 8–11. 8. Pseudolestes mirabilis, ventral view of terminal abdominal segments and caudal gills of showing gill tufts on S10 (after Yu &amp; Bu 2011). 9. Devadatta argyoides, detail of extruded right gill tuft, posterior view (redrawn from Watson 1966). 10. Pseudolestes mirabilis, habitus with detail of mask (after Yu &amp; Bu 2011). 11. Devadatta argyoides, habitus with details of right antenna and mask (original drawing)." figureDoi="http://doi.org/10.5281/zenodo.13618323" httpUri="https://zenodo.org/record/13618323/files/figure.png" pageId="23" pageNumber="232">Fig.11</figureCitation>
) from peninsular
<collectingCountry id="63CD762BE022D745FD12EE67FCA9FDBD" box="[764,867,583,609]" name="Malaysia" pageId="23" pageNumber="232">Malaysia</collectingCountry>
where it occurs in leaf packs and under dead wood near the head of forest streams The habitus was figured by
<bibRefCitation id="7F4B4B4AE022D745FCACEE4BFC7CFD59" author="Orr, A. G." box="[834,950,619,645]" pageId="23" pageNumber="232" refId="ref25717" refString="Orr, A. G. (2003) A guide to the Dragonflies of Borneo: their identification and biology. Natural History Publications (Borneo), Kota Kinabalu, x + 195 pp." type="book" year="2003">Orr (2003</bibRefCitation>
,
<bibRefCitation id="7F4B4B4AE022D745FC2DEE4AFC37FD59" author="Orr, A. G." box="[963,1021,618,645]" pageId="23" pageNumber="232" refId="ref25754" refString="Orr, A. G. (2005) Dragonflies of Peninsular Malaysia and Singapore. Natural History Publications (Borneo), Kota Kinabalu, vi + 127 pp." type="book" year="2005">2005</bibRefCitation>
) and structural details are shown by
<bibRefCitation id="7F4B4B4AE022D745FF79EEAFFE7BFD75" author="Novelo-Gutierrez, R." box="[151,433,654,681]" pageId="23" pageNumber="232" pagination="73 - 87" refId="ref25567" refString="Novelo-Gutierrez, R. (1995) The larva of Amphipteryx and a reclassification of Amphipterygidae sensu lato based upon the larvae (Zygoptera). Odonatologica, 24, 73 - 87." type="journal article" year="1995">Novelo-Gutierrez (1995)</bibRefCitation>
. In life, the unique gill tufts (
<figureCitation id="83E12A3EE022D745FD1CEEAFFCFCFD75" box="[754,822,655,681]" captionStart="FIGURES 8–11. 8" captionStartId="4.[151,264,1914,1938]" captionTargetBox="[200,1382,177,1900]" captionTargetId="figure-21@4.[193,1394,169,1908]" captionTargetPageId="4" captionText="FIGURES 8–11. 8. Pseudolestes mirabilis, ventral view of terminal abdominal segments and caudal gills of showing gill tufts on S10 (after Yu &amp; Bu 2011). 9. Devadatta argyoides, detail of extruded right gill tuft, posterior view (redrawn from Watson 1966). 10. Pseudolestes mirabilis, habitus with detail of mask (after Yu &amp; Bu 2011). 11. Devadatta argyoides, habitus with details of right antenna and mask (original drawing)." figureDoi="http://doi.org/10.5281/zenodo.13618323" httpUri="https://zenodo.org/record/13618323/files/figure.png" pageId="23" pageNumber="232">Fig. 9</figureCitation>
), illustrated by
<bibRefCitation id="7F4B4B4AE022D745FC0CEEAFFB42FD75" author="Watson, J. A. L." box="[994,1160,655,681]" pageId="23" pageNumber="232" pagination="9 - 21" refId="ref27538" refString="Watson, J. A. L. (1966) The structure and function of gill tufts in larval Amphipterygidae (Odonata: Zygoptera). Journal of Morphology, 120, 9 - 21. https: // doi. org / 10.1002 / jmor. 1051200103" type="journal article" year="1966">Watson (1966)</bibRefCitation>
, on the venter of the S10 pulsate regularly as they are retracted and extended, especially in low oxygen conditions induced in the laboratory (
<bibRefCitation id="7F4B4B4AE022D745FF71EEF7FEC6FD2D" author="Orr, A. G." box="[159,268,727,753]" pageId="23" pageNumber="232" pagination="24 - 27" refId="ref25785" refString="Orr, A. G. (2008) Eight days in Peninsular Malaysia - and the benefits of hindsight. Agrion, 12, 24 - 27." type="journal article" year="2008">Orr 2008</bibRefCitation>
). These structures, and the unique hardened tricorn caudal gill, the elongated antennal scape longer than the pedicel and the short, broad prementum are expected in all species, but interspecific variation may occur in proportions of body parts and the shape of the caudal gills. The gill tufts may be retracted in living or preserved specimens, but in this case two small tumescent orifices are apparent. It is unlikely the larvae of any species can be mistaken for those of any other family.
</paragraph>
</subSubSection>
</treatment>
</document>

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<subSubSection id="53C06530E022D745FF79EFF2FE41FC30" box="[151,395,978,1004]" pageId="23" pageNumber="232" type="nomenclature">
<paragraph id="1B6536BBE022D745FF79EFF2FE41FC30" blockId="23.[151,395,978,1004]" box="[151,395,978,1004]" pageId="23" pageNumber="232">
<heading id="402D81D7E022D745FF79EFF2FE41FC30" bold="true" box="[151,395,978,1004]" fontSize="11" level="1" pageId="23" pageNumber="232" reason="1">
<emphasis id="29AEEAA9E022D745FF79EFF2FE41FC30" bold="true" box="[151,395,978,1004]" pageId="23" pageNumber="232">
Family
<taxonomicName id="DCDA4D38E022D745FF19EFF2FE41FC30" ID-CoL="9Y5" baseAuthorityName="Jacobson &amp; Bianchi" baseAuthorityYear="1905" box="[247,395,978,1004]" class="Insecta" family="Euphaeidae" kingdom="Animalia" order="Odonata" pageId="23" pageNumber="232" phylum="Arthropoda" rank="family">Euphaeidae</taxonomicName>
</emphasis>
</heading>
</paragraph>
</subSubSection>
<subSubSection id="53C06530E022D745FF79E83BFC94F849" pageId="23" pageNumber="232" type="discussion">
<paragraph id="1B6536BBE022D745FF79E83BFEBFFB19" blockId="23.[151,1437,1051,1941]" pageId="23" pageNumber="232">
The larvae of 6 of the 8 genera and approximately one third of the total of 79 species are known but not necessarily individually described. The family is exclusively Oriental with the exception of the monotypic genus
<emphasis id="29AEEAA9E022D745FADEE81FFA56FB85" box="[1328,1436,1087,1113]" italics="true" pageId="23" pageNumber="232">
<taxonomicName id="DCDA4D38E022D745FADEE81FFA52FB85" authorityName="Charpentier" authorityYear="1840" box="[1328,1432,1087,1113]" class="Insecta" family="Euphaeidae" genus="Epallage" kingdom="Animalia" order="Odonata" pageId="23" pageNumber="232" phylum="Arthropoda" rank="genus">Epallage</taxonomicName>
,
</emphasis>
ranges from south-east Europe to Kashmir, politically within Indian borders, but placed by
<bibRefCitation id="7F4B4B4AE022D745FB7AE843FA9CFBA1" author="Holt, B. G. &amp; Lessard, J. - P. &amp; Borregaard, M. K. &amp; Fritz, S. A. &amp; Araujo, M. B. &amp; Dimitrov, D. &amp; Fabre, P. - H. &amp; Graham, C. H. &amp; Graves, G. R. &amp; Jonsson, K. A. &amp; Nogues-Bravo, D. &amp; Wang, Z. &amp; Whittaker, R. J. &amp; Fjeldsa, J. &amp; Rahbek, C." box="[1172,1366,1123,1149]" pageId="23" pageNumber="232" pagination="74 - 78" refId="ref23468" refString="Holt, B. G., Lessard, J. - P., Borregaard, M. K., Fritz, S. A., Araujo, M. B., Dimitrov, D., Fabre, P. - H., Graham, C. H., Graves, G. R., Jonsson, K. A., Nogues-Bravo, D., Wang, Z., Whittaker, R. J., Fjeldsa, J. &amp; Rahbek, C. (2013) An update of Wallace's zoogeographic regions of the world. Science, 339, 74 - 78. https: // doi. org / 10.1126 / science. 1228282" type="journal article" year="2013">
Holt
<emphasis id="29AEEAA9E022D745FB23E843FB34FBA1" box="[1229,1278,1123,1149]" italics="true" pageId="23" pageNumber="232">et al</emphasis>
. (2013)
</bibRefCitation>
in the Saharo-Arabian realm. Within the Oriental region the family is conspicuously absent from
<collectingRegion id="D91EF859E022D745FB7EE8A7FB3FFB7D" box="[1168,1269,1159,1185]" country="Indonesia" name="Sulawesi Utara" pageId="23" pageNumber="232">Sulawesi</collectingRegion>
and Wetar and Timor in
<collectingCountry id="63CD762BE022D745FEEEE88BFEBBFB19" box="[256,369,1195,1221]" name="Indonesia" pageId="23" pageNumber="232">Indonesia</collectingCountry>
.
</paragraph>
<paragraph id="1B6536BBE022D745FF29E8EFFDE9F8AD" blockId="23.[151,1437,1051,1941]" pageId="23" pageNumber="232">
Euphaeid larvae are unmistakable by virtue of the series of long paired fleshy gills on the venter of abdominal S2–8 (
<figureCitation id="83E12A3EE022D745FF0EE8D3FEEBFAD1" box="[224,289,1267,1293]" captionStart="FIGURES 1–7. 1" captionStartId="3.[151,275,1843,1869]" captionTargetBox="[252,1341,160,1825]" captionTargetId="figure-15@3.[243,1344,149,1836]" captionTargetPageId="3" captionText="FIGURES 1–7. 1. Ventral view of abdomen of Euphaea sp. showing abdominal gills and saccoid gills (original drawing). 2. Head and antennae of Calopterygidae (Neurobasis chinensis) (original drawing). 3. Head and antennae of Chlorocyphidae (Heliocypha biseriata) (original drawing). 4. Antenna of Podolestes orientalis Selys, 1852, scape arrowed (from Choong &amp; Orr 2010). 5. Mask of Calopterygidae (Calopteryx japonica Selys redrawn from Miyakawa 1983). 6. Mask of Chlorocyphidae (Libellago hyalina, from Orr et al. 2024). 7. Habitus of L. hyalina, with detail of right caudal spike (from Orr et al. 2024)." figureDoi="http://doi.org/10.5281/zenodo.13618321" httpUri="https://zenodo.org/record/13618321/files/figure.png" pageId="23" pageNumber="232">Fig. 1</figureCitation>
). These are present even on very young larvae and begin to develop early in embryogenesis (
<bibRefCitation id="7F4B4B4AE022D745FADCE8D3FF3CFAED" author="Suzuki, K. &amp; Watanabe, Y. &amp; Tojo, K." pageId="23" pageNumber="232" pagination="280 - 293" refId="ref26853" refString="Suzuki, K., Watanabe, Y. &amp; Tojo, K. (2020) Embryogenesis of the damselfly Euphaea yayeyamana Oguma (Insecta: Odonata: Euphaeidae), with special reference to the formation of their larval abdominal &quot; gill-like &quot; appendages. Entomological Science, 23, 280 - 293. https: // doi. org / 10.1111 / ens. 12421" type="journal article" year="2020">
Suzuki
<emphasis id="29AEEAA9E022D745FA66E8D3FF67FAED" italics="true" pageId="23" pageNumber="232">et al</emphasis>
. 2020
</bibRefCitation>
,
<bibRefCitation id="7F4B4B4AE022D745FEEFE937FEBFFAED" author="Ng, C. Y." box="[257,373,1303,1329]" pageId="23" pageNumber="232" refId="ref25282" refString="Ng, C. Y. (2024 d) Freshwater Macroinvertebrates in Hong Kong: Odonata: Euphaea decorata. Available from: https: // www. biosch. hku. hk / ecology / staffhp / dd / macroinvertebrates / Odonata / Euphaeidae / Euphaea _ decorata. html (accessed 20 March 2024)" type="url" year="2024">Ng 2024d</bibRefCitation>
). The habitus (examples
<figureCitation id="83E12A3EE022D745FD62E937FCC1FAED" box="[652,779,1302,1329]" captionStart="FIGURES 57–60. 57" captionStartId="11.[151,264,1933,1957]" captionTargetBox="[193,1384,155,1907]" captionTargetId="figure-15@11.[188,1400,151,1917]" captionTargetPageId="11" captionText="FIGURES 57–60. 57. Dysphaea gloriosa, habitus (drawn from Nguyen et al. 2024). 58. Bayadera strigata, habitus (drawn from Yang &amp; Orr 2024). 59. Euphaea decorata, habitus. (drawn from Xu 2017). 60. Anisopleura furcata, habitus (drawn from Keetapithchayakul et al. 2024)." figureDoi="http://doi.org/10.5281/zenodo.13618341" httpUri="https://zenodo.org/record/13618341/files/figure.png" pageId="23" pageNumber="232">Figs 57–60</figureCitation>
) is overall flattened, reminiscent of Plecoptera, with broad flattened femora adapted to clinging to the underside of stones and large boulders where they are generally to be found in swiftly flowing clear water and rapids. All have saccoid caudal gills (
<figureCitation id="83E12A3EE022D745FBF7E97FFB57FAA5" box="[1049,1181,1375,1401]" captionStart="FIGURES 67–74. 67" captionStartId="13.[151,264,1814,1838]" captionTargetBox="[256,1340,160,1799]" captionTargetId="figure-15@13.[246,1342,151,1807]" captionTargetPageId="13" captionText="FIGURES 67–74. 67. Anisopleura subplatystyla, caudal gills dorsal view (original drawing). 68. Bayadera strigata, caudal gills dorsal view, (drawn from Yang &amp; Orr 2024). 69. Euphaea decorata, caudal gills dorsal view (drawn from Xu 2017). 70. Euphaea superba, caudal gills dorsal view (redrawn from Wu et al. 2017). 71. Bayadera strigata: male genital apophyses; ventral (left), right lateral (right), (drawn from Yang &amp; Orr 2024). 72. Euphaea decorata, male genital apophyses; ventral view (drawn from Xu 2017). 73. Euphaea superba, terminal processes of palpal lobe ventral view (movable hook truncated), (redrawn from Wu et al. 2017). 74. Anisopleura furcata, left labial palp showing terminal processes of palpal lobe, dorsal view (redrawn from Keetapithchayakul et al. 2024)." figureDoi="http://doi.org/10.5281/zenodo.13618345" httpUri="https://zenodo.org/record/13618345/files/figure.png" pageId="23" pageNumber="232">Figs 67–70</figureCitation>
), the form of which is of some diagnostic value, but these may be lost and/or shrivelled in exuviae. Euphaeid larvae were first described by
<bibRefCitation id="7F4B4B4AE022D745FF55E987FE9CFA1D" author="Hagen, H." box="[187,342,1447,1473]" pageId="23" pageNumber="232" pagination="lxv - lxvii" refId="ref23289" refString="Hagen, H. (1880) Essai d'un synopsis des larves de Calopterygines. [Comptes-rendus de la Socie´te´Entomologique de Belgique. Seance du 1 er Mai 1880]. Annales de la Societe entomologique de Belgique, 23, lxv - lxvii." type="journal article" year="1880">Hagen (1880)</bibRefCitation>
with reference to
<taxonomicName id="DCDA4D38E022D745FDCBE987FC85FA1D" authority="Hagen" authorityName="Hagen" box="[549,847,1447,1473]" class="Insecta" family="Euphaeidae" genus="Euphaea" kingdom="Animalia" order="Odonata" pageId="23" pageNumber="232" phylum="Arthropoda" rank="species" species="splendens">
<emphasis id="29AEEAA9E022D745FDCBE987FD34FA1D" box="[549,766,1447,1473]" italics="true" pageId="23" pageNumber="232">Euphaea splendens</emphasis>
Hagen
</taxonomicName>
from
<collectingCountry id="63CD762BE022D745FC7BE987FBC9FA1D" box="[917,1027,1447,1473]" name="Sri Lanka" pageId="23" pageNumber="232">Sri Lanka</collectingCountry>
and tentatively?
<taxonomicName id="DCDA4D38E022D745FB28E987FF2AFA39" authority="Hagen" authorityName="Hagen" authorityYear="1880" class="Insecta" family="Euphaeidae" genus="Anisopleura" kingdom="Animalia" order="Odonata" pageId="23" pageNumber="232" phylum="Arthropoda" rank="species" species="comes">
<emphasis id="29AEEAA9E022D745FB28E987FA56FA1C" box="[1222,1436,1447,1473]" italics="true" pageId="23" pageNumber="232">Anisopleura comes</emphasis>
Hagen
</taxonomicName>
and?
<taxonomicName id="DCDA4D38E022D745FEC9E9EBFD8AFA39" authority="Rambur" authorityName="Rambur" authorityYear="1842" box="[295,576,1483,1509]" class="Insecta" family="Euphaeidae" genus="Euphaea" kingdom="Animalia" order="Odonata" pageId="23" pageNumber="232" phylum="Arthropoda" rank="species" species="dispar">
<emphasis id="29AEEAA9E022D745FEC9E9EBFE11FA39" box="[295,475,1483,1509]" italics="true" pageId="23" pageNumber="232">Euphaea dispar</emphasis>
Rambur
</taxonomicName>
, however he incorrectly stated that the abdominal gills were born on S1–8. This was corrected by J.W Folsom, published in
<bibRefCitation id="7F4B4B4AE022D745FD2EE9CFFCA5F9D5" author="Packard, A. S." box="[704,879,1519,1545]" pageId="23" pageNumber="232" refId="ref26228" refString="Packard, A. S. (1898) A textbook of entomology, including the anatomy, physiology, embryology and metamorphoses of insects. Macmillan, London, xvii + 729 pp. [https: // www. biodiversitylibrary. org / item / 16837 # page / 493 / mode / 1 up]" type="book" year="1898">Packard (1898)</bibRefCitation>
, who also noted the tracheation of the abdominal gills and depicted a convincing habitus and detail of a single gill of
<taxonomicName id="DCDA4D38E022D745FC42EA33FBF1F9F1" box="[940,1083,1555,1581]" class="Insecta" family="Euphaeidae" genus="Euphaea" kingdom="Animalia" order="Odonata" pageId="23" pageNumber="232" phylum="Arthropoda" rank="species" species="splendens">
<emphasis id="29AEEAA9E022D745FC42EA33FBF1F9F1" box="[940,1083,1555,1581]" italics="true" pageId="23" pageNumber="232">E. splendens</emphasis>
</taxonomicName>
. A respiratory function for the abdominal gills was deduced by
<bibRefCitation id="7F4B4B4AE022D745FDFFEA17FD7AF98D" author="Hagen, H." box="[529,688,1591,1617]" pageId="23" pageNumber="232" pagination="lxv - lxvii" refId="ref23289" refString="Hagen, H. (1880) Essai d'un synopsis des larves de Calopterygines. [Comptes-rendus de la Socie´te´Entomologique de Belgique. Seance du 1 er Mai 1880]. Annales de la Societe entomologique de Belgique, 23, lxv - lxvii." type="journal article" year="1880">Hagen (1880)</bibRefCitation>
,
<bibRefCitation id="7F4B4B4AE022D745FD51EA17FCA7F98D" author="Packard, A. S." box="[703,877,1591,1617]" pageId="23" pageNumber="232" refId="ref26228" refString="Packard, A. S. (1898) A textbook of entomology, including the anatomy, physiology, embryology and metamorphoses of insects. Macmillan, London, xvii + 729 pp. [https: // www. biodiversitylibrary. org / item / 16837 # page / 493 / mode / 1 up]" type="book" year="1898">Packard (1898)</bibRefCitation>
and
<bibRefCitation id="7F4B4B4AE022D745FC49EA17FB9DF98D" author="Tillyard, R. J." box="[935,1111,1591,1617]" pageId="23" pageNumber="232" refId="ref27017" refString="Tillyard, R. J. (1917) The biology of dragonflies (Odonata or Paraneuroptera). University Press, Cambridge, xii + 396 pp., pls. 1 - 4 and 2 folded attachments excl. https: // doi. org / 10.5962 / bhl. title. 27401" type="book" year="1917">Tillyard (1917)</bibRefCitation>
, which was later queried by several authors following
<bibRefCitation id="7F4B4B4AE022D745FE59EA7BFDABF9A9" author="Fraser, F. C." box="[439,609,1627,1653]" pageId="23" pageNumber="232" refId="ref23008" refString="Fraser, F. C. (1933) The fauna of British India, including Ceylon and Burma. Odonata. Vol. I. Taylor &amp; Francis Ltd., London, xiii + 423 pp., 1 folded map excl." type="book" year="1933">Fraser’s (1933)</bibRefCitation>
assertion that they were grasping organs, before the respiratory hypothesis was settled conclusively by
<bibRefCitation id="7F4B4B4AE022D745FE23EA5FFDB2F945" author="Norling, U." box="[461,632,1663,1689]" pageId="23" pageNumber="232" pagination="343 - 389" refId="ref25522" refString="Norling, U. (1982) Structure and ontogeny of the lateral abdominal gills and the caudal gills in Euphaeidae (Odonata: Zygoptera) larvae. Zoologische Jahrbucher. Abteilung fur Anatomie und Ontogenie der Tiere, 107, 343 - 389." type="journal article" year="1982">Norling (1982)</bibRefCitation>
. Intrageneric variation, inadequate species sampling and poor artwork in some available descriptions sometimes makes a universal distinction between
<taxonomicName id="DCDA4D38E022D745FBEEEA83FB43F961" authorityName="Selys" authorityYear="1853" box="[1024,1161,1699,1725]" class="Insecta" family="Euphaeidae" genus="Anisopleura" kingdom="Animalia" order="Odonata" pageId="23" pageNumber="232" phylum="Arthropoda" rank="genus">
<emphasis id="29AEEAA9E022D745FBEEEA83FB43F961" box="[1024,1161,1699,1725]" italics="true" pageId="23" pageNumber="232">Anisopleura</emphasis>
</taxonomicName>
,
<taxonomicName id="DCDA4D38E022D745FB78EA83FAC9F961" authorityName="Selys" authorityYear="1853" box="[1174,1283,1699,1725]" class="Insecta" family="Euphaeidae" genus="Bayadera" kingdom="Animalia" order="Odonata" pageId="23" pageNumber="232" phylum="Arthropoda" rank="genus">
<emphasis id="29AEEAA9E022D745FB78EA83FAC9F961" box="[1174,1283,1699,1725]" italics="true" pageId="23" pageNumber="232">Bayadera</emphasis>
</taxonomicName>
and
<taxonomicName id="DCDA4D38E022D745FAD7EA83FA57F961" authorityName="Selys" authorityYear="1840" box="[1337,1437,1699,1725]" class="Insecta" family="Euphaeidae" genus="Euphaea" kingdom="Animalia" order="Odonata" pageId="23" pageNumber="232" phylum="Arthropoda" rank="genus">
<emphasis id="29AEEAA9E022D745FAD7EA83FA57F961" box="[1337,1437,1699,1725]" italics="true" pageId="23" pageNumber="232">Euphaea</emphasis>
</taxonomicName>
tentative, although certain species within these genera are distinctive. Important diagnostic characters include the shape of the postocular lobes of the head, relative length of antennae, distribution of tubercles and claviform setae on the head, development of male genital apophyses, armature of outer mandibles (
<figureCitation id="83E12A3EE022D745FBECEB2FFB95F8F5" box="[1026,1119,1807,1833]" captionStart="FIGURES 61–66. 61" captionStartId="12.[151,264,1841,1865]" captionTargetBox="[273,1314,156,1828]" captionTargetId="figure-21@12.[268,1319,151,1834]" captionTargetPageId="12" captionText="FIGURES 61–66. 61. Dysphaea gloriosa: detail of spines on genae and postocular lobe; underside (left), upperside (right) (modified from Nguyen et al. 2024). 62. Heterophaea barbata: dorsal view of head, showing (a) outer mandibular spines (b) genal spines partially visible and (c) right antenna (from Orr &amp; Hämäläinen 2024). 63. Anisopleura subplatystyla, dorsal view of head (original drawing). 64. Bayadera strigata, dorsal view of head (drawn from Yang &amp; Orr 2024). 65. Euphaea decorata, dorsal view of head. (drawn from Xu 2017). 66. Bayadera strigata, left labial palp with detail showing wavy outer margin of outer process of palpal lobe (arrowed), (drawn from Yang &amp; Orr 2024)." figureDoi="http://doi.org/10.5281/zenodo.13618343" httpUri="https://zenodo.org/record/13618343/files/figure.png" pageId="23" pageNumber="232">Fig. 62a</figureCitation>
), subocular spines (
<figureCitation id="83E12A3EE022D745FAD9EB2FFA5EF8F5" box="[1335,1428,1807,1833]" captionStart="FIGURES 61–66. 61" captionStartId="12.[151,264,1841,1865]" captionTargetBox="[273,1314,156,1828]" captionTargetId="figure-21@12.[268,1319,151,1834]" captionTargetPageId="12" captionText="FIGURES 61–66. 61. Dysphaea gloriosa: detail of spines on genae and postocular lobe; underside (left), upperside (right) (modified from Nguyen et al. 2024). 62. Heterophaea barbata: dorsal view of head, showing (a) outer mandibular spines (b) genal spines partially visible and (c) right antenna (from Orr &amp; Hämäläinen 2024). 63. Anisopleura subplatystyla, dorsal view of head (original drawing). 64. Bayadera strigata, dorsal view of head (drawn from Yang &amp; Orr 2024). 65. Euphaea decorata, dorsal view of head. (drawn from Xu 2017). 66. Bayadera strigata, left labial palp with detail showing wavy outer margin of outer process of palpal lobe (arrowed), (drawn from Yang &amp; Orr 2024)." figureDoi="http://doi.org/10.5281/zenodo.13618343" httpUri="https://zenodo.org/record/13618343/files/figure.png" pageId="23" pageNumber="232">Fig. 62b</figureCitation>
) and presence of supracoxal and coxal spurs (
<bibRefCitation id="7F4B4B4AE022D745FD6BEB13FC12F891" author="Keetapithchayakul, T. S. &amp; Sripanya, J. &amp; Phlai-Ngam, S. &amp; Tungpairojwong, N." box="[645,984,1843,1869]" pageId="23" pageNumber="232" pagination="98 - 110" refId="ref24014" refString="Keetapithchayakul, T. S., Sripanya, J., Phlai-Ngam, S. &amp; Tungpairojwong, N. (2020) Description of the final stadium larva of Bayadera serrata Davies &amp; Yang, 1996 (Odonata: Euphaeidae) from Thailand. Zootaxa, 4894 (1), 98 - 110. https: // doi. org / 10.11646 / zootaxa. 4894.1.5" type="journal article" year="2020">
Keetapithchayakul
<emphasis id="29AEEAA9E022D745FC8EEB13FC5DF891" box="[864,919,1843,1869]" italics="true" pageId="23" pageNumber="232">et al.</emphasis>
2020
</bibRefCitation>
,
<bibRefCitation id="7F4B4B4AE022D745FC0FEB13FB6CF891" author="Yang, G. - H. &amp; Orr, A. G." box="[993,1190,1843,1869]" pageId="23" pageNumber="232" pagination="11 - 18" refId="ref27987" refString="Yang, G. - H. &amp; Orr, A. G. (2024) Descriptions of larvae of Bayadera strigata Davies &amp; Yang, 1996 (Odonata: Euphaeidae). International Journal of Odonatology, 27, 11 - 18. https: // doi. org / 10.48156 / 1388.2024.1917250" type="journal article" year="2024">Yang &amp; Orr 2024</bibRefCitation>
,
<bibRefCitation id="7F4B4B4AE022D745FB5EEB13FF3CF8AD" author="Keetapithchayakul, T. S. &amp; Orr, A. G. &amp; Wongkamhaeng, K." pageId="23" pageNumber="232" pagination="529 - 542" refId="ref24083" refString="Keetapithchayakul, T. S., Orr, A. G. &amp; Wongkamhaeng, K. (2024) Description of the final stadium larva of Anisopleura furcata Selys, 1891 (Odonata: Euphaeidae) from Thailand. Zootaxa, 5415 (4), 529 - 542. https: // doi. org / 10.11646 / zootaxa. 5415.4.2" type="journal article" year="2024">
Keetapithchayakul
<emphasis id="29AEEAA9E022D745FA66EB13FF67F8AD" italics="true" pageId="23" pageNumber="232">et al</emphasis>
. 2024
</bibRefCitation>
,
<bibRefCitation id="7F4B4B4AE022D745FEEFEB77FDDDF8AD" author="Orr, A. G. &amp; Hamalainen, M." box="[257,535,1879,1905]" pageId="23" pageNumber="232" pagination="169 - 182" refId="ref26097" refString="Orr, A. G. &amp; Hamalainen, M. (2024) The larva of the endangered Heterophaea barbata (Martin, 1902), from Luzon Island, the Philippines (Odonata: Euphaeidae). Odonatologica, 53, 169 - 182. https: // doi. org / 10.60024 / odon. v 53 i 1 - 2. a 9" type="journal article" year="2024">Orr &amp; Hämäläinen 2024</bibRefCitation>
).
</paragraph>
<paragraph id="1B6536BBE022D745FF29EB5BFC94F849" blockId="23.[151,1437,1051,1941]" box="[199,862,1915,1941]" pageId="23" pageNumber="232">
Larva unknown:
<taxonomicName id="DCDA4D38E022D745FE68EB5BFDC6F849" authorityName="Ris" authorityYear="1930" box="[390,524,1915,1941]" class="Insecta" family="Euphaeidae" genus="Cyclophaea" kingdom="Animalia" order="Odonata" pageId="23" pageNumber="232" phylum="Arthropoda" rank="genus">
<emphasis id="29AEEAA9E022D745FE68EB5BFDC6F849" box="[390,524,1915,1941]" italics="true" pageId="23" pageNumber="232">Cyclophaea</emphasis>
</taxonomicName>
(
<specimenCount id="0DDCFD32E022D745FDF0EB5BFD99F849" box="[542,595,1915,1941]" count="1" pageId="23" pageNumber="232" type="generic">1 sp.</specimenCount>
),
<taxonomicName id="DCDA4D38E022D745FD87EB5BFCC5F849" authorityName="Asahina" authorityYear="1978" box="[617,783,1915,1941]" class="Insecta" family="Euphaeidae" genus="Schmidtiphaea" kingdom="Animalia" order="Odonata" pageId="23" pageNumber="232" phylum="Arthropoda" rank="genus">
<emphasis id="29AEEAA9E022D745FD87EB5BFCC5F849" box="[617,783,1915,1941]" italics="true" pageId="23" pageNumber="232">Schmidtiphaea</emphasis>
</taxonomicName>
(
<specimenCount id="0DDCFD32E022D745FCCFEB5BFC9FF849" box="[801,853,1915,1941]" count="1" pageId="23" pageNumber="232" type="generic">1 sp.</specimenCount>
)
</paragraph>
</subSubSection>
<subSubSection id="53C06530E02ED749FF79ECB7FE8CFF6D" box="[151,326,151,177]" pageId="27" pageNumber="236" type="multiple">
<paragraph id="1B6536BBE02ED749FF79ECB7FE8CFF6D" blockId="27.[151,326,151,177]" box="[151,326,151,177]" pageId="27" pageNumber="236">
<heading id="402D81D7E02ED749FF79ECB7FE8CFF6D" bold="true" box="[151,326,151,177]" fontSize="11" level="1" pageId="27" pageNumber="236" reason="1">
<emphasis id="29AEEAA9E02ED749FF79ECB7FE8CFF6D" bold="true" box="[151,326,151,177]" pageId="27" pageNumber="236">Key to genera</emphasis>
</heading>
</paragraph>
</subSubSection>
<subSubSection id="53C06530E02ED749FF79ECFCFA56FCAB" pageId="27" pageNumber="236" type="key">
<key id="6F4B618AE02ED749FF79ECFCFA56FCAB" pageId="27" pageNumber="236">
<keyStep id="A02E2E1EE02ED749FF79ECFCFA56FE95" pageId="27" pageNumber="236">
<paragraph id="1B6536BBE02ED749FF79ECFCFA57FEF1" blockId="27.[151,1437,220,887]" pageId="27" pageNumber="236">
<keyLead id="A02B958EE02ED749FF79ECFCFA57FEF1" pageId="27" pageNumber="236">
1 Head massive, nearly twice width of pronotum; genae swollen and projecting well beyond eyes in dorsal view bearing numerous short heavy spines; strong spines present on bulbous postocular lobe (
<figureCitation id="83E12A3EE02ED749FC91ECD9FC00FED3" box="[895,970,249,271]" captionStart="FIGURES 57–60. 57" captionStartId="11.[151,264,1933,1957]" captionTargetBox="[193,1384,155,1907]" captionTargetId="figure-15@11.[188,1400,151,1917]" captionTargetPageId="11" captionText="FIGURES 57–60. 57. Dysphaea gloriosa, habitus (drawn from Nguyen et al. 2024). 58. Bayadera strigata, habitus (drawn from Yang &amp; Orr 2024). 59. Euphaea decorata, habitus. (drawn from Xu 2017). 60. Anisopleura furcata, habitus (drawn from Keetapithchayakul et al. 2024)." figureDoi="http://doi.org/10.5281/zenodo.13618341" httpUri="https://zenodo.org/record/13618341/files/figure.png" pageId="27" pageNumber="236">Figs 57</figureCitation>
,
<figureCitation id="83E12A3EE02ED749FC3AECD9FC27FED3" box="[980,1005,249,271]" captionStart="FIGURES 61–66. 61" captionStartId="12.[151,264,1841,1865]" captionTargetBox="[273,1314,156,1828]" captionTargetId="figure-21@12.[268,1319,151,1834]" captionTargetPageId="12" captionText="FIGURES 61–66. 61. Dysphaea gloriosa: detail of spines on genae and postocular lobe; underside (left), upperside (right) (modified from Nguyen et al. 2024). 62. Heterophaea barbata: dorsal view of head, showing (a) outer mandibular spines (b) genal spines partially visible and (c) right antenna (from Orr &amp; Hämäläinen 2024). 63. Anisopleura subplatystyla, dorsal view of head (original drawing). 64. Bayadera strigata, dorsal view of head (drawn from Yang &amp; Orr 2024). 65. Euphaea decorata, dorsal view of head. (drawn from Xu 2017). 66. Bayadera strigata, left labial palp with detail showing wavy outer margin of outer process of palpal lobe (arrowed), (drawn from Yang &amp; Orr 2024)." figureDoi="http://doi.org/10.5281/zenodo.13618343" httpUri="https://zenodo.org/record/13618343/files/figure.png" pageId="27" pageNumber="236">61</figureCitation>
)..........................
<taxonomicName id="DCDA4D38E02ED749FAD5ECD9FA56FED3" authorityName="Selys" authorityYear="1853" box="[1339,1436,249,271]" class="Insecta" family="Euphaeidae" genus="Dysphaea" kingdom="Animalia" order="Odonata" pageId="27" pageNumber="236" phylum="Arthropoda" rank="genus">
<emphasis id="29AEEAA9E02ED749FAD5ECD9FA56FED3" bold="true" box="[1339,1436,249,271]" italics="true" pageId="27" pageNumber="236">Dysphaea</emphasis>
</taxonomicName>
[widespread west of Wallace’s line]
</keyLead>
</paragraph>
<paragraph id="1B6536BBE02ED749FF79ED14FA56FE95" blockId="27.[151,1437,220,887]" box="[151,1436,308,330]" pageId="27" pageNumber="236">
<keyLead id="A02B958EE02ED749FF79ED14FA56FE95" box="[151,1436,308,330]" pageId="27" pageNumber="236">1’ Head narrower in relation to pronotum, genae not swollen, no strong spines on postocular lobe which is not bulbous....... 2</keyLead>
</paragraph>
</keyStep>
<keyStep id="A02E2E1EE02ED749FF79ED72FA56FE25" pageId="27" pageNumber="236">
<paragraph id="1B6536BBE02ED749FF79ED72FA56FE63" blockId="27.[151,1437,220,887]" pageId="27" pageNumber="236">
<keyLead id="A02B958EE02ED749FF79ED72FA56FE63" pageId="27" pageNumber="236">
2 (1’) Genae with 6–7 long subocular spines (greater than one third width of eye viewed ventrally); antennae with 12–15 antennomeres including pedicel and scape; outer face of mandibles with 2 or 3 rows of strong spines [4 ventral, 5 dorsal, 0–4 central] (
<figureCitation id="83E12A3EE02ED749FA9BED4FFF37FE7E" captionStart="FIGURES 61–66. 61" captionStartId="12.[151,264,1841,1865]" captionTargetBox="[273,1314,156,1828]" captionTargetId="figure-21@12.[268,1319,151,1834]" captionTargetPageId="12" captionText="FIGURES 61–66. 61. Dysphaea gloriosa: detail of spines on genae and postocular lobe; underside (left), upperside (right) (modified from Nguyen et al. 2024). 62. Heterophaea barbata: dorsal view of head, showing (a) outer mandibular spines (b) genal spines partially visible and (c) right antenna (from Orr &amp; Hämäläinen 2024). 63. Anisopleura subplatystyla, dorsal view of head (original drawing). 64. Bayadera strigata, dorsal view of head (drawn from Yang &amp; Orr 2024). 65. Euphaea decorata, dorsal view of head. (drawn from Xu 2017). 66. Bayadera strigata, left labial palp with detail showing wavy outer margin of outer process of palpal lobe (arrowed), (drawn from Yang &amp; Orr 2024)." figureDoi="http://doi.org/10.5281/zenodo.13618343" httpUri="https://zenodo.org/record/13618343/files/figure.png" pageId="27" pageNumber="236">Fig. 62</figureCitation>
).......................................................................................
<taxonomicName id="DCDA4D38E02ED749FAF2EDACFA56FE7E" authorityName="Cowley" authorityYear="1934" box="[1308,1436,396,418]" class="Insecta" family="Euphaeidae" genus="Heterophaea" kingdom="Animalia" order="Odonata" pageId="27" pageNumber="236" phylum="Arthropoda" rank="genus">
<emphasis id="29AEEAA9E02ED749FAF2EDACFA56FE7E" bold="true" box="[1308,1436,396,418]" italics="true" pageId="27" pageNumber="236">Heterophaea</emphasis>
</taxonomicName>
[endemic to north-eastern Luzon,
<collectingCountry id="63CD762BE02ED749FAC6ED89FA5DFE63" box="[1320,1431,425,447]" name="Philippines" pageId="27" pageNumber="236">Philippines</collectingCountry>
]
</keyLead>
</paragraph>
<paragraph id="1B6536BBE02ED749FF79EDE7FA56FE25" blockId="27.[151,1437,220,887]" pageId="27" pageNumber="236">
<keyLead id="A02B958EE02ED749FF79EDE7FA56FE25" pageId="27" pageNumber="236">2’ Genae with 5 or fewer long subocular spines; antennae with 7, rarely 8 antennomeres; outer margin of mandible with only one row of spines, or just one or two long spines............................................................... 3</keyLead>
</paragraph>
</keyStep>
<keyStep id="A02E2E1EE02ED749FF79EE22FA56FD75" pageId="27" pageNumber="236">
<paragraph id="1B6536BBE02ED749FF79EE22FA56FD8E" blockId="27.[151,1437,220,887]" pageId="27" pageNumber="236">
<keyLead id="A02B958EE02ED749FF79EE22FA56FD8E" pageId="27" pageNumber="236">
3 (2’) Antennae distinctly shorter than head from occipital margin to base of labrum; postocular lobes shallow, weakly rounded (
<figureCitation id="83E12A3EE02ED749FA9BEE21FF37FDE9" captionStart="FIGURES 61–66. 61" captionStartId="12.[151,264,1841,1865]" captionTargetBox="[273,1314,156,1828]" captionTargetId="figure-21@12.[268,1319,151,1834]" captionTargetPageId="12" captionText="FIGURES 61–66. 61. Dysphaea gloriosa: detail of spines on genae and postocular lobe; underside (left), upperside (right) (modified from Nguyen et al. 2024). 62. Heterophaea barbata: dorsal view of head, showing (a) outer mandibular spines (b) genal spines partially visible and (c) right antenna (from Orr &amp; Hämäläinen 2024). 63. Anisopleura subplatystyla, dorsal view of head (original drawing). 64. Bayadera strigata, dorsal view of head (drawn from Yang &amp; Orr 2024). 65. Euphaea decorata, dorsal view of head. (drawn from Xu 2017). 66. Bayadera strigata, left labial palp with detail showing wavy outer margin of outer process of palpal lobe (arrowed), (drawn from Yang &amp; Orr 2024)." figureDoi="http://doi.org/10.5281/zenodo.13618343" httpUri="https://zenodo.org/record/13618343/files/figure.png" pageId="27" pageNumber="236">Fig. 63</figureCitation>
); labrum, frons and vertex with smattering of variably developed tubercles, low or raised and finger-like; caudal gills tapered to long, fine filament (
<figureCitation id="83E12A3EE02ED749FE54EE1CFDCAFD8E" box="[442,512,572,594]" captionStart="FIGURES 67–74. 67" captionStartId="13.[151,264,1814,1838]" captionTargetBox="[256,1340,160,1799]" captionTargetId="figure-15@13.[246,1342,151,1807]" captionTargetPageId="13" captionText="FIGURES 67–74. 67. Anisopleura subplatystyla, caudal gills dorsal view (original drawing). 68. Bayadera strigata, caudal gills dorsal view, (drawn from Yang &amp; Orr 2024). 69. Euphaea decorata, caudal gills dorsal view (drawn from Xu 2017). 70. Euphaea superba, caudal gills dorsal view (redrawn from Wu et al. 2017). 71. Bayadera strigata: male genital apophyses; ventral (left), right lateral (right), (drawn from Yang &amp; Orr 2024). 72. Euphaea decorata, male genital apophyses; ventral view (drawn from Xu 2017). 73. Euphaea superba, terminal processes of palpal lobe ventral view (movable hook truncated), (redrawn from Wu et al. 2017). 74. Anisopleura furcata, left labial palp showing terminal processes of palpal lobe, dorsal view (redrawn from Keetapithchayakul et al. 2024)." figureDoi="http://doi.org/10.5281/zenodo.13618345" httpUri="https://zenodo.org/record/13618345/files/figure.png" pageId="27" pageNumber="236">Fig. 67</figureCitation>
).................................................................
<taxonomicName id="DCDA4D38E02ED749FACDEE1CFA56FD8E" authorityName="Selys" authorityYear="1853" box="[1315,1436,572,594]" class="Insecta" family="Euphaeidae" genus="Anisopleura" kingdom="Animalia" order="Odonata" pageId="27" pageNumber="236" phylum="Arthropoda" rank="genus">
<emphasis id="29AEEAA9E02ED749FACDEE1CFA56FD8E" bold="true" box="[1315,1436,572,594]" italics="true" pageId="27" pageNumber="236">Anisopleura</emphasis>
</taxonomicName>
</keyLead>
</paragraph>
<paragraph id="1B6536BBE02ED749FF79EE7AFA56FD75" blockId="27.[151,1437,220,887]" pageId="27" pageNumber="236">
<keyLead id="A02B958EE02ED749FF79EE7AFA56FD75" pageId="27" pageNumber="236">
3’ Antennae at least as long as or longer than head from occipital margin to base of labrum postocular lobes deep and strongly rounded at corners, sometimes appearing square by virtue of tuft of setae (
<figureCitation id="83E12A3EE02ED749FC73EE57FBC0FD51" box="[925,1034,631,653]" captionStart="FIGURES 61–66. 61" captionStartId="12.[151,264,1841,1865]" captionTargetBox="[273,1314,156,1828]" captionTargetId="figure-21@12.[268,1319,151,1834]" captionTargetPageId="12" captionText="FIGURES 61–66. 61. Dysphaea gloriosa: detail of spines on genae and postocular lobe; underside (left), upperside (right) (modified from Nguyen et al. 2024). 62. Heterophaea barbata: dorsal view of head, showing (a) outer mandibular spines (b) genal spines partially visible and (c) right antenna (from Orr &amp; Hämäläinen 2024). 63. Anisopleura subplatystyla, dorsal view of head (original drawing). 64. Bayadera strigata, dorsal view of head (drawn from Yang &amp; Orr 2024). 65. Euphaea decorata, dorsal view of head. (drawn from Xu 2017). 66. Bayadera strigata, left labial palp with detail showing wavy outer margin of outer process of palpal lobe (arrowed), (drawn from Yang &amp; Orr 2024)." figureDoi="http://doi.org/10.5281/zenodo.13618343" httpUri="https://zenodo.org/record/13618343/files/figure.png" pageId="27" pageNumber="236">Figs 64, 65</figureCitation>
); tubercles on head lacking or very sparse; caudal gills terminating in long filament or a short blunt cone, sometimes constricted basally......................... 4
</keyLead>
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<keyStep id="A02E2E1EE02ED749FF79EE92FA56FCAB" pageId="27" pageNumber="236">
<paragraph id="1B6536BBE02ED749FF79EE92FA57FCDE" blockId="27.[151,1437,220,887]" pageId="27" pageNumber="236">
<keyLead id="A02B958EE02ED749FF79EE92FA57FCDE" pageId="27" pageNumber="236">
4 (3’) Caudal gills terminating in a short blunt cone (
<figureCitation id="83E12A3EE02ED749FD57EE91FCCDFD1B" box="[697,775,689,711]" captionStart="FIGURES 57–60. 57" captionStartId="11.[151,264,1933,1957]" captionTargetBox="[193,1384,155,1907]" captionTargetId="figure-15@11.[188,1400,151,1917]" captionTargetPageId="11" captionText="FIGURES 57–60. 57. Dysphaea gloriosa, habitus (drawn from Nguyen et al. 2024). 58. Bayadera strigata, habitus (drawn from Yang &amp; Orr 2024). 59. Euphaea decorata, habitus. (drawn from Xu 2017). 60. Anisopleura furcata, habitus (drawn from Keetapithchayakul et al. 2024)." figureDoi="http://doi.org/10.5281/zenodo.13618341" httpUri="https://zenodo.org/record/13618341/files/figure.png" pageId="27" pageNumber="236">Figs 58</figureCitation>
,
<figureCitation id="83E12A3EE02ED749FCFAEE91FCE7FD1B" box="[788,813,689,711]" captionStart="FIGURES 67–74. 67" captionStartId="13.[151,264,1814,1838]" captionTargetBox="[256,1340,160,1799]" captionTargetId="figure-15@13.[246,1342,151,1807]" captionTargetPageId="13" captionText="FIGURES 67–74. 67. Anisopleura subplatystyla, caudal gills dorsal view (original drawing). 68. Bayadera strigata, caudal gills dorsal view, (drawn from Yang &amp; Orr 2024). 69. Euphaea decorata, caudal gills dorsal view (drawn from Xu 2017). 70. Euphaea superba, caudal gills dorsal view (redrawn from Wu et al. 2017). 71. Bayadera strigata: male genital apophyses; ventral (left), right lateral (right), (drawn from Yang &amp; Orr 2024). 72. Euphaea decorata, male genital apophyses; ventral view (drawn from Xu 2017). 73. Euphaea superba, terminal processes of palpal lobe ventral view (movable hook truncated), (redrawn from Wu et al. 2017). 74. Anisopleura furcata, left labial palp showing terminal processes of palpal lobe, dorsal view (redrawn from Keetapithchayakul et al. 2024)." figureDoi="http://doi.org/10.5281/zenodo.13618345" httpUri="https://zenodo.org/record/13618345/files/figure.png" pageId="27" pageNumber="236">68</figureCitation>
) occasionally basally constricted; antennae clearly longer than head from occipital margin to base of labrum (
<figureCitation id="83E12A3EE02ED749FD5AEEEFFD35FD39" box="[692,767,719,741]" captionStart="FIGURES 61–66. 61" captionStartId="12.[151,264,1841,1865]" captionTargetBox="[273,1314,156,1828]" captionTargetId="figure-21@12.[268,1319,151,1834]" captionTargetPageId="12" captionText="FIGURES 61–66. 61. Dysphaea gloriosa: detail of spines on genae and postocular lobe; underside (left), upperside (right) (modified from Nguyen et al. 2024). 62. Heterophaea barbata: dorsal view of head, showing (a) outer mandibular spines (b) genal spines partially visible and (c) right antenna (from Orr &amp; Hämäläinen 2024). 63. Anisopleura subplatystyla, dorsal view of head (original drawing). 64. Bayadera strigata, dorsal view of head (drawn from Yang &amp; Orr 2024). 65. Euphaea decorata, dorsal view of head. (drawn from Xu 2017). 66. Bayadera strigata, left labial palp with detail showing wavy outer margin of outer process of palpal lobe (arrowed), (drawn from Yang &amp; Orr 2024)." figureDoi="http://doi.org/10.5281/zenodo.13618343" httpUri="https://zenodo.org/record/13618343/files/figure.png" pageId="27" pageNumber="236">Fig. 64</figureCitation>
); outer margin of outer process of labial palp wavy (
<figureCitation id="83E12A3EE02ED749FAE1EEEFFA93FD39" box="[1295,1369,719,741]" captionStart="FIGURES 61–66. 61" captionStartId="12.[151,264,1841,1865]" captionTargetBox="[273,1314,156,1828]" captionTargetId="figure-21@12.[268,1319,151,1834]" captionTargetPageId="12" captionText="FIGURES 61–66. 61. Dysphaea gloriosa: detail of spines on genae and postocular lobe; underside (left), upperside (right) (modified from Nguyen et al. 2024). 62. Heterophaea barbata: dorsal view of head, showing (a) outer mandibular spines (b) genal spines partially visible and (c) right antenna (from Orr &amp; Hämäläinen 2024). 63. Anisopleura subplatystyla, dorsal view of head (original drawing). 64. Bayadera strigata, dorsal view of head (drawn from Yang &amp; Orr 2024). 65. Euphaea decorata, dorsal view of head. (drawn from Xu 2017). 66. Bayadera strigata, left labial palp with detail showing wavy outer margin of outer process of palpal lobe (arrowed), (drawn from Yang &amp; Orr 2024)." figureDoi="http://doi.org/10.5281/zenodo.13618343" httpUri="https://zenodo.org/record/13618343/files/figure.png" pageId="27" pageNumber="236">Fig. 66</figureCitation>
); male gonapophyses down-curved and sausage shaped, overlapping S10 (
<figureCitation id="83E12A3EE02ED749FC86EECCFC67FCDD" box="[872,941,748,770]" captionStart="FIGURES 67–74. 67" captionStartId="13.[151,264,1814,1838]" captionTargetBox="[256,1340,160,1799]" captionTargetId="figure-15@13.[246,1342,151,1807]" captionTargetPageId="13" captionText="FIGURES 67–74. 67. Anisopleura subplatystyla, caudal gills dorsal view (original drawing). 68. Bayadera strigata, caudal gills dorsal view, (drawn from Yang &amp; Orr 2024). 69. Euphaea decorata, caudal gills dorsal view (drawn from Xu 2017). 70. Euphaea superba, caudal gills dorsal view (redrawn from Wu et al. 2017). 71. Bayadera strigata: male genital apophyses; ventral (left), right lateral (right), (drawn from Yang &amp; Orr 2024). 72. Euphaea decorata, male genital apophyses; ventral view (drawn from Xu 2017). 73. Euphaea superba, terminal processes of palpal lobe ventral view (movable hook truncated), (redrawn from Wu et al. 2017). 74. Anisopleura furcata, left labial palp showing terminal processes of palpal lobe, dorsal view (redrawn from Keetapithchayakul et al. 2024)." figureDoi="http://doi.org/10.5281/zenodo.13618345" httpUri="https://zenodo.org/record/13618345/files/figure.png" pageId="27" pageNumber="236">Fig. 71</figureCitation>
)................................
<emphasis id="29AEEAA9E02ED749FAD0EECCFA57FCDE" bold="true" box="[1342,1437,748,770]" pageId="27" pageNumber="236">
<emphasis id="29AEEAA9E02ED749FAD0EECCFA84FCDD" bold="true" box="[1342,1358,748,769]" italics="true" pageId="27" pageNumber="236">B</emphasis>
a
<emphasis id="29AEEAA9E02ED749FAB4EECCFA57FCDE" bold="true" box="[1370,1437,748,770]" italics="true" pageId="27" pageNumber="236">yadera</emphasis>
</emphasis>
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<paragraph id="1B6536BBE02ED749FF79EF2AFA56FCAB" blockId="27.[151,1437,220,887]" pageId="27" pageNumber="236">
<keyLead id="A02B958EE02ED749FF79EF2AFA56FCAB" pageId="27" pageNumber="236">
4’ Tip of caudal gills with long filament usually arising abruptly from sac of gill (
<figureCitation id="83E12A3EE02ED749FC30EF29FBE3FCC3" box="[990,1065,777,799]" captionStart="FIGURES 57–60. 57" captionStartId="11.[151,264,1933,1957]" captionTargetBox="[193,1384,155,1907]" captionTargetId="figure-15@11.[188,1400,151,1917]" captionTargetPageId="11" captionText="FIGURES 57–60. 57. Dysphaea gloriosa, habitus (drawn from Nguyen et al. 2024). 58. Bayadera strigata, habitus (drawn from Yang &amp; Orr 2024). 59. Euphaea decorata, habitus. (drawn from Xu 2017). 60. Anisopleura furcata, habitus (drawn from Keetapithchayakul et al. 2024)." figureDoi="http://doi.org/10.5281/zenodo.13618341" httpUri="https://zenodo.org/record/13618341/files/figure.png" pageId="27" pageNumber="236">Figs 59</figureCitation>
,
<figureCitation id="83E12A3EE02ED749FBDAEF29FB87FCC3" box="[1076,1101,777,799]" captionStart="FIGURES 67–74. 67" captionStartId="13.[151,264,1814,1838]" captionTargetBox="[256,1340,160,1799]" captionTargetId="figure-15@13.[246,1342,151,1807]" captionTargetPageId="13" captionText="FIGURES 67–74. 67. Anisopleura subplatystyla, caudal gills dorsal view (original drawing). 68. Bayadera strigata, caudal gills dorsal view, (drawn from Yang &amp; Orr 2024). 69. Euphaea decorata, caudal gills dorsal view (drawn from Xu 2017). 70. Euphaea superba, caudal gills dorsal view (redrawn from Wu et al. 2017). 71. Bayadera strigata: male genital apophyses; ventral (left), right lateral (right), (drawn from Yang &amp; Orr 2024). 72. Euphaea decorata, male genital apophyses; ventral view (drawn from Xu 2017). 73. Euphaea superba, terminal processes of palpal lobe ventral view (movable hook truncated), (redrawn from Wu et al. 2017). 74. Anisopleura furcata, left labial palp showing terminal processes of palpal lobe, dorsal view (redrawn from Keetapithchayakul et al. 2024)." figureDoi="http://doi.org/10.5281/zenodo.13618345" httpUri="https://zenodo.org/record/13618345/files/figure.png" pageId="27" pageNumber="236">69</figureCitation>
) or sac elongate with a short blunt filament (
<taxonomicName id="DCDA4D38E02ED749FED1EF07FE6CFCE0" authorityName="Kimmins" authorityYear="1936" box="[319,422,807,828]" class="Insecta" family="Euphaeidae" genus="Euphaea" kingdom="Animalia" order="Odonata" pageId="27" pageNumber="236" phylum="Arthropoda" rank="species" species="superba">
<emphasis id="29AEEAA9E02ED749FED1EF07FE6CFCE0" box="[319,422,807,828]" italics="true" pageId="27" pageNumber="236">E. superba</emphasis>
</taxonomicName>
<figureCitation id="83E12A3EE02ED749FE45EF07FE38FCE0" box="[427,498,807,829]" captionStart="FIGURES 67–74. 67" captionStartId="13.[151,264,1814,1838]" captionTargetBox="[256,1340,160,1799]" captionTargetId="figure-15@13.[246,1342,151,1807]" captionTargetPageId="13" captionText="FIGURES 67–74. 67. Anisopleura subplatystyla, caudal gills dorsal view (original drawing). 68. Bayadera strigata, caudal gills dorsal view, (drawn from Yang &amp; Orr 2024). 69. Euphaea decorata, caudal gills dorsal view (drawn from Xu 2017). 70. Euphaea superba, caudal gills dorsal view (redrawn from Wu et al. 2017). 71. Bayadera strigata: male genital apophyses; ventral (left), right lateral (right), (drawn from Yang &amp; Orr 2024). 72. Euphaea decorata, male genital apophyses; ventral view (drawn from Xu 2017). 73. Euphaea superba, terminal processes of palpal lobe ventral view (movable hook truncated), (redrawn from Wu et al. 2017). 74. Anisopleura furcata, left labial palp showing terminal processes of palpal lobe, dorsal view (redrawn from Keetapithchayakul et al. 2024)." figureDoi="http://doi.org/10.5281/zenodo.13618345" httpUri="https://zenodo.org/record/13618345/files/figure.png" pageId="27" pageNumber="236">Fig. 70</figureCitation>
); antennae about equal to head from occipital margin to base of labrum or slightly longer (
<figureCitation id="83E12A3EE02ED749FAA7EF07FA5AFCE1" box="[1353,1424,807,829]" captionStart="FIGURES 61–66. 61" captionStartId="12.[151,264,1841,1865]" captionTargetBox="[273,1314,156,1828]" captionTargetId="figure-21@12.[268,1319,151,1834]" captionTargetPageId="12" captionText="FIGURES 61–66. 61. Dysphaea gloriosa: detail of spines on genae and postocular lobe; underside (left), upperside (right) (modified from Nguyen et al. 2024). 62. Heterophaea barbata: dorsal view of head, showing (a) outer mandibular spines (b) genal spines partially visible and (c) right antenna (from Orr &amp; Hämäläinen 2024). 63. Anisopleura subplatystyla, dorsal view of head (original drawing). 64. Bayadera strigata, dorsal view of head (drawn from Yang &amp; Orr 2024). 65. Euphaea decorata, dorsal view of head. (drawn from Xu 2017). 66. Bayadera strigata, left labial palp with detail showing wavy outer margin of outer process of palpal lobe (arrowed), (drawn from Yang &amp; Orr 2024)." figureDoi="http://doi.org/10.5281/zenodo.13618343" httpUri="https://zenodo.org/record/13618343/files/figure.png" pageId="27" pageNumber="236">Fig. 65</figureCitation>
); outer margin of outer process of labial palp smooth; male gonapophyses small and triangular or vestigial, not overlapping S10 (
<figureCitation id="83E12A3EE02ED749FF07EF41FEE5FCAB" box="[233,303,865,887]" captionStart="FIGURES 67–74. 67" captionStartId="13.[151,264,1814,1838]" captionTargetBox="[256,1340,160,1799]" captionTargetId="figure-15@13.[246,1342,151,1807]" captionTargetPageId="13" captionText="FIGURES 67–74. 67. Anisopleura subplatystyla, caudal gills dorsal view (original drawing). 68. Bayadera strigata, caudal gills dorsal view, (drawn from Yang &amp; Orr 2024). 69. Euphaea decorata, caudal gills dorsal view (drawn from Xu 2017). 70. Euphaea superba, caudal gills dorsal view (redrawn from Wu et al. 2017). 71. Bayadera strigata: male genital apophyses; ventral (left), right lateral (right), (drawn from Yang &amp; Orr 2024). 72. Euphaea decorata, male genital apophyses; ventral view (drawn from Xu 2017). 73. Euphaea superba, terminal processes of palpal lobe ventral view (movable hook truncated), (redrawn from Wu et al. 2017). 74. Anisopleura furcata, left labial palp showing terminal processes of palpal lobe, dorsal view (redrawn from Keetapithchayakul et al. 2024)." figureDoi="http://doi.org/10.5281/zenodo.13618345" httpUri="https://zenodo.org/record/13618345/files/figure.png" pageId="27" pageNumber="236">Fig. 72</figureCitation>
)......................................................................................
<taxonomicName id="DCDA4D38E02ED749FAADEF41FA56FCAB" authorityName="Selys" authorityYear="1840" box="[1347,1436,865,887]" class="Insecta" family="Euphaeidae" genus="Euphaea" kingdom="Animalia" order="Odonata" pageId="27" pageNumber="236" phylum="Arthropoda" rank="genus">
<emphasis id="29AEEAA9E02ED749FAADEF41FA56FCAB" bold="true" box="[1347,1436,865,887]" italics="true" pageId="27" pageNumber="236">Euphaea</emphasis>
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<paragraph id="1B6536BBE023D744FF79E91AFEE9FA89" blockId="22.[151,1437,1338,1905]" box="[151,291,1338,1365]" pageId="22" pageNumber="231">
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<taxonomicName id="DCDA4D38E023D744FF79E91AFEE9FA89" ID-CoL="78PK" authorityName="Rambur" authorityYear="1842" box="[151,291,1338,1365]" class="Insecta" family="Chlorocyphidae" genus="Rhinocypha" kingdom="Animalia" order="Odonata" pageId="22" pageNumber="231" phylum="Arthropoda" rank="genus">
<emphasis id="29AEEAA9E023D744FF79E91AFEE9FA89" bold="true" box="[151,291,1338,1365]" italics="true" pageId="22" pageNumber="231">Rhinocypha</emphasis>
</taxonomicName>
</heading>
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<subSubSection id="53C06530E023D744FF79E97FFE00F8AD" pageId="22" pageNumber="231" type="discussion">
<paragraph id="1B6536BBE023D744FF79E97FFE00F8AD" blockId="22.[151,1437,1338,1905]" pageId="22" pageNumber="231">
The 45 species of this diverse genus occur throughout most of the Oriental Realm with two species confined to the western Oceanic Realm (sensu
<bibRefCitation id="7F4B4B4AE023D744FE1CE9A3FD6EFA41" author="Holt, B. G. &amp; Lessard, J. - P. &amp; Borregaard, M. K. &amp; Fritz, S. A. &amp; Araujo, M. B. &amp; Dimitrov, D. &amp; Fabre, P. - H. &amp; Graham, C. H. &amp; Graves, G. R. &amp; Jonsson, K. A. &amp; Nogues-Bravo, D. &amp; Wang, Z. &amp; Whittaker, R. J. &amp; Fjeldsa, J. &amp; Rahbek, C." box="[498,676,1411,1437]" pageId="22" pageNumber="231" pagination="74 - 78" refId="ref23468" refString="Holt, B. G., Lessard, J. - P., Borregaard, M. K., Fritz, S. A., Araujo, M. B., Dimitrov, D., Fabre, P. - H., Graham, C. H., Graves, G. R., Jonsson, K. A., Nogues-Bravo, D., Wang, Z., Whittaker, R. J., Fjeldsa, J. &amp; Rahbek, C. (2013) An update of Wallace's zoogeographic regions of the world. Science, 339, 74 - 78. https: // doi. org / 10.1126 / science. 1228282" type="journal article" year="2013">
Holt
<emphasis id="29AEEAA9E023D744FDC4E9A3FDABFA41" box="[554,609,1411,1437]" italics="true" pageId="22" pageNumber="231">et al.</emphasis>
2013
</bibRefCitation>
). The larvae of just three species have been described and illustrated. These are
<taxonomicName id="DCDA4D38E023D744FEE5E987FD9BFA1D" authority="Selys" authorityName="Selys" authorityYear="1879" box="[267,593,1447,1473]" class="Insecta" family="Chlorocyphidae" genus="Rhinocypha" kingdom="Animalia" order="Odonata" pageId="22" pageNumber="231" phylum="Arthropoda" rank="species" species="ignipennis">
<emphasis id="29AEEAA9E023D744FEE5E987FDC7FA1C" box="[267,525,1447,1473]" italics="true" pageId="22" pageNumber="231">Rhinocypha ignipennis</emphasis>
Selys
</taxonomicName>
(
<bibRefCitation id="7F4B4B4AE023D744FD8EE987FD20FA1D" author="Fraser, F. C." box="[608,746,1447,1473]" pageId="22" pageNumber="231" pagination="13 - 14" refId="ref22861" refString="Fraser, F. C. (1920) Description of a Rhinocyphine larva from Shillong. Memoirs of the department of agriculture in India, 7, 13 - 14, 1 pl. excl." type="journal article" year="1920">Fraser 1920</bibRefCitation>
,
<bibRefCitation id="7F4B4B4AE023D744FD1BE987FCFBFA1D" author="Fraser, F. C." box="[757,817,1447,1473]" pageId="22" pageNumber="231" pagination="683 - 691" refId="ref22898" refString="Fraser, F. C. (1928) Indian dragonflies, Part XXX. Journal of the Bombay Natural History Society, 32, 683 - 691, pls. 1 - 3, excl." type="journal article" year="1928">1928</bibRefCitation>
), and
<taxonomicName id="DCDA4D38E023D744FC9BE987FB5CFA1D" authority="Oguma" authorityName="Oguma" box="[885,1174,1447,1473]" class="Insecta" family="Chlorocyphidae" genus="Rhinocypha" kingdom="Animalia" order="Odonata" pageId="22" pageNumber="231" phylum="Arthropoda" rank="species" species="ogasawarensis">
<emphasis id="29AEEAA9E023D744FC9BE987FBF3FA1C" box="[885,1081,1447,1472]" italics="true" pageId="22" pageNumber="231">R. ogasawarensis</emphasis>
Oguma
</taxonomicName>
,
<taxonomicName id="DCDA4D38E023D744FB4CE987FAA1FA1D" authority="Asahina." authorityName="Asahina." box="[1186,1387,1447,1473]" class="Insecta" family="Chlorocyphidae" genus="Rhinocypha" kingdom="Animalia" order="Odonata" pageId="22" pageNumber="231" phylum="Arthropoda" rank="species" species="uenoi">
<emphasis id="29AEEAA9E023D744FB4CE987FB35FA1C" box="[1186,1279,1447,1472]" italics="true" pageId="22" pageNumber="231">R. uenoi</emphasis>
Asahina.
</taxonomicName>
The latter two species are both figured in detail by
<bibRefCitation id="7F4B4B4AE023D744FD44E9EBFC89FA39" author="Ishida, K." box="[682,835,1483,1509]" pageId="22" pageNumber="231" refId="ref23597" refString="Ishida, K. (1996) Monograph of Odonata larvae in Japan. Hokkaido University Press, Sapporo, 446 pp. [in Japanese]" type="book" year="1996">Ishida (1996)</bibRefCitation>
, with habitus and details of prementum, labial palps, caudal gills and gonoapophyses being shown. These however were not fully mature specimens and habitus drawings based on exuviae with details of the mask and prementum were provided by
<bibRefCitation id="7F4B4B4AE023D744FC02EA33FB20F9F1" author="Sugimura, M. &amp; Ishida, S. &amp; Kojima, K. &amp; Ishida, K. &amp; Aoki, T." box="[1004,1258,1555,1581]" pageId="22" pageNumber="231" refId="ref26769" refString="Sugimura, M., Ishida, S., Kojima, K., Ishida, K. &amp; Aoki, T. (2001) Dragonflies of the Japanese Archipelago in Color. Hokkaido University Press, Sapporo, 641 pp. (English edition)" type="book" year="2001">
Sugimura
<emphasis id="29AEEAA9E023D744FBB1EA33FB5DF9F1" box="[1119,1175,1555,1581]" italics="true" pageId="22" pageNumber="231">et al.</emphasis>
(2001)
</bibRefCitation>
, together with a key which notes some differences in the antennae and caudal spikes of the two species.
<taxonomicName id="DCDA4D38E023D744FB84EA17FA56F98C" box="[1130,1436,1591,1617]" class="Insecta" family="Chlorocyphidae" genus="Rhinocypha" kingdom="Animalia" order="Odonata" pageId="22" pageNumber="231" phylum="Arthropoda" rank="species" species="ogasawarensis">
<emphasis id="29AEEAA9E023D744FB84EA17FA56F98C" box="[1130,1436,1591,1617]" italics="true" pageId="22" pageNumber="231">Rhinocypha ogasawarensis</emphasis>
</taxonomicName>
was also earlier briefly described and partially figured from an exuvia by
<bibRefCitation id="7F4B4B4AE023D744FC35EA7BFB47F9A9" author="Asahina, S." box="[987,1165,1626,1653]" pageId="22" pageNumber="231" pagination="52 - 55" refId="ref21577" refString="Asahina, S. (1956) An annotated list of Odonata of Japan with brief descriptions of their larva. Shin-konchu, 9 (12), 52 - 55." type="journal article" year="1956">Asahina (1956)</bibRefCitation>
. Although not shown in detail by
<bibRefCitation id="7F4B4B4AE023D744FF10EA5FFE5CF945" author="Fraser, F. C." box="[254,406,1663,1689]" pageId="22" pageNumber="231" pagination="683 - 691" refId="ref22898" refString="Fraser, F. C. (1928) Indian dragonflies, Part XXX. Journal of the Bombay Natural History Society, 32, 683 - 691, pls. 1 - 3, excl." type="journal article" year="1928">Fraser (1928)</bibRefCitation>
, it is clear that
<taxonomicName id="DCDA4D38E023D744FDD3EA5FFD1AF944" authorityName="Selys" authorityYear="1879" box="[573,720,1663,1688]" class="Insecta" family="Chlorocyphidae" genus="Rhinocypha" kingdom="Animalia" order="Odonata" pageId="22" pageNumber="231" phylum="Arthropoda" rank="species" species="ignipennis">
<emphasis id="29AEEAA9E023D744FDD3EA5FFD1AF944" box="[573,720,1663,1688]" italics="true" pageId="22" pageNumber="231">R. ignipennis</emphasis>
</taxonomicName>
conforms to a typical chlorocyphid pattern with long thin spikes and a short sagittate head. The Japanese species have unusually short caudal spikes, as noted above, and the head is more elongate, at least in figures provided by
<bibRefCitation id="7F4B4B4AE023D744FD74EAE7FCF9F93D" author="Ishida, K." box="[666,819,1735,1761]" pageId="22" pageNumber="231" refId="ref23597" refString="Ishida, K. (1996) Monograph of Odonata larvae in Japan. Hokkaido University Press, Sapporo, 446 pp. [in Japanese]" type="book" year="1996">Ishida (1996)</bibRefCitation>
. Despite being closely related (
<bibRefCitation id="7F4B4B4AE023D744FB7FEAE7FA9EF93D" author="Wang, L. - J. &amp; Cherng, J. - J. &amp; Chang, Y. - J. &amp; Jiang, J. - L." box="[1169,1364,1735,1761]" pageId="22" pageNumber="231" pagination="93 - 107" refId="ref27350" refString="Wang, L. - J., Cherng, J. - J., Chang, Y. - J. &amp; Jiang, J. - L. (2013) Description of Rhinocypha taiwana sp. nov. from Taiwan, with a preliminary molecular phylogenetic analysis of the Rhinocypha drusilla - group (Odonata: Chlorocyphidae). International Journal of Odonatology, 16, 93 - 107. https: // doi. org / 10.1080 / 13887890.2012.763152" type="journal article" year="2013">
Wang
<emphasis id="29AEEAA9E023D744FB35EAE7FAC6F93D" box="[1243,1292,1735,1761]" italics="true" pageId="22" pageNumber="231">et al</emphasis>
. 2013
</bibRefCitation>
) these two species differ markedly in the shape of the prementum, and in minor details of the palpal lobe armature (
<bibRefCitation id="7F4B4B4AE023D744FAB8EACBFF1AF8F5" author="Ishida, K." pageId="22" pageNumber="231" refId="ref23597" refString="Ishida, K. (1996) Monograph of Odonata larvae in Japan. Hokkaido University Press, Sapporo, 446 pp. [in Japanese]" type="book" year="1996">Ishida 1996</bibRefCitation>
) and antennae (
<bibRefCitation id="7F4B4B4AE023D744FE6CEB2FFDB9F8F5" author="Sugimura, M. &amp; Ishida, S. &amp; Kojima, K. &amp; Ishida, K. &amp; Aoki, T." box="[386,627,1807,1833]" pageId="22" pageNumber="231" refId="ref26769" refString="Sugimura, M., Ishida, S., Kojima, K., Ishida, K. &amp; Aoki, T. (2001) Dragonflies of the Japanese Archipelago in Color. Hokkaido University Press, Sapporo, 641 pp. (English edition)" type="book" year="2001">
Sugimura
<emphasis id="29AEEAA9E023D744FE16EB2FFDFAF8F5" box="[504,560,1807,1833]" italics="true" pageId="22" pageNumber="231">et al.</emphasis>
2001
</bibRefCitation>
), dissimilarities which perhaps reflect ecological differences. The genus is very diverse, and as noted by
<bibRefCitation id="7F4B4B4AE023D744FDE8EB13FD3EF891" author="Dijkstra, K-D. B. &amp; Kalkman, V. J. &amp; Dow, R. A. &amp; Stokvis, F. R. &amp; van Tol, J." box="[518,756,1843,1869]" pageId="22" pageNumber="231" pagination="68 - 96" refId="ref22480" refString="Dijkstra, K-D. B., Kalkman, V. J., Dow, R. A., Stokvis, F. R. &amp; van Tol, J. (2014) Redefining the damselfly families: a comprehensive molecular phylogeny of Zygoptera (Odonata). Systematic entomology, 39, 68 - 96. https: // doi. org / 10.1111 / syen. 12035" type="journal article" year="2014">
Dijkstra
<emphasis id="29AEEAA9E023D744FD86EB13FD51F891" box="[616,667,1843,1869]" italics="true" pageId="22" pageNumber="231">et al</emphasis>
. (2014)
</bibRefCitation>
‘non monophyletic’, hence a good deal of variation may be expected among the larvae.
</paragraph>
</subSubSection>
</treatment>
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135
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<emphasis id="29AEEAA9E023D744FF79ED06FECEFE9D" bold="true" box="[151,260,294,321]" italics="true" pageId="22" pageNumber="231">Libellago</emphasis>
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The genus includes 25 species, most, but not all, are small for the family, and range throughout most of the Oriental realm. Larger species occur especially on
<collectingRegion id="D91EF859E023D744FD97ED4FFD28FE55" box="[633,738,367,393]" country="Indonesia" name="Sulawesi Utara" pageId="22" pageNumber="231">Sulawesi</collectingRegion>
. The larva of
<taxonomicName id="DCDA4D38E023D744FC6AED4FFB8DFE55" box="[900,1095,367,393]" class="Insecta" family="Chlorocyphidae" genus="Libellago" kingdom="Animalia" order="Odonata" pageId="22" pageNumber="231" phylum="Arthropoda" rank="species" species="lineata">
<emphasis id="29AEEAA9E023D744FC6AED4FFB8DFE55" box="[900,1095,367,393]" italics="true" pageId="22" pageNumber="231">Libellago lineata</emphasis>
</taxonomicName>
was described
<bibRefCitation id="7F4B4B4AE023D744FB16ED4FFA52FE55" author="Fraser, F. C." box="[1272,1432,367,393]" pageId="22" pageNumber="231" pagination="197 - 198" refId="ref22755" refString="Fraser, F. C. (1919 a) The larva of Micromerus lineatus Burm. Records of the Indian Museum, 16, 197 - 198, pl. 23 excl. https: // doi. org / 10.26515 / rzsi / v 16 / i 2 / 1919 / 162807" type="journal article" year="1919">Fraser (1919a</bibRefCitation>
,
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) with the habitus illustrated, with some omissions, and a rough figure of part of the mask, which does not conform to the detailed description; however the entire prementum and in-life habitus are figured by Orr
<emphasis id="29AEEAA9E023D744FA8FED97FA57FE0D" box="[1377,1437,439,465]" italics="true" pageId="22" pageNumber="231">et al.</emphasis>
(2024).
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,
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) figures the habitus of
<taxonomicName id="DCDA4D38E023D744FD5DEDFBFCB7FE29" authority="(Selys)" baseAuthorityName="Selys" box="[691,893,475,501]" class="Insecta" family="Chlorocyphidae" genus="Libellago" kingdom="Animalia" order="Odonata" pageId="22" pageNumber="231" phylum="Arthropoda" rank="species" species="hyalina">
<emphasis id="29AEEAA9E023D744FD5DEDFBFCECFE29" box="[691,806,475,501]" italics="true" pageId="22" pageNumber="231">L. hyalina</emphasis>
(Selys)
</taxonomicName>
and
<bibRefCitation id="7F4B4B4AE023D744FC58EDFBFB57FE29" author="Ngiam, R. W. J. &amp; Ng, M. F. C." box="[950,1181,475,501]" pageId="22" pageNumber="231" refId="ref25398" refString="Ngiam, R. W. J. &amp; Ng, M. F. C. (2022) A photographic field guide to the dragonflies &amp; damselflies of Singapore. John Beaufoy Publishing, Oxford, 340 pp." type="book" year="2022">Ngiam &amp; Ng (2022)</bibRefCitation>
show a photograph of the dorsal habitus of
<taxonomicName id="DCDA4D38E023D744FE6AEDDFFDBCFDC5" authority="(Selys)" baseAuthorityName="Selys" box="[388,630,511,537]" class="Insecta" family="Chlorocyphidae" genus="Libellago" kingdom="Animalia" order="Odonata" pageId="22" pageNumber="231" phylum="Arthropoda" rank="species" species="aurantiaca">
<emphasis id="29AEEAA9E023D744FE6AEDDFFDD7FDC4" box="[388,541,511,536]" italics="true" pageId="22" pageNumber="231">L. aurantiaca</emphasis>
(Selys)
</taxonomicName>
. Orr
<emphasis id="29AEEAA9E023D744FD5EEDDFFD28FDC5" box="[688,738,511,537]" italics="true" pageId="22" pageNumber="231">et al</emphasis>
. (2024) provide a detailed description of
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<emphasis id="29AEEAA9E023D744FB5BEDDFFAEDFDC5" box="[1205,1319,511,537]" italics="true" pageId="22" pageNumber="231">L. hyalina</emphasis>
</taxonomicName>
as well as photographs of living
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<emphasis id="29AEEAA9E023D744FE62EE03FE3CFDE1" box="[396,502,547,573]" italics="true" pageId="22" pageNumber="231">L. lineata</emphasis>
</taxonomicName>
. All species are small and have characteristic thin tapered caudal spikes with a slight basal swelling. The spikes are long in
<taxonomicName id="DCDA4D38E023D744FDA7EE67FD76FDBD" box="[585,700,583,609]" class="Insecta" family="Chlorocyphidae" genus="Libellago" kingdom="Animalia" order="Odonata" pageId="22" pageNumber="231" phylum="Arthropoda" rank="species" species="hyalina">
<emphasis id="29AEEAA9E023D744FDA7EE67FD76FDBD" box="[585,700,583,609]" italics="true" pageId="22" pageNumber="231">L. hyalina</emphasis>
</taxonomicName>
and
<taxonomicName id="DCDA4D38E023D744FD1AEE67FCABFDBD" box="[756,865,583,609]" class="Insecta" family="Chlorocyphidae" genus="Libellago" kingdom="Animalia" order="Odonata" pageId="22" pageNumber="231" phylum="Arthropoda" rank="species" species="lineata">
<emphasis id="29AEEAA9E023D744FD1AEE67FCABFDBD" box="[756,865,583,609]" italics="true" pageId="22" pageNumber="231">L. lineata</emphasis>
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but shorter in
<taxonomicName id="DCDA4D38E023D744FBE9EE67FB6BFDBC" box="[1031,1185,583,608]" class="Insecta" family="Chlorocyphidae" genus="Libellago" kingdom="Animalia" order="Odonata" pageId="22" pageNumber="231" phylum="Arthropoda" rank="species" species="aurantiaca">
<emphasis id="29AEEAA9E023D744FBE9EE67FB6BFDBC" box="[1031,1185,583,608]" italics="true" pageId="22" pageNumber="231">L. aurantiaca</emphasis>
</taxonomicName>
. The antennae tend to be relatively shorter than in other genera studied. The prementum in
<taxonomicName id="DCDA4D38E023D744FC44EE4BFBD7FD59" box="[938,1053,619,645]" class="Insecta" family="Chlorocyphidae" genus="Libellago" kingdom="Animalia" order="Odonata" pageId="22" pageNumber="231" phylum="Arthropoda" rank="species" species="hyalina">
<emphasis id="29AEEAA9E023D744FC44EE4BFBD7FD59" box="[938,1053,619,645]" italics="true" pageId="22" pageNumber="231">L. hyalina</emphasis>
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and
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<emphasis id="29AEEAA9E023D744FBB9EE4BFB0EFD59" box="[1111,1220,619,645]" italics="true" pageId="22" pageNumber="231">L. lineata</emphasis>
</taxonomicName>
differs sufficiently to separate the two species, but in both is shorter and broader than in other genera and has distinctly serrate lateral anterior margins not recorded in other genera except possibly the much larger
<taxonomicName id="DCDA4D38E023D744FBF2EE93FB5DFD11" authorityName="Fraser" authorityYear="1949" box="[1052,1175,691,717]" class="Insecta" family="Chlorocyphidae" genus="Paracypha" kingdom="Animalia" order="Odonata" pageId="22" pageNumber="231" phylum="Arthropoda" rank="genus">
<emphasis id="29AEEAA9E023D744FBF2EE93FB5DFD11" box="[1052,1175,691,717]" italics="true" pageId="22" pageNumber="231">Paracypha</emphasis>
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. All three species lack a dark spot on the tip of the pedicel, present in all other known chlorocyphids. In mainland Asia and much of Sundaland their small size also makes these larvae immediately recognisable as belonging to
<taxonomicName id="DCDA4D38E023D744FB35EEDBFA8CFCC9" box="[1243,1350,763,789]" class="Insecta" family="Chlorocyphidae" genus="Libellago" kingdom="Animalia" order="Odonata" pageId="22" pageNumber="231" phylum="Arthropoda" rank="genus">
<emphasis id="29AEEAA9E023D744FB35EEDBFA8CFCC9" box="[1243,1350,763,789]" italics="true" pageId="22" pageNumber="231">Libellago</emphasis>
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, but in
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much larger members of the genus occur (
<bibRefCitation id="7F4B4B4AE023D744FD37EF3FFCA5FCE5" author="van Tol, J." box="[729,879,799,825]" pageId="22" pageNumber="231" pagination="209 - 248" refId="ref27249" refString="van Tol, J. (2007) The Odonata of Sulawesi and adjacent islands. Part 7. Libellago and Sclerocypha (Chlorocyphidae). International Journal of Odonatology, 10, 209 - 248. https: // doi. org / 10.1080 / 13887890.2007.9748301" type="journal article" year="2007">van Tol 2007</bibRefCitation>
,
<bibRefCitation id="7F4B4B4AE023D744FC94EF3FFBF9FCE5" author="Dow, R. A. &amp; Choong, C. Y. &amp; Grinang, J. &amp; Lupiyaningdyah, P. &amp; Ngiam, R. W. J. &amp; Kalkman, V. J." box="[890,1075,799,825]" pageId="22" pageNumber="231" pagination="1 - 122" refId="ref22552" refString="Dow, R. A., Choong, C. Y., Grinang, J., Lupiyaningdyah, P., Ngiam, R. W. J. &amp; Kalkman, V. J. (2024) Checklist of the Odonata (Insecta) of Sundaland and Wallacea (Malaysia, Singapore, Brunei, Indonesia and Timor Leste). Zootaxa, 5460 (1), 1 - 122. https: // doi. org / 10.11646 / zootaxa. 5460.1.1" type="journal article" year="2024">
Dow
<emphasis id="29AEEAA9E023D744FC59EF3FFC22FCE5" box="[951,1000,799,825]" italics="true" pageId="22" pageNumber="231">et al</emphasis>
. 2024
</bibRefCitation>
). It is not known if these species share the presently recognised generic traits.
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This monotypic genus (considered by some to be a synonym of
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which is restricted to Meghahalya (
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) and the Himalayas, ranging to
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in the west. The larva was described but not figured by
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;
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figures it in detail, and it is mentioned in the key of
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. Its size separates it from most, but not all, sympatric chlorocyphid species, and the extensive spines along the lateral margin of the prementum may be a good generic character.
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illustrates uneven caudal spikes, suggesting one has regenerated, without detail, but a specimen in the collection of S.G. Butler suggests that the head shape and the spikes may also provide distinctive characters (S.G. Butler pers comm.).
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<paragraph id="1B6536BBE024D743FF79EB2EFEE6F8F5" blockId="17.[151,1437,1806,2013]" box="[151,300,1806,1833]" pageId="17" pageNumber="226">
<heading id="402D81D7E024D743FF79EB2EFEE6F8F5" bold="true" box="[151,300,1806,1833]" fontSize="11" level="1" pageId="17" pageNumber="226" reason="1">
<taxonomicName id="DCDA4D38E024D743FF79EB2EFEE6F8F5" ID-CoL="74D8" authorityName="McLachlan" authorityYear="1870" box="[151,300,1806,1833]" class="Insecta" family="Calopterygidae" genus="Psolodesmus" kingdom="Animalia" order="Odonata" pageId="17" pageNumber="226" phylum="Arthropoda" rank="genus">
<emphasis id="29AEEAA9E024D743FF79EB2EFEE6F8F5" bold="true" box="[151,300,1806,1833]" italics="true" pageId="17" pageNumber="226">Psolodesmus</emphasis>
</taxonomicName>
</heading>
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<subSubSection id="53C06530E024D740FF79EB13FB8DFE71" lastPageId="18" lastPageNumber="227" pageId="17" pageNumber="226" type="discussion">
<paragraph id="1B6536BBE024D740FF79EB13FB8DFE71" blockId="17.[151,1437,1806,2013]" lastBlockId="18.[151,1437,151,429]" lastPageId="18" lastPageNumber="227" pageId="17" pageNumber="226">
Just two closely related species belong to this genus: one,
<taxonomicName id="DCDA4D38E024D743FCAFEB13FB24F891" authority="McLachlan" authorityName="McLachlan" authorityYear="1870" box="[833,1262,1843,1869]" class="Insecta" family="Calopterygidae" genus="Psolodesmus" kingdom="Animalia" order="Odonata" pageId="17" pageNumber="226" phylum="Arthropoda" rank="species" species="mandarinus">
<emphasis id="29AEEAA9E024D743FCAFEB13FBAEF891" box="[833,1124,1843,1869]" italics="true" pageId="17" pageNumber="226">Psolodesmus mandarinus</emphasis>
McLachlan
</taxonomicName>
is restricted to
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and the other,
<taxonomicName id="DCDA4D38E024D743FE7BEB77FDB9F8AD" authority="Oguma" authorityName="Oguma" authorityYear="1913" box="[405,627,1879,1905]" class="Insecta" family="Calopterygidae" genus="Psolodesmus" kingdom="Animalia" order="Odonata" pageId="17" pageNumber="226" phylum="Arthropoda" rank="species" species="kuroiwae">
<emphasis id="29AEEAA9E024D743FE7BEB77FDD3F8AD" box="[405,537,1879,1905]" italics="true" pageId="17" pageNumber="226">P. kuroiwae</emphasis>
Oguma
</taxonomicName>
to the southern Japanese Yaeyama Islands. The larva of
<taxonomicName id="DCDA4D38E024D743FB14EB77FE4EF849" authority="Williamson" authorityName="Williamson" authorityYear="1904" class="Insecta" family="Calopterygidae" genus="Psolodesmus" kingdom="Animalia" order="Odonata" pageId="17" pageNumber="226" phylum="Arthropoda" rank="subSpecies" species="mandarinus" subSpecies="dorothea">
<emphasis id="29AEEAA9E024D743FB14EB77FF31F849" italics="true" pageId="17" pageNumber="226">P. mandarinus dorothea</emphasis>
Williamson
</taxonomicName>
was described in detail by
<bibRefCitation id="7F4B4B4AE024D743FD56EB5BFC0AF849" author="Matsuki, K. &amp; Lien, J. C." box="[696,960,1915,1941]" pageId="17" pageNumber="226" pagination="15 - 26" refId="ref24810" refString="Matsuki, K. &amp; Lien, J. C. (1978) Descriptions of the larvae of three families of Zygoptera breeding in the streams of Taiwan (Synlestidae, Euphaeidae &amp; Calopterygidae). Tombo - Acta Odonatologica Japonica, 21, 15 - 26." type="journal article" year="1978">Matsuki &amp; Lien (1978)</bibRefCitation>
, who also figure details of the prementum of
<taxonomicName id="DCDA4D38E024D743FF54EBBFFE38F865" box="[186,498,1951,1977]" class="Insecta" family="Calopterygidae" genus="Psolodesmus" kingdom="Animalia" order="Odonata" pageId="17" pageNumber="226" phylum="Arthropoda" rank="subSpecies" species="mandarinus" subSpecies="mandarinus">
<emphasis id="29AEEAA9E024D743FF54EBBFFE38F865" box="[186,498,1951,1977]" italics="true" pageId="17" pageNumber="226">P. mandarinus mandarinus</emphasis>
</taxonomicName>
, claiming that small differences in the premental cleft distinguish larvae of the two subspecies. Otherwise there is little difference between them, but those of
<emphasis id="29AEEAA9E024D743FBDEEBE3FB30F801" box="[1072,1274,1987,2013]" italics="true" pageId="17" pageNumber="226">
P. m.
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</emphasis>
are described as inhabiting low hills “living in fountain head and upper section of slow streams, but occasionally breeding in artificially made ditches”. The larva of
<taxonomicName id="DCDA4D38E027D740FDB0EC9BFD29FF09" authorityName="Oguma" authorityYear="1913" box="[606,739,187,213]" class="Insecta" family="Calopterygidae" genus="Psolodesmus" kingdom="Animalia" order="Odonata" pageId="18" pageNumber="227" phylum="Arthropoda" rank="species" species="kuroiwae">
<emphasis id="29AEEAA9E027D740FDB0EC9BFD29FF09" box="[606,739,187,213]" italics="true" pageId="18" pageNumber="227">P. kuroiwae</emphasis>
</taxonomicName>
was described and illustrated by
<bibRefCitation id="7F4B4B4AE027D740FB88EC9BFAE0FF09" author="Watanabe, K." box="[1126,1322,187,213]" pageId="18" pageNumber="227" pagination="39 - 41" refId="ref27502" refString="Watanabe, K. (1984) Description of the larval stage of Psolodesmus mandarinus kuroiwae and Gomphus yayeyamensis. Tombo - Acta Odonatologica Japonica, 27, 39 - 41. [in Japanese]" type="journal article" year="1984">Watanabe (1984)</bibRefCitation>
,
<bibRefCitation id="7F4B4B4AE027D740FAD7EC9BFEE7FF25" author="Ishida, S. &amp; Ishida, K." pageId="18" pageNumber="227" pagination="33 - 124" refId="ref23624" refString="Ishida, S. &amp; Ishida, K. (1985) Odonata. In: Kawai, T. (Ed.), An illustrated book of aquatic insects of Japan. Tokai University Press, Tokyo, pp. 33 - 124. [in Japanese]" type="book chapter" year="1985">Ishida &amp; Ishida (1985)</bibRefCitation>
and
<bibRefCitation id="7F4B4B4AE027D740FE8CECFFFE32FF25" author="Ishida, K." box="[354,504,223,249]" pageId="18" pageNumber="227" refId="ref23597" refString="Ishida, K. (1996) Monograph of Odonata larvae in Japan. Hokkaido University Press, Sapporo, 446 pp. [in Japanese]" type="book" year="1996">Ishida (1996)</bibRefCitation>
and differs very little from
<taxonomicName id="DCDA4D38E027D740FCC2ECFFFC07FF25" authorityName="McLachlan" authorityYear="1870" box="[812,973,223,249]" class="Insecta" family="Calopterygidae" genus="Psolodesmus" kingdom="Animalia" order="Odonata" pageId="18" pageNumber="227" phylum="Arthropoda" rank="species" species="mandarinus">
<emphasis id="29AEEAA9E027D740FCC2ECFFFC07FF25" box="[812,973,223,249]" italics="true" pageId="18" pageNumber="227">P. mandarinus</emphasis>
</taxonomicName>
. In all cases the prementum is very broad anteriorly with the anterior processes flanking the central narrow tear-drop cleft touching apically or overlapping, and bearing a single pair of strong setae near the base (
<figureCitation id="83E12A3EE027D740FD16ED07FC86FE9D" box="[760,844,295,321]" captionStart="FIGURES 27–34. 27" captionStartId="7.[151,264,1845,1869]" captionTargetBox="[252,1326,159,1834]" captionTargetId="figure-15@7.[251,1336,153,1839]" captionTargetPageId="7" captionText="FIGURES 27–34. 27. Prementum of Calopterygidae showing measurements used in family key (based on Calopteryx japonica, redrawn from Miyakawa 1983). 28. Neurobasis chinensis, prementum and left labial palp (redrawn from Lieftinck 1965). 29. Matronoides cyaneipennis, prementum and labial palps (ventral view). 30. Matrona basilaris, prementum (dorsal view, drawn from Wang et al. 2017). 31. Atrocalopteryx atrata, prementum (dorsal view, redrawn from Miyakawa 1983). 32. Mnais tenuis, prementum (dorsal view, redrawn from Matsuki &amp; Lien 1977). 33. Echo uniformis, prementum and left labial palp dorsal view (redrawn from Lieftinck 1965). 34. Psolodesmus mandarinus, prementum (dorsal view, redrawn from Matsuki &amp; Lien 1978)." figureDoi="http://doi.org/10.5281/zenodo.13618333" httpUri="https://zenodo.org/record/13618333/files/figure.png" pageId="18" pageNumber="227">Fig. 34</figureCitation>
). The overall build of the larva is heavy and the short lateral caudal lamellae have a strong row of teeth along the central outer ridge (
<figureCitation id="83E12A3EE027D740FBE2ED6BFB95FEB9" box="[1036,1119,331,357]" captionStart="FIGURES 42–51. 42" captionStartId="9.[151,264,1592,1616]" captionTargetBox="[165,1413,192,1557]" captionTargetId="figure-15@9.[151,1435,181,1568]" captionTargetPageId="9" captionText="FIGURES 42–51. 42. Archineura incarnata: caudal gills; left lateral above, median below (drawn from Yang et al. 2022). 43. Echo modesta: caudal gills; left lateral above, median below, with baso-dorsal shield arrowed (original illustration). 44. Psolodesmus mandarinus, dorsal view of caudal gills, with baso-dorsal shield of median gill arrowed (redrawn from Matsuki &amp; Lien 1978). 45. Mnais andersoni, left lateral caudal gill with upper margin of median gill visible; baso-dorsal shield of median gill arrowed (original illustration). 46. Mnais gregoryi: caudal gills; left lateral above, median below, with baso-dorsal shield arrowed (drawn from Yang et al. 2021). 47. Caliphaea angka: caudal gills; left lateral above, median below (drawn from Yang et al. 2021). 48. Matrona basilaris: caudal gills; left lateral above, median below (drawn from Wang et al. 2017). 49. Vestalaria venusta: caudal gills; left lateral above, median below (drawn from Wang et al. 2017). 50. Vestalis amoena group: caudal gills; left lateral above, median below (redrawn from Lieftinck 1965). 51. Vestalis melania: caudal gills; left lateral above, median below (redrawn from Guadalquiver et al. 2022)." figureDoi="http://doi.org/10.5281/zenodo.13618337" httpUri="https://zenodo.org/record/13618337/files/figure.png" pageId="18" pageNumber="227">Fig. 44</figureCitation>
), with the dorsal and ventral margins smooth (cf
<taxonomicName id="DCDA4D38E027D740FE6EED4FFE70FE55" box="[384,442,367,393]" class="Insecta" family="Calopterygidae" genus="Echo" kingdom="Animalia" order="Odonata" pageId="18" pageNumber="227" phylum="Arthropoda" rank="genus">
<emphasis id="29AEEAA9E027D740FE6EED4FFE70FE55" box="[384,442,367,393]" italics="true" pageId="18" pageNumber="227">Echo</emphasis>
</taxonomicName>
). The central lamella is significantly shorter than the lateral ones and has a long thin, narrowly triangular dorsal guard (also slightly different from
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<emphasis id="29AEEAA9E027D740FCA8EDB3FC4AFE71" box="[838,896,403,429]" italics="true" pageId="18" pageNumber="227">Echo</emphasis>
</taxonomicName>
in both respects).
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The genus ranges from the island of New
<collectingCountry id="63CD762BE024D743FD64E81FFD11FB85" box="[650,731,1087,1113]" name="Guinea-Bissau" pageId="17" pageNumber="226">Guinea</collectingCountry>
to subtropical
<collectingCountry id="63CD762BE024D743FC62E81FFC18FB85" box="[908,978,1087,1113]" name="China" pageId="17" pageNumber="226">China</collectingCountry>
, Indomalaya and the Himalayan region as far as north
<collectingCountry id="63CD762BE024D743FEA9E843FE2DFBA1" box="[327,487,1123,1149]" name="Pakistan" pageId="17" pageNumber="226">west Pakistan</collectingCountry>
. It includes nine species in the Oriental region, the remaining four being found in New
<collectingCountry id="63CD762BE024D743FF38E8A7FEE1FB7D" box="[214,299,1159,1185]" name="Guinea-Bissau" pageId="17" pageNumber="226">Guinea</collectingCountry>
. The habitus shown for
<taxonomicName id="DCDA4D38E024D743FDA6E8A7FC92FB7D" authority="(Linnaeus)" baseAuthorityName="Linnaeus" box="[584,856,1159,1185]" class="Insecta" family="Calopterygidae" genus="Neurobasis" kingdom="Animalia" order="Odonata" pageId="17" pageNumber="226" phylum="Arthropoda" rank="species" species="chinensis">
<emphasis id="29AEEAA9E024D743FDA6E8A7FD1EFB7D" box="[584,724,1159,1185]" italics="true" pageId="17" pageNumber="226">N. chinensis</emphasis>
(Linnaeus)
</taxonomicName>
(
<figureCitation id="83E12A3EE024D743FC85E8A7FC75FB7D" box="[875,959,1158,1185]" captionStart="FIGURES 52–56. 52" captionStartId="10.[151,264,1941,1965]" captionTargetBox="[191,1388,163,1929]" captionTargetId="figure-21@10.[190,1397,151,1933]" captionTargetPageId="10" captionText="FIGURES 52–56. 52. Caliphaea angka, habitus (based on Yang et al. 2021). 53. Vestalis gracilis, habitus (redrawn from Rattanachan et al. 2022). 54. Neurobasis chinensis, habitus (original drawing). 55. Matrona basilaris, habitus (drawn from Wang et al. 2017). 56. Echo uniformis, habitus (redrawn from Lieftinck 1965)." figureDoi="http://doi.org/10.5281/zenodo.13618339" httpUri="https://zenodo.org/record/13618339/files/figure.png" pageId="17" pageNumber="226">Fig. 54</figureCitation>
) and deep cleft prementum (
<figureCitation id="83E12A3EE024D743FAFDE8A7FAADFB7D" box="[1299,1383,1159,1185]" captionStart="FIGURES 27–34. 27" captionStartId="7.[151,264,1845,1869]" captionTargetBox="[252,1326,159,1834]" captionTargetId="figure-15@7.[251,1336,153,1839]" captionTargetPageId="7" captionText="FIGURES 27–34. 27. Prementum of Calopterygidae showing measurements used in family key (based on Calopteryx japonica, redrawn from Miyakawa 1983). 28. Neurobasis chinensis, prementum and left labial palp (redrawn from Lieftinck 1965). 29. Matronoides cyaneipennis, prementum and labial palps (ventral view). 30. Matrona basilaris, prementum (dorsal view, drawn from Wang et al. 2017). 31. Atrocalopteryx atrata, prementum (dorsal view, redrawn from Miyakawa 1983). 32. Mnais tenuis, prementum (dorsal view, redrawn from Matsuki &amp; Lien 1977). 33. Echo uniformis, prementum and left labial palp dorsal view (redrawn from Lieftinck 1965). 34. Psolodesmus mandarinus, prementum (dorsal view, redrawn from Matsuki &amp; Lien 1978)." figureDoi="http://doi.org/10.5281/zenodo.13618333" httpUri="https://zenodo.org/record/13618333/files/figure.png" pageId="17" pageNumber="226">Fig. 28</figureCitation>
) are both typical for the genus. Larvae, tentatively mentioned by
<bibRefCitation id="7F4B4B4AE024D743FCB8E88BFC3CFB19" author="Hagen, H." box="[854,1014,1195,1221]" pageId="17" pageNumber="226" pagination="lxv - lxvii" refId="ref23289" refString="Hagen, H. (1880) Essai d'un synopsis des larves de Calopterygines. [Comptes-rendus de la Socie´te´Entomologique de Belgique. Seance du 1 er Mai 1880]. Annales de la Societe entomologique de Belgique, 23, lxv - lxvii." type="journal article" year="1880">Hagen (1880)</bibRefCitation>
, were first described in detail for
<taxonomicName id="DCDA4D38E024D743FA6DE88BFE0BFB35" authority="(Needham 1911)" baseAuthorityName="Needham" baseAuthorityYear="1911" class="Insecta" family="Calopterygidae" genus="Neurobasis" kingdom="Animalia" order="Odonata" pageId="17" pageNumber="226" phylum="Arthropoda" rank="species" species="chinensis">
<emphasis id="29AEEAA9E024D743FA6DE88BFF35FB35" italics="true" pageId="17" pageNumber="226">N. chinensis</emphasis>
(
<bibRefCitation id="7F4B4B4AE024D743FEE2E8EFFE73FB35" author="Needham, J. G." box="[268,441,1231,1257]" pageId="17" pageNumber="226" pagination="145 - 154" refId="ref24959" refString="Needham, J. G. (1911) Descriptions of dragonfly nymphs of the subfamily Calopteryginae (Odonata). Entomological News, 22, 145 - 154." type="journal article" year="1911">Needham 1911</bibRefCitation>
)
</taxonomicName>
, and likewise by
<bibRefCitation id="7F4B4B4AE024D743FD90E8EFFCD6FB35" author="Kumar, A." box="[638,796,1231,1257]" pageId="17" pageNumber="226" pagination="83 - 118" refId="ref24196" refString="Kumar, A. (1973) Descriptions of the last instar larvae of Odonata from the Dehra Dun Valley (India), with notes on biology I. (suborder Zygoptera). Oriental Insects, 7, 83 - 118. https: // doi. org / 10.1080 / 00305316.1973.10434207" type="journal article" year="1973">Kumar (1973)</bibRefCitation>
who also illustrated a lateral habitus and briefly discussed behaviour and ecology, which is given in more detail by
<bibRefCitation id="7F4B4B4AE024D743FCE5E8D3FBECFAD1" author="Kumar, A. &amp; Prasad, M." box="[779,1062,1267,1293]" pageId="17" pageNumber="226" pagination="225 - 230" refId="ref24294" refString="Kumar, A. &amp; Prasad, M. (1977 a) Last instar larvae of two Odonata species from Western Himalayas. Entomon, 2, 225 - 230." type="journal article" year="1977">Kumar &amp; Prasad (1977a)</bibRefCitation>
.
<bibRefCitation id="7F4B4B4AE024D743FBDEE8D3FB0AFAD1" author="Fraser, F. C." box="[1072,1216,1267,1293]" pageId="17" pageNumber="226" refId="ref23008" refString="Fraser, F. C. (1933) The fauna of British India, including Ceylon and Burma. Odonata. Vol. I. Taylor &amp; Francis Ltd., London, xiii + 423 pp., 1 folded map excl." type="book" year="1933">Fraser (1933</bibRefCitation>
, p. 8) illustrated the habitus (rather stouter than in reality).
<bibRefCitation id="7F4B4B4AE024D743FDA0E937FCC2FAED" author="Lieftinck, M. A." box="[590,776,1302,1329]" pageId="17" pageNumber="226" pagination="155 - 166" refId="ref24556" refString="Lieftinck, M. A. (1955) Notes on Australasian species of Neurobasis Selys (Odonata, Agriidae). Nova Guinea, New Series, 6, 155 - 166, pl. 3 excl." type="journal article" year="1955">Lieftinck (1955)</bibRefCitation>
described and illustrated the habitus and structural details of
<taxonomicName id="DCDA4D38E024D743FF56E91BFEB3FA89" authority="Hagen" authorityName="Hagen" box="[184,377,1339,1365]" class="Insecta" family="Calopterygidae" genus="Neurobasis" kingdom="Animalia" order="Odonata" pageId="17" pageNumber="226" phylum="Arthropoda" rank="species" species="florida">
<emphasis id="29AEEAA9E024D743FF56E91BFEECFA89" box="[184,294,1339,1365]" italics="true" pageId="17" pageNumber="226">N. florida</emphasis>
Hagen
</taxonomicName>
as
<taxonomicName id="DCDA4D38E024D743FE4DE91BFDE7FA89" baseAuthorityName="Needham" baseAuthorityYear="1911" box="[419,557,1339,1365]" class="Insecta" family="Calopterygidae" genus="Neurobasis" kingdom="Animalia" order="Odonata" pageId="17" pageNumber="226" phylum="Arthropoda" rank="species" species="chinensis">
<emphasis id="29AEEAA9E024D743FE4DE91BFDE7FA89" box="[419,557,1339,1365]" italics="true" pageId="17" pageNumber="226">N. chinensis</emphasis>
</taxonomicName>
, and the habitus only was shown for
<taxonomicName id="DCDA4D38E024D743FC0EE91BFA53FA89" authority="(Orr &amp; Hamalainen 2007)" baseAuthorityName="Orr &amp; Hamalainen" baseAuthorityYear="2007" box="[992,1433,1339,1365]" class="Insecta" family="Calopterygidae" genus="Neurobasis" kingdom="Animalia" order="Odonata" pageId="17" pageNumber="226" phylum="Arthropoda" rank="species" species="longipes">
<emphasis id="29AEEAA9E024D743FC0EE91BFBA8FA89" box="[992,1122,1339,1365]" italics="true" pageId="17" pageNumber="226">N. longipes</emphasis>
(
<bibRefCitation id="7F4B4B4AE024D743FB9DE91BFA5BFA89" author="Orr, A. G. &amp; Hamalainen, M." box="[1139,1425,1339,1365]" pageId="17" pageNumber="226" refId="ref26055" refString="Orr, A. G. &amp; Hamalainen, M. (2007) The metalwing demoiselles of the eastern tropics - their identification and biology. Natural History Publications (Borneo), Kota Kinabalu, x + 115 pp." type="book" year="2007">Orr &amp; Hämäläinen 2007</bibRefCitation>
)
</taxonomicName>
.
<taxonomicName id="DCDA4D38E024D743FF79E97FFE4CFAA5" baseAuthorityName="Needham" baseAuthorityYear="1911" box="[151,390,1375,1401]" class="Insecta" family="Calopterygidae" genus="Neurobasis" kingdom="Animalia" order="Odonata" pageId="17" pageNumber="226" phylum="Arthropoda" rank="species" species="chinensis">
<emphasis id="29AEEAA9E024D743FF79E97FFE4CFAA5" box="[151,390,1375,1401]" italics="true" pageId="17" pageNumber="226">Neurobasis chinensis</emphasis>
</taxonomicName>
is also depicted on web pages such as (
<bibRefCitation id="7F4B4B4AE024D743FCA6E97FFC73FAA5" author="Ng, C. Y." box="[840,953,1375,1401]" pageId="17" pageNumber="226" refId="ref25100" refString="Ng, C. Y. (2024 a) Freshwater Macroinvertebrates in Hong Kong: Odonata: Neurobasis chinensis. Available from: https: // www. biosch. hku. hk / ecology / staffhp / dd / macroinvertebrates / Odonata / Calopterygidae / Neurobasis _ chinensis / Neurobasis _ chinensis. html (accessed 22 December 23)" type="url" year="2024">Ng 2024a</bibRefCitation>
) and there is at least one photograph of an unidentified New
<collectingCountry id="63CD762BE024D743FE8FE9A3FE78FA41" box="[353,434,1411,1437]" name="Guinea-Bissau" pageId="17" pageNumber="226">Guinea</collectingCountry>
species in life (
<bibRefCitation id="7F4B4B4AE024D743FDB1E9A3FCBCFA41" author="Orr, A. G. &amp; Hamalainen, M." box="[607,886,1411,1437]" pageId="17" pageNumber="226" refId="ref26055" refString="Orr, A. G. &amp; Hamalainen, M. (2007) The metalwing demoiselles of the eastern tropics - their identification and biology. Natural History Publications (Borneo), Kota Kinabalu, x + 115 pp." type="book" year="2007">Orr &amp; Hämäläinen 2007</bibRefCitation>
). For all species known the spidery larvae inhabit open vegetation, leaf packs and root masses in clear streams and clamber around slowly with careful deliberate movements, perhaps ensuring they do not lose their grip in swift flowing water. In some locations
<taxonomicName id="DCDA4D38E024D743FB19E9EBFA4BFA39" baseAuthorityName="Needham" baseAuthorityYear="1911" box="[1271,1409,1483,1509]" class="Insecta" family="Calopterygidae" genus="Neurobasis" kingdom="Animalia" order="Odonata" pageId="17" pageNumber="226" phylum="Arthropoda" rank="species" species="chinensis">
<emphasis id="29AEEAA9E024D743FB19E9EBFA4BFA39" box="[1271,1409,1483,1509]" italics="true" pageId="17" pageNumber="226">N. chinensis</emphasis>
</taxonomicName>
is recorded in slower flowing water and artificial drains (
<bibRefCitation id="7F4B4B4AE024D743FD1CE9CFFC48F9D5" author="Kumar, A." box="[754,898,1519,1545]" pageId="17" pageNumber="226" pagination="83 - 118" refId="ref24196" refString="Kumar, A. (1973) Descriptions of the last instar larvae of Odonata from the Dehra Dun Valley (India), with notes on biology I. (suborder Zygoptera). Oriental Insects, 7, 83 - 118. https: // doi. org / 10.1080 / 00305316.1973.10434207" type="journal article" year="1973">Kumar 1973</bibRefCitation>
,
<bibRefCitation id="7F4B4B4AE024D743FC62E9CFFB59F9D5" author="Kumar, A. &amp; Prasad, M." box="[908,1171,1519,1545]" pageId="17" pageNumber="226" pagination="163 - 171" refId="ref24326" refString="Kumar, A. &amp; Prasad, M. (1977 b) Reproductive behaviour in Neurobasis chinensis chinensis (Linnaeus) (Zygoptera: Calopterygidae). Odonatologica, 6, 163 - 171." type="journal article" year="1977">Kumar &amp; Prasad 1977b</bibRefCitation>
) It is not known if there are consistent interspecific differences in the larvae except that
<taxonomicName id="DCDA4D38E024D743FCB8EA33FC1FF9F1" baseAuthorityName="Orr &amp; Hamalainen" baseAuthorityYear="2007" box="[854,981,1555,1581]" class="Insecta" family="Calopterygidae" genus="Neurobasis" kingdom="Animalia" order="Odonata" pageId="17" pageNumber="226" phylum="Arthropoda" rank="species" species="longipes">
<emphasis id="29AEEAA9E024D743FCB8EA33FC1FF9F1" box="[854,981,1555,1581]" italics="true" pageId="17" pageNumber="226">N. longipes</emphasis>
</taxonomicName>
evidently has slightly longer legs (
<bibRefCitation id="7F4B4B4AE024D743FAB6EA33FEA8F98D" author="Orr, A. G. &amp; Hamalainen, M." pageId="17" pageNumber="226" refId="ref26055" refString="Orr, A. G. &amp; Hamalainen, M. (2007) The metalwing demoiselles of the eastern tropics - their identification and biology. Natural History Publications (Borneo), Kota Kinabalu, x + 115 pp." type="book" year="2007">Orr &amp; Hämäläinen 2007</bibRefCitation>
). To settle this point detailed direct comparisons of multiple specimens of all species are needed, but as only a few species occur sympatrically, location is often a reliable guide to species identity, and it may also be possible to identify appropriately preserved specimens by DNA analysis. The subgenus
<taxonomicName id="DCDA4D38E024D743FB6FEA5FFB26F945" box="[1153,1260,1663,1689]" pageId="17" pageNumber="226" rank="subGenus" subGenus="Sinobasis">
<emphasis id="29AEEAA9E024D743FB6FEA5FFB26F945" box="[1153,1260,1663,1689]" italics="true" pageId="17" pageNumber="226">Sinobasis</emphasis>
</taxonomicName>
, represented by the poorly known species
<taxonomicName id="DCDA4D38E024D743FE59EA83FD98F961" authorityName="Sjostedt" authorityYear="1926" box="[439,594,1699,1725]" class="Insecta" family="Calopterygidae" genus="Neurobasis" kingdom="Animalia" order="Odonata" pageId="17" pageNumber="226" phylum="Arthropoda" rank="species" species="anderssoni">
<emphasis id="29AEEAA9E024D743FE59EA83FD98F961" box="[439,594,1699,1725]" italics="true" pageId="17" pageNumber="226">N. anderssoni</emphasis>
</taxonomicName>
in southern
<collectingCountry id="63CD762BE024D743FD34EA83FCD7F961" box="[730,797,1699,1725]" name="China" pageId="17" pageNumber="226">China</collectingCountry>
has a heavier build in the adult than subgenus
<taxonomicName id="DCDA4D38E024D743FAF0EA83FE71F93D" authority="(Orr &amp; Hamalainen 2007)" baseAuthorityName="Orr &amp; Hamalainen" baseAuthorityYear="2007" class="Insecta" family="Calopterygidae" genus="Neurobasis" kingdom="Animalia" order="Odonata" pageId="17" pageNumber="226" phylum="Arthropoda" rank="genus">
<emphasis id="29AEEAA9E024D743FAF0EA83FA57F961" box="[1310,1437,1699,1725]" italics="true" pageId="17" pageNumber="226">Neurobasis</emphasis>
(
<bibRefCitation id="7F4B4B4AE024D743FF71EAE7FE79F93D" author="Orr, A. G. &amp; Hamalainen, M." box="[159,435,1735,1761]" pageId="17" pageNumber="226" refId="ref26055" refString="Orr, A. G. &amp; Hamalainen, M. (2007) The metalwing demoiselles of the eastern tropics - their identification and biology. Natural History Publications (Borneo), Kota Kinabalu, x + 115 pp." type="book" year="2007">Orr &amp; Hämäläinen 2007</bibRefCitation>
)
</taxonomicName>
and it seems likely that its larva would also be distinctive.
</paragraph>
</subSubSection>
</treatment>
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88
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Just one larval specimen of this monotypic genus is known (
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).
<taxonomicName id="DCDA4D38E025D742FBF0E97FFA56FAA5" authority="Forster" authorityName="Forster" box="[1054,1436,1375,1401]" class="Insecta" family="Calopterygidae" genus="Matronoides" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="16" pageNumber="225" phylum="Arthropoda" rank="species" species="cyaneipennis">
<emphasis id="29AEEAA9E025D742FBF0E97FFA8FFAA4" box="[1054,1349,1375,1401]" italics="true" pageId="16" pageNumber="225">Matronoides cyaneipennis</emphasis>
Förster
</taxonomicName>
is endemic to altitudes over
<quantity id="DC229B5EE025D742FE31E9A3FDE2FA41" box="[479,552,1411,1437]" metricMagnitude="2" metricUnit="m" metricValue="9.0" pageId="16" pageNumber="225" unit="m" value="900.0">900 m</quantity>
in
<collectingCountry id="63CD762BE025D742FDBFE9A3FD27FA41" box="[593,749,1411,1437]" name="Malaysia" pageId="16" pageNumber="225">North Borneo</collectingCountry>
and in some localities may occur together with
<taxonomicName id="DCDA4D38E025D742FAF0E9A3FE8FFA1D" authority="Hagen" authorityName="Hagen" authorityYear="1887" class="Insecta" family="Calopterygidae" genus="Neurobasis" kingdom="Animalia" order="Odonata" pageId="16" pageNumber="225" phylum="Arthropoda" rank="species" species="longipes">
<emphasis id="29AEEAA9E025D742FAF0E9A3FF3CFA1D" italics="true" pageId="16" pageNumber="225">Neurobasis longipes</emphasis>
Hagen
</taxonomicName>
(R.A. Dow pers. comm.). The adults are reliably seen on Silau Silau Creek in the Kinabalu National Park,
<collectingRegion id="D91EF859E025D742FF32E9EBFEE9FA39" box="[220,291,1483,1509]" country="Malaysia" name="Sabah" pageId="16" pageNumber="225">Sabah</collectingRegion>
, and presumably diligent searching among leaf packs and submerged bankside root masses would produce more material. The larva is of a heavier build than those its sister genus
<taxonomicName id="DCDA4D38E025D742FBCCE9CFFB6BF9D5" baseAuthorityName="Orr &amp; Hamalainen" baseAuthorityYear="2007" box="[1058,1185,1519,1545]" class="Insecta" family="Calopterygidae" genus="Neurobasis" kingdom="Animalia" order="Odonata" pageId="16" pageNumber="225" phylum="Arthropoda" rank="genus">
<emphasis id="29AEEAA9E025D742FBCCE9CFFB6BF9D5" box="[1058,1185,1519,1545]" italics="true" pageId="16" pageNumber="225">Neurobasis</emphasis>
</taxonomicName>
, but the prementum is relatively longer and narrower than in any known
<taxonomicName id="DCDA4D38E025D742FD3FEA33FC63F9F1" box="[721,937,1555,1581]" pageId="16" pageNumber="225">
<emphasis id="29AEEAA9E025D742FD3FEA33FC9AF9F1" box="[721,848,1555,1581]" italics="true" pageId="16" pageNumber="225">Neurobasis</emphasis>
species
</taxonomicName>
(
<figureCitation id="83E12A3EE025D742FC57EA33FBC5F9F1" box="[953,1039,1555,1581]" captionStart="FIGURES 27–34. 27" captionStartId="7.[151,264,1845,1869]" captionTargetBox="[252,1326,159,1834]" captionTargetId="figure-15@7.[251,1336,153,1839]" captionTargetPageId="7" captionText="FIGURES 27–34. 27. Prementum of Calopterygidae showing measurements used in family key (based on Calopteryx japonica, redrawn from Miyakawa 1983). 28. Neurobasis chinensis, prementum and left labial palp (redrawn from Lieftinck 1965). 29. Matronoides cyaneipennis, prementum and labial palps (ventral view). 30. Matrona basilaris, prementum (dorsal view, drawn from Wang et al. 2017). 31. Atrocalopteryx atrata, prementum (dorsal view, redrawn from Miyakawa 1983). 32. Mnais tenuis, prementum (dorsal view, redrawn from Matsuki &amp; Lien 1977). 33. Echo uniformis, prementum and left labial palp dorsal view (redrawn from Lieftinck 1965). 34. Psolodesmus mandarinus, prementum (dorsal view, redrawn from Matsuki &amp; Lien 1978)." figureDoi="http://doi.org/10.5281/zenodo.13618333" httpUri="https://zenodo.org/record/13618333/files/figure.png" pageId="16" pageNumber="225">Fig. 29</figureCitation>
). There is also heavier serration on the outer rib of the external caudal lamellae.
</paragraph>
</subSubSection>
</treatment>
</document>

194
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<subSubSection id="53C06530E025D742FF79ED4FFF36FE55" box="[151,252,367,393]" pageId="16" pageNumber="225" type="nomenclature">
<paragraph id="1B6536BBE025D742FF79ED4FFF36FE55" blockId="16.[151,1437,367,1293]" box="[151,252,367,393]" pageId="16" pageNumber="225">
<heading id="402D81D7E025D742FF79ED4FFF36FE55" bold="true" box="[151,252,367,393]" fontSize="11" level="1" pageId="16" pageNumber="225" reason="1">
<taxonomicName id="DCDA4D38E025D742FF79ED4FFF36FE55" ID-CoL="5M27" authorityName="Selys" authorityYear="1853" box="[151,252,367,393]" class="Insecta" family="Calopterygidae" genus="Matrona" kingdom="Animalia" order="Odonata" pageId="16" pageNumber="225" phylum="Arthropoda" rank="genus">
<emphasis id="29AEEAA9E025D742FF79ED4FFF36FE55" bold="true" box="[151,252,367,393]" italics="true" pageId="16" pageNumber="225">Matrona</emphasis>
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<subSubSection id="53C06530E025D742FF79EDB3FEA7FAD1" pageId="16" pageNumber="225" type="discussion">
<paragraph id="1B6536BBE025D742FF79EDB3FEA7FAD1" blockId="16.[151,1437,367,1293]" pageId="16" pageNumber="225">
The genus
<taxonomicName id="DCDA4D38E025D742FEFDEDB3FEBEFE70" authorityName="Selys" authorityYear="1853" box="[275,372,403,428]" class="Insecta" family="Calopterygidae" genus="Matrona" kingdom="Animalia" order="Odonata" pageId="16" pageNumber="225" phylum="Arthropoda" rank="genus">
<emphasis id="29AEEAA9E025D742FEFDEDB3FEBEFE70" box="[275,372,403,428]" italics="true" pageId="16" pageNumber="225">Matrona</emphasis>
</taxonomicName>
ranges from islands of southern
<collectingCountry id="63CD762BE025D742FD0AEDB3FCEEFE71" box="[740,804,403,429]" name="Japan" pageId="16" pageNumber="225">Japan</collectingCountry>
to
<collectingCountry id="63CD762BE025D742FCA6EDB3FC44FE71" box="[840,910,403,429]" name="China" pageId="16" pageNumber="225">China</collectingCountry>
, Indochina,
<collectingCountry id="63CD762BE025D742FBF9EDB3FBB7FE71" box="[1047,1149,403,429]" name="Thailand" pageId="16" pageNumber="225">Thailand</collectingCountry>
,
<collectingCountry id="63CD762BE025D742FB66EDB3FB3FFE71" box="[1160,1269,403,429]" name="Myanmar" pageId="16" pageNumber="225">Myanmar</collectingCountry>
and north-east
<collectingCountry id="63CD762BE025D742FF79ED97FF1BFE0D" box="[151,209,439,465]" name="India" pageId="16" pageNumber="225">India</collectingCountry>
and includes nine species. It has recently undergone substantial revision (Yu
<emphasis id="29AEEAA9E025D742FBC0ED97FB95FE0D" box="[1070,1119,439,465]" italics="true" pageId="16" pageNumber="225">et al</emphasis>
. 2015) hence although there are several descriptions of the larvae of supposedly
<taxonomicName id="DCDA4D38E025D742FD00EDFBFC71FE29" authorityName="Selys" authorityYear="1853" box="[750,955,475,501]" class="Insecta" family="Calopterygidae" genus="Matrona" kingdom="Animalia" order="Odonata" pageId="16" pageNumber="225" phylum="Arthropoda" rank="species" species="basilaris">
<emphasis id="29AEEAA9E025D742FD00EDFBFC71FE29" box="[750,955,475,501]" italics="true" pageId="16" pageNumber="225">Matrona basilaris</emphasis>
</taxonomicName>
, based on their range some are obviously of other species, and some may be of uncertain identity.
<bibRefCitation id="7F4B4B4AE025D742FCCCEDDFFC03FDC5" author="Fraser, F. C." box="[802,969,511,537]" pageId="16" pageNumber="225" pagination="459 - 467" refId="ref22809" refString="Fraser, F. C. (1919 b) Descriptions of new Indian Odonate larvae and exuviae. Records of the Indian Museum, 16, 459 - 467, pls. 32 - 37 excl. https: // doi. org / 10.5962 / bhl. part. 25934" type="journal article" year="1919">Fraser (1919b)</bibRefCitation>
described
<taxonomicName id="DCDA4D38E025D742FBA9EDDFFAF8FDC5" authority="Selys" authorityName="Selys" authorityYear="1879" box="[1095,1330,511,537]" class="Insecta" family="Calopterygidae" genus="Matrona" kingdom="Animalia" order="Odonata" pageId="16" pageNumber="225" phylum="Arthropoda" rank="species" species="nigripectus">
<emphasis id="29AEEAA9E025D742FBA9EDDFFB26FDC4" box="[1095,1260,511,536]" italics="true" pageId="16" pageNumber="225">M. nigripectus</emphasis>
Selys
</taxonomicName>
larva (as
<taxonomicName id="DCDA4D38E025D742FF79EE03FF33FDE1" authorityName="Selys" authorityYear="1853" box="[151,249,547,573]" class="Insecta" family="Calopterygidae" genus="Matrona" kingdom="Animalia" order="Odonata" pageId="16" pageNumber="225" phylum="Arthropoda" rank="species" species="basilaris">
<emphasis id="29AEEAA9E025D742FF79EE03FF33FDE1" box="[151,249,547,573]" italics="true" pageId="16" pageNumber="225">basilaris</emphasis>
</taxonomicName>
) from the Shillong area in
<collectingRegion id="D91EF859E025D742FDDDEE03FD78FDE1" box="[563,690,547,573]" country="India" name="Meghalaya" pageId="16" pageNumber="225">Meghalaya</collectingRegion>
,
<collectingCountry id="63CD762BE025D742FD51EE03FD33FDE1" box="[703,761,547,573]" name="India" pageId="16" pageNumber="225">India</collectingCountry>
at about
<quantity id="DC229B5EE025D742FC8BEE02FC35FDE1" box="[869,1023,546,573]" metricMagnitude="3" metricUnit="m" metricValue="1.45" metricValueMax="1.5" metricValueMin="1.4" pageId="16" pageNumber="225" unit="m" value="1450.0" valueMax="1500.0" valueMin="1400.0">1400-1500 m</quantity>
, providing a habitus drawing of fair quality and a rather surreal depiction of structural details which nevertheless can be interpreted accurately enough to recognise similarities with other species.
<bibRefCitation id="7F4B4B4AE025D742FD92EE4BFCB7FD59" author="Matsuki, K. &amp; Lien, J. C." box="[636,893,619,645]" pageId="16" pageNumber="225" pagination="15 - 26" refId="ref24810" refString="Matsuki, K. &amp; Lien, J. C. (1978) Descriptions of the larvae of three families of Zygoptera breeding in the streams of Taiwan (Synlestidae, Euphaeidae &amp; Calopterygidae). Tombo - Acta Odonatologica Japonica, 21, 15 - 26." type="journal article" year="1978">Matsuki &amp; Lien (1978)</bibRefCitation>
and
<bibRefCitation id="7F4B4B4AE025D742FC5CEE4BFB8DFD59" author="Ishida, K." box="[946,1095,619,645]" pageId="16" pageNumber="225" refId="ref23597" refString="Ishida, K. (1996) Monograph of Odonata larvae in Japan. Hokkaido University Press, Sapporo, 446 pp. [in Japanese]" type="book" year="1996">Ishida (1996)</bibRefCitation>
provided detailed descriptions and illustrations including the habitus for
<taxonomicName id="DCDA4D38E025D742FD81EEAFFC38FD75" authority="Hamalainen &amp; Yeh" authorityName="Hamalainen &amp; Yeh" authorityYear="2000" box="[623,1010,655,681]" class="Insecta" family="Calopterygidae" genus="Matrona" kingdom="Animalia" order="Odonata" pageId="16" pageNumber="225" phylum="Arthropoda" rank="species" species="cyanoptera">
<emphasis id="29AEEAA9E025D742FD81EEAFFCD9FD74" box="[623,787,655,680]" italics="true" pageId="16" pageNumber="225">M. cyanoptera</emphasis>
Hämäläinen &amp; Yeh
</taxonomicName>
and
<taxonomicName id="DCDA4D38E025D742FBC7EEAFFAC2FD75" authority="Forster" authorityName="Forster" box="[1065,1288,655,681]" class="Insecta" family="Calopterygidae" genus="Calopteryx" kingdom="Animalia" order="Odonata" pageId="16" pageNumber="225" phylum="Arthropoda" rank="species" species="japonica">
<emphasis id="29AEEAA9E025D742FBC7EEAFFB7BFD74" box="[1065,1201,655,680]" italics="true" pageId="16" pageNumber="225">M. japonica</emphasis>
Förster
</taxonomicName>
respectively.
<bibRefCitation id="7F4B4B4AE025D742FF79EE93FE9FFD11" author="Needham, J. G." box="[151,341,691,717]" pageId="16" pageNumber="225" pagination="1 - 20" refId="ref24989" refString="Needham, J. G. (1930) A manual of the dragonflies of China: A monographic study of the Chinese Odonata. [Zoologia Sinica, Series A. Invertebrates of China, Vol. 11. Fascicle 1]. The Fan Memorial Institute of Biology, Peiping, xi + 344 pp., pls. 1 - 20 excl." type="book chapter" year="1930">Needham (1930)</bibRefCitation>
, illustrates the basic habitus sketch of ‘
<taxonomicName id="DCDA4D38E025D742FCEBEE93FC46FD11" authorityName="Selys" authorityYear="1853" box="[773,908,691,717]" class="Insecta" family="Calopterygidae" genus="Matrona" kingdom="Animalia" order="Odonata" pageId="16" pageNumber="225" phylum="Arthropoda" rank="species" species="basilaris">
<emphasis id="29AEEAA9E025D742FCEBEE93FC46FD11" box="[773,908,691,717]" italics="true" pageId="16" pageNumber="225">M. basilaris</emphasis>
</taxonomicName>
’ but as it is evidently based on a specimen from mainland
<collectingCountry id="63CD762BE025D742FEEAEEF7FE8DFD2D" box="[260,327,727,753]" name="China" pageId="16" pageNumber="225">China</collectingCountry>
(Hangchow,
<collectingRegion id="D91EF859E025D742FE37EEF7FD8BFD2D" box="[473,577,727,753]" country="China" name="Zhejiang" pageId="16" pageNumber="225">Zhejiang</collectingRegion>
), its identity must be treated with caution, especially as it depicts exceptionally long legs. Most recently
<bibRefCitation id="7F4B4B4AE025D742FE59EEDBFD5BFCC9" author="Wang, R. &amp; Yu, X. &amp; Xue, J. &amp; Ning, X." box="[439,657,763,789]" pageId="16" pageNumber="225" pagination="580 - 592" refId="ref27434" refString="Wang, R., Yu, X., Xue, J. &amp; Ning, X. (2017) Descriptions of larvae of Vestalaria venusta (Hamalainen, 2004) and Matrona basilaris Selys, 1853 (Odonata: Calopterygidae). Zootaxa, 4306 (4), 580 - 592. https: // doi. org / 10.11646 / zootaxa. 4306.4.8" type="journal article" year="2017">
Wang
<emphasis id="29AEEAA9E025D742FDECEEDBFDFCFCC9" box="[514,566,763,789]" italics="true" pageId="16" pageNumber="225">et al</emphasis>
. (2017)
</bibRefCitation>
described and illustrated photographically
<taxonomicName id="DCDA4D38E025D742FB6AEEDBFAC4FCC9" authorityName="Selys" authorityYear="1853" box="[1156,1294,763,789]" class="Insecta" family="Calopterygidae" genus="Matrona" kingdom="Animalia" order="Odonata" pageId="16" pageNumber="225" phylum="Arthropoda" rank="species" species="basilaris">
<emphasis id="29AEEAA9E025D742FB6AEEDBFAC4FCC9" box="[1156,1294,763,789]" italics="true" pageId="16" pageNumber="225">M. basilaris</emphasis>
</taxonomicName>
larvae from
<collectingRegion id="D91EF859E025D742FF79EF3FFF2AFCE5" box="[151,224,799,825]" country="China" name="Anhui" pageId="16" pageNumber="225">Anhui</collectingRegion>
,
<collectingCountry id="63CD762BE025D742FF00EF3FFEFFFCE5" box="[238,309,799,825]" name="China" pageId="16" pageNumber="225">China</collectingCountry>
. This specimen is a confirmed DNA match with adult
<taxonomicName id="DCDA4D38E025D742FC41EF3FFBF3FCE5" authorityName="Selys" authorityYear="1853" box="[943,1081,799,825]" class="Insecta" family="Calopterygidae" genus="Matrona" kingdom="Animalia" order="Odonata" pageId="16" pageNumber="225" phylum="Arthropoda" rank="species" species="basilaris">
<emphasis id="29AEEAA9E025D742FC41EF3FFBF3FCE5" box="[943,1081,799,825]" italics="true" pageId="16" pageNumber="225">M. basilaris</emphasis>
</taxonomicName>
hence their description can be considered definitive (
<figureCitation id="83E12A3EE025D742FE7CEF63FE23FC81" box="[402,489,835,861]" captionStart="FIGURES 27–34. 27" captionStartId="7.[151,264,1845,1869]" captionTargetBox="[252,1326,159,1834]" captionTargetId="figure-15@7.[251,1336,153,1839]" captionTargetPageId="7" captionText="FIGURES 27–34. 27. Prementum of Calopterygidae showing measurements used in family key (based on Calopteryx japonica, redrawn from Miyakawa 1983). 28. Neurobasis chinensis, prementum and left labial palp (redrawn from Lieftinck 1965). 29. Matronoides cyaneipennis, prementum and labial palps (ventral view). 30. Matrona basilaris, prementum (dorsal view, drawn from Wang et al. 2017). 31. Atrocalopteryx atrata, prementum (dorsal view, redrawn from Miyakawa 1983). 32. Mnais tenuis, prementum (dorsal view, redrawn from Matsuki &amp; Lien 1977). 33. Echo uniformis, prementum and left labial palp dorsal view (redrawn from Lieftinck 1965). 34. Psolodesmus mandarinus, prementum (dorsal view, redrawn from Matsuki &amp; Lien 1978)." figureDoi="http://doi.org/10.5281/zenodo.13618333" httpUri="https://zenodo.org/record/13618333/files/figure.png" pageId="16" pageNumber="225">Figs 30</figureCitation>
,
<figureCitation id="83E12A3EE025D742FE18EF63FDDEFC81" box="[502,532,835,861]" captionStart="FIGURES 42–51. 42" captionStartId="9.[151,264,1592,1616]" captionTargetBox="[165,1413,192,1557]" captionTargetId="figure-15@9.[151,1435,181,1568]" captionTargetPageId="9" captionText="FIGURES 42–51. 42. Archineura incarnata: caudal gills; left lateral above, median below (drawn from Yang et al. 2022). 43. Echo modesta: caudal gills; left lateral above, median below, with baso-dorsal shield arrowed (original illustration). 44. Psolodesmus mandarinus, dorsal view of caudal gills, with baso-dorsal shield of median gill arrowed (redrawn from Matsuki &amp; Lien 1978). 45. Mnais andersoni, left lateral caudal gill with upper margin of median gill visible; baso-dorsal shield of median gill arrowed (original illustration). 46. Mnais gregoryi: caudal gills; left lateral above, median below, with baso-dorsal shield arrowed (drawn from Yang et al. 2021). 47. Caliphaea angka: caudal gills; left lateral above, median below (drawn from Yang et al. 2021). 48. Matrona basilaris: caudal gills; left lateral above, median below (drawn from Wang et al. 2017). 49. Vestalaria venusta: caudal gills; left lateral above, median below (drawn from Wang et al. 2017). 50. Vestalis amoena group: caudal gills; left lateral above, median below (redrawn from Lieftinck 1965). 51. Vestalis melania: caudal gills; left lateral above, median below (redrawn from Guadalquiver et al. 2022)." figureDoi="http://doi.org/10.5281/zenodo.13618337" httpUri="https://zenodo.org/record/13618337/files/figure.png" pageId="16" pageNumber="225">48</figureCitation>
,
<figureCitation id="83E12A3EE025D742FDCEEF62FD8AFC81" box="[544,576,834,861]" captionStart="FIGURES 52–56. 52" captionStartId="10.[151,264,1941,1965]" captionTargetBox="[191,1388,163,1929]" captionTargetId="figure-21@10.[190,1397,151,1933]" captionTargetPageId="10" captionText="FIGURES 52–56. 52. Caliphaea angka, habitus (based on Yang et al. 2021). 53. Vestalis gracilis, habitus (redrawn from Rattanachan et al. 2022). 54. Neurobasis chinensis, habitus (original drawing). 55. Matrona basilaris, habitus (drawn from Wang et al. 2017). 56. Echo uniformis, habitus (redrawn from Lieftinck 1965)." figureDoi="http://doi.org/10.5281/zenodo.13618339" httpUri="https://zenodo.org/record/13618339/files/figure.png" pageId="16" pageNumber="225">55</figureCitation>
). All species are evidently very similar and direct comparisons of specimens are needed to determine any interspecific differences, preferably paired with confirmatory molecular determination. The revision by Yu
<emphasis id="29AEEAA9E025D742FE9AEFABFE6FFC79" box="[372,421,907,933]" italics="true" pageId="16" pageNumber="225">et al</emphasis>
. (2015) also erected a second subgenus,
<taxonomicName id="DCDA4D38E025D742FC9FEFABFC18FC78" authorityName="Selys" authorityYear="1853" box="[881,978,907,932]" class="Insecta" family="Calopterygidae" genus="Matrona" kingdom="Animalia" order="Odonata" pageId="16" pageNumber="225" phylum="Arthropoda" rank="genus">
<emphasis id="29AEEAA9E025D742FC9FEFABFC18FC78" box="[881,978,907,932]" italics="true" pageId="16" pageNumber="225">Matrona</emphasis>
</taxonomicName>
(
<taxonomicName id="DCDA4D38E025D742FC0DEFABFB88FC78" authorityName="Yu, Xue &amp; Hämäläinen" authorityYear="2015" box="[995,1090,907,932]" class="Insecta" family="Calopterygidae" genus="Divortia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="16" pageNumber="225" phylum="Arthropoda" rank="genus">
<emphasis id="29AEEAA9E025D742FC0DEFABFB88FC78" box="[995,1090,907,932]" italics="true" pageId="16" pageNumber="225">Divortia</emphasis>
</taxonomicName>
but so far no confirmed larvae belonging to this taxon have been documented; but workers should be alert to the possibility that they may display clear differences from subgenus
<taxonomicName id="DCDA4D38E025D742FDECEFF3FD15FC31" box="[514,735,979,1005]" class="Insecta" family="Calopterygidae" genus="Matrona" kingdom="Animalia" order="Odonata" pageId="16" pageNumber="225" phylum="Arthropoda" rank="subGenus" subGenus="Matrona">
<emphasis id="29AEEAA9E025D742FDECEFF3FDA9FC30" box="[514,611,979,1004]" italics="true" pageId="16" pageNumber="225">Matrona</emphasis>
(
<emphasis id="29AEEAA9E025D742FD9DEFF3FD1EFC30" box="[627,724,979,1004]" italics="true" pageId="16" pageNumber="225">Matrona</emphasis>
)
</taxonomicName>
. The habits and ecology of the species are variously described: for
<taxonomicName id="DCDA4D38E025D742FF2CEFD7FEA3FBCC" authorityName="Hamalainen &amp; Yeh" authorityYear="2000" box="[194,361,1015,1040]" class="Insecta" family="Calopterygidae" genus="Matrona" kingdom="Animalia" order="Odonata" pageId="16" pageNumber="225" phylum="Arthropoda" rank="species" species="cyanoptera">
<emphasis id="29AEEAA9E025D742FF2CEFD7FEA3FBCC" box="[194,361,1015,1040]" italics="true" pageId="16" pageNumber="225">M. cyanoptera</emphasis>
</taxonomicName>
, “Distributed in the plains and low hills throughout
<collectingCountry id="63CD762BE025D742FC3EEFD7FBECFBCD" box="[976,1062,1015,1041]" name="Taiwan" pageId="16" pageNumber="225">Taiwan</collectingCountry>
; the larva living in [the] middle section of streams resting on tree roots branches and stems in unshaded slow streams”, (
<bibRefCitation id="7F4B4B4AE025D742FB9EE83BFAADFBE9" author="Matsuki, K. &amp; Lien, J. C." box="[1136,1383,1051,1077]" pageId="16" pageNumber="225" pagination="15 - 26" refId="ref24810" refString="Matsuki, K. &amp; Lien, J. C. (1978) Descriptions of the larvae of three families of Zygoptera breeding in the streams of Taiwan (Synlestidae, Euphaeidae &amp; Calopterygidae). Tombo - Acta Odonatologica Japonica, 21, 15 - 26." type="journal article" year="1978">Matsuki &amp; Lien 1978</bibRefCitation>
); for
<taxonomicName id="DCDA4D38E025D742FF79E81FFED5FB85" authorityName="Selys" authorityYear="1853" box="[151,287,1087,1113]" class="Insecta" family="Calopterygidae" genus="Matrona" kingdom="Animalia" order="Odonata" pageId="16" pageNumber="225" phylum="Arthropoda" rank="species" species="basilaris">
<emphasis id="29AEEAA9E025D742FF79E81FFED5FB85" box="[151,287,1087,1113]" italics="true" pageId="16" pageNumber="225">M. basilaris</emphasis>
</taxonomicName>
, “...found both in small montane streams with sandy substrates and open rivers with stony substrates). Some [larva] even occurred in very small puddles covered with dense vegetation formed by discontinuous streams. They were usually concealed among stones and gravel, or in water plants, depending on their body colour. The habitat usually had many adults of
<taxonomicName id="DCDA4D38E025D742FDC9E88BFCDDFB19" authority="Oguma" authorityName="Oguma" authorityYear="1913" box="[551,791,1195,1221]" class="Insecta" family="Calopterygidae" genus="Mnais" kingdom="Animalia" order="Odonata" pageId="16" pageNumber="225" phylum="Arthropoda" rank="species" species="tenuis">
<emphasis id="29AEEAA9E025D742FDC9E88BFD72FB18" box="[551,696,1195,1220]" italics="true" pageId="16" pageNumber="225">Mnais tenuis</emphasis>
Oguma
</taxonomicName>
”, (
<bibRefCitation id="7F4B4B4AE025D742FCD8E88BFC35FB19" author="Wang, R. &amp; Yu, X. &amp; Xue, J. &amp; Ning, X." box="[822,1023,1195,1221]" pageId="16" pageNumber="225" pagination="580 - 592" refId="ref27434" refString="Wang, R., Yu, X., Xue, J. &amp; Ning, X. (2017) Descriptions of larvae of Vestalaria venusta (Hamalainen, 2004) and Matrona basilaris Selys, 1853 (Odonata: Calopterygidae). Zootaxa, 4306 (4), 580 - 592. https: // doi. org / 10.11646 / zootaxa. 4306.4.8" type="journal article" year="2017">
Wang
<emphasis id="29AEEAA9E025D742FC6FE88BFC79FB19" box="[897,947,1195,1221]" italics="true" pageId="16" pageNumber="225">et al</emphasis>
. 2017
</bibRefCitation>
); and for
<taxonomicName id="DCDA4D38E025D742FB83E88BFAD8FB18" authorityName="Selys" authorityYear="1879" box="[1133,1298,1195,1220]" class="Insecta" family="Calopterygidae" genus="Matrona" kingdom="Animalia" order="Odonata" pageId="16" pageNumber="225" phylum="Arthropoda" rank="species" species="nigripectus">
<emphasis id="29AEEAA9E025D742FB83E88BFAD8FB18" box="[1133,1298,1195,1220]" italics="true" pageId="16" pageNumber="225">M. nigripectus</emphasis>
</taxonomicName>
, “Generally concealed amongst debris, dead twigs etc or lying under cover of rocks. Movements sluggish, ...resembles
<taxonomicName id="DCDA4D38E025D742FAAEE8EFFA56FB34" authorityName="Fabricius" authorityYear="1790" box="[1344,1436,1231,1256]" class="Insecta" family="Nepidae" genus="Ranatra" kingdom="Animalia" order="Hemiptera" pageId="16" pageNumber="225" phylum="Arthropoda" rank="genus">
<emphasis id="29AEEAA9E025D742FAAEE8EFFA56FB34" box="[1344,1436,1231,1256]" italics="true" pageId="16" pageNumber="225">Ranatra</emphasis>
</taxonomicName>
...” (
<bibRefCitation id="7F4B4B4AE025D742FF29E8D3FEAAFAD1" author="Fraser, F. C." box="[199,352,1267,1293]" pageId="16" pageNumber="225" pagination="459 - 467" refId="ref22809" refString="Fraser, F. C. (1919 b) Descriptions of new Indian Odonate larvae and exuviae. Records of the Indian Museum, 16, 459 - 467, pls. 32 - 37 excl. https: // doi. org / 10.5962 / bhl. part. 25934" type="journal article" year="1919">Fraser 1919b</bibRefCitation>
).
</paragraph>
</subSubSection>
</treatment>
</document>

222
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Within the Oriental realm there are nine described species in the genus
<taxonomicName id="DCDA4D38E025D742FC52EA83FBC8F960" authorityName="Selys" authorityYear="1853" box="[956,1026,1699,1724]" class="Insecta" family="Calopterygidae" genus="Mnais" kingdom="Animalia" order="Odonata" pageId="16" pageNumber="225" phylum="Arthropoda" rank="genus">
<emphasis id="29AEEAA9E025D742FC52EA83FBC8F960" box="[956,1026,1699,1724]" italics="true" pageId="16" pageNumber="225">Mnais</emphasis>
</taxonomicName>
but this number is likely to increase as suggested by
<bibRefCitation id="7F4B4B4AE025D742FEA0EAE7FE21F93D" author="Zhang, H. - M." box="[334,491,1735,1761]" pageId="16" pageNumber="225" refId="ref28336" refString="Zhang, H. - M. (2019) Dragonflies and Damselflies of China. Vols. 1 &amp; 2. Chong Qing University Press, ChongQing, 14 + 1460 pp. [bilingual, in Chinese and English]" type="book" year="2019">Zhang (2019)</bibRefCitation>
. The genus ranges from
<collectingCountry id="63CD762BE025D742FD13EAE7FC8AF93D" box="[765,832,1735,1761]" name="China" pageId="16" pageNumber="225">China</collectingCountry>
and
<collectingCountry id="63CD762BE025D742FC99EAE7FC7DF93D" box="[887,951,1735,1761]" name="Japan" pageId="16" pageNumber="225">Japan</collectingCountry>
to Indochina,
<collectingCountry id="63CD762BE025D742FBB6EAE7FB76F93D" box="[1112,1212,1735,1761]" name="Thailand" pageId="16" pageNumber="225">Thailand</collectingCountry>
and
<collectingCountry id="63CD762BE025D742FB1CEAE7FAABF93D" box="[1266,1377,1735,1761]" name="Myanmar" pageId="16" pageNumber="225">Myanmar</collectingCountry>
. Two species,
<taxonomicName id="DCDA4D38E025D742FF1BEACBFE11F8D9" authority="Selys" authorityName="Selys" authorityYear="1869" box="[245,475,1771,1797]" class="Insecta" family="Calopterygidae" genus="Mnais" kingdom="Animalia" order="Odonata" pageId="16" pageNumber="225" phylum="Arthropoda" rank="species" species="costalis">
<emphasis id="29AEEAA9E025D742FF1BEACBFE52F8D9" box="[245,408,1771,1797]" italics="true" pageId="16" pageNumber="225">Mnais costalis</emphasis>
Selys
</taxonomicName>
and
<taxonomicName id="DCDA4D38E025D742FDFEEACBFD11F8D9" authority="Selys" authorityName="Selys" authorityYear="1853" box="[528,731,1771,1797]" class="Insecta" family="Calopterygidae" genus="Mnais" kingdom="Animalia" order="Odonata" pageId="16" pageNumber="225" phylum="Arthropoda" rank="species" species="pruinosa">
<emphasis id="29AEEAA9E025D742FDFEEACBFD52F8D8" box="[528,664,1771,1796]" italics="true" pageId="16" pageNumber="225">M. pruinosa</emphasis>
Selys
</taxonomicName>
are known only from the main islands of
<collectingCountry id="63CD762BE025D742FB45EACBFB21F8D9" box="[1195,1259,1771,1797]" name="Japan" pageId="16" pageNumber="225">Japan</collectingCountry>
and fall outside the Oriental region, but their larval morphology, figured in detail by
<bibRefCitation id="7F4B4B4AE025D742FC43EB2FFB25F8F5" author="Hirose, K. &amp; Rokuyama, Y." box="[941,1263,1807,1833]" pageId="16" pageNumber="225" pagination="23 - 27" refId="ref23425" refString="Hirose, K. &amp; Rokuyama, Y. (1966) Two types of the larva of Mnais strigata and their distribution in the streams. Tombo - Acta Odonatologica Japonica, 9, 23 - 27. [in Japanese]" type="journal article" year="1966">Hirose &amp; Rokuyama (1966)</bibRefCitation>
,
<bibRefCitation id="7F4B4B4AE025D742FB13EB2FFA5DF8F5" author="Ishida, K." box="[1277,1431,1807,1833]" pageId="16" pageNumber="225" refId="ref23597" refString="Ishida, K. (1996) Monograph of Odonata larvae in Japan. Hokkaido University Press, Sapporo, 446 pp. [in Japanese]" type="book" year="1996">Ishida (1996)</bibRefCitation>
, and revisited by
<bibRefCitation id="7F4B4B4AE025D742FEB8EB13FD80F891" author="Hayashi, F. &amp; Dobata, S. &amp; Futahashi, R." box="[342,586,1843,1869]" pageId="16" pageNumber="225" pagination="13 - 24" refId="ref23371" refString="Hayashi, F., Dobata, S. &amp; Futahashi, R. (2004) Larval morphology of Japanese Mnais damselflies (Odonata: Calopterygidae) distinguished by nuclear DNA (ITS l) sequences. Tombo - Acta Odonatologica Japonica, 47, 13 - 24. [in Japanese]" type="journal article" year="2004">
Hayashi
<emphasis id="29AEEAA9E025D742FE54EB13FE3EF891" box="[442,500,1843,1869]" italics="true" pageId="16" pageNumber="225">et al.</emphasis>
(2004)
</bibRefCitation>
, are considered here as they give insight into the range of variation within the genus. The larva of
<taxonomicName id="DCDA4D38E025D742FE4CEB77FDAFF8AD" authority="Oguma" authorityName="Oguma" authorityYear="1913" box="[418,613,1879,1905]" class="Insecta" family="Calopterygidae" genus="Mnais" kingdom="Animalia" order="Odonata" pageId="16" pageNumber="225" phylum="Arthropoda" rank="species" species="tenuis">
<emphasis id="29AEEAA9E025D742FE4CEB77FDC1F8AC" box="[418,523,1879,1904]" italics="true" pageId="16" pageNumber="225">M. tenuis</emphasis>
Oguma
</taxonomicName>
was described from
<collectingCountry id="63CD762BE025D742FCBFEB77FC69F8AD" box="[849,931,1879,1905]" name="Taiwan" pageId="16" pageNumber="225">Taiwan</collectingCountry>
(as
<taxonomicName id="DCDA4D38E025D742FC3DEB77FB79F8AC" box="[979,1203,1879,1905]" class="Insecta" family="Calopterygidae" genus="Mnais" kingdom="Animalia" order="Odonata" pageId="16" pageNumber="225" phylum="Arthropoda" rank="subSpecies" species="andersoni" subSpecies="tenuis">
<emphasis id="29AEEAA9E025D742FC3DEB77FB79F8AC" box="[979,1203,1879,1905]" italics="true" pageId="16" pageNumber="225">M. andersoni tenuis</emphasis>
</taxonomicName>
) by
<bibRefCitation id="7F4B4B4AE025D742FB06EB77FF2CF849" author="Matsuki, K. &amp; Lien, J. C." pageId="16" pageNumber="225" pagination="15 - 26" refId="ref24810" refString="Matsuki, K. &amp; Lien, J. C. (1978) Descriptions of the larvae of three families of Zygoptera breeding in the streams of Taiwan (Synlestidae, Euphaeidae &amp; Calopterygidae). Tombo - Acta Odonatologica Japonica, 21, 15 - 26." type="journal article" year="1978">Matsuki &amp; Lien (1978)</bibRefCitation>
. The larva of
<taxonomicName id="DCDA4D38E025D742FE66EB5BFDE7F849" authority="Ris" authorityName="Ris" authorityYear="1916" box="[392,557,1915,1941]" class="Insecta" family="Calopterygidae" genus="Mnais" kingdom="Animalia" order="Odonata" pageId="16" pageNumber="225" phylum="Arthropoda" rank="species" species="mneme">
<emphasis id="29AEEAA9E025D742FE66EB5BFE34F848" box="[392,510,1915,1940]" italics="true" pageId="16" pageNumber="225">M. mneme</emphasis>
Ris
</taxonomicName>
was described and figured by
<bibRefCitation id="7F4B4B4AE025D742FC7EEB5BFB6BF849" author="Matsuki, K. &amp; Saito, Y." box="[912,1185,1915,1941]" pageId="16" pageNumber="225" pagination="43 - 45" refId="ref24859" refString="Matsuki, K. &amp; Saito, Y. (2000) Description of the larva of Mnais mneme (Ris, 1916) (Calopterygidae) from Hong Kong. Tombo - Acta Odonatologica Japonica, 42, 43 - 45. [in Japanese]" type="journal article" year="2000">Matsuki &amp; Saito (2000)</bibRefCitation>
and photographs of a young larva and some details of a more mature specimen of the same species are provided by (
<bibRefCitation id="7F4B4B4AE025D742FB43EBBFFAEBF865" author="Ng, C. Y." box="[1197,1313,1951,1977]" pageId="16" pageNumber="225" refId="ref25162" refString="Ng, C. Y. (2024 b) Freshwater Macroinvertebrates in Hong Kong: Odonata: Mnais mneme. Available from: https: // www. biosch. hku. hk / ecology / staffhp / dd / macroinvertebrates / Odonata / Calopterygidae / Mnias _ mneme / Mnias _ mneme. html (accessed 16 May 2024)" type="url" year="2024">Ng 2024b</bibRefCitation>
). The larva of
<taxonomicName id="DCDA4D38E025D742FF5BEBE3FE81F801" authorityName="McLachlan in Selys" authorityYear="1873" box="[181,331,1987,2013]" class="Insecta" family="Calopterygidae" genus="Mnais" kingdom="Animalia" order="Odonata" pageId="16" pageNumber="225" phylum="Arthropoda" rank="species" species="andersoni">
<emphasis id="29AEEAA9E025D742FF5BEBE3FE81F801" box="[181,331,1987,2013]" italics="true" pageId="16" pageNumber="225">M. andersoni</emphasis>
</taxonomicName>
is known (Orr &amp; Keetapithchayakul unpublished data), with the caudal gills figured (
<figureCitation id="83E12A3EE025D742FAE4EBE3FA94F801" box="[1290,1374,1986,2013]" captionStart="FIGURES 42–51. 42" captionStartId="9.[151,264,1592,1616]" captionTargetBox="[165,1413,192,1557]" captionTargetId="figure-15@9.[151,1435,181,1568]" captionTargetPageId="9" captionText="FIGURES 42–51. 42. Archineura incarnata: caudal gills; left lateral above, median below (drawn from Yang et al. 2022). 43. Echo modesta: caudal gills; left lateral above, median below, with baso-dorsal shield arrowed (original illustration). 44. Psolodesmus mandarinus, dorsal view of caudal gills, with baso-dorsal shield of median gill arrowed (redrawn from Matsuki &amp; Lien 1978). 45. Mnais andersoni, left lateral caudal gill with upper margin of median gill visible; baso-dorsal shield of median gill arrowed (original illustration). 46. Mnais gregoryi: caudal gills; left lateral above, median below, with baso-dorsal shield arrowed (drawn from Yang et al. 2021). 47. Caliphaea angka: caudal gills; left lateral above, median below (drawn from Yang et al. 2021). 48. Matrona basilaris: caudal gills; left lateral above, median below (drawn from Wang et al. 2017). 49. Vestalaria venusta: caudal gills; left lateral above, median below (drawn from Wang et al. 2017). 50. Vestalis amoena group: caudal gills; left lateral above, median below (redrawn from Lieftinck 1965). 51. Vestalis melania: caudal gills; left lateral above, median below (redrawn from Guadalquiver et al. 2022)." figureDoi="http://doi.org/10.5281/zenodo.13618337" httpUri="https://zenodo.org/record/13618337/files/figure.png" pageId="16" pageNumber="225">Fig. 45</figureCitation>
). The larva of
<taxonomicName id="DCDA4D38E024D743FF1BECB7FE0DFF6D" authority="Fraser" authorityName="Fraser" authorityYear="1924" box="[245,455,151,177]" class="Insecta" family="Calopterygidae" genus="Mnais" kingdom="Animalia" order="Odonata" pageId="17" pageNumber="226" phylum="Arthropoda" rank="species" species="gregoryi">
<emphasis id="29AEEAA9E024D743FF1BECB7FEB3FF6C" box="[245,377,151,176]" italics="true" pageId="17" pageNumber="226">M. gregoryi</emphasis>
Fraser
</taxonomicName>
was described in detail by
<bibRefCitation id="7F4B4B4AE024D743FD19ECB7FC08FF6D" author="Yang, G. - H. &amp; Zhang, H. - M. &amp; Orr, A. G." box="[759,962,151,177]" pageId="17" pageNumber="226" pagination="276 - 284" refId="ref27918" refString="Yang, G. - H., Zhang, H. - M. &amp; Orr, A. G. (2021) Descriptions of larvae of Caliphaea angka Hamalainen, 2003 and Mnais gregoryi Fraser, 1924 (Odonata: Calopterygidae). Zootaxa, 4926 (2), 276 - 284. https: // doi. org / 10.11646 / zootaxa. 4926.2.7" type="journal article" year="2021">
Yang
<emphasis id="29AEEAA9E024D743FCD6ECB7FCA0FF6D" box="[824,874,151,177]" italics="true" pageId="17" pageNumber="226">et al</emphasis>
. (2021)
</bibRefCitation>
from specimens collected at ca
<quantity id="DC229B5EE024D743FAC4ECB7FAB5FF6D" box="[1322,1407,151,177]" metricMagnitude="3" metricUnit="m" metricValue="2.3" pageId="17" pageNumber="226" unit="m" value="2300.0">2300 m</quantity>
in
<collectingRegion id="D91EF859E024D743FF79EC9BFF3AFF09" box="[151,240,187,213]" country="China" name="Yunnan" pageId="17" pageNumber="226">Yunnan</collectingRegion>
,
<collectingCountry id="63CD762BE024D743FF15EC9BFE88FF09" box="[251,322,187,213]" name="China" pageId="17" pageNumber="226">China</collectingCountry>
.
</paragraph>
<paragraph id="1B6536BBE024D743FF79ED23FD19FD75" blockId="17.[151,1437,259,681]" pageId="17" pageNumber="226">
In all species the head is fairly broad and semi-pentangular; the body moderately stout to elongate (
<taxonomicName id="DCDA4D38E024D743FAE8ED23FA41FEC0" authorityName="Fraser" authorityYear="1924" box="[1286,1419,259,284]" class="Insecta" family="Calopterygidae" genus="Mnais" kingdom="Animalia" order="Odonata" pageId="17" pageNumber="226" phylum="Arthropoda" rank="species" species="gregoryi">
<emphasis id="29AEEAA9E024D743FAE8ED23FA41FEC0" box="[1286,1419,259,284]" italics="true" pageId="17" pageNumber="226">M. gregoryi</emphasis>
</taxonomicName>
).
<bibRefCitation id="7F4B4B4AE024D743FF79ED07FE28FE9D" author="Hirose, K. &amp; Rokuyama, Y." box="[151,482,295,321]" pageId="17" pageNumber="226" pagination="23 - 27" refId="ref23425" refString="Hirose, K. &amp; Rokuyama, Y. (1966) Two types of the larva of Mnais strigata and their distribution in the streams. Tombo - Acta Odonatologica Japonica, 9, 23 - 27. [in Japanese]" type="journal article" year="1966">Hirose &amp; Rokuyama’s (1966)</bibRefCitation>
and
<bibRefCitation id="7F4B4B4AE024D743FDF6ED07FD0BFE9D" author="Ishida, K." box="[536,705,295,321]" pageId="17" pageNumber="226" refId="ref23597" refString="Ishida, K. (1996) Monograph of Odonata larvae in Japan. Hokkaido University Press, Sapporo, 446 pp. [in Japanese]" type="book" year="1996">Ishida’s (1996)</bibRefCitation>
illustrations of the habitus show stouter bodies in
<taxonomicName id="DCDA4D38E024D743FB1CED07FAA7FE9D" authorityName="Selys" authorityYear="1869" box="[1266,1389,295,321]" class="Insecta" family="Calopterygidae" genus="Mnais" kingdom="Animalia" order="Odonata" pageId="17" pageNumber="226" phylum="Arthropoda" rank="species" species="costalis">
<emphasis id="29AEEAA9E024D743FB1CED07FAA7FE9D" box="[1266,1389,295,321]" italics="true" pageId="17" pageNumber="226">M. costalis</emphasis>
</taxonomicName>
and
<taxonomicName id="DCDA4D38E024D743FF79ED6BFEE8FEB8" authorityName="Selys" authorityYear="1853" box="[151,290,331,356]" class="Insecta" family="Calopterygidae" genus="Mnais" kingdom="Animalia" order="Odonata" pageId="17" pageNumber="226" phylum="Arthropoda" rank="species" species="pruinosa">
<emphasis id="29AEEAA9E024D743FF79ED6BFEE8FEB8" box="[151,290,331,356]" italics="true" pageId="17" pageNumber="226">M. pruinosa</emphasis>
</taxonomicName>
, and this is confirmed by photographs.
<bibRefCitation id="7F4B4B4AE024D743FD0AED6BFC26FEB9" author="Matsuki, K. &amp; Lien, J. C." box="[740,1004,331,357]" pageId="17" pageNumber="226" pagination="15 - 26" refId="ref24810" refString="Matsuki, K. &amp; Lien, J. C. (1978) Descriptions of the larvae of three families of Zygoptera breeding in the streams of Taiwan (Synlestidae, Euphaeidae &amp; Calopterygidae). Tombo - Acta Odonatologica Japonica, 21, 15 - 26." type="journal article" year="1978">Matsuki &amp; Lien (1978)</bibRefCitation>
depict a much thinner, but still robust habitus for
<taxonomicName id="DCDA4D38E024D743FEF6ED4FFE4AFE54" authorityName="Oguma" authorityYear="1913" box="[280,384,367,392]" class="Insecta" family="Calopterygidae" genus="Mnais" kingdom="Animalia" order="Odonata" pageId="17" pageNumber="226" phylum="Arthropoda" rank="species" species="tenuis">
<emphasis id="29AEEAA9E024D743FEF6ED4FFE4AFE54" box="[280,384,367,392]" italics="true" pageId="17" pageNumber="226">M. tenuis</emphasis>
</taxonomicName>
. The prementum is broad apically, with wide anterior lobes that may overlap at the tips and a comparatively short very narrow tear-drop shaped median cleft (
<figureCitation id="83E12A3EE024D743FC85EDB3FC0AFE71" box="[875,960,403,429]" captionStart="FIGURES 27–34. 27" captionStartId="7.[151,264,1845,1869]" captionTargetBox="[252,1326,159,1834]" captionTargetId="figure-15@7.[251,1336,153,1839]" captionTargetPageId="7" captionText="FIGURES 27–34. 27. Prementum of Calopterygidae showing measurements used in family key (based on Calopteryx japonica, redrawn from Miyakawa 1983). 28. Neurobasis chinensis, prementum and left labial palp (redrawn from Lieftinck 1965). 29. Matronoides cyaneipennis, prementum and labial palps (ventral view). 30. Matrona basilaris, prementum (dorsal view, drawn from Wang et al. 2017). 31. Atrocalopteryx atrata, prementum (dorsal view, redrawn from Miyakawa 1983). 32. Mnais tenuis, prementum (dorsal view, redrawn from Matsuki &amp; Lien 1977). 33. Echo uniformis, prementum and left labial palp dorsal view (redrawn from Lieftinck 1965). 34. Psolodesmus mandarinus, prementum (dorsal view, redrawn from Matsuki &amp; Lien 1978)." figureDoi="http://doi.org/10.5281/zenodo.13618333" httpUri="https://zenodo.org/record/13618333/files/figure.png" pageId="17" pageNumber="226">Fig. 32</figureCitation>
). There is normally one pair of strong seta at the base of the lobes. The caudal gills of most known species (
<figureCitation id="83E12A3EE024D743FC8AED97FC7FFE0D" box="[868,949,438,465]" captionStart="FIGURES 42–51. 42" captionStartId="9.[151,264,1592,1616]" captionTargetBox="[165,1413,192,1557]" captionTargetId="figure-15@9.[151,1435,181,1568]" captionTargetPageId="9" captionText="FIGURES 42–51. 42. Archineura incarnata: caudal gills; left lateral above, median below (drawn from Yang et al. 2022). 43. Echo modesta: caudal gills; left lateral above, median below, with baso-dorsal shield arrowed (original illustration). 44. Psolodesmus mandarinus, dorsal view of caudal gills, with baso-dorsal shield of median gill arrowed (redrawn from Matsuki &amp; Lien 1978). 45. Mnais andersoni, left lateral caudal gill with upper margin of median gill visible; baso-dorsal shield of median gill arrowed (original illustration). 46. Mnais gregoryi: caudal gills; left lateral above, median below, with baso-dorsal shield arrowed (drawn from Yang et al. 2021). 47. Caliphaea angka: caudal gills; left lateral above, median below (drawn from Yang et al. 2021). 48. Matrona basilaris: caudal gills; left lateral above, median below (drawn from Wang et al. 2017). 49. Vestalaria venusta: caudal gills; left lateral above, median below (drawn from Wang et al. 2017). 50. Vestalis amoena group: caudal gills; left lateral above, median below (redrawn from Lieftinck 1965). 51. Vestalis melania: caudal gills; left lateral above, median below (redrawn from Guadalquiver et al. 2022)." figureDoi="http://doi.org/10.5281/zenodo.13618337" httpUri="https://zenodo.org/record/13618337/files/figure.png" pageId="17" pageNumber="226">Fig. 45</figureCitation>
) are typically an elongate oval shape, about 2.5–3 times as long as broad, somewhat flattened, triquetral with no spines or tubercules on the margins or central rib in known species; the central gill is membranous and typically almost as long as the lateral gills; at its base there is a broad dorsal triangular shield that is typically as long as it is wide, but may be longer (
<taxonomicName id="DCDA4D38E024D743FB58EE03FAF6FDE0" authorityName="Fraser" authorityYear="1924" box="[1206,1340,547,572]" class="Insecta" family="Calopterygidae" genus="Mnais" kingdom="Animalia" order="Odonata" pageId="17" pageNumber="226" phylum="Arthropoda" rank="species" species="gregoryi">
<emphasis id="29AEEAA9E024D743FB58EE03FAF6FDE0" box="[1206,1340,547,572]" italics="true" pageId="17" pageNumber="226">M. gregoryi</emphasis>
</taxonomicName>
).
<taxonomicName id="DCDA4D38E024D743FAB8EE03FF3CFDBC" authorityName="Fraser" authorityYear="1924" class="Insecta" family="Calopterygidae" genus="Mnais" kingdom="Animalia" order="Odonata" pageId="17" pageNumber="226" phylum="Arthropoda" rank="species" species="gregoryi">
<emphasis id="29AEEAA9E024D743FAB8EE03FF3CFDBC" italics="true" pageId="17" pageNumber="226">Mnais gregoryi</emphasis>
</taxonomicName>
differs considerably from other species in the shape of its caudal gills, the laterals being long and tapered to a point, about five times as long as broad, whereas the median gill, which tapers to a broader rounded point, is about two thirds the length of the laterals (
<figureCitation id="83E12A3EE024D743FD9CEEAFFD0CFD75" box="[626,710,655,681]" captionStart="FIGURES 42–51. 42" captionStartId="9.[151,264,1592,1616]" captionTargetBox="[165,1413,192,1557]" captionTargetId="figure-15@9.[151,1435,181,1568]" captionTargetPageId="9" captionText="FIGURES 42–51. 42. Archineura incarnata: caudal gills; left lateral above, median below (drawn from Yang et al. 2022). 43. Echo modesta: caudal gills; left lateral above, median below, with baso-dorsal shield arrowed (original illustration). 44. Psolodesmus mandarinus, dorsal view of caudal gills, with baso-dorsal shield of median gill arrowed (redrawn from Matsuki &amp; Lien 1978). 45. Mnais andersoni, left lateral caudal gill with upper margin of median gill visible; baso-dorsal shield of median gill arrowed (original illustration). 46. Mnais gregoryi: caudal gills; left lateral above, median below, with baso-dorsal shield arrowed (drawn from Yang et al. 2021). 47. Caliphaea angka: caudal gills; left lateral above, median below (drawn from Yang et al. 2021). 48. Matrona basilaris: caudal gills; left lateral above, median below (drawn from Wang et al. 2017). 49. Vestalaria venusta: caudal gills; left lateral above, median below (drawn from Wang et al. 2017). 50. Vestalis amoena group: caudal gills; left lateral above, median below (redrawn from Lieftinck 1965). 51. Vestalis melania: caudal gills; left lateral above, median below (redrawn from Guadalquiver et al. 2022)." figureDoi="http://doi.org/10.5281/zenodo.13618337" httpUri="https://zenodo.org/record/13618337/files/figure.png" pageId="17" pageNumber="226">Fig. 46</figureCitation>
).
</paragraph>
<paragraph id="1B6536BBE024D743FF79EEF7FE6EFC31" blockId="17.[151,1437,727,1005]" pageId="17" pageNumber="226">
The larvae are often found in quiet disturbed areas near the head of streams clinging to vegetation. In tropical and subtropical regions, they are chiefly montane, with
<taxonomicName id="DCDA4D38E024D743FD0FEEDBFCACFCC8" authorityName="Fraser" authorityYear="1924" box="[737,870,763,788]" class="Insecta" family="Calopterygidae" genus="Mnais" kingdom="Animalia" order="Odonata" pageId="17" pageNumber="226" phylum="Arthropoda" rank="species" species="gregoryi">
<emphasis id="29AEEAA9E024D743FD0FEEDBFCACFCC8" box="[737,870,763,788]" italics="true" pageId="17" pageNumber="226">M. gregoryi</emphasis>
</taxonomicName>
reaching
<quantity id="DC229B5EE024D743FC36EEDAFBECFCC9" box="[984,1062,762,789]" metricMagnitude="3" metricUnit="m" metricValue="2.5" pageId="17" pageNumber="226" unit="m" value="2500.0">2500m</quantity>
in
<collectingRegion id="D91EF859E024D743FBA5EEDBFB6EFCC9" box="[1099,1188,763,789]" country="China" name="Yunnan" pageId="17" pageNumber="226">Yunnan</collectingRegion>
,
<collectingCountry id="63CD762BE024D743FB5EEEDBFB39FCC9" box="[1200,1267,763,789]" name="China" pageId="17" pageNumber="226">China</collectingCountry>
(
<bibRefCitation id="7F4B4B4AE024D743FAECEEDBFA5BFCC9" author="Zhang, H. - M." box="[1282,1425,763,789]" pageId="17" pageNumber="226" refId="ref28336" refString="Zhang, H. - M. (2019) Dragonflies and Damselflies of China. Vols. 1 &amp; 2. Chong Qing University Press, ChongQing, 14 + 1460 pp. [bilingual, in Chinese and English]" type="book" year="2019">Zhang 2019</bibRefCitation>
).
<bibRefCitation id="7F4B4B4AE024D743FF79EF3FFE5DFCE5" author="Matsuki, K. &amp; Lien, J. C." box="[151,407,799,825]" pageId="17" pageNumber="226" pagination="15 - 26" refId="ref24810" refString="Matsuki, K. &amp; Lien, J. C. (1978) Descriptions of the larvae of three families of Zygoptera breeding in the streams of Taiwan (Synlestidae, Euphaeidae &amp; Calopterygidae). Tombo - Acta Odonatologica Japonica, 21, 15 - 26." type="journal article" year="1978">Matsuki &amp; Lien (1978)</bibRefCitation>
described
<taxonomicName id="DCDA4D38E024D743FDE0EF3FFDBFFCE4" authorityName="Oguma" authorityYear="1913" box="[526,629,799,824]" class="Insecta" family="Calopterygidae" genus="Mnais" kingdom="Animalia" order="Odonata" pageId="17" pageNumber="226" phylum="Arthropoda" rank="species" species="tenuis">
<emphasis id="29AEEAA9E024D743FDE0EF3FFDBFFCE4" box="[526,629,799,824]" italics="true" pageId="17" pageNumber="226">M. tenuis</emphasis>
</taxonomicName>
as inhabiting “...low hills in northern parts of
<collectingCountry id="63CD762BE024D743FB80EF3FFB0EFCE5" box="[1134,1220,799,825]" name="Taiwan" pageId="17" pageNumber="226">Taiwan</collectingCountry>
; the larvae living in fountainhead of streams, resting on tree-roots, branches and leaves in shaded slow streams.”, and
<bibRefCitation id="7F4B4B4AE024D743FB3DEF63FA56FC81" author="Yang, G. - H. &amp; Zhang, H. - M. &amp; Orr, A. G." box="[1235,1436,835,861]" pageId="17" pageNumber="226" pagination="276 - 284" refId="ref27918" refString="Yang, G. - H., Zhang, H. - M. &amp; Orr, A. G. (2021) Descriptions of larvae of Caliphaea angka Hamalainen, 2003 and Mnais gregoryi Fraser, 1924 (Odonata: Calopterygidae). Zootaxa, 4926 (2), 276 - 284. https: // doi. org / 10.11646 / zootaxa. 4926.2.7" type="journal article" year="2021">
Yang
<emphasis id="29AEEAA9E024D743FAFAEF63FA8EFC81" box="[1300,1348,835,861]" italics="true" pageId="17" pageNumber="226">et al</emphasis>
. (2021)
</bibRefCitation>
illustrate the habitat of
<taxonomicName id="DCDA4D38E024D743FE79EF47FDD1FC5C" authorityName="Fraser" authorityYear="1924" box="[407,539,871,896]" class="Insecta" family="Calopterygidae" genus="Mnais" kingdom="Animalia" order="Odonata" pageId="17" pageNumber="226" phylum="Arthropoda" rank="species" species="gregoryi">
<emphasis id="29AEEAA9E024D743FE79EF47FDD1FC5C" box="[407,539,871,896]" italics="true" pageId="17" pageNumber="226">M. gregoryi</emphasis>
</taxonomicName>
larva as a very shallow open stony montane stream with hydrophytes. They state that larvae are to be found clinging to these plants. The larvae of
<taxonomicName id="DCDA4D38E024D743FC92EFABFBD8FC79" authorityName="McLachlan in Selys" authorityYear="1873" box="[892,1042,907,933]" class="Insecta" family="Calopterygidae" genus="Mnais" kingdom="Animalia" order="Odonata" pageId="17" pageNumber="226" phylum="Arthropoda" rank="species" species="andersoni">
<emphasis id="29AEEAA9E024D743FC92EFABFBD8FC79" box="[892,1042,907,933]" italics="true" pageId="17" pageNumber="226">M. andersoni</emphasis>
</taxonomicName>
were found to be abundant among grass choking the head of a small open stream in a highly disturbed area at ca
<quantity id="DC229B5EE024D743FBF8EF8FFBA9FC15" box="[1046,1123,943,969]" metricMagnitude="3" metricUnit="m" metricValue="1.1" pageId="17" pageNumber="226" unit="m" value="1100.0">1100m</quantity>
near
<collectingRegion id="D91EF859E024D743FB4AEF8FFAE7FC15" box="[1188,1325,943,969]" country="Thailand" name="Chiang Mai" pageId="17" pageNumber="226">Chiang Mai</collectingRegion>
,
<collectingCountry id="63CD762BE024D743FAD7EF8FFA57FC15" box="[1337,1437,943,969]" name="Thailand" pageId="17" pageNumber="226">Thailand</collectingCountry>
(Orr, unpublished data).
</paragraph>
</subSubSection>
</treatment>
</document>

309
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<mods:title id="E64ECD64B671AC6719E50398C46DC786">A review of present knowledge of larvae of the Calopterygoidea (Zygoptera) of the Oriental realm, including keys to families and known genera</mods:title>
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<subSubSection id="53C06530E026D741FF79EB0CFE75F89A" box="[151,447,1836,1862]" pageId="19" pageNumber="228" type="nomenclature">
<paragraph id="1B6536BBE026D741FF79EB0CFE75F89A" blockId="19.[151,447,1836,1862]" box="[151,447,1836,1862]" pageId="19" pageNumber="228">
<heading id="402D81D7E026D741FF79EB0CFE75F89A" bold="true" box="[151,447,1836,1862]" fontSize="11" level="1" pageId="19" pageNumber="228" reason="1">
<emphasis id="29AEEAA9E026D741FF79EB0CFE75F89A" bold="true" box="[151,447,1836,1862]" pageId="19" pageNumber="228">
Family
<taxonomicName id="DCDA4D38E026D741FF19EB0CFE75F89A" ID-CoL="85T" authorityName="Cowley" authorityYear="1937" box="[247,447,1836,1862]" class="Insecta" family="Chlorocyphidae" kingdom="Animalia" order="Odonata" pageId="19" pageNumber="228" phylum="Arthropoda" rank="family">Chlorocyphidae</taxonomicName>
</emphasis>
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<paragraph id="1B6536BBE026D741FF79EB54FB06F826" blockId="19.[151,1436,1908,2043]" pageId="19" pageNumber="228">The larvae of just six of the 16 recognised Oriental genera have been described and illustrated, representing in total just 15 out of 110 species present in the region. A few additional species of uncertain identity are represented by photographs posted on the internet, but none provides useful or reliable information, although they do give some indication of the range of additional morphological variation we might expect within the family.</paragraph>
<paragraph id="1B6536BBE021D746FF29ECB7FAAEFC81" blockId="20.[151,1437,151,2013]" pageId="20" pageNumber="229">
The first account of a chorocyphid larvae was by
<bibRefCitation id="7F4B4B4AE021D746FD1AECB7FC5DFF6D" author="Fraser, F. C." box="[756,919,151,177]" pageId="20" pageNumber="229" pagination="197 - 198" refId="ref22755" refString="Fraser, F. C. (1919 a) The larva of Micromerus lineatus Burm. Records of the Indian Museum, 16, 197 - 198, pl. 23 excl. https: // doi. org / 10.26515 / rzsi / v 16 / i 2 / 1919 / 162807" type="journal article" year="1919">Fraser (1919a)</bibRefCitation>
who described
<taxonomicName id="DCDA4D38E021D746FBA6ECB7FA56FF6D" authority="(Burmeister)" baseAuthorityName="Burmeister" box="[1096,1436,151,177]" class="Insecta" family="Chlorocyphidae" genus="Libellago" kingdom="Animalia" order="Odonata" pageId="20" pageNumber="229" phylum="Arthropoda" rank="species" species="lineata">
<emphasis id="29AEEAA9E021D746FBA6ECB7FACDFF6D" box="[1096,1287,151,177]" italics="true" pageId="20" pageNumber="229">Libellago lineata</emphasis>
(Burmeister)
</taxonomicName>
in considerable detail (as
<taxonomicName id="DCDA4D38E021D746FE41EC9BFD59FF09" box="[431,659,187,213]" class="Insecta" family="Chlorocyphidae" genus="Micromerus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="20" pageNumber="229" phylum="Arthropoda" rank="species" species="lineatus">
<emphasis id="29AEEAA9E021D746FE41EC9BFD59FF09" box="[431,659,187,213]" italics="true" pageId="20" pageNumber="229">Micromerus lineatus</emphasis>
</taxonomicName>
), providing a reasonably accurate illustration of the habitus and partial detail of the mask. He omitted the vestigial central gill (epiproct), and other details, but stated that the exuviae were very common and that the larvae cling to submerged roots, twigs and other debris in swift water and rarely to aquatic vegetation. He asserted “they are purely rectal breathers and if the larva be viewed in muddy water, strong currents of particles are seen issuing to and from the rectum”. The following year
<bibRefCitation id="7F4B4B4AE021D746FBF5ED6BFB0CFEB9" author="Laidlaw, F. F." box="[1051,1222,331,357]" pageId="20" pageNumber="229" pagination="185 - 187" refId="ref24408" refString="Laidlaw, F. F. (1920) Notes on some interesting larvae of dragonflies (Odonata) in the collection of the Indian Museum. Records of the Indian Museum, 19, 185 - 187. https: // doi. org / 10.26515 / rzsi / v 19 / i 5 / 1920 / 163534" type="journal article" year="1920">Laidlaw (1920)</bibRefCitation>
described the larva of
<taxonomicName id="DCDA4D38E021D746FF56ED4FFDE8FE55" authority="(Selys)" baseAuthorityName="Selys" box="[184,546,367,393]" class="Insecta" family="Chlorocyphidae" genus="Paracypha" kingdom="Animalia" order="Odonata" pageId="20" pageNumber="229" phylum="Arthropoda" rank="species" species="unimaculata">
<emphasis id="29AEEAA9E021D746FF56ED4FFE0DFE55" box="[184,455,367,393]" italics="true" pageId="20" pageNumber="229">Paracypha unimaculata</emphasis>
(Selys)
</taxonomicName>
, (as
<taxonomicName id="DCDA4D38E021D746FDB5ED4FFCBEFE55" authorityName="" baseAuthorityName="Kumar" baseAuthorityYear="1973" box="[603,884,367,393]" class="Insecta" family="Chlorocyphidae" genus="Rhinocypha" kingdom="Animalia" order="Odonata" pageId="20" pageNumber="229" phylum="Arthropoda" rank="species" species="unimaculata">
<emphasis id="29AEEAA9E021D746FDB5ED4FFCBEFE55" box="[603,884,367,393]" italics="true" pageId="20" pageNumber="229">Rhinocypha unimaculata</emphasis>
</taxonomicName>
) concluding that it showed the affinity between
<taxonomicName id="DCDA4D38E021D746FF79EDB3FECBFE71" box="[151,257,403,429]" class="Insecta" family="Chlorocyphidae" genus="Libellago" kingdom="Animalia" order="Odonata" pageId="20" pageNumber="229" phylum="Arthropoda" rank="genus">
<emphasis id="29AEEAA9E021D746FF79EDB3FECBFE71" box="[151,257,403,429]" italics="true" pageId="20" pageNumber="229">Libellago</emphasis>
</taxonomicName>
and
<taxonomicName id="DCDA4D38E021D746FED7EDB3FE77FE71" authorityName="Rambur" authorityYear="1842" box="[313,445,403,429]" class="Insecta" family="Chlorocyphidae" genus="Rhinocypha" kingdom="Animalia" order="Odonata" pageId="20" pageNumber="229" phylum="Arthropoda" rank="genus">
<emphasis id="29AEEAA9E021D746FED7EDB3FE77FE71" box="[313,445,403,429]" italics="true" pageId="20" pageNumber="229">Rhinocypha</emphasis>
</taxonomicName>
(sensu lato), and correctly surmised that the African members of the family would have similar larvae.
<bibRefCitation id="7F4B4B4AE021D746FED0ED97FE1EFE0D" author="Fraser, F. C." box="[318,468,439,465]" pageId="20" pageNumber="229" pagination="13 - 14" refId="ref22861" refString="Fraser, F. C. (1920) Description of a Rhinocyphine larva from Shillong. Memoirs of the department of agriculture in India, 7, 13 - 14, 1 pl. excl." type="journal article" year="1920">Fraser (1920)</bibRefCitation>
reached the same conclusion from the study of
<taxonomicName id="DCDA4D38E021D746FC04ED97FAFAFE0D" authority="Selys" authorityName="Selys" authorityYear="1879" box="[1002,1328,439,465]" class="Insecta" family="Chlorocyphidae" genus="Rhinocypha" kingdom="Animalia" order="Odonata" pageId="20" pageNumber="229" phylum="Arthropoda" rank="species" species="ignipennis">
<emphasis id="29AEEAA9E021D746FC04ED97FB20FE0C" box="[1002,1258,439,465]" italics="true" pageId="20" pageNumber="229">Rhinocypha ignipennis</emphasis>
Selys
</taxonomicName>
, although his illustration of the habitus is very rough and lacks wing buds, which might be expected even in quite young larvae a few millimetres long, and he states that the specimen has no caudal gills or any evidence of them having been present, which is surely also an error as these structures are known develop very early (see
<bibRefCitation id="7F4B4B4AE021D746FBCCEE03FB44FDE1" author="Ng, C. Y." box="[1058,1166,547,573]" pageId="20" pageNumber="229" refId="ref25224" refString="Ng, C. Y. (2024 c) Freshwater Macroinvertebrates in Hong Kong: Odonata: Rhinocypha perforata. Available from: https: // www. biosch. hku. hk / ecology / staffhp / dd / macroinvertebrates / Odonata / Chlorocyphidae / Rhinocypha _ perforata. html (accessed 16 May 2024)" type="url" year="2024">Ng 2024c</bibRefCitation>
for a depiction of a very young larva of
<emphasis id="29AEEAA9E021D746FEA8EE67FE29FDBC" box="[326,483,583,609]" italics="true" pageId="20" pageNumber="229">
<taxonomicName id="DCDA4D38E021D746FEA8EE67FE0CFDBD" authorityName="Fraser" authorityYear="1949" box="[326,454,583,609]" class="Insecta" family="Chlorocyphidae" genus="Heliocypha" kingdom="Animalia" order="Odonata" pageId="20" pageNumber="229" phylum="Arthropoda" rank="genus">Heliocypha</taxonomicName>
p.
</emphasis>
<taxonomicName id="DCDA4D38E021D746FE02EE67FD19FDBD" authority="Percheron" authorityName="Percheron" authorityYear="1835" box="[492,723,583,609]" class="Insecta" family="Chlorocyphidae" genus="Heliocypha" kingdom="Animalia" order="Odonata" pageId="20" pageNumber="229" phylum="Arthropoda" rank="species" species="perforata">
<emphasis id="29AEEAA9E021D746FE02EE67FD9FFDBD" box="[492,597,583,609]" italics="true" pageId="20" pageNumber="229">perforata</emphasis>
Percheron
</taxonomicName>
). Subsequently
<bibRefCitation id="7F4B4B4AE021D746FC6BEE67FBD7FDBD" author="Fraser, F. C." box="[901,1053,583,609]" pageId="20" pageNumber="229" pagination="683 - 691" refId="ref22898" refString="Fraser, F. C. (1928) Indian dragonflies, Part XXX. Journal of the Bombay Natural History Society, 32, 683 - 691, pls. 1 - 3, excl." type="journal article" year="1928">Fraser (1928)</bibRefCitation>
provided an excellent drawing of the mature habitus of this species and
<taxonomicName id="DCDA4D38E021D746FDD7EE4BFC41FD59" authority="(Hagen)" baseAuthorityName="Hagen" box="[569,907,619,645]" class="Insecta" family="Chlorocyphidae" genus="Heliocypha" kingdom="Animalia" order="Odonata" pageId="20" pageNumber="229" phylum="Arthropoda" rank="species" species="bisignata">
<emphasis id="29AEEAA9E021D746FDD7EE4BFCECFD59" box="[569,806,619,645]" italics="true" pageId="20" pageNumber="229">Heliocypha bisignata</emphasis>
(Hagen)
</taxonomicName>
, as well as a new depiction of
<taxonomicName id="DCDA4D38E021D746FB36EE4BFA5FFD59" box="[1240,1429,619,645]" class="Insecta" family="Chlorocyphidae" genus="Libellago" kingdom="Animalia" order="Odonata" pageId="20" pageNumber="229" phylum="Arthropoda" rank="species" species="lineata">
<emphasis id="29AEEAA9E021D746FB36EE4BFA5FFD59" box="[1240,1429,619,645]" italics="true" pageId="20" pageNumber="229">Libellago lineata</emphasis>
</taxonomicName>
, again unaccountably lacking the vestigial central gill which is shown in the other species, and commented further on the habits of the family: “Very sluggish in habits and cryptic in colouring, they are difficult creatures to find. They cling to roots or broken-up debris in slow-running streams or at the bottom of pools in swifter running ones. I have found the easiest way to take them is to dredge out a quantity of such debris and spread it on the foreshore exposed to the sun...”. Under these circumstances the larvae stir as they try to escape the heat and the drying plant matter and can be easily seen. They may exhibit thanatosis when originally scooped out, making them very hard to detect.
</paragraph>
<paragraph id="1B6536BBE021D746FF29EF47FD3FFA39" blockId="20.[151,1437,151,2013]" pageId="20" pageNumber="229">
Chlorocyphid larvae are unmistakable by virtue of having two long spikes representing the lateral caudal gills, with a vestigial dorsal median gill (also termed epiproct) in the form of a short conical spine; all known species also have an unusual arrow-head shaped head with a prominent tubercle on the subocular lobe and a first antennal segment (scape) is at least 40% the total length of the antenna (
<figureCitation id="83E12A3EE021D746FC69EFF3FC39FC31" box="[903,1011,979,1005]" captionStart="FIGURES 1–7. 1" captionStartId="3.[151,275,1843,1869]" captionTargetBox="[252,1341,160,1825]" captionTargetId="figure-15@3.[243,1344,149,1836]" captionTargetPageId="3" captionText="FIGURES 1–7. 1. Ventral view of abdomen of Euphaea sp. showing abdominal gills and saccoid gills (original drawing). 2. Head and antennae of Calopterygidae (Neurobasis chinensis) (original drawing). 3. Head and antennae of Chlorocyphidae (Heliocypha biseriata) (original drawing). 4. Antenna of Podolestes orientalis Selys, 1852, scape arrowed (from Choong &amp; Orr 2010). 5. Mask of Calopterygidae (Calopteryx japonica Selys redrawn from Miyakawa 1983). 6. Mask of Chlorocyphidae (Libellago hyalina, from Orr et al. 2024). 7. Habitus of L. hyalina, with detail of right caudal spike (from Orr et al. 2024)." figureDoi="http://doi.org/10.5281/zenodo.13618321" httpUri="https://zenodo.org/record/13618321/files/figure.png" pageId="20" pageNumber="229">Figs 3, 7</figureCitation>
). The labrum of chlorocyphids is of an unusual form (
<figureCitation id="83E12A3EE021D746FE8DEFD7FE7CFBCD" box="[355,438,1015,1041]" captionStart="FIGURES 35–41. 35" captionStartId="8.[151,264,1776,1800]" captionTargetBox="[257,1332,159,1764]" captionTargetId="figure-21@8.[253,1335,153,1771]" captionTargetPageId="8" captionText="FIGURES 35–41. 35. Archineura incarnata, prementum (dorsal view, drawn from Yang et al. 2022). 36. Vestalis gracilis, prementum (dorsal view) and right antenna with supplementary antennomere between pedicel and scape (a) arrowed (redrawn from Rattanachan et al. 2022). 37. Vestalaria venusta: prementum (dorsal view) and right antenna (drawn from Wang et al. 2017). 38. Caliphaea angka: prementum (dorsal view) and right antenna (drawn from Yang et al. 2021). 39. Heliocypha biseriata, frontal view of labrum showing dorsal and ventral sections and ‘moustache’ of long, heavy setae on the latter (from Orr et al. 2024). 40. Heliocypha biseriata: details of labial palp in relation to prementum (a) dorsal view—base of movable hook dashed to reveal palpal lobe beneath and open base of premental median cleft arrowed, (b) frontal view, (c) inner view (from Orr et al. 2024). 41. Heliocypha biseriata, detail of fore tarsus with pectinate setae magnified (from Orr et al. 2024)." figureDoi="http://doi.org/10.5281/zenodo.13618335" httpUri="https://zenodo.org/record/13618335/files/figure.png" pageId="20" pageNumber="229">Fig. 39</figureCitation>
) with the anterior extremity folded under and forming with the upper posterior part an elongate, rounded boat-like cavity into which the tips of the mandibles fit. The lower section bears a ‘moustache’ consisting of a single row of long, thick setae (Orr
<emphasis id="29AEEAA9E021D746FD3AE81FFCC7FB85" box="[724,781,1087,1113]" italics="true" pageId="20" pageNumber="229">et al.</emphasis>
2024). The palpal lobe of the mask (
<figureCitation id="83E12A3EE021D746FB42E81FFA8FFB85" box="[1196,1349,1087,1113]" captionStart="FIGURES 35–41. 35" captionStartId="8.[151,264,1776,1800]" captionTargetBox="[257,1332,159,1764]" captionTargetId="figure-21@8.[253,1335,153,1771]" captionTargetPageId="8" captionText="FIGURES 35–41. 35. Archineura incarnata, prementum (dorsal view, drawn from Yang et al. 2022). 36. Vestalis gracilis, prementum (dorsal view) and right antenna with supplementary antennomere between pedicel and scape (a) arrowed (redrawn from Rattanachan et al. 2022). 37. Vestalaria venusta: prementum (dorsal view) and right antenna (drawn from Wang et al. 2017). 38. Caliphaea angka: prementum (dorsal view) and right antenna (drawn from Yang et al. 2021). 39. Heliocypha biseriata, frontal view of labrum showing dorsal and ventral sections and ‘moustache’ of long, heavy setae on the latter (from Orr et al. 2024). 40. Heliocypha biseriata: details of labial palp in relation to prementum (a) dorsal view—base of movable hook dashed to reveal palpal lobe beneath and open base of premental median cleft arrowed, (b) frontal view, (c) inner view (from Orr et al. 2024). 41. Heliocypha biseriata, detail of fore tarsus with pectinate setae magnified (from Orr et al. 2024)." figureDoi="http://doi.org/10.5281/zenodo.13618335" httpUri="https://zenodo.org/record/13618335/files/figure.png" pageId="20" pageNumber="229">Figs 40a, b, c</figureCitation>
) is also unusual being deeply divided into upper and lower branches each with a shorter process originating about the midpoint. This was noted by
<bibRefCitation id="7F4B4B4AE021D746FE31E8A7FD4DFB7D" author="Fraser, F. C." box="[479,647,1159,1185]" pageId="20" pageNumber="229" pagination="197 - 198" refId="ref22755" refString="Fraser, F. C. (1919 a) The larva of Micromerus lineatus Burm. Records of the Indian Museum, 16, 197 - 198, pl. 23 excl. https: // doi. org / 10.26515 / rzsi / v 16 / i 2 / 1919 / 162807" type="journal article" year="1919">Fraser (1919a)</bibRefCitation>
. It has been suggested that the spur on the upper process is secondary, with the other branches representing a highly modified form of the three terminal processes normally found on the palpal lobe of other
<taxonomicName id="DCDA4D38E021D746FE44E8EFFDA9FB35" box="[426,611,1231,1257]" pageId="20" pageNumber="210" rank="superFamily" superFamily="Calopterygoidea">Calopterygoidea</taxonomicName>
larvae (Orr
<emphasis id="29AEEAA9E021D746FD1FE8EFFCEEFB35" box="[753,804,1231,1257]" italics="true" pageId="20" pageNumber="229">et al</emphasis>
. 2024). The prementum is moderately long and narrow with a modest median cleft that is frequently fused; the median lobe always has a clearly serrated margin (
<figureCitation id="83E12A3EE021D746FAA3E8D3FA5DFAD1" box="[1357,1431,1267,1293]" captionStart="FIGURES 1–7. 1" captionStartId="3.[151,275,1843,1869]" captionTargetBox="[252,1341,160,1825]" captionTargetId="figure-15@3.[243,1344,149,1836]" captionTargetPageId="3" captionText="FIGURES 1–7. 1. Ventral view of abdomen of Euphaea sp. showing abdominal gills and saccoid gills (original drawing). 2. Head and antennae of Calopterygidae (Neurobasis chinensis) (original drawing). 3. Head and antennae of Chlorocyphidae (Heliocypha biseriata) (original drawing). 4. Antenna of Podolestes orientalis Selys, 1852, scape arrowed (from Choong &amp; Orr 2010). 5. Mask of Calopterygidae (Calopteryx japonica Selys redrawn from Miyakawa 1983). 6. Mask of Chlorocyphidae (Libellago hyalina, from Orr et al. 2024). 7. Habitus of L. hyalina, with detail of right caudal spike (from Orr et al. 2024)." figureDoi="http://doi.org/10.5281/zenodo.13618321" httpUri="https://zenodo.org/record/13618321/files/figure.png" pageId="20" pageNumber="229">Figs 6</figureCitation>
,
<figureCitation id="83E12A3EE021D746FF79E937FF0EFAED" box="[151,196,1303,1329]" captionStart="FIGURES 35–41. 35" captionStartId="8.[151,264,1776,1800]" captionTargetBox="[257,1332,159,1764]" captionTargetId="figure-21@8.[253,1335,153,1771]" captionTargetPageId="8" captionText="FIGURES 35–41. 35. Archineura incarnata, prementum (dorsal view, drawn from Yang et al. 2022). 36. Vestalis gracilis, prementum (dorsal view) and right antenna with supplementary antennomere between pedicel and scape (a) arrowed (redrawn from Rattanachan et al. 2022). 37. Vestalaria venusta: prementum (dorsal view) and right antenna (drawn from Wang et al. 2017). 38. Caliphaea angka: prementum (dorsal view) and right antenna (drawn from Yang et al. 2021). 39. Heliocypha biseriata, frontal view of labrum showing dorsal and ventral sections and ‘moustache’ of long, heavy setae on the latter (from Orr et al. 2024). 40. Heliocypha biseriata: details of labial palp in relation to prementum (a) dorsal view—base of movable hook dashed to reveal palpal lobe beneath and open base of premental median cleft arrowed, (b) frontal view, (c) inner view (from Orr et al. 2024). 41. Heliocypha biseriata, detail of fore tarsus with pectinate setae magnified (from Orr et al. 2024)." figureDoi="http://doi.org/10.5281/zenodo.13618335" httpUri="https://zenodo.org/record/13618335/files/figure.png" pageId="20" pageNumber="229">40a</figureCitation>
). All species examined bear two rows of pectinate setae on the underside of the tarsi (
<figureCitation id="83E12A3EE021D746FB6CE937FB1EFAED" box="[1154,1236,1303,1329]" captionStart="FIGURES 35–41. 35" captionStartId="8.[151,264,1776,1800]" captionTargetBox="[257,1332,159,1764]" captionTargetId="figure-21@8.[253,1335,153,1771]" captionTargetPageId="8" captionText="FIGURES 35–41. 35. Archineura incarnata, prementum (dorsal view, drawn from Yang et al. 2022). 36. Vestalis gracilis, prementum (dorsal view) and right antenna with supplementary antennomere between pedicel and scape (a) arrowed (redrawn from Rattanachan et al. 2022). 37. Vestalaria venusta: prementum (dorsal view) and right antenna (drawn from Wang et al. 2017). 38. Caliphaea angka: prementum (dorsal view) and right antenna (drawn from Yang et al. 2021). 39. Heliocypha biseriata, frontal view of labrum showing dorsal and ventral sections and ‘moustache’ of long, heavy setae on the latter (from Orr et al. 2024). 40. Heliocypha biseriata: details of labial palp in relation to prementum (a) dorsal view—base of movable hook dashed to reveal palpal lobe beneath and open base of premental median cleft arrowed, (b) frontal view, (c) inner view (from Orr et al. 2024). 41. Heliocypha biseriata, detail of fore tarsus with pectinate setae magnified (from Orr et al. 2024)." figureDoi="http://doi.org/10.5281/zenodo.13618335" httpUri="https://zenodo.org/record/13618335/files/figure.png" pageId="20" pageNumber="229">Fig. 41</figureCitation>
) that require high magnification to resolve (
<bibRefCitation id="7F4B4B4AE021D746FE5DE91BFD8EFA89" author="Kumar, A." box="[435,580,1339,1365]" pageId="20" pageNumber="229" pagination="83 - 118" refId="ref24196" refString="Kumar, A. (1973) Descriptions of the last instar larvae of Odonata from the Dehra Dun Valley (India), with notes on biology I. (suborder Zygoptera). Oriental Insects, 7, 83 - 118. https: // doi. org / 10.1080 / 00305316.1973.10434207" type="journal article" year="1973">Kumar 1973</bibRefCitation>
, Kumar &amp; Prasad 1977, Orr
<emphasis id="29AEEAA9E021D746FC6CE91BFC78FA89" box="[898,946,1339,1365]" italics="true" pageId="20" pageNumber="229">et al</emphasis>
. 2024). Finally, all chlorocyphids examined bear a dense dorsal pile of long, very fine setae that entraps particles of detritus and silt and presumably aids in camouflage. This is only clearly visible in submerged living specimens, as even when wet-preserved the filaments collapse into a dense tangled mat covering the body surface as well as the caudal spikes (Orr
<emphasis id="29AEEAA9E021D746FB45E987FB29FA1D" box="[1195,1251,1447,1473]" italics="true" pageId="20" pageNumber="229">et al.</emphasis>
2024). All these features may represent autapomorphies for the family.
</paragraph>
<paragraph id="1B6536BBE021D746FF29E9CFFC6FF801" blockId="20.[151,1437,151,2013]" pageId="20" pageNumber="229">
Despite being the most speciose of all Oriental calopterygoids, with the highest number of recognised genera, the family is remarkable for the lack of inter-generic variation among its larvae. Apart from characters already mentioned, the body is squat, although a little more elongate in the comparatively large
<taxonomicName id="DCDA4D38E021D746FB62EA17FA56F98D" box="[1164,1436,1591,1617]" class="Insecta" family="Chlorocyphidae" genus="Paracypha" kingdom="Animalia" order="Odonata" pageId="20" pageNumber="229" phylum="Arthropoda" rank="species" species="unimaculata">
<emphasis id="29AEEAA9E021D746FB62EA17FA56F98D" box="[1164,1436,1591,1617]" italics="true" pageId="20" pageNumber="229">Paracypha unimaculata</emphasis>
</taxonomicName>
(figured by
<bibRefCitation id="7F4B4B4AE021D746FEF4EA7BFE60F9A9" author="Kumar, A." box="[282,426,1627,1653]" pageId="20" pageNumber="229" pagination="83 - 118" refId="ref24196" refString="Kumar, A. (1973) Descriptions of the last instar larvae of Odonata from the Dehra Dun Valley (India), with notes on biology I. (suborder Zygoptera). Oriental Insects, 7, 83 - 118. https: // doi. org / 10.1080 / 00305316.1973.10434207" type="journal article" year="1973">Kumar 1973</bibRefCitation>
, as
<taxonomicName id="DCDA4D38E021D746FE3CEA7BFD20F9A9" authorityName="" baseAuthorityName="Kumar" baseAuthorityYear="1973" box="[466,746,1627,1653]" class="Insecta" family="Chlorocyphidae" genus="Rhinocypha" kingdom="Animalia" order="Odonata" pageId="20" pageNumber="229" phylum="Arthropoda" rank="species" species="unimaculata">
<emphasis id="29AEEAA9E021D746FE3CEA7BFD20F9A9" box="[466,746,1627,1653]" italics="true" pageId="20" pageNumber="229">Rhinocypha unimaculata</emphasis>
</taxonomicName>
).
<bibRefCitation id="7F4B4B4AE021D746FCEFEA7BFBD3F9A9" author="Kumar, A. &amp; Prasad, M." box="[769,1049,1627,1653]" pageId="20" pageNumber="229" pagination="547 - 554" refId="ref24362" refString="Kumar, A. &amp; Prasad, M. (1977 c) On the larvae of Rhinocypha (Odonata: Cholorocyphidae) from Garhwal hills. Oriental Insects, 11, 547 - 554. https: // doi. org / 10.1080 / 00305316.1977.11090927" type="journal article" year="1977">Kumar &amp; Prasad (1977c)</bibRefCitation>
use slight differences in the palpal lobe to discriminate between taxa, as well as differences in size, and the spines on the margins of the prementum which are unusually well developed in
<taxonomicName id="DCDA4D38E021D746FDA6EA83FD25F961" authorityName="" baseAuthorityName="Kumar" baseAuthorityYear="1973" box="[584,751,1699,1725]" class="Insecta" family="Chlorocyphidae" genus="Rhinocypha" kingdom="Animalia" order="Odonata" pageId="20" pageNumber="229" phylum="Arthropoda" rank="species" species="unimaculata">
<emphasis id="29AEEAA9E021D746FDA6EA83FD25F961" box="[584,751,1699,1725]" italics="true" pageId="20" pageNumber="229">P. unimaculata</emphasis>
</taxonomicName>
. All species also have some spines on the gena below the eye, but these show some variation which may manifest at the intra-generic level, as is also the case with the relative length and shape of the prementum (Orr
<emphasis id="29AEEAA9E021D746FD82EACBFD6CF8D9" box="[620,678,1771,1797]" italics="true" pageId="20" pageNumber="229">et al.</emphasis>
2024). Variation in the lateral caudal gills is a potentially difficult character, in that if lost early in development they may regenerate, but not to their normal extent (as evidenced by the depiction of a specimen with lateral gills of unequal lengths in
<taxonomicName id="DCDA4D38E021D746FC95EB13FB1AF891" authority="(Kumar 1973))" authorityName="" baseAuthorityName="Kumar" baseAuthorityYear="1973" box="[891,1232,1843,1869]" class="Insecta" family="Chlorocyphidae" genus="Rhinocypha" kingdom="Animalia" order="Odonata" pageId="20" pageNumber="229" phylum="Arthropoda" rank="species" species="unimaculata">
<emphasis id="29AEEAA9E021D746FC95EB13FBEBF891" box="[891,1057,1843,1869]" italics="true" pageId="20" pageNumber="229">P. unimaculata</emphasis>
(
<bibRefCitation id="7F4B4B4AE021D746FBC1EB13FB77F891" author="Kumar, A." box="[1071,1213,1843,1869]" pageId="20" pageNumber="229" pagination="83 - 118" refId="ref24196" refString="Kumar, A. (1973) Descriptions of the last instar larvae of Odonata from the Dehra Dun Valley (India), with notes on biology I. (suborder Zygoptera). Oriental Insects, 7, 83 - 118. https: // doi. org / 10.1080 / 00305316.1973.10434207" type="journal article" year="1973">Kumar 1973</bibRefCitation>
))
</taxonomicName>
and
<bibRefCitation id="7F4B4B4AE021D746FAEBEB13FA56F891" author="Fraser, F. C." box="[1285,1436,1843,1869]" pageId="20" pageNumber="229" pagination="683 - 691" refId="ref22898" refString="Fraser, F. C. (1928) Indian dragonflies, Part XXX. Journal of the Bombay Natural History Society, 32, 683 - 691, pls. 1 - 3, excl." type="journal article" year="1928">Fraser (1928)</bibRefCitation>
states that this phenomenon is common, stating “autonomy [sic = autotomy] of the caudal gills is accomplished with the greatest ease, so it is difficult to obtain one with all intact; probably these are more for defence than respiratory use, and they certainly carry on quite well without them”. Nevertheless where the structures are of equal length and consistent within a species they can probably be assumed to be intact.
</paragraph>
<paragraph id="1B6536BBE020D747FF29ECB7FC6DFDBD" blockId="21.[151,1437,151,933]" pageId="21" pageNumber="230">
Comparing four species representing the genera
<taxonomicName id="DCDA4D38E020D747FD1BECB7FCB6FF6D" authorityName="Laidlaw" authorityYear="1950" box="[757,892,151,177]" class="Insecta" family="Chlorocyphidae" genus="Aristocypha" kingdom="Animalia" order="Odonata" pageId="21" pageNumber="230" phylum="Arthropoda" rank="genus">
<emphasis id="29AEEAA9E020D747FD1BECB7FCB6FF6D" box="[757,892,151,177]" italics="true" pageId="21" pageNumber="230">Aristocypha</emphasis>
</taxonomicName>
,
<taxonomicName id="DCDA4D38E020D747FC63ECB7FBC4FF6D" authorityName="Fraser" authorityYear="1949" box="[909,1038,151,177]" class="Insecta" family="Chlorocyphidae" genus="Heliocypha" kingdom="Animalia" order="Odonata" pageId="21" pageNumber="230" phylum="Arthropoda" rank="genus">
<emphasis id="29AEEAA9E020D747FC63ECB7FBC4FF6D" box="[909,1038,151,177]" italics="true" pageId="21" pageNumber="230">Heliocypha</emphasis>
</taxonomicName>
,
<taxonomicName id="DCDA4D38E020D747FBF0ECB7FB6DFF6D" authorityName="Laidlaw" authorityYear="1950" box="[1054,1191,151,177]" class="Insecta" family="Chlorocyphidae" genus="Sundacypha" kingdom="Animalia" order="Odonata" pageId="21" pageNumber="230" phylum="Arthropoda" rank="genus">
<emphasis id="29AEEAA9E020D747FBF0ECB7FB6DFF6D" box="[1054,1191,151,177]" italics="true" pageId="21" pageNumber="230">Sundacypha</emphasis>
</taxonomicName>
and
<taxonomicName id="DCDA4D38E020D747FB0DECB7FECCFF09" authority="Orr et al. (2024)" authorityName="Orr" authorityYear="2024" class="Insecta" family="Chlorocyphidae" genus="Libellago" kingdom="Animalia" order="Odonata" pageId="21" pageNumber="230" phylum="Arthropoda" rank="genus">
<emphasis id="29AEEAA9E020D747FB0DECB7FA84FF6D" box="[1251,1358,151,177]" italics="true" pageId="21" pageNumber="230">Libellago</emphasis>
Orr
<emphasis id="29AEEAA9E020D747FA66ECB7FF67FF09" italics="true" pageId="21" pageNumber="230">et al</emphasis>
. (2024)
</taxonomicName>
concluded that the species studied could be separated readily based on characters such as the form of the caudal spikes, the subocular spines on the genae and the form of the prementum, relative antennal length and other small but consistent characters. However, taking into account these results with those of
<bibRefCitation id="7F4B4B4AE020D747FB94ED23FA5DFEC1" author="Kumar, A. &amp; Prasad, M." box="[1146,1431,259,285]" pageId="21" pageNumber="230" pagination="547 - 554" refId="ref24362" refString="Kumar, A. &amp; Prasad, M. (1977 c) On the larvae of Rhinocypha (Odonata: Cholorocyphidae) from Garhwal hills. Oriental Insects, 11, 547 - 554. https: // doi. org / 10.1080 / 00305316.1977.11090927" type="journal article" year="1977">Kumar &amp; Prasad (1977c)</bibRefCitation>
, definitive generic differences could not always be established with confidence, even in the limited range of known species, with intra-generic variation blurring supposed distinctions. It appears that the caudal spikes are slightly broader and taper to a point more abruptly from near the apex in
<taxonomicName id="DCDA4D38E020D747FC66ED4FFBC5FE55" authorityName="Laidlaw" authorityYear="1950" box="[904,1039,367,393]" class="Insecta" family="Chlorocyphidae" genus="Aristocypha" kingdom="Animalia" order="Odonata" pageId="21" pageNumber="230" phylum="Arthropoda" rank="genus">
<emphasis id="29AEEAA9E020D747FC66ED4FFBC5FE55" box="[904,1039,367,393]" italics="true" pageId="21" pageNumber="230">Aristocypha</emphasis>
</taxonomicName>
than
<taxonomicName id="DCDA4D38E020D747FBBCED4FFB19FE55" authorityName="Fraser" authorityYear="1949" box="[1106,1235,367,393]" class="Insecta" family="Chlorocyphidae" genus="Heliocypha" kingdom="Animalia" order="Odonata" pageId="21" pageNumber="230" phylum="Arthropoda" rank="genus">
<emphasis id="29AEEAA9E020D747FBBCED4FFB19FE55" box="[1106,1235,367,393]" italics="true" pageId="21" pageNumber="230">Heliocypha</emphasis>
</taxonomicName>
but more species need to be examined to confirm this. The caudal spikes of known
<taxonomicName id="DCDA4D38E020D747FC9BEDB3FC2AFE71" box="[885,992,403,429]" class="Insecta" family="Chlorocyphidae" genus="Libellago" kingdom="Animalia" order="Odonata" pageId="21" pageNumber="230" phylum="Arthropoda" rank="genus">
<emphasis id="29AEEAA9E020D747FC9BEDB3FC2AFE71" box="[885,992,403,429]" italics="true" pageId="21" pageNumber="230">Libellago</emphasis>
</taxonomicName>
and
<taxonomicName id="DCDA4D38E020D747FBF8EDB3FB55FE71" authorityName="Laidlaw" authorityYear="1950" box="[1046,1183,403,429]" class="Insecta" family="Chlorocyphidae" genus="Sundacypha" kingdom="Animalia" order="Odonata" pageId="21" pageNumber="230" phylum="Arthropoda" rank="genus">
<emphasis id="29AEEAA9E020D747FBF8EDB3FB55FE71" box="[1046,1183,403,429]" italics="true" pageId="21" pageNumber="230">Sundacypha</emphasis>
</taxonomicName>
are also distinctive. In one of the best depictions of chlorocyphid larvae
<bibRefCitation id="7F4B4B4AE020D747FD2CED97FC92FE0D" author="Ishida, K." box="[706,856,439,465]" pageId="21" pageNumber="230" refId="ref23597" refString="Ishida, K. (1996) Monograph of Odonata larvae in Japan. Hokkaido University Press, Sapporo, 446 pp. [in Japanese]" type="book" year="1996">Ishida (1996)</bibRefCitation>
illustrates unusually short, broad, somewhat foliate caudal spikes in
<taxonomicName id="DCDA4D38E020D747FEBDEDFBFD15FE29" authority="Oguma" authorityName="Oguma" box="[339,735,475,501]" class="Insecta" family="Chlorocyphidae" genus="Rhinocypha" kingdom="Animalia" order="Odonata" pageId="21" pageNumber="230" phylum="Arthropoda" rank="species" species="ogasawarensis">
<emphasis id="29AEEAA9E020D747FEBDEDFBFD4FFE28" box="[339,645,475,501]" italics="true" pageId="21" pageNumber="230">Rhinocypha ogasawarensis</emphasis>
Oguma
</taxonomicName>
and
<taxonomicName id="DCDA4D38E020D747FCF8EDFBFB82FE29" authority="Asahina" authorityName="Asahina" box="[790,1096,475,501]" class="Insecta" family="Chlorocyphidae" genus="Rhinocypha" kingdom="Animalia" order="Odonata" pageId="21" pageNumber="230" phylum="Arthropoda" rank="species" species="uenoi">
<emphasis id="29AEEAA9E020D747FCF8EDFBFC28FE28" box="[790,994,475,501]" italics="true" pageId="21" pageNumber="230">Rhinocypha uenoi</emphasis>
Asahina
</taxonomicName>
, but this cannot be considered a generic character as
<taxonomicName id="DCDA4D38E020D747FE63EDDFFDEBFDC4" authorityName="Selys" authorityYear="1879" box="[397,545,511,536]" class="Insecta" family="Chlorocyphidae" genus="Rhinocypha" kingdom="Animalia" order="Odonata" pageId="21" pageNumber="230" phylum="Arthropoda" rank="species" species="ignipennis">
<emphasis id="29AEEAA9E020D747FE63EDDFFDEBFDC4" box="[397,545,511,536]" italics="true" pageId="21" pageNumber="230">R. ignipennis</emphasis>
</taxonomicName>
has normal, long lateral spikes (
<bibRefCitation id="7F4B4B4AE020D747FC66EDDFFBD8FDC5" author="Fraser, F. C." box="[904,1042,511,537]" pageId="21" pageNumber="230" pagination="683 - 691" refId="ref22898" refString="Fraser, F. C. (1928) Indian dragonflies, Part XXX. Journal of the Bombay Natural History Society, 32, 683 - 691, pls. 1 - 3, excl." type="journal article" year="1928">Fraser 1928</bibRefCitation>
). However it must be noted that the generic classification of
<taxonomicName id="DCDA4D38E020D747FE43EE03FDF9FDE1" authorityName="Rambur" authorityYear="1842" box="[429,563,547,573]" class="Insecta" family="Chlorocyphidae" genus="Rhinocypha" kingdom="Animalia" order="Odonata" pageId="21" pageNumber="230" phylum="Arthropoda" rank="genus">
<emphasis id="29AEEAA9E020D747FE43EE03FDF9FDE1" box="[429,563,547,573]" italics="true" pageId="21" pageNumber="230">Rhinocypha</emphasis>
</taxonomicName>
and allied genera is unsettled (
<bibRefCitation id="7F4B4B4AE020D747FC7BEE03FBBDFDE1" author="Dijkstra, K-D. B. &amp; Kalkman, V. J. &amp; Dow, R. A. &amp; Stokvis, F. R. &amp; van Tol, J." box="[917,1143,547,573]" pageId="21" pageNumber="230" pagination="68 - 96" refId="ref22480" refString="Dijkstra, K-D. B., Kalkman, V. J., Dow, R. A., Stokvis, F. R. &amp; van Tol, J. (2014) Redefining the damselfly families: a comprehensive molecular phylogeny of Zygoptera (Odonata). Systematic entomology, 39, 68 - 96. https: // doi. org / 10.1111 / syen. 12035" type="journal article" year="2014">
Dijkstra
<emphasis id="29AEEAA9E020D747FC17EE03FBE6FDE1" box="[1017,1068,547,573]" italics="true" pageId="21" pageNumber="230">et al</emphasis>
. 2014
</bibRefCitation>
), and this non-conformity may reflect a higher level phylogenetic difference between these taxa.
</paragraph>
<paragraph id="1B6536BBE020D747FF29EE4BFE1FFC81" blockId="21.[151,1437,151,933]" pageId="21" pageNumber="230">
Despite these difficulties, over broad swathes of the Asian mainland assemblages of just four or five genera are likely to be encountered, and they may to some extent be differentiated at least to genus on size and other characters mentioned, as shown by
<bibRefCitation id="7F4B4B4AE020D747FE52EE93FD2FFD11" author="Kumar, A. &amp; Prasad, M." box="[444,741,691,717]" pageId="21" pageNumber="230" pagination="547 - 554" refId="ref24362" refString="Kumar, A. &amp; Prasad, M. (1977 c) On the larvae of Rhinocypha (Odonata: Cholorocyphidae) from Garhwal hills. Oriental Insects, 11, 547 - 554. https: // doi. org / 10.1080 / 00305316.1977.11090927" type="journal article" year="1977">Kumar &amp; Prasad (1977c)</bibRefCitation>
. However, it is unlikely that a comprehensive morphologybased regional key to chorocyphid genera will ever be possible. Perhaps the most promising way of determining chlorocyphid larvae over the whole region, short of breeding specimens to adulthood, is by molecular association with adults, but even this may fail at the specific level, at least when using the most widely available COI barcoding techniques (Orr
<emphasis id="29AEEAA9E020D747FEA3EF63FEB4FC81" box="[333,382,835,861]" italics="true" pageId="21" pageNumber="230">et al</emphasis>
. 2024).
</paragraph>
<paragraph id="1B6536BBE020D747FF29EF47FA9BFC78" blockId="21.[151,1437,151,933]" pageId="21" pageNumber="230">
Larva unknown:
<taxonomicName id="DCDA4D38E020D747FE69EF47FDCAFC5D" authorityName="Fraser" authorityYear="1928" box="[391,512,871,897]" class="Insecta" family="Chlorocyphidae" genus="Calocypha" kingdom="Animalia" order="Odonata" pageId="21" pageNumber="230" phylum="Arthropoda" rank="genus">
<emphasis id="29AEEAA9E020D747FE69EF47FDCAFC5D" box="[391,512,871,897]" italics="true" pageId="21" pageNumber="230">Calocypha</emphasis>
</taxonomicName>
(
<specimenCount id="0DDCFD32E020D747FDFDEF47FD82FC5D" box="[531,584,871,897]" count="1" pageId="21" pageNumber="230" type="generic">1 sp.</specimenCount>
),
<taxonomicName id="DCDA4D38E020D747FDB1EF47FD79FC5D" authorityName="Needham &amp; Gyger" authorityYear="1939" box="[607,691,871,897]" class="Insecta" family="Chlorocyphidae" genus="Cyrano" kingdom="Animalia" order="Odonata" pageId="21" pageNumber="230" phylum="Arthropoda" rank="genus">
<emphasis id="29AEEAA9E020D747FDB1EF47FD79FC5D" box="[607,691,871,897]" italics="true" pageId="21" pageNumber="230">Cyrano</emphasis>
</taxonomicName>
(2 spp.),
<taxonomicName id="DCDA4D38E020D747FCF0EF47FC76FC5D" authorityName="Ris" authorityYear="1916" box="[798,956,871,897]" class="Insecta" family="Chlorocyphidae" genus="Disparocypha" kingdom="Animalia" order="Odonata" pageId="21" pageNumber="230" phylum="Arthropoda" rank="genus">
<emphasis id="29AEEAA9E020D747FCF0EF47FC76FC5D" box="[798,956,871,897]" italics="true" pageId="21" pageNumber="230">Disparocypha</emphasis>
</taxonomicName>
(
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),
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<emphasis id="29AEEAA9E020D747FBF4EF47FB60FC5D" box="[1050,1194,871,897]" italics="true" pageId="21" pageNumber="230">Heterocypha</emphasis>
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(
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),
<taxonomicName id="DCDA4D38E020D747FAE9EF47FAB7FC5D" authorityName="Fraser" authorityYear="1949" box="[1287,1405,871,897]" class="Insecta" family="Chlorocyphidae" genus="Indocypha" kingdom="Animalia" order="Odonata" pageId="21" pageNumber="230" phylum="Arthropoda" rank="genus">
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</taxonomicName>
(7 spp.),
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<emphasis id="29AEEAA9E020D747FF32EFABFEBEFC79" box="[220,372,907,933]" italics="true" pageId="21" pageNumber="230">Melanocypha</emphasis>
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(
<specimenCount id="0DDCFD32E020D747FE69EFABFE76FC79" box="[391,444,907,933]" count="1" pageId="21" pageNumber="230" type="generic">1 sp.</specimenCount>
),
<taxonomicName id="DCDA4D38E020D747FE3FEFABFD90FC79" authorityName="Lieftinck" authorityYear="1950" box="[465,602,907,933]" class="Insecta" family="Chlorocyphidae" genus="Pachycypha" kingdom="Animalia" order="Odonata" pageId="21" pageNumber="230" phylum="Arthropoda" rank="genus">
<emphasis id="29AEEAA9E020D747FE3FEFABFD90FC79" box="[465,602,907,933]" italics="true" pageId="21" pageNumber="230">Pachycypha</emphasis>
</taxonomicName>
(
<specimenCount id="0DDCFD32E020D747FD82EFABFD6BFC79" box="[620,673,907,933]" count="1" pageId="21" pageNumber="230" type="generic">1 sp.</specimenCount>
),
<taxonomicName id="DCDA4D38E020D747FD59EFABFCF1FC79" authorityName="Laidlaw" authorityYear="1915" box="[695,827,907,933]" class="Insecta" family="Chlorocyphidae" genus="Rhinoneura" kingdom="Animalia" order="Odonata" pageId="21" pageNumber="230" phylum="Arthropoda" rank="genus">
<emphasis id="29AEEAA9E020D747FD59EFABFCF1FC79" box="[695,827,907,933]" italics="true" pageId="21" pageNumber="230">Rhinoneura</emphasis>
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(2 spp.),
<taxonomicName id="DCDA4D38E020D747FC4BEFABFBE4FC79" authorityName="Fraser" authorityYear="1949" box="[933,1070,907,933]" class="Insecta" family="Chlorocyphidae" genus="Sclerocypha" kingdom="Animalia" order="Odonata" pageId="21" pageNumber="230" phylum="Arthropoda" rank="genus">
<emphasis id="29AEEAA9E020D747FC4BEFABFBE4FC79" box="[933,1070,907,933]" italics="true" pageId="21" pageNumber="230">Sclerocypha</emphasis>
</taxonomicName>
(
<specimenCount id="0DDCFD32E020D747FBAFEFABFBBCFC79" box="[1089,1142,907,933]" count="1" pageId="21" pageNumber="230" type="generic">1 sp.</specimenCount>
),
<emphasis id="29AEEAA9E020D747FB65EFABFB36FC79" box="[1163,1276,907,933]" italics="true" pageId="21" pageNumber="230">Watuwilla</emphasis>
(
<specimenCount id="0DDCFD32E020D747FAE0EFABFA88FC79" box="[1294,1346,907,933]" count="1" pageId="21" pageNumber="230" type="generic">1 sp.</specimenCount>
).
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The five species of the genus range from
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through Indochina,
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and
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to north-east
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and are fairly closely related. The larva is known for just one species,
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<emphasis id="29AEEAA9E027D740FC55EE67FB41FDBC" box="[955,1163,583,609]" italics="true" pageId="18" pageNumber="227">Vestalaria venusta</emphasis>
(Hämäläinen) (
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Wang
<emphasis id="29AEEAA9E027D740FA66EE67FF67FD59" italics="true" pageId="18" pageNumber="227">et al</emphasis>
. 2017
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)
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. It differs most clearly from known
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<emphasis id="29AEEAA9E027D740FD4BEE4BFD31FD59" box="[677,763,619,645]" italics="true" pageId="18" pageNumber="227">Vestalis</emphasis>
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larvae by having relatively much longer acuminate caudal gills (
<figureCitation id="83E12A3EE027D740FF39EEAFFEE0FD75" box="[215,298,655,681]" captionStart="FIGURES 42–51. 42" captionStartId="9.[151,264,1592,1616]" captionTargetBox="[165,1413,192,1557]" captionTargetId="figure-15@9.[151,1435,181,1568]" captionTargetPageId="9" captionText="FIGURES 42–51. 42. Archineura incarnata: caudal gills; left lateral above, median below (drawn from Yang et al. 2022). 43. Echo modesta: caudal gills; left lateral above, median below, with baso-dorsal shield arrowed (original illustration). 44. Psolodesmus mandarinus, dorsal view of caudal gills, with baso-dorsal shield of median gill arrowed (redrawn from Matsuki &amp; Lien 1978). 45. Mnais andersoni, left lateral caudal gill with upper margin of median gill visible; baso-dorsal shield of median gill arrowed (original illustration). 46. Mnais gregoryi: caudal gills; left lateral above, median below, with baso-dorsal shield arrowed (drawn from Yang et al. 2021). 47. Caliphaea angka: caudal gills; left lateral above, median below (drawn from Yang et al. 2021). 48. Matrona basilaris: caudal gills; left lateral above, median below (drawn from Wang et al. 2017). 49. Vestalaria venusta: caudal gills; left lateral above, median below (drawn from Wang et al. 2017). 50. Vestalis amoena group: caudal gills; left lateral above, median below (redrawn from Lieftinck 1965). 51. Vestalis melania: caudal gills; left lateral above, median below (redrawn from Guadalquiver et al. 2022)." figureDoi="http://doi.org/10.5281/zenodo.13618337" httpUri="https://zenodo.org/record/13618337/files/figure.png" pageId="18" pageNumber="227">Fig. 49</figureCitation>
), the laterals about half the length of the remainder of the body, and in lacking the small antennomere between pedicel and scape (
<figureCitation id="83E12A3EE027D740FE34EE93FDFAFD11" box="[474,560,691,717]" captionStart="FIGURES 35–41. 35" captionStartId="8.[151,264,1776,1800]" captionTargetBox="[257,1332,159,1764]" captionTargetId="figure-21@8.[253,1335,153,1771]" captionTargetPageId="8" captionText="FIGURES 35–41. 35. Archineura incarnata, prementum (dorsal view, drawn from Yang et al. 2022). 36. Vestalis gracilis, prementum (dorsal view) and right antenna with supplementary antennomere between pedicel and scape (a) arrowed (redrawn from Rattanachan et al. 2022). 37. Vestalaria venusta: prementum (dorsal view) and right antenna (drawn from Wang et al. 2017). 38. Caliphaea angka: prementum (dorsal view) and right antenna (drawn from Yang et al. 2021). 39. Heliocypha biseriata, frontal view of labrum showing dorsal and ventral sections and ‘moustache’ of long, heavy setae on the latter (from Orr et al. 2024). 40. Heliocypha biseriata: details of labial palp in relation to prementum (a) dorsal view—base of movable hook dashed to reveal palpal lobe beneath and open base of premental median cleft arrowed, (b) frontal view, (c) inner view (from Orr et al. 2024). 41. Heliocypha biseriata, detail of fore tarsus with pectinate setae magnified (from Orr et al. 2024)." figureDoi="http://doi.org/10.5281/zenodo.13618335" httpUri="https://zenodo.org/record/13618335/files/figure.png" pageId="18" pageNumber="227">Fig. 37</figureCitation>
), the prementum is somewhat less incised but it is unclear how reliable this character is, especially given the known variation in
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; the other caveat is that other
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species
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may exhibit unexpected characters.
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<subSubSection id="53C06530E027D740FF79EF62FF24FC81" box="[151,238,834,861]" pageId="18" pageNumber="227" type="nomenclature">
<paragraph id="1B6536BBE027D740FF79EF62FF24FC81" blockId="18.[151,1437,834,2013]" box="[151,238,834,861]" pageId="18" pageNumber="227">
<heading id="402D81D7E027D740FF79EF62FF24FC81" bold="true" box="[151,238,834,861]" fontSize="11" level="1" pageId="18" pageNumber="227" reason="1">
<taxonomicName id="DCDA4D38E027D740FF79EF62FF24FC81" ID-CoL="86KH" authorityName="Selys" authorityYear="1853" box="[151,238,834,861]" class="Insecta" family="Calopterygidae" genus="Vestalis" kingdom="Animalia" order="Odonata" pageId="18" pageNumber="227" phylum="Arthropoda" rank="genus">
<emphasis id="29AEEAA9E027D740FF79EF62FF24FC81" bold="true" box="[151,238,834,861]" italics="true" pageId="18" pageNumber="227">Vestalis</emphasis>
</taxonomicName>
</heading>
</paragraph>
</subSubSection>
<subSubSection id="53C06530E027D741FF79EF47FEA0FF09" lastPageId="19" lastPageNumber="228" pageId="18" pageNumber="227" type="discussion">
<paragraph id="1B6536BBE027D740FF79EF47FAA6F8D9" blockId="18.[151,1437,834,2013]" pageId="18" pageNumber="227">
With 16 species occurring from
<collectingRegion id="D91EF859E027D740FDE7EF47FDA8FC5D" box="[521,610,871,897]" country="China" name="Yunnan" pageId="18" pageNumber="227">Yunnan</collectingRegion>
, southernmost
<collectingCountry id="63CD762BE027D740FCE0EF47FC9EFC5D" box="[782,852,871,897]" name="China" pageId="18" pageNumber="227">China</collectingCountry>
, the
<collectingCountry id="63CD762BE027D740FC62EF47FBC0FC5D" box="[908,1034,871,897]" name="Philippines" pageId="18" pageNumber="227">Philippines</collectingCountry>
and Bali in the east to the western Ghats and Sri-Lanka in the south-west the genus is the most speciose and diverse of the family in the Oriental region. The first description of a larva was
<taxonomicName id="DCDA4D38E027D740FD6CEF8FFC1FFC15" authority="(Burmeister)" baseAuthorityName="Burmeister" box="[642,981,943,969]" class="Insecta" family="Calopterygidae" genus="Vestalis" kingdom="Animalia" order="Odonata" pageId="18" pageNumber="227" phylum="Arthropoda" rank="species" species="luctuosa">
<emphasis id="29AEEAA9E027D740FD6CEF8FFCF4FC15" box="[642,830,943,969]" italics="true" pageId="18" pageNumber="227">Vestalis luctuosa</emphasis>
(Burmeister)
</taxonomicName>
from Java by
<bibRefCitation id="7F4B4B4AE027D740FB97EF8FFB3EFC15" author="Ris, F." box="[1145,1268,943,969]" pageId="18" pageNumber="227" pagination="157 - 183" refId="ref26559" refString="Ris, F. (1912) Uber Odonaten von Java und Krakatau gesammelt von Edward Jacobson. Tijdschrift voor Entomologie, 55, 157 - 183, pls. 6 - 8 excl." type="journal article" year="1912">Ris (1912)</bibRefCitation>
, who provided diagnostic drawings of exquisite detail, including the tiny supplementary antennomere following the pedicel (
<figureCitation id="83E12A3EE027D740FA81EFF3FF09FBCD" captionStart="FIGURES 35–41. 35" captionStartId="8.[151,264,1776,1800]" captionTargetBox="[257,1332,159,1764]" captionTargetId="figure-21@8.[253,1335,153,1771]" captionTargetPageId="8" captionText="FIGURES 35–41. 35. Archineura incarnata, prementum (dorsal view, drawn from Yang et al. 2022). 36. Vestalis gracilis, prementum (dorsal view) and right antenna with supplementary antennomere between pedicel and scape (a) arrowed (redrawn from Rattanachan et al. 2022). 37. Vestalaria venusta: prementum (dorsal view) and right antenna (drawn from Wang et al. 2017). 38. Caliphaea angka: prementum (dorsal view) and right antenna (drawn from Yang et al. 2021). 39. Heliocypha biseriata, frontal view of labrum showing dorsal and ventral sections and ‘moustache’ of long, heavy setae on the latter (from Orr et al. 2024). 40. Heliocypha biseriata: details of labial palp in relation to prementum (a) dorsal view—base of movable hook dashed to reveal palpal lobe beneath and open base of premental median cleft arrowed, (b) frontal view, (c) inner view (from Orr et al. 2024). 41. Heliocypha biseriata, detail of fore tarsus with pectinate setae magnified (from Orr et al. 2024)." figureDoi="http://doi.org/10.5281/zenodo.13618335" httpUri="https://zenodo.org/record/13618335/files/figure.png" pageId="18" pageNumber="227">Fig. 36a</figureCitation>
), noted with approbation by
<bibRefCitation id="7F4B4B4AE027D740FDFEEFD7FD06FBCD" author="Lieftinck, M. A." box="[528,716,1014,1041]" pageId="18" pageNumber="227" pagination="325 - 364" refId="ref24671" refString="Lieftinck, M. A. (1965) The species-group of Vestalis amoena Selys, 1853, in Sundaland (Odonata, Calopterygidae). Tijdschrift voor Entomologie, 108, 325 - 364." type="journal article" year="1965">Lieftinck (1965)</bibRefCitation>
, who illustrated the habitus.
<bibRefCitation id="7F4B4B4AE027D740FBF4EFD7FB1DFBCD" author="Lieftinck, M. A." box="[1050,1239,1014,1041]" pageId="18" pageNumber="227" pagination="325 - 364" refId="ref24671" refString="Lieftinck, M. A. (1965) The species-group of Vestalis amoena Selys, 1853, in Sundaland (Odonata, Calopterygidae). Tijdschrift voor Entomologie, 108, 325 - 364." type="journal article" year="1965">Lieftinck (1965)</bibRefCitation>
, also described a larva of the
<taxonomicName id="DCDA4D38E027D740FEF1E83BFE2FFBE9" authority="Hagen" authorityName="Hagen" box="[287,485,1051,1077]" class="Insecta" family="Calopterygidae" genus="Vestalis" kingdom="Animalia" order="Odonata" pageId="18" pageNumber="227" phylum="Arthropoda" rank="species" species="amoena">
<emphasis id="29AEEAA9E027D740FEF1E83BFE5EFBE8" box="[287,404,1051,1076]" italics="true" pageId="18" pageNumber="227">V. amoena</emphasis>
Hagen
</taxonomicName>
group, with the species not identified with certainty. Lieftinck notes that the median caudal gill of the
<taxonomicName id="DCDA4D38E027D740FE8FE81FFE70FB84" box="[353,442,1087,1112]" class="Insecta" family="Calopterygidae" genus="Vestalis" kingdom="Animalia" order="Odonata" pageId="18" pageNumber="227" phylum="Arthropoda" rank="species" species="amoena">
<emphasis id="29AEEAA9E027D740FE8FE81FFE70FB84" box="[353,442,1087,1112]" italics="true" pageId="18" pageNumber="227">amoena</emphasis>
</taxonomicName>
group is truncated (
<figureCitation id="83E12A3EE027D740FD4FE81FFD3EFB85" box="[673,756,1086,1113]" captionStart="FIGURES 42–51. 42" captionStartId="9.[151,264,1592,1616]" captionTargetBox="[165,1413,192,1557]" captionTargetId="figure-15@9.[151,1435,181,1568]" captionTargetPageId="9" captionText="FIGURES 42–51. 42. Archineura incarnata: caudal gills; left lateral above, median below (drawn from Yang et al. 2022). 43. Echo modesta: caudal gills; left lateral above, median below, with baso-dorsal shield arrowed (original illustration). 44. Psolodesmus mandarinus, dorsal view of caudal gills, with baso-dorsal shield of median gill arrowed (redrawn from Matsuki &amp; Lien 1978). 45. Mnais andersoni, left lateral caudal gill with upper margin of median gill visible; baso-dorsal shield of median gill arrowed (original illustration). 46. Mnais gregoryi: caudal gills; left lateral above, median below, with baso-dorsal shield arrowed (drawn from Yang et al. 2021). 47. Caliphaea angka: caudal gills; left lateral above, median below (drawn from Yang et al. 2021). 48. Matrona basilaris: caudal gills; left lateral above, median below (drawn from Wang et al. 2017). 49. Vestalaria venusta: caudal gills; left lateral above, median below (drawn from Wang et al. 2017). 50. Vestalis amoena group: caudal gills; left lateral above, median below (redrawn from Lieftinck 1965). 51. Vestalis melania: caudal gills; left lateral above, median below (redrawn from Guadalquiver et al. 2022)." figureDoi="http://doi.org/10.5281/zenodo.13618337" httpUri="https://zenodo.org/record/13618337/files/figure.png" pageId="18" pageNumber="227">Fig. 50</figureCitation>
) whereas in
<taxonomicName id="DCDA4D38E027D740FC69E81FFC2CFB85" box="[903,998,1087,1113]" class="Insecta" family="Calopterygidae" genus="Vestalis" kingdom="Animalia" order="Odonata" pageId="18" pageNumber="227" phylum="Arthropoda" rank="species" species="luctuosa">
<emphasis id="29AEEAA9E027D740FC69E81FFC2CFB85" box="[903,998,1087,1113]" italics="true" pageId="18" pageNumber="227">luctuosa</emphasis>
</taxonomicName>
it is acuminate (
<figureCitation id="83E12A3EE027D740FB48E81FFB31FB85" box="[1190,1275,1086,1113]" captionStart="FIGURES 42–51. 42" captionStartId="9.[151,264,1592,1616]" captionTargetBox="[165,1413,192,1557]" captionTargetId="figure-15@9.[151,1435,181,1568]" captionTargetPageId="9" captionText="FIGURES 42–51. 42. Archineura incarnata: caudal gills; left lateral above, median below (drawn from Yang et al. 2022). 43. Echo modesta: caudal gills; left lateral above, median below, with baso-dorsal shield arrowed (original illustration). 44. Psolodesmus mandarinus, dorsal view of caudal gills, with baso-dorsal shield of median gill arrowed (redrawn from Matsuki &amp; Lien 1978). 45. Mnais andersoni, left lateral caudal gill with upper margin of median gill visible; baso-dorsal shield of median gill arrowed (original illustration). 46. Mnais gregoryi: caudal gills; left lateral above, median below, with baso-dorsal shield arrowed (drawn from Yang et al. 2021). 47. Caliphaea angka: caudal gills; left lateral above, median below (drawn from Yang et al. 2021). 48. Matrona basilaris: caudal gills; left lateral above, median below (drawn from Wang et al. 2017). 49. Vestalaria venusta: caudal gills; left lateral above, median below (drawn from Wang et al. 2017). 50. Vestalis amoena group: caudal gills; left lateral above, median below (redrawn from Lieftinck 1965). 51. Vestalis melania: caudal gills; left lateral above, median below (redrawn from Guadalquiver et al. 2022)." figureDoi="http://doi.org/10.5281/zenodo.13618337" httpUri="https://zenodo.org/record/13618337/files/figure.png" pageId="18" pageNumber="227">Fig. 51</figureCitation>
). In particular he states “A peculiar feature of the antennae... [of all
<taxonomicName id="DCDA4D38E027D740FD14E843FC9AFBA1" authorityName="Selys" authorityYear="1853" box="[762,848,1123,1149]" class="Insecta" family="Calopterygidae" genus="Vestalis" kingdom="Animalia" order="Odonata" pageId="18" pageNumber="227" phylum="Arthropoda" rank="genus">
<emphasis id="29AEEAA9E027D740FD14E843FC9AFBA1" box="[762,848,1123,1149]" italics="true" pageId="18" pageNumber="227">Vestalis</emphasis>
</taxonomicName>
] is the presence of a vestigial—rarely incompletely developed—extra joint between the pedicel and the second segment [a
<emphasis id="29AEEAA9E027D740FC2FE8A7FBC3FB7D" box="[961,1033,1159,1185]" italics="true" pageId="18" pageNumber="227">lapsus</emphasis>
, he means pedicel and scape, since the pedicel normally is the second segment]. This intercalated minute segment is present in all [
<taxonomicName id="DCDA4D38E027D740FB56E88BFADBFB18" box="[1208,1297,1195,1220]" class="Insecta" family="Calopterygidae" genus="Vestalis" kingdom="Animalia" order="Odonata" pageId="18" pageNumber="227" phylum="Arthropoda" rank="species" species="amoena">
<emphasis id="29AEEAA9E027D740FB56E88BFADBFB18" box="[1208,1297,1195,1220]" italics="true" pageId="18" pageNumber="227">amoena</emphasis>
</taxonomicName>
] individuals examined...”. However, in another
<emphasis id="29AEEAA9E027D740FDF2E8EFFDAEFB35" box="[540,612,1231,1257]" italics="true" pageId="18" pageNumber="227">lapsus</emphasis>
, his illustration (
<figureCitation id="83E12A3EE027D740FCCEE8EFFCB6FB35" box="[800,892,1231,1257]" captionStart="FIGURES 12–17. 12" captionStartId="5.[151,264,1903,1927]" captionTargetBox="[190,1400,160,1874]" captionTargetId="figure-15@5.[178,1409,151,1889]" captionTargetPageId="5" captionText="FIGURES 12–17. 12. Mesopodagrion tibetanum, habitus with detail of right profile of head (based on Yu 2016). 13. M. tibetanum, detail of mask (redrawn from Yu 2016). 14. Podolestes orientalis, habitus (from Choong &amp; Orr 2010). 15. Podolestes orientalis, detail of mask (from Choong &amp; Orr 2010). 16. Rhipidolestes aculeatus, habitus and detail of lateral and median gill (modified from Asahina 1994). 17. Rhipidolestes aculeatus, detail of mask (modified from Ishida 1996)." figureDoi="http://doi.org/10.5281/zenodo.13618327" httpUri="https://zenodo.org/record/13618327/files/figure.png" pageId="18" pageNumber="227">Fig. 13c</figureCitation>
in
<bibRefCitation id="7F4B4B4AE027D740FC71E8EFFB8FFB35" author="Lieftinck, M. A." box="[927,1093,1230,1257]" pageId="18" pageNumber="227" pagination="325 - 364" refId="ref24671" refString="Lieftinck, M. A. (1965) The species-group of Vestalis amoena Selys, 1853, in Sundaland (Odonata, Calopterygidae). Tijdschrift voor Entomologie, 108, 325 - 364." type="journal article" year="1965">Lieftinck 1965</bibRefCitation>
) of the antenna shows no trace of this vestigial segment, leading later authors (
<bibRefCitation id="7F4B4B4AE027D740FD4AE8D3FC64FAD1" author="Rattanachan, K. &amp; Sangpradub, N. &amp; Keetapithchayakul, K. S." box="[676,942,1267,1293]" pageId="18" pageNumber="227" pagination="1 - 6" refId="ref26457" refString="Rattanachan, K., Sangpradub, N. &amp; Keetapithchayakul, K. S. (2022) Description of the larva of Vestalis gracilis (Rambur, 1842) (Zygoptera: Calopterygidae) from Thailand. International Journal of Odonatology, 25, 1 - 6. https: // doi. org / 10.48156 / 1388.2022.1917151" type="journal article" year="2022">
Rattanachan
<emphasis id="29AEEAA9E027D740FCD8E8D3FCADFAD1" box="[822,871,1267,1293]" italics="true" pageId="18" pageNumber="227">et al</emphasis>
. 2022
</bibRefCitation>
) to state incorrectly that
<taxonomicName id="DCDA4D38E027D740FB2FE8D3FAFEFAD0" box="[1217,1332,1267,1292]" class="Insecta" family="Calopterygidae" genus="Vestalis" kingdom="Animalia" order="Odonata" pageId="18" pageNumber="227" phylum="Arthropoda" rank="species" species="amoena">
<emphasis id="29AEEAA9E027D740FB2FE8D3FAFEFAD0" box="[1217,1332,1267,1292]" italics="true" pageId="18" pageNumber="227">V. amoena</emphasis>
</taxonomicName>
lacks the supplementary third antennomere. I have examined specimens of
<taxonomicName id="DCDA4D38E027D740FC6EE937FC3CFAEC" box="[896,1014,1303,1328]" class="Insecta" family="Calopterygidae" genus="Vestalis" kingdom="Animalia" order="Odonata" pageId="18" pageNumber="227" phylum="Arthropoda" rank="species" species="amoena">
<emphasis id="29AEEAA9E027D740FC6EE937FC3CFAEC" box="[896,1014,1303,1328]" italics="true" pageId="18" pageNumber="227">V. amoena</emphasis>
</taxonomicName>
,
<taxonomicName id="DCDA4D38E027D740FBEAE937FB30FAED" authority="Lieftinck" authorityName="Lieftinck" authorityYear="1965" box="[1028,1274,1303,1329]" class="Insecta" family="Calopterygidae" genus="Vestalis" kingdom="Animalia" order="Odonata" pageId="18" pageNumber="227" phylum="Arthropoda" rank="species" species="amaryllis">
<emphasis id="29AEEAA9E027D740FBEAE937FB41FAED" box="[1028,1163,1303,1329]" italics="true" pageId="18" pageNumber="227">V. amaryllis</emphasis>
Lieftinck
</taxonomicName>
(supposition), and
<taxonomicName id="DCDA4D38E027D740FF28E91BFE73FA89" authority="Lieftinck" authorityName="Lieftinck" authorityYear="1965" box="[198,441,1339,1365]" class="Insecta" family="Calopterygidae" genus="Vestalis" kingdom="Animalia" order="Odonata" pageId="18" pageNumber="227" phylum="Arthropoda" rank="species" species="amnicola">
<emphasis id="29AEEAA9E027D740FF28E91BFE83FA89" box="[198,329,1339,1365]" italics="true" pageId="18" pageNumber="227">V. amnicola</emphasis>
Lieftinck
</taxonomicName>
, from
<collectingCountry id="63CD762BE027D740FE11E91BFD81FA89" box="[511,587,1339,1365]" name="Brunei" pageId="18" pageNumber="227">Brunei</collectingCountry>
(
<bibRefCitation id="7F4B4B4AE027D740FDB7E91BFD09FA89" author="Orr, A. G." box="[601,707,1339,1365]" pageId="18" pageNumber="227" pagination="167 - 220" refId="ref25667" refString="Orr, A. G. (2001) An annotated checklist of the Odonata of Brunei with ecological notes and descriptions of hitherto unknown males and larvae. International Journal of Odonatology, 4, 167 - 220. https: // doi. org / 10.1080 / 13887890.2001.9748168" type="journal article" year="2001">Orr 2001</bibRefCitation>
, Orr unpublished data), all belonging to the
<taxonomicName id="DCDA4D38E027D740FB42E91BFAD5FA88" box="[1196,1311,1339,1364]" class="Insecta" family="Calopterygidae" genus="Vestalis" kingdom="Animalia" order="Odonata" pageId="18" pageNumber="227" phylum="Arthropoda" rank="species" species="amoena">
<emphasis id="29AEEAA9E027D740FB42E91BFAD5FA88" box="[1196,1311,1339,1364]" italics="true" pageId="18" pageNumber="227">V. amoena</emphasis>
</taxonomicName>
group, and all have the supplementary third antennomere and in consequent 8 segments in total.
<bibRefCitation id="7F4B4B4AE027D740FB86E97FFAE8FAA5" author="Lieftinck, M. A." box="[1128,1314,1374,1401]" pageId="18" pageNumber="227" pagination="325 - 364" refId="ref24671" refString="Lieftinck, M. A. (1965) The species-group of Vestalis amoena Selys, 1853, in Sundaland (Odonata, Calopterygidae). Tijdschrift voor Entomologie, 108, 325 - 364." type="journal article" year="1965">Lieftinck (1965)</bibRefCitation>
also notes how in general build
<taxonomicName id="DCDA4D38E027D740FE65E9A3FE2BFA41" authorityName="Selys" authorityYear="1853" box="[395,481,1411,1437]" class="Insecta" family="Calopterygidae" genus="Vestalis" kingdom="Animalia" order="Odonata" pageId="18" pageNumber="227" phylum="Arthropoda" rank="genus">
<emphasis id="29AEEAA9E027D740FE65E9A3FE2BFA41" box="[395,481,1411,1437]" italics="true" pageId="18" pageNumber="227">Vestalis</emphasis>
</taxonomicName>
larvae are intermediate between the spidery long legged
<taxonomicName id="DCDA4D38E027D740FB83E9A3FB26FA41" baseAuthorityName="Orr &amp; Hamalainen" baseAuthorityYear="2007" box="[1133,1260,1411,1437]" class="Insecta" family="Calopterygidae" genus="Neurobasis" kingdom="Animalia" order="Odonata" pageId="18" pageNumber="227" phylum="Arthropoda" rank="genus">
<emphasis id="29AEEAA9E027D740FB83E9A3FB26FA41" box="[1133,1260,1411,1437]" italics="true" pageId="18" pageNumber="227">Neurobasis</emphasis>
</taxonomicName>
and the stouter
<taxonomicName id="DCDA4D38E027D740FF79E987FF1BFA1D" box="[151,209,1447,1473]" class="Insecta" family="Calopterygidae" genus="Echo" kingdom="Animalia" order="Odonata" pageId="18" pageNumber="227" phylum="Arthropoda" rank="genus">
<emphasis id="29AEEAA9E027D740FF79E987FF1BFA1D" box="[151,209,1447,1473]" italics="true" pageId="18" pageNumber="227">Echo</emphasis>
</taxonomicName>
larvae. Larvae of the
<taxonomicName id="DCDA4D38E027D740FE27E987FDF4FA1C" box="[457,574,1447,1472]" class="Insecta" family="Calopterygidae" genus="Vestalis" kingdom="Animalia" order="Odonata" pageId="18" pageNumber="227" phylum="Arthropoda" rank="species" species="amoena">
<emphasis id="29AEEAA9E027D740FE27E987FDF4FA1C" box="[457,574,1447,1472]" italics="true" pageId="18" pageNumber="227">V. amoena</emphasis>
</taxonomicName>
group are variously found depending on their habits “among leafy trash in a shady brook”, (
<bibRefCitation id="7F4B4B4AE027D740FEAFE9EBFE20FA39" author="Lieftinck, M. A." box="[321,490,1482,1509]" pageId="18" pageNumber="227" pagination="325 - 364" refId="ref24671" refString="Lieftinck, M. A. (1965) The species-group of Vestalis amoena Selys, 1853, in Sundaland (Odonata, Calopterygidae). Tijdschrift voor Entomologie, 108, 325 - 364." type="journal article" year="1965">Lieftinck 1965</bibRefCitation>
), to the underside of stones at the edge of a fast-flowing stream and leaf packs among riffles (
<bibRefCitation id="7F4B4B4AE027D740FF07E9CFFE9CF9D5" author="Orr, A. G." box="[233,342,1519,1545]" pageId="18" pageNumber="227" pagination="167 - 220" refId="ref25667" refString="Orr, A. G. (2001) An annotated checklist of the Odonata of Brunei with ecological notes and descriptions of hitherto unknown males and larvae. International Journal of Odonatology, 4, 167 - 220. https: // doi. org / 10.1080 / 13887890.2001.9748168" type="journal article" year="2001">Orr 2001</bibRefCitation>
). It seems likely that the sylvan
<taxonomicName id="DCDA4D38E027D740FD28E9CFFC69F9D5" authority="Laidlaw" authorityName="Laidlaw" authorityYear="1915" box="[710,931,1519,1545]" class="Insecta" family="Calopterygidae" genus="Vestalis" kingdom="Animalia" order="Odonata" pageId="18" pageNumber="227" phylum="Arthropoda" rank="species" species="beryllae">
<emphasis id="29AEEAA9E027D740FD28E9CFFCF5F9D5" box="[710,831,1519,1545]" italics="true" pageId="18" pageNumber="227">V. beryllae</emphasis>
Laidlaw
</taxonomicName>
breeds in boggy springs among leaf litter as the adults are always to be found in such places (
<bibRefCitation id="7F4B4B4AE027D740FD2BEA33FCF8F9F1" author="Orr, A. G." box="[709,818,1555,1581]" pageId="18" pageNumber="227" pagination="167 - 220" refId="ref25667" refString="Orr, A. G. (2001) An annotated checklist of the Odonata of Brunei with ecological notes and descriptions of hitherto unknown males and larvae. International Journal of Odonatology, 4, 167 - 220. https: // doi. org / 10.1080 / 13887890.2001.9748168" type="journal article" year="2001">Orr 2001</bibRefCitation>
). Other descriptions and illustrations of
<taxonomicName id="DCDA4D38E027D740FB14EA33FA9AF9F1" authorityName="Selys" authorityYear="1853" box="[1274,1360,1555,1581]" class="Insecta" family="Calopterygidae" genus="Vestalis" kingdom="Animalia" order="Odonata" pageId="18" pageNumber="227" phylum="Arthropoda" rank="genus">
<emphasis id="29AEEAA9E027D740FB14EA33FA9AF9F1" box="[1274,1360,1555,1581]" italics="true" pageId="18" pageNumber="227">Vestalis</emphasis>
</taxonomicName>
larvae include:
<taxonomicName id="DCDA4D38E027D740FF14EA17FEBAF98C" box="[250,368,1591,1616]" class="Insecta" family="Calopterygidae" genus="Vestalis" kingdom="Animalia" order="Odonata" pageId="18" pageNumber="227" phylum="Arthropoda" rank="species" species="amoena">
<emphasis id="29AEEAA9E027D740FF14EA17FEBAF98C" box="[250,368,1591,1616]" italics="true" pageId="18" pageNumber="227">V. amoena</emphasis>
</taxonomicName>
group, photograph of habitus;
<taxonomicName id="DCDA4D38E027D740FD3FEA17FC71F98D" authority="(Rambur)" baseAuthorityName="Rambur" box="[721,955,1591,1617]" class="Insecta" family="Calopterygidae" genus="Vestalis" kingdom="Animalia" order="Odonata" pageId="18" pageNumber="227" phylum="Arthropoda" rank="species" species="gracilis">
<emphasis id="29AEEAA9E027D740FD3FEA17FC89F98D" box="[721,835,1591,1617]" italics="true" pageId="18" pageNumber="227">V. gracilis</emphasis>
(Rambur)
</taxonomicName>
, description and illustrations of diagnostic structures and habitus with note “usually concealed among vegetation in riffle zones”, (
<bibRefCitation id="7F4B4B4AE027D740FB92EA7BFA5AF9A9" author="Rattanachan, K. &amp; Sangpradub, N. &amp; Keetapithchayakul, K. S." box="[1148,1424,1627,1653]" pageId="18" pageNumber="227" pagination="1 - 6" refId="ref26457" refString="Rattanachan, K., Sangpradub, N. &amp; Keetapithchayakul, K. S. (2022) Description of the larva of Vestalis gracilis (Rambur, 1842) (Zygoptera: Calopterygidae) from Thailand. International Journal of Odonatology, 25, 1 - 6. https: // doi. org / 10.48156 / 1388.2022.1917151" type="journal article" year="2022">
Rattanachan
<emphasis id="29AEEAA9E027D740FAFFEA7BFA8EF9A9" box="[1297,1348,1627,1653]" italics="true" pageId="18" pageNumber="227">et al</emphasis>
. 2022
</bibRefCitation>
);
<taxonomicName id="DCDA4D38E027D740FF79EA5FFE9EF945" authority="Selys" authorityName="Selys" authorityYear="1873" box="[151,340,1663,1689]" class="Insecta" family="Calopterygidae" genus="Vestalis" kingdom="Animalia" order="Odonata" pageId="18" pageNumber="227" phylum="Arthropoda" rank="species" species="melania">
<emphasis id="29AEEAA9E027D740FF79EA5FFEC7F945" box="[151,269,1663,1689]" italics="true" pageId="18" pageNumber="227">V. melania</emphasis>
Selys
</taxonomicName>
, description and illustrations of diagnostic structures and habitus (
<figureCitation id="83E12A3EE027D740FBD3EA5FFB45F945" box="[1085,1167,1662,1689]" captionStart="FIGURES 52–56. 52" captionStartId="10.[151,264,1941,1965]" captionTargetBox="[191,1388,163,1929]" captionTargetId="figure-21@10.[190,1397,151,1933]" captionTargetPageId="10" captionText="FIGURES 52–56. 52. Caliphaea angka, habitus (based on Yang et al. 2021). 53. Vestalis gracilis, habitus (redrawn from Rattanachan et al. 2022). 54. Neurobasis chinensis, habitus (original drawing). 55. Matrona basilaris, habitus (drawn from Wang et al. 2017). 56. Echo uniformis, habitus (redrawn from Lieftinck 1965)." figureDoi="http://doi.org/10.5281/zenodo.13618339" httpUri="https://zenodo.org/record/13618339/files/figure.png" pageId="18" pageNumber="227">Fig. 53</figureCitation>
) with note “found in an unshaded, narrow, montane stream with a sandy substrate and dense marginal and submerged vegetation”, as well as comparisons with other species (
<bibRefCitation id="7F4B4B4AE027D740FDC8EAE7FC8FF93D" author="Guadalquiver, D. M. E. &amp; Nuneza, O. M. &amp; Tabugo, S. R. M. &amp; Villanueva, R. J. T." box="[550,837,1735,1761]" pageId="18" pageNumber="227" pagination="21612 - 21618" refId="ref23206" refString="Guadalquiver, D. M. E., Nuneza, O. M., Tabugo, S. R. M. &amp; Villanueva, R. J. T. (2022) Description of the larva of Vestalis melania (Selys, 1873) (Odonata: Calopterygidae) identified through DNA barcoding. Journal of Threatened Taxa, 14, 21612 - 21618. https: // doi. org / 10.11609 / jott. 7830.14.8.21612 - 21618" type="journal article" year="2022">
Guadalquiver
<emphasis id="29AEEAA9E027D740FD26EAE7FD30F93D" box="[712,762,1735,1761]" italics="true" pageId="18" pageNumber="227">et al</emphasis>
. 2022
</bibRefCitation>
);
<taxonomicName id="DCDA4D38E027D740FCB7EAE7FBF1F93D" authority="Fraser" authorityName="Fraser" authorityYear="1929" box="[857,1083,1735,1761]" class="Insecta" family="Calopterygidae" genus="Vestalis" kingdom="Animalia" order="Odonata" pageId="18" pageNumber="227" phylum="Arthropoda" rank="species" species="nigrescens">
<emphasis id="29AEEAA9E027D740FCB7EAE7FC26F93C" box="[857,1004,1735,1760]" italics="true" pageId="18" pageNumber="227">V. nigrescens</emphasis>
Fraser
</taxonomicName>
, stated by
<bibRefCitation id="7F4B4B4AE027D740FB5DEAE7FAB5F93D" author="St Quentin, D." box="[1203,1407,1735,1761]" pageId="18" pageNumber="227" pagination="113 - 124" refId="ref26637" refString="St Quentin, D. (1973) Contributions to the ecology of the larvae some Odonata from Ceylon. Bulletin of the Fisheries Research Station, Sri Lanka (Ceylon), 24, 113 - 124." type="journal article" year="1973">St Quentin (1973)</bibRefCitation>
to be similar to
<taxonomicName id="DCDA4D38E027D740FEC4EACBFE6FF8D9" box="[298,421,1771,1797]" class="Insecta" family="Calopterygidae" genus="Vestalis" kingdom="Animalia" order="Odonata" pageId="18" pageNumber="227" phylum="Arthropoda" rank="species" species="luctuosa">
<emphasis id="29AEEAA9E027D740FEC4EACBFE6FF8D9" box="[298,421,1771,1797]" italics="true" pageId="18" pageNumber="227">V. luctuosa</emphasis>
</taxonomicName>
, with acuminate caudal lamellae, and inhabiting the banks of stony bottomed streams.
</paragraph>
<paragraph id="1B6536BBE027D741FF29EB2FFEA0FF09" blockId="18.[151,1437,834,2013]" lastBlockId="19.[151,1437,151,213]" lastPageId="19" lastPageNumber="228" pageId="18" pageNumber="227">
The species, essentially alike in general appearance, fall into two categories: those with a truncate median caudal gill (
<figureCitation id="83E12A3EE027D740FF21EB13FEEFF891" box="[207,293,1842,1869]" captionStart="FIGURES 42–51. 42" captionStartId="9.[151,264,1592,1616]" captionTargetBox="[165,1413,192,1557]" captionTargetId="figure-15@9.[151,1435,181,1568]" captionTargetPageId="9" captionText="FIGURES 42–51. 42. Archineura incarnata: caudal gills; left lateral above, median below (drawn from Yang et al. 2022). 43. Echo modesta: caudal gills; left lateral above, median below, with baso-dorsal shield arrowed (original illustration). 44. Psolodesmus mandarinus, dorsal view of caudal gills, with baso-dorsal shield of median gill arrowed (redrawn from Matsuki &amp; Lien 1978). 45. Mnais andersoni, left lateral caudal gill with upper margin of median gill visible; baso-dorsal shield of median gill arrowed (original illustration). 46. Mnais gregoryi: caudal gills; left lateral above, median below, with baso-dorsal shield arrowed (drawn from Yang et al. 2021). 47. Caliphaea angka: caudal gills; left lateral above, median below (drawn from Yang et al. 2021). 48. Matrona basilaris: caudal gills; left lateral above, median below (drawn from Wang et al. 2017). 49. Vestalaria venusta: caudal gills; left lateral above, median below (drawn from Wang et al. 2017). 50. Vestalis amoena group: caudal gills; left lateral above, median below (redrawn from Lieftinck 1965). 51. Vestalis melania: caudal gills; left lateral above, median below (redrawn from Guadalquiver et al. 2022)." figureDoi="http://doi.org/10.5281/zenodo.13618337" httpUri="https://zenodo.org/record/13618337/files/figure.png" pageId="18" pageNumber="227">Fig. 50</figureCitation>
), (
<taxonomicName id="DCDA4D38E027D740FEABEB13FE54F890" box="[325,414,1843,1868]" class="Insecta" family="Calopterygidae" genus="Vestalis" kingdom="Animalia" order="Odonata" pageId="18" pageNumber="227" phylum="Arthropoda" rank="species" species="amoena">
<emphasis id="29AEEAA9E027D740FEABEB13FE54F890" box="[325,414,1843,1868]" italics="true" pageId="18" pageNumber="227">amoena</emphasis>
</taxonomicName>
group and
<taxonomicName id="DCDA4D38E027D740FDCAEB13FDB0F891" box="[548,634,1843,1869]" class="Insecta" family="Calopterygidae" genus="Vestalis" kingdom="Animalia" order="Odonata" pageId="18" pageNumber="227" phylum="Arthropoda" rank="species" species="gracilis">
<emphasis id="29AEEAA9E027D740FDCAEB13FDB0F891" box="[548,634,1843,1869]" italics="true" pageId="18" pageNumber="227">gracilis</emphasis>
</taxonomicName>
) and those with an acuminate central caudal gill (
<figureCitation id="83E12A3EE027D740FB2EEB13FADFF891" box="[1216,1301,1842,1869]" captionStart="FIGURES 42–51. 42" captionStartId="9.[151,264,1592,1616]" captionTargetBox="[165,1413,192,1557]" captionTargetId="figure-15@9.[151,1435,181,1568]" captionTargetPageId="9" captionText="FIGURES 42–51. 42. Archineura incarnata: caudal gills; left lateral above, median below (drawn from Yang et al. 2022). 43. Echo modesta: caudal gills; left lateral above, median below, with baso-dorsal shield arrowed (original illustration). 44. Psolodesmus mandarinus, dorsal view of caudal gills, with baso-dorsal shield of median gill arrowed (redrawn from Matsuki &amp; Lien 1978). 45. Mnais andersoni, left lateral caudal gill with upper margin of median gill visible; baso-dorsal shield of median gill arrowed (original illustration). 46. Mnais gregoryi: caudal gills; left lateral above, median below, with baso-dorsal shield arrowed (drawn from Yang et al. 2021). 47. Caliphaea angka: caudal gills; left lateral above, median below (drawn from Yang et al. 2021). 48. Matrona basilaris: caudal gills; left lateral above, median below (drawn from Wang et al. 2017). 49. Vestalaria venusta: caudal gills; left lateral above, median below (drawn from Wang et al. 2017). 50. Vestalis amoena group: caudal gills; left lateral above, median below (redrawn from Lieftinck 1965). 51. Vestalis melania: caudal gills; left lateral above, median below (redrawn from Guadalquiver et al. 2022)." figureDoi="http://doi.org/10.5281/zenodo.13618337" httpUri="https://zenodo.org/record/13618337/files/figure.png" pageId="18" pageNumber="227">Fig. 51</figureCitation>
), (
<taxonomicName id="DCDA4D38E027D740FAD8EB13FA5FF891" box="[1334,1429,1843,1869]" class="Insecta" family="Calopterygidae" genus="Vestalis" kingdom="Animalia" order="Odonata" pageId="18" pageNumber="227" phylum="Arthropoda" rank="species" species="luctuosa">
<emphasis id="29AEEAA9E027D740FAD8EB13FA5FF891" box="[1334,1429,1843,1869]" italics="true" pageId="18" pageNumber="227">luctuosa</emphasis>
</taxonomicName>
,
<taxonomicName id="DCDA4D38E027D740FF79EB77FF3BF8AD" authorityName="Selys" authorityYear="1873" box="[151,241,1879,1905]" class="Insecta" family="Calopterygidae" genus="Vestalis" kingdom="Animalia" order="Odonata" pageId="18" pageNumber="227" phylum="Arthropoda" rank="species" species="melania">
<emphasis id="29AEEAA9E027D740FF79EB77FF3BF8AD" box="[151,241,1879,1905]" italics="true" pageId="18" pageNumber="227">melania</emphasis>
</taxonomicName>
and
<taxonomicName id="DCDA4D38E027D740FEC7EB77FE6AF8AC" authorityName="Fraser" authorityYear="1929" box="[297,416,1879,1904]" class="Insecta" family="Calopterygidae" genus="Vestalis" kingdom="Animalia" order="Odonata" pageId="18" pageNumber="227" phylum="Arthropoda" rank="species" species="nigrescens">
<emphasis id="29AEEAA9E027D740FEC7EB77FE6AF8AC" box="[297,416,1879,1904]" italics="true" pageId="18" pageNumber="227">nigrescens</emphasis>
</taxonomicName>
). There are also differences in the shape of the mask, which tends to be relatively broader apically with shorter, broader premental lobes in
<taxonomicName id="DCDA4D38E027D740FD29EB5BFCECF849" box="[711,806,1915,1941]" class="Insecta" family="Calopterygidae" genus="Vestalis" kingdom="Animalia" order="Odonata" pageId="18" pageNumber="227" phylum="Arthropoda" rank="species" species="luctuosa">
<emphasis id="29AEEAA9E027D740FD29EB5BFCECF849" box="[711,806,1915,1941]" italics="true" pageId="18" pageNumber="227">luctuosa</emphasis>
</taxonomicName>
and perhaps
<taxonomicName id="DCDA4D38E027D740FC50EB5BFBD2F849" authorityName="Selys" authorityYear="1873" box="[958,1048,1915,1941]" class="Insecta" family="Calopterygidae" genus="Vestalis" kingdom="Animalia" order="Odonata" pageId="18" pageNumber="227" phylum="Arthropoda" rank="species" species="melania">
<emphasis id="29AEEAA9E027D740FC50EB5BFBD2F849" box="[958,1048,1915,1941]" italics="true" pageId="18" pageNumber="227">melania</emphasis>
</taxonomicName>
and with subtle differences in the form of the premental cleft, about which
<bibRefCitation id="7F4B4B4AE027D740FDB1EBBFFCD2F865" author="Lieftinck, M. A." box="[607,792,1950,1977]" pageId="18" pageNumber="227" pagination="325 - 364" refId="ref24671" refString="Lieftinck, M. A. (1965) The species-group of Vestalis amoena Selys, 1853, in Sundaland (Odonata, Calopterygidae). Tijdschrift voor Entomologie, 108, 325 - 364." type="journal article" year="1965">Lieftinck (1965)</bibRefCitation>
, who first noted a possible difference, remained sceptical. With good information available for fewer than half the known species, morphology based larval identification will generally depend on locality, with the probability that eight antennomeres with a tiny third segment is positive proof of generic identity.
</paragraph>
</subSubSection>
</treatment>
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Family
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This monotypic family is known only from
<collectingRegion id="D91EF859E029D74EFD6CE92FFD19FAF5" box="[642,723,1295,1321]" country="China" name="Hainan" pageId="28" pageNumber="237">Hainan</collectingRegion>
island where the larvae of
<taxonomicName id="DCDA4D38E029D74EFBECE92FFA8DFAF5" authority="Kirby" authorityName="Kirby" authorityYear="1900" box="[1026,1351,1295,1321]" class="Insecta" family="Pseudolestidae" genus="Pseudolestes" kingdom="Animalia" order="Odonata" pageId="28" pageNumber="237" phylum="Arthropoda" rank="species" species="mirabilis">
<emphasis id="29AEEAA9E029D74EFBECE92FFB35FAF5" box="[1026,1279,1295,1321]" italics="true" pageId="28" pageNumber="237">Pseudolestes mirabilis</emphasis>
Kirby
</taxonomicName>
inhabit small clear, rocky streams; the females lay in dead wood (
<bibRefCitation id="7F4B4B4AE029D74EFCC7E914FC7DFA93" author="Zhang, H. - M." box="[809,951,1332,1359]" pageId="28" pageNumber="237" refId="ref28336" refString="Zhang, H. - M. (2019) Dragonflies and Damselflies of China. Vols. 1 &amp; 2. Chong Qing University Press, ChongQing, 14 + 1460 pp. [bilingual, in Chinese and English]" type="book" year="2019">Zhang 2019</bibRefCitation>
). The larvae (
<figureCitation id="83E12A3EE029D74EFBBCE914FB0FFA93" box="[1106,1221,1332,1359]" captionStart="FIGURES 8–11. 8" captionStartId="4.[151,264,1914,1938]" captionTargetBox="[200,1382,177,1900]" captionTargetId="figure-21@4.[193,1394,169,1908]" captionTargetPageId="4" captionText="FIGURES 8–11. 8. Pseudolestes mirabilis, ventral view of terminal abdominal segments and caudal gills of showing gill tufts on S10 (after Yu &amp; Bu 2011). 9. Devadatta argyoides, detail of extruded right gill tuft, posterior view (redrawn from Watson 1966). 10. Pseudolestes mirabilis, habitus with detail of mask (after Yu &amp; Bu 2011). 11. Devadatta argyoides, habitus with details of right antenna and mask (original drawing)." figureDoi="http://doi.org/10.5281/zenodo.13618323" httpUri="https://zenodo.org/record/13618323/files/figure.png" pageId="28" pageNumber="237">Figs 8, 10</figureCitation>
) are unmistakable, being provided with paired dendritic gill tufts on the venter of S10 (
<bibRefCitation id="7F4B4B4AE029D74EFC69E97AFBE7FAA8" author="Yu, X. &amp; Bu, W." box="[903,1069,1370,1396]" pageId="28" pageNumber="237" pagination="105 - 110" refId="ref28152" refString="Yu, X. &amp; Bu, W. (2011) A description of the remarkable larva of Pseudolestes mirabilis Kirby (Odonata: Pseudolestidae). International Journal of Odonatology, 14 (2), 105 - 110. https: // doi. org / 10.1080 / 13887890.2011.592486" type="journal article" year="2011">Yu &amp; Bu 2011</bibRefCitation>
), a character shared in the region only with
<taxonomicName id="DCDA4D38E029D74EFEE2E95FFE4BFA45" authorityName="Kirby" authorityYear="1890" box="[268,385,1407,1433]" class="Insecta" family="Amphipterygidae" genus="Devadatta" kingdom="Animalia" order="Odonata" pageId="28" pageNumber="237" phylum="Arthropoda" rank="genus">
<emphasis id="29AEEAA9E029D74EFEE2E95FFE4BFA45" box="[268,385,1407,1433]" italics="true" pageId="28" pageNumber="237">Devadatta</emphasis>
</taxonomicName>
but it is unclear if the structures are homologous. Larva of
<taxonomicName id="DCDA4D38E029D74EFBC1E95FFB0AFA45" authorityName="Kirby" authorityYear="1900" box="[1071,1216,1407,1433]" class="Insecta" family="Pseudolestidae" genus="Pseudolestes" kingdom="Animalia" order="Odonata" pageId="28" pageNumber="237" phylum="Arthropoda" rank="genus">
<emphasis id="29AEEAA9E029D74EFBC1E95FFB0AFA45" box="[1071,1216,1407,1433]" italics="true" pageId="28" pageNumber="237">Pseudolestes</emphasis>
</taxonomicName>
also differs clearly from
<taxonomicName id="DCDA4D38E029D74EFF3BE984FE80FA62" authorityName="Kirby" authorityYear="1890" box="[213,330,1444,1470]" class="Insecta" family="Amphipterygidae" genus="Devadatta" kingdom="Animalia" order="Odonata" pageId="28" pageNumber="237" phylum="Arthropoda" rank="genus">
<emphasis id="29AEEAA9E029D74EFF3BE984FE80FA62" box="[213,330,1444,1470]" italics="true" pageId="28" pageNumber="237">Devadatta</emphasis>
</taxonomicName>
in the structure of the hardened balloon like caudal gills, the shape of the prementum and the form of the antennae. The tufts can be retracted and extruded and presumably play a role in respiration (
<bibRefCitation id="7F4B4B4AE029D74EFB0BE9EAFA44FA38" author="Yu, X. &amp; Bu, W." box="[1253,1422,1482,1508]" pageId="28" pageNumber="237" pagination="123 - 126" refId="ref28204" refString="Yu, X. &amp; Bu, W. (2014) Notes on the retractability of gill tufts in Pseudolestes mirabilis (Zygoptera: Pseudolestidae). International Journal of Odonatology, 17 (2 - 3), 123 - 126. https: // doi. org / 10.1080 / 13887890.2014.933447" type="journal article" year="2014">Yu &amp; Bu 2014</bibRefCitation>
).
</paragraph>
</subSubSection>
</treatment>
</document>

230
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<subSubSection id="53C06530E029D74EFF79ECB7FE5AFF6D" box="[151,400,151,177]" pageId="28" pageNumber="237" type="nomenclature">
<paragraph id="1B6536BBE029D74EFF79ECB7FE5AFF6D" blockId="28.[151,400,151,177]" box="[151,400,151,177]" pageId="28" pageNumber="237">
<heading id="402D81D7E029D74EFF79ECB7FE5AFF6D" bold="true" box="[151,400,151,177]" fontSize="11" level="1" pageId="28" pageNumber="237" reason="1">
<emphasis id="29AEEAA9E029D74EFF79ECB7FE5AFF6D" bold="true" box="[151,400,151,177]" pageId="28" pageNumber="237">
Family
<taxonomicName id="DCDA4D38E029D74EFF19ECB7FE5AFF6D" ID-CoL="9WSX7" authorityName="Kennedy" authorityYear="1925" box="[247,400,151,177]" class="Insecta" family="Philosinidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="28" pageNumber="237" phylum="Arthropoda" rank="family">Philosinidae</taxonomicName>
</emphasis>
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<subSubSection id="53C06530E029D74EFF79ECFFFAE2FCC9" pageId="28" pageNumber="237" type="discussion">
<paragraph id="1B6536BBE029D74EFF79ECFFFAE2FCC9" blockId="28.[151,1437,223,790]" pageId="28" pageNumber="237">
This small family ranges from southern
<collectingCountry id="63CD762BE029D74EFDB0ECFFFD6BFF25" box="[606,673,223,249]" name="China" pageId="28" pageNumber="237">China</collectingCountry>
to Indochina,
<collectingCountry id="63CD762BE029D74EFCABECFFFC63FF25" box="[837,937,223,249]" name="Thailand" pageId="28" pageNumber="237">Thailand</collectingCountry>
and much of Sundaland. The larvae of both genera are described (
<bibRefCitation id="7F4B4B4AE029D74EFE7AED23FD7FFEC1" author="Needham, J. G. &amp; Gyger, M. K." box="[404,693,259,285]" pageId="28" pageNumber="237" pagination="239 - 314" refId="ref25055" refString="Needham, J. G. &amp; Gyger, M. K. (1939) The Odonata of the Philippines, II. Suborder Zygoptera. Philippine Journal of Science, 70, 239 - 314, pls. 11 - 22 excl." type="journal article" year="1939">Needham &amp; Gyger 1939</bibRefCitation>
,
<bibRefCitation id="7F4B4B4AE029D74EFD2DED23FCA5FEC2" author="Lieftinck, M. A." box="[707,879,259,286]" pageId="28" pageNumber="237" pagination="59 - 121" refId="ref24595" refString="Lieftinck, M. A. (1956) Revision of the genus Argiolestes Selys (Odonata) in New Guinea and the Moluccas, with notes on the larval forms of the family Megapodagrionidae. Nova Guinea, New Series, 7, 59 - 121." type="journal article" year="1956">Lieftinck 1956</bibRefCitation>
,
<bibRefCitation id="7F4B4B4AE029D74EFC92ED23FC22FEC1" author="Orr, A. G." box="[892,1000,259,285]" pageId="28" pageNumber="237" refId="ref25717" refString="Orr, A. G. (2003) A guide to the Dragonflies of Borneo: their identification and biology. Natural History Publications (Borneo), Kota Kinabalu, x + 195 pp." type="book" year="2003">Orr 2003</bibRefCitation>
,
<bibRefCitation id="7F4B4B4AE029D74EFC18ED23FB0CFEC1" author="Zhang, H. - M. &amp; Kalkman, V. J. &amp; Tong, X. - L." box="[1014,1222,259,285]" pageId="28" pageNumber="237" pagination="55 - 68" refId="ref28378" refString="Zhang, H. - M., Kalkman, V. J. &amp; Tong, X. - L. (2011) A synopsis of the genus Philosina with descriptions of the larvae of P. alba and P. buchi (Odonata: Megapodagrionidae), International Journal of Odonatology, 14, 55 - 68. https: // doi. org / 10.1080 / 13887890.2011.568189" type="journal article" year="2011">
Zhang
<emphasis id="29AEEAA9E029D74EFBA9ED24FB48FEC1" box="[1095,1154,259,285]" italics="true" pageId="28" pageNumber="237">et al.</emphasis>
2011
</bibRefCitation>
,
<bibRefCitation id="7F4B4B4AE029D74EFB3DED23FF1BFE9D" author="Kawashima, I. &amp; Sasamoto, A. &amp; Toan, P. Q. &amp; Cuong, D. M." pageId="28" pageNumber="237" pagination="93 - 99" refId="ref23943" refString="Kawashima, I., Sasamoto, A., Toan, P. Q. &amp; Cuong, D. M. (2011) First discovery and description of female and larva of Rhinagrion hainanense Wilson and Reels, 2001 (= R. yokoii Sasamoto, 2003) (Zygoptera: Megapodagrionidae) from Vietnam. Tombo - Acta Odonatologica Japonica, 53, 93 - 99." type="journal article" year="2011">
Kawashima
<emphasis id="29AEEAA9E029D74EFA8FED24FA5FFEC1" box="[1377,1429,259,285]" italics="true" pageId="28" pageNumber="237">et al</emphasis>
. 2011
</bibRefCitation>
,
<bibRefCitation id="7F4B4B4AE029D74EFF32ED07FDE8FE9D" author="Novelo-Gutierrez, R. &amp; Sites, R. W. &amp; Vitheepradit, A." box="[220,546,295,322]" pageId="28" pageNumber="237" pagination="562 - 568" refId="ref25601" refString="Novelo-Gutierrez, R., Sites, R. W. &amp; Vitheepradit, A. (2014) New province record of Rhinagrion for Thailand and description of the larva of R. mima (Odonata: Zygoptera: Philosinidae). Zootaxa, 5852 (5), 562 - 568. https: // doi. org / 10.11646 / zootaxa. 3852.5.4" type="journal article" year="2014">
Novelo-Gutiérrez
<emphasis id="29AEEAA9E029D74EFE46ED08FE13FE9D" box="[424,473,295,321]" italics="true" pageId="28" pageNumber="237">et al</emphasis>
. 2014
</bibRefCitation>
), with both species of
<taxonomicName id="DCDA4D38E029D74EFCF2ED07FC4DFE9D" box="[796,903,295,321]" class="Insecta" family="Megapodagrionidae" genus="Philosina" kingdom="Animalia" order="Odonata" pageId="28" pageNumber="237" phylum="Arthropoda" rank="genus">
<emphasis id="29AEEAA9E029D74EFCF2ED07FC4DFE9D" box="[796,903,295,321]" italics="true" pageId="28" pageNumber="237">Philosina</emphasis>
</taxonomicName>
,
<taxonomicName id="DCDA4D38E029D74EFC7BED08FBF2FE9D" authority="Wilson" authorityName="Wilson" authorityYear="1999" box="[917,1080,295,321]" class="Insecta" family="Megapodagrionidae" genus="Philosina" kingdom="Animalia" order="Odonata" pageId="28" pageNumber="237" phylum="Arthropoda" rank="species" species="alba">
<emphasis id="29AEEAA9E029D74EFC7BED08FC28FE9D" box="[917,994,295,321]" italics="true" pageId="28" pageNumber="237">P. alba</emphasis>
Wilson
</taxonomicName>
and
<taxonomicName id="DCDA4D38E029D74EFB80ED08FB3CFE9D" authority="Ris" authorityName="Ris" authorityYear="1917" box="[1134,1270,295,321]" class="Insecta" family="Megapodagrionidae" genus="Philosina" kingdom="Animalia" order="Odonata" pageId="28" pageNumber="237" phylum="Arthropoda" rank="species" species="buchi">
<emphasis id="29AEEAA9E029D74EFB80ED08FB0DFE9D" box="[1134,1223,295,321]" italics="true" pageId="28" pageNumber="237">P. buchi</emphasis>
Ris
</taxonomicName>
, and five of 10 species of
<taxonomicName id="DCDA4D38E029D74EFEE1ED6BFE45FEB9" authorityName="Calvert" authorityYear="1913" box="[271,399,331,357]" class="Insecta" family="Megapodagrionidae" genus="Rhinagrion" kingdom="Animalia" order="Odonata" pageId="28" pageNumber="237" phylum="Arthropoda" rank="genus">
<emphasis id="29AEEAA9E029D74EFEE1ED6BFE45FEB9" box="[271,399,331,357]" italics="true" pageId="28" pageNumber="237">Rhinagrion</emphasis>
</taxonomicName>
known. Not much interspecific variation is expected in the latter. Distinctive features of the family include: prementum with strongly developed median lobe with deep, apically closed, median cleft (
<figureCitation id="83E12A3EE029D74EFAAEED4FFF7CFE71" captionStart="FIGURES 18–26. 18" captionStartId="6.[151,264,1811,1835]" captionTargetBox="[265,1336,167,1790]" captionTargetId="figure-21@6.[248,1339,153,1804]" captionTargetPageId="6" captionText="FIGURES 18–26. 18. Rhinagrion borneense, habitus (original drawing). 19. Philosina alba, habitus (modified from Zhang et al. 2011). 20. Rhinagrion mima, caudal gills: above heavily tracheated median gill; below right lateral gill with basal spines along mid-rib (modified from Lieftinck 1956). 21. Philosina alba caudal gills: above heavily tracheated median gill; below right lateral gill with basal spines along mid-rib (modified from Zhang et al. 2011) 22. Rhinagrion borneense, detail of mask (original drawing). 23. Philosina buchi, detail of mask (redrawn from Zhang et al. 2011). 24. Philoganga vetusta, habitus (adapted from Xu 2016). 25. Philoganga vetusta, details of caudal gills and antenna (redrawn from Xu 2016). 26. Philoganga montana, detail of mask (original drawing)." figureDoi="http://doi.org/10.5281/zenodo.13618331" httpUri="https://zenodo.org/record/13618331/files/figure.png" pageId="28" pageNumber="237">Figs 22, 23</figureCitation>
); median caudal gill densely tracheated, slightly (
<figureCitation id="83E12A3EE029D74EFD09EDB3FCF3FE71" box="[743,825,403,429]" captionStart="FIGURES 18–26. 18" captionStartId="6.[151,264,1811,1835]" captionTargetBox="[265,1336,167,1790]" captionTargetId="figure-21@6.[248,1339,153,1804]" captionTargetPageId="6" captionText="FIGURES 18–26. 18. Rhinagrion borneense, habitus (original drawing). 19. Philosina alba, habitus (modified from Zhang et al. 2011). 20. Rhinagrion mima, caudal gills: above heavily tracheated median gill; below right lateral gill with basal spines along mid-rib (modified from Lieftinck 1956). 21. Philosina alba caudal gills: above heavily tracheated median gill; below right lateral gill with basal spines along mid-rib (modified from Zhang et al. 2011) 22. Rhinagrion borneense, detail of mask (original drawing). 23. Philosina buchi, detail of mask (redrawn from Zhang et al. 2011). 24. Philoganga vetusta, habitus (adapted from Xu 2016). 25. Philoganga vetusta, details of caudal gills and antenna (redrawn from Xu 2016). 26. Philoganga montana, detail of mask (original drawing)." figureDoi="http://doi.org/10.5281/zenodo.13618331" httpUri="https://zenodo.org/record/13618331/files/figure.png" pageId="28" pageNumber="237">Fig. 20</figureCitation>
) or significantly shorter (
<figureCitation id="83E12A3EE029D74EFBB7EDB3FB61FE71" box="[1113,1195,403,429]" captionStart="FIGURES 18–26. 18" captionStartId="6.[151,264,1811,1835]" captionTargetBox="[265,1336,167,1790]" captionTargetId="figure-21@6.[248,1339,153,1804]" captionTargetPageId="6" captionText="FIGURES 18–26. 18. Rhinagrion borneense, habitus (original drawing). 19. Philosina alba, habitus (modified from Zhang et al. 2011). 20. Rhinagrion mima, caudal gills: above heavily tracheated median gill; below right lateral gill with basal spines along mid-rib (modified from Lieftinck 1956). 21. Philosina alba caudal gills: above heavily tracheated median gill; below right lateral gill with basal spines along mid-rib (modified from Zhang et al. 2011) 22. Rhinagrion borneense, detail of mask (original drawing). 23. Philosina buchi, detail of mask (redrawn from Zhang et al. 2011). 24. Philoganga vetusta, habitus (adapted from Xu 2016). 25. Philoganga vetusta, details of caudal gills and antenna (redrawn from Xu 2016). 26. Philoganga montana, detail of mask (original drawing)." figureDoi="http://doi.org/10.5281/zenodo.13618331" httpUri="https://zenodo.org/record/13618331/files/figure.png" pageId="28" pageNumber="237">Fig. 21</figureCitation>
) and thinner than the lateral caudal lamellae; lateral lamellae fleshy and undulating each with basally expanded midrib armed with spines on outer face. In
<taxonomicName id="DCDA4D38E029D74EFEBCEDFBFE18FE29" authorityName="Calvert" authorityYear="1913" box="[338,466,475,501]" class="Insecta" family="Megapodagrionidae" genus="Rhinagrion" kingdom="Animalia" order="Odonata" pageId="28" pageNumber="237" phylum="Arthropoda" rank="genus">
<emphasis id="29AEEAA9E029D74EFEBCEDFBFE18FE29" box="[338,466,475,501]" italics="true" pageId="28" pageNumber="237">Rhinagrion</emphasis>
</taxonomicName>
, in life the lateral gills form a tube shielding the median lamellae (
<bibRefCitation id="7F4B4B4AE029D74EFB41EDFBFA5FFE29" author="Kalkman, V. J. &amp; Choong, C. Y. &amp; Orr, A. G. &amp; Schutte, K." box="[1199,1429,475,501]" pageId="28" pageNumber="237" pagination="119 - 135" refId="ref23830" 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, 119 - 135. https: // doi. org / 10.1080 / 13887890.2010.9748366" type="journal article" year="2010">
Kalkman
<emphasis id="29AEEAA9E029D74EFAF0EDFCFA9FFE29" box="[1310,1365,475,501]" italics="true" pageId="28" pageNumber="237">et al.</emphasis>
2010
</bibRefCitation>
) and it is believed the same may be true of
<taxonomicName id="DCDA4D38E029D74EFD82EDDFFC72FDC5" authority="(Zhang et al. 2011)" baseAuthorityName="Zhang" baseAuthorityYear="2011" box="[620,952,511,538]" class="Insecta" family="Megapodagrionidae" genus="Philosina" kingdom="Animalia" order="Odonata" pageId="28" pageNumber="237" phylum="Arthropoda" rank="genus">
<emphasis id="29AEEAA9E029D74EFD82EDDFFD1DFDC5" box="[620,727,511,537]" italics="true" pageId="28" pageNumber="237">Philosina</emphasis>
(
<bibRefCitation id="7F4B4B4AE029D74EFD08EDDFFC7AFDC5" author="Zhang, H. - M. &amp; Kalkman, V. J. &amp; Tong, X. - L." box="[742,944,511,538]" pageId="28" pageNumber="237" pagination="55 - 68" refId="ref28378" refString="Zhang, H. - M., Kalkman, V. J. &amp; Tong, X. - L. (2011) A synopsis of the genus Philosina with descriptions of the larvae of P. alba and P. buchi (Odonata: Megapodagrionidae), International Journal of Odonatology, 14, 55 - 68. https: // doi. org / 10.1080 / 13887890.2011.568189" type="journal article" year="2011">
Zhang
<emphasis id="29AEEAA9E029D74EFCD8EE20FCADFDC5" box="[822,871,511,537]" italics="true" pageId="28" pageNumber="237">et al</emphasis>
. 2011
</bibRefCitation>
)
</taxonomicName>
. The same authors claimed that
<taxonomicName id="DCDA4D38E029D74EFAF3EDDFFA57FDC5" authorityName="Calvert" authorityYear="1913" box="[1309,1437,511,537]" class="Insecta" family="Megapodagrionidae" genus="Rhinagrion" kingdom="Animalia" order="Odonata" pageId="28" pageNumber="237" phylum="Arthropoda" rank="genus">
<emphasis id="29AEEAA9E029D74EFAF3EDDFFA57FDC5" box="[1309,1437,511,537]" italics="true" pageId="28" pageNumber="237">Rhinagrion</emphasis>
</taxonomicName>
lack a subocular comb on the gena, present in
<taxonomicName id="DCDA4D38E029D74EFD75EE03FCCCFDE1" box="[667,774,547,573]" class="Insecta" family="Megapodagrionidae" genus="Philosina" kingdom="Animalia" order="Odonata" pageId="28" pageNumber="237" phylum="Arthropoda" rank="genus">
<emphasis id="29AEEAA9E029D74EFD75EE03FCCCFDE1" box="[667,774,547,573]" italics="true" pageId="28" pageNumber="237">Philosina</emphasis>
</taxonomicName>
, but this was shown to be incorrect by
<bibRefCitation id="7F4B4B4AE029D74EFB52EE03FECFFDBD" author="Novelo-Gutierrez, R. &amp; Sites, R. W. &amp; Vitheepradit, A." pageId="28" pageNumber="237" pagination="562 - 568" refId="ref25601" refString="Novelo-Gutierrez, R., Sites, R. W. &amp; Vitheepradit, A. (2014) New province record of Rhinagrion for Thailand and description of the larva of R. mima (Odonata: Zygoptera: Philosinidae). Zootaxa, 5852 (5), 562 - 568. https: // doi. org / 10.11646 / zootaxa. 3852.5.4" type="journal article" year="2014">
Novelo-Gutiérrez
<emphasis id="29AEEAA9E029D74EFA66EE04FF67FDBD" italics="true" pageId="28" pageNumber="237">et al</emphasis>
. (2014)
</bibRefCitation>
in
<taxonomicName id="DCDA4D38E029D74EFEC9EE68FE26FDBD" authority="(Karsch)" baseAuthorityName="Karsch" box="[295,492,583,609]" class="Insecta" family="Megapodagrionidae" genus="Rhinagrion" kingdom="Animalia" order="Odonata" pageId="28" pageNumber="237" phylum="Arthropoda" rank="species" species="mima">
<emphasis id="29AEEAA9E029D74EFEC9EE68FE49FDBD" box="[295,387,584,609]" italics="true" pageId="28" pageNumber="237">R. mima</emphasis>
(Karsch)
</taxonomicName>
and a comb is also present in
<taxonomicName id="DCDA4D38E029D74EFCDBEE68FBD4FDBD" authority="(Selys)" baseAuthorityName="Selys" box="[821,1054,583,609]" class="Insecta" family="Megapodagrionidae" genus="Rhinagrion" kingdom="Animalia" order="Odonata" pageId="28" pageNumber="237" phylum="Arthropoda" rank="species" species="borneense">
<emphasis id="29AEEAA9E029D74EFCDBEE68FC0DFDBD" box="[821,967,583,609]" italics="true" pageId="28" pageNumber="237">R. borneense</emphasis>
(Selys)
</taxonomicName>
, (Orr, unpublished data). The only consistent differences between the genera appears to be relative leg-length, and the lateral caudal gills are relatively longer in
<taxonomicName id="DCDA4D38E029D74EFEECEEAFFEA7FD75" box="[258,365,655,681]" class="Insecta" family="Megapodagrionidae" genus="Philosina" kingdom="Animalia" order="Odonata" pageId="28" pageNumber="237" phylum="Arthropoda" rank="genus">
<emphasis id="29AEEAA9E029D74EFEECEEAFFEA7FD75" box="[258,365,655,681]" italics="true" pageId="28" pageNumber="237">Philosina</emphasis>
</taxonomicName>
and distinctly longer than the median one (
<figureCitation id="83E12A3EE029D74EFCBEEEAFFC1BFD75" box="[848,977,655,681]" captionStart="FIGURES 18–26. 18" captionStartId="6.[151,264,1811,1835]" captionTargetBox="[265,1336,167,1790]" captionTargetId="figure-21@6.[248,1339,153,1804]" captionTargetPageId="6" captionText="FIGURES 18–26. 18. Rhinagrion borneense, habitus (original drawing). 19. Philosina alba, habitus (modified from Zhang et al. 2011). 20. Rhinagrion mima, caudal gills: above heavily tracheated median gill; below right lateral gill with basal spines along mid-rib (modified from Lieftinck 1956). 21. Philosina alba caudal gills: above heavily tracheated median gill; below right lateral gill with basal spines along mid-rib (modified from Zhang et al. 2011) 22. Rhinagrion borneense, detail of mask (original drawing). 23. Philosina buchi, detail of mask (redrawn from Zhang et al. 2011). 24. Philoganga vetusta, habitus (adapted from Xu 2016). 25. Philoganga vetusta, details of caudal gills and antenna (redrawn from Xu 2016). 26. Philoganga montana, detail of mask (original drawing)." figureDoi="http://doi.org/10.5281/zenodo.13618331" httpUri="https://zenodo.org/record/13618331/files/figure.png" pageId="28" pageNumber="237">Figs 19, 21</figureCitation>
), whereas the median gill is only slightly shorter than the laterals in
<taxonomicName id="DCDA4D38E029D74EFE2DEE93FD89FD11" authorityName="Calvert" authorityYear="1913" box="[451,579,691,717]" class="Insecta" family="Megapodagrionidae" genus="Rhinagrion" kingdom="Animalia" order="Odonata" pageId="28" pageNumber="237" phylum="Arthropoda" rank="genus">
<emphasis id="29AEEAA9E029D74EFE2DEE93FD89FD11" box="[451,579,691,717]" italics="true" pageId="28" pageNumber="237">Rhinagrion</emphasis>
</taxonomicName>
(
<figureCitation id="83E12A3EE029D74EFDBCEE93FD1FFD11" box="[594,725,691,717]" captionStart="FIGURES 18–26. 18" captionStartId="6.[151,264,1811,1835]" captionTargetBox="[265,1336,167,1790]" captionTargetId="figure-21@6.[248,1339,153,1804]" captionTargetPageId="6" captionText="FIGURES 18–26. 18. Rhinagrion borneense, habitus (original drawing). 19. Philosina alba, habitus (modified from Zhang et al. 2011). 20. Rhinagrion mima, caudal gills: above heavily tracheated median gill; below right lateral gill with basal spines along mid-rib (modified from Lieftinck 1956). 21. Philosina alba caudal gills: above heavily tracheated median gill; below right lateral gill with basal spines along mid-rib (modified from Zhang et al. 2011) 22. Rhinagrion borneense, detail of mask (original drawing). 23. Philosina buchi, detail of mask (redrawn from Zhang et al. 2011). 24. Philoganga vetusta, habitus (adapted from Xu 2016). 25. Philoganga vetusta, details of caudal gills and antenna (redrawn from Xu 2016). 26. Philoganga montana, detail of mask (original drawing)." figureDoi="http://doi.org/10.5281/zenodo.13618331" httpUri="https://zenodo.org/record/13618331/files/figure.png" pageId="28" pageNumber="237">Figs 18, 20</figureCitation>
). Also, in
<taxonomicName id="DCDA4D38E029D74EFCA6EE93FC79FD11" box="[840,947,691,717]" class="Insecta" family="Megapodagrionidae" genus="Philosina" kingdom="Animalia" order="Odonata" pageId="28" pageNumber="237" phylum="Arthropoda" rank="genus">
<emphasis id="29AEEAA9E029D74EFCA6EE93FC79FD11" box="[840,947,691,717]" italics="true" pageId="28" pageNumber="237">Philosina</emphasis>
</taxonomicName>
there are three variably developed setae on the anterior labial palp (
<figureCitation id="83E12A3EE029D74EFE70EEF7FE3AFD2D" box="[414,496,727,753]" captionStart="FIGURES 18–26. 18" captionStartId="6.[151,264,1811,1835]" captionTargetBox="[265,1336,167,1790]" captionTargetId="figure-21@6.[248,1339,153,1804]" captionTargetPageId="6" captionText="FIGURES 18–26. 18. Rhinagrion borneense, habitus (original drawing). 19. Philosina alba, habitus (modified from Zhang et al. 2011). 20. Rhinagrion mima, caudal gills: above heavily tracheated median gill; below right lateral gill with basal spines along mid-rib (modified from Lieftinck 1956). 21. Philosina alba caudal gills: above heavily tracheated median gill; below right lateral gill with basal spines along mid-rib (modified from Zhang et al. 2011) 22. Rhinagrion borneense, detail of mask (original drawing). 23. Philosina buchi, detail of mask (redrawn from Zhang et al. 2011). 24. Philoganga vetusta, habitus (adapted from Xu 2016). 25. Philoganga vetusta, details of caudal gills and antenna (redrawn from Xu 2016). 26. Philoganga montana, detail of mask (original drawing)." figureDoi="http://doi.org/10.5281/zenodo.13618331" httpUri="https://zenodo.org/record/13618331/files/figure.png" pageId="28" pageNumber="237">Fig. 23</figureCitation>
), whereas in published accounts of
<taxonomicName id="DCDA4D38E029D74EFC99EEF7FC3DFD2D" authorityName="Calvert" authorityYear="1913" box="[887,1015,727,753]" class="Insecta" family="Megapodagrionidae" genus="Rhinagrion" kingdom="Animalia" order="Odonata" pageId="28" pageNumber="237" phylum="Arthropoda" rank="genus">
<emphasis id="29AEEAA9E029D74EFC99EEF7FC3DFD2D" box="[887,1015,727,753]" italics="true" pageId="28" pageNumber="237">Rhinagrion</emphasis>
</taxonomicName>
, there is only one, but in
<taxonomicName id="DCDA4D38E029D74EFAE5EEF8FA56FD2D" box="[1291,1436,727,753]" class="Insecta" family="Megapodagrionidae" genus="Rhinagrion" kingdom="Animalia" order="Odonata" pageId="28" pageNumber="237" phylum="Arthropoda" rank="species" species="borneense">
<emphasis id="29AEEAA9E029D74EFAE5EEF8FA56FD2D" box="[1291,1436,727,753]" italics="true" pageId="28" pageNumber="237">R. borneense</emphasis>
</taxonomicName>
(
<figureCitation id="83E12A3EE029D74EFF71EEDBFF3BFCC9" box="[159,241,763,789]" captionStart="FIGURES 18–26. 18" captionStartId="6.[151,264,1811,1835]" captionTargetBox="[265,1336,167,1790]" captionTargetId="figure-21@6.[248,1339,153,1804]" captionTargetPageId="6" captionText="FIGURES 18–26. 18. Rhinagrion borneense, habitus (original drawing). 19. Philosina alba, habitus (modified from Zhang et al. 2011). 20. Rhinagrion mima, caudal gills: above heavily tracheated median gill; below right lateral gill with basal spines along mid-rib (modified from Lieftinck 1956). 21. Philosina alba caudal gills: above heavily tracheated median gill; below right lateral gill with basal spines along mid-rib (modified from Zhang et al. 2011) 22. Rhinagrion borneense, detail of mask (original drawing). 23. Philosina buchi, detail of mask (redrawn from Zhang et al. 2011). 24. Philoganga vetusta, habitus (adapted from Xu 2016). 25. Philoganga vetusta, details of caudal gills and antenna (redrawn from Xu 2016). 26. Philoganga montana, detail of mask (original drawing)." figureDoi="http://doi.org/10.5281/zenodo.13618331" httpUri="https://zenodo.org/record/13618331/files/figure.png" pageId="28" pageNumber="237">Fig. 22</figureCitation>
) there are two strong setae (Orr, unpublished data) hence this character is considered unreliable.
</paragraph>
</subSubSection>
<subSubSection id="53C06530E029D74EFF79EF47FE8CFC5D" box="[151,326,871,897]" pageId="28" pageNumber="237" type="multiple">
<paragraph id="1B6536BBE029D74EFF79EF47FE8CFC5D" blockId="28.[151,326,871,897]" box="[151,326,871,897]" pageId="28" pageNumber="237">
<heading id="402D81D7E029D74EFF79EF47FE8CFC5D" bold="true" box="[151,326,871,897]" fontSize="11" level="1" pageId="28" pageNumber="237" reason="1">
<emphasis id="29AEEAA9E029D74EFF79EF47FE8CFC5D" bold="true" box="[151,326,871,897]" pageId="28" pageNumber="237">Key to genera</emphasis>
</heading>
</paragraph>
</subSubSection>
<subSubSection id="53C06530E029D74EFF79EF8CFA56FBAE" pageId="28" pageNumber="237" type="key">
<key id="6F4B618AE029D74EFF79EF8CFA56FBAE" pageId="28" pageNumber="237">
<keyStep id="A02E2E1EE029D74EFF79EF8CFA56FBAE" pageId="28" pageNumber="237">
<paragraph id="1B6536BBE029D74EFF79EF8CFA56FBC6" blockId="28.[151,1437,940,1138]" pageId="28" pageNumber="237">
<keyLead id="A02B958EE029D74EFF79EF8CFA56FBC6" pageId="28" pageNumber="237">
1 Legs very long; hind femur reaching abdominal S8–9; lateral caudal gill ovoid elongate, exceeding half length of abdomen; median caudal gill significantly shorter than lateral gills (
<figureCitation id="83E12A3EE029D74EFCE9EFE9FCBDFC03" box="[775,887,969,991]" captionStart="FIGURES 18–26. 18" captionStartId="6.[151,264,1811,1835]" captionTargetBox="[265,1336,167,1790]" captionTargetId="figure-21@6.[248,1339,153,1804]" captionTargetPageId="6" captionText="FIGURES 18–26. 18. Rhinagrion borneense, habitus (original drawing). 19. Philosina alba, habitus (modified from Zhang et al. 2011). 20. Rhinagrion mima, caudal gills: above heavily tracheated median gill; below right lateral gill with basal spines along mid-rib (modified from Lieftinck 1956). 21. Philosina alba caudal gills: above heavily tracheated median gill; below right lateral gill with basal spines along mid-rib (modified from Zhang et al. 2011) 22. Rhinagrion borneense, detail of mask (original drawing). 23. Philosina buchi, detail of mask (redrawn from Zhang et al. 2011). 24. Philoganga vetusta, habitus (adapted from Xu 2016). 25. Philoganga vetusta, details of caudal gills and antenna (redrawn from Xu 2016). 26. Philoganga montana, detail of mask (original drawing)." figureDoi="http://doi.org/10.5281/zenodo.13618331" httpUri="https://zenodo.org/record/13618331/files/figure.png" pageId="28" pageNumber="237">Figs 19, 21</figureCitation>
). Total length excluding caudal gills
<quantity id="DC229B5EE029D74EFB37EFEAFA8AFC03" box="[1241,1344,970,991]" metricMagnitude="-2" metricUnit="m" metricValue="1.7" metricValueMax="2.0" metricValueMin="1.4" pageId="28" pageNumber="237" unit="mm" value="17.0" valueMax="20.0" valueMin="14.0">14–20 mm</quantity>
...................................................................................................
<taxonomicName id="DCDA4D38E029D74EFAD0EFC6FA57FC20" box="[1342,1437,998,1020]" class="Insecta" family="Megapodagrionidae" genus="Philosina" kingdom="Animalia" order="Odonata" pageId="28" pageNumber="237" phylum="Arthropoda" rank="genus">
<emphasis id="29AEEAA9E029D74EFAD0EFC6FA57FC20" bold="true" box="[1342,1437,998,1020]" italics="true" pageId="28" pageNumber="237">Philosina</emphasis>
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[southern
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and northern Indochina]
</keyLead>
</paragraph>
<paragraph id="1B6536BBE029D74EFF79E802FA56FBAE" blockId="28.[151,1437,940,1138]" pageId="28" pageNumber="237">
<keyLead id="A02B958EE029D74EFF79E802FA56FBAE" pageId="28" pageNumber="237">
1’ Legs relatively shorter; hind femur reaching abdominal S6; lateral caudal gill squarish, nor exceeding half length of abdomen; median caudal gill slightly shorter than lateral gills (
<figureCitation id="83E12A3EE029D74EFD39E81FFC8DFB88" box="[727,839,1087,1109]" captionStart="FIGURES 18–26. 18" captionStartId="6.[151,264,1811,1835]" captionTargetBox="[265,1336,167,1790]" captionTargetId="figure-21@6.[248,1339,153,1804]" captionTargetPageId="6" captionText="FIGURES 18–26. 18. Rhinagrion borneense, habitus (original drawing). 19. Philosina alba, habitus (modified from Zhang et al. 2011). 20. Rhinagrion mima, caudal gills: above heavily tracheated median gill; below right lateral gill with basal spines along mid-rib (modified from Lieftinck 1956). 21. Philosina alba caudal gills: above heavily tracheated median gill; below right lateral gill with basal spines along mid-rib (modified from Zhang et al. 2011) 22. Rhinagrion borneense, detail of mask (original drawing). 23. Philosina buchi, detail of mask (redrawn from Zhang et al. 2011). 24. Philoganga vetusta, habitus (adapted from Xu 2016). 25. Philoganga vetusta, details of caudal gills and antenna (redrawn from Xu 2016). 26. Philoganga montana, detail of mask (original drawing)." figureDoi="http://doi.org/10.5281/zenodo.13618331" httpUri="https://zenodo.org/record/13618331/files/figure.png" pageId="28" pageNumber="237">Figs 18, 20</figureCitation>
). Total length excluding gills
<quantity id="DC229B5EE029D74EFB8AE81FFB01FB88" box="[1124,1227,1087,1108]" metricMagnitude="-2" metricUnit="m" metricValue="1.3" metricValueMax="1.4" metricValueMin="1.2" pageId="28" pageNumber="237" unit="mm" value="13.0" valueMax="14.0" valueMin="12.0">12–14 mm</quantity>
.......
<taxonomicName id="DCDA4D38E029D74EFAC7E81EFA56FB88" authorityName="Calvert" authorityYear="1913" box="[1321,1436,1086,1108]" class="Insecta" family="Megapodagrionidae" genus="Rhinagrion" kingdom="Animalia" order="Odonata" pageId="28" pageNumber="237" phylum="Arthropoda" rank="genus">
<emphasis id="29AEEAA9E029D74EFAC7E81EFA56FB88" bold="true" box="[1321,1436,1086,1108]" italics="true" pageId="28" pageNumber="237">Rhinagrion</emphasis>
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[Southern
<collectingCountry id="63CD762BE029D74EFBEBE87CFBF4FBAE" box="[1029,1086,1116,1138]" name="China" pageId="28" pageNumber="237">China</collectingCountry>
to
<collectingCountry id="63CD762BE029D74EFBB2E87CFB7BFBAE" box="[1116,1201,1116,1138]" name="Thailand" pageId="28" pageNumber="237">Thailand</collectingCountry>
and most of Sundaland]
</keyLead>
</paragraph>
</keyStep>
</key>
</subSubSection>
</treatment>
</document>

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Family
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Only larvae of the genus
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<emphasis id="29AEEAA9E029D74EFE5FEA5FFD82F945" box="[433,584,1663,1689]" italics="true" pageId="28" pageNumber="237">Rhipidolestes</emphasis>
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are known (
<figureCitation id="83E12A3EE029D74EFD3AEA5FFCECF945" box="[724,806,1663,1689]" captionStart="FIGURES 12–17. 12" captionStartId="5.[151,264,1903,1927]" captionTargetBox="[190,1400,160,1874]" captionTargetId="figure-15@5.[178,1409,151,1889]" captionTargetPageId="5" captionText="FIGURES 12–17. 12. Mesopodagrion tibetanum, habitus with detail of right profile of head (based on Yu 2016). 13. M. tibetanum, detail of mask (redrawn from Yu 2016). 14. Podolestes orientalis, habitus (from Choong &amp; Orr 2010). 15. Podolestes orientalis, detail of mask (from Choong &amp; Orr 2010). 16. Rhipidolestes aculeatus, habitus and detail of lateral and median gill (modified from Asahina 1994). 17. Rhipidolestes aculeatus, detail of mask (modified from Ishida 1996)." figureDoi="http://doi.org/10.5281/zenodo.13618327" httpUri="https://zenodo.org/record/13618327/files/figure.png" pageId="28" pageNumber="237">Fig. 16</figureCitation>
), from three species from southern Japanese islands and
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(Tabaru 1975,
<bibRefCitation id="7F4B4B4AE029D74EFE75EA83FDF6F961" author="Asahina, S." box="[411,572,1699,1725]" pageId="28" pageNumber="237" pagination="7 - 11" refId="ref21647" refString="Asahina, S. (1994) A Revision of the genus Rhipidolestes from Taiwan and Japan - A new species from Koshikijima Islands and descriptions of larval forms. Gekkan Mushi, 284, 7 - 11." type="journal article" year="1994">Asahina 1994</bibRefCitation>
,
<bibRefCitation id="7F4B4B4AE029D74EFDA4EA83FD1EF961" author="Ishida, K." box="[586,724,1699,1725]" pageId="28" pageNumber="237" refId="ref23597" refString="Ishida, K. (1996) Monograph of Odonata larvae in Japan. Hokkaido University Press, Sapporo, 446 pp. [in Japanese]" type="book" year="1996">Ishida 1996</bibRefCitation>
). Working in a small forested mountain stream in Kumamoto Prefecture, Kyushu, Tabaru (1975) determined that
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<emphasis id="29AEEAA9E029D74EFD0FEAE8FCA7F93D" box="[737,877,1735,1761]" italics="true" pageId="28" pageNumber="237">R. aculeatus</emphasis>
Ris
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(a species also present in Taiwan) completes 13 larval stadia with a three–four year developmental period. He observed: “The larvae are active but they seem to be nocturnal insect[s] taking small aquatic animals in the water. They avoid the1ight and hide themselves in the daytime under the bryophytes or among the plant leaves. They feign death...” The habitus of every instar is depicted photographically, but no detailed description is provided (although a later morphological description of the F larva is promised, but evidently did not eventuate).
</paragraph>
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The genus includes at least 23 species (undescribed species are illustrated in
<bibRefCitation id="7F4B4B4AE029D74EFBCDEBBFFB67F865" author="Zhang, H. - M." box="[1059,1197,1951,1977]" pageId="28" pageNumber="237" refId="ref28336" refString="Zhang, H. - M. (2019) Dragonflies and Damselflies of China. Vols. 1 &amp; 2. Chong Qing University Press, ChongQing, 14 + 1460 pp. [bilingual, in Chinese and English]" type="book" year="2019">Zhang 2019</bibRefCitation>
), a majority of which occur in the Oriental realm, but a significant number of which are confined to the Sino-Japanese realm; the exact numbers are difficult to establish owing to a lack of clarity in the northern boundary of the Oriental realm defined by
<bibRefCitation id="7F4B4B4AE028D74FFF54EC9BFE4AFF09" author="Holt, B. G. &amp; Lessard, J. - P. &amp; Borregaard, M. K. &amp; Fritz, S. A. &amp; Araujo, M. B. &amp; Dimitrov, D. &amp; Fabre, P. - H. &amp; Graham, C. H. &amp; Graves, G. R. &amp; Jonsson, K. A. &amp; Nogues-Bravo, D. &amp; Wang, Z. &amp; Whittaker, R. J. &amp; Fjeldsa, J. &amp; Rahbek, C." box="[186,384,187,213]" pageId="29" pageNumber="238" pagination="74 - 78" refId="ref23468" refString="Holt, B. G., Lessard, J. - P., Borregaard, M. K., Fritz, S. A., Araujo, M. B., Dimitrov, D., Fabre, P. - H., Graham, C. H., Graves, G. R., Jonsson, K. A., Nogues-Bravo, D., Wang, Z., Whittaker, R. J., Fjeldsa, J. &amp; Rahbek, C. (2013) An update of Wallace's zoogeographic regions of the world. Science, 339, 74 - 78. https: // doi. org / 10.1126 / science. 1228282" type="journal article" year="2013">
Holt
<emphasis id="29AEEAA9E028D74FFF1DEC9BFEEFFF09" box="[243,293,187,213]" italics="true" pageId="29" pageNumber="238">et al</emphasis>
. (2013)
</bibRefCitation>
. Elsewhere in the Oriental realm the genus ranges from northern
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and Indo-China to Southern
<collectingCountry id="63CD762BE028D74FFEEDECFFFE8CFF25" box="[259,326,223,249]" name="China" pageId="29" pageNumber="238">China</collectingCountry>
where the adults are found around small slow flowing shady forest streams and seepages from low altitudes to about
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, with species showing differing altitudinal preferences (
<bibRefCitation id="7F4B4B4AE028D74FFBDFED23FB0AFEC1" author="Zhang, H. - M." box="[1073,1216,259,285]" pageId="29" pageNumber="238" refId="ref28336" refString="Zhang, H. - M. (2019) Dragonflies and Damselflies of China. Vols. 1 &amp; 2. Chong Qing University Press, ChongQing, 14 + 1460 pp. [bilingual, in Chinese and English]" type="book" year="2019">Zhang 2019</bibRefCitation>
). The larval habitat is generally expected to be near the adult haunts. The larvae bear a superficial resemblance to those of
<taxonomicName id="DCDA4D38E028D74FFAFDED07FA53FE9D" baseAuthorityName="Jacobson &amp; Bianchi" baseAuthorityYear="1905" box="[1299,1433,295,321]" class="Insecta" family="Euphaeidae" kingdom="Animalia" order="Odonata" pageId="29" pageNumber="238" phylum="Arthropoda" rank="family">Euphaeidae</taxonomicName>
, especially
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<emphasis id="29AEEAA9E028D74FFEFEED6BFEB7FEB9" box="[272,381,331,357]" italics="true" pageId="29" pageNumber="238">Bayadera</emphasis>
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but are smaller and are immediately recognised by the lack of ventral abdominal gills. The antennae are relatively short and stout, especially in the basal segments. The prementum is broad and short with a moderately produced, rounded anterior median lobe with a moderately developed median cleft; the labial palps are short (
<figureCitation id="83E12A3EE028D74FFF30ED97FEFBFE0D" box="[222,305,439,465]" captionStart="FIGURES 12–17. 12" captionStartId="5.[151,264,1903,1927]" captionTargetBox="[190,1400,160,1874]" captionTargetId="figure-15@5.[178,1409,151,1889]" captionTargetPageId="5" captionText="FIGURES 12–17. 12. Mesopodagrion tibetanum, habitus with detail of right profile of head (based on Yu 2016). 13. M. tibetanum, detail of mask (redrawn from Yu 2016). 14. Podolestes orientalis, habitus (from Choong &amp; Orr 2010). 15. Podolestes orientalis, detail of mask (from Choong &amp; Orr 2010). 16. Rhipidolestes aculeatus, habitus and detail of lateral and median gill (modified from Asahina 1994). 17. Rhipidolestes aculeatus, detail of mask (modified from Ishida 1996)." figureDoi="http://doi.org/10.5281/zenodo.13618327" httpUri="https://zenodo.org/record/13618327/files/figure.png" pageId="29" pageNumber="238">Fig. 17</figureCitation>
). As
<bibRefCitation id="7F4B4B4AE028D74FFE84ED97FD8BFE0D" author="Bybee, S. M. &amp; Kalkman, V. J. &amp; Erickson, R. J. &amp; Breinholt, J. W. &amp; Suvorov, A. &amp; Dijkstra, K-D. B. &amp; Cordero-Riviera, A. &amp; Skevington, J. H. &amp; Abbott, J. C. &amp; Sanchez Herrera, M. &amp; Lemmon, A. R. &amp; Moriaty Lemmon, E. &amp; Ware, J. L." box="[362,577,439,465]" pageId="29" pageNumber="238" pagination="1 - 15" refId="ref22012" refString="Bybee, S. M., Kalkman, V. J., Erickson, R. J., Breinholt, J. W., Suvorov, A., Dijkstra, K-D. B., Cordero-Riviera, A., Skevington, J. H., Abbott, J. C., Sanchez Herrera, M., Lemmon, A. R., Moriaty Lemmon, E. &amp; Ware, J. L. (2021) Phylogeny and classification of Odonata using targeted genomics. Molecular Phylogenetics and Evolution, 160, 1 - 15. https: // doi. org / 10.1016 / j. ympev. 2021.107115" type="journal article" year="2021">
Bybee
<emphasis id="29AEEAA9E028D74FFE56ED97FE23FE0D" box="[440,489,439,465]" italics="true" pageId="29" pageNumber="238">et al</emphasis>
. (2021)
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were not fully confident in the composition of the family it is possible the as yet unknown larvae of other genera differ significantly.
</paragraph>
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Unknown larvae:
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<emphasis id="29AEEAA9E028D74FFE7EEDDFFDEDFDC5" box="[400,551,511,537]" italics="true" pageId="29" pageNumber="238">Agriomorpha</emphasis>
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(2 spp.),
<taxonomicName id="DCDA4D38E028D74FFD7FEDDFFC83FDC5" authorityName="Kennedy" authorityYear="1925" box="[657,841,511,537]" class="Insecta" family="Megapodagrionidae" genus="Burmargiolestes" kingdom="Animalia" order="Odonata" pageId="29" pageNumber="238" phylum="Arthropoda" rank="genus">
<emphasis id="29AEEAA9E028D74FFD7FEDDFFC83FDC5" box="[657,841,511,537]" italics="true" pageId="29" pageNumber="238">Burmargiolestes</emphasis>
</taxonomicName>
(2 spp.),
<taxonomicName id="DCDA4D38E028D74FFC5AEDDFFBACFDC5" authorityName="Kimmins" authorityYear="1936" box="[948,1126,511,537]" class="Insecta" family="Megapodagrionidae" genus="Bornargiolestes" kingdom="Animalia" order="Odonata" pageId="29" pageNumber="238" phylum="Arthropoda" rank="genus">
<emphasis id="29AEEAA9E028D74FFC5AEDDFFBACFDC5" box="[948,1126,511,537]" italics="true" pageId="29" pageNumber="238">Bornargiolestes</emphasis>
</taxonomicName>
(3 spp.)
</paragraph>
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The genus is known from three species occurring in northern Indochina,
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and southern
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. The larva of one species,
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, originally placed in
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, has been confirmed by breeding and is presently being described (T.S. Keetapithchayakul pers comm.). I refrain from pre-empting the results of that study. However it may be noted that this species does share clear synapomorphies with
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spp.
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, which it resembles closely, but is nevertheless distinct from known members of that genus.
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<taxonomicName id="DCDA4D38E02CD74BFF79E986FEC3FA1D" ID-CoL="496W" authorityName="Selys" authorityYear="1853" box="[151,265,1446,1473]" class="Insecta" family="Euphaeidae" genus="Dysphaea" kingdom="Animalia" order="Odonata" pageId="25" pageNumber="234" phylum="Arthropoda" rank="genus">
<emphasis id="29AEEAA9E02CD74BFF79E986FEC3FA1D" bold="true" box="[151,265,1446,1473]" italics="true" pageId="25" pageNumber="234">Dysphaea</emphasis>
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The genus is represented by nine species distributed widely from Sundaland to southern
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, including
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, Indochina.
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,
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across north-west
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,
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and the eastern Himalayas to the Western Ghats. Larva of this common genus have proved remarkably elusive.
<bibRefCitation id="7F4B4B4AE02CD74BFCAAEA33FC31F9F1" author="Lieftinck, M. A." box="[836,1019,1554,1581]" pageId="25" pageNumber="234" pagination="657 - 716" refId="ref24498" refString="Lieftinck, M. A. (1950) Further studies on Southeast Asiatic species of Macromia Rambur, with notes on their ecology, habits and life history, and with descriptions of larvae and two new species. (Odon., Epophthalmiinae). Treubia, 20 (3), 657 - 716." type="journal article" year="1950">Lieftinck (1950)</bibRefCitation>
recorded larvae of
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<emphasis id="29AEEAA9E02CD74BFB20EA33FA90F9F1" box="[1230,1370,1555,1581]" italics="true" pageId="25" pageNumber="234">D. dimidiata</emphasis>
Selys
</taxonomicName>
from West Java and noted their ecology in general terms, but surprisingly provided no description or even a general comment of their unusual morphology, either in that publication or in any later paper. Recently larvae and exuviae of
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<emphasis id="29AEEAA9E02CD74BFF59EA5FFE41F945" box="[183,395,1663,1689]" italics="true" pageId="25" pageNumber="234">Dysphaea gloriosa</emphasis>
Fraser
</taxonomicName>
(
<figureCitation id="83E12A3EE02CD74BFE04EA5FFDF7F945" box="[490,573,1662,1689]" captionStart="FIGURES 57–60. 57" captionStartId="11.[151,264,1933,1957]" captionTargetBox="[193,1384,155,1907]" captionTargetId="figure-15@11.[188,1400,151,1917]" captionTargetPageId="11" captionText="FIGURES 57–60. 57. Dysphaea gloriosa, habitus (drawn from Nguyen et al. 2024). 58. Bayadera strigata, habitus (drawn from Yang &amp; Orr 2024). 59. Euphaea decorata, habitus. (drawn from Xu 2017). 60. Anisopleura furcata, habitus (drawn from Keetapithchayakul et al. 2024)." figureDoi="http://doi.org/10.5281/zenodo.13618341" httpUri="https://zenodo.org/record/13618341/files/figure.png" pageId="25" pageNumber="234">Fig. 57</figureCitation>
) were discovered in
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and described (
<bibRefCitation id="7F4B4B4AE02CD74BFBAFEA5FFAEBF945" author="Nguyen, M. T. &amp; Ignatius, M. J. &amp; Sunprasit, S. &amp; Phan, Q. T. &amp; Keetapithchayakul, T. S." box="[1089,1313,1663,1689]" pageId="25" pageNumber="234" pagination="385 - 396" refId="ref25439" refString="Nguyen, M. T., Ignatius, M. J., Sunprasit, S., Phan, Q. T. &amp; Keetapithchayakul, T. S. (2024) Description of final stadium larva of Dysphaea gloriosa Fraser, 1938, (Odonata: Euphaeidae) with notes on its habitat and biology. Zootaxa, 5447 (3), 385 - 396. https: // doi. org / 10.11646 / zootaxa. 5447.3.4" type="journal article" year="2024">
Nguyen
<emphasis id="29AEEAA9E02CD74BFB4DEA5FFB1FF945" box="[1187,1237,1663,1689]" italics="true" pageId="25" pageNumber="234">et al</emphasis>
. 2024
</bibRefCitation>
). The head is extremely large, much wider than the prothorax, with small eyes; the genae are swollen beyond the outer margin of the eyes and armed with rows of heavy spines The postocular lobes are bulbous and also bear strong marginal spines (
<figureCitation id="83E12A3EE02CD74BFF03EACBFE8AF8D9" box="[237,320,1771,1797]" captionStart="FIGURES 61–66. 61" captionStartId="12.[151,264,1841,1865]" captionTargetBox="[273,1314,156,1828]" captionTargetId="figure-21@12.[268,1319,151,1834]" captionTargetPageId="12" captionText="FIGURES 61–66. 61. Dysphaea gloriosa: detail of spines on genae and postocular lobe; underside (left), upperside (right) (modified from Nguyen et al. 2024). 62. Heterophaea barbata: dorsal view of head, showing (a) outer mandibular spines (b) genal spines partially visible and (c) right antenna (from Orr &amp; Hämäläinen 2024). 63. Anisopleura subplatystyla, dorsal view of head (original drawing). 64. Bayadera strigata, dorsal view of head (drawn from Yang &amp; Orr 2024). 65. Euphaea decorata, dorsal view of head. (drawn from Xu 2017). 66. Bayadera strigata, left labial palp with detail showing wavy outer margin of outer process of palpal lobe (arrowed), (drawn from Yang &amp; Orr 2024)." figureDoi="http://doi.org/10.5281/zenodo.13618343" httpUri="https://zenodo.org/record/13618343/files/figure.png" pageId="25" pageNumber="234">Fig. 61</figureCitation>
). The saccoid gills are elongate and tapered to a long fine filament. The genus perhaps has the most distinctive of all euphaeid larvae and it is expected that all species will be recognisable.
</paragraph>
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<emphasis id="29AEEAA9E02CD74BFF29EB13FEFDF891" box="[199,311,1843,1869]" italics="true" pageId="25" pageNumber="234">Dysphaea</emphasis>
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have been observed ovipositing in areas of torrential water flow (
<bibRefCitation id="7F4B4B4AE02CD74BFBC6EB13FB5FF891" author="Orr, A. G." box="[1064,1173,1843,1869]" pageId="25" pageNumber="234" pagination="167 - 220" refId="ref25667" refString="Orr, A. G. (2001) An annotated checklist of the Odonata of Brunei with ecological notes and descriptions of hitherto unknown males and larvae. International Journal of Odonatology, 4, 167 - 220. https: // doi. org / 10.1080 / 13887890.2001.9748168" type="journal article" year="2001">Orr 2001</bibRefCitation>
). The larvae have been collected clinging under rocks in swift flowing rapids (
<bibRefCitation id="7F4B4B4AE02CD74BFD13EB77FC62F8AD" author="Lieftinck, M. A." box="[765,936,1878,1905]" pageId="25" pageNumber="234" pagination="657 - 716" refId="ref24498" refString="Lieftinck, M. A. (1950) Further studies on Southeast Asiatic species of Macromia Rambur, with notes on their ecology, habits and life history, and with descriptions of larvae and two new species. (Odon., Epophthalmiinae). Treubia, 20 (3), 657 - 716." type="journal article" year="1950">Lieftinck 1950</bibRefCitation>
,
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Nguyen
<emphasis id="29AEEAA9E02CD74BFBFCEB77FB8EF8AD" box="[1042,1092,1879,1905]" italics="true" pageId="25" pageNumber="234">et al</emphasis>
. 2024
</bibRefCitation>
).
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<emphasis id="29AEEAA9E02DD74AFF79ECFEFEEFFF25" bold="true" box="[151,293,222,249]" italics="true" pageId="24" pageNumber="233">Anisopleura</emphasis>
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This largely montane genus is confined to northern parts of the Oriental realm where eleven species are recognised from the eastern Himalayas, through northern
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The first account of larvae the genus was provided by
<bibRefCitation id="7F4B4B4AE02DD74AFCCFED6AFC16FEB8" author="Needham, J. G." box="[801,988,330,356]" pageId="24" pageNumber="233" pagination="145 - 154" refId="ref24959" refString="Needham, J. G. (1911) Descriptions of dragonfly nymphs of the subfamily Calopteryginae (Odonata). Entomological News, 22, 145 - 154." type="journal article" year="1911">Needham (1911)</bibRefCitation>
who described and illustrated presumed
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<emphasis id="29AEEAA9E02DD74AFF79ED4DFEA4FE5B" box="[151,366,365,391]" italics="true" pageId="24" pageNumber="233">Anisopleura comes</emphasis>
Hagen
</taxonomicName>
, his identification being based on examination of the venation of the as yet unexpanded wing and is consistent with present knowledge. Details of the habitus and labial palp are figured and this information was largely repeated in
<bibRefCitation id="7F4B4B4AE02DD74AFE49ED95FDA2FE13" author="Needham, J. G." box="[423,616,437,463]" pageId="24" pageNumber="233" pagination="1 - 20" refId="ref24989" refString="Needham, J. G. (1930) A manual of the dragonflies of China: A monographic study of the Chinese Odonata. [Zoologia Sinica, Series A. Invertebrates of China, Vol. 11. Fascicle 1]. The Fan Memorial Institute of Biology, Peiping, xi + 344 pp., pls. 1 - 20 excl." type="book chapter" year="1930">Needham (1930)</bibRefCitation>
.
<bibRefCitation id="7F4B4B4AE02DD74AFD9AED95FCD0FE13" author="Fraser, F. C." box="[628,794,437,463]" pageId="24" pageNumber="233" pagination="47 - 59" refId="ref22935" refString="Fraser, F. C. (1929 a) Indian dragonflies, Part XXXI. Journal of the Bombay Natural History Society, 33, 47 - 59, pls. 1 - 3 excl." type="journal article" year="1929">Fraser (1929a)</bibRefCitation>
figured the habitus and mask of
<taxonomicName id="DCDA4D38E02DD74AFB7AED95FA52FE13" authority="Fraser" authorityName="Fraser" authorityYear="1927" box="[1172,1432,437,463]" class="Insecta" family="Euphaeidae" genus="Anisopleura" kingdom="Animalia" order="Odonata" pageId="24" pageNumber="233" phylum="Arthropoda" rank="species" species="subplatystyla">
<emphasis id="29AEEAA9E02DD74AFB7AED95FA82FE13" box="[1172,1352,437,463]" italics="true" pageId="24" pageNumber="233">A. subplatystyla</emphasis>
Fraser
</taxonomicName>
, both apparently fairly accurately, but he unaccountably noted in the legend a lack of ventral abdominal gills, which is quite wrong.
<bibRefCitation id="7F4B4B4AE02DD74AFEA8EDDDFDABFDCB" author="Kumar, A. &amp; Prasad, M." box="[326,609,508,535]" pageId="24" pageNumber="233" pagination="225 - 230" refId="ref24294" refString="Kumar, A. &amp; Prasad, M. (1977 a) Last instar larvae of two Odonata species from Western Himalayas. Entomon, 2, 225 - 230." type="journal article" year="1977">Kumar &amp; Prasad (1977a)</bibRefCitation>
describe and figured the larva of
<taxonomicName id="DCDA4D38E02DD74AFC32EDDDFB6BFDCA" authority="Selys" authorityName="Selys" authorityYear="1853" box="[988,1185,508,535]" class="Insecta" family="Euphaeidae" genus="Anisopleura" kingdom="Animalia" order="Odonata" pageId="24" pageNumber="233" phylum="Arthropoda" rank="species" species="lestoides">
<emphasis id="29AEEAA9E02DD74AFC32EDDDFB97FDCA" box="[988,1117,508,535]" italics="true" pageId="24" pageNumber="233">A. lestoides</emphasis>
Selys
</taxonomicName>
in detail and although the artwork is somewhat stylised it appears accurate in essentials.
<bibRefCitation id="7F4B4B4AE02DD74AFC60EE00FB3DFDE6" author="Keetapithchayakul, T. S. &amp; Orr, A. G. &amp; Wongkamhaeng, K." box="[910,1271,544,571]" pageId="24" pageNumber="233" pagination="529 - 542" refId="ref24083" refString="Keetapithchayakul, T. S., Orr, A. G. &amp; Wongkamhaeng, K. (2024) Description of the final stadium larva of Anisopleura furcata Selys, 1891 (Odonata: Euphaeidae) from Thailand. Zootaxa, 5415 (4), 529 - 542. https: // doi. org / 10.11646 / zootaxa. 5415.4.2" type="journal article" year="2024">
Keetapithchayakul
<emphasis id="29AEEAA9E02DD74AFB87EE01FB56FDE6" box="[1129,1180,544,570]" italics="true" pageId="24" pageNumber="233">et al</emphasis>
. (2024)
</bibRefCitation>
described and figured the rather atypical
<taxonomicName id="DCDA4D38E02DD74AFE2DEE64FDBDFD82" authority="Selys" authorityName="Selys" authorityYear="1891" box="[451,631,580,606]" class="Insecta" family="Euphaeidae" genus="Anisopleura" kingdom="Animalia" order="Odonata" pageId="24" pageNumber="233" phylum="Arthropoda" rank="species" species="furcata">
<emphasis id="29AEEAA9E02DD74AFE2DEE64FDF9FD82" box="[451,563,580,606]" italics="true" pageId="24" pageNumber="233">A. furcata</emphasis>
Selys
</taxonomicName>
(
<figureCitation id="83E12A3EE02DD74AFD69EE64FD16FD82" box="[647,732,580,606]" captionStart="FIGURES 57–60. 57" captionStartId="11.[151,264,1933,1957]" captionTargetBox="[193,1384,155,1907]" captionTargetId="figure-15@11.[188,1400,151,1917]" captionTargetPageId="11" captionText="FIGURES 57–60. 57. Dysphaea gloriosa, habitus (drawn from Nguyen et al. 2024). 58. Bayadera strigata, habitus (drawn from Yang &amp; Orr 2024). 59. Euphaea decorata, habitus. (drawn from Xu 2017). 60. Anisopleura furcata, habitus (drawn from Keetapithchayakul et al. 2024)." figureDoi="http://doi.org/10.5281/zenodo.13618341" httpUri="https://zenodo.org/record/13618341/files/figure.png" pageId="24" pageNumber="233">Fig. 60</figureCitation>
). This was the first modern, detailed description and depiction of a larva of the genus.
</paragraph>
<paragraph id="1B6536BBE02DD74AFF29EEACFB07FB8F" blockId="24.[151,1437,222,1357]" pageId="24" pageNumber="233">
Larvae of the genus are distinguished by several characters; the antennae are relatively short, being shorter than the length of the head from the occipital margin to the base of the labrum (
<figureCitation id="83E12A3EE02DD74AFBD6EE8FFB44FD15" box="[1080,1166,687,713]" captionStart="FIGURES 61–66. 61" captionStartId="12.[151,264,1841,1865]" captionTargetBox="[273,1314,156,1828]" captionTargetId="figure-21@12.[268,1319,151,1834]" captionTargetPageId="12" captionText="FIGURES 61–66. 61. Dysphaea gloriosa: detail of spines on genae and postocular lobe; underside (left), upperside (right) (modified from Nguyen et al. 2024). 62. Heterophaea barbata: dorsal view of head, showing (a) outer mandibular spines (b) genal spines partially visible and (c) right antenna (from Orr &amp; Hämäläinen 2024). 63. Anisopleura subplatystyla, dorsal view of head (original drawing). 64. Bayadera strigata, dorsal view of head (drawn from Yang &amp; Orr 2024). 65. Euphaea decorata, dorsal view of head. (drawn from Xu 2017). 66. Bayadera strigata, left labial palp with detail showing wavy outer margin of outer process of palpal lobe (arrowed), (drawn from Yang &amp; Orr 2024)." figureDoi="http://doi.org/10.5281/zenodo.13618343" httpUri="https://zenodo.org/record/13618343/files/figure.png" pageId="24" pageNumber="233">Fig. 63</figureCitation>
); the labrum, frons and vertex typically bear numerous small low tubercles, sparse or absent in other
<taxonomicName id="DCDA4D38E02DD74AFC10EEF3FBA8FD31" authorityName="Selys" authorityYear="1840" box="[1022,1122,723,749]" class="Insecta" family="Euphaeidae" genus="Euphaea" kingdom="Animalia" order="Odonata" pageId="24" pageNumber="233" phylum="Arthropoda" rank="genus">
<emphasis id="29AEEAA9E02DD74AFC10EEF3FBA8FD31" box="[1022,1122,723,749]" italics="true" pageId="24" pageNumber="233">Euphaea</emphasis>
</taxonomicName>
and
<taxonomicName id="DCDA4D38E02DD74AFB77EEF3FACCFD31" authorityName="Selys" authorityYear="1853" box="[1177,1286,723,749]" class="Insecta" family="Euphaeidae" genus="Bayadera" kingdom="Animalia" order="Odonata" pageId="24" pageNumber="233" phylum="Arthropoda" rank="genus">
<emphasis id="29AEEAA9E02DD74AFB77EEF3FACCFD31" box="[1177,1286,723,749]" italics="true" pageId="24" pageNumber="233">Bayadera</emphasis>
</taxonomicName>
, which, in
<taxonomicName id="DCDA4D38E02DD74AFA6AEEF3FF2DFCCD" authorityName="Selys" authorityYear="1891" class="Insecta" family="Euphaeidae" genus="Anisopleura" kingdom="Animalia" order="Odonata" pageId="24" pageNumber="233" phylum="Arthropoda" rank="species" species="furcata">
<emphasis id="29AEEAA9E02DD74AFA6AEEF3FF2DFCCD" italics="true" pageId="24" pageNumber="233">A. furcata</emphasis>
</taxonomicName>
are elongated giving the head a strongly warty appearance. Similar but differently arranged structures are also present on a larva suspected to be
<taxonomicName id="DCDA4D38E02DD74AFDABEF3BFC74FCE8" authority="Hamalainen &amp; Karube" authorityName="Hamalainen &amp; Karube" box="[581,958,794,821]" class="Insecta" family="Euphaeidae" genus="Anisopleura" kingdom="Animalia" order="Odonata" pageId="24" pageNumber="233" phylum="Arthropoda" rank="species" species="bipugio">
<emphasis id="29AEEAA9E02DD74AFDABEF3BFD73FCE8" box="[581,697,794,821]" italics="true" pageId="24" pageNumber="233">A. bipugio</emphasis>
Hämäläinen &amp; Karube
</taxonomicName>
, the probable nearest relative of
<taxonomicName id="DCDA4D38E02DD74AFAC9EF3BFA5FFCE8" authorityName="Selys" authorityYear="1891" box="[1319,1429,794,821]" class="Insecta" family="Euphaeidae" genus="Anisopleura" kingdom="Animalia" order="Odonata" pageId="24" pageNumber="233" phylum="Arthropoda" rank="species" species="furcata">
<emphasis id="29AEEAA9E02DD74AFAC9EF3BFA5FFCE8" box="[1319,1429,794,821]" italics="true" pageId="24" pageNumber="233">A. furcata</emphasis>
</taxonomicName>
, but otherwise the tubercles in this species show a degree of development unique in the genus as far as is known. In known species the outer edge of the mandibles bears one or two long spurs, as opposed to more numerous groups of smaller teeth in
<taxonomicName id="DCDA4D38E02DD74AFE9CEFA5FE1CFC43" authorityName="Selys" authorityYear="1840" box="[370,470,901,927]" class="Insecta" family="Euphaeidae" genus="Euphaea" kingdom="Animalia" order="Odonata" pageId="24" pageNumber="233" phylum="Arthropoda" rank="genus">
<emphasis id="29AEEAA9E02DD74AFE9CEFA5FE1CFC43" box="[370,470,901,927]" italics="true" pageId="24" pageNumber="233">Euphaea</emphasis>
</taxonomicName>
and
<emphasis id="29AEEAA9E02DD74AFDFEEFA5FD4FFC43" box="[528,645,901,927]" italics="true" pageId="24" pageNumber="233">
<taxonomicName id="DCDA4D38E02DD74AFDFEEFA5FD4BFC43" authorityName="Selys" authorityYear="1853" box="[528,641,901,927]" class="Insecta" family="Euphaeidae" genus="Bayadera" kingdom="Animalia" order="Odonata" pageId="24" pageNumber="233" phylum="Arthropoda" rank="genus">Bayadera</taxonomicName>
.
</emphasis>
The inner apical process on the labial palp is short and adpressed or strongly reduced (
<figureCitation id="83E12A3EE02DD74AFE89EF89FE77FC1F" box="[359,445,937,963]" captionStart="FIGURES 67–74. 67" captionStartId="13.[151,264,1814,1838]" captionTargetBox="[256,1340,160,1799]" captionTargetId="figure-15@13.[246,1342,151,1807]" captionTargetPageId="13" captionText="FIGURES 67–74. 67. Anisopleura subplatystyla, caudal gills dorsal view (original drawing). 68. Bayadera strigata, caudal gills dorsal view, (drawn from Yang &amp; Orr 2024). 69. Euphaea decorata, caudal gills dorsal view (drawn from Xu 2017). 70. Euphaea superba, caudal gills dorsal view (redrawn from Wu et al. 2017). 71. Bayadera strigata: male genital apophyses; ventral (left), right lateral (right), (drawn from Yang &amp; Orr 2024). 72. Euphaea decorata, male genital apophyses; ventral view (drawn from Xu 2017). 73. Euphaea superba, terminal processes of palpal lobe ventral view (movable hook truncated), (redrawn from Wu et al. 2017). 74. Anisopleura furcata, left labial palp showing terminal processes of palpal lobe, dorsal view (redrawn from Keetapithchayakul et al. 2024)." figureDoi="http://doi.org/10.5281/zenodo.13618345" httpUri="https://zenodo.org/record/13618345/files/figure.png" pageId="24" pageNumber="233">Fig. 74</figureCitation>
), a condition rare in other genera. The postocular lobes are shallow and the habitus is comparatively slender, tapering caudad (
<figureCitation id="83E12A3EE02DD74AFD8FEFEDFD7FFC3B" box="[609,693,973,999]" captionStart="FIGURES 61–66. 61" captionStartId="12.[151,264,1841,1865]" captionTargetBox="[273,1314,156,1828]" captionTargetId="figure-21@12.[268,1319,151,1834]" captionTargetPageId="12" captionText="FIGURES 61–66. 61. Dysphaea gloriosa: detail of spines on genae and postocular lobe; underside (left), upperside (right) (modified from Nguyen et al. 2024). 62. Heterophaea barbata: dorsal view of head, showing (a) outer mandibular spines (b) genal spines partially visible and (c) right antenna (from Orr &amp; Hämäläinen 2024). 63. Anisopleura subplatystyla, dorsal view of head (original drawing). 64. Bayadera strigata, dorsal view of head (drawn from Yang &amp; Orr 2024). 65. Euphaea decorata, dorsal view of head. (drawn from Xu 2017). 66. Bayadera strigata, left labial palp with detail showing wavy outer margin of outer process of palpal lobe (arrowed), (drawn from Yang &amp; Orr 2024)." figureDoi="http://doi.org/10.5281/zenodo.13618343" httpUri="https://zenodo.org/record/13618343/files/figure.png" pageId="24" pageNumber="233">Fig. 63</figureCitation>
). There are 2–4 strong spines on the genae below the eye margin. The caudal gills are elongate and evenly tapered to a long filament (a condition not known in
<taxonomicName id="DCDA4D38E02DD74AFB32EFD1FA83FBD7" authorityName="Selys" authorityYear="1853" box="[1244,1353,1009,1035]" class="Insecta" family="Euphaeidae" genus="Bayadera" kingdom="Animalia" order="Odonata" pageId="24" pageNumber="233" phylum="Arthropoda" rank="genus">
<emphasis id="29AEEAA9E02DD74AFB32EFD1FA83FBD7" box="[1244,1353,1009,1035]" italics="true" pageId="24" pageNumber="233">Bayadera</emphasis>
</taxonomicName>
). Male genital apophyses (known in detail only in
<taxonomicName id="DCDA4D38E02DD74AFD62E835FD37FBF3" authorityName="Selys" authorityYear="1891" box="[652,765,1045,1071]" class="Insecta" family="Euphaeidae" genus="Anisopleura" kingdom="Animalia" order="Odonata" pageId="24" pageNumber="233" phylum="Arthropoda" rank="species" species="furcata">
<emphasis id="29AEEAA9E02DD74AFD62E835FD37FBF3" box="[652,765,1045,1071]" italics="true" pageId="24" pageNumber="233">A. furcata</emphasis>
</taxonomicName>
) are moderately well developed, serrate ventrally and just overlying S10, but larger than in
<taxonomicName id="DCDA4D38E02DD74AFDE4E818FDA4FB8E" authorityName="Selys" authorityYear="1840" box="[522,622,1080,1106]" class="Insecta" family="Euphaeidae" genus="Euphaea" kingdom="Animalia" order="Odonata" pageId="24" pageNumber="233" phylum="Arthropoda" rank="genus">
<emphasis id="29AEEAA9E02DD74AFDE4E818FDA4FB8E" box="[522,622,1080,1106]" italics="true" pageId="24" pageNumber="233">Euphaea</emphasis>
</taxonomicName>
and not large and sausage like as in known
<taxonomicName id="DCDA4D38E02DD74AFBB4E818FB0DFB8E" authorityName="Selys" authorityYear="1853" box="[1114,1223,1080,1106]" class="Insecta" family="Euphaeidae" genus="Bayadera" kingdom="Animalia" order="Odonata" pageId="24" pageNumber="233" phylum="Arthropoda" rank="genus">
<emphasis id="29AEEAA9E02DD74AFBB4E818FB0DFB8E" box="[1114,1223,1080,1106]" italics="true" pageId="24" pageNumber="233">Bayadera</emphasis>
</taxonomicName>
.
</paragraph>
<paragraph id="1B6536BBE02DD74AFF29E87CFC47FA90" blockId="24.[151,1437,222,1357]" pageId="24" pageNumber="233">
Larvae of
<taxonomicName id="DCDA4D38E02DD74AFED8E87CFE75FBAA" authorityName="Selys" authorityYear="1853" box="[310,447,1116,1142]" class="Insecta" family="Euphaeidae" genus="Anisopleura" kingdom="Animalia" order="Odonata" pageId="24" pageNumber="233" phylum="Arthropoda" rank="genus">
<emphasis id="29AEEAA9E02DD74AFED8E87CFE75FBAA" box="[310,447,1116,1142]" italics="true" pageId="24" pageNumber="233">Anisopleura</emphasis>
</taxonomicName>
have been found in a variety of small stream habitats from
<quantity id="DC229B5EE02DD74AFBADE87CFB03FBAA" box="[1091,1225,1116,1142]" metricMagnitude="3" metricUnit="m" metricValue="1.25" metricValueMax="1.6" metricValueMin="0.9" pageId="24" pageNumber="233" unit="m" value="1250.0" valueMax="1600.0" valueMin="900.0">900–1600m</quantity>
in
<collectingCountry id="63CD762BE02DD74AFB07E87CFAF0FBAA" box="[1257,1338,1116,1142]" name="Bhutan" pageId="24" pageNumber="233">Bhutan</collectingCountry>
(
<bibRefCitation id="7F4B4B4AE02DD74AFAA8E87CFEC5FB46" author="Rasaily, B. &amp; Kalkman, V. J. &amp; Katel, O. &amp; Dorji, C." pageId="24" pageNumber="233" pagination="1 - 20" refId="ref26410" refString="Rasaily, B., Kalkman, V. J., Katel, O. &amp; Dorji, C. (2021) Composition of the dragonfly fauna at different altitudes in Bhutan based on larval samples. IDF-Report, 160, 1 - 20." type="journal article" year="2021">
Rasaily
<emphasis id="29AEEAA9E02DD74AFF79E8A0FF02FB46" box="[151,200,1152,1178]" italics="true" pageId="24" pageNumber="233">et al</emphasis>
. 2021
</bibRefCitation>
) including open, disturbed places with dense ground vegetation; those of
<taxonomicName id="DCDA4D38E02DD74AFBA9E8A0FB7CFB46" authorityName="Selys" authorityYear="1891" box="[1095,1206,1152,1178]" class="Insecta" family="Euphaeidae" genus="Anisopleura" kingdom="Animalia" order="Odonata" pageId="24" pageNumber="233" phylum="Arthropoda" rank="species" species="furcata">
<emphasis id="29AEEAA9E02DD74AFBA9E8A0FB7CFB46" box="[1095,1206,1152,1178]" italics="true" pageId="24" pageNumber="233">A. furcata</emphasis>
</taxonomicName>
were collected from under stones at the headwaters of clear streams with forest cover from
<quantity id="DC229B5EE02DD74AFC29E883FB94FB61" box="[967,1118,1187,1213]" metricMagnitude="3" metricUnit="m" metricValue="1.15" metricValueMax="1.2" metricValueMin="1.1" pageId="24" pageNumber="233" unit="m" value="1150.0" valueMax="1200.0" valueMin="1100.0">1100–1200m</quantity>
. They occurred in places of torrential flow and lower gradient trickles
<bibRefCitation id="7F4B4B4AE02DD74AFD67E8E7FC30FB3D" author="Keetapithchayakul, T. S. &amp; Orr, A. G. &amp; Wongkamhaeng, K." box="[649,1018,1223,1250]" pageId="24" pageNumber="233" pagination="529 - 542" refId="ref24083" refString="Keetapithchayakul, T. S., Orr, A. G. &amp; Wongkamhaeng, K. (2024) Description of the final stadium larva of Anisopleura furcata Selys, 1891 (Odonata: Euphaeidae) from Thailand. Zootaxa, 5415 (4), 529 - 542. https: // doi. org / 10.11646 / zootaxa. 5415.4.2" type="journal article" year="2024">
Keetapithchayakul
<emphasis id="29AEEAA9E02DD74AFC88E8E8FC51FB3D" box="[870,923,1223,1249]" italics="true" pageId="24" pageNumber="233">et al</emphasis>
. (2024)
</bibRefCitation>
, including sites where
<taxonomicName id="DCDA4D38E02DD74AFAE4E8E7FEBBFAD9" authority="Hamalainen" authorityName="Hamalainen" authorityYear="2002" class="Insecta" family="Euphaeidae" genus="Cryptophaea" kingdom="Animalia" order="Odonata" pageId="24" pageNumber="233" phylum="Arthropoda" rank="species" species="saukra">
<emphasis id="29AEEAA9E02DD74AFAE4E8E7FF29FAD9" italics="true" pageId="24" pageNumber="233">Cryptophaea saukra</emphasis>
Hämäläinen
</taxonomicName>
occurs. Similarly
<bibRefCitation id="7F4B4B4AE02DD74AFDD2E8CBFC99FAD9" author="Kumar, A. &amp; Prasad, M." box="[572,851,1259,1285]" pageId="24" pageNumber="233" pagination="225 - 230" refId="ref24294" refString="Kumar, A. &amp; Prasad, M. (1977 a) Last instar larvae of two Odonata species from Western Himalayas. Entomon, 2, 225 - 230." type="journal article" year="1977">Kumar &amp; Prasad (1977a)</bibRefCitation>
report the larvae of
<taxonomicName id="DCDA4D38E02DD74AFBDDE8CBFB79FAD9" authorityName="Selys" authorityYear="1853" box="[1075,1203,1259,1285]" class="Insecta" family="Euphaeidae" genus="Anisopleura" kingdom="Animalia" order="Odonata" pageId="24" pageNumber="233" phylum="Arthropoda" rank="species" species="lestoides">
<emphasis id="29AEEAA9E02DD74AFBDDE8CBFB79FAD9" box="[1075,1203,1259,1285]" italics="true" pageId="24" pageNumber="233">A. lestoides</emphasis>
</taxonomicName>
in rocky hill streams with swift water, sometimes together with
<taxonomicName id="DCDA4D38E02DD74AFD82E92EFD34FAF5" box="[620,766,1294,1321]" class="Insecta" family="Euphaeidae" genus="Bayadera" kingdom="Animalia" order="Odonata" pageId="24" pageNumber="233" phylum="Arthropoda" rank="species" species="undetermined">
<emphasis id="29AEEAA9E02DD74AFD82E92EFD13FAF4" box="[620,729,1294,1320]" italics="true" pageId="24" pageNumber="233">Bayadera</emphasis>
sp.
</taxonomicName>
<bibRefCitation id="7F4B4B4AE02DD74AFCEDE92EFBA8FAF5" author="Keetapithchayakul, T. S. &amp; Orr, A. G. &amp; Wongkamhaeng, K." box="[771,1122,1294,1321]" pageId="24" pageNumber="233" pagination="529 - 542" refId="ref24083" refString="Keetapithchayakul, T. S., Orr, A. G. &amp; Wongkamhaeng, K. (2024) Description of the final stadium larva of Anisopleura furcata Selys, 1891 (Odonata: Euphaeidae) from Thailand. Zootaxa, 5415 (4), 529 - 542. https: // doi. org / 10.11646 / zootaxa. 5415.4.2" type="journal article" year="2024">
Keetapithchayakul
<emphasis id="29AEEAA9E02DD74AFC35E92FFBC1FAF4" box="[987,1035,1294,1320]" italics="true" pageId="24" pageNumber="233">et al</emphasis>
. (2024)
</bibRefCitation>
report low level infestations by phoretic chironomid larvae on
<taxonomicName id="DCDA4D38E02DD74AFDFDE913FD49FA90" authorityName="Selys" authorityYear="1891" box="[531,643,1330,1357]" class="Insecta" family="Euphaeidae" genus="Anisopleura" kingdom="Animalia" order="Odonata" pageId="24" pageNumber="233" phylum="Arthropoda" rank="species" species="furcata">
<emphasis id="29AEEAA9E02DD74AFDFDE913FD49FA90" box="[531,643,1330,1357]" italics="true" pageId="24" pageNumber="233">A. furcata</emphasis>
</taxonomicName>
(see
<bibRefCitation id="7F4B4B4AE02DD74AFD53E913FC4BFA90" author="Boonsoong, B." box="[701,897,1330,1357]" pageId="24" pageNumber="233" pagination="163 - 169" refId="ref21686" refString="Boonsoong, B. (2016) Phoretic associations between Nanocladius asiaticus (Diptera, Chironomidae) and its hosts Gestroiella (Heteroptera, Naucoridae) and Euphaea masoni (Odonata, Euphaeidae) in streams in Western Thailand. Annales de Limnolgie - International Journal of Limnology, 52, 163 - 169. https: // doi. org / 10.1051 / limn / 2015025" type="journal article" year="2016">Boonsoong 2016</bibRefCitation>
).
</paragraph>
</subSubSection>
</treatment>
</document>

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<mods:title id="CDF76836E3EE58431096DC7013822679">A review of present knowledge of larvae of the Calopterygoidea (Zygoptera) of the Oriental realm, including keys to families and known genera</mods:title>
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<subSubSection id="53C06530E02DD74AFF79E959FECFFA48" box="[151,261,1401,1428]" pageId="24" pageNumber="233" type="nomenclature">
<paragraph id="1B6536BBE02DD74AFF79E959FECFFA48" blockId="24.[151,1437,1401,2036]" box="[151,261,1401,1428]" pageId="24" pageNumber="233">
<heading id="402D81D7E02DD74AFF79E959FECFFA48" bold="true" box="[151,261,1401,1428]" fontSize="11" level="1" pageId="24" pageNumber="233" reason="1">
<taxonomicName id="DCDA4D38E02DD74AFF79E959FECFFA48" ID-CoL="38YJ" authorityName="Selys" authorityYear="1853" box="[151,261,1401,1428]" class="Insecta" family="Euphaeidae" genus="Bayadera" kingdom="Animalia" order="Odonata" pageId="24" pageNumber="233" phylum="Arthropoda" rank="genus">
<emphasis id="29AEEAA9E02DD74AFF79E959FECFFA48" bold="true" box="[151,261,1401,1428]" italics="true" pageId="24" pageNumber="233">Bayadera</emphasis>
</taxonomicName>
</heading>
</paragraph>
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<subSubSection id="53C06530E02DD74BFF79E9BDFF27FBA1" lastPageId="25" lastPageNumber="234" pageId="24" pageNumber="233" type="discussion">
<paragraph id="1B6536BBE02DD74AFF79E9BDFCA8F952" blockId="24.[151,1437,1401,2036]" pageId="24" pageNumber="233">
The genus presently includes 17 species ranging from
<collectingRegion id="D91EF859E02DD74AFCC2E9BDFBCAFA6B" box="[812,1024,1437,1463]" country="India" name="Himachal Pradesh" pageId="24" pageNumber="233">Himachal Pradesh</collectingRegion>
(
<collectingCountry id="63CD762BE02DD74AFBFBE9BDFB98FA6B" box="[1045,1106,1437,1463]" name="India" pageId="24" pageNumber="233">India</collectingCountry>
) in the Western Himalayan region, through Northern
<collectingCountry id="63CD762BE02DD74AFE56E9E2FDEDFA00" box="[440,551,1474,1500]" name="Myanmar" pageId="24" pageNumber="233">Myanmar</collectingCountry>
,
<collectingCountry id="63CD762BE02DD74AFDDCE9E1FD52FA07" box="[562,664,1473,1499]" name="Thailand" pageId="24" pageNumber="233">Thailand</collectingCountry>
, Indochina, mainland
<collectingCountry id="63CD762BE02DD74AFC7CE9E1FC12FA07" box="[914,984,1473,1499]" name="China" pageId="24" pageNumber="233">China</collectingCountry>
,
<collectingRegion id="D91EF859E02DD74AFC0DE9E1FBFEFA07" box="[995,1076,1473,1499]" country="China" name="Hainan" pageId="24" pageNumber="233">Hainan</collectingRegion>
and
<collectingCountry id="63CD762BE02DD74AFB85E9E1FB0BFA07" box="[1131,1217,1473,1499]" name="Taiwan" pageId="24" pageNumber="233">Taiwan</collectingCountry>
. In mainland
<collectingCountry id="63CD762BE02DD74AFAB7E9E1FA56FA07" box="[1369,1436,1473,1499]" name="China" pageId="24" pageNumber="233">China</collectingCountry>
<taxonomicName id="DCDA4D38E02DD74AFF79E9C5FE6FFA23" authorityName="Ris" authorityYear="1912" box="[151,421,1509,1535]" class="Insecta" family="Euphaeidae" genus="Bayadera" kingdom="Animalia" order="Odonata" pageId="24" pageNumber="233" phylum="Arthropoda" rank="species" species="melanopteryx">
<emphasis id="29AEEAA9E02DD74AFF79E9C5FE6FFA23" box="[151,421,1509,1535]" italics="true" pageId="24" pageNumber="233">Bayadera melanopteryx</emphasis>
</taxonomicName>
ranges north into the Sino-Japanese realm of
<bibRefCitation id="7F4B4B4AE02DD74AFC5CE9C5FBB1FA23" author="Holt, B. G. &amp; Lessard, J. - P. &amp; Borregaard, M. K. &amp; Fritz, S. A. &amp; Araujo, M. B. &amp; Dimitrov, D. &amp; Fabre, P. - H. &amp; Graham, C. H. &amp; Graves, G. R. &amp; Jonsson, K. A. &amp; Nogues-Bravo, D. &amp; Wang, Z. &amp; Whittaker, R. J. &amp; Fjeldsa, J. &amp; Rahbek, C." box="[946,1147,1509,1535]" pageId="24" pageNumber="233" pagination="74 - 78" refId="ref23468" refString="Holt, B. G., Lessard, J. - P., Borregaard, M. K., Fritz, S. A., Araujo, M. B., Dimitrov, D., Fabre, P. - H., Graham, C. H., Graves, G. R., Jonsson, K. A., Nogues-Bravo, D., Wang, Z., Whittaker, R. J., Fjeldsa, J. &amp; Rahbek, C. (2013) An update of Wallace's zoogeographic regions of the world. Science, 339, 74 - 78. https: // doi. org / 10.1126 / science. 1228282" type="journal article" year="2013">
Holt
<emphasis id="29AEEAA9E02DD74AFC02E9C6FBECFA23" box="[1004,1062,1509,1535]" italics="true" pageId="24" pageNumber="233">et al.</emphasis>
(2013)
</bibRefCitation>
, formerly included in the Palaearctic realm. The larvae of more species have been described than other most euphaeid genera and there is clearly significant intrageneric variation in several characters, although the quality of illustrations and descriptions leave some uncertainty regarding diagnostic details, particularly the form of the caudal gills.
<bibRefCitation id="7F4B4B4AE02DD74AFB41EA70FECCF952" author="Keetapithchayakul, T. S. &amp; Sripanya, J. &amp; Phlai-Ngam, S. &amp; Tungpairojwong, N." pageId="24" pageNumber="233" pagination="98 - 110" refId="ref24014" refString="Keetapithchayakul, T. S., Sripanya, J., Phlai-Ngam, S. &amp; Tungpairojwong, N. (2020) Description of the final stadium larva of Bayadera serrata Davies &amp; Yang, 1996 (Odonata: Euphaeidae) from Thailand. Zootaxa, 4894 (1), 98 - 110. https: // doi. org / 10.11646 / zootaxa. 4894.1.5" type="journal article" year="2020">
Keetapithchayakul
<emphasis id="29AEEAA9E02DD74AFA66EA71FF67F952" italics="true" pageId="24" pageNumber="233">et al</emphasis>
. (2020)
</bibRefCitation>
analysed differences in larvae of species then known.
</paragraph>
<paragraph id="1B6536BBE02DD74AFF29EAB8FB53F82F" blockId="24.[151,1437,1401,2036]" pageId="24" pageNumber="233">
The larva of
<taxonomicName id="DCDA4D38E02DD74AFEB6EAB8FDC8F96E" authority="Selys" authorityName="Selys" authorityYear="1853" box="[344,514,1688,1714]" class="Insecta" family="Euphaeidae" genus="Bayadera" kingdom="Animalia" order="Odonata" pageId="24" pageNumber="233" phylum="Arthropoda" rank="species" species="indica">
<emphasis id="29AEEAA9E02DD74AFEB6EAB8FE74F96E" box="[344,446,1688,1714]" italics="true" pageId="24" pageNumber="233">B. indica</emphasis>
Selys
</taxonomicName>
from the Himalayas was first mentioned by
<bibRefCitation id="7F4B4B4AE02DD74AFC16EAB8FB59F96E" author="Hagen, H." box="[1016,1171,1688,1714]" pageId="24" pageNumber="233" pagination="lxv - lxvii" refId="ref23289" refString="Hagen, H. (1880) Essai d'un synopsis des larves de Calopterygines. [Comptes-rendus de la Socie´te´Entomologique de Belgique. Seance du 1 er Mai 1880]. Annales de la Societe entomologique de Belgique, 23, lxv - lxvii." type="journal article" year="1880">Hagen (1880)</bibRefCitation>
as
<taxonomicName id="DCDA4D38E02DD74AFB54EAB8FA5AF96D" authorityName="Hagen" authorityYear="1880" box="[1210,1424,1688,1714]" class="Insecta" family="Euphaeidae" genus="Anisopleura" kingdom="Animalia" order="Odonata" pageId="24" pageNumber="233" phylum="Arthropoda" rank="species" species="comes">
<emphasis id="29AEEAA9E02DD74AFB54EAB8FA5AF96D" box="[1210,1424,1688,1714]" italics="true" pageId="24" pageNumber="233">Anisopleura comes</emphasis>
</taxonomicName>
? (see
<bibRefCitation id="7F4B4B4AE02DD74AFF24EA9BFEBEF909" author="Needham, J. G." box="[202,372,1723,1749]" pageId="24" pageNumber="233" pagination="145 - 154" refId="ref24959" refString="Needham, J. G. (1911) Descriptions of dragonfly nymphs of the subfamily Calopteryginae (Odonata). Entomological News, 22, 145 - 154." type="journal article" year="1911">Needham 1911</bibRefCitation>
) who described the caudal gills as conical and short and noted that the second antennomere was shorter than the third. The same species was described and figured accurately and in some detail by
<bibRefCitation id="7F4B4B4AE02DD74AFB35EAFFFA5DF925" author="Needham, J. G." box="[1243,1431,1759,1785]" pageId="24" pageNumber="233" pagination="145 - 154" refId="ref24959" refString="Needham, J. G. (1911) Descriptions of dragonfly nymphs of the subfamily Calopteryginae (Odonata). Entomological News, 22, 145 - 154." type="journal article" year="1911">Needham (1911)</bibRefCitation>
, information partly repeated in
<bibRefCitation id="7F4B4B4AE02DD74AFE1DEB23FD7FF8C1" author="Needham, J. G." box="[499,693,1795,1821]" pageId="24" pageNumber="233" pagination="1 - 20" refId="ref24989" refString="Needham, J. G. (1930) A manual of the dragonflies of China: A monographic study of the Chinese Odonata. [Zoologia Sinica, Series A. Invertebrates of China, Vol. 11. Fascicle 1]. The Fan Memorial Institute of Biology, Peiping, xi + 344 pp., pls. 1 - 20 excl." type="book chapter" year="1930">Needham (1930)</bibRefCitation>
. Evidently unaware of this work,
<bibRefCitation id="7F4B4B4AE02DD74AFBD6EB23FB1BF8C1" author="Fraser, F. C." box="[1080,1233,1795,1821]" pageId="24" pageNumber="233" pagination="1 - 4" refId="ref23053" refString="Fraser, F. C. (1934) The fauna of British India, including Ceylon and Burma. Odonata. Vol. II. Taylor &amp; Francis Ltd., London, xxiii + 398 pp., pls. 1 - 4 excl." type="book chapter" year="1934">Fraser (1934)</bibRefCitation>
stated incorrectly that the larva was unknown for the genus, which is the more curious as earlier he had (
<bibRefCitation id="7F4B4B4AE02DD74AFB78EB07FAFAF89C" author="Fraser, F. C." box="[1174,1328,1830,1857]" pageId="24" pageNumber="233" pagination="288 - 301" refId="ref22972" refString="Fraser, F. C. (1929 b) Indian dragonflies, Part XXXII. Journal of the Bombay Natural History Society 33, 288 - 301, pls. 1 - 4 excl." type="journal article" year="1929">Fraser 1929b</bibRefCitation>
) asserted (incorrectly) that
<taxonomicName id="DCDA4D38E02DD74AFEB7EB6AFE0CF8B8" authorityName="Selys" authorityYear="1853" box="[345,454,1866,1892]" class="Insecta" family="Euphaeidae" genus="Bayadera" kingdom="Animalia" order="Odonata" pageId="24" pageNumber="233" phylum="Arthropoda" rank="genus">
<emphasis id="29AEEAA9E02DD74AFEB7EB6AFE0CF8B8" box="[345,454,1866,1892]" italics="true" pageId="24" pageNumber="233">Bayadera</emphasis>
</taxonomicName>
larvae lacked abdominal gills. The larva of
<taxonomicName id="DCDA4D38E02DD74AFC47EB6BFBC7F8B8" authorityName="Selys" authorityYear="1853" box="[937,1037,1866,1892]" class="Insecta" family="Euphaeidae" genus="Bayadera" kingdom="Animalia" order="Odonata" pageId="24" pageNumber="233" phylum="Arthropoda" rank="species" species="indica">
<emphasis id="29AEEAA9E02DD74AFC47EB6BFBC7F8B8" box="[937,1037,1866,1892]" italics="true" pageId="24" pageNumber="233">B. indica</emphasis>
</taxonomicName>
was also figured in detail by
<bibRefCitation id="7F4B4B4AE02DD74AFAA0EB6BFF2EF854" author="Kumar, A." pageId="24" pageNumber="233" pagination="83 - 118" refId="ref24196" refString="Kumar, A. (1973) Descriptions of the last instar larvae of Odonata from the Dehra Dun Valley (India), with notes on biology I. (suborder Zygoptera). Oriental Insects, 7, 83 - 118. https: // doi. org / 10.1080 / 00305316.1973.10434207" type="journal article" year="1973">Kumar (1973)</bibRefCitation>
, with structural details well presented and generally agreeing with
<bibRefCitation id="7F4B4B4AE02DD74AFC27EB4EFB42F854" author="Needham, J. G." box="[969,1160,1902,1928]" pageId="24" pageNumber="233" pagination="145 - 154" refId="ref24959" refString="Needham, J. G. (1911) Descriptions of dragonfly nymphs of the subfamily Calopteryginae (Odonata). Entomological News, 22, 145 - 154." type="journal article" year="1911">Needham (1911)</bibRefCitation>
. However
<bibRefCitation id="7F4B4B4AE02DD74AFB10EB4FFA56F854" author="Kumar, A." box="[1278,1436,1902,1929]" pageId="24" pageNumber="233" pagination="83 - 118" refId="ref24196" refString="Kumar, A. (1973) Descriptions of the last instar larvae of Odonata from the Dehra Dun Valley (India), with notes on biology I. (suborder Zygoptera). Oriental Insects, 7, 83 - 118. https: // doi. org / 10.1080 / 00305316.1973.10434207" type="journal article" year="1973">Kumar (1973)</bibRefCitation>
depicts the habitus rather roughly, with a longer filament on a caudal gills reminiscent of
<taxonomicName id="DCDA4D38E02DD74AFB67EBB2FAD8F870" authorityName="Selys" authorityYear="1853" box="[1161,1298,1938,1964]" class="Insecta" family="Euphaeidae" genus="Anisopleura" kingdom="Animalia" order="Odonata" pageId="24" pageNumber="233" phylum="Arthropoda" rank="genus">
<emphasis id="29AEEAA9E02DD74AFB67EBB2FAD8F870" box="[1161,1298,1938,1964]" italics="true" pageId="24" pageNumber="233">Anisopleura</emphasis>
</taxonomicName>
or
<taxonomicName id="DCDA4D38E02DD74AFAD7EBB2FA57F870" authorityName="Selys" authorityYear="1840" box="[1337,1437,1938,1964]" class="Insecta" family="Euphaeidae" genus="Euphaea" kingdom="Animalia" order="Odonata" pageId="24" pageNumber="233" phylum="Arthropoda" rank="genus">
<emphasis id="29AEEAA9E02DD74AFAD7EBB2FA57F870" box="[1337,1437,1938,1964]" italics="true" pageId="24" pageNumber="233">Euphaea</emphasis>
</taxonomicName>
and inconsistent with
<bibRefCitation id="7F4B4B4AE02DD74AFE61EB95FD94F80C" author="Needham, J. G." box="[399,606,1973,2000]" pageId="24" pageNumber="233" pagination="145 - 154" refId="ref24959" refString="Needham, J. G. (1911) Descriptions of dragonfly nymphs of the subfamily Calopteryginae (Odonata). Entomological News, 22, 145 - 154." type="journal article" year="1911">Needham’s (1911)</bibRefCitation>
detailed drawing of the lateral gill and also the habitus drawing seems to show the antennae too short for
<taxonomicName id="DCDA4D38E02DD74AFE10EBF9FDA1F82F" authorityName="Selys" authorityYear="1853" box="[510,619,2009,2035]" class="Insecta" family="Euphaeidae" genus="Bayadera" kingdom="Animalia" order="Odonata" pageId="24" pageNumber="233" phylum="Arthropoda" rank="genus">
<emphasis id="29AEEAA9E02DD74AFE10EBF9FDA1F82F" box="[510,619,2009,2035]" italics="true" pageId="24" pageNumber="233">Bayadera</emphasis>
</taxonomicName>
, whereas in the detail of the head they are longer.
</paragraph>
<paragraph id="1B6536BBE02CD74BFF29ECB7FCE4FE71" blockId="25.[151,1437,151,1149]" pageId="25" pageNumber="234">
<bibRefCitation id="7F4B4B4AE02CD74BFF29ECB7FE07FF6D" author="Matsuki, K. &amp; Lien, J. C." box="[199,461,151,177]" pageId="25" pageNumber="234" pagination="15 - 26" refId="ref24810" refString="Matsuki, K. &amp; Lien, J. C. (1978) Descriptions of the larvae of three families of Zygoptera breeding in the streams of Taiwan (Synlestidae, Euphaeidae &amp; Calopterygidae). Tombo - Acta Odonatologica Japonica, 21, 15 - 26." type="journal article" year="1978">Matsuki &amp; Lien (1978)</bibRefCitation>
described and illustrated the larva of
<taxonomicName id="DCDA4D38E02CD74BFC9BECB7FB95FF6D" authority="Fraser" authorityName="Fraser" authorityYear="1934" box="[885,1119,151,177]" class="Insecta" family="Euphaeidae" genus="Bayadera" kingdom="Animalia" order="Odonata" pageId="25" pageNumber="234" phylum="Arthropoda" rank="species" species="brevicauda">
<emphasis id="29AEEAA9E02CD74BFC9BECB7FBD8FF6D" box="[885,1042,151,177]" italics="true" pageId="25" pageNumber="234">B. brevicauda</emphasis>
Fraser
</taxonomicName>
from
<collectingCountry id="63CD762BE02CD74BFB4AECB7FB33FF6D" box="[1188,1273,151,177]" name="Taiwan" pageId="25" pageNumber="234">Taiwan</collectingCountry>
, showing very squared caudal gills with a short narrow tip constricted at the base, which they describe as “more or less sausage shaped with a short tail”.
<bibRefCitation id="7F4B4B4AE02CD74BFE57ECFFFD85FF25" author="Ishida, K." box="[441,591,223,249]" pageId="25" pageNumber="234" refId="ref23597" refString="Ishida, K. (1996) Monograph of Odonata larvae in Japan. Hokkaido University Press, Sapporo, 446 pp. [in Japanese]" type="book" year="1996">Ishida (1996)</bibRefCitation>
figured
<taxonomicName id="DCDA4D38E02CD74BFD5EECFFFC7EFF25" authority="Asahina" authorityName="Asahina" authorityYear="1964" box="[688,948,223,249]" class="Insecta" family="Euphaeidae" genus="Bayadera" kingdom="Animalia" order="Odonata" pageId="25" pageNumber="234" phylum="Arthropoda" rank="species" species="ishigakiana">
<emphasis id="29AEEAA9E02CD74BFD5EECFFFC98FF25" box="[688,850,223,249]" italics="true" pageId="25" pageNumber="234">B. ishigakiana</emphasis>
Asahina
</taxonomicName>
in detail, illustrating a rather attenuated tip to the gills, which arise abruptly and do not taper from the main sac as in other species of the genus, although this form, while similar is less extreme in an illustration by Sugimura
<emphasis id="29AEEAA9E02CD74BFC83ED07FC6FFE9D" box="[877,933,295,321]" italics="true" pageId="25" pageNumber="234">et al.</emphasis>
(1999).
<bibRefCitation id="7F4B4B4AE02CD74BFBECED07FB0BFE9D" author="Wan, F. X. &amp; Jiang, Y. H. &amp; Wan, J." box="[1026,1217,295,321]" pageId="25" pageNumber="234" pagination="143 - 147" refId="ref27296" refString="Wan, F. X., Jiang, Y. H. &amp; Wan, J. (2011) Description of Bayadera bidentata Needham and Asiagomphus cuneatus Needham larvae from Huangshan Mountain, China (Zygoptera: Euphaeidae, Anisoptera: Gomphidae). Odonatologica, 40, 143 - 147." type="journal article" year="2011">
Wan
<emphasis id="29AEEAA9E02CD74BFBD4ED07FBA1FE9D" box="[1082,1131,295,321]" italics="true" pageId="25" pageNumber="234">et al</emphasis>
. (2011)
</bibRefCitation>
describe and figure diagnostic details for
<taxonomicName id="DCDA4D38E02CD74BFE6BED6BFD4BFEB9" authority="Needham" authorityName="Needham" authorityYear="1930" box="[389,641,331,357]" class="Insecta" family="Euphaeidae" genus="Bayadera" kingdom="Animalia" order="Odonata" pageId="25" pageNumber="234" phylum="Arthropoda" rank="species" species="bidentata">
<emphasis id="29AEEAA9E02CD74BFE6BED6BFE5CFEB8" box="[389,406,331,356]" italics="true" pageId="25" pageNumber="234">B</emphasis>
.
<emphasis id="29AEEAA9E02CD74BFE4DED6BFDC7FEB9" box="[419,525,331,357]" italics="true" pageId="25" pageNumber="234">bidentata</emphasis>
Needham
</taxonomicName>
; The caudal gills are depicted as sausage shaped and tipped with a short, abrupt cone, and a similar form was found in
<taxonomicName id="DCDA4D38E02CD74BFD75ED4FFC75FE55" authority="Davies &amp; Yang" authorityName="Davies &amp; Yang" authorityYear="1996" box="[667,959,367,393]" class="Insecta" family="Euphaeidae" genus="Bayadera" kingdom="Animalia" order="Odonata" pageId="25" pageNumber="234" phylum="Arthropoda" rank="species" species="serrata">
<emphasis id="29AEEAA9E02CD74BFD75ED4FFCC6FE54" box="[667,780,367,392]" italics="true" pageId="25" pageNumber="234">B. serrata</emphasis>
Davies &amp; Yang
</taxonomicName>
by
<bibRefCitation id="7F4B4B4AE02CD74BFC04ED4FFA85FE55" author="Keetapithchayakul, T. S. &amp; Sripanya, J. &amp; Phlai-Ngam, S. &amp; Tungpairojwong, N." box="[1002,1359,367,393]" pageId="25" pageNumber="234" pagination="98 - 110" refId="ref24014" refString="Keetapithchayakul, T. S., Sripanya, J., Phlai-Ngam, S. &amp; Tungpairojwong, N. (2020) Description of the final stadium larva of Bayadera serrata Davies &amp; Yang, 1996 (Odonata: Euphaeidae) from Thailand. Zootaxa, 4894 (1), 98 - 110. https: // doi. org / 10.11646 / zootaxa. 4894.1.5" type="journal article" year="2020">
Keetapithchayakul
<emphasis id="29AEEAA9E02CD74BFB2AED4FFB3CFE55" box="[1220,1270,367,393]" italics="true" pageId="25" pageNumber="234">et al</emphasis>
. (2020)
</bibRefCitation>
and in
<taxonomicName id="DCDA4D38E02CD74BFF79EDB3FE75FE71" authority="Davies &amp; Yang" authorityName="Davies &amp; Yang" authorityYear="1996" box="[151,447,403,429]" class="Insecta" family="Euphaeidae" genus="Bayadera" kingdom="Animalia" order="Odonata" pageId="25" pageNumber="234" phylum="Arthropoda" rank="species" species="strigata">
<emphasis id="29AEEAA9E02CD74BFF79EDB3FEC7FE70" box="[151,269,403,428]" italics="true" pageId="25" pageNumber="234">B. strigata</emphasis>
Davies &amp; Yang
</taxonomicName>
(
<figureCitation id="83E12A3EE02CD74BFE20EDB3FDD5FE71" box="[462,543,403,429]" captionStart="FIGURES 67–74. 67" captionStartId="13.[151,264,1814,1838]" captionTargetBox="[256,1340,160,1799]" captionTargetId="figure-15@13.[246,1342,151,1807]" captionTargetPageId="13" captionText="FIGURES 67–74. 67. Anisopleura subplatystyla, caudal gills dorsal view (original drawing). 68. Bayadera strigata, caudal gills dorsal view, (drawn from Yang &amp; Orr 2024). 69. Euphaea decorata, caudal gills dorsal view (drawn from Xu 2017). 70. Euphaea superba, caudal gills dorsal view (redrawn from Wu et al. 2017). 71. Bayadera strigata: male genital apophyses; ventral (left), right lateral (right), (drawn from Yang &amp; Orr 2024). 72. Euphaea decorata, male genital apophyses; ventral view (drawn from Xu 2017). 73. Euphaea superba, terminal processes of palpal lobe ventral view (movable hook truncated), (redrawn from Wu et al. 2017). 74. Anisopleura furcata, left labial palp showing terminal processes of palpal lobe, dorsal view (redrawn from Keetapithchayakul et al. 2024)." figureDoi="http://doi.org/10.5281/zenodo.13618345" httpUri="https://zenodo.org/record/13618345/files/figure.png" pageId="25" pageNumber="234">Fig. 68</figureCitation>
) by
<bibRefCitation id="7F4B4B4AE02CD74BFDBFEDB3FCE0FE71" author="Yang, G. - H. &amp; Orr, A. G." box="[593,810,403,429]" pageId="25" pageNumber="234" pagination="11 - 18" refId="ref27987" refString="Yang, G. - H. &amp; Orr, A. G. (2024) Descriptions of larvae of Bayadera strigata Davies &amp; Yang, 1996 (Odonata: Euphaeidae). International Journal of Odonatology, 27, 11 - 18. https: // doi. org / 10.48156 / 1388.2024.1917250" type="journal article" year="2024">Yang &amp; Orr (2024)</bibRefCitation>
.
</paragraph>
<paragraph id="1B6536BBE02CD74BFF29ED97FAC2FC81" blockId="25.[151,1437,151,1149]" pageId="25" pageNumber="234">
Larvae of
<taxonomicName id="DCDA4D38E02CD74BFED5ED97FE62FE0D" authorityName="Selys" authorityYear="1853" box="[315,424,439,465]" class="Insecta" family="Euphaeidae" genus="Bayadera" kingdom="Animalia" order="Odonata" pageId="25" pageNumber="234" phylum="Arthropoda" rank="genus">
<emphasis id="29AEEAA9E02CD74BFED5ED97FE62FE0D" box="[315,424,439,465]" italics="true" pageId="25" pageNumber="234">Bayadera</emphasis>
</taxonomicName>
tend to be slighter in build than
<taxonomicName id="DCDA4D38E02CD74BFCF7ED97FCB7FE0D" authorityName="Selys" authorityYear="1840" box="[793,893,439,465]" class="Insecta" family="Euphaeidae" genus="Euphaea" kingdom="Animalia" order="Odonata" pageId="25" pageNumber="234" phylum="Arthropoda" rank="genus">
<emphasis id="29AEEAA9E02CD74BFCF7ED97FCB7FE0D" box="[793,893,439,465]" italics="true" pageId="25" pageNumber="234">Euphaea</emphasis>
</taxonomicName>
, with longer legs (
<figureCitation id="83E12A3EE02CD74BFBBFED97FB6FFE0D" box="[1105,1189,438,465]" captionStart="FIGURES 57–60. 57" captionStartId="11.[151,264,1933,1957]" captionTargetBox="[193,1384,155,1907]" captionTargetId="figure-15@11.[188,1400,151,1917]" captionTargetPageId="11" captionText="FIGURES 57–60. 57. Dysphaea gloriosa, habitus (drawn from Nguyen et al. 2024). 58. Bayadera strigata, habitus (drawn from Yang &amp; Orr 2024). 59. Euphaea decorata, habitus. (drawn from Xu 2017). 60. Anisopleura furcata, habitus (drawn from Keetapithchayakul et al. 2024)." figureDoi="http://doi.org/10.5281/zenodo.13618341" httpUri="https://zenodo.org/record/13618341/files/figure.png" pageId="25" pageNumber="234">Fig. 58</figureCitation>
), and the hind margin of the head is slightly more rounded, but these differences cannot be considered diagnostic. All species known share the following characteristics: the antennae are conspicuously long, being longer than the distance between the posterior occipital margin and the base of the labrum. The postocular lobes are strongly developed, slightly bulbous, and smoothly rounded (
<figureCitation id="83E12A3EE02CD74BFE71EE67FE39FDBD" box="[415,499,583,609]" captionStart="FIGURES 61–66. 61" captionStartId="12.[151,264,1841,1865]" captionTargetBox="[273,1314,156,1828]" captionTargetId="figure-21@12.[268,1319,151,1834]" captionTargetPageId="12" captionText="FIGURES 61–66. 61. Dysphaea gloriosa: detail of spines on genae and postocular lobe; underside (left), upperside (right) (modified from Nguyen et al. 2024). 62. Heterophaea barbata: dorsal view of head, showing (a) outer mandibular spines (b) genal spines partially visible and (c) right antenna (from Orr &amp; Hämäläinen 2024). 63. Anisopleura subplatystyla, dorsal view of head (original drawing). 64. Bayadera strigata, dorsal view of head (drawn from Yang &amp; Orr 2024). 65. Euphaea decorata, dorsal view of head. (drawn from Xu 2017). 66. Bayadera strigata, left labial palp with detail showing wavy outer margin of outer process of palpal lobe (arrowed), (drawn from Yang &amp; Orr 2024)." figureDoi="http://doi.org/10.5281/zenodo.13618343" httpUri="https://zenodo.org/record/13618343/files/figure.png" pageId="25" pageNumber="234">Fig. 64</figureCitation>
). In species for which information is available (
<taxonomicName id="DCDA4D38E02CD74BFBE8EE67FBA0FDBD" authorityName="Selys" authorityYear="1853" box="[1030,1130,583,609]" class="Insecta" family="Euphaeidae" genus="Bayadera" kingdom="Animalia" order="Odonata" pageId="25" pageNumber="234" phylum="Arthropoda" rank="species" species="indica">
<emphasis id="29AEEAA9E02CD74BFBE8EE67FBA0FDBD" box="[1030,1130,583,609]" italics="true" pageId="25" pageNumber="234">B. indica</emphasis>
</taxonomicName>
,
<emphasis id="29AEEAA9E02CD74BFB96EE67FAA0FDBC" box="[1144,1386,583,608]" italics="true" pageId="25" pageNumber="234">
<taxonomicName id="DCDA4D38E02CD74BFB96EE67FB20FDBC" authorityName="Davies &amp; Yang" authorityYear="1996" box="[1144,1258,583,608]" class="Insecta" family="Euphaeidae" genus="Bayadera" kingdom="Animalia" order="Odonata" pageId="25" pageNumber="234" phylum="Arthropoda" rank="species" species="serrata">B. serrata</taxonomicName>
,
<taxonomicName id="DCDA4D38E02CD74BFB1BEE67FAA0FDBC" authorityName="Davies &amp; Yang" authorityYear="1996" box="[1269,1386,583,608]" class="Insecta" family="Euphaeidae" genus="Bayadera" kingdom="Animalia" order="Odonata" pageId="25" pageNumber="234" phylum="Arthropoda" rank="species" species="strigata">B. strigata</taxonomicName>
</emphasis>
) the male genital apophyses are large and rounded, curving to overlap S10 (
<figureCitation id="83E12A3EE02CD74BFC53EE4BFBD8FD59" box="[957,1042,619,645]" captionStart="FIGURES 67–74. 67" captionStartId="13.[151,264,1814,1838]" captionTargetBox="[256,1340,160,1799]" captionTargetId="figure-15@13.[246,1342,151,1807]" captionTargetPageId="13" captionText="FIGURES 67–74. 67. Anisopleura subplatystyla, caudal gills dorsal view (original drawing). 68. Bayadera strigata, caudal gills dorsal view, (drawn from Yang &amp; Orr 2024). 69. Euphaea decorata, caudal gills dorsal view (drawn from Xu 2017). 70. Euphaea superba, caudal gills dorsal view (redrawn from Wu et al. 2017). 71. Bayadera strigata: male genital apophyses; ventral (left), right lateral (right), (drawn from Yang &amp; Orr 2024). 72. Euphaea decorata, male genital apophyses; ventral view (drawn from Xu 2017). 73. Euphaea superba, terminal processes of palpal lobe ventral view (movable hook truncated), (redrawn from Wu et al. 2017). 74. Anisopleura furcata, left labial palp showing terminal processes of palpal lobe, dorsal view (redrawn from Keetapithchayakul et al. 2024)." figureDoi="http://doi.org/10.5281/zenodo.13618345" httpUri="https://zenodo.org/record/13618345/files/figure.png" pageId="25" pageNumber="234">Fig. 71</figureCitation>
), in which respect they differ from
<taxonomicName id="DCDA4D38E02CD74BFF79EEAFFF31FD75" authorityName="Selys" authorityYear="1840" box="[151,251,655,681]" class="Insecta" family="Euphaeidae" genus="Euphaea" kingdom="Animalia" order="Odonata" pageId="25" pageNumber="234" phylum="Arthropoda" rank="genus">
<emphasis id="29AEEAA9E02CD74BFF79EEAFFF31FD75" box="[151,251,655,681]" italics="true" pageId="25" pageNumber="234">Euphaea</emphasis>
</taxonomicName>
. There are few or no tubercles on the labrum and anterior part of the head, as in
<taxonomicName id="DCDA4D38E02CD74BFB9CEEAFFB31FD75" authorityName="Selys" authorityYear="1853" box="[1138,1275,655,681]" class="Insecta" family="Euphaeidae" genus="Anisopleura" kingdom="Animalia" order="Odonata" pageId="25" pageNumber="234" phylum="Arthropoda" rank="genus">
<emphasis id="29AEEAA9E02CD74BFB9CEEAFFB31FD75" box="[1138,1275,655,681]" italics="true" pageId="25" pageNumber="234">Anisopleura</emphasis>
</taxonomicName>
and claviform setae characteristic of all euphaids are often especially dense in this area. The inner apical process of the labial palp is free and securiform except in
<taxonomicName id="DCDA4D38E02CD74BFDECEEF7FD41FD2D" authorityName="Needham" authorityYear="1930" box="[514,651,727,753]" class="Insecta" family="Euphaeidae" genus="Bayadera" kingdom="Animalia" order="Odonata" pageId="25" pageNumber="234" phylum="Arthropoda" rank="species" species="bidentata">
<emphasis id="29AEEAA9E02CD74BFDECEEF7FD41FD2D" box="[514,651,727,753]" italics="true" pageId="25" pageNumber="234">B. bidentata</emphasis>
</taxonomicName>
in which it is almost atrophied to an angulated thorn like process, but also supplemented by another blunt process about the mid-inner margin of the palp that is unknown in any other member of the family. In species examined sufficiently closely (
<taxonomicName id="DCDA4D38E02CD74BFC9EEF3FFC1CFCE5" authorityName="Selys" authorityYear="1853" box="[880,982,799,825]" class="Insecta" family="Euphaeidae" genus="Bayadera" kingdom="Animalia" order="Odonata" pageId="25" pageNumber="234" phylum="Arthropoda" rank="species" species="indica">
<emphasis id="29AEEAA9E02CD74BFC9EEF3FFC1CFCE5" box="[880,982,799,825]" italics="true" pageId="25" pageNumber="234">B. indica</emphasis>
</taxonomicName>
,
<emphasis id="29AEEAA9E02CD74BFC0BEF3FFB16FCE4" box="[997,1244,799,824]" italics="true" pageId="25" pageNumber="234">
<taxonomicName id="DCDA4D38E02CD74BFC0BEF3FFB93FCE4" authorityName="Davies &amp; Yang" authorityYear="1996" box="[997,1113,799,824]" class="Insecta" family="Euphaeidae" genus="Bayadera" kingdom="Animalia" order="Odonata" pageId="25" pageNumber="234" phylum="Arthropoda" rank="species" species="serrata">B. serrata</taxonomicName>
,
<taxonomicName id="DCDA4D38E02CD74BFB8BEF3FFB16FCE4" authorityName="Davies &amp; Yang" authorityYear="1996" box="[1125,1244,799,824]" class="Insecta" family="Euphaeidae" genus="Bayadera" kingdom="Animalia" order="Odonata" pageId="25" pageNumber="234" phylum="Arthropoda" rank="species" species="strigata">B. strigata</taxonomicName>
</emphasis>
) the outer apical process of the lobe of the labial palp has a wavy margin unknown in other described genera (
<figureCitation id="83E12A3EE02CD74BFB49EF63FB31FC81" box="[1191,1275,835,861]" captionStart="FIGURES 61–66. 61" captionStartId="12.[151,264,1841,1865]" captionTargetBox="[273,1314,156,1828]" captionTargetId="figure-21@12.[268,1319,151,1834]" captionTargetPageId="12" captionText="FIGURES 61–66. 61. Dysphaea gloriosa: detail of spines on genae and postocular lobe; underside (left), upperside (right) (modified from Nguyen et al. 2024). 62. Heterophaea barbata: dorsal view of head, showing (a) outer mandibular spines (b) genal spines partially visible and (c) right antenna (from Orr &amp; Hämäläinen 2024). 63. Anisopleura subplatystyla, dorsal view of head (original drawing). 64. Bayadera strigata, dorsal view of head (drawn from Yang &amp; Orr 2024). 65. Euphaea decorata, dorsal view of head. (drawn from Xu 2017). 66. Bayadera strigata, left labial palp with detail showing wavy outer margin of outer process of palpal lobe (arrowed), (drawn from Yang &amp; Orr 2024)." figureDoi="http://doi.org/10.5281/zenodo.13618343" httpUri="https://zenodo.org/record/13618343/files/figure.png" pageId="25" pageNumber="234">Fig. 66</figureCitation>
).
</paragraph>
<paragraph id="1B6536BBE02CD74BFF79EF47FB0DFC15" blockId="25.[151,1437,151,1149]" pageId="25" pageNumber="234">
Most species lack long spines on the genae (longer than about one third of the width of the ventrally visible compound eye), but
<taxonomicName id="DCDA4D38E02CD74BFF10EFABFEA6FC78" authorityName="Davies &amp; Yang" authorityYear="1996" box="[254,364,907,932]" class="Insecta" family="Euphaeidae" genus="Bayadera" kingdom="Animalia" order="Odonata" pageId="25" pageNumber="234" phylum="Arthropoda" rank="species" species="serrata">
<emphasis id="29AEEAA9E02CD74BFF10EFABFEA6FC78" box="[254,364,907,932]" italics="true" pageId="25" pageNumber="234">B. serrata</emphasis>
</taxonomicName>
has 1–3 spines
<bibRefCitation id="7F4B4B4AE02CD74BFDF6EFABFCB0FC79" author="Keetapithchayakul, T. S. &amp; Sripanya, J. &amp; Phlai-Ngam, S. &amp; Tungpairojwong, N." box="[536,890,907,933]" pageId="25" pageNumber="234" pagination="98 - 110" refId="ref24014" refString="Keetapithchayakul, T. S., Sripanya, J., Phlai-Ngam, S. &amp; Tungpairojwong, N. (2020) Description of the final stadium larva of Bayadera serrata Davies &amp; Yang, 1996 (Odonata: Euphaeidae) from Thailand. Zootaxa, 4894 (1), 98 - 110. https: // doi. org / 10.11646 / zootaxa. 4894.1.5" type="journal article" year="2020">
Keetapithchayakul
<emphasis id="29AEEAA9E02CD74BFD1EEFABFCEAFC79" box="[752,800,907,933]" italics="true" pageId="25" pageNumber="234">et al</emphasis>
. (2020)
</bibRefCitation>
, hence a lack of spines or just one spine generally indicates that the larva is
<taxonomicName id="DCDA4D38E02CD74BFE5BEF8FFDE8FC15" authorityName="Selys" authorityYear="1853" box="[437,546,943,969]" class="Insecta" family="Euphaeidae" genus="Bayadera" kingdom="Animalia" order="Odonata" pageId="25" pageNumber="234" phylum="Arthropoda" rank="genus">
<emphasis id="29AEEAA9E02CD74BFE5BEF8FFDE8FC15" box="[437,546,943,969]" italics="true" pageId="25" pageNumber="234">Bayadera</emphasis>
</taxonomicName>
, but presence of multiple spines does not rule out the genus.
</paragraph>
<paragraph id="1B6536BBE02CD74BFF29EFF3FF27FBA1" blockId="25.[151,1437,151,1149]" pageId="25" pageNumber="234">
<taxonomicName id="DCDA4D38E02CD74BFF29EFF3FEFEFC31" ID-CoL="38YJ" authorityName="Selys" authorityYear="1853" box="[199,308,979,1005]" class="Insecta" family="Euphaeidae" genus="Bayadera" kingdom="Animalia" order="Odonata" pageId="25" pageNumber="234" phylum="Arthropoda" rank="genus">
<emphasis id="29AEEAA9E02CD74BFF29EFF3FEFEFC31" box="[199,308,979,1005]" italics="true" pageId="25" pageNumber="234">Bayadera</emphasis>
</taxonomicName>
<materialsCitation id="ABB23CE6E02CD74BFED3EFF3FA53FBE9" elevation="1500" location="At" municipality="The" pageId="25" pageNumber="234" specimenCount="1">
larvae are typically found clinging under stones in clear, moderately fast flowing water in shallow streams.
<location id="1E056060E02CD74BFF13EFD7FED3FBCD" LSID="urn:lsid:plazi:treatment:937387ADE02DD74BFF79E959FF27FBA1:1E056060E02CD74BFF13EFD7FED3FBCD" box="[253,281,1015,1041]" municipality="The" name="At" pageId="25" pageNumber="234">At</location>
higher altitudes they frequently occur together with
<taxonomicName id="DCDA4D38E02CD74BFC68EFD7FBC5FBCD" ID-CoL="WFM" authorityName="Selys" authorityYear="1853" box="[902,1039,1015,1041]" class="Insecta" family="Euphaeidae" genus="Anisopleura" kingdom="Animalia" order="Odonata" pageId="25" pageNumber="234" phylum="Arthropoda" rank="genus">
<emphasis id="29AEEAA9E02CD74BFC68EFD7FBC5FBCD" box="[902,1039,1015,1041]" italics="true" pageId="25" pageNumber="234">Anisopleura</emphasis>
</taxonomicName>
larvae, apparently in very similar microhabitats.
<collectingMunicipality id="FB01ACC1E02CD74BFED1E83BFEA1FBE9" box="[319,363,1051,1077]" pageId="25" pageNumber="234">The</collectingMunicipality>
various species have different altitudinal preferences ranging from
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(
<bibRefCitation id="7F4B4B4AE02CD74BFAECE83BFA5BFBE9" author="Zhang, H. - M." box="[1282,1425,1051,1077]" pageId="25" pageNumber="234" refId="ref28336" refString="Zhang, H. - M. (2019) Dragonflies and Damselflies of China. Vols. 1 &amp; 2. Chong Qing University Press, ChongQing, 14 + 1460 pp. [bilingual, in Chinese and English]" type="book" year="2019">Zhang 2019</bibRefCitation>
)
</materialsCitation>
.
<materialsCitation id="ABB23CE6E02CD74BFF79E81FFF23FBA1" country="Bhutan" elevation="900" location="Bhutan" pageId="25" pageNumber="234" specimenCount="1">
<bibRefCitation id="7F4B4B4AE02CD74BFF79E81FFE42FB85" author="Rasaily, B. &amp; Kalkman, V. J. &amp; Katel, O. &amp; Dorji, C." box="[151,392,1087,1113]" pageId="25" pageNumber="234" pagination="1 - 20" refId="ref26410" refString="Rasaily, B., Kalkman, V. J., Katel, O. &amp; Dorji, C. (2021) Composition of the dragonfly fauna at different altitudes in Bhutan based on larval samples. IDF-Report, 160, 1 - 20." type="journal article" year="2021">
Rasaily
<emphasis id="29AEEAA9E02CD74BFF18E81FFEE6FB85" box="[246,300,1087,1113]" italics="true" pageId="25" pageNumber="234">et al</emphasis>
. (2021)
</bibRefCitation>
report
<taxonomicName id="DCDA4D38E02CD74BFE0CE81FFD85FB85" ID-CoL="38YJ" authorityName="Selys" authorityYear="1853" box="[482,591,1087,1113]" class="Insecta" family="Euphaeidae" genus="Bayadera" kingdom="Animalia" order="Odonata" pageId="25" pageNumber="234" phylum="Arthropoda" rank="genus">
<emphasis id="29AEEAA9E02CD74BFE0CE81FFD85FB85" box="[482,591,1087,1113]" italics="true" pageId="25" pageNumber="234">Bayadera</emphasis>
</taxonomicName>
larva at many locations in
<collectingCountry id="63CD762BE02CD74BFC79E81FFC21FB85" box="[919,1003,1087,1113]" name="Bhutan" pageId="25" pageNumber="234">Bhutan</collectingCountry>
, some quite disturbed, from ca
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.
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Family
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The single genus
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<emphasis id="29AEEAA9E02ED749FE85E9B7FE24FA6D" box="[363,494,1431,1457]" italics="true" pageId="27" pageNumber="236">Philoganga</emphasis>
</taxonomicName>
is known from four large species ranging from the eastern Himalaya, northern
<collectingCountry id="63CD762BE02ED749FF79E99CFECCFA0A" box="[151,262,1468,1494]" name="Myanmar" pageId="27" pageNumber="236">Myanmar</collectingCountry>
,
<collectingCountry id="63CD762BE02ED749FEFDE99BFEBDFA09" box="[275,375,1467,1493]" name="Thailand" pageId="27" pageNumber="236">Thailand</collectingCountry>
and Indochina to southern
<collectingCountry id="63CD762BE02ED749FD59E99BFD34FA09" box="[695,766,1467,1493]" name="China" pageId="27" pageNumber="236">China</collectingCountry>
. The larvae of three species are known, and are generally similar,
<bibRefCitation id="7F4B4B4AE02ED749FF1AE9C0FE44FA25" author="Fraser, F. C." box="[244,398,1503,1530]" pageId="27" pageNumber="236" pagination="137 - 143" refId="ref23100" refString="Fraser, F. C. (1938) Additions to the Amphipterygidae (Order Odonata). Proceedings of the Royal Entomological Society of London, Series B, 7, 137 - 143. https: // doi. org / 10.1111 / j. 1365 - 3113.1938. tb 01267. x" type="journal article" year="1938">Fraser (1938)</bibRefCitation>
described and figured
<taxonomicName id="DCDA4D38E02ED749FD79E9C0FCB5FA25" authority="(Hagen)" baseAuthorityName="Hagen" box="[663,895,1503,1529]" class="Insecta" family="Lestoideidae" genus="Philoganga" kingdom="Animalia" order="Odonata" pageId="27" pageNumber="236" phylum="Arthropoda" rank="species" species="montana">
<emphasis id="29AEEAA9E02ED749FD79E9C0FCDDFA25" box="[663,791,1504,1529]" italics="true" pageId="27" pageNumber="236">P. montana</emphasis>
(Hagen)
</taxonomicName>
,
<bibRefCitation id="7F4B4B4AE02ED749FC62E9FFFBDFFA25" author="Chao, H. - F." box="[908,1045,1503,1529]" pageId="27" pageNumber="236" refId="ref22196" refString="Chao, H. - F. (1948) The taxonomic position and the larval characters of genus Philoganga Kirby (Odonata). Science, 30, 19. [in Chinese]" type="journal volume" year="1948">Chao (1948</bibRefCitation>
,
<bibRefCitation id="7F4B4B4AE02ED749FBCCE9FFFB91FA26" author="Chao, H. - F." box="[1058,1115,1503,1530]" pageId="27" pageNumber="236" pagination="137 - 143" refId="ref22231" refString="Chao, H. - F. (1953) A study of Chinese dragonflies of the genus Philoganga Kirby (Odonata, Zygoptera, Amphipterygidae). Acta Entomologica Sinica, 3, 137 - 143, plate 1, excl. [in Chinese, with English summary]" type="journal article" year="1953">1953</bibRefCitation>
) discussed Chinese species, including
<taxonomicName id="DCDA4D38E02ED749FEE2EA24FE1EF9C1" authority="Navas" authorityName="Navas" authorityYear="1936" box="[268,468,1539,1565]" class="Insecta" family="Lestoideidae" genus="Philoganga" kingdom="Animalia" order="Odonata" pageId="27" pageNumber="236" phylum="Arthropoda" rank="species" species="robusta">
<emphasis id="29AEEAA9E02ED749FEE2EA24FE4AF9C1" box="[268,384,1539,1565]" italics="true" pageId="27" pageNumber="236">P. robusta</emphasis>
Navás
</taxonomicName>
, and the most detailed descriptions are available for
<taxonomicName id="DCDA4D38E02ED749FBADEA24FB28F9C1" authority="Ris" authorityName="Ris" authorityYear="1912" box="[1091,1250,1539,1565]" class="Insecta" family="Lestoideidae" genus="Philoganga" kingdom="Animalia" order="Odonata" pageId="27" pageNumber="236" phylum="Arthropoda" rank="species" species="vetusta">
<emphasis id="29AEEAA9E02ED749FBADEA24FB78F9C1" box="[1091,1202,1540,1565]" italics="true" pageId="27" pageNumber="236">P. vetusta</emphasis>
Ris
</taxonomicName>
(
<bibRefCitation id="7F4B4B4AE02ED749FB1BEA23FA52F9C1" author="Asahina, S." box="[1269,1432,1539,1565]" pageId="27" pageNumber="236" pagination="323 - 326" refId="ref21609" refString="Asahina, S. (1967) Notes on two amphipterygid dragonflies from Southeast Asia. Deutsche Entomologische Zeitschrift, 14, 323 - 326. https: // doi. org / 10.1002 / mmnd. 19670140310" type="journal article" year="1967">Asahina 1967</bibRefCitation>
,
<bibRefCitation id="7F4B4B4AE02ED749FF79EA08FF37F99D" author="Xu, Q. - H." box="[151,253,1575,1602]" pageId="27" pageNumber="236" pagination="69 - 74" refId="ref27726" refString="Xu, Q. - H. (2016) Description of the final stadium larva of Philoganga vetusta Ris, with discussion of the taxonomic characters of the larvae of the genus Philoganga Kirby (Odonata: Philogangidae). International Journal of Odonatology, 19, 69 - 74. https: // doi. org / 10.1080 / 13887890.2016.1181009" type="journal article" year="2016">Xu 2016</bibRefCitation>
,
<bibRefCitation id="7F4B4B4AE02ED749FEE4EA08FEB4F99D" author="Ng, C. Y." box="[266,382,1575,1602]" pageId="27" pageNumber="236" refId="ref25340" refString="Ng, C. Y. (2024 e) Freshwater Macroinvertebrates in Hong Kong: Odonata: Philoganga vetusta. Available from: https: // www. biosch. hku. hk / ecology / staffhp / dd / macroinvertebrates / Odonata / Diphlebiidae / Philoganga _ vetusta. html (accessed 20 March 2024)" type="url" year="2024">Ng 2024e</bibRefCitation>
). The larvae (
<figureCitation id="83E12A3EE02ED749FDF2EA07FDA5F99D" box="[540,623,1575,1601]" captionStart="FIGURES 18–26. 18" captionStartId="6.[151,264,1811,1835]" captionTargetBox="[265,1336,167,1790]" captionTargetId="figure-21@6.[248,1339,153,1804]" captionTargetPageId="6" captionText="FIGURES 18–26. 18. Rhinagrion borneense, habitus (original drawing). 19. Philosina alba, habitus (modified from Zhang et al. 2011). 20. Rhinagrion mima, caudal gills: above heavily tracheated median gill; below right lateral gill with basal spines along mid-rib (modified from Lieftinck 1956). 21. Philosina alba caudal gills: above heavily tracheated median gill; below right lateral gill with basal spines along mid-rib (modified from Zhang et al. 2011) 22. Rhinagrion borneense, detail of mask (original drawing). 23. Philosina buchi, detail of mask (redrawn from Zhang et al. 2011). 24. Philoganga vetusta, habitus (adapted from Xu 2016). 25. Philoganga vetusta, details of caudal gills and antenna (redrawn from Xu 2016). 26. Philoganga montana, detail of mask (original drawing)." figureDoi="http://doi.org/10.5281/zenodo.13618331" httpUri="https://zenodo.org/record/13618331/files/figure.png" pageId="27" pageNumber="236">Fig. 24</figureCitation>
) are found among stones in clear wooded streams in hill country up to
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.
<bibRefCitation id="7F4B4B4AE02ED749FF1CEA6CFE43F9B9" author="Fraser, F. C." box="[242,393,1611,1638]" pageId="27" pageNumber="236" pagination="137 - 143" refId="ref23100" refString="Fraser, F. C. (1938) Additions to the Amphipterygidae (Order Odonata). Proceedings of the Royal Entomological Society of London, Series B, 7, 137 - 143. https: // doi. org / 10.1111 / j. 1365 - 3113.1938. tb 01267. x" type="journal article" year="1938">Fraser (1938)</bibRefCitation>
reported
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<emphasis id="29AEEAA9E02ED749FE1DEA6CFDA5F9B9" box="[499,623,1612,1637]" italics="true" pageId="27" pageNumber="236">P. montana</emphasis>
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as inhabiting “...torrential mountain streams, clinging to the underside of stones or rocks or hiding up in clefts of rocks.”. Mature larvae are easily distinguished by their massive head, eight segmented antennae (
<figureCitation id="83E12A3EE02ED749FE63EAB3FE29F972" box="[397,483,1683,1710]" captionStart="FIGURES 18–26. 18" captionStartId="6.[151,264,1811,1835]" captionTargetBox="[265,1336,167,1790]" captionTargetId="figure-21@6.[248,1339,153,1804]" captionTargetPageId="6" captionText="FIGURES 18–26. 18. Rhinagrion borneense, habitus (original drawing). 19. Philosina alba, habitus (modified from Zhang et al. 2011). 20. Rhinagrion mima, caudal gills: above heavily tracheated median gill; below right lateral gill with basal spines along mid-rib (modified from Lieftinck 1956). 21. Philosina alba caudal gills: above heavily tracheated median gill; below right lateral gill with basal spines along mid-rib (modified from Zhang et al. 2011) 22. Rhinagrion borneense, detail of mask (original drawing). 23. Philosina buchi, detail of mask (redrawn from Zhang et al. 2011). 24. Philoganga vetusta, habitus (adapted from Xu 2016). 25. Philoganga vetusta, details of caudal gills and antenna (redrawn from Xu 2016). 26. Philoganga montana, detail of mask (original drawing)." figureDoi="http://doi.org/10.5281/zenodo.13618331" httpUri="https://zenodo.org/record/13618331/files/figure.png" pageId="27" pageNumber="236">Fig. 25</figureCitation>
), (although
<bibRefCitation id="7F4B4B4AE02ED749FD83EAB3FCD7F971" author="Asahina, S." box="[621,797,1683,1709]" pageId="27" pageNumber="236" pagination="323 - 326" refId="ref21609" refString="Asahina, S. (1967) Notes on two amphipterygid dragonflies from Southeast Asia. Deutsche Entomologische Zeitschrift, 14, 323 - 326. https: // doi. org / 10.1002 / mmnd. 19670140310" type="journal article" year="1967">Asahina (1967)</bibRefCitation>
reported some specimens of
<taxonomicName id="DCDA4D38E02ED749FB9EEAB4FB17F971" authorityName="Ris" authorityYear="1912" box="[1136,1245,1684,1709]" class="Insecta" family="Lestoideidae" genus="Philoganga" kingdom="Animalia" order="Odonata" pageId="27" pageNumber="236" phylum="Arthropoda" rank="species" species="vetusta">
<emphasis id="29AEEAA9E02ED749FB9EEAB4FB17F971" box="[1136,1245,1684,1709]" italics="true" pageId="27" pageNumber="236">P. vetusta</emphasis>
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with only seven antennal segments), long tapered saccoid terminal gills and large size. As far as is known all species bear a cluster of long strong subocular spines on the anterior gena; the prementum (
<figureCitation id="83E12A3EE02ED749FC76EAFBFC23F929" box="[920,1001,1755,1781]" captionStart="FIGURES 18–26. 18" captionStartId="6.[151,264,1811,1835]" captionTargetBox="[265,1336,167,1790]" captionTargetId="figure-21@6.[248,1339,153,1804]" captionTargetPageId="6" captionText="FIGURES 18–26. 18. Rhinagrion borneense, habitus (original drawing). 19. Philosina alba, habitus (modified from Zhang et al. 2011). 20. Rhinagrion mima, caudal gills: above heavily tracheated median gill; below right lateral gill with basal spines along mid-rib (modified from Lieftinck 1956). 21. Philosina alba caudal gills: above heavily tracheated median gill; below right lateral gill with basal spines along mid-rib (modified from Zhang et al. 2011) 22. Rhinagrion borneense, detail of mask (original drawing). 23. Philosina buchi, detail of mask (redrawn from Zhang et al. 2011). 24. Philoganga vetusta, habitus (adapted from Xu 2016). 25. Philoganga vetusta, details of caudal gills and antenna (redrawn from Xu 2016). 26. Philoganga montana, detail of mask (original drawing)." figureDoi="http://doi.org/10.5281/zenodo.13618331" httpUri="https://zenodo.org/record/13618331/files/figure.png" pageId="27" pageNumber="236">Fig. 26</figureCitation>
) is broad, with rounded lateral margins bearing short spines in their anterior half, a slightly produced anterior median lobe with short median cleft. I was able to make direct comparisons with an exuvia of
<taxonomicName id="DCDA4D38E02ED749FD26EB04FC8EF8E1" box="[712,836,1828,1853]" class="Insecta" family="Lestoideidae" genus="Philoganga" kingdom="Animalia" order="Odonata" pageId="27" pageNumber="236" phylum="Arthropoda" rank="species" species="montana">
<emphasis id="29AEEAA9E02ED749FD26EB04FC8EF8E1" box="[712,836,1828,1853]" italics="true" pageId="27" pageNumber="236">P. montana</emphasis>
</taxonomicName>
collected in
<collectingCountry id="63CD762BE02ED749FC3EEB03FBD9F8E1" box="[976,1043,1827,1853]" name="Nepal" pageId="27" pageNumber="236">Nepal</collectingCountry>
by S.G. Butler (see
<bibRefCitation id="7F4B4B4AE02ED749FB1FEB04FF19F8BD" author="Kemp, R. G. &amp; Butler, S. G." pageId="27" pageNumber="236" pagination="85 - 86" refId="ref24147" refString="Kemp, R. G. &amp; Butler, S. G. (2001) Some dragonfly records from Phewa Tal, Pokhara, Nepal with notes on Philoganga montana (Selys) (Zygoptera: Amphipterygidae). Notulae odonatologicae, 5, 85 - 86." type="journal article" year="2001">Kemp &amp; Butler 2001</bibRefCitation>
), which had 8-segmented antennae, and, aside from its huge size (total length excluding antennae and caudal gills
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) it differed only slightly from the allopatric
<taxonomicName id="DCDA4D38E02ED749FCE9EB4CFCB8F859" authorityName="Ris" authorityYear="1912" box="[775,882,1900,1925]" class="Insecta" family="Lestoideidae" genus="Philoganga" kingdom="Animalia" order="Odonata" pageId="27" pageNumber="236" phylum="Arthropoda" rank="species" species="vetusta">
<emphasis id="29AEEAA9E02ED749FCE9EB4CFCB8F859" box="[775,882,1900,1925]" italics="true" pageId="27" pageNumber="236">P. vetusta</emphasis>
</taxonomicName>
.
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Family
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The family has just one genus including two species, with the larva of
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<emphasis id="29AEEAA9E02ED749FC0BE82BFADBFBF9" box="[997,1297,1035,1061]" italics="true" pageId="27" pageNumber="236">Mesopodagrion tibetanum</emphasis>
McLachlan
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having been described (
<bibRefCitation id="7F4B4B4AE02ED749FE70E80FFDC8FB95" author="Yu, X." box="[414,514,1071,1097]" pageId="27" pageNumber="236" pagination="275 - 282" refId="ref28104" refString="Yu, X. (2016) A description of the larva of Mesopodagrion tibetanum australe (Odonata: &quot; Megapodagrionidae &quot;). International Journal of Odonatology, 19 (4), 275 - 282. https: // doi. org / 10.1080 / 13887890.2016.1259663" type="journal article" year="2016">Yu 2016</bibRefCitation>
). They inhabit mountain streams from
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in southern
<collectingCountry id="63CD762BE02ED749FB50E80FFACEFB95" box="[1214,1284,1071,1097]" name="China" pageId="27" pageNumber="236">China</collectingCountry>
, and northern
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and
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. They bear unique lateral tufted projections arising from the postocular lobes (
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) and the prementum is shout and broad, with convex lateral margins only slightly expanded anteriorly and with a shallow but distinct anterior median lobe with short median cleft (
<figureCitation id="83E12A3EE02ED749FCC8E8BBFCB0FB69" box="[806,890,1179,1205]" captionStart="FIGURES 12–17. 12" captionStartId="5.[151,264,1903,1927]" captionTargetBox="[190,1400,160,1874]" captionTargetId="figure-15@5.[178,1409,151,1889]" captionTargetPageId="5" captionText="FIGURES 12–17. 12. Mesopodagrion tibetanum, habitus with detail of right profile of head (based on Yu 2016). 13. M. tibetanum, detail of mask (redrawn from Yu 2016). 14. Podolestes orientalis, habitus (from Choong &amp; Orr 2010). 15. Podolestes orientalis, detail of mask (from Choong &amp; Orr 2010). 16. Rhipidolestes aculeatus, habitus and detail of lateral and median gill (modified from Asahina 1994). 17. Rhipidolestes aculeatus, detail of mask (modified from Ishida 1996)." figureDoi="http://doi.org/10.5281/zenodo.13618327" httpUri="https://zenodo.org/record/13618327/files/figure.png" pageId="27" pageNumber="236">Fig. 13</figureCitation>
). As in
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, the caudal gills are foliate and arranged in a horizontal fan (
<figureCitation id="83E12A3EE02ED749FDE2E89FFD97FB05" box="[524,605,1215,1241]" captionStart="FIGURES 12–17. 12" captionStartId="5.[151,264,1903,1927]" captionTargetBox="[190,1400,160,1874]" captionTargetId="figure-15@5.[178,1409,151,1889]" captionTargetPageId="5" captionText="FIGURES 12–17. 12. Mesopodagrion tibetanum, habitus with detail of right profile of head (based on Yu 2016). 13. M. tibetanum, detail of mask (redrawn from Yu 2016). 14. Podolestes orientalis, habitus (from Choong &amp; Orr 2010). 15. Podolestes orientalis, detail of mask (from Choong &amp; Orr 2010). 16. Rhipidolestes aculeatus, habitus and detail of lateral and median gill (modified from Asahina 1994). 17. Rhipidolestes aculeatus, detail of mask (modified from Ishida 1996)." figureDoi="http://doi.org/10.5281/zenodo.13618327" httpUri="https://zenodo.org/record/13618327/files/figure.png" pageId="27" pageNumber="236">Fig. 12</figureCitation>
) but there are numerous points of difference between
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and
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(
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), the only argiolestid genus known from mainland Asia.
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This monotypic genus is known only from north-east Luzon,
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is the largest species of the family and the distinctive larvae were recently described on the basis of four exuviae (
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), determined by supposition. They are notable for their size and for having extremely numerous and well-developed spines on the genae, and 2–3 rows of spines on the outer face of the mandible, sometimes arranged in a crown-like circlet. The head bears numerous small tubercles and the postocular lobes are shallow, somewhat reminiscent of
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<emphasis id="29AEEAA9E02FD748FEAEEAE7FE03F93D" box="[320,457,1735,1761]" italics="true" pageId="26" pageNumber="235">Anisopleura</emphasis>
</taxonomicName>
but no close relationship is inferred (
<figureCitation id="83E12A3EE02FD748FC80EAE7FC08F93D" box="[878,962,1735,1761]" captionStart="FIGURES 61–66. 61" captionStartId="12.[151,264,1841,1865]" captionTargetBox="[273,1314,156,1828]" captionTargetId="figure-21@12.[268,1319,151,1834]" captionTargetPageId="12" captionText="FIGURES 61–66. 61. Dysphaea gloriosa: detail of spines on genae and postocular lobe; underside (left), upperside (right) (modified from Nguyen et al. 2024). 62. Heterophaea barbata: dorsal view of head, showing (a) outer mandibular spines (b) genal spines partially visible and (c) right antenna (from Orr &amp; Hämäläinen 2024). 63. Anisopleura subplatystyla, dorsal view of head (original drawing). 64. Bayadera strigata, dorsal view of head (drawn from Yang &amp; Orr 2024). 65. Euphaea decorata, dorsal view of head. (drawn from Xu 2017). 66. Bayadera strigata, left labial palp with detail showing wavy outer margin of outer process of palpal lobe (arrowed), (drawn from Yang &amp; Orr 2024)." figureDoi="http://doi.org/10.5281/zenodo.13618343" httpUri="https://zenodo.org/record/13618343/files/figure.png" pageId="26" pageNumber="235">Fig. 62</figureCitation>
). The antennae are 12–15 segmented (V.J. Kalkman pers comm.), more than any other known odonate larva. Various parts of the body bear dense lines of dark claviform setae and long filiform setae. The habitat is presumed to be under stones and boulder in clear swift flowing streams at around
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600–
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.
</paragraph>
</subSubSection>
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287
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<paragraph id="1B6536BBE02FD748FF79ECB6FF35FF6D" blockId="26.[151,1437,150,1473]" box="[151,255,150,177]" pageId="26" pageNumber="235">
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<taxonomicName id="DCDA4D38E02FD748FF79ECB6FF35FF6D" ID-CoL="4G8X" authorityName="Selys" authorityYear="1840" box="[151,255,150,177]" class="Insecta" family="Euphaeidae" genus="Euphaea" kingdom="Animalia" order="Odonata" pageId="26" pageNumber="235" phylum="Arthropoda" rank="genus">
<emphasis id="29AEEAA9E02FD748FF79ECB6FF35FF6D" bold="true" box="[151,255,150,177]" italics="true" pageId="26" pageNumber="235">Euphaea</emphasis>
</taxonomicName>
</heading>
</paragraph>
</subSubSection>
<subSubSection id="53C06530E02FD748FF79EC9BFB08FA1D" pageId="26" pageNumber="235" type="discussion">
<paragraph id="1B6536BBE02FD748FF79EC9BFC70FDE1" blockId="26.[151,1437,150,1473]" pageId="26" pageNumber="235">
With 35 species ranging from
<collectingCountry id="63CD762BE02FD748FE19EC9BFDFDFF09" box="[503,567,187,213]" name="Japan" pageId="26" pageNumber="235">Japan</collectingCountry>
to
<collectingCountry id="63CD762BE02FD748FD8EEC9BFD1AFF09" box="[608,720,187,213]" name="Sri Lanka" pageId="26" pageNumber="235">Sri Lanka</collectingCountry>
in the west to the Alor Archipelago in the Lesser Sundas and the
<collectingCountry id="63CD762BE02FD748FF2FECFFFEF5FF25" box="[193,319,223,249]" name="Philippines" pageId="26" pageNumber="235">Philippines</collectingCountry>
in the east,
<taxonomicName id="DCDA4D38E02FD748FE27ECFFFDE7FF25" authorityName="Selys" authorityYear="1840" box="[457,557,223,249]" class="Insecta" family="Euphaeidae" genus="Euphaea" kingdom="Animalia" order="Odonata" pageId="26" pageNumber="235" phylum="Arthropoda" rank="genus">
<emphasis id="29AEEAA9E02FD748FE27ECFFFDE7FF25" box="[457,557,223,249]" italics="true" pageId="26" pageNumber="235">Euphaea</emphasis>
</taxonomicName>
is by far the most speciose genus of the family and the most morphologically diverse in the adult stage. A habitus of the larvae of
<taxonomicName id="DCDA4D38E02FD748FD36ED23FC72FEC1" authority="Hagen" authorityName="Hagen" box="[728,952,259,285]" class="Insecta" family="Euphaeidae" genus="Euphaea" kingdom="Animalia" order="Odonata" pageId="26" pageNumber="235" phylum="Arthropoda" rank="species" species="splendens">
<emphasis id="29AEEAA9E02FD748FD36ED23FCAFFEC1" box="[728,869,259,285]" italics="true" pageId="26" pageNumber="235">E. splendens</emphasis>
Hagen
</taxonomicName>
, first mentioned by
<bibRefCitation id="7F4B4B4AE02FD748FB7BED23FAF8FEC1" author="Hagen, H." box="[1173,1330,259,285]" pageId="26" pageNumber="235" pagination="lxv - lxvii" refId="ref23289" refString="Hagen, H. (1880) Essai d'un synopsis des larves de Calopterygines. [Comptes-rendus de la Socie´te´Entomologique de Belgique. Seance du 1 er Mai 1880]. Annales de la Societe entomologique de Belgique, 23, lxv - lxvii." type="journal article" year="1880">Hagen (1880)</bibRefCitation>
, was first illustrated with fair accuracy in
<bibRefCitation id="7F4B4B4AE02FD748FDE9ED07FD7FFE9D" author="Packard, A. S." box="[519,693,295,321]" pageId="26" pageNumber="235" refId="ref26228" refString="Packard, A. S. (1898) A textbook of entomology, including the anatomy, physiology, embryology and metamorphoses of insects. Macmillan, London, xvii + 729 pp. [https: // www. biodiversitylibrary. org / item / 16837 # page / 493 / mode / 1 up]" type="book" year="1898">Packard (1898)</bibRefCitation>
and
<bibRefCitation id="7F4B4B4AE02FD748FD1EED07FCA3FE9D" author="Ris, F." box="[752,873,295,321]" pageId="26" pageNumber="235" pagination="157 - 183" refId="ref26559" refString="Ris, F. (1912) Uber Odonaten von Java und Krakatau gesammelt von Edward Jacobson. Tijdschrift voor Entomologie, 55, 157 - 183, pls. 6 - 8 excl." type="journal article" year="1912">Ris (1912)</bibRefCitation>
described and figured very accurately diagnostic features of
<taxonomicName id="DCDA4D38E02FD748FEFDED6BFE37FEB9" authority="Rambur" authorityName="Rambur" authorityYear="1842" box="[275,509,331,357]" class="Insecta" family="Euphaeidae" genus="Euphaea" kingdom="Animalia" order="Odonata" pageId="26" pageNumber="235" phylum="Arthropoda" rank="species" species="variegata">
<emphasis id="29AEEAA9E02FD748FEFDED6BFE57FEB8" box="[275,413,331,356]" italics="true" pageId="26" pageNumber="235">E. variegata</emphasis>
Rambur
</taxonomicName>
larva from Java.
<bibRefCitation id="7F4B4B4AE02FD748FD53ED6BFCADFEB9" author="Tillyard, R. J." box="[701,871,331,357]" pageId="26" pageNumber="235" refId="ref27017" refString="Tillyard, R. J. (1917) The biology of dragonflies (Odonata or Paraneuroptera). University Press, Cambridge, xii + 396 pp., pls. 1 - 4 and 2 folded attachments excl. https: // doi. org / 10.5962 / bhl. title. 27401" type="book" year="1917">Tillyard (1917)</bibRefCitation>
depicted the lateral habitus of
<taxonomicName id="DCDA4D38E02FD748FB55ED6BFA56FEB8" authorityName="Rambur" authorityYear="1842" box="[1211,1436,331,357]" class="Insecta" family="Euphaeidae" genus="Euphaea" kingdom="Animalia" order="Odonata" pageId="26" pageNumber="235" phylum="Arthropoda" rank="species" species="variegata">
<emphasis id="29AEEAA9E02FD748FB55ED6BFAD5FEB9" box="[1211,1311,331,357]" italics="true" pageId="26" pageNumber="235">Euphaea</emphasis>
?
<emphasis id="29AEEAA9E02FD748FADFED6BFA56FEB8" box="[1329,1436,331,356]" italics="true" pageId="26" pageNumber="235">variegata</emphasis>
</taxonomicName>
(as
<taxonomicName id="DCDA4D38E02FD748FF2EED4FFE4AFE55" box="[192,384,367,393]" class="Insecta" family="Euphaeidae" genus="Pseudophaea" kingdom="Animalia" order="Odonata" pageId="26" pageNumber="235" phylum="Arthropoda" rank="species" species="undetermined">
<emphasis id="29AEEAA9E02FD748FF2EED4FFE9DFE55" box="[192,343,367,393]" italics="true" pageId="26" pageNumber="235">Pseudophaea</emphasis>
sp.
</taxonomicName>
) based on a photograph sent by F. Ris.
<bibRefCitation id="7F4B4B4AE02FD748FCA8ED4FFC27FE55" author="Fraser, F. C." box="[838,1005,367,393]" pageId="26" pageNumber="235" pagination="288 - 301" refId="ref22972" refString="Fraser, F. C. (1929 b) Indian dragonflies, Part XXXII. Journal of the Bombay Natural History Society 33, 288 - 301, pls. 1 - 4 excl." type="journal article" year="1929">Fraser (1929b)</bibRefCitation>
crudely figured the ventral habitus of
<taxonomicName id="DCDA4D38E02FD748FF79EDB3FEEFFE71" box="[151,293,403,429]" class="Insecta" family="Euphaeidae" genus="Euphaea" kingdom="Animalia" order="Odonata" pageId="26" pageNumber="235" phylum="Arthropoda" rank="species" species="splendens">
<emphasis id="29AEEAA9E02FD748FF79EDB3FEEFFE71" box="[151,293,403,429]" italics="true" pageId="26" pageNumber="235">E. splendens</emphasis>
</taxonomicName>
(as
<emphasis id="29AEEAA9E02FD748FEB8EDB3FE3EFE71" box="[342,500,403,429]" italics="true" pageId="26" pageNumber="235">
<taxonomicName id="DCDA4D38E02FD748FEB8EDB3FE3AFE71" authorityName="Kirby" authorityYear="1890" box="[342,496,403,429]" class="Insecta" family="Euphaeidae" genus="Pseudophaea" higherTaxonomySource="CoL,GBIF" kingdom="Animalia" order="Odonata" pageId="26" pageNumber="235" phylum="Arthropoda" rank="genus">Pseudophaea</taxonomicName>
,
</emphasis>
wrongly depicting exaggerated abdominal gills on S1–7) and the dorsal habitus of
<taxonomicName id="DCDA4D38E02FD748FF79ED97FEB1FE0D" authority="(Laidlaw)" baseAuthorityName="Laidlaw" baseAuthorityYear="1920" box="[151,379,439,465]" class="Insecta" family="Euphaeidae" genus="Euphaea" kingdom="Animalia" order="Odonata" pageId="26" pageNumber="235" phylum="Arthropoda" rank="species" species="fraseri">
<emphasis id="29AEEAA9E02FD748FF79ED97FEC9FE0D" box="[151,259,439,465]" italics="true" pageId="26" pageNumber="235">E. fraseri</emphasis>
(Laidlaw)
</taxonomicName>
which also suggests the abdominal gills are on S 1–7 which are incorrectly described as ‘S’ shaped in both species, which may have been an artefact of preservation. A slightly improved version of the
<taxonomicName id="DCDA4D38E02FD748FA6AEDFBFF28FDC5" baseAuthorityName="Laidlaw" baseAuthorityYear="1920" class="Insecta" family="Euphaeidae" genus="Euphaea" kingdom="Animalia" order="Odonata" pageId="26" pageNumber="235" phylum="Arthropoda" rank="species" species="fraseri">
<emphasis id="29AEEAA9E02FD748FA6AEDFBFF28FDC5" italics="true" pageId="26" pageNumber="235">E. fraseri</emphasis>
</taxonomicName>
habitus was depicted in
<bibRefCitation id="7F4B4B4AE02FD748FDEEEDDFFD57FDC5" author="Fraser, F. C." box="[512,669,511,537]" pageId="26" pageNumber="235" pagination="1 - 4" refId="ref23053" refString="Fraser, F. C. (1934) The fauna of British India, including Ceylon and Burma. Odonata. Vol. II. Taylor &amp; Francis Ltd., London, xxiii + 398 pp., pls. 1 - 4 excl." type="book chapter" year="1934">Fraser (1934)</bibRefCitation>
. Some structural details, photographs of the habitus and ecological information were provided for
<taxonomicName id="DCDA4D38E02FD748FE1CEE03FDB5FDE1" box="[498,639,547,573]" class="Insecta" family="Euphaeidae" genus="Euphaea" kingdom="Animalia" order="Odonata" pageId="26" pageNumber="235" phylum="Arthropoda" rank="species" species="splendens">
<emphasis id="29AEEAA9E02FD748FE1CEE03FDB5FDE1" box="[498,639,547,573]" italics="true" pageId="26" pageNumber="235">E. splendens</emphasis>
</taxonomicName>
larva by
<bibRefCitation id="7F4B4B4AE02FD748FD06EE03FC7CFDE1" author="St Quentin, D." box="[744,950,547,573]" pageId="26" pageNumber="235" pagination="113 - 124" refId="ref26637" refString="St Quentin, D. (1973) Contributions to the ecology of the larvae some Odonata from Ceylon. Bulletin of the Fisheries Research Station, Sri Lanka (Ceylon), 24, 113 - 124." type="journal article" year="1973">St Quentin (1973)</bibRefCitation>
.
</paragraph>
<paragraph id="1B6536BBE02FD748FF29EE67FB6FFC79" blockId="26.[151,1437,150,1473]" pageId="26" pageNumber="235">
Other species that have been described and/or illustrated include:
<taxonomicName id="DCDA4D38E02FD748FC2FEE67FB53FDBD" authority="Hagen" authorityName="Hagen" box="[961,1177,583,609]" class="Insecta" family="Euphaeidae" genus="Euphaea" kingdom="Animalia" order="Odonata" pageId="26" pageNumber="235" phylum="Arthropoda" rank="species" species="decorata">
<emphasis id="29AEEAA9E02FD748FC2FEE67FB8DFDBD" box="[961,1095,583,609]" italics="true" pageId="26" pageNumber="235">E. decorata</emphasis>
Hagen
</taxonomicName>
(
<figureCitation id="83E12A3EE02FD748FB42EE67FAC8FDBD" box="[1196,1282,582,609]" captionStart="FIGURES 57–60. 57" captionStartId="11.[151,264,1933,1957]" captionTargetBox="[193,1384,155,1907]" captionTargetId="figure-15@11.[188,1400,151,1917]" captionTargetPageId="11" captionText="FIGURES 57–60. 57. Dysphaea gloriosa, habitus (drawn from Nguyen et al. 2024). 58. Bayadera strigata, habitus (drawn from Yang &amp; Orr 2024). 59. Euphaea decorata, habitus. (drawn from Xu 2017). 60. Anisopleura furcata, habitus (drawn from Keetapithchayakul et al. 2024)." figureDoi="http://doi.org/10.5281/zenodo.13618341" httpUri="https://zenodo.org/record/13618341/files/figure.png" pageId="26" pageNumber="235">Fig. 59</figureCitation>
), habitus and diagnostic details figured (
<bibRefCitation id="7F4B4B4AE02FD748FE26EE4BFD49FD59" author="Tam, T. - W. &amp; Leung, K. K. &amp; Kwan, B. S. P. &amp; Wu, K. K. Y. &amp; Tang, S. S. H. &amp; So, I. W. Y. &amp; Cheng, J. C. Y. &amp; Yuen, E. F. M. &amp; Tsang, Y. - M. T. &amp; Hui, W. L." box="[456,643,619,645]" pageId="26" pageNumber="235" refId="ref27108" refString="Tam, T. - W., Leung, K. K., Kwan, B. S. P., Wu, K. K. Y., Tang, S. S. H., So, I. W. Y., Cheng, J. C. Y., Yuen, E. F. M., Tsang, Y. - M. T. &amp; Hui, W. L. (2011) The dragonflies of Hong Kong. Agriculture, Fisheries and Conservation Department, Hong Kong, 368 pp." type="book" year="2011">
Tam
<emphasis id="29AEEAA9E02FD748FDEBEE4BFDF2FD59" box="[517,568,619,645]" italics="true" pageId="26" pageNumber="235">et al</emphasis>
. 2011
</bibRefCitation>
,
<bibRefCitation id="7F4B4B4AE02FD748FD7EEE4BFD33FD59" author="Xu, Q. - H." box="[656,761,619,645]" pageId="26" pageNumber="235" pagination="595 - 599" refId="ref27788" refString="Xu, Q. - H. (2017) The final stadium larva of Euphaea decorata Hagen in Selys, 1853, from Fujian, China (Odonata: Zygoptera: Euphaeidae). Zootaxa, 4244 (4), 595 - 599. https: // doi. org / 10.11646 / zootaxa. 4244.4.10" type="journal article" year="2017">Xu 2017</bibRefCitation>
);
<taxonomicName id="DCDA4D38E02FD748FCE0EE4BFC2BFD59" authority="Hagen" authorityName="Hagen" box="[782,993,619,645]" class="Insecta" family="Euphaeidae" genus="Euphaea" kingdom="Animalia" order="Odonata" pageId="26" pageNumber="235" phylum="Arthropoda" rank="species" species="formosa">
<emphasis id="29AEEAA9E02FD748FCE0EE4BFC46FD59" box="[782,908,619,645]" italics="true" pageId="26" pageNumber="235">E. formosa</emphasis>
Hagen
</taxonomicName>
, diagnostic details figured (
<bibRefCitation id="7F4B4B4AE02FD748FAF1EE4BFEC4FD75" author="Matsuki, K. &amp; Lien, J. C." pageId="26" pageNumber="235" pagination="15 - 26" refId="ref24810" refString="Matsuki, K. &amp; Lien, J. C. (1978) Descriptions of the larvae of three families of Zygoptera breeding in the streams of Taiwan (Synlestidae, Euphaeidae &amp; Calopterygidae). Tombo - Acta Odonatologica Japonica, 21, 15 - 26." type="journal article" year="1978">Matsuki &amp; Lien 1978</bibRefCitation>
);
<taxonomicName id="DCDA4D38E02FD748FECCEEAFFE01FD75" authority="Selys" authorityName="Selys" authorityYear="1859" box="[290,459,655,681]" class="Insecta" family="Euphaeidae" genus="Euphaea" kingdom="Animalia" order="Odonata" pageId="26" pageNumber="235" phylum="Arthropoda" rank="species" species="impar">
<emphasis id="29AEEAA9E02FD748FECCEEAFFE4EFD74" box="[290,388,655,680]" italics="true" pageId="26" pageNumber="235">E. impar</emphasis>
Selys
</taxonomicName>
, photograph of habitus (
<bibRefCitation id="7F4B4B4AE02FD748FD32EEAFFC7EFD75" author="Ngiam, R. W. J. &amp; Ng, M. F. C." box="[732,948,655,681]" pageId="26" pageNumber="235" refId="ref25398" refString="Ngiam, R. W. J. &amp; Ng, M. F. C. (2022) A photographic field guide to the dragonflies &amp; damselflies of Singapore. John Beaufoy Publishing, Oxford, 340 pp." type="book" year="2022">Ngiam &amp; Ng 2022</bibRefCitation>
);
<taxonomicName id="DCDA4D38E02FD748FC26EEAFFB5FFD75" authority="Selys" authorityName="Selys" authorityYear="1859" box="[968,1173,655,681]" class="Insecta" family="Euphaeidae" genus="Euphaea" kingdom="Animalia" order="Odonata" pageId="26" pageNumber="235" phylum="Arthropoda" rank="species" species="ochracea">
<emphasis id="29AEEAA9E02FD748FC26EEAFFB85FD75" box="[968,1103,655,681]" italics="true" pageId="26" pageNumber="235">E. ochracea</emphasis>
Selys
</taxonomicName>
, habitus and diagnostic details, the former suggests the filament of the caudal gills is constricted at the base, which seems likely to be an artefact of preservation or artistic licence (
<bibRefCitation id="7F4B4B4AE02FD748FD91EEF7FCE8FD2D" author="Matsuki, K." box="[639,802,727,753]" pageId="26" pageNumber="235" pagination="23 - 24" refId="ref24774" refString="Matsuki, K. (2001) Description of the last instar larva Euphaea ochracea Selys, 1859 from Thailand (Euphaeidae). Tombo - Acta Odonatologica Japonica, 43, 23 - 24." type="journal article" year="2001">Matsuki 2001</bibRefCitation>
);
<taxonomicName id="DCDA4D38E02FD748FCD6EEF7FC2CFD2D" authority="Selys" authorityName="Selys" authorityYear="1853" box="[824,998,727,753]" class="Insecta" family="Euphaeidae" genus="Euphaea" kingdom="Animalia" order="Odonata" pageId="26" pageNumber="235" phylum="Arthropoda" rank="species" species="opaca">
<emphasis id="29AEEAA9E02FD748FCD6EEF7FC54FD2C" box="[824,926,727,752]" italics="true" pageId="26" pageNumber="235">E. opaca</emphasis>
Selys
</taxonomicName>
, photograph habitus (
<bibRefCitation id="7F4B4B4AE02FD748FB31EEF7FAA5FD2D" author="Zhang, H. - M." box="[1247,1391,727,753]" pageId="26" pageNumber="235" refId="ref28336" refString="Zhang, H. - M. (2019) Dragonflies and Damselflies of China. Vols. 1 &amp; 2. Chong Qing University Press, ChongQing, 14 + 1460 pp. [bilingual, in Chinese and English]" type="book" year="2019">Zhang 2019</bibRefCitation>
);
<taxonomicName id="DCDA4D38E02FD748FA6AEEF7FE9FFCC9" authority="Hagen" authorityName="Hagen" class="Insecta" family="Euphaeidae" genus="Euphaea" kingdom="Animalia" order="Odonata" pageId="26" pageNumber="235" phylum="Arthropoda" rank="species" species="refulgens">
<emphasis id="29AEEAA9E02FD748FA6AEEF7FF34FCC9" italics="true" pageId="26" pageNumber="235">E. refulgens</emphasis>
Hagen
</taxonomicName>
, habitus figured (
<bibRefCitation id="7F4B4B4AE02FD748FDCCEEDBFC8DFCC9" author="Needham, J. G. &amp; Gyger, M. K." box="[546,839,763,789]" pageId="26" pageNumber="235" pagination="239 - 314" refId="ref25055" refString="Needham, J. G. &amp; Gyger, M. K. (1939) The Odonata of the Philippines, II. Suborder Zygoptera. Philippine Journal of Science, 70, 239 - 314, pls. 11 - 22 excl." type="journal article" year="1939">Needham &amp; Gyger 1939</bibRefCitation>
);
<taxonomicName id="DCDA4D38E02FD748FCB0EEDBFB8DFCC9" authority="Selys" authorityName="Selys" authorityYear="1873" box="[862,1095,763,789]" class="Insecta" family="Euphaeidae" genus="Euphaea" kingdom="Animalia" order="Odonata" pageId="26" pageNumber="235" phylum="Arthropoda" rank="species" species="subcostalis">
<emphasis id="29AEEAA9E02FD748FCB0EEDBFC37FCC9" box="[862,1021,763,789]" italics="true" pageId="26" pageNumber="235">E. subcostalis</emphasis>
Selys
</taxonomicName>
, habitus figured (as
<taxonomicName id="DCDA4D38E02FD748FAD7EEDBFF73FCE5" class="Insecta" family="Euphaeidae" genus="Euphaea" kingdom="Animalia" order="Odonata" pageId="26" pageNumber="235" phylum="Arthropoda" rank="species" species="undetermined">
<emphasis id="29AEEAA9E02FD748FAD7EEDBFA57FCC9" box="[1337,1437,763,789]" italics="true" pageId="26" pageNumber="235">Euphaea</emphasis>
sp.
</taxonomicName>
),
<bibRefCitation id="7F4B4B4AE02FD748FF3FEF3FFE85FCE5" author="Orr, A. G." box="[209,335,799,825]" pageId="26" pageNumber="235" refId="ref25717" refString="Orr, A. G. (2003) A guide to the Dragonflies of Borneo: their identification and biology. Natural History Publications (Borneo), Kota Kinabalu, x + 195 pp." type="book" year="2003">Orr (2003)</bibRefCitation>
;
<taxonomicName id="DCDA4D38E02FD748FEB0EF3FFD5AFCE5" authority="Kimmins 1936" authorityName="Kimmins" authorityYear="1936" box="[350,656,799,825]" class="Insecta" family="Euphaeidae" genus="Euphaea" kingdom="Animalia" order="Odonata" pageId="26" pageNumber="235" phylum="Arthropoda" rank="species" species="superba">
<emphasis id="29AEEAA9E02FD748FEB0EF3FFE13FCE5" box="[350,473,799,825]" italics="true" pageId="26" pageNumber="235">E. superba</emphasis>
Kimmins 1936
</taxonomicName>
, habitus and diagnostic details figured (
<bibRefCitation id="7F4B4B4AE02FD748FBB5EF3FFAC5FCE5" author="Wu, X. - J. &amp; Liu, Z. &amp; Chen, Y. &amp; Wang, B. - X." box="[1115,1295,799,825]" pageId="26" pageNumber="235" pagination="585 - 592" refId="ref27587" refString="Wu, X. - J., Liu, Z., Chen, Y. &amp; Wang, B. - X. (2019) Description of larva of Euphaea superba Kimmins, 1936 (Odonata: Zygoptera: Euphaeidae) from China. Zootaxa, 4545 (4), 585 - 592. https: // doi. org / 10.11646 / zootaxa. 4545.4.9" type="journal article" year="2019">
Wu
<emphasis id="29AEEAA9E02FD748FB60EF3FFB08FCE5" box="[1166,1218,799,825]" italics="true" pageId="26" pageNumber="235">et al</emphasis>
. 2019
</bibRefCitation>
);
<taxonomicName id="DCDA4D38E02FD748FACBEF3FFF1CFC81" authority="Selys" authorityName="Selys" authorityYear="1859" class="Insecta" family="Euphaeidae" genus="Euphaea" kingdom="Animalia" order="Odonata" pageId="26" pageNumber="235" phylum="Arthropoda" rank="species" species="tricolor">
<emphasis id="29AEEAA9E02FD748FACBEF3FFA57FCE5" box="[1317,1437,799,825]" italics="true" pageId="26" pageNumber="235">E. tricolor</emphasis>
Selys
</taxonomicName>
, habitus in life,
<bibRefCitation id="7F4B4B4AE02FD748FE62EF63FDC2FC81" author="Orr, A. G." box="[396,520,835,861]" pageId="26" pageNumber="235" refId="ref25717" refString="Orr, A. G. (2003) A guide to the Dragonflies of Borneo: their identification and biology. Natural History Publications (Borneo), Kota Kinabalu, x + 195 pp." type="book" year="2003">Orr (2003)</bibRefCitation>
;
<taxonomicName id="DCDA4D38E02FD748FDFAEF63FCA2FC81" authority="Oguma" authorityName="Oguma" box="[532,872,835,861]" class="Insecta" family="Euphaeidae" genus="Euphaea" kingdom="Animalia" order="Odonata" pageId="26" pageNumber="235" phylum="Arthropoda" rank="species" species="yayeyamana">
<emphasis id="29AEEAA9E02FD748FDFAEF63FCC6FC80" box="[532,780,835,861]" italics="true" pageId="26" pageNumber="235">Euphaea yayeyamana</emphasis>
Oguma
</taxonomicName>
, habitus and diagnostic details (
<bibRefCitation id="7F4B4B4AE02FD748FB21EF63FA52FC81" author="Ishida, S. &amp; Ishida, K. &amp; Kojima, K. &amp; Sugimura, M." box="[1231,1432,835,861]" pageId="26" pageNumber="235" refId="ref23674" refString="Ishida, S., Ishida, K., Kojima, K. &amp; Sugimura, M. (1988) Illustrated guide for identification of the Japanese Odonata. Tokai University Press, Tokyo, viii + 141 pp., 177 pls. exc. [in Japanese]" type="book" year="1988">
Ishida
<emphasis id="29AEEAA9E02FD748FAF3EF63FA85FC81" box="[1309,1359,835,861]" italics="true" pageId="26" pageNumber="235">et al</emphasis>
. 1988
</bibRefCitation>
,
<bibRefCitation id="7F4B4B4AE02FD748FF79EF47FED4FC5D" author="Ishida, K." box="[151,286,871,897]" pageId="26" pageNumber="235" refId="ref23597" refString="Ishida, K. (1996) Monograph of Odonata larvae in Japan. Hokkaido University Press, Sapporo, 446 pp. [in Japanese]" type="book" year="1996">Ishida 1996</bibRefCitation>
, Sugimura
<emphasis id="29AEEAA9E02FD748FE70EF47FE1CFC5D" box="[414,470,871,897]" italics="true" pageId="26" pageNumber="235">et al.</emphasis>
1999). The larva of
<taxonomicName id="DCDA4D38E02FD748FD53EF47FCB8FC5D" authority="Selys" authorityName="Selys" authorityYear="1879" box="[701,882,871,897]" class="Insecta" family="Euphaeidae" genus="Euphaea" kingdom="Animalia" order="Odonata" pageId="26" pageNumber="235" phylum="Arthropoda" rank="species" species="masoni">
<emphasis id="29AEEAA9E02FD748FD53EF47FCE4FC5C" box="[701,814,871,896]" italics="true" pageId="26" pageNumber="235">E. masoni</emphasis>
Selys
</taxonomicName>
was partially figured by
<bibRefCitation id="7F4B4B4AE02FD748FB62EF47FAAAFC5D" author="Boonsoong, B." box="[1164,1376,871,897]" pageId="26" pageNumber="235" pagination="163 - 169" refId="ref21686" refString="Boonsoong, B. (2016) Phoretic associations between Nanocladius asiaticus (Diptera, Chironomidae) and its hosts Gestroiella (Heteroptera, Naucoridae) and Euphaea masoni (Odonata, Euphaeidae) in streams in Western Thailand. Annales de Limnolgie - International Journal of Limnology, 52, 163 - 169. https: // doi. org / 10.1051 / limn / 2015025" type="journal article" year="2016">Boonsoong (2016)</bibRefCitation>
, who documented its phoretic associations with chironomid larvae in streams in western
<collectingCountry id="63CD762BE02FD748FBD7EFABFB6AFC79" box="[1081,1184,907,933]" name="Thailand" pageId="26" pageNumber="235">Thailand</collectingCountry>
.
</paragraph>
<paragraph id="1B6536BBE02FD748FF29EF8FFB08FA1D" blockId="26.[151,1437,150,1473]" pageId="26" pageNumber="235">
Larvae in the genus are typically squat and heavily built, but may vary in some characters, especially the form of the caudal gills. The head tends to be rather square with strong postocular lobes and the antennae are about the same length as or a little longer than as the distance from the posterior occipital margin to the base of the labrum (
<figureCitation id="83E12A3EE02FD748FF71E83BFF38FBE9" box="[159,242,1050,1077]" captionStart="FIGURES 61–66. 61" captionStartId="12.[151,264,1841,1865]" captionTargetBox="[273,1314,156,1828]" captionTargetId="figure-21@12.[268,1319,151,1834]" captionTargetPageId="12" captionText="FIGURES 61–66. 61. Dysphaea gloriosa: detail of spines on genae and postocular lobe; underside (left), upperside (right) (modified from Nguyen et al. 2024). 62. Heterophaea barbata: dorsal view of head, showing (a) outer mandibular spines (b) genal spines partially visible and (c) right antenna (from Orr &amp; Hämäläinen 2024). 63. Anisopleura subplatystyla, dorsal view of head (original drawing). 64. Bayadera strigata, dorsal view of head (drawn from Yang &amp; Orr 2024). 65. Euphaea decorata, dorsal view of head. (drawn from Xu 2017). 66. Bayadera strigata, left labial palp with detail showing wavy outer margin of outer process of palpal lobe (arrowed), (drawn from Yang &amp; Orr 2024)." figureDoi="http://doi.org/10.5281/zenodo.13618343" httpUri="https://zenodo.org/record/13618343/files/figure.png" pageId="26" pageNumber="235">Fig. 65</figureCitation>
); the inner apical process on the labial palp is well-developed, free and securiform in species examined (
<figureCitation id="83E12A3EE02FD748FA81E83BFF7CFB85" captionStart="FIGURES 67–74. 67" captionStartId="13.[151,264,1814,1838]" captionTargetBox="[256,1340,160,1799]" captionTargetId="figure-15@13.[246,1342,151,1807]" captionTargetPageId="13" captionText="FIGURES 67–74. 67. Anisopleura subplatystyla, caudal gills dorsal view (original drawing). 68. Bayadera strigata, caudal gills dorsal view, (drawn from Yang &amp; Orr 2024). 69. Euphaea decorata, caudal gills dorsal view (drawn from Xu 2017). 70. Euphaea superba, caudal gills dorsal view (redrawn from Wu et al. 2017). 71. Bayadera strigata: male genital apophyses; ventral (left), right lateral (right), (drawn from Yang &amp; Orr 2024). 72. Euphaea decorata, male genital apophyses; ventral view (drawn from Xu 2017). 73. Euphaea superba, terminal processes of palpal lobe ventral view (movable hook truncated), (redrawn from Wu et al. 2017). 74. Anisopleura furcata, left labial palp showing terminal processes of palpal lobe, dorsal view (redrawn from Keetapithchayakul et al. 2024)." figureDoi="http://doi.org/10.5281/zenodo.13618345" httpUri="https://zenodo.org/record/13618345/files/figure.png" pageId="26" pageNumber="235">Fig. 74</figureCitation>
), as in most
<taxonomicName id="DCDA4D38E02FD748FEAEE81FFE67FB85" authorityName="Selys" authorityYear="1853" box="[320,429,1087,1113]" class="Insecta" family="Euphaeidae" genus="Bayadera" kingdom="Animalia" order="Odonata" pageId="26" pageNumber="235" phylum="Arthropoda" rank="genus">
<emphasis id="29AEEAA9E02FD748FEAEE81FFE67FB85" box="[320,429,1087,1113]" italics="true" pageId="26" pageNumber="235">Bayadera</emphasis>
</taxonomicName>
, from which they differ in not possessing a wavy outer margin to the outer process. Strong subocular spines on the gena range from 2–5 (
<bibRefCitation id="7F4B4B4AE02FD748FD71E843FC3DFBA1" author="Keetapithchayakul, T. S. &amp; Sripanya, J. &amp; Phlai-Ngam, S. &amp; Tungpairojwong, N." box="[671,1015,1123,1149]" pageId="26" pageNumber="235" pagination="98 - 110" refId="ref24014" refString="Keetapithchayakul, T. S., Sripanya, J., Phlai-Ngam, S. &amp; Tungpairojwong, N. (2020) Description of the final stadium larva of Bayadera serrata Davies &amp; Yang, 1996 (Odonata: Euphaeidae) from Thailand. Zootaxa, 4894 (1), 98 - 110. https: // doi. org / 10.11646 / zootaxa. 4894.1.5" type="journal article" year="2020">
Keetapithchayakul
<emphasis id="29AEEAA9E02FD748FC94E843FC66FBA1" box="[890,940,1123,1149]" italics="true" pageId="26" pageNumber="235">et al</emphasis>
. 2020
</bibRefCitation>
). the male genital apophyses are very short and triangular or atrophied; the short, saccoid caudal gills often bear a long filament tapering abruptly from the main sac, but sometimes are more elongate with a pale cone-like process (especially in
<taxonomicName id="DCDA4D38E02FD748FB61E88BFAC2FB19" authorityName="Kimmins" authorityYear="1936" box="[1167,1288,1195,1221]" class="Insecta" family="Euphaeidae" genus="Euphaea" kingdom="Animalia" order="Odonata" pageId="26" pageNumber="235" phylum="Arthropoda" rank="species" species="superba">
<emphasis id="29AEEAA9E02FD748FB61E88BFAC2FB19" box="[1167,1288,1195,1221]" italics="true" pageId="26" pageNumber="235">E. superba</emphasis>
</taxonomicName>
, and to some extent in
<taxonomicName id="DCDA4D38E02FD748FF10E8EFFEB8FB35" authorityName="Selys" authorityYear="1859" box="[254,370,1231,1257]" class="Insecta" family="Euphaeidae" genus="Euphaea" kingdom="Animalia" order="Odonata" pageId="26" pageNumber="235" phylum="Arthropoda" rank="species" species="tricolor">
<emphasis id="29AEEAA9E02FD748FF10E8EFFEB8FB35" box="[254,370,1231,1257]" italics="true" pageId="26" pageNumber="235">E. tricolor</emphasis>
</taxonomicName>
and
<taxonomicName id="DCDA4D38E02FD748FE49E8EFFDC3FB34" authorityName="Selys" authorityYear="1853" box="[423,521,1231,1256]" class="Insecta" family="Euphaeidae" genus="Euphaea" kingdom="Animalia" order="Odonata" pageId="26" pageNumber="235" phylum="Arthropoda" rank="species" species="opaca">
<emphasis id="29AEEAA9E02FD748FE49E8EFFDC3FB34" box="[423,521,1231,1256]" italics="true" pageId="26" pageNumber="235">E. opaca</emphasis>
</taxonomicName>
) which is more characteristic of
<taxonomicName id="DCDA4D38E02FD748FC9BE8EFFC28FB35" authorityName="Selys" authorityYear="1853" box="[885,994,1231,1257]" class="Insecta" family="Euphaeidae" genus="Bayadera" kingdom="Animalia" order="Odonata" pageId="26" pageNumber="235" phylum="Arthropoda" rank="genus">
<emphasis id="29AEEAA9E02FD748FC9BE8EFFC28FB35" box="[885,994,1231,1257]" italics="true" pageId="26" pageNumber="235">Bayadera</emphasis>
</taxonomicName>
. Larval habitats of
<taxonomicName id="DCDA4D38E02FD748FB57E8EFFABEFB35" box="[1209,1396,1231,1257]" pageId="26" pageNumber="235">
<emphasis id="29AEEAA9E02FD748FB57E8EFFAD7FB35" box="[1209,1309,1231,1257]" italics="true" pageId="26" pageNumber="235">Euphaea</emphasis>
species
</taxonomicName>
are quite variable, given the geographic and taxonomic range of the genus, but most occur at lower elevations (below
<quantity id="DC229B5EE02FD748FF79E937FF2DFAED" box="[151,231,1303,1329]" metricMagnitude="3" metricUnit="m" metricValue="1.0" pageId="26" pageNumber="235" unit="m" value="1000.0">1000m</quantity>
) in forested or sometimes open streams where the larvae tend to be found under stones in riffles and towards the centre of streams. In
<collectingCountry id="63CD762BE02FD748FE5FE91BFDCAFA89" box="[433,512,1339,1365]" name="Brunei" pageId="26" pageNumber="235">Brunei</collectingCountry>
, for example, four species occur below
<quantity id="DC229B5EE02FD748FC2AE91BFBCEFA89" box="[964,1028,1339,1365]" metricMagnitude="2" metricUnit="m" metricValue="1.0" pageId="26" pageNumber="235" unit="m" value="100.0">100m</quantity>
asl with habitat segregation evident between sister species;
<taxonomicName id="DCDA4D38E02FD748FE72E97FFDDBFAA5" authorityName="Selys" authorityYear="1859" box="[412,529,1375,1401]" class="Insecta" family="Euphaeidae" genus="Euphaea" kingdom="Animalia" order="Odonata" pageId="26" pageNumber="235" phylum="Arthropoda" rank="species" species="tricolor">
<emphasis id="29AEEAA9E02FD748FE72E97FFDDBFAA5" box="[412,529,1375,1401]" italics="true" pageId="26" pageNumber="235">E. tricolor</emphasis>
</taxonomicName>
inhabits the main stream on the Sungei Belalong with larvae almost aways found under stones in a shallow riffle midstream whereas
<taxonomicName id="DCDA4D38E02FD748FD38E9A3FCBBFA41" authorityName="Selys" authorityYear="1873" box="[726,881,1411,1437]" class="Insecta" family="Euphaeidae" genus="Euphaea" kingdom="Animalia" order="Odonata" pageId="26" pageNumber="235" phylum="Arthropoda" rank="species" species="subcostalis">
<emphasis id="29AEEAA9E02FD748FD38E9A3FCBBFA41" box="[726,881,1411,1437]" italics="true" pageId="26" pageNumber="235">E. subcostalis</emphasis>
</taxonomicName>
is only found on small steep side streams leading into the main flow and the microhabitats of the adults also virtually abut each other (
<bibRefCitation id="7F4B4B4AE02FD748FBA7E987FB7FFA1D" author="Orr, A. G." box="[1097,1205,1447,1473]" pageId="26" pageNumber="235" pagination="167 - 220" refId="ref25667" refString="Orr, A. G. (2001) An annotated checklist of the Odonata of Brunei with ecological notes and descriptions of hitherto unknown males and larvae. International Journal of Odonatology, 4, 167 - 220. https: // doi. org / 10.1080 / 13887890.2001.9748168" type="journal article" year="2001">Orr 2001</bibRefCitation>
).
</paragraph>
</subSubSection>
</treatment>
</document>

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data/93/73/87/937387ADE034D750FF79EA82FA56F88C.xml Normal file
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@ -0,0 +1,343 @@
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<mods:title id="657CC22718B7A3DEEF506722E102354E">A review of present knowledge of larvae of the Calopterygoidea (Zygoptera) of the Oriental realm, including keys to families and known genera</mods:title>
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<mods:namePart id="26EFDBF7806175AD9F6A7DCFD7EA44E8">Orr, Albert G. W.</mods:namePart>
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<mods:date id="B836917A0CD5209CCCEC8717F2F1EE9A">2024</mods:date>
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<mods:number id="63D15485A1B7A7DD77628E882AB5BF5C">5497</mods:number>
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<subSubSection id="53C06530E034D753FF79EA82FDC2F960" box="[151,520,1698,1724]" pageId="1" pageNumber="210" type="nomenclature">
<paragraph id="1B6536BBE034D753FF79EA82FDC2F960" blockId="1.[151,520,1698,1724]" box="[151,520,1698,1724]" pageId="1" pageNumber="210">
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Superfamily
<taxonomicName id="DCDA4D38E034D753FED7EA82FDC2F960" authorityName="Sélys" authorityYear="1850" box="[313,520,1698,1724]" class="Insecta" kingdom="Animalia" order="Odonata" pageId="1" pageNumber="210" phylum="Arthropoda" rank="superFamily" superFamily="Calopterygoidea">Calopterygoidea</taxonomicName>
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<subSubSection id="53C06530E034D750FF79EACBFC68FDE1" lastPageId="2" lastPageNumber="211" pageId="1" pageNumber="210" type="description">
<paragraph id="1B6536BBE034D750FF79EACBFD7EFE71" blockId="1.[151,1436,1771,2013]" lastBlockId="2.[151,1436,151,429]" lastPageId="2" lastPageNumber="211" pageId="1" pageNumber="210">
The superfamily
<taxonomicName id="DCDA4D38E034D753FEB7EACBFDDFF8D9" box="[345,533,1771,1797]" pageId="1" pageNumber="210" rank="superFamily" superFamily="Calopterygoidea">Calopterygoidea</taxonomicName>
, as it is presently understood, is a relatively new assemblage of taxa which has been recognised in recent years through the morphological studies of several authors, such as
<bibRefCitation id="7F4B4B4AE034D753FB34EB2FFAA0F8F5" author="Rehn, A. C." box="[1242,1386,1807,1833]" pageId="1" pageNumber="210" pagination="181 - 239" refId="ref26517" refString="Rehn, A. C. (2003) Phylogenetic analysis of higher-level relationships of Odonata. Systematic Entomology, 28, 181 - 239. https: // doi. org / 10.1046 / j. 1365 - 3113.2003.00210. x" type="journal article" year="2003">Rehn (2003)</bibRefCitation>
and molecular studies, notably by
<bibRefCitation id="7F4B4B4AE034D753FE0BEB13FD04F891" author="Dijkstra, K-D. B. &amp; Kalkman, V. J. &amp; Dow, R. A. &amp; Stokvis, F. R. &amp; van Tol, J." box="[485,718,1843,1869]" pageId="1" pageNumber="210" pagination="68 - 96" refId="ref22480" refString="Dijkstra, K-D. B., Kalkman, V. J., Dow, R. A., Stokvis, F. R. &amp; van Tol, J. (2014) Redefining the damselfly families: a comprehensive molecular phylogeny of Zygoptera (Odonata). Systematic entomology, 39, 68 - 96. https: // doi. org / 10.1111 / syen. 12035" type="journal article" year="2014">
Dijkstra
<emphasis id="29AEEAA9E034D753FDA8EB13FDBCF891" box="[582,630,1843,1869]" italics="true" pageId="1" pageNumber="210">et al</emphasis>
. (2014)
</bibRefCitation>
and
<bibRefCitation id="7F4B4B4AE034D753FCEDEB13FC16F891" author="Bybee, S. M. &amp; Kalkman, V. J. &amp; Erickson, R. J. &amp; Breinholt, J. W. &amp; Suvorov, A. &amp; Dijkstra, K-D. B. &amp; Cordero-Riviera, A. &amp; Skevington, J. H. &amp; Abbott, J. C. &amp; Sanchez Herrera, M. &amp; Lemmon, A. R. &amp; Moriaty Lemmon, E. &amp; Ware, J. L." box="[771,988,1843,1869]" pageId="1" pageNumber="210" pagination="1 - 15" refId="ref22012" refString="Bybee, S. M., Kalkman, V. J., Erickson, R. J., Breinholt, J. W., Suvorov, A., Dijkstra, K-D. B., Cordero-Riviera, A., Skevington, J. H., Abbott, J. C., Sanchez Herrera, M., Lemmon, A. R., Moriaty Lemmon, E. &amp; Ware, J. L. (2021) Phylogeny and classification of Odonata using targeted genomics. Molecular Phylogenetics and Evolution, 160, 1 - 15. https: // doi. org / 10.1016 / j. ympev. 2021.107115" type="journal article" year="2021">
Bybee
<emphasis id="29AEEAA9E034D753FCBFEB13FC48F891" box="[849,898,1843,1869]" italics="true" pageId="1" pageNumber="210">et al</emphasis>
. (2021)
</bibRefCitation>
. One unexpected result of the molecular studies has been the inclusion within the group of the family
<taxonomicName id="DCDA4D38E034D753FCBBEB77FC24F8AD" authorityName="Fraser" authorityYear="1960" box="[853,1006,1879,1905]" class="Insecta" family="Argiolestidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="1" pageNumber="210" phylum="Arthropoda" rank="family">Argiolestidae</taxonomicName>
, defined by
<bibRefCitation id="7F4B4B4AE034D753FB93EB77FF2FF849" author="Kalkman, V. J. &amp; Theischinger, G." pageId="1" pageNumber="210" pagination="1 - 52" refId="ref23895" refString="Kalkman, V. J. &amp; Theischinger, G. (2013) Generic revision of Argiolestidae (Odonata), with four new genera. International Journal of Odonatology, 16, 1 - 52. https: // doi. org / 10.1080 / 13887890.2012.749450" type="journal article" year="2013">Kalkman &amp; Theischinger (2013)</bibRefCitation>
. However both molecular studies assess the
<taxonomicName id="DCDA4D38E034D753FD35EB5BFC5EF849" box="[731,916,1915,1941]" pageId="1" pageNumber="210" rank="superFamily" superFamily="Calopterygoidea">Calopterygoidea</taxonomicName>
as paraphyletic (Djikstra
<emphasis id="29AEEAA9E034D753FB56EB5BFB23F849" box="[1208,1257,1915,1941]" italics="true" pageId="1" pageNumber="210">et al</emphasis>
. 2014) or ‘nonmonophyletic’ (
<bibRefCitation id="7F4B4B4AE034D753FEABEBBFFDC1F865" author="Bybee, S. M. &amp; Kalkman, V. J. &amp; Erickson, R. J. &amp; Breinholt, J. W. &amp; Suvorov, A. &amp; Dijkstra, K-D. B. &amp; Cordero-Riviera, A. &amp; Skevington, J. H. &amp; Abbott, J. C. &amp; Sanchez Herrera, M. &amp; Lemmon, A. R. &amp; Moriaty Lemmon, E. &amp; Ware, J. L." box="[325,523,1951,1977]" pageId="1" pageNumber="210" pagination="1 - 15" refId="ref22012" refString="Bybee, S. M., Kalkman, V. J., Erickson, R. J., Breinholt, J. W., Suvorov, A., Dijkstra, K-D. B., Cordero-Riviera, A., Skevington, J. H., Abbott, J. C., Sanchez Herrera, M., Lemmon, A. R., Moriaty Lemmon, E. &amp; Ware, J. L. (2021) Phylogeny and classification of Odonata using targeted genomics. Molecular Phylogenetics and Evolution, 160, 1 - 15. https: // doi. org / 10.1016 / j. ympev. 2021.107115" type="journal article" year="2021">
Bybee
<emphasis id="29AEEAA9E034D753FE7DEBBFFE09F865" box="[403,451,1951,1977]" italics="true" pageId="1" pageNumber="210">et al</emphasis>
. 2021
</bibRefCitation>
), but while the former authors left many minor taxa in the category ‘
<emphasis id="29AEEAA9E034D753FB1DEBBFFA5FF865" box="[1267,1429,1951,1977]" italics="true" pageId="1" pageNumber="210">incertae sedis’</emphasis>
, the latter did assign taxa to named families within
<taxonomicName id="DCDA4D38E034D753FD35EBE3FC5EF801" box="[731,916,1987,2013]" pageId="1" pageNumber="210" rank="superFamily" superFamily="Calopterygoidea">Calopterygoidea</taxonomicName>
(as ‘Calopterygoidea’ see above). Worldwide it includes 26 families and one
<emphasis id="29AEEAA9E037D750FE12ECB7FD52FF6D" box="[508,664,151,177]" italics="true" pageId="2" pageNumber="211">incertae sedis</emphasis>
group (
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Bybee
<emphasis id="29AEEAA9E037D750FCADECB7FCBFFF6D" box="[835,885,151,177]" italics="true" pageId="2" pageNumber="211">et al</emphasis>
. 2021
</bibRefCitation>
). It is the family arrangement of
<bibRefCitation id="7F4B4B4AE037D750FAD6ECB7FECDFF09" author="Bybee, S. M. &amp; Kalkman, V. J. &amp; Erickson, R. J. &amp; Breinholt, J. W. &amp; Suvorov, A. &amp; Dijkstra, K-D. B. &amp; Cordero-Riviera, A. &amp; Skevington, J. H. &amp; Abbott, J. C. &amp; Sanchez Herrera, M. &amp; Lemmon, A. R. &amp; Moriaty Lemmon, E. &amp; Ware, J. L." pageId="2" pageNumber="211" pagination="1 - 15" refId="ref22012" refString="Bybee, S. M., Kalkman, V. J., Erickson, R. J., Breinholt, J. W., Suvorov, A., Dijkstra, K-D. B., Cordero-Riviera, A., Skevington, J. H., Abbott, J. C., Sanchez Herrera, M., Lemmon, A. R., Moriaty Lemmon, E. &amp; Ware, J. L. (2021) Phylogeny and classification of Odonata using targeted genomics. Molecular Phylogenetics and Evolution, 160, 1 - 15. https: // doi. org / 10.1016 / j. ympev. 2021.107115" type="journal article" year="2021">
Bybee
<emphasis id="29AEEAA9E037D750FA66ECB7FF7EFF09" italics="true" pageId="2" pageNumber="211">et al.</emphasis>
(2021)
</bibRefCitation>
that is followed here. This comprises 11 families in the Oriental realm, and for 10 of them, at least
<specimenCount id="0DDCFD32E037D750FA9AEC9BFF05FF25" count="1" pageId="2" pageNumber="211" type="larva">one larva</specimenCount>
is known. The key is expected to generally be reliable at the family level, with the possible exception of some genera of the weakly defined
<taxonomicName id="DCDA4D38E037D750FE0AED23FD52FEC1" box="[484,664,259,285]" family="Rhipidolestidae" pageId="2" pageNumber="237" rank="family">Rhipidolestidae</taxonomicName>
. Larvae of the newly erected family
<taxonomicName id="DCDA4D38E037D750FBDCED23FB29FEC1" box="[1074,1251,259,285]" family="Priscagrionidae" pageId="2" pageNumber="211" rank="family">Priscagrionidae</taxonomicName>
, first recognised in
<bibRefCitation id="7F4B4B4AE037D750FF56ED07FE54FE9D" author="Bybee, S. M. &amp; Kalkman, V. J. &amp; Erickson, R. J. &amp; Breinholt, J. W. &amp; Suvorov, A. &amp; Dijkstra, K-D. B. &amp; Cordero-Riviera, A. &amp; Skevington, J. H. &amp; Abbott, J. C. &amp; Sanchez Herrera, M. &amp; Lemmon, A. R. &amp; Moriaty Lemmon, E. &amp; Ware, J. L." box="[184,414,295,321]" pageId="2" pageNumber="211" pagination="1 - 15" refId="ref22012" refString="Bybee, S. M., Kalkman, V. J., Erickson, R. J., Breinholt, J. W., Suvorov, A., Dijkstra, K-D. B., Cordero-Riviera, A., Skevington, J. H., Abbott, J. C., Sanchez Herrera, M., Lemmon, A. R., Moriaty Lemmon, E. &amp; Ware, J. L. (2021) Phylogeny and classification of Odonata using targeted genomics. Molecular Phylogenetics and Evolution, 160, 1 - 15. https: // doi. org / 10.1016 / j. ympev. 2021.107115" type="journal article" year="2021">
Bybee
<emphasis id="29AEEAA9E037D750FEE4ED07FE8AFE9D" box="[266,320,295,321]" italics="true" pageId="2" pageNumber="211">et al</emphasis>
. (2021)
</bibRefCitation>
, remain unknown. Within the region larvae of the large families
<taxonomicName id="DCDA4D38E037D750FB70ED07FA83FE9D" box="[1182,1353,295,321]" class="Insecta" family="Calopterygidae" kingdom="Animalia" order="Odonata" pageId="2" pageNumber="211" phylum="Arthropoda" rank="family">Calopterygidae</taxonomicName>
(11/12 genera),
<taxonomicName id="DCDA4D38E037D750FF17ED6BFEB6FEB9" baseAuthorityName="Jacobson &amp; Bianchi" baseAuthorityYear="1905" box="[249,380,331,357]" class="Insecta" family="Euphaeidae" kingdom="Animalia" order="Odonata" pageId="2" pageNumber="211" phylum="Arthropoda" rank="family">Euphaeidae</taxonomicName>
(5/8 genera) and
<taxonomicName id="DCDA4D38E037D750FDADED6BFD3CFEB9" authorityName="Cowley" authorityYear="1937" box="[579,758,331,357]" class="Insecta" family="Chlorocyphidae" kingdom="Animalia" order="Odonata" pageId="2" pageNumber="211" phylum="Arthropoda" rank="family">Chlorocyphidae</taxonomicName>
(5/16 genera) are unevenly known. The generic divisions of the
<taxonomicName id="DCDA4D38E037D750FF2FED4FFEBEFE55" authorityName="Cowley" authorityYear="1937" box="[193,372,367,393]" class="Insecta" family="Chlorocyphidae" kingdom="Animalia" order="Odonata" pageId="2" pageNumber="211" phylum="Arthropoda" rank="family">Chlorocyphidae</taxonomicName>
are not fully resolved, and only small morphological differences have been found between the larvae of even quite distant genera in the family.
</paragraph>
<paragraph id="1B6536BBE037D750FF79EDDEFDBEFDC4" blockId="2.[151,930,510,573]" box="[151,628,510,537]" pageId="2" pageNumber="211">
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<emphasis id="29AEEAA9E037D750FF79EDDEFDBEFDC4" bold="true" box="[151,628,510,537]" pageId="2" pageNumber="211">
Family key Oriental
<taxonomicName id="DCDA4D38E037D750FE75EDDEFDA5FDC4" box="[411,623,510,536]" pageId="2" pageNumber="210" rank="superFamily" superFamily="Calopterygoidea">Calopterygoidea</taxonomicName>
:
</emphasis>
</heading>
</paragraph>
<paragraph id="1B6536BBE037D750FF79EE03FC68FDE1" blockId="2.[151,930,510,573]" box="[151,930,547,573]" pageId="2" pageNumber="211">
larva unknown:
<taxonomicName id="DCDA4D38E037D750FEA3EE03FE31FDE1" box="[333,507,547,573]" family="Priscagrionidae" pageId="2" pageNumber="211" rank="family">Priscagrionidae</taxonomicName>
(genera
<taxonomicName id="DCDA4D38E037D750FDB3EE03FD29FDE0" authorityName="Zhou &amp; Wilson" authorityYear="2001" box="[605,739,547,572]" class="Insecta" family="Megapodagrionidae" genus="Priscagrion" kingdom="Animalia" order="Odonata" pageId="2" pageNumber="211" phylum="Arthropoda" rank="genus">
<emphasis id="29AEEAA9E037D750FDB3EE03FD29FDE0" box="[605,739,547,572]" italics="true" pageId="2" pageNumber="211">Priscagrion</emphasis>
</taxonomicName>
and
<taxonomicName id="DCDA4D38E037D750FCF7EE03FC53FDE1" authorityName="Wilson &amp; Zhou" authorityYear="2000" box="[793,921,547,573]" class="Insecta" family="Megapodagrionidae" genus="Sinocnemis" kingdom="Animalia" order="Odonata" pageId="2" pageNumber="211" phylum="Arthropoda" rank="genus">
<emphasis id="29AEEAA9E037D750FCF7EE03FC53FDE1" box="[793,921,547,573]" italics="true" pageId="2" pageNumber="211">Sinocnemis</emphasis>
</taxonomicName>
)
</paragraph>
</subSubSection>
<subSubSection id="53C06530E037D750FF79EE48FA56F88C" pageId="2" pageNumber="211" type="key">
<key id="6F4B618AE037D750FF79EE48FA56F88C" pageId="2" pageNumber="211">
<keyStep id="A02E2E1EE037D750FF79EE48FA56FD29" pageId="2" pageNumber="211">
<paragraph id="1B6536BBE037D750FF79EE48FA56FD67" blockId="2.[151,1437,616,1872]" pageId="2" pageNumber="211">
<keyLead id="A02B958EE037D750FF79EE48FA56FD67" pageId="2" pageNumber="211">
1 Abdomen with two ventral rows of long fleshy, filamentous gills on S2–8; build generally robust; caudal gills saccoid with terminal filament or blunt process (
<figureCitation id="83E12A3EE037D750FDDAEEA5FDA6FD46" box="[564,620,645,667]" captionStart="FIGURES 1–7. 1" captionStartId="3.[151,275,1843,1869]" captionTargetBox="[252,1341,160,1825]" captionTargetId="figure-15@3.[243,1344,149,1836]" captionTargetPageId="3" captionText="FIGURES 1–7. 1. Ventral view of abdomen of Euphaea sp. showing abdominal gills and saccoid gills (original drawing). 2. Head and antennae of Calopterygidae (Neurobasis chinensis) (original drawing). 3. Head and antennae of Chlorocyphidae (Heliocypha biseriata) (original drawing). 4. Antenna of Podolestes orientalis Selys, 1852, scape arrowed (from Choong &amp; Orr 2010). 5. Mask of Calopterygidae (Calopteryx japonica Selys redrawn from Miyakawa 1983). 6. Mask of Chlorocyphidae (Libellago hyalina, from Orr et al. 2024). 7. Habitus of L. hyalina, with detail of right caudal spike (from Orr et al. 2024)." figureDoi="http://doi.org/10.5281/zenodo.13618321" httpUri="https://zenodo.org/record/13618321/files/figure.png" pageId="2" pageNumber="211">Fig. 1</figureCitation>
).........................................................
<taxonomicName id="DCDA4D38E037D750FACDEEA5FA56FD47" baseAuthorityName="Jacobson &amp; Bianchi" baseAuthorityYear="1905" box="[1315,1436,645,667]" class="Insecta" family="Euphaeidae" kingdom="Animalia" order="Odonata" pageId="2" pageNumber="211" phylum="Arthropoda" rank="family">
<emphasis id="29AEEAA9E037D750FACDEEA5FA56FD47" bold="true" box="[1315,1436,645,667]" pageId="2" pageNumber="211">Euphaeidae</emphasis>
</taxonomicName>
[absent from
<collectingRegion id="D91EF859E037D750FADDEE83FA5CFD67" box="[1331,1430,675,699]" country="Indonesia" name="Sulawesi Utara" pageId="2" pageNumber="211">Sulawesi</collectingRegion>
]
</keyLead>
</paragraph>
<paragraph id="1B6536BBE037D750FF79EEE2FA56FD29" blockId="2.[151,1437,616,1872]" pageId="2" pageNumber="211">
<keyLead id="A02B958EE037D750FF79EEE2FA56FD29" pageId="2" pageNumber="211">1’ Abdomen lacking long fleshy gills on venter of abdomen, build very slender to robust, caudal gills of variable form, including saccoid............................................................................................. 2</keyLead>
</paragraph>
</keyStep>
<keyStep id="A02E2E1EE037D750FF79EEDDFA56FC91" pageId="2" pageNumber="211">
<paragraph id="1B6536BBE037D750FF79EEDDFA56FCF3" blockId="2.[151,1437,616,1872]" pageId="2" pageNumber="211">
<keyLead id="A02B958EE037D750FF79EEDDFA56FCF3" pageId="2" pageNumber="211">
2 (1’) Antennal scape at least one third total antennal length; antennal sockets set in raised prominence; (
<figureCitation id="83E12A3EE037D750FB6BEEDDFB13FCCE" box="[1157,1241,765,787]" captionStart="FIGURES 1–7. 1" captionStartId="3.[151,275,1843,1869]" captionTargetBox="[252,1341,160,1825]" captionTargetId="figure-15@3.[243,1344,149,1836]" captionTargetPageId="3" captionText="FIGURES 1–7. 1. Ventral view of abdomen of Euphaea sp. showing abdominal gills and saccoid gills (original drawing). 2. Head and antennae of Calopterygidae (Neurobasis chinensis) (original drawing). 3. Head and antennae of Chlorocyphidae (Heliocypha biseriata) (original drawing). 4. Antenna of Podolestes orientalis Selys, 1852, scape arrowed (from Choong &amp; Orr 2010). 5. Mask of Calopterygidae (Calopteryx japonica Selys redrawn from Miyakawa 1983). 6. Mask of Chlorocyphidae (Libellago hyalina, from Orr et al. 2024). 7. Habitus of L. hyalina, with detail of right caudal spike (from Orr et al. 2024)." figureDoi="http://doi.org/10.5281/zenodo.13618321" httpUri="https://zenodo.org/record/13618321/files/figure.png" pageId="2" pageNumber="211">Figs 2, 3</figureCitation>
); very small to large species with long legs; moderately robust to very slender build.................................................. 3
</keyLead>
</paragraph>
<paragraph id="1B6536BBE037D750FF79EF18FA56FC91" blockId="2.[151,1437,616,1872]" box="[151,1436,824,846]" pageId="2" pageNumber="211">
<keyLead id="A02B958EE037D750FF79EF18FA56FC91" box="[151,1436,824,846]" pageId="2" pageNumber="211">
2’ Antennal scape less than one third total antennal length (
<figureCitation id="83E12A3EE037D750FCECEF18FCF6FC91" box="[770,828,824,846]" captionStart="FIGURES 1–7. 1" captionStartId="3.[151,275,1843,1869]" captionTargetBox="[252,1341,160,1825]" captionTargetId="figure-15@3.[243,1344,149,1836]" captionTargetPageId="3" captionText="FIGURES 1–7. 1. Ventral view of abdomen of Euphaea sp. showing abdominal gills and saccoid gills (original drawing). 2. Head and antennae of Calopterygidae (Neurobasis chinensis) (original drawing). 3. Head and antennae of Chlorocyphidae (Heliocypha biseriata) (original drawing). 4. Antenna of Podolestes orientalis Selys, 1852, scape arrowed (from Choong &amp; Orr 2010). 5. Mask of Calopterygidae (Calopteryx japonica Selys redrawn from Miyakawa 1983). 6. Mask of Chlorocyphidae (Libellago hyalina, from Orr et al. 2024). 7. Habitus of L. hyalina, with detail of right caudal spike (from Orr et al. 2024)." figureDoi="http://doi.org/10.5281/zenodo.13618321" httpUri="https://zenodo.org/record/13618321/files/figure.png" pageId="2" pageNumber="211">Fig. 4</figureCitation>
),................................................ 4
</keyLead>
</paragraph>
</keyStep>
<keyStep id="A02E2E1EE037D750FF79EF75FA57FC3C" pageId="2" pageNumber="211">
<paragraph id="1B6536BBE037D750FF79EF75FA56FC54" blockId="2.[151,1437,616,1872]" pageId="2" pageNumber="211">
<keyLead id="A02B958EE037D750FF79EF75FA56FC54" pageId="2" pageNumber="211">
3(2) Prementum with long lateral lobes and often deep median cleft (
<figureCitation id="83E12A3EE037D750FCD3EF75FCBDFCB7" box="[829,887,853,875]" captionStart="FIGURES 1–7. 1" captionStartId="3.[151,275,1843,1869]" captionTargetBox="[252,1341,160,1825]" captionTargetId="figure-15@3.[243,1344,149,1836]" captionTargetPageId="3" captionText="FIGURES 1–7. 1. Ventral view of abdomen of Euphaea sp. showing abdominal gills and saccoid gills (original drawing). 2. Head and antennae of Calopterygidae (Neurobasis chinensis) (original drawing). 3. Head and antennae of Chlorocyphidae (Heliocypha biseriata) (original drawing). 4. Antenna of Podolestes orientalis Selys, 1852, scape arrowed (from Choong &amp; Orr 2010). 5. Mask of Calopterygidae (Calopteryx japonica Selys redrawn from Miyakawa 1983). 6. Mask of Chlorocyphidae (Libellago hyalina, from Orr et al. 2024). 7. Habitus of L. hyalina, with detail of right caudal spike (from Orr et al. 2024)." figureDoi="http://doi.org/10.5281/zenodo.13618321" httpUri="https://zenodo.org/record/13618321/files/figure.png" pageId="2" pageNumber="211">Fig. 5</figureCitation>
); antennal scape mostly longer than pedicel and flagellum together (
<figureCitation id="83E12A3EE037D750FED0EF52FEB3FC5B" box="[318,377,882,904]" captionStart="FIGURES 1–7. 1" captionStartId="3.[151,275,1843,1869]" captionTargetBox="[252,1341,160,1825]" captionTargetId="figure-15@3.[243,1344,149,1836]" captionTargetPageId="3" captionText="FIGURES 1–7. 1. Ventral view of abdomen of Euphaea sp. showing abdominal gills and saccoid gills (original drawing). 2. Head and antennae of Calopterygidae (Neurobasis chinensis) (original drawing). 3. Head and antennae of Chlorocyphidae (Heliocypha biseriata) (original drawing). 4. Antenna of Podolestes orientalis Selys, 1852, scape arrowed (from Choong &amp; Orr 2010). 5. Mask of Calopterygidae (Calopteryx japonica Selys redrawn from Miyakawa 1983). 6. Mask of Chlorocyphidae (Libellago hyalina, from Orr et al. 2024). 7. Habitus of L. hyalina, with detail of right caudal spike (from Orr et al. 2024)." figureDoi="http://doi.org/10.5281/zenodo.13618321" httpUri="https://zenodo.org/record/13618321/files/figure.png" pageId="2" pageNumber="211">Fig. 2</figureCitation>
); three well developed caudal gills of variable form; medium to large size...............
<taxonomicName id="DCDA4D38E037D750FB11EF52FA56FC54" box="[1279,1436,882,904]" class="Insecta" family="Calopterygidae" kingdom="Animalia" order="Odonata" pageId="2" pageNumber="211" phylum="Arthropoda" rank="family">
<emphasis id="29AEEAA9E037D750FB11EF52FA56FC54" bold="true" box="[1279,1436,882,904]" pageId="2" pageNumber="211">Calopterygidae</emphasis>
</taxonomicName>
</keyLead>
</paragraph>
<paragraph id="1B6536BBE037D750FF79EFB0FA57FC3C" blockId="2.[151,1437,616,1872]" pageId="2" pageNumber="211">
<keyLead id="A02B958EE037D750FF79EFB0FA57FC3C" pageId="2" pageNumber="211">
3’ Prementum without long lateral lobes or deep median cleft (
<figureCitation id="83E12A3EE037D750FCAEEFB0FCB4FC7A" box="[832,894,912,934]" captionStart="FIGURES 1–7. 1" captionStartId="3.[151,275,1843,1869]" captionTargetBox="[252,1341,160,1825]" captionTargetId="figure-15@3.[243,1344,149,1836]" captionTargetPageId="3" captionText="FIGURES 1–7. 1. Ventral view of abdomen of Euphaea sp. showing abdominal gills and saccoid gills (original drawing). 2. Head and antennae of Calopterygidae (Neurobasis chinensis) (original drawing). 3. Head and antennae of Chlorocyphidae (Heliocypha biseriata) (original drawing). 4. Antenna of Podolestes orientalis Selys, 1852, scape arrowed (from Choong &amp; Orr 2010). 5. Mask of Calopterygidae (Calopteryx japonica Selys redrawn from Miyakawa 1983). 6. Mask of Chlorocyphidae (Libellago hyalina, from Orr et al. 2024). 7. Habitus of L. hyalina, with detail of right caudal spike (from Orr et al. 2024)." figureDoi="http://doi.org/10.5281/zenodo.13618321" httpUri="https://zenodo.org/record/13618321/files/figure.png" pageId="2" pageNumber="211">Fig. 6</figureCitation>
); caudal gills reduced to two long, thin lateral spikes, triangular in section, between 0.5–0.9 times length of abdomen, (
<figureCitation id="83E12A3EE037D750FC84EF8DFC6DFC1E" box="[874,935,941,963]" captionStart="FIGURES 1–7. 1" captionStartId="3.[151,275,1843,1869]" captionTargetBox="[252,1341,160,1825]" captionTargetId="figure-15@3.[243,1344,149,1836]" captionTargetPageId="3" captionText="FIGURES 1–7. 1. Ventral view of abdomen of Euphaea sp. showing abdominal gills and saccoid gills (original drawing). 2. Head and antennae of Calopterygidae (Neurobasis chinensis) (original drawing). 3. Head and antennae of Chlorocyphidae (Heliocypha biseriata) (original drawing). 4. Antenna of Podolestes orientalis Selys, 1852, scape arrowed (from Choong &amp; Orr 2010). 5. Mask of Calopterygidae (Calopteryx japonica Selys redrawn from Miyakawa 1983). 6. Mask of Chlorocyphidae (Libellago hyalina, from Orr et al. 2024). 7. Habitus of L. hyalina, with detail of right caudal spike (from Orr et al. 2024)." figureDoi="http://doi.org/10.5281/zenodo.13618321" httpUri="https://zenodo.org/record/13618321/files/figure.png" pageId="2" pageNumber="211">Fig. 7</figureCitation>
), vestigial central gill barely apparent as a conical epiproct; build moderately stout, small to very small............................................
<taxonomicName id="DCDA4D38E037D750FB17EFEAFA57FC3C" authorityName="Cowley" authorityYear="1937" box="[1273,1437,970,992]" class="Insecta" family="Chlorocyphidae" kingdom="Animalia" order="Odonata" pageId="2" pageNumber="211" phylum="Arthropoda" rank="family">
<emphasis id="29AEEAA9E037D750FB17EFEAFA57FC3C" bold="true" box="[1273,1437,970,992]" pageId="2" pageNumber="211">Chlorocyphidae</emphasis>
</taxonomicName>
</keyLead>
</paragraph>
</keyStep>
<keyStep id="A02E2E1EE037D750FF79EFC8FA56FBE4" pageId="2" pageNumber="211">
<paragraph id="1B6536BBE037D750FF79EFC8FA56FBC7" blockId="2.[151,1437,616,1872]" pageId="2" pageNumber="211">
<keyLead id="A02B958EE037D750FF79EFC8FA56FBC7" pageId="2" pageNumber="211">
4 (2’). Caudal gills bulbous and sclerotised, either ovate or conical with blunt trifid tips; venter of S10 with paired retractable gill tufts (
<figureCitation id="83E12A3EE037D750FF07E825FEF5FBC6" box="[233,319,1029,1051]" captionStart="FIGURES 8–11. 8" captionStartId="4.[151,264,1914,1938]" captionTargetBox="[200,1382,177,1900]" captionTargetId="figure-21@4.[193,1394,169,1908]" captionTargetPageId="4" captionText="FIGURES 8–11. 8. Pseudolestes mirabilis, ventral view of terminal abdominal segments and caudal gills of showing gill tufts on S10 (after Yu &amp; Bu 2011). 9. Devadatta argyoides, detail of extruded right gill tuft, posterior view (redrawn from Watson 1966). 10. Pseudolestes mirabilis, habitus with detail of mask (after Yu &amp; Bu 2011). 11. Devadatta argyoides, habitus with details of right antenna and mask (original drawing)." figureDoi="http://doi.org/10.5281/zenodo.13618323" httpUri="https://zenodo.org/record/13618323/files/figure.png" pageId="2" pageNumber="211">Figs 8, 9</figureCitation>
), sometimes just evident as tiny tumescent orifices when retracted; stoutly built............................ 5
</keyLead>
</paragraph>
<paragraph id="1B6536BBE037D750FF79E802FA56FBE4" blockId="2.[151,1437,616,1872]" box="[151,1436,1058,1080]" pageId="2" pageNumber="211">
<keyLead id="A02B958EE037D750FF79E802FA56FBE4" box="[151,1436,1058,1080]" pageId="2" pageNumber="211">4’ Caudal gills not as above, either soft and saccoid, or else laterally or ventrally flattened. Ventral gill tufts on S10 absent.... 6</keyLead>
</paragraph>
</keyStep>
<keyStep id="A02E2E1EE037D750FF79E81FFA56FB34" pageId="2" pageNumber="211">
<paragraph id="1B6536BBE037D750FF79E81FFA56FB4C" blockId="2.[151,1437,616,1872]" pageId="2" pageNumber="211">
<keyLead id="A02B958EE037D750FF79E81FFA56FB4C" pageId="2" pageNumber="211">
5(4) Caudal gills subspherical hardened balloons with slight acuminate tip (
<figureCitation id="83E12A3EE037D750FC67E860FC1AFB89" box="[905,976,1088,1110]" captionStart="FIGURES 8–11. 8" captionStartId="4.[151,264,1914,1938]" captionTargetBox="[200,1382,177,1900]" captionTargetId="figure-21@4.[193,1394,169,1908]" captionTargetPageId="4" captionText="FIGURES 8–11. 8. Pseudolestes mirabilis, ventral view of terminal abdominal segments and caudal gills of showing gill tufts on S10 (after Yu &amp; Bu 2011). 9. Devadatta argyoides, detail of extruded right gill tuft, posterior view (redrawn from Watson 1966). 10. Pseudolestes mirabilis, habitus with detail of mask (after Yu &amp; Bu 2011). 11. Devadatta argyoides, habitus with details of right antenna and mask (original drawing)." figureDoi="http://doi.org/10.5281/zenodo.13618323" httpUri="https://zenodo.org/record/13618323/files/figure.png" pageId="2" pageNumber="211">Fig. 10</figureCitation>
), antennae normally proportioned with scape ca half length of pedicel; prementum long, narrow and gently tapered...................................
<taxonomicName id="DCDA4D38E037D750FAE8E87DFA57FBAF" authorityName="Fraser" authorityYear="1957" box="[1286,1437,1117,1139]" class="Insecta" family="Pseudolestidae" kingdom="Animalia" order="Odonata" pageId="2" pageNumber="211" phylum="Arthropoda" rank="family">
<emphasis id="29AEEAA9E037D750FAE8E87DFA57FBAF" bold="true" box="[1286,1437,1117,1139]" pageId="2" pageNumber="211">Pseudolestidae</emphasis>
</taxonomicName>
[endemic to
<collectingRegion id="D91EF859E037D750FAA1E85AFA5DFB4C" box="[1359,1431,1146,1168]" country="China" name="Hainan" pageId="2" pageNumber="211">Hainan</collectingRegion>
]
</keyLead>
</paragraph>
<paragraph id="1B6536BBE037D750FF79E8B7FA56FB34" blockId="2.[151,1437,616,1872]" pageId="2" pageNumber="211">
<keyLead id="A02B958EE037D750FF79E8B7FA56FB34" pageId="2" pageNumber="211">
5’ Caudal gills hardened, conical, median tricorn (
<figureCitation id="83E12A3EE037D750FD5CE8B8FD30FB71" box="[690,762,1176,1198]" captionStart="FIGURES 8–11. 8" captionStartId="4.[151,264,1914,1938]" captionTargetBox="[200,1382,177,1900]" captionTargetId="figure-21@4.[193,1394,169,1908]" captionTargetPageId="4" captionText="FIGURES 8–11. 8. Pseudolestes mirabilis, ventral view of terminal abdominal segments and caudal gills of showing gill tufts on S10 (after Yu &amp; Bu 2011). 9. Devadatta argyoides, detail of extruded right gill tuft, posterior view (redrawn from Watson 1966). 10. Pseudolestes mirabilis, habitus with detail of mask (after Yu &amp; Bu 2011). 11. Devadatta argyoides, habitus with details of right antenna and mask (original drawing)." figureDoi="http://doi.org/10.5281/zenodo.13618323" httpUri="https://zenodo.org/record/13618323/files/figure.png" pageId="2" pageNumber="211">Fig. 11</figureCitation>
); first three segments of antennae abnormally thick with scape about twice length of pedicel; prementum broad, squared................................................
<taxonomicName id="DCDA4D38E037D750FAFAE895FA56FB17" authorityName="Dijkstra, Kalkman, Dow, Stokvis &amp; van Tol" authorityYear="2014" box="[1300,1436,1205,1227]" class="Insecta" family="Devadattidae" higherTaxonomySource="CoL,GBIF" kingdom="Animalia" order="Odonata" pageId="2" pageNumber="211" phylum="Arthropoda" rank="family">
<emphasis id="29AEEAA9E037D750FAFAE895FA56FB17" bold="true" box="[1300,1436,1205,1227]" pageId="2" pageNumber="211">Devadattidae</emphasis>
</taxonomicName>
[mainland south-east Asia from southern
<collectingRegion id="D91EF859E037D750FB59E8F3FACAFB34" box="[1207,1280,1235,1256]" country="China" name="Yunnan" pageId="2" pageNumber="211">Yunnan</collectingRegion>
and Sundaland]
</keyLead>
</paragraph>
</keyStep>
<keyStep id="A02E2E1EE037D750FF79E8D0FA56FAFE" pageId="2" pageNumber="211">
<paragraph id="1B6536BBE037D750FF79E8D0FA56FAD9" blockId="2.[151,1437,616,1872]" box="[151,1436,1264,1286]" pageId="2" pageNumber="211">
<keyLead id="A02B958EE037D750FF79E8D0FA56FAD9" box="[151,1436,1264,1286]" pageId="2" pageNumber="211">6 (4’) Caudal gills flattened, foliate and arranged horizontally in a broad fan........................................... 7</keyLead>
</paragraph>
<paragraph id="1B6536BBE037D750FF79E92DFA56FAFE" blockId="2.[151,1437,616,1872]" box="[151,1436,1293,1315]" pageId="2" pageNumber="211">
<keyLead id="A02B958EE037D750FF79E92DFA56FAFE" box="[151,1436,1293,1315]" pageId="2" pageNumber="211">6’ Caudal gills either saccoid or broadly flattened and arranged vertically........................................... 8</keyLead>
</paragraph>
</keyStep>
<keyStep id="A02E2E1EE037D750FF79E90AFA56FA2C" pageId="2" pageNumber="211">
<paragraph id="1B6536BBE037D750FF79E90AFA56FA44" blockId="2.[151,1437,616,1872]" pageId="2" pageNumber="211">
<keyLead id="A02B958EE037D750FF79E90AFA56FA44" pageId="2" pageNumber="211">
7(6) Stoutly built with broad, acutely tipped, foliate caudal gills ca 0.6 length of abdomen (
<figureCitation id="83E12A3EE037D750FBF9E90AFB94FAE3" box="[1047,1118,1322,1344]" captionStart="FIGURES 12–17. 12" captionStartId="5.[151,264,1903,1927]" captionTargetBox="[190,1400,160,1874]" captionTargetId="figure-15@5.[178,1409,151,1889]" captionTargetPageId="5" captionText="FIGURES 12–17. 12. Mesopodagrion tibetanum, habitus with detail of right profile of head (based on Yu 2016). 13. M. tibetanum, detail of mask (redrawn from Yu 2016). 14. Podolestes orientalis, habitus (from Choong &amp; Orr 2010). 15. Podolestes orientalis, detail of mask (from Choong &amp; Orr 2010). 16. Rhipidolestes aculeatus, habitus and detail of lateral and median gill (modified from Asahina 1994). 17. Rhipidolestes aculeatus, detail of mask (modified from Ishida 1996)." figureDoi="http://doi.org/10.5281/zenodo.13618327" httpUri="https://zenodo.org/record/13618327/files/figure.png" pageId="2" pageNumber="211">Fig. 12</figureCitation>
); head robust with distinct lateral cephalic lobes; antennae short, no longer than long axis of head from occipital rim to base of labrum; prementum broad and short with short labial palps (
<figureCitation id="83E12A3EE037D750FE14E945FDF5FAA6" box="[506,575,1381,1403]" captionStart="FIGURES 12–17. 12" captionStartId="5.[151,264,1903,1927]" captionTargetBox="[190,1400,160,1874]" captionTargetId="figure-15@5.[178,1409,151,1889]" captionTargetPageId="5" captionText="FIGURES 12–17. 12. Mesopodagrion tibetanum, habitus with detail of right profile of head (based on Yu 2016). 13. M. tibetanum, detail of mask (redrawn from Yu 2016). 14. Podolestes orientalis, habitus (from Choong &amp; Orr 2010). 15. Podolestes orientalis, detail of mask (from Choong &amp; Orr 2010). 16. Rhipidolestes aculeatus, habitus and detail of lateral and median gill (modified from Asahina 1994). 17. Rhipidolestes aculeatus, detail of mask (modified from Ishida 1996)." figureDoi="http://doi.org/10.5281/zenodo.13618327" httpUri="https://zenodo.org/record/13618327/files/figure.png" pageId="2" pageNumber="211">Fig. 13</figureCitation>
)......................................................
<taxonomicName id="DCDA4D38E037D750FB3CE945FA57FAA7" authorityName="Kalkman &amp; Abbott" authorityYear="2021" box="[1234,1437,1381,1403]" class="Insecta" family="Mesopodagrionidae" higherTaxonomySource="CoL" kingdom="Animalia" order="Odonata" pageId="2" pageNumber="211" phylum="Arthropoda" rank="family">
<emphasis id="29AEEAA9E037D750FB3CE945FA57FAA7" bold="true" box="[1234,1437,1381,1403]" pageId="2" pageNumber="211">Mesopodagrionidae</emphasis>
</taxonomicName>
[southern
<collectingCountry id="63CD762BE037D750FBF2E9A2FB92FA44" box="[1052,1112,1410,1432]" name="China" pageId="2" pageNumber="211">China</collectingCountry>
, northern Indochina &amp;
<collectingCountry id="63CD762BE037D750FAD9E9A3FA5DFA44" box="[1335,1431,1411,1432]" name="Myanmar" pageId="2" pageNumber="211">Myanmar</collectingCountry>
]
</keyLead>
</paragraph>
<paragraph id="1B6536BBE037D750FF79E980FA56FA2C" blockId="2.[151,1437,616,1872]" pageId="2" pageNumber="211">
<keyLead id="A02B958EE037D750FF79E980FA56FA2C" pageId="2" pageNumber="211">
7’ Variable in build but head never with lateral cephalic lobes; antenna moderately long or short, caudal gills always broad and arranged in a horizontal fan, but vary considerably in relative size;
<taxonomicName id="DCDA4D38E037D750FC8EE99DFC02FA0E" authorityName="Selys" authorityYear="1862" box="[864,968,1469,1490]" class="Insecta" family="Megapodagrionidae" genus="Podolestes" kingdom="Animalia" order="Odonata" pageId="2" pageNumber="211" phylum="Arthropoda" rank="genus">
<emphasis id="29AEEAA9E037D750FC8EE99DFC02FA0E" box="[864,968,1469,1490]" italics="true" pageId="2" pageNumber="211">Podolestes</emphasis>
</taxonomicName>
(
<figureCitation id="83E12A3EE037D750FC3AE99DFBF5FA0F" box="[980,1087,1469,1491]" captionStart="FIGURES 12–17. 12" captionStartId="5.[151,264,1903,1927]" captionTargetBox="[190,1400,160,1874]" captionTargetId="figure-15@5.[178,1409,151,1889]" captionTargetPageId="5" captionText="FIGURES 12–17. 12. Mesopodagrion tibetanum, habitus with detail of right profile of head (based on Yu 2016). 13. M. tibetanum, detail of mask (redrawn from Yu 2016). 14. Podolestes orientalis, habitus (from Choong &amp; Orr 2010). 15. Podolestes orientalis, detail of mask (from Choong &amp; Orr 2010). 16. Rhipidolestes aculeatus, habitus and detail of lateral and median gill (modified from Asahina 1994). 17. Rhipidolestes aculeatus, detail of mask (modified from Ishida 1996)." figureDoi="http://doi.org/10.5281/zenodo.13618327" httpUri="https://zenodo.org/record/13618327/files/figure.png" pageId="2" pageNumber="211">Figs 14, 15</figureCitation>
) may be broadly sympatric with last family but has antennae longer than the long axis of head; caudal lamellae relatively much larger...........
<taxonomicName id="DCDA4D38E037D750FAFAE9FAFA56FA2C" authorityName="Fraser" authorityYear="1960" box="[1300,1436,1498,1520]" class="Insecta" family="Argiolestidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="2" pageNumber="211" phylum="Arthropoda" rank="family">
<emphasis id="29AEEAA9E037D750FAFAE9FAFA56FA2C" bold="true" box="[1300,1436,1498,1520]" pageId="2" pageNumber="211">Argiolestidae</emphasis>
</taxonomicName>
</keyLead>
</paragraph>
</keyStep>
<keyStep id="A02E2E1EE037D750FF79E9D8FA56F9B9" pageId="2" pageNumber="211">
<paragraph id="1B6536BBE037D750FF79E9D8FA57F994" blockId="2.[151,1437,616,1872]" pageId="2" pageNumber="211">
<keyLead id="A02B958EE037D750FF79E9D8FA57F994" pageId="2" pageNumber="211">
8(6’) Caudal gills flattened, fleshy short and broad, arranged vertically, median gill thin walled with complex tracheation, (
<figureCitation id="83E12A3EE037D750FA9CE9D8FED5F9F6" captionStart="FIGURES 18–26. 18" captionStartId="6.[151,264,1811,1835]" captionTargetBox="[265,1336,167,1790]" captionTargetId="figure-21@6.[248,1339,153,1804]" captionTargetPageId="6" captionText="FIGURES 18–26. 18. Rhinagrion borneense, habitus (original drawing). 19. Philosina alba, habitus (modified from Zhang et al. 2011). 20. Rhinagrion mima, caudal gills: above heavily tracheated median gill; below right lateral gill with basal spines along mid-rib (modified from Lieftinck 1956). 21. Philosina alba caudal gills: above heavily tracheated median gill; below right lateral gill with basal spines along mid-rib (modified from Zhang et al. 2011) 22. Rhinagrion borneense, detail of mask (original drawing). 23. Philosina buchi, detail of mask (redrawn from Zhang et al. 2011). 24. Philoganga vetusta, habitus (adapted from Xu 2016). 25. Philoganga vetusta, details of caudal gills and antenna (redrawn from Xu 2016). 26. Philoganga montana, detail of mask (original drawing)." figureDoi="http://doi.org/10.5281/zenodo.13618331" httpUri="https://zenodo.org/record/13618331/files/figure.png" pageId="2" pageNumber="211">Figs 18–21</figureCitation>
)....................................................................................
<taxonomicName id="DCDA4D38E037D750FAF1EA35FA56F9F7" authorityName="Kennedy" authorityYear="1925" box="[1311,1436,1557,1579]" class="Insecta" family="Philosinidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="2" pageNumber="211" phylum="Arthropoda" rank="family">
<emphasis id="29AEEAA9E037D750FAF1EA35FA56F9F7" bold="true" box="[1311,1436,1557,1579]" pageId="2" pageNumber="211">Philosinidae</emphasis>
</taxonomicName>
[West of Wallace’s line]
</keyLead>
</paragraph>
<paragraph id="1B6536BBE037D750FF79EA70FA56F9B9" blockId="2.[151,1437,616,1872]" box="[151,1436,1616,1638]" pageId="2" pageNumber="211">
<keyLead id="A02B958EE037D750FF79EA70FA56F9B9" box="[151,1436,1616,1638]" pageId="2" pageNumber="211">8’ Caudal gills saccoid, terminating in filament with median gill placed over two lower, lateral gills...................... 9</keyLead>
</paragraph>
</keyStep>
<keyStep id="A02E2E1EE037D750FF79EA4DFA56F88C" pageId="2" pageNumber="211">
<paragraph id="1B6536BBE037D750FF79EA4DFA56F907" blockId="2.[151,1437,616,1872]" pageId="2" pageNumber="211">
<keyLead id="A02B958EE037D750FF79EA4DFA56F907" pageId="2" pageNumber="211">
9 (8’) Head nearly straight sided behind eyes producing an overall squared profile (
<figureCitation id="83E12A3EE037D750FC53EA4DFBCEF95E" box="[957,1028,1645,1667]" captionStart="FIGURES 18–26. 18" captionStartId="6.[151,264,1811,1835]" captionTargetBox="[265,1336,167,1790]" captionTargetId="figure-21@6.[248,1339,153,1804]" captionTargetPageId="6" captionText="FIGURES 18–26. 18. Rhinagrion borneense, habitus (original drawing). 19. Philosina alba, habitus (modified from Zhang et al. 2011). 20. Rhinagrion mima, caudal gills: above heavily tracheated median gill; below right lateral gill with basal spines along mid-rib (modified from Lieftinck 1956). 21. Philosina alba caudal gills: above heavily tracheated median gill; below right lateral gill with basal spines along mid-rib (modified from Zhang et al. 2011) 22. Rhinagrion borneense, detail of mask (original drawing). 23. Philosina buchi, detail of mask (redrawn from Zhang et al. 2011). 24. Philoganga vetusta, habitus (adapted from Xu 2016). 25. Philoganga vetusta, details of caudal gills and antenna (redrawn from Xu 2016). 26. Philoganga montana, detail of mask (original drawing)." figureDoi="http://doi.org/10.5281/zenodo.13618331" httpUri="https://zenodo.org/record/13618331/files/figure.png" pageId="2" pageNumber="211">Fig. 24</figureCitation>
); antennae mostly 8 segmented and clearly longer than head from rear occipital margin to base of frons; caudal gills elongate and terminating uniformly in a point (
<figureCitation id="83E12A3EE037D750FA9BEAAAFF37F961" captionStart="FIGURES 18–26. 18" captionStartId="6.[151,264,1811,1835]" captionTargetBox="[265,1336,167,1790]" captionTargetId="figure-21@6.[248,1339,153,1804]" captionTargetPageId="6" captionText="FIGURES 18–26. 18. Rhinagrion borneense, habitus (original drawing). 19. Philosina alba, habitus (modified from Zhang et al. 2011). 20. Rhinagrion mima, caudal gills: above heavily tracheated median gill; below right lateral gill with basal spines along mid-rib (modified from Lieftinck 1956). 21. Philosina alba caudal gills: above heavily tracheated median gill; below right lateral gill with basal spines along mid-rib (modified from Zhang et al. 2011) 22. Rhinagrion borneense, detail of mask (original drawing). 23. Philosina buchi, detail of mask (redrawn from Zhang et al. 2011). 24. Philoganga vetusta, habitus (adapted from Xu 2016). 25. Philoganga vetusta, details of caudal gills and antenna (redrawn from Xu 2016). 26. Philoganga montana, detail of mask (original drawing)." figureDoi="http://doi.org/10.5281/zenodo.13618331" httpUri="https://zenodo.org/record/13618331/files/figure.png" pageId="2" pageNumber="211">Fig. 25</figureCitation>
); mature larvae and exuviae huge,
<quantity id="DC229B5EE037D750FDA2EA88FD79F961" box="[588,691,1704,1725]" metricMagnitude="-2" metricUnit="m" metricValue="3.5" metricValueMax="4.0" metricValueMin="3.0" pageId="2" pageNumber="211" unit="mm" value="35.0" valueMax="40.0" valueMin="30.0">30–40 mm</quantity>
in total length including gills; general build robust but elongate..................................................................................................
<taxonomicName id="DCDA4D38E037D750FAE5EAE5FA56F907" authorityName="Kennedy" authorityYear="1920" box="[1291,1436,1733,1755]" class="Insecta" family="Philogangidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="2" pageNumber="211" phylum="Arthropoda" rank="family">
<emphasis id="29AEEAA9E037D750FAE5EAE5FA56F907" bold="true" box="[1291,1436,1733,1755]" pageId="2" pageNumber="211">Philogangidae</emphasis>
</taxonomicName>
</keyLead>
</paragraph>
<paragraph id="1B6536BBE037D750FF79EAC2FA56F88C" blockId="2.[151,1437,616,1872]" pageId="2" pageNumber="211">
<keyLead id="A02B958EE037D750FF79EAC2FA56F88C" pageId="2" pageNumber="211">
9’ Head with postocular lobes rounded (
<figureCitation id="83E12A3EE037D750FDA6EAC2FD45F924" box="[584,655,1762,1784]" captionStart="FIGURES 12–17. 12" captionStartId="5.[151,264,1903,1927]" captionTargetBox="[190,1400,160,1874]" captionTargetId="figure-15@5.[178,1409,151,1889]" captionTargetPageId="5" captionText="FIGURES 12–17. 12. Mesopodagrion tibetanum, habitus with detail of right profile of head (based on Yu 2016). 13. M. tibetanum, detail of mask (redrawn from Yu 2016). 14. Podolestes orientalis, habitus (from Choong &amp; Orr 2010). 15. Podolestes orientalis, detail of mask (from Choong &amp; Orr 2010). 16. Rhipidolestes aculeatus, habitus and detail of lateral and median gill (modified from Asahina 1994). 17. Rhipidolestes aculeatus, detail of mask (modified from Ishida 1996)." figureDoi="http://doi.org/10.5281/zenodo.13618327" httpUri="https://zenodo.org/record/13618327/files/figure.png" pageId="2" pageNumber="211">Fig. 16</figureCitation>
); antennae short and robust, about same length as head from rear margin to frons; caudal gills ovate saccoid with distinct dorsal ridge and terminating in short, well defined filament; prementum broad and short with short labial palps (
<figureCitation id="83E12A3EE037D750FE27EB3DFDDBF8EE" box="[457,529,1821,1843]" captionStart="FIGURES 12–17. 12" captionStartId="5.[151,264,1903,1927]" captionTargetBox="[190,1400,160,1874]" captionTargetId="figure-15@5.[178,1409,151,1889]" captionTargetPageId="5" captionText="FIGURES 12–17. 12. Mesopodagrion tibetanum, habitus with detail of right profile of head (based on Yu 2016). 13. M. tibetanum, detail of mask (redrawn from Yu 2016). 14. Podolestes orientalis, habitus (from Choong &amp; Orr 2010). 15. Podolestes orientalis, detail of mask (from Choong &amp; Orr 2010). 16. Rhipidolestes aculeatus, habitus and detail of lateral and median gill (modified from Asahina 1994). 17. Rhipidolestes aculeatus, detail of mask (modified from Ishida 1996)." figureDoi="http://doi.org/10.5281/zenodo.13618327" httpUri="https://zenodo.org/record/13618327/files/figure.png" pageId="2" pageNumber="211">Fig. 17</figureCitation>
); general build stout, tapering caudad and less than
<quantity id="DC229B5EE037D750FBE0EB3DFB98F8EE" box="[1038,1106,1821,1843]" metricMagnitude="-2" metricUnit="m" metricValue="1.5" pageId="2" pageNumber="211" unit="mm" value="15.0">15 mm</quantity>
in length (
<quantity id="DC229B5EE037D750FB53EB3DFAEDF8EE" box="[1213,1319,1821,1842]" metricMagnitude="-2" metricUnit="m" metricValue="1.1" metricValueMax="1.2" metricValueMin="1.0" pageId="2" pageNumber="211" unit="mm" value="11.0" valueMax="12.0" valueMin="10.0">10–12 mm</quantity>
in recorded specimens)..............................................................................
<taxonomicName id="DCDA4D38E037D750FB12EB1AFA56F88C" box="[1276,1436,1850,1872]" family="Rhipidolestidae" pageId="2" pageNumber="237" rank="family">
<emphasis id="29AEEAA9E037D750FB12EB1AFA56F88C" bold="true" box="[1276,1436,1850,1872]" pageId="2" pageNumber="211">Rhipidolestidae</emphasis>
</taxonomicName>
</keyLead>
</paragraph>
</keyStep>
</key>
</subSubSection>
</treatment>
</document>

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<subSubSection id="53C06530E03AD75DFF79E88AFF1EFB19" box="[151,212,1194,1221]" pageId="15" pageNumber="224" type="nomenclature">
<paragraph id="1B6536BBE03AD75DFF79E88AFF1EFB19" blockId="15.[151,1437,1194,1941]" box="[151,212,1194,1221]" pageId="15" pageNumber="224">
<heading id="402D81D7E03AD75DFF79E88AFF1EFB19" bold="true" box="[151,212,1194,1221]" fontSize="11" level="1" pageId="15" pageNumber="224" reason="1">
<taxonomicName id="DCDA4D38E03AD75DFF79E88AFF1EFB19" ID-CoL="49PY" authorityName="Selys" authorityYear="1853" box="[151,212,1194,1221]" class="Insecta" family="Calopterygidae" genus="Echo" kingdom="Animalia" order="Odonata" pageId="15" pageNumber="224" phylum="Arthropoda" rank="genus">
<emphasis id="29AEEAA9E03AD75DFF79E88AFF1EFB19" bold="true" box="[151,212,1194,1221]" italics="true" pageId="15" pageNumber="224">Echo</emphasis>
</taxonomicName>
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</subSubSection>
<subSubSection id="53C06530E03AD742FF79E8EFFEFFFE9D" lastPageId="16" lastPageNumber="225" pageId="15" pageNumber="224" type="discussion">
<paragraph id="1B6536BBE03AD75DFF79E8EFFC95F849" blockId="15.[151,1437,1194,1941]" pageId="15" pageNumber="224">
The genus includes five species ranging from southern
<collectingCountry id="63CD762BE03AD75DFD15E8EFFCF4FB35" box="[763,830,1231,1257]" name="China" pageId="15" pageNumber="224">China</collectingCountry>
and the eastern Himalayas to the Malay Peninsula and Sumatra. Only the larva of
<taxonomicName id="DCDA4D38E03AD75DFE28E8D3FD7FFAD1" authority="Selys" authorityName="Selys" authorityYear="1879" box="[454,693,1267,1293]" class="Insecta" family="Calopterygidae" genus="Echo" kingdom="Animalia" order="Odonata" pageId="15" pageNumber="224" phylum="Arthropoda" rank="species" species="uniformis">
<emphasis id="29AEEAA9E03AD75DFE28E8D3FDBBFAD1" box="[454,625,1267,1293]" italics="true" pageId="15" pageNumber="224">Echo uniformis</emphasis>
Selys
</taxonomicName>
from Sumatra has been briefly described and illustrated (
<bibRefCitation id="7F4B4B4AE03AD75DFADDE8D3FF18FAED" author="Lieftinck, M. A." pageId="15" pageNumber="224" pagination="325 - 364" refId="ref24671" refString="Lieftinck, M. A. (1965) The species-group of Vestalis amoena Selys, 1853, in Sundaland (Odonata, Calopterygidae). Tijdschrift voor Entomologie, 108, 325 - 364." type="journal article" year="1965">Lieftinck 1965</bibRefCitation>
,
<bibRefCitation id="7F4B4B4AE03AD75DFF30E937FEBDFAED" author="van Tol, J." box="[222,375,1303,1329]" pageId="15" pageNumber="224" pagination="1 - 263" refId="ref27215" refString="van Tol, J. (1992) An annotated index to names of Odonata used in publications by M. A. Lieftinck. Zoologische Verhandelingen, 279, 1 - 263." type="journal article" year="1992">van Tol 1992</bibRefCitation>
), with drawings showing the habitus and the prementum with labial palps. Nevertheless, both illustrations and the brief text convey critical information that separates the genus from any other, including its sister genus
<taxonomicName id="DCDA4D38E03AD75DFF31E97FFEBBFAA5" authorityName="McLachlan" authorityYear="1870" box="[223,369,1375,1401]" class="Insecta" family="Calopterygidae" genus="Psolodesmus" kingdom="Animalia" order="Odonata" pageId="15" pageNumber="224" phylum="Arthropoda" rank="genus">
<emphasis id="29AEEAA9E03AD75DFF31E97FFEBBFAA5" box="[223,369,1375,1401]" italics="true" pageId="15" pageNumber="224">Psolodesmus</emphasis>
</taxonomicName>
(according to
<bibRefCitation id="7F4B4B4AE03AD75DFDFDE97FFD3AFAA5" author="Dumont, H. J. &amp; Vierstraete, A. &amp; Vanfleteren, J. R." box="[531,752,1375,1401]" pageId="15" pageNumber="224" pagination="365 - 372" refId="ref22641" refString="Dumont, H. J., Vierstraete, A. &amp; Vanfleteren, J. R. (2007) A revised molecular phylogeny of the Calopteryginae (Zygoptera: Calopterygidae). Odonatologica, 36, 365 - 372." type="journal article" year="2007">
Dumont
<emphasis id="29AEEAA9E03AD75DFD98E97FFD6DFAA5" box="[630,679,1375,1401]" italics="true" pageId="15" pageNumber="224">et al</emphasis>
. 2007
</bibRefCitation>
), which is confined to
<collectingCountry id="63CD762BE03AD75DFC03E97FFBF5FAA5" box="[1005,1087,1375,1401]" name="Taiwan" pageId="15" pageNumber="224">Taiwan</collectingCountry>
and southern Japanese Islands. The prementum (
<figureCitation id="83E12A3EE03AD75DFEB0E9A3FE79FA41" box="[350,435,1411,1437]" captionStart="FIGURES 27–34. 27" captionStartId="7.[151,264,1845,1869]" captionTargetBox="[252,1326,159,1834]" captionTargetId="figure-15@7.[251,1336,153,1839]" captionTargetPageId="7" captionText="FIGURES 27–34. 27. Prementum of Calopterygidae showing measurements used in family key (based on Calopteryx japonica, redrawn from Miyakawa 1983). 28. Neurobasis chinensis, prementum and left labial palp (redrawn from Lieftinck 1965). 29. Matronoides cyaneipennis, prementum and labial palps (ventral view). 30. Matrona basilaris, prementum (dorsal view, drawn from Wang et al. 2017). 31. Atrocalopteryx atrata, prementum (dorsal view, redrawn from Miyakawa 1983). 32. Mnais tenuis, prementum (dorsal view, redrawn from Matsuki &amp; Lien 1977). 33. Echo uniformis, prementum and left labial palp dorsal view (redrawn from Lieftinck 1965). 34. Psolodesmus mandarinus, prementum (dorsal view, redrawn from Matsuki &amp; Lien 1978)." figureDoi="http://doi.org/10.5281/zenodo.13618333" httpUri="https://zenodo.org/record/13618333/files/figure.png" pageId="15" pageNumber="224">Fig. 33</figureCitation>
) is very similar to that of
<taxonomicName id="DCDA4D38E03AD75DFD04E9A3FCB6FA41" authorityName="McLachlan" authorityYear="1870" box="[746,892,1411,1437]" class="Insecta" family="Calopterygidae" genus="Psolodesmus" kingdom="Animalia" order="Odonata" pageId="15" pageNumber="224" phylum="Arthropoda" rank="genus">
<emphasis id="29AEEAA9E03AD75DFD04E9A3FCB6FA41" box="[746,892,1411,1437]" italics="true" pageId="15" pageNumber="224">Psolodesmus</emphasis>
</taxonomicName>
, but has two setae on the anterior lobes of the prementum (one in
<taxonomicName id="DCDA4D38E03AD75DFE9DE987FDCFFA1D" authorityName="McLachlan" authorityYear="1870" box="[371,517,1447,1473]" class="Insecta" family="Calopterygidae" genus="Psolodesmus" kingdom="Animalia" order="Odonata" pageId="15" pageNumber="224" phylum="Arthropoda" rank="genus">
<emphasis id="29AEEAA9E03AD75DFE9DE987FDCFFA1D" box="[371,517,1447,1473]" italics="true" pageId="15" pageNumber="224">Psolodesmus</emphasis>
</taxonomicName>
). Both genera have a similar robust build (
<figureCitation id="83E12A3EE03AD75DFC0AE987FBF2FA1D" box="[996,1080,1446,1473]" captionStart="FIGURES 52–56. 52" captionStartId="10.[151,264,1941,1965]" captionTargetBox="[191,1388,163,1929]" captionTargetId="figure-21@10.[190,1397,151,1933]" captionTargetPageId="10" captionText="FIGURES 52–56. 52. Caliphaea angka, habitus (based on Yang et al. 2021). 53. Vestalis gracilis, habitus (redrawn from Rattanachan et al. 2022). 54. Neurobasis chinensis, habitus (original drawing). 55. Matrona basilaris, habitus (drawn from Wang et al. 2017). 56. Echo uniformis, habitus (redrawn from Lieftinck 1965)." figureDoi="http://doi.org/10.5281/zenodo.13618339" httpUri="https://zenodo.org/record/13618339/files/figure.png" pageId="15" pageNumber="224">Fig. 56</figureCitation>
). Also in
<taxonomicName id="DCDA4D38E03AD75DFB4CE987FAFEFA1D" authorityName="McLachlan" authorityYear="1870" box="[1186,1332,1447,1473]" class="Insecta" family="Calopterygidae" genus="Psolodesmus" kingdom="Animalia" order="Odonata" pageId="15" pageNumber="224" phylum="Arthropoda" rank="genus">
<emphasis id="29AEEAA9E03AD75DFB4CE987FAFEFA1D" box="[1186,1332,1447,1473]" italics="true" pageId="15" pageNumber="224">Psolodesmus</emphasis>
</taxonomicName>
the short outer, triquetral caudal gills bear very strong teeth along the outer central ridge but not on the dorsal and ventral margins, whereas in
<taxonomicName id="DCDA4D38E03AD75DFE94E9CFFDC9F9D5" authorityName="Selys" authorityYear="1879" box="[378,515,1519,1545]" class="Insecta" family="Calopterygidae" genus="Echo" kingdom="Animalia" order="Odonata" pageId="15" pageNumber="224" phylum="Arthropoda" rank="species" species="uniformis">
<emphasis id="29AEEAA9E03AD75DFE94E9CFFDC9F9D5" box="[378,515,1519,1545]" italics="true" pageId="15" pageNumber="224">E. uniformis</emphasis>
</taxonomicName>
all three ridges bear dentition, according to
<bibRefCitation id="7F4B4B4AE03AD75DFC06E9CFFB7BF9D5" author="Lieftinck, M. A." box="[1000,1201,1518,1545]" pageId="15" pageNumber="224" pagination="325 - 364" refId="ref24671" refString="Lieftinck, M. A. (1965) The species-group of Vestalis amoena Selys, 1853, in Sundaland (Odonata, Calopterygidae). Tijdschrift voor Entomologie, 108, 325 - 364." type="journal article" year="1965">Lieftinck’s (1965)</bibRefCitation>
text, and illustration. The larva of
<taxonomicName id="DCDA4D38E03AD75DFEC9EA33FDCDF9F1" authority="Laidlaw" authorityName="Laidlaw" authorityYear="1902" box="[295,519,1555,1581]" class="Insecta" family="Calopterygidae" genus="Echo" kingdom="Animalia" order="Odonata" pageId="15" pageNumber="224" phylum="Arthropoda" rank="species" species="modesta">
<emphasis id="29AEEAA9E03AD75DFEC9EA33FE69F9F1" box="[295,419,1555,1581]" italics="true" pageId="15" pageNumber="224">E. modesta</emphasis>
Laidlaw
</taxonomicName>
has been collected on Langkawi Island,
<collectingCountry id="63CD762BE03AD75DFC21EA33FBFCF9F1" box="[975,1078,1555,1581]" name="Malaysia" pageId="15" pageNumber="224">Malaysia</collectingCountry>
(S.G. Butler pers. comm.), and, an F-
<specimenCount id="0DDCFD32E03AD75DFF39EA17FE99F98D" box="[215,339,1591,1617]" count="1" pageId="15" pageNumber="224" type="generic">1 specimen</specimenCount>
was collected by AGO in a leaf pack in a very small and shallow rocky branch of a small forest stream in
<collectingRegion id="D91EF859E03AD75DFEEAEA7BFE6EF9A9" box="[260,420,1627,1653]" country="Thailand" name="Kanchanaburi" pageId="15" pageNumber="224">Kanchanaburi</collectingRegion>
,
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in 2003. Adult
<taxonomicName id="DCDA4D38E03AD75DFD53EA7BFCF2F9A9" authorityName="Laidlaw" authorityYear="1902" box="[701,824,1627,1653]" class="Insecta" family="Calopterygidae" genus="Echo" kingdom="Animalia" order="Odonata" pageId="15" pageNumber="224" phylum="Arthropoda" rank="species" species="modesta">
<emphasis id="29AEEAA9E03AD75DFD53EA7BFCF2F9A9" box="[701,824,1627,1653]" italics="true" pageId="15" pageNumber="224">E. modesta</emphasis>
</taxonomicName>
were present and no other species with which the larva might be confused occurs in the area. The species has also been bred, but not yet described (T.S. Keetapithchayakul pers. comm.), confirming the above identifications. Both specimens have strong dentition dorsally and laterally on the margins of the caudal gills as well as lighter denticulations on the outer central ridge (
<figureCitation id="83E12A3EE03AD75DFB67EAE7FB14F93D" box="[1161,1246,1735,1761]" captionStart="FIGURES 42–51. 42" captionStartId="9.[151,264,1592,1616]" captionTargetBox="[165,1413,192,1557]" captionTargetId="figure-15@9.[151,1435,181,1568]" captionTargetPageId="9" captionText="FIGURES 42–51. 42. Archineura incarnata: caudal gills; left lateral above, median below (drawn from Yang et al. 2022). 43. Echo modesta: caudal gills; left lateral above, median below, with baso-dorsal shield arrowed (original illustration). 44. Psolodesmus mandarinus, dorsal view of caudal gills, with baso-dorsal shield of median gill arrowed (redrawn from Matsuki &amp; Lien 1978). 45. Mnais andersoni, left lateral caudal gill with upper margin of median gill visible; baso-dorsal shield of median gill arrowed (original illustration). 46. Mnais gregoryi: caudal gills; left lateral above, median below, with baso-dorsal shield arrowed (drawn from Yang et al. 2021). 47. Caliphaea angka: caudal gills; left lateral above, median below (drawn from Yang et al. 2021). 48. Matrona basilaris: caudal gills; left lateral above, median below (drawn from Wang et al. 2017). 49. Vestalaria venusta: caudal gills; left lateral above, median below (drawn from Wang et al. 2017). 50. Vestalis amoena group: caudal gills; left lateral above, median below (redrawn from Lieftinck 1965). 51. Vestalis melania: caudal gills; left lateral above, median below (redrawn from Guadalquiver et al. 2022)." figureDoi="http://doi.org/10.5281/zenodo.13618337" httpUri="https://zenodo.org/record/13618337/files/figure.png" pageId="15" pageNumber="224">Fig. 43</figureCitation>
); the central gill, which is almost as long as the lateral ones, has dorsally at its base a broad triangular sclerotised and slightly serrate guard plate for about one third its length, which is also well developed on
<taxonomicName id="DCDA4D38E03AD75DFC0CEB2FFBA7F8F5" authorityName="Selys" authorityYear="1879" box="[994,1133,1807,1833]" class="Insecta" family="Calopterygidae" genus="Echo" kingdom="Animalia" order="Odonata" pageId="15" pageNumber="224" phylum="Arthropoda" rank="species" species="uniformis">
<emphasis id="29AEEAA9E03AD75DFC0CEB2FFBA7F8F5" box="[994,1133,1807,1833]" italics="true" pageId="15" pageNumber="224">E. uniformis</emphasis>
</taxonomicName>
. The other genus in which the prementum is similar is
<taxonomicName id="DCDA4D38E03AD75DFE21EB13FDDFF890" authorityName="Selys" authorityYear="1853" box="[463,533,1843,1868]" class="Insecta" family="Calopterygidae" genus="Mnais" kingdom="Animalia" order="Odonata" pageId="15" pageNumber="224" phylum="Arthropoda" rank="genus">
<emphasis id="29AEEAA9E03AD75DFE21EB13FDDFF890" box="[463,533,1843,1868]" italics="true" pageId="15" pageNumber="224">Mnais</emphasis>
</taxonomicName>
, but as in
<taxonomicName id="DCDA4D38E03AD75DFD66EB13FCD0F891" authorityName="McLachlan" authorityYear="1870" box="[648,794,1843,1869]" class="Insecta" family="Calopterygidae" genus="Psolodesmus" kingdom="Animalia" order="Odonata" pageId="15" pageNumber="224" phylum="Arthropoda" rank="genus">
<emphasis id="29AEEAA9E03AD75DFD66EB13FCD0F891" box="[648,794,1843,1869]" italics="true" pageId="15" pageNumber="224">Psolodesmus</emphasis>
</taxonomicName>
it differs in bearing only one pair of setae on the lobes of the prementum. In addition the known larvae of
<taxonomicName id="DCDA4D38E03AD75DFD43EB77FD39F8AC" authorityName="Selys" authorityYear="1853" box="[685,755,1879,1904]" class="Insecta" family="Calopterygidae" genus="Mnais" kingdom="Animalia" order="Odonata" pageId="15" pageNumber="224" phylum="Arthropoda" rank="genus">
<emphasis id="29AEEAA9E03AD75DFD43EB77FD39F8AC" box="[685,755,1879,1904]" italics="true" pageId="15" pageNumber="224">Mnais</emphasis>
</taxonomicName>
are less heavily built and the margins of the outer caudal gills are smooth, without dentition, and usually longer and narrower.
</paragraph>
<paragraph id="1B6536BBE03AD742FF79EBE3FEFFFE9D" blockId="15.[151,1436,1987,2013]" lastBlockId="16.[151,1437,151,321]" lastPageId="16" lastPageNumber="225" pageId="15" pageNumber="224">
Nevertheless, in southern
<collectingCountry id="63CD762BE03AD75DFE53EBE3FDCAF801" box="[445,512,1987,2013]" name="China" pageId="15" pageNumber="224">China</collectingCountry>
there is a distinct possibility that species of
<taxonomicName id="DCDA4D38E03AD75DFC16EBE3FBF8F801" box="[1016,1074,1987,2013]" class="Insecta" family="Calopterygidae" genus="Echo" kingdom="Animalia" order="Odonata" pageId="15" pageNumber="224" phylum="Arthropoda" rank="genus">
<emphasis id="29AEEAA9E03AD75DFC16EBE3FBF8F801" box="[1016,1074,1987,2013]" italics="true" pageId="15" pageNumber="224">Echo</emphasis>
</taxonomicName>
and
<taxonomicName id="DCDA4D38E03AD75DFB87EBE3FB65F800" authorityName="Selys" authorityYear="1853" box="[1129,1199,1987,2012]" class="Insecta" family="Calopterygidae" genus="Mnais" kingdom="Animalia" order="Odonata" pageId="15" pageNumber="224" phylum="Arthropoda" rank="genus">
<emphasis id="29AEEAA9E03AD75DFB87EBE3FB65F800" box="[1129,1199,1987,2012]" italics="true" pageId="15" pageNumber="224">Mnais</emphasis>
</taxonomicName>
may overlap (
<bibRefCitation id="7F4B4B4AE03AD742FABDEBE3FF19FF6D" author="Zhang, H. - M." lastPageId="16" lastPageNumber="225" pageId="15" pageNumber="224" refId="ref28336" refString="Zhang, H. - M. (2019) Dragonflies and Damselflies of China. Vols. 1 &amp; 2. Chong Qing University Press, ChongQing, 14 + 1460 pp. [bilingual, in Chinese and English]" type="book" year="2019">Zhang 2019</bibRefCitation>
), and it is unclear if any of those
<taxonomicName id="DCDA4D38E025D742FDA4ECB7FD11FF6D" box="[586,731,151,177]" pageId="16" pageNumber="225">
<emphasis id="29AEEAA9E025D742FDA4ECB7FD4EFF6D" box="[586,644,151,177]" italics="true" pageId="16" pageNumber="225">Echo</emphasis>
species
</taxonomicName>
(
<taxonomicName id="DCDA4D38E025D742FD02ECB7FC3EFF6D" authority="Zhang et al." authorityName="Zhang" box="[748,1012,151,177]" class="Insecta" family="Calopterygidae" genus="Echo" kingdom="Animalia" order="Odonata" pageId="16" pageNumber="225" phylum="Arthropoda" rank="species" species="candens">
<emphasis id="29AEEAA9E025D742FD02ECB7FCADFF6D" box="[748,871,151,177]" italics="true" pageId="16" pageNumber="225">E. candens</emphasis>
Zhang
<emphasis id="29AEEAA9E025D742FC53ECB7FC24FF6D" box="[957,1006,151,177]" italics="true" pageId="16" pageNumber="225">et al</emphasis>
.
</taxonomicName>
,
<taxonomicName id="DCDA4D38E025D742FBEDECB7FB1DFF6D" authority="Selys" authorityName="Selys" authorityYear="1853" box="[1027,1239,151,177]" class="Insecta" family="Calopterygidae" genus="Echo" kingdom="Animalia" order="Odonata" pageId="16" pageNumber="225" phylum="Arthropoda" rank="species" species="margarita">
<emphasis id="29AEEAA9E025D742FBEDECB7FB5EFF6C" box="[1027,1172,151,176]" italics="true" pageId="16" pageNumber="225">E. margarita</emphasis>
Selys
</taxonomicName>
and
<taxonomicName id="DCDA4D38E025D742FAE0ECB7FEAFFF09" authority="Yu &amp; Hamalainen" authorityName="Yu &amp; Hamalainen" class="Insecta" family="Calopterygidae" genus="Echo" kingdom="Animalia" order="Odonata" pageId="16" pageNumber="225" phylum="Arthropoda" rank="species" species="perornata">
<emphasis id="29AEEAA9E025D742FAE0ECB7FA56FF6C" box="[1294,1436,151,176]" italics="true" pageId="16" pageNumber="225">E. perornata</emphasis>
Yu &amp; Hämäläinen
</taxonomicName>
) have the same larval form and same distinguishing characters as
<taxonomicName id="DCDA4D38E025D742FBBAEC9BFB15FF09" authorityName="Selys" authorityYear="1879" box="[1108,1247,187,213]" class="Insecta" family="Calopterygidae" genus="Echo" kingdom="Animalia" order="Odonata" pageId="16" pageNumber="225" phylum="Arthropoda" rank="species" species="uniformis">
<emphasis id="29AEEAA9E025D742FBBAEC9BFB15FF09" box="[1108,1247,187,213]" italics="true" pageId="16" pageNumber="225">E. uniformis</emphasis>
</taxonomicName>
and
<taxonomicName id="DCDA4D38E025D742FAF6EC9BFA5FFF09" authorityName="Laidlaw" authorityYear="1902" box="[1304,1429,187,213]" class="Insecta" family="Calopterygidae" genus="Echo" kingdom="Animalia" order="Odonata" pageId="16" pageNumber="225" phylum="Arthropoda" rank="species" species="modesta">
<emphasis id="29AEEAA9E025D742FAF6EC9BFA5FFF09" box="[1304,1429,187,213]" italics="true" pageId="16" pageNumber="225">E. modesta</emphasis>
</taxonomicName>
, or indeed, if all
<taxonomicName id="DCDA4D38E025D742FEBCECFFFE38FF25" box="[338,498,223,249]" pageId="16" pageNumber="225">
<emphasis id="29AEEAA9E025D742FEBCECFFFE52FF24" box="[338,408,223,248]" italics="true" pageId="16" pageNumber="225">Mnais</emphasis>
species
</taxonomicName>
differ in the way we presently recognise. A possible hint as to the differences we may expect has been suggested (
<bibRefCitation id="7F4B4B4AE025D742FDE7ED23FD2BFEC1" author="Orr, A. G. &amp; Butler, S. G." box="[521,737,259,285]" pageId="16" pageNumber="225" pagination="155 - 168" refId="ref25814" refString="Orr, A. G. &amp; Butler, S. G. (2024) A riddle wrapped in a mystery inside an enigma: notes on an enigmatic larva collected in Nepal, with a reappraisal of Fraser's so called ' Caliphaea confusa ' larva from Meghalaya, India (Odonata: Calopterygidae). Odonatologica, 53, 155 - 168. https: // doi. org / 10.60024 / odon. v 53 i 1 - 2. a 8" type="journal article" year="2024">Orr &amp; Butler 2024</bibRefCitation>
), but until we have certified voucher specimens, this question remains open.
</paragraph>
</subSubSection>
</treatment>
</document>

128
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<subSubSection id="53C06530E03AD75DFF79EE66FEC7FDBD" box="[151,269,582,609]" pageId="15" pageNumber="224" type="nomenclature">
<paragraph id="1B6536BBE03AD75DFF79EE66FEC7FDBD" blockId="15.[151,1437,582,1149]" box="[151,269,582,609]" pageId="15" pageNumber="224">
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<taxonomicName id="DCDA4D38E03AD75DFF79EE66FEC7FDBD" ID-CoL="3F9W" authorityName="Hagen in Selys" authorityYear="1859" box="[151,269,582,609]" class="Insecta" family="Calopterygidae" genus="Caliphaea" kingdom="Animalia" order="Odonata" pageId="15" pageNumber="224" phylum="Arthropoda" rank="genus">
<emphasis id="29AEEAA9E03AD75DFF79EE66FEC7FDBD" bold="true" box="[151,269,582,609]" italics="true" pageId="15" pageNumber="224">Caliphaea</emphasis>
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<paragraph id="1B6536BBE03AD75DFF79EE4BFA41FBA1" blockId="15.[151,1437,582,1149]" pageId="15" pageNumber="224">
The genus includes five species inhabiting mountainous regions from
<collectingCountry id="63CD762BE03AD75DFC5FEE4BFC3EFD59" box="[945,1012,619,645]" name="Nepal" pageId="15" pageNumber="224">Nepal</collectingCountry>
to north-east
<collectingCountry id="63CD762BE03AD75DFB7CEE4BFB04FD59" box="[1170,1230,619,645]" name="India" pageId="15" pageNumber="224">India</collectingCountry>
, south and central
<collectingCountry id="63CD762BE03AD75DFF79EEAFFF17FD75" box="[151,221,655,681]" name="China" pageId="15" pageNumber="224">China</collectingCountry>
, northern
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,
<collectingCountry id="63CD762BE03AD75DFE25EEAFFDFBFD75" box="[459,561,655,681]" name="Thailand" pageId="15" pageNumber="224">Thailand</collectingCountry>
,
<collectingCountry id="63CD762BE03AD75DFDD0EEAFFDBEFD75" box="[574,628,655,681]" name="Laos" pageId="15" pageNumber="224">Laos</collectingCountry>
and
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. The stout, long-legged larva has been described in detail only for
<taxonomicName id="DCDA4D38E03AD75DFF16EE93FD8CFD11" authority="Hamalainen" authorityName="Hamalainen" authorityYear="2003" box="[248,582,691,717]" class="Insecta" family="Calopterygidae" genus="Caliphaea" kingdom="Animalia" order="Odonata" pageId="15" pageNumber="224" phylum="Arthropoda" rank="species" species="angka">
<emphasis id="29AEEAA9E03AD75DFF16EE93FE7DFD11" box="[248,439,691,717]" italics="true" pageId="15" pageNumber="224">Caliphaea angka</emphasis>
Hämäläinen
</taxonomicName>
(
<bibRefCitation id="7F4B4B4AE03AD75DFDBAEE93FCD8FD11" author="Yang, G. - H. &amp; Zhang, H. - M. &amp; Orr, A. G." box="[596,786,691,717]" pageId="15" pageNumber="224" pagination="276 - 284" refId="ref27918" refString="Yang, G. - H., Zhang, H. - M. &amp; Orr, A. G. (2021) Descriptions of larvae of Caliphaea angka Hamalainen, 2003 and Mnais gregoryi Fraser, 1924 (Odonata: Calopterygidae). Zootaxa, 4926 (2), 276 - 284. https: // doi. org / 10.11646 / zootaxa. 4926.2.7" type="journal article" year="2021">
Yang
<emphasis id="29AEEAA9E03AD75DFD79EE93FD02FD11" box="[663,712,691,717]" italics="true" pageId="15" pageNumber="224">et al</emphasis>
. 2021
</bibRefCitation>
), and was collected at
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“in small sluggish montane streams, with numerous hydro-phytes”. Apart from its distinctive habitus (
<figureCitation id="83E12A3EE03AD75DFC21EEF7FBEBFD2D" box="[975,1057,726,753]" captionStart="FIGURES 52–56. 52" captionStartId="10.[151,264,1941,1965]" captionTargetBox="[191,1388,163,1929]" captionTargetId="figure-21@10.[190,1397,151,1933]" captionTargetPageId="10" captionText="FIGURES 52–56. 52. Caliphaea angka, habitus (based on Yang et al. 2021). 53. Vestalis gracilis, habitus (redrawn from Rattanachan et al. 2022). 54. Neurobasis chinensis, habitus (original drawing). 55. Matrona basilaris, habitus (drawn from Wang et al. 2017). 56. Echo uniformis, habitus (redrawn from Lieftinck 1965)." figureDoi="http://doi.org/10.5281/zenodo.13618339" httpUri="https://zenodo.org/record/13618339/files/figure.png" pageId="15" pageNumber="224">Fig. 52</figureCitation>
), with very short ovoid caudal gill bearing a row of 5–6 tubercles along the outer ridge, with the median gill much smaller and thin and membranous (
<figureCitation id="83E12A3EE03AD75DFF71EF3FFF3FFCE5" box="[159,245,799,825]" captionStart="FIGURES 42–51. 42" captionStartId="9.[151,264,1592,1616]" captionTargetBox="[165,1413,192,1557]" captionTargetId="figure-15@9.[151,1435,181,1568]" captionTargetPageId="9" captionText="FIGURES 42–51. 42. Archineura incarnata: caudal gills; left lateral above, median below (drawn from Yang et al. 2022). 43. Echo modesta: caudal gills; left lateral above, median below, with baso-dorsal shield arrowed (original illustration). 44. Psolodesmus mandarinus, dorsal view of caudal gills, with baso-dorsal shield of median gill arrowed (redrawn from Matsuki &amp; Lien 1978). 45. Mnais andersoni, left lateral caudal gill with upper margin of median gill visible; baso-dorsal shield of median gill arrowed (original illustration). 46. Mnais gregoryi: caudal gills; left lateral above, median below, with baso-dorsal shield arrowed (drawn from Yang et al. 2021). 47. Caliphaea angka: caudal gills; left lateral above, median below (drawn from Yang et al. 2021). 48. Matrona basilaris: caudal gills; left lateral above, median below (drawn from Wang et al. 2017). 49. Vestalaria venusta: caudal gills; left lateral above, median below (drawn from Wang et al. 2017). 50. Vestalis amoena group: caudal gills; left lateral above, median below (redrawn from Lieftinck 1965). 51. Vestalis melania: caudal gills; left lateral above, median below (redrawn from Guadalquiver et al. 2022)." figureDoi="http://doi.org/10.5281/zenodo.13618337" httpUri="https://zenodo.org/record/13618337/files/figure.png" pageId="15" pageNumber="224">Fig. 47</figureCitation>
); the prementum is distinctive, with a very narrow deep median cleft and the lateral lobes well separated apically, and the scape of the antenna is less than half its total length (
<figureCitation id="83E12A3EE03AD75DFC72EF63FC3AFC81" box="[924,1008,835,861]" captionStart="FIGURES 35–41. 35" captionStartId="8.[151,264,1776,1800]" captionTargetBox="[257,1332,159,1764]" captionTargetId="figure-21@8.[253,1335,153,1771]" captionTargetPageId="8" captionText="FIGURES 35–41. 35. Archineura incarnata, prementum (dorsal view, drawn from Yang et al. 2022). 36. Vestalis gracilis, prementum (dorsal view) and right antenna with supplementary antennomere between pedicel and scape (a) arrowed (redrawn from Rattanachan et al. 2022). 37. Vestalaria venusta: prementum (dorsal view) and right antenna (drawn from Wang et al. 2017). 38. Caliphaea angka: prementum (dorsal view) and right antenna (drawn from Yang et al. 2021). 39. Heliocypha biseriata, frontal view of labrum showing dorsal and ventral sections and ‘moustache’ of long, heavy setae on the latter (from Orr et al. 2024). 40. Heliocypha biseriata: details of labial palp in relation to prementum (a) dorsal view—base of movable hook dashed to reveal palpal lobe beneath and open base of premental median cleft arrowed, (b) frontal view, (c) inner view (from Orr et al. 2024). 41. Heliocypha biseriata, detail of fore tarsus with pectinate setae magnified (from Orr et al. 2024)." figureDoi="http://doi.org/10.5281/zenodo.13618335" httpUri="https://zenodo.org/record/13618335/files/figure.png" pageId="15" pageNumber="224">Fig. 38</figureCitation>
). The best-known species of the genus is
<taxonomicName id="DCDA4D38E03AD75DFF5CEF47FEB7FC5D" authority="Hagen" authorityName="Hagen" box="[178,381,871,897]" class="Insecta" family="Calopterygidae" genus="Caliphaea" kingdom="Animalia" order="Odonata" pageId="15" pageNumber="224" phylum="Arthropoda" rank="species" species="confusa">
<emphasis id="29AEEAA9E03AD75DFF5CEF47FEE6FC5D" box="[178,300,871,897]" italics="true" pageId="15" pageNumber="224">C. confusa</emphasis>
Hagen
</taxonomicName>
but the larva has not yet been formally described or illustrated, the purported description by
<bibRefCitation id="7F4B4B4AE03AD75DFF79EFABFEE4FC79" author="Fraser, F. C." box="[151,302,907,933]" pageId="15" pageNumber="224" pagination="81 - 93" refId="ref23155" refString="Fraser, F. C. (1943) New oriental Odonata larvae. Proceedings of the Royal entomological society of London, Series B, 12, 81 - 93. https: // doi. org / 10.1111 / j. 1365 - 3113.1943. tb 00749. x" type="journal article" year="1943">Fraser (1943)</bibRefCitation>
having been shown to be based on a misidentification (
<bibRefCitation id="7F4B4B4AE03AD75DFC78EFABFBA0FC79" author="Orr, A. G. &amp; Butler, S. G." box="[918,1130,907,933]" pageId="15" pageNumber="224" pagination="155 - 168" refId="ref25814" refString="Orr, A. G. &amp; Butler, S. G. (2024) A riddle wrapped in a mystery inside an enigma: notes on an enigmatic larva collected in Nepal, with a reappraisal of Fraser's so called ' Caliphaea confusa ' larva from Meghalaya, India (Odonata: Calopterygidae). Odonatologica, 53, 155 - 168. https: // doi. org / 10.60024 / odon. v 53 i 1 - 2. a 8" type="journal article" year="2024">Orr &amp; Butler 2024</bibRefCitation>
). Larva of this species have been collected in
<collectingCountry id="63CD762BE03AD75DFEB1EF8FFE68FC15" box="[351,418,943,969]" name="Nepal" pageId="15" pageNumber="224">Nepal</collectingCountry>
(S.G. Butler pers comm.), and its similarity to
<taxonomicName id="DCDA4D38E03AD75DFC54EF8FFBEAFC15" authorityName="Hamalainen" authorityYear="2003" box="[954,1056,943,969]" class="Insecta" family="Calopterygidae" genus="Caliphaea" kingdom="Animalia" order="Odonata" pageId="15" pageNumber="224" phylum="Arthropoda" rank="species" species="angka">
<emphasis id="29AEEAA9E03AD75DFC54EF8FFBEAFC15" box="[954,1056,943,969]" italics="true" pageId="15" pageNumber="224">C. angka</emphasis>
</taxonomicName>
leaves no doubt as to its identity, given that it is the only species of the genus in those localities. It differs slightly in the shape of the prementum and the outer caudal gills are slightly more elongate. The species in the genus are all closely related and all larvae are expected to have similar characters. In addition, I collected in 2003 a single mature specimen of
<taxonomicName id="DCDA4D38E03AD75DFB07E83BFA94FBE9" authorityName="Hagen in Selys" authorityYear="1859" box="[1257,1374,1051,1077]" class="Insecta" family="Calopterygidae" genus="Caliphaea" kingdom="Animalia" order="Odonata" pageId="15" pageNumber="224" phylum="Arthropoda" rank="genus">
<emphasis id="29AEEAA9E03AD75DFB07E83BFA94FBE9" box="[1257,1374,1051,1077]" italics="true" pageId="15" pageNumber="224">Caliphaea</emphasis>
</taxonomicName>
from Doi Inthanon,
<collectingCountry id="63CD762BE03AD75DFED9E81FFE51FB85" box="[311,411,1087,1113]" name="Thailand" pageId="15" pageNumber="224">Thailand</collectingCountry>
at ca
<quantity id="DC229B5EE03AD75DFE34E81FFDEDFB85" box="[474,551,1087,1113]" metricMagnitude="3" metricUnit="m" metricValue="1.3" pageId="15" pageNumber="224" unit="m" value="1300.0">1300m</quantity>
which may be
<taxonomicName id="DCDA4D38E03AD75DFD21E81FFC1AFB85" authority="Asahina" authorityName="Asahina" authorityYear="1976" box="[719,976,1087,1113]" class="Insecta" family="Calopterygidae" genus="Caliphaea" kingdom="Animalia" order="Odonata" pageId="15" pageNumber="224" phylum="Arthropoda" rank="species" species="thailandica">
<emphasis id="29AEEAA9E03AD75DFD21E81FFCA4FB85" box="[719,878,1087,1113]" italics="true" pageId="15" pageNumber="224">C. thailandica</emphasis>
Asahina
</taxonomicName>
(Orr unpublished data). It differs in some respects, notably general build, from Himalayan
<taxonomicName id="DCDA4D38E03AD75DFD55E843FCF8FBA1" box="[699,818,1123,1149]" class="Insecta" family="Calopterygidae" genus="Caliphaea" kingdom="Animalia" order="Odonata" pageId="15" pageNumber="224" phylum="Arthropoda" rank="species" species="confusa">
<emphasis id="29AEEAA9E03AD75DFD55E843FCF8FBA1" box="[699,818,1123,1149]" italics="true" pageId="15" pageNumber="224">C. confusa</emphasis>
</taxonomicName>
and
<taxonomicName id="DCDA4D38E03AD75DFC87E843FC07FBA1" authorityName="Hamalainen" authorityYear="2003" box="[873,973,1123,1149]" class="Insecta" family="Calopterygidae" genus="Caliphaea" kingdom="Animalia" order="Odonata" pageId="15" pageNumber="224" phylum="Arthropoda" rank="species" species="angka">
<emphasis id="29AEEAA9E03AD75DFC87E843FC07FBA1" box="[873,973,1123,1149]" italics="true" pageId="15" pageNumber="224">C. angka</emphasis>
</taxonomicName>
but agrees with them in key characters.
</paragraph>
</subSubSection>
</treatment>
</document>

108
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<paragraph id="1B6536BBE03BD75CFF79E9CFFED7F9D6" blockId="14.[151,1437,1519,1870]" box="[151,285,1519,1546]" pageId="14" pageNumber="223">
<heading id="402D81D7E03BD75CFF79E9CFFED7F9D6" bold="true" box="[151,285,1519,1546]" fontSize="11" level="1" pageId="14" pageNumber="223" reason="1">
<taxonomicName id="DCDA4D38E03BD75CFF79E9CFFED7F9D6" ID-CoL="326K" authorityName="Kirby" authorityYear="1894" box="[151,285,1519,1546]" class="Insecta" family="Calopterygidae" genus="Archineura" kingdom="Animalia" order="Odonata" pageId="14" pageNumber="223" phylum="Arthropoda" rank="genus">
<emphasis id="29AEEAA9E03BD75CFF79E9CFFED7F9D6" bold="true" box="[151,285,1519,1546]" italics="true" pageId="14" pageNumber="223">Archineura</emphasis>
</taxonomicName>
</heading>
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<subSubSection id="53C06530E03BD75CFF79EA33FEE1F891" pageId="14" pageNumber="223" type="discussion">
<paragraph id="1B6536BBE03BD75CFF79EA33FEE1F891" blockId="14.[151,1437,1519,1870]" pageId="14" pageNumber="223">
The genus includes just three species, two from southern
<collectingCountry id="63CD762BE03BD75CFCE0EA33FC9BF9F1" box="[782,849,1555,1581]" name="China" pageId="14" pageNumber="223">China</collectingCountry>
and one from northern
<collectingCountry id="63CD762BE03BD75CFBBAEA33FB7CF9F1" box="[1108,1206,1555,1581]" name="Vietnam" pageId="14" pageNumber="223">Vietnam</collectingCountry>
,
<taxonomicName id="DCDA4D38E03BD75CFB51EA33FF33F98D" authority="(Martin)" baseAuthorityName="Martin" baseAuthorityYear="1904" class="Insecta" family="Calopterygidae" genus="Archineura" kingdom="Animalia" order="Odonata" pageId="14" pageNumber="223" phylum="Arthropoda" rank="species" species="maxima">
<emphasis id="29AEEAA9E03BD75CFB51EA33FA57F9F1" box="[1215,1437,1555,1581]" italics="true" pageId="14" pageNumber="223">Archineura maxima</emphasis>
(Martin)
</taxonomicName>
, which is likely already extinct (
<bibRefCitation id="7F4B4B4AE03BD75CFD8CEA17FCE5F98E" author="Hamalainen, M." box="[610,815,1591,1618]" pageId="14" pageNumber="223" pagination="157 - 168" refId="ref23338" refString="Hamalainen, M. (2015) From Echo maxima to Archineura maxima - a slow taxonomic process (Odonata: Calopterygidae). Notulae odonatologicae, 8, 157 - 168." type="journal article" year="2015">Hämäläinen 2015</bibRefCitation>
). The larva of
<taxonomicName id="DCDA4D38E03BD75CFC21EA17FB09F98D" authority="(Karsch)" baseAuthorityName="Karsch" box="[975,1219,1591,1617]" class="Insecta" family="Calopterygidae" genus="Archineura" kingdom="Animalia" order="Odonata" pageId="14" pageNumber="223" phylum="Arthropoda" rank="species" species="incarnata">
<emphasis id="29AEEAA9E03BD75CFC21EA17FB90F98D" box="[975,1114,1591,1617]" italics="true" pageId="14" pageNumber="223">A. incarnata</emphasis>
(Karsch)
</taxonomicName>
has been described in detail (
<bibRefCitation id="7F4B4B4AE03BD75CFEEAEA7BFE0CF9A9" author="Yang, G. - H. &amp; Orr, A. G. &amp; Zhang, H. - M." box="[260,454,1627,1654]" pageId="14" pageNumber="223" pagination="441 - 447" refId="ref27846" refString="Yang, G. - H., Orr, A. G. &amp; Zhang, H. - M. (2022) First description of the larva of Archineura incarnata (Karsch, 1891) with notes on the biology (Odonata: Calopterygidae). Zootaxa, 5134 (3), 441 - 447. https: // doi. org / 10.11646 / zootaxa. 5134.3.8" type="journal article" year="2022">
Yang
<emphasis id="29AEEAA9E03BD75CFEA7EA7CFEB6F9A9" box="[329,380,1627,1653]" italics="true" pageId="14" pageNumber="223">et al</emphasis>
. 2022
</bibRefCitation>
) and by virtue of its size and uniquely elongate, tuberculate caudal lamellae (
<figureCitation id="83E12A3EE03BD75CFAAFEA7BFA5EF9A9" box="[1345,1428,1627,1653]" captionStart="FIGURES 42–51. 42" captionStartId="9.[151,264,1592,1616]" captionTargetBox="[165,1413,192,1557]" captionTargetId="figure-15@9.[151,1435,181,1568]" captionTargetPageId="9" captionText="FIGURES 42–51. 42. Archineura incarnata: caudal gills; left lateral above, median below (drawn from Yang et al. 2022). 43. Echo modesta: caudal gills; left lateral above, median below, with baso-dorsal shield arrowed (original illustration). 44. Psolodesmus mandarinus, dorsal view of caudal gills, with baso-dorsal shield of median gill arrowed (redrawn from Matsuki &amp; Lien 1978). 45. Mnais andersoni, left lateral caudal gill with upper margin of median gill visible; baso-dorsal shield of median gill arrowed (original illustration). 46. Mnais gregoryi: caudal gills; left lateral above, median below, with baso-dorsal shield arrowed (drawn from Yang et al. 2021). 47. Caliphaea angka: caudal gills; left lateral above, median below (drawn from Yang et al. 2021). 48. Matrona basilaris: caudal gills; left lateral above, median below (drawn from Wang et al. 2017). 49. Vestalaria venusta: caudal gills; left lateral above, median below (drawn from Wang et al. 2017). 50. Vestalis amoena group: caudal gills; left lateral above, median below (redrawn from Lieftinck 1965). 51. Vestalis melania: caudal gills; left lateral above, median below (redrawn from Guadalquiver et al. 2022)." figureDoi="http://doi.org/10.5281/zenodo.13618337" httpUri="https://zenodo.org/record/13618337/files/figure.png" pageId="14" pageNumber="223">Fig. 42</figureCitation>
) cannot be mistaken with any other genus. According to
<bibRefCitation id="7F4B4B4AE03BD75CFCE9EAA0FC1FF945" author="Yang, G. - H. &amp; Orr, A. G. &amp; Zhang, H. - M." box="[775,981,1663,1690]" pageId="14" pageNumber="223" pagination="441 - 447" refId="ref27846" refString="Yang, G. - H., Orr, A. G. &amp; Zhang, H. - M. (2022) First description of the larva of Archineura incarnata (Karsch, 1891) with notes on the biology (Odonata: Calopterygidae). Zootaxa, 5134 (3), 441 - 447. https: // doi. org / 10.11646 / zootaxa. 5134.3.8" type="journal article" year="2022">
Yang
<emphasis id="29AEEAA9E03BD75CFCA6EAA0FCB3F945" box="[840,889,1663,1689]" italics="true" pageId="14" pageNumber="223">et al</emphasis>
. (2022)
</bibRefCitation>
, “they are typical claspers and are found on the under surface large stones and boulders in the middle of streams, but not at the stream edge where most other larvae of the family are found...” and, “If a rock provides a suitable microhabitat and is sufficiently large, 2–
<specimenCount id="0DDCFD32E03BD75CFAABEAE7FA56F93D" box="[1349,1436,1735,1761]" count="3" pageId="14" pageNumber="223" type="larva">3 larvae</specimenCount>
can be found together.”, these notes being based on both presumed larvae of
<taxonomicName id="DCDA4D38E03BD75CFC0AEACBFB30F8D9" authority="(Fraser)" baseAuthorityName="Fraser" box="[996,1274,1771,1797]" class="Insecta" family="Calopterygidae" genus="Archineura" kingdom="Animalia" order="Odonata" pageId="14" pageNumber="223" phylum="Arthropoda" rank="species" species="hetaerinoides">
<emphasis id="29AEEAA9E03BD75CFC0AEACBFB51F8D9" box="[996,1179,1771,1797]" italics="true" pageId="14" pageNumber="223">A. hetaerinoides</emphasis>
(Fraser)
</taxonomicName>
from Guangxi and
<taxonomicName id="DCDA4D38E03BD75CFF2BEB2FFE9AF8F5" box="[197,336,1807,1833]" class="Insecta" family="Calopterygidae" genus="Archineura" kingdom="Animalia" order="Odonata" pageId="14" pageNumber="223" phylum="Arthropoda" rank="species" species="incarnata">
<emphasis id="29AEEAA9E03BD75CFF2BEB2FFE9AF8F5" box="[197,336,1807,1833]" italics="true" pageId="14" pageNumber="223">A. incarnata</emphasis>
</taxonomicName>
from Guangdong. The presumed
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<emphasis id="29AEEAA9E03BD75CFD2BEB2FFCB1F8F5" box="[709,891,1807,1833]" italics="true" pageId="14" pageNumber="223">A. hetaerinoides</emphasis>
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larva is very similar to
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<emphasis id="29AEEAA9E03BD75CFB90EB2FFAC3F8F5" box="[1150,1289,1807,1833]" italics="true" pageId="14" pageNumber="223">A. incarnata</emphasis>
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(H-M Zhang pers comm.).
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Family
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Just four genera and 15 species of this mainly Australian and Oceanian realm (sensu
<bibRefCitation id="7F4B4B4AE03BD75CFB6AECFFFA8BFF25" author="Holt, B. G. &amp; Lessard, J. - P. &amp; Borregaard, M. K. &amp; Fritz, S. A. &amp; Araujo, M. B. &amp; Dimitrov, D. &amp; Fabre, P. - H. &amp; Graham, C. H. &amp; Graves, G. R. &amp; Jonsson, K. A. &amp; Nogues-Bravo, D. &amp; Wang, Z. &amp; Whittaker, R. J. &amp; Fjeldsa, J. &amp; Rahbek, C." box="[1156,1345,223,250]" pageId="14" pageNumber="223" pagination="74 - 78" refId="ref23468" refString="Holt, B. G., Lessard, J. - P., Borregaard, M. K., Fritz, S. A., Araujo, M. B., Dimitrov, D., Fabre, P. - H., Graham, C. H., Graves, G. R., Jonsson, K. A., Nogues-Bravo, D., Wang, Z., Whittaker, R. J., Fjeldsa, J. &amp; Rahbek, C. (2013) An update of Wallace's zoogeographic regions of the world. Science, 339, 74 - 78. https: // doi. org / 10.1126 / science. 1228282" type="journal article" year="2013">
Holt
<emphasis id="29AEEAA9E03BD75CFB2EECC0FB3FFF25" box="[1216,1269,223,249]" italics="true" pageId="14" pageNumber="223">et al</emphasis>
. 2013
</bibRefCitation>
) family occur in the Oriental realm, and only one genus,
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<emphasis id="29AEEAA9E03BD75CFD34ED23FC99FEC1" box="[730,851,259,285]" italics="true" pageId="14" pageNumber="223">Podolestes</emphasis>
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(9 spp.), occurs on the Asian mainland (
<bibRefCitation id="7F4B4B4AE03BD75CFADAED23FE44FE9D" author="Kalkman, V. J. &amp; Theischinger, G." pageId="14" pageNumber="223" pagination="1 - 52" refId="ref23895" refString="Kalkman, V. J. &amp; Theischinger, G. (2013) Generic revision of Argiolestidae (Odonata), with four new genera. International Journal of Odonatology, 16, 1 - 52. https: // doi. org / 10.1080 / 13887890.2012.749450" type="journal article" year="2013">Kalkman &amp; Theischinger 2013</bibRefCitation>
). The larva is known and described only for
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<emphasis id="29AEEAA9E03BD75CFC7AED08FBD7FE9D" box="[916,1053,295,321]" italics="true" pageId="14" pageNumber="223">P. orientalis</emphasis>
Selys
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(
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), but other species of the genus are probably similar. Based on their well-known Australian relatives all argiolestids are believed to possess broad, rounded or foliate caudal gills arranged in a horizontal fan, and most are of moderately stout build.
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<emphasis id="29AEEAA9E03BD75CFEF5EDB3FE5EFE71" box="[283,404,403,429]" italics="true" pageId="14" pageNumber="223">Podolestes</emphasis>
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(
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), is known west of Sulawesi and the
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and in eastern mainland Asia where it might be broadly sympatric with species of
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which argiolestid larvae most resemble. However,
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larvae have much larger gill lamellae, a different head shape and finer, longer antennae; the prementum is long and narrow, expanded anteriorly with almost no median lobe (
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). They inhabit pools in lowland swamp forest, as opposed to swift upland streams. Even in situations where the adults are abundant, the larvae can be very difficult to locate, but they have been discovered among leaf litter in shallow forest pools and in bankside root masses in very slowly flowing streams. According to
<bibRefCitation id="7F4B4B4AE03BD75CFD02EE4BFC05FD59" author="Ngiam, R. W. J. &amp; Ng, M. F. C." box="[748,975,619,646]" pageId="14" pageNumber="223" refId="ref25398" refString="Ngiam, R. W. J. &amp; Ng, M. F. C. (2022) A photographic field guide to the dragonflies &amp; damselflies of Singapore. John Beaufoy Publishing, Oxford, 340 pp." type="book" year="2022">Ngiam &amp; Ng (2022)</bibRefCitation>
the larva “Hunts among fallen leaf packs at shallow edges of forest pools, with abdomen raised and gills aimed at the surface, possibly to maximise oxygen uptake in poorly oxygenated pools”.
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Larva unknown:
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<emphasis id="29AEEAA9E03BD75CFE68EEF7FDC9FD2D" box="[390,515,727,753]" italics="true" pageId="14" pageNumber="223">Argiolestes</emphasis>
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(
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),
<taxonomicName id="DCDA4D38E03BD75CFD8EEEF7FCD1FD2D" authorityName="Kennedy" authorityYear="1925" box="[608,795,727,753]" class="Insecta" family="Megapodagrionidae" genus="Celebargiolestes" kingdom="Animalia" order="Odonata" pageId="14" pageNumber="223" phylum="Arthropoda" rank="genus">
<emphasis id="29AEEAA9E03BD75CFD8EEEF7FCD1FD2D" box="[608,795,727,753]" italics="true" pageId="14" pageNumber="223">Celebargiolestes</emphasis>
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(4 spp.),
<taxonomicName id="DCDA4D38E03BD75CFC68EEF7FB8EFD2D" authorityName="Kalkman &amp; Theischinger" authorityYear="2013" box="[902,1092,727,753]" class="Insecta" family="Argiolestidae" genus="Luzonargiolestes" higherTaxonomySource="CoL,GBIF" kingdom="Animalia" order="Odonata" pageId="14" pageNumber="223" phylum="Arthropoda" rank="genus">
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(2 spp.)
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<mods:identifier id="1BEB64ABB6ED321A7F9D43E0A4321372" type="ISSN">1175-5326</mods:identifier>
<mods:identifier id="136B6CCFD17107EE1BAC2D1404B0E5E7" type="Zenodo-Dep">13618317</mods:identifier>
<mods:identifier id="80319CDD4EF2E2C8624472DB21749AF3" type="ZooBank">B3C66D95-3585-4920-BE93-A44D33FB2FBB</mods:identifier>
</mods:mods>
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<subSubSection id="53C06530E03BD75CFF79EF63FE7DFC81" box="[151,439,835,861]" pageId="14" pageNumber="223" type="nomenclature">
<paragraph id="1B6536BBE03BD75CFF79EF63FE7DFC81" blockId="14.[151,439,835,861]" box="[151,439,835,861]" pageId="14" pageNumber="223">
<heading id="402D81D7E03BD75CFF79EF63FE7DFC81" bold="true" box="[151,439,835,861]" fontSize="11" level="1" pageId="14" pageNumber="223" reason="1">
<emphasis id="29AEEAA9E03BD75CFF79EF63FE7DFC81" bold="true" box="[151,439,835,861]" pageId="14" pageNumber="223">
Family
<taxonomicName id="DCDA4D38E03BD75CFF19EF63FE7DFC81" ID-CoL="7LD" authorityName="Selys" authorityYear="1850" box="[247,439,835,861]" class="Insecta" family="Calopterygidae" kingdom="Animalia" order="Odonata" pageId="14" pageNumber="223" phylum="Arthropoda" rank="family">Calopterygidae</taxonomicName>
</emphasis>
</heading>
</paragraph>
</subSubSection>
<subSubSection id="53C06530E03BD75CFF79EFABFD4CFA89" pageId="14" pageNumber="223" type="discussion">
<paragraph id="1B6536BBE03BD75CFF79EFABFAF6FAED" blockId="14.[151,1437,907,1366]" pageId="14" pageNumber="223">
The larvae of 11 of the 12 genera present in the region have been described and approximately one third of the 68 species are known but not necessarily described. The larvae
<emphasis id="29AEEAA9E03BD75CFCBEEF8FFCACFC15" box="[848,870,943,969]" italics="true" pageId="14" pageNumber="223">of</emphasis>
<taxonomicName id="DCDA4D38E03BD75CFC81EF8FFA96FC15" authority="Sjostedt" authorityName="Sjostedt" authorityYear="1926" box="[879,1372,943,969]" class="Insecta" family="Calopterygidae" genus="Neurobasis" kingdom="Animalia" order="Odonata" pageId="14" pageNumber="223" phylum="Arthropoda" rank="species" species="anderssoni" subGenus="Sinobasis">
<emphasis id="29AEEAA9E03BD75CFC81EF8FFC24FC15" box="[879,1006,943,969]" italics="true" pageId="14" pageNumber="223">Neurobasis</emphasis>
(
<emphasis id="29AEEAA9E03BD75CFBEEEF8FFBA1FC15" box="[1024,1131,943,969]" italics="true" pageId="14" pageNumber="223">Sinobasis</emphasis>
)
<emphasis id="29AEEAA9E03BD75CFB90EF8FFB33FC15" box="[1150,1273,943,969]" italics="true" pageId="14" pageNumber="223">anderssoni</emphasis>
Sjöstedt
</taxonomicName>
from
<collectingCountry id="63CD762BE03BD75CFF79EFF3FF10FC31" box="[151,218,979,1005]" name="China" pageId="14" pageNumber="223">China</collectingCountry>
is unknown and likely to be distinctive. All species are long legged and elongate, but some genera are stouter; in all species the first antennal segment (scape) is greatly enlarged, broad at the base and longer than or as long as the remaining six segments and often longer than the head is wide; intergeneric variation occurs mainly in the form of the prementum and of the caudal gills, which vary greatly in relative length and in structure. We can mostly be confident of generic level classification, but too few species of the pairs of similar genera
<taxonomicName id="DCDA4D38E03BD75CFB63E843FAFEFBA1" authorityName="Dumont, Vanfleteren, De Jonckheere &amp; Weekers" authorityYear="2005" box="[1165,1332,1123,1149]" class="Insecta" family="Calopterygidae" genus="Atrocalopteryx" kingdom="Animalia" order="Odonata" pageId="14" pageNumber="223" phylum="Arthropoda" rank="genus">
<emphasis id="29AEEAA9E03BD75CFB63E843FAFEFBA1" box="[1165,1332,1123,1149]" italics="true" pageId="14" pageNumber="223">Atrocalopteryx</emphasis>
</taxonomicName>
/
<taxonomicName id="DCDA4D38E03BD75CFAD5E844FA56FBA1" authorityName="Selys" authorityYear="1853" box="[1339,1436,1124,1149]" class="Insecta" family="Calopterygidae" genus="Matrona" kingdom="Animalia" order="Odonata" pageId="14" pageNumber="223" phylum="Arthropoda" rank="genus">
<emphasis id="29AEEAA9E03BD75CFAD5E844FA56FBA1" box="[1339,1436,1124,1149]" italics="true" pageId="14" pageNumber="223">Matrona</emphasis>
</taxonomicName>
and
<taxonomicName id="DCDA4D38E03BD75CFF28E8A7FED6FB7D" authorityName="Selys" authorityYear="1853" box="[198,284,1159,1185]" class="Insecta" family="Calopterygidae" genus="Vestalis" kingdom="Animalia" order="Odonata" pageId="14" pageNumber="223" phylum="Arthropoda" rank="genus">
<emphasis id="29AEEAA9E03BD75CFF28E8A7FED6FB7D" box="[198,284,1159,1185]" italics="true" pageId="14" pageNumber="223">Vestalis</emphasis>
</taxonomicName>
/
<taxonomicName id="DCDA4D38E03BD75CFECDE8A7FE5FFB7D" authorityName="May" authorityYear="1935" box="[291,405,1159,1185]" class="Insecta" family="Calopterygidae" genus="Vestalaria" kingdom="Animalia" order="Odonata" pageId="14" pageNumber="223" phylum="Arthropoda" rank="genus">
<emphasis id="29AEEAA9E03BD75CFECDE8A7FE5FFB7D" box="[291,405,1159,1185]" italics="true" pageId="14" pageNumber="223">Vestalaria</emphasis>
</taxonomicName>
are known to be sure the keys accurately resolve all species. A distinctive form with strongly angulated postocular lobes described by
<bibRefCitation id="7F4B4B4AE03BD75CFDB0E88CFD3FFB19" author="Fraser, F. C." box="[606,757,1195,1222]" pageId="14" pageNumber="223" pagination="81 - 93" refId="ref23155" refString="Fraser, F. C. (1943) New oriental Odonata larvae. Proceedings of the Royal entomological society of London, Series B, 12, 81 - 93. https: // doi. org / 10.1111 / j. 1365 - 3113.1943. tb 00749. x" type="journal article" year="1943">Fraser (1943)</bibRefCitation>
as
<taxonomicName id="DCDA4D38E03BD75CFCF5E88BFBF4FB1A" authority="Hagen" authorityName="Hagen" box="[795,1086,1195,1222]" class="Insecta" family="Calopterygidae" genus="Caliphaea" kingdom="Animalia" order="Odonata" pageId="14" pageNumber="223" phylum="Arthropoda" rank="species" species="confusa">
<emphasis id="29AEEAA9E03BD75CFCF5E88BFC24FB19" box="[795,1006,1195,1221]" italics="true" pageId="14" pageNumber="223">Caliphaea confusa</emphasis>
Hagen
</taxonomicName>
may in fact be
<taxonomicName id="DCDA4D38E03BD75CFB05E88BFF1EFB35" authority="Selys" authorityName="Selys" authorityYear="1853" class="Insecta" family="Calopterygidae" genus="Echo" kingdom="Animalia" order="Odonata" pageId="14" pageNumber="223" phylum="Arthropoda" rank="species" species="margarita">
<emphasis id="29AEEAA9E03BD75CFB05E88BFA57FB19" box="[1259,1437,1195,1221]" italics="true" pageId="14" pageNumber="223">Echo margarita</emphasis>
Selys
</taxonomicName>
and another enigmatic larva collected in
<collectingCountry id="63CD762BE03BD75CFD4DE8EFFD2CFB35" box="[675,742,1231,1257]" name="Nepal" pageId="14" pageNumber="223">Nepal</collectingCountry>
cannot at present be assigned to a genus (
<bibRefCitation id="7F4B4B4AE03BD75CFB54E8EFFA5BFB35" author="Orr, A. G. &amp; Butler, S. G." box="[1210,1425,1231,1257]" pageId="14" pageNumber="223" pagination="155 - 168" refId="ref25814" refString="Orr, A. G. &amp; Butler, S. G. (2024) A riddle wrapped in a mystery inside an enigma: notes on an enigmatic larva collected in Nepal, with a reappraisal of Fraser's so called ' Caliphaea confusa ' larva from Meghalaya, India (Odonata: Calopterygidae). Odonatologica, 53, 155 - 168. https: // doi. org / 10.60024 / odon. v 53 i 1 - 2. a 8" type="journal article" year="2024">Orr &amp; Butler 2024</bibRefCitation>
). Almost all species in the family inhabit clear free flowing streams, typically in forest towards the equator and at lower elevations. Measurements (A, B, C, D) used in subsequent key descriptions are indicated in
<figureCitation id="83E12A3EE03BD75CFB0BE937FAFDFAED" box="[1253,1335,1303,1329]" captionStart="FIGURES 27–34. 27" captionStartId="7.[151,264,1845,1869]" captionTargetBox="[252,1326,159,1834]" captionTargetId="figure-15@7.[251,1336,153,1839]" captionTargetPageId="7" captionText="FIGURES 27–34. 27. Prementum of Calopterygidae showing measurements used in family key (based on Calopteryx japonica, redrawn from Miyakawa 1983). 28. Neurobasis chinensis, prementum and left labial palp (redrawn from Lieftinck 1965). 29. Matronoides cyaneipennis, prementum and labial palps (ventral view). 30. Matrona basilaris, prementum (dorsal view, drawn from Wang et al. 2017). 31. Atrocalopteryx atrata, prementum (dorsal view, redrawn from Miyakawa 1983). 32. Mnais tenuis, prementum (dorsal view, redrawn from Matsuki &amp; Lien 1977). 33. Echo uniformis, prementum and left labial palp dorsal view (redrawn from Lieftinck 1965). 34. Psolodesmus mandarinus, prementum (dorsal view, redrawn from Matsuki &amp; Lien 1978)." figureDoi="http://doi.org/10.5281/zenodo.13618333" httpUri="https://zenodo.org/record/13618333/files/figure.png" pageId="14" pageNumber="223">Fig. 27</figureCitation>
.
</paragraph>
<paragraph id="1B6536BBE03BD75CFF29E91CFD4CFA89" blockId="14.[151,1437,907,1366]" box="[199,646,1339,1366]" pageId="14" pageNumber="223">
Larva unknown:
<taxonomicName id="DCDA4D38E03BD75CFE68E91BFDE3FA89" authorityName="Asahina" authorityYear="1976" box="[390,553,1339,1365]" class="Insecta" family="Calopterygidae" genus="Noguchiphaea" kingdom="Animalia" order="Odonata" pageId="14" pageNumber="223" phylum="Arthropoda" rank="genus">
<emphasis id="29AEEAA9E03BD75CFE68E91BFDE3FA89" box="[390,553,1339,1365]" italics="true" pageId="14" pageNumber="223">Noguchiphaea</emphasis>
</taxonomicName>
(3 spp.)
</paragraph>
</subSubSection>
<subSubSection id="53C06530E026D741FF79ED04FE8CFEE2" box="[151,326,292,318]" pageId="19" pageNumber="228" type="multiple">
<paragraph id="1B6536BBE026D741FF79ED04FE8CFEE2" blockId="19.[151,326,292,318]" box="[151,326,292,318]" pageId="19" pageNumber="228">
<heading id="402D81D7E026D741FF79ED04FE8CFEE2" bold="true" box="[151,326,292,318]" fontSize="11" level="1" pageId="19" pageNumber="228" reason="1">
<emphasis id="29AEEAA9E026D741FF79ED04FE8CFEE2" bold="true" box="[151,326,292,318]" pageId="19" pageNumber="228">Key to genera</emphasis>
</heading>
</paragraph>
</subSubSection>
<subSubSection id="53C06530E026D741FF79ED49FA56F93E" pageId="19" pageNumber="228" type="key">
<key id="6F4B618AE026D741FF79ED49FA56F93E" pageId="19" pageNumber="228">
<keyStep id="A02E2E1EE026D741FF79ED49FA56FE0B" pageId="19" pageNumber="228">
<paragraph id="1B6536BBE026D741FF79ED49FA56FE47" blockId="19.[151,1437,361,1762]" pageId="19" pageNumber="228">
<keyLead id="A02B958EE026D741FF79ED49FA56FE47" pageId="19" pageNumber="228">
1 Premental cleft deep, diamond shaped or ovoid, ratio A/B at least 0.5 (
<figureCitation id="83E12A3EE026D741FC7BED49FB9AFEA2" box="[917,1104,361,383]" captionStart="FIGURES 27–34. 27" captionStartId="7.[151,264,1845,1869]" captionTargetBox="[252,1326,159,1834]" captionTargetId="figure-15@7.[251,1336,153,1839]" captionTargetPageId="7" captionText="FIGURES 27–34. 27. Prementum of Calopterygidae showing measurements used in family key (based on Calopteryx japonica, redrawn from Miyakawa 1983). 28. Neurobasis chinensis, prementum and left labial palp (redrawn from Lieftinck 1965). 29. Matronoides cyaneipennis, prementum and labial palps (ventral view). 30. Matrona basilaris, prementum (dorsal view, drawn from Wang et al. 2017). 31. Atrocalopteryx atrata, prementum (dorsal view, redrawn from Miyakawa 1983). 32. Mnais tenuis, prementum (dorsal view, redrawn from Matsuki &amp; Lien 1977). 33. Echo uniformis, prementum and left labial palp dorsal view (redrawn from Lieftinck 1965). 34. Psolodesmus mandarinus, prementum (dorsal view, redrawn from Matsuki &amp; Lien 1978)." figureDoi="http://doi.org/10.5281/zenodo.13618333" httpUri="https://zenodo.org/record/13618333/files/figure.png" pageId="19" pageNumber="228">Figs 28, 29, 30, 31</figureCitation>
); thinly built long legged species; lateral caudal gills at least half length of remainder of body (
<figureCitation id="83E12A3EE026D741FCC8EDA6FC5EFE40" box="[806,916,390,412]" captionStart="FIGURES 52–56. 52" captionStartId="10.[151,264,1941,1965]" captionTargetBox="[191,1388,163,1929]" captionTargetId="figure-21@10.[190,1397,151,1933]" captionTargetPageId="10" captionText="FIGURES 52–56. 52. Caliphaea angka, habitus (based on Yang et al. 2021). 53. Vestalis gracilis, habitus (redrawn from Rattanachan et al. 2022). 54. Neurobasis chinensis, habitus (original drawing). 55. Matrona basilaris, habitus (drawn from Wang et al. 2017). 56. Echo uniformis, habitus (redrawn from Lieftinck 1965)." figureDoi="http://doi.org/10.5281/zenodo.13618339" httpUri="https://zenodo.org/record/13618339/files/figure.png" pageId="19" pageNumber="228">Figs 54, 55</figureCitation>
)......................................... 2
</keyLead>
</paragraph>
<paragraph id="1B6536BBE026D741FF79ED84FA56FE0B" blockId="19.[151,1437,361,1762]" pageId="19" pageNumber="228">
<keyLead id="A02B958EE026D741FF79ED84FA56FE0B" pageId="19" pageNumber="228">
1’ Premental cleft, ratio A/B: 0.25–0.33 and tear-drop shaped (
<figureCitation id="83E12A3EE026D741FCF9ED84FC6DFE65" box="[791,935,420,442]" captionStart="FIGURES 27–34. 27" captionStartId="7.[151,264,1845,1869]" captionTargetBox="[252,1326,159,1834]" captionTargetId="figure-15@7.[251,1336,153,1839]" captionTargetPageId="7" captionText="FIGURES 27–34. 27. Prementum of Calopterygidae showing measurements used in family key (based on Calopteryx japonica, redrawn from Miyakawa 1983). 28. Neurobasis chinensis, prementum and left labial palp (redrawn from Lieftinck 1965). 29. Matronoides cyaneipennis, prementum and labial palps (ventral view). 30. Matrona basilaris, prementum (dorsal view, drawn from Wang et al. 2017). 31. Atrocalopteryx atrata, prementum (dorsal view, redrawn from Miyakawa 1983). 32. Mnais tenuis, prementum (dorsal view, redrawn from Matsuki &amp; Lien 1977). 33. Echo uniformis, prementum and left labial palp dorsal view (redrawn from Lieftinck 1965). 34. Psolodesmus mandarinus, prementum (dorsal view, redrawn from Matsuki &amp; Lien 1978)." figureDoi="http://doi.org/10.5281/zenodo.13618333" httpUri="https://zenodo.org/record/13618333/files/figure.png" pageId="19" pageNumber="228">Figs 32, 33, 34</figureCitation>
,
<figureCitation id="83E12A3EE026D741FC41ED83FBC5FE65" box="[943,1039,419,442]" captionStart="FIGURES 35–41. 35" captionStartId="8.[151,264,1776,1800]" captionTargetBox="[257,1332,159,1764]" captionTargetId="figure-21@8.[253,1335,153,1771]" captionTargetPageId="8" captionText="FIGURES 35–41. 35. Archineura incarnata, prementum (dorsal view, drawn from Yang et al. 2022). 36. Vestalis gracilis, prementum (dorsal view) and right antenna with supplementary antennomere between pedicel and scape (a) arrowed (redrawn from Rattanachan et al. 2022). 37. Vestalaria venusta: prementum (dorsal view) and right antenna (drawn from Wang et al. 2017). 38. Caliphaea angka: prementum (dorsal view) and right antenna (drawn from Yang et al. 2021). 39. Heliocypha biseriata, frontal view of labrum showing dorsal and ventral sections and ‘moustache’ of long, heavy setae on the latter (from Orr et al. 2024). 40. Heliocypha biseriata: details of labial palp in relation to prementum (a) dorsal view—base of movable hook dashed to reveal palpal lobe beneath and open base of premental median cleft arrowed, (b) frontal view, (c) inner view (from Orr et al. 2024). 41. Heliocypha biseriata, detail of fore tarsus with pectinate setae magnified (from Orr et al. 2024)." figureDoi="http://doi.org/10.5281/zenodo.13618335" httpUri="https://zenodo.org/record/13618335/files/figure.png" pageId="19" pageNumber="228">35, 36, 37</figureCitation>
), or cleft narrow and open anteriorly (
<figureCitation id="83E12A3EE026D741FA9BED84FF37FE0A" captionStart="FIGURES 35–41. 35" captionStartId="8.[151,264,1776,1800]" captionTargetBox="[257,1332,159,1764]" captionTargetId="figure-21@8.[253,1335,153,1771]" captionTargetPageId="8" captionText="FIGURES 35–41. 35. Archineura incarnata, prementum (dorsal view, drawn from Yang et al. 2022). 36. Vestalis gracilis, prementum (dorsal view) and right antenna with supplementary antennomere between pedicel and scape (a) arrowed (redrawn from Rattanachan et al. 2022). 37. Vestalaria venusta: prementum (dorsal view) and right antenna (drawn from Wang et al. 2017). 38. Caliphaea angka: prementum (dorsal view) and right antenna (drawn from Yang et al. 2021). 39. Heliocypha biseriata, frontal view of labrum showing dorsal and ventral sections and ‘moustache’ of long, heavy setae on the latter (from Orr et al. 2024). 40. Heliocypha biseriata: details of labial palp in relation to prementum (a) dorsal view—base of movable hook dashed to reveal palpal lobe beneath and open base of premental median cleft arrowed, (b) frontal view, (c) inner view (from Orr et al. 2024). 41. Heliocypha biseriata, detail of fore tarsus with pectinate setae magnified (from Orr et al. 2024)." figureDoi="http://doi.org/10.5281/zenodo.13618335" httpUri="https://zenodo.org/record/13618335/files/figure.png" pageId="19" pageNumber="228">Fig. 38</figureCitation>
); often moderately heavy in build; lateral caudal gills rarely more than half length of remainder of body............... 5
</keyLead>
</paragraph>
</keyStep>
<keyStep id="A02E2E1EE026D741FF79EDFEFA56FD97" pageId="19" pageNumber="228">
<paragraph id="1B6536BBE026D741FF79EDFEFA56FDCD" blockId="19.[151,1437,361,1762]" pageId="19" pageNumber="228">
<keyLead id="A02B958EE026D741FF79EDFEFA56FDCD" pageId="19" pageNumber="228">
2 (1) Prementum very long and narrow, ratio B/D: 2.2–2.8 and premental cleft deep, ratio A/B: ca 0.6 (
<figureCitation id="83E12A3EE026D741FB97EDFEFB2DFE2F" box="[1145,1255,478,500]" captionStart="FIGURES 27–34. 27" captionStartId="7.[151,264,1845,1869]" captionTargetBox="[252,1326,159,1834]" captionTargetId="figure-15@7.[251,1336,153,1839]" captionTargetPageId="7" captionText="FIGURES 27–34. 27. Prementum of Calopterygidae showing measurements used in family key (based on Calopteryx japonica, redrawn from Miyakawa 1983). 28. Neurobasis chinensis, prementum and left labial palp (redrawn from Lieftinck 1965). 29. Matronoides cyaneipennis, prementum and labial palps (ventral view). 30. Matrona basilaris, prementum (dorsal view, drawn from Wang et al. 2017). 31. Atrocalopteryx atrata, prementum (dorsal view, redrawn from Miyakawa 1983). 32. Mnais tenuis, prementum (dorsal view, redrawn from Matsuki &amp; Lien 1977). 33. Echo uniformis, prementum and left labial palp dorsal view (redrawn from Lieftinck 1965). 34. Psolodesmus mandarinus, prementum (dorsal view, redrawn from Matsuki &amp; Lien 1978)." figureDoi="http://doi.org/10.5281/zenodo.13618333" httpUri="https://zenodo.org/record/13618333/files/figure.png" pageId="19" pageNumber="228">Figs 28, 29</figureCitation>
); single strong seta on each lobe of prementum.............................................................................. 3
</keyLead>
</paragraph>
<paragraph id="1B6536BBE026D741FF79EE39FA56FD97" blockId="19.[151,1437,361,1762]" pageId="19" pageNumber="228">
<keyLead id="A02B958EE026D741FF79EE39FA56FD97" pageId="19" pageNumber="228">
2’ Prementum broader and shorter, ratio B/D: 1.6-2 and premental cleft depth ratio A/B ca. 0.5 (
<figureCitation id="83E12A3EE026D741FB8FEE39FB05FDF2" box="[1121,1231,537,559]" captionStart="FIGURES 27–34. 27" captionStartId="7.[151,264,1845,1869]" captionTargetBox="[252,1326,159,1834]" captionTargetId="figure-15@7.[251,1336,153,1839]" captionTargetPageId="7" captionText="FIGURES 27–34. 27. Prementum of Calopterygidae showing measurements used in family key (based on Calopteryx japonica, redrawn from Miyakawa 1983). 28. Neurobasis chinensis, prementum and left labial palp (redrawn from Lieftinck 1965). 29. Matronoides cyaneipennis, prementum and labial palps (ventral view). 30. Matrona basilaris, prementum (dorsal view, drawn from Wang et al. 2017). 31. Atrocalopteryx atrata, prementum (dorsal view, redrawn from Miyakawa 1983). 32. Mnais tenuis, prementum (dorsal view, redrawn from Matsuki &amp; Lien 1977). 33. Echo uniformis, prementum and left labial palp dorsal view (redrawn from Lieftinck 1965). 34. Psolodesmus mandarinus, prementum (dorsal view, redrawn from Matsuki &amp; Lien 1978)." figureDoi="http://doi.org/10.5281/zenodo.13618333" httpUri="https://zenodo.org/record/13618333/files/figure.png" pageId="19" pageNumber="228">Figs 30, 31</figureCitation>
); two strong setae on each lobe of prementum................................................................................ 4
</keyLead>
</paragraph>
</keyStep>
<keyStep id="A02E2E1EE026D741FF79EE74FA56FD20" pageId="19" pageNumber="228">
<paragraph id="1B6536BBE026D741FF79EE74FA56FD7F" blockId="19.[151,1437,361,1762]" pageId="19" pageNumber="228">
<keyLead id="A02B958EE026D741FF79EE74FA56FD7F" pageId="19" pageNumber="228">
3 Lateral caudal gills about half length of rest of body; middle gill less than half length of laterals; hindlegs extended just reaching tip of S10. large species (body length excluding antennae and caudal gills
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).............
<taxonomicName id="DCDA4D38E026D741FB76EE50FA56FD5A" box="[1176,1436,624,646]" class="Insecta" family="Calopterygidae" genus="Matronoides" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="19" pageNumber="228" phylum="Arthropoda" rank="species" species="cyaneipennis">
<emphasis id="29AEEAA9E026D741FB76EE50FA56FD5A" bold="true" box="[1176,1436,624,646]" italics="true" pageId="19" pageNumber="228">Matronoides cyaneipennis</emphasis>
</taxonomicName>
[endemic to
<collectingCountry id="63CD762BE026D741FB4CEEAEFAEDFD78" box="[1186,1319,654,676]" name="Malaysia" pageId="19" pageNumber="228">North Borneo</collectingCountry>
over
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<paragraph id="1B6536BBE026D741FF79EE8CFA56FD20" blockId="19.[151,1437,361,1762]" pageId="19" pageNumber="228">
<keyLead id="A02B958EE026D741FF79EE8CFA56FD20" pageId="19" pageNumber="228">
3’ Lateral gills about 0.7 length of rest of body; middle gill more than half length of laterals; very slender in build with long spindly legs; hindlegs extended reaching well beyond S10 of abdomen (
<figureCitation id="83E12A3EE026D741FC71EEE9FC22FD03" box="[927,1000,713,735]" captionStart="FIGURES 52–56. 52" captionStartId="10.[151,264,1941,1965]" captionTargetBox="[191,1388,163,1929]" captionTargetId="figure-21@10.[190,1397,151,1933]" captionTargetPageId="10" captionText="FIGURES 52–56. 52. Caliphaea angka, habitus (based on Yang et al. 2021). 53. Vestalis gracilis, habitus (redrawn from Rattanachan et al. 2022). 54. Neurobasis chinensis, habitus (original drawing). 55. Matrona basilaris, habitus (drawn from Wang et al. 2017). 56. Echo uniformis, habitus (redrawn from Lieftinck 1965)." figureDoi="http://doi.org/10.5281/zenodo.13618339" httpUri="https://zenodo.org/record/13618339/files/figure.png" pageId="19" pageNumber="228">Fig. 54</figureCitation>
); widespread in clear streams throughout the region.......................................................................................
<taxonomicName id="DCDA4D38E026D741FAC2EEC6FA56FD20" baseAuthorityName="Orr &amp; Hamalainen" baseAuthorityYear="2007" box="[1324,1436,742,764]" class="Insecta" family="Calopterygidae" genus="Neurobasis" kingdom="Animalia" order="Odonata" pageId="19" pageNumber="228" phylum="Arthropoda" rank="genus">
<emphasis id="29AEEAA9E026D741FAC2EEC6FA56FD20" bold="true" box="[1324,1436,742,764]" italics="true" pageId="19" pageNumber="228">Neurobasis</emphasis>
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</keyStep>
<keyStep id="A02E2E1EE026D741FF79EF24FA57FC70" pageId="19" pageNumber="228">
<paragraph id="1B6536BBE026D741FF79EF24FA57FC88" blockId="19.[151,1437,361,1762]" pageId="19" pageNumber="228">
<keyLead id="A02B958EE026D741FF79EF24FA57FC88" pageId="19" pageNumber="228">
4 (2’) Prementum (
<figureCitation id="83E12A3EE026D741FEB0EF24FE6EFCC5" box="[350,420,772,794]" captionStart="FIGURES 27–34. 27" captionStartId="7.[151,264,1845,1869]" captionTargetBox="[252,1326,159,1834]" captionTargetId="figure-15@7.[251,1336,153,1839]" captionTargetPageId="7" captionText="FIGURES 27–34. 27. Prementum of Calopterygidae showing measurements used in family key (based on Calopteryx japonica, redrawn from Miyakawa 1983). 28. Neurobasis chinensis, prementum and left labial palp (redrawn from Lieftinck 1965). 29. Matronoides cyaneipennis, prementum and labial palps (ventral view). 30. Matrona basilaris, prementum (dorsal view, drawn from Wang et al. 2017). 31. Atrocalopteryx atrata, prementum (dorsal view, redrawn from Miyakawa 1983). 32. Mnais tenuis, prementum (dorsal view, redrawn from Matsuki &amp; Lien 1977). 33. Echo uniformis, prementum and left labial palp dorsal view (redrawn from Lieftinck 1965). 34. Psolodesmus mandarinus, prementum (dorsal view, redrawn from Matsuki &amp; Lien 1978)." figureDoi="http://doi.org/10.5281/zenodo.13618333" httpUri="https://zenodo.org/record/13618333/files/figure.png" pageId="19" pageNumber="228">Fig. 30</figureCitation>
) elongate, ratio B/D: 1.8–1.9 and cleft boad, ratio C/D: 0.3; anterior lobes of prementum long and narrow (total length from base: width at midpoint = 9.5); abdomen moderately broad; hind femur reaching S6; lateral caudal gill about half length of remainder of body (
<figureCitation id="83E12A3EE026D741FDC9EF1EFDB8FC8F" box="[551,626,830,852]" captionStart="FIGURES 42–51. 42" captionStartId="9.[151,264,1592,1616]" captionTargetBox="[165,1413,192,1557]" captionTargetId="figure-15@9.[151,1435,181,1568]" captionTargetPageId="9" captionText="FIGURES 42–51. 42. Archineura incarnata: caudal gills; left lateral above, median below (drawn from Yang et al. 2022). 43. Echo modesta: caudal gills; left lateral above, median below, with baso-dorsal shield arrowed (original illustration). 44. Psolodesmus mandarinus, dorsal view of caudal gills, with baso-dorsal shield of median gill arrowed (redrawn from Matsuki &amp; Lien 1978). 45. Mnais andersoni, left lateral caudal gill with upper margin of median gill visible; baso-dorsal shield of median gill arrowed (original illustration). 46. Mnais gregoryi: caudal gills; left lateral above, median below, with baso-dorsal shield arrowed (drawn from Yang et al. 2021). 47. Caliphaea angka: caudal gills; left lateral above, median below (drawn from Yang et al. 2021). 48. Matrona basilaris: caudal gills; left lateral above, median below (drawn from Wang et al. 2017). 49. Vestalaria venusta: caudal gills; left lateral above, median below (drawn from Wang et al. 2017). 50. Vestalis amoena group: caudal gills; left lateral above, median below (redrawn from Lieftinck 1965). 51. Vestalis melania: caudal gills; left lateral above, median below (redrawn from Guadalquiver et al. 2022)." figureDoi="http://doi.org/10.5281/zenodo.13618337" httpUri="https://zenodo.org/record/13618337/files/figure.png" pageId="19" pageNumber="228">Figs 48</figureCitation>
,
<figureCitation id="83E12A3EE026D741FD92EF1EFD5DFC88" box="[636,663,830,852]" captionStart="FIGURES 52–56. 52" captionStartId="10.[151,264,1941,1965]" captionTargetBox="[191,1388,163,1929]" captionTargetId="figure-21@10.[190,1397,151,1933]" captionTargetPageId="10" captionText="FIGURES 52–56. 52. Caliphaea angka, habitus (based on Yang et al. 2021). 53. Vestalis gracilis, habitus (redrawn from Rattanachan et al. 2022). 54. Neurobasis chinensis, habitus (original drawing). 55. Matrona basilaris, habitus (drawn from Wang et al. 2017). 56. Echo uniformis, habitus (redrawn from Lieftinck 1965)." figureDoi="http://doi.org/10.5281/zenodo.13618339" httpUri="https://zenodo.org/record/13618339/files/figure.png" pageId="19" pageNumber="228">55</figureCitation>
).........................................................
<taxonomicName id="DCDA4D38E026D741FAA8EF1FFA57FC88" authorityName="Selys" authorityYear="1853" box="[1350,1437,831,852]" class="Insecta" family="Calopterygidae" genus="Matrona" kingdom="Animalia" order="Odonata" pageId="19" pageNumber="228" phylum="Arthropoda" rank="genus">
<emphasis id="29AEEAA9E026D741FAA8EF1FFA57FC88" bold="true" box="[1350,1437,831,852]" italics="true" pageId="19" pageNumber="228">Matrona</emphasis>
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</paragraph>
<paragraph id="1B6536BBE026D741FF79EF7CFA57FC70" blockId="19.[151,1437,361,1762]" pageId="19" pageNumber="228">
<keyLead id="A02B958EE026D741FF79EF7CFA57FC70" pageId="19" pageNumber="228">
4’ Prementum (
<figureCitation id="83E12A3EE026D741FEB3EF7CFE68FCAD" box="[349,418,860,882]" captionStart="FIGURES 27–34. 27" captionStartId="7.[151,264,1845,1869]" captionTargetBox="[252,1326,159,1834]" captionTargetId="figure-15@7.[251,1336,153,1839]" captionTargetPageId="7" captionText="FIGURES 27–34. 27. Prementum of Calopterygidae showing measurements used in family key (based on Calopteryx japonica, redrawn from Miyakawa 1983). 28. Neurobasis chinensis, prementum and left labial palp (redrawn from Lieftinck 1965). 29. Matronoides cyaneipennis, prementum and labial palps (ventral view). 30. Matrona basilaris, prementum (dorsal view, drawn from Wang et al. 2017). 31. Atrocalopteryx atrata, prementum (dorsal view, redrawn from Miyakawa 1983). 32. Mnais tenuis, prementum (dorsal view, redrawn from Matsuki &amp; Lien 1977). 33. Echo uniformis, prementum and left labial palp dorsal view (redrawn from Lieftinck 1965). 34. Psolodesmus mandarinus, prementum (dorsal view, redrawn from Matsuki &amp; Lien 1978)." figureDoi="http://doi.org/10.5281/zenodo.13618333" httpUri="https://zenodo.org/record/13618333/files/figure.png" pageId="19" pageNumber="228">Fig. 31</figureCitation>
) shorter (ratio B/D: 1.65) and more solid; cleft narrow, ratio C/D: 0.24 anterior lobes of prementum shorter and broader (total length from base: width at midpoint = 5.6); hind femur at most reaching S5; lateral caudal gill about 0.6 length of remainder of body..................................................................
<taxonomicName id="DCDA4D38E026D741FAE4EFB6FA57FC70" authorityName="Dumont, Vanfleteren, De Jonckheere &amp; Weekers" authorityYear="2005" box="[1290,1437,918,940]" class="Insecta" family="Calopterygidae" genus="Atrocalopteryx" kingdom="Animalia" order="Odonata" pageId="19" pageNumber="228" phylum="Arthropoda" rank="genus">
<emphasis id="29AEEAA9E026D741FAE4EFB6FA57FC70" bold="true" box="[1290,1437,918,940]" italics="true" pageId="19" pageNumber="228">Atrocalopteryx</emphasis>
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</paragraph>
</keyStep>
<keyStep id="A02E2E1EE026D741FF79EF93FA56FBFE" pageId="19" pageNumber="228">
<paragraph id="1B6536BBE026D741FF79EF93FA57FBD8" blockId="19.[151,1437,361,1762]" pageId="19" pageNumber="228">
<keyLead id="A02B958EE026D741FF79EF93FA57FBD8" pageId="19" pageNumber="228">
5 (1’) Lateral caudal gills, long, narrow, clearly triquetral, slightly more than half length of remainder of body and armed with wellspaced tubercles along margins, with posterior tubercles long and finger-like (
<figureCitation id="83E12A3EE026D741FC34EFF1FBE9FC3A" box="[986,1059,977,999]" captionStart="FIGURES 42–51. 42" captionStartId="9.[151,264,1592,1616]" captionTargetBox="[165,1413,192,1557]" captionTargetId="figure-15@9.[151,1435,181,1568]" captionTargetPageId="9" captionText="FIGURES 42–51. 42. Archineura incarnata: caudal gills; left lateral above, median below (drawn from Yang et al. 2022). 43. Echo modesta: caudal gills; left lateral above, median below, with baso-dorsal shield arrowed (original illustration). 44. Psolodesmus mandarinus, dorsal view of caudal gills, with baso-dorsal shield of median gill arrowed (redrawn from Matsuki &amp; Lien 1978). 45. Mnais andersoni, left lateral caudal gill with upper margin of median gill visible; baso-dorsal shield of median gill arrowed (original illustration). 46. Mnais gregoryi: caudal gills; left lateral above, median below, with baso-dorsal shield arrowed (drawn from Yang et al. 2021). 47. Caliphaea angka: caudal gills; left lateral above, median below (drawn from Yang et al. 2021). 48. Matrona basilaris: caudal gills; left lateral above, median below (drawn from Wang et al. 2017). 49. Vestalaria venusta: caudal gills; left lateral above, median below (drawn from Wang et al. 2017). 50. Vestalis amoena group: caudal gills; left lateral above, median below (redrawn from Lieftinck 1965). 51. Vestalis melania: caudal gills; left lateral above, median below (redrawn from Guadalquiver et al. 2022)." figureDoi="http://doi.org/10.5281/zenodo.13618337" httpUri="https://zenodo.org/record/13618337/files/figure.png" pageId="19" pageNumber="228">Fig. 42</figureCitation>
); premental cleft short (A/B: 0.24); no inner strong setae on relatively short lobes of prementum (
<figureCitation id="83E12A3EE026D741FCFDEFCEFC91FBD8" box="[787,859,1006,1028]" captionStart="FIGURES 35–41. 35" captionStartId="8.[151,264,1776,1800]" captionTargetBox="[257,1332,159,1764]" captionTargetId="figure-21@8.[253,1335,153,1771]" captionTargetPageId="8" captionText="FIGURES 35–41. 35. Archineura incarnata, prementum (dorsal view, drawn from Yang et al. 2022). 36. Vestalis gracilis, prementum (dorsal view) and right antenna with supplementary antennomere between pedicel and scape (a) arrowed (redrawn from Rattanachan et al. 2022). 37. Vestalaria venusta: prementum (dorsal view) and right antenna (drawn from Wang et al. 2017). 38. Caliphaea angka: prementum (dorsal view) and right antenna (drawn from Yang et al. 2021). 39. Heliocypha biseriata, frontal view of labrum showing dorsal and ventral sections and ‘moustache’ of long, heavy setae on the latter (from Orr et al. 2024). 40. Heliocypha biseriata: details of labial palp in relation to prementum (a) dorsal view—base of movable hook dashed to reveal palpal lobe beneath and open base of premental median cleft arrowed, (b) frontal view, (c) inner view (from Orr et al. 2024). 41. Heliocypha biseriata, detail of fore tarsus with pectinate setae magnified (from Orr et al. 2024)." figureDoi="http://doi.org/10.5281/zenodo.13618335" httpUri="https://zenodo.org/record/13618335/files/figure.png" pageId="19" pageNumber="228">Fig. 35</figureCitation>
).......................................
<taxonomicName id="DCDA4D38E026D741FAC4EFCEFA57FBD8" authorityName="Kirby" authorityYear="1894" box="[1322,1437,1006,1028]" class="Insecta" family="Calopterygidae" genus="Archineura" kingdom="Animalia" order="Odonata" pageId="19" pageNumber="228" phylum="Arthropoda" rank="genus">
<emphasis id="29AEEAA9E026D741FAC4EFCEFA57FBD8" bold="true" box="[1322,1437,1006,1028]" italics="true" pageId="19" pageNumber="228">Archineura</emphasis>
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<paragraph id="1B6536BBE026D741FF79E82BFA56FBFE" blockId="19.[151,1437,361,1762]" box="[151,1436,1035,1058]" pageId="19" pageNumber="228">
<keyLead id="A02B958EE026D741FF79E82BFA56FBFE" box="[151,1436,1035,1058]" pageId="19" pageNumber="228">5’ Lateral caudal gills without tubercles, or if tubercles present then less than one third length of remainder of body.......... 6</keyLead>
</paragraph>
</keyStep>
<keyStep id="A02E2E1EE026D741FF79E809FA56FBA5" pageId="19" pageNumber="228">
<paragraph id="1B6536BBE026D741FF79E809FA56FB80" blockId="19.[151,1437,361,1762]" pageId="19" pageNumber="228">
<keyLead id="A02B958EE026D741FF79E809FA56FB80" pageId="19" pageNumber="228">
6 (5’) Lateral caudal gills laminate, foliate and at least one third length of remainder of body (
<figureCitation id="83E12A3EE026D741FBDFE809FBB3FBE3" box="[1073,1145,1065,1087]" captionStart="FIGURES 42–51. 42" captionStartId="9.[151,264,1592,1616]" captionTargetBox="[165,1413,192,1557]" captionTargetId="figure-15@9.[151,1435,181,1568]" captionTargetPageId="9" captionText="FIGURES 42–51. 42. Archineura incarnata: caudal gills; left lateral above, median below (drawn from Yang et al. 2022). 43. Echo modesta: caudal gills; left lateral above, median below, with baso-dorsal shield arrowed (original illustration). 44. Psolodesmus mandarinus, dorsal view of caudal gills, with baso-dorsal shield of median gill arrowed (redrawn from Matsuki &amp; Lien 1978). 45. Mnais andersoni, left lateral caudal gill with upper margin of median gill visible; baso-dorsal shield of median gill arrowed (original illustration). 46. Mnais gregoryi: caudal gills; left lateral above, median below, with baso-dorsal shield arrowed (drawn from Yang et al. 2021). 47. Caliphaea angka: caudal gills; left lateral above, median below (drawn from Yang et al. 2021). 48. Matrona basilaris: caudal gills; left lateral above, median below (drawn from Wang et al. 2017). 49. Vestalaria venusta: caudal gills; left lateral above, median below (drawn from Wang et al. 2017). 50. Vestalis amoena group: caudal gills; left lateral above, median below (redrawn from Lieftinck 1965). 51. Vestalis melania: caudal gills; left lateral above, median below (redrawn from Guadalquiver et al. 2022)." figureDoi="http://doi.org/10.5281/zenodo.13618337" httpUri="https://zenodo.org/record/13618337/files/figure.png" pageId="19" pageNumber="228">Fig. 50</figureCitation>
); prementum ratio B/D: 1.37– 1.6. (
<figureCitation id="83E12A3EE026D741FEFDE866FE4AFB87" box="[275,384,1094,1116]" captionStart="FIGURES 35–41. 35" captionStartId="8.[151,264,1776,1800]" captionTargetBox="[257,1332,159,1764]" captionTargetId="figure-21@8.[253,1335,153,1771]" captionTargetPageId="8" captionText="FIGURES 35–41. 35. Archineura incarnata, prementum (dorsal view, drawn from Yang et al. 2022). 36. Vestalis gracilis, prementum (dorsal view) and right antenna with supplementary antennomere between pedicel and scape (a) arrowed (redrawn from Rattanachan et al. 2022). 37. Vestalaria venusta: prementum (dorsal view) and right antenna (drawn from Wang et al. 2017). 38. Caliphaea angka: prementum (dorsal view) and right antenna (drawn from Yang et al. 2021). 39. Heliocypha biseriata, frontal view of labrum showing dorsal and ventral sections and ‘moustache’ of long, heavy setae on the latter (from Orr et al. 2024). 40. Heliocypha biseriata: details of labial palp in relation to prementum (a) dorsal view—base of movable hook dashed to reveal palpal lobe beneath and open base of premental median cleft arrowed, (b) frontal view, (c) inner view (from Orr et al. 2024). 41. Heliocypha biseriata, detail of fore tarsus with pectinate setae magnified (from Orr et al. 2024)." figureDoi="http://doi.org/10.5281/zenodo.13618335" httpUri="https://zenodo.org/record/13618335/files/figure.png" pageId="19" pageNumber="228">Figs 36, 37</figureCitation>
)..................................................................................... 7
</keyLead>
</paragraph>
<paragraph id="1B6536BBE026D741FF79E844FA56FBA5" blockId="19.[151,1437,361,1762]" box="[151,1436,1124,1146]" pageId="19" pageNumber="228">
<keyLead id="A02B958EE026D741FF79E844FA56FBA5" box="[151,1436,1124,1146]" pageId="19" pageNumber="228">6’ Lateral caudal gills less than one third length of remainder of body; prementum ratio B/D: 1.1–1.3...................... 8</keyLead>
</paragraph>
</keyStep>
<keyStep id="A02E2E1EE026D741FF79E8A1FA57FA9A" pageId="19" pageNumber="228">
<paragraph id="1B6536BBE026D741FF79E8A1FA57FB33" blockId="19.[151,1437,361,1762]" pageId="19" pageNumber="228">
<keyLead id="A02B958EE026D741FF79E8A1FA57FB33" pageId="19" pageNumber="228">
7 (6) Two strong spines on each lobe of prementum (
<figureCitation id="83E12A3EE026D741FD4AE8A1FD21FB4B" box="[676,747,1153,1175]" captionStart="FIGURES 35–41. 35" captionStartId="8.[151,264,1776,1800]" captionTargetBox="[257,1332,159,1764]" captionTargetId="figure-21@8.[253,1335,153,1771]" captionTargetPageId="8" captionText="FIGURES 35–41. 35. Archineura incarnata, prementum (dorsal view, drawn from Yang et al. 2022). 36. Vestalis gracilis, prementum (dorsal view) and right antenna with supplementary antennomere between pedicel and scape (a) arrowed (redrawn from Rattanachan et al. 2022). 37. Vestalaria venusta: prementum (dorsal view) and right antenna (drawn from Wang et al. 2017). 38. Caliphaea angka: prementum (dorsal view) and right antenna (drawn from Yang et al. 2021). 39. Heliocypha biseriata, frontal view of labrum showing dorsal and ventral sections and ‘moustache’ of long, heavy setae on the latter (from Orr et al. 2024). 40. Heliocypha biseriata: details of labial palp in relation to prementum (a) dorsal view—base of movable hook dashed to reveal palpal lobe beneath and open base of premental median cleft arrowed, (b) frontal view, (c) inner view (from Orr et al. 2024). 41. Heliocypha biseriata, detail of fore tarsus with pectinate setae magnified (from Orr et al. 2024)." figureDoi="http://doi.org/10.5281/zenodo.13618335" httpUri="https://zenodo.org/record/13618335/files/figure.png" pageId="19" pageNumber="228">Fig. 36</figureCitation>
); depth of premental cleft ratio A/B 0.33 or greater; prementum abruptly expanded anteriorly; lateral gills 0.33–0.36 length of remainder of body (
<figureCitation id="83E12A3EE026D741FC74E8BEFC28FB68" box="[922,994,1182,1204]" captionStart="FIGURES 52–56. 52" captionStartId="10.[151,264,1941,1965]" captionTargetBox="[191,1388,163,1929]" captionTargetId="figure-21@10.[190,1397,151,1933]" captionTargetPageId="10" captionText="FIGURES 52–56. 52. Caliphaea angka, habitus (based on Yang et al. 2021). 53. Vestalis gracilis, habitus (redrawn from Rattanachan et al. 2022). 54. Neurobasis chinensis, habitus (original drawing). 55. Matrona basilaris, habitus (drawn from Wang et al. 2017). 56. Echo uniformis, habitus (redrawn from Lieftinck 1965)." figureDoi="http://doi.org/10.5281/zenodo.13618339" httpUri="https://zenodo.org/record/13618339/files/figure.png" pageId="19" pageNumber="228">Fig. 53</figureCitation>
); antennal scape ca 0.8 width of head; antenna with very short segment between pedicel and scape, 8 segmented (
<figureCitation id="83E12A3EE026D741FC8EE89CFC6CFB0E" box="[864,934,1212,1234]" captionStart="FIGURES 35–41. 35" captionStartId="8.[151,264,1776,1800]" captionTargetBox="[257,1332,159,1764]" captionTargetId="figure-21@8.[253,1335,153,1771]" captionTargetPageId="8" captionText="FIGURES 35–41. 35. Archineura incarnata, prementum (dorsal view, drawn from Yang et al. 2022). 36. Vestalis gracilis, prementum (dorsal view) and right antenna with supplementary antennomere between pedicel and scape (a) arrowed (redrawn from Rattanachan et al. 2022). 37. Vestalaria venusta: prementum (dorsal view) and right antenna (drawn from Wang et al. 2017). 38. Caliphaea angka: prementum (dorsal view) and right antenna (drawn from Yang et al. 2021). 39. Heliocypha biseriata, frontal view of labrum showing dorsal and ventral sections and ‘moustache’ of long, heavy setae on the latter (from Orr et al. 2024). 40. Heliocypha biseriata: details of labial palp in relation to prementum (a) dorsal view—base of movable hook dashed to reveal palpal lobe beneath and open base of premental median cleft arrowed, (b) frontal view, (c) inner view (from Orr et al. 2024). 41. Heliocypha biseriata, detail of fore tarsus with pectinate setae magnified (from Orr et al. 2024)." figureDoi="http://doi.org/10.5281/zenodo.13618335" httpUri="https://zenodo.org/record/13618335/files/figure.png" pageId="19" pageNumber="228">Fig. 36</figureCitation>
)..................................
<taxonomicName id="DCDA4D38E026D741FABCE89BFA57FB0D" authorityName="Selys" authorityYear="1853" box="[1362,1437,1211,1233]" class="Insecta" family="Calopterygidae" genus="Vestalis" kingdom="Animalia" order="Odonata" pageId="19" pageNumber="228" phylum="Arthropoda" rank="genus">
<emphasis id="29AEEAA9E026D741FABCE89BFA57FB0D" bold="true" box="[1362,1437,1211,1233]" italics="true" pageId="19" pageNumber="228">Vestalis</emphasis>
</taxonomicName>
[widespread west of Wallace’s line]
</keyLead>
</paragraph>
<paragraph id="1B6536BBE026D741FF79E8D6FA57FA9A" blockId="19.[151,1437,361,1762]" pageId="19" pageNumber="228">
<keyLead id="A02B958EE026D741FF79E8D6FA57FA9A" pageId="19" pageNumber="228">
7’ One strong spine on each lobe of prementum (
<figureCitation id="83E12A3EE026D741FD79E8D6FD14FAD7" box="[663,734,1270,1292]" captionStart="FIGURES 35–41. 35" captionStartId="8.[151,264,1776,1800]" captionTargetBox="[257,1332,159,1764]" captionTargetId="figure-21@8.[253,1335,153,1771]" captionTargetPageId="8" captionText="FIGURES 35–41. 35. Archineura incarnata, prementum (dorsal view, drawn from Yang et al. 2022). 36. Vestalis gracilis, prementum (dorsal view) and right antenna with supplementary antennomere between pedicel and scape (a) arrowed (redrawn from Rattanachan et al. 2022). 37. Vestalaria venusta: prementum (dorsal view) and right antenna (drawn from Wang et al. 2017). 38. Caliphaea angka: prementum (dorsal view) and right antenna (drawn from Yang et al. 2021). 39. Heliocypha biseriata, frontal view of labrum showing dorsal and ventral sections and ‘moustache’ of long, heavy setae on the latter (from Orr et al. 2024). 40. Heliocypha biseriata: details of labial palp in relation to prementum (a) dorsal view—base of movable hook dashed to reveal palpal lobe beneath and open base of premental median cleft arrowed, (b) frontal view, (c) inner view (from Orr et al. 2024). 41. Heliocypha biseriata, detail of fore tarsus with pectinate setae magnified (from Orr et al. 2024)." figureDoi="http://doi.org/10.5281/zenodo.13618335" httpUri="https://zenodo.org/record/13618335/files/figure.png" pageId="19" pageNumber="228">Fig. 37</figureCitation>
); premental cleft ratio A/B less than 0.33; prementum gradually expanded anteriorly; lateral gills 0.46–0.5 length remainder of body; antennal scape ca width of head or longer, antenna lacking short segment between pedicel and scape, 7 segmented.....................................................
<taxonomicName id="DCDA4D38E026D741FAD4E910FA57FA9A" authorityName="May" authorityYear="1935" box="[1338,1437,1328,1350]" class="Insecta" family="Calopterygidae" genus="Vestalaria" kingdom="Animalia" order="Odonata" pageId="19" pageNumber="228" phylum="Arthropoda" rank="genus">
<emphasis id="29AEEAA9E026D741FAD4E910FA57FA9A" bold="true" box="[1338,1437,1328,1350]" italics="true" pageId="19" pageNumber="228">Vestalaria</emphasis>
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</paragraph>
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<keyStep id="A02E2E1EE026D741FF79E96EFA56FA05" pageId="19" pageNumber="228">
<paragraph id="1B6536BBE026D741FF79E96EFA56FA42" blockId="19.[151,1437,361,1762]" pageId="19" pageNumber="228">
<keyLead id="A02B958EE026D741FF79E96EFA56FA42" pageId="19" pageNumber="228">
8 (6’) Premental cleft long and narrow, lobes of prementum not meeting anteriorly; ratio A/B: 0.33 (
<figureCitation id="83E12A3EE026D741FB8DE96EFB61FABF" box="[1123,1195,1358,1380]" captionStart="FIGURES 35–41. 35" captionStartId="8.[151,264,1776,1800]" captionTargetBox="[257,1332,159,1764]" captionTargetId="figure-21@8.[253,1335,153,1771]" captionTargetPageId="8" captionText="FIGURES 35–41. 35. Archineura incarnata, prementum (dorsal view, drawn from Yang et al. 2022). 36. Vestalis gracilis, prementum (dorsal view) and right antenna with supplementary antennomere between pedicel and scape (a) arrowed (redrawn from Rattanachan et al. 2022). 37. Vestalaria venusta: prementum (dorsal view) and right antenna (drawn from Wang et al. 2017). 38. Caliphaea angka: prementum (dorsal view) and right antenna (drawn from Yang et al. 2021). 39. Heliocypha biseriata, frontal view of labrum showing dorsal and ventral sections and ‘moustache’ of long, heavy setae on the latter (from Orr et al. 2024). 40. Heliocypha biseriata: details of labial palp in relation to prementum (a) dorsal view—base of movable hook dashed to reveal palpal lobe beneath and open base of premental median cleft arrowed, (b) frontal view, (c) inner view (from Orr et al. 2024). 41. Heliocypha biseriata, detail of fore tarsus with pectinate setae magnified (from Orr et al. 2024)." figureDoi="http://doi.org/10.5281/zenodo.13618335" httpUri="https://zenodo.org/record/13618335/files/figure.png" pageId="19" pageNumber="228">Fig. 38</figureCitation>
); lateral caudal gills very short, no more than 0.25 length of remainder of body (
<figureCitation id="83E12A3EE026D741FD05E94CFCF9FA5D" box="[747,819,1387,1410]" captionStart="FIGURES 52–56. 52" captionStartId="10.[151,264,1941,1965]" captionTargetBox="[191,1388,163,1929]" captionTargetId="figure-21@10.[190,1397,151,1933]" captionTargetPageId="10" captionText="FIGURES 52–56. 52. Caliphaea angka, habitus (based on Yang et al. 2021). 53. Vestalis gracilis, habitus (redrawn from Rattanachan et al. 2022). 54. Neurobasis chinensis, habitus (original drawing). 55. Matrona basilaris, habitus (drawn from Wang et al. 2017). 56. Echo uniformis, habitus (redrawn from Lieftinck 1965)." figureDoi="http://doi.org/10.5281/zenodo.13618339" httpUri="https://zenodo.org/record/13618339/files/figure.png" pageId="19" pageNumber="228">Fig. 52</figureCitation>
); ovoid, margins smooth, with outer central rib bearing 5–7 well defined denticles (
<figureCitation id="83E12A3EE026D741FE7CE9A9FE10FA42" box="[402,474,1417,1439]" captionStart="FIGURES 42–51. 42" captionStartId="9.[151,264,1592,1616]" captionTargetBox="[165,1413,192,1557]" captionTargetId="figure-15@9.[151,1435,181,1568]" captionTargetPageId="9" captionText="FIGURES 42–51. 42. Archineura incarnata: caudal gills; left lateral above, median below (drawn from Yang et al. 2022). 43. Echo modesta: caudal gills; left lateral above, median below, with baso-dorsal shield arrowed (original illustration). 44. Psolodesmus mandarinus, dorsal view of caudal gills, with baso-dorsal shield of median gill arrowed (redrawn from Matsuki &amp; Lien 1978). 45. Mnais andersoni, left lateral caudal gill with upper margin of median gill visible; baso-dorsal shield of median gill arrowed (original illustration). 46. Mnais gregoryi: caudal gills; left lateral above, median below, with baso-dorsal shield arrowed (drawn from Yang et al. 2021). 47. Caliphaea angka: caudal gills; left lateral above, median below (drawn from Yang et al. 2021). 48. Matrona basilaris: caudal gills; left lateral above, median below (drawn from Wang et al. 2017). 49. Vestalaria venusta: caudal gills; left lateral above, median below (drawn from Wang et al. 2017). 50. Vestalis amoena group: caudal gills; left lateral above, median below (redrawn from Lieftinck 1965). 51. Vestalis melania: caudal gills; left lateral above, median below (redrawn from Guadalquiver et al. 2022)." figureDoi="http://doi.org/10.5281/zenodo.13618337" httpUri="https://zenodo.org/record/13618337/files/figure.png" pageId="19" pageNumber="228">Fig. 47</figureCitation>
); antennal scape about half total length of antenna (
<figureCitation id="83E12A3EE026D741FC24E9A9FBDBFA42" box="[970,1041,1417,1439]" captionStart="FIGURES 35–41. 35" captionStartId="8.[151,264,1776,1800]" captionTargetBox="[257,1332,159,1764]" captionTargetId="figure-21@8.[253,1335,153,1771]" captionTargetPageId="8" captionText="FIGURES 35–41. 35. Archineura incarnata, prementum (dorsal view, drawn from Yang et al. 2022). 36. Vestalis gracilis, prementum (dorsal view) and right antenna with supplementary antennomere between pedicel and scape (a) arrowed (redrawn from Rattanachan et al. 2022). 37. Vestalaria venusta: prementum (dorsal view) and right antenna (drawn from Wang et al. 2017). 38. Caliphaea angka: prementum (dorsal view) and right antenna (drawn from Yang et al. 2021). 39. Heliocypha biseriata, frontal view of labrum showing dorsal and ventral sections and ‘moustache’ of long, heavy setae on the latter (from Orr et al. 2024). 40. Heliocypha biseriata: details of labial palp in relation to prementum (a) dorsal view—base of movable hook dashed to reveal palpal lobe beneath and open base of premental median cleft arrowed, (b) frontal view, (c) inner view (from Orr et al. 2024). 41. Heliocypha biseriata, detail of fore tarsus with pectinate setae magnified (from Orr et al. 2024)." figureDoi="http://doi.org/10.5281/zenodo.13618335" httpUri="https://zenodo.org/record/13618335/files/figure.png" pageId="19" pageNumber="228">Fig. 38</figureCitation>
)........................
<taxonomicName id="DCDA4D38E026D741FAD9E9A8FA56FA42" authorityName="Hagen in Selys" authorityYear="1859" box="[1335,1436,1416,1438]" class="Insecta" family="Calopterygidae" genus="Caliphaea" kingdom="Animalia" order="Odonata" pageId="19" pageNumber="228" phylum="Arthropoda" rank="genus">
<emphasis id="29AEEAA9E026D741FAD9E9A8FA56FA42" bold="true" box="[1335,1436,1416,1438]" italics="true" pageId="19" pageNumber="228">Caliphaea</emphasis>
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</keyLead>
</paragraph>
<paragraph id="1B6536BBE026D741FF79E986FA56FA05" blockId="19.[151,1437,361,1762]" pageId="19" pageNumber="228">
<keyLead id="A02B958EE026D741FF79E986FA56FA05" pageId="19" pageNumber="228">
8’ Premental cleft narrow, pear-drop outline, tips of lobes of prementum touching or overlapping cleft, relatively short, ratio A/B:0.25–0.30 (
<figureCitation id="83E12A3EE026D741FEBBE9E4FE0CFA05" box="[341,454,1476,1498]" captionStart="FIGURES 27–34. 27" captionStartId="7.[151,264,1845,1869]" captionTargetBox="[252,1326,159,1834]" captionTargetId="figure-15@7.[251,1336,153,1839]" captionTargetPageId="7" captionText="FIGURES 27–34. 27. Prementum of Calopterygidae showing measurements used in family key (based on Calopteryx japonica, redrawn from Miyakawa 1983). 28. Neurobasis chinensis, prementum and left labial palp (redrawn from Lieftinck 1965). 29. Matronoides cyaneipennis, prementum and labial palps (ventral view). 30. Matrona basilaris, prementum (dorsal view, drawn from Wang et al. 2017). 31. Atrocalopteryx atrata, prementum (dorsal view, redrawn from Miyakawa 1983). 32. Mnais tenuis, prementum (dorsal view, redrawn from Matsuki &amp; Lien 1977). 33. Echo uniformis, prementum and left labial palp dorsal view (redrawn from Lieftinck 1965). 34. Psolodesmus mandarinus, prementum (dorsal view, redrawn from Matsuki &amp; Lien 1978)." figureDoi="http://doi.org/10.5281/zenodo.13618333" httpUri="https://zenodo.org/record/13618333/files/figure.png" pageId="19" pageNumber="228">Figs 32–34</figureCitation>
); lateral caudal gills ranging from 0.25–0.33 total length of remainder of body.................. 9
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<keyStep id="A02E2E1EE026D741FF79E9C1FA56F9ED" pageId="19" pageNumber="228">
<paragraph id="1B6536BBE026D741FF79E9C1FA57F9CF" blockId="19.[151,1437,361,1762]" pageId="19" pageNumber="228">
<keyLead id="A02B958EE026D741FF79E9C1FA57F9CF" pageId="19" pageNumber="228">
9 (8’) Lateral gills smooth, without denticles, elongate ovoid (
<figureCitation id="83E12A3EE026D741FD15E9C1FC8BFA2B" box="[763,833,1505,1527]" captionStart="FIGURES 42–51. 42" captionStartId="9.[151,264,1592,1616]" captionTargetBox="[165,1413,192,1557]" captionTargetId="figure-15@9.[151,1435,181,1568]" captionTargetPageId="9" captionText="FIGURES 42–51. 42. Archineura incarnata: caudal gills; left lateral above, median below (drawn from Yang et al. 2022). 43. Echo modesta: caudal gills; left lateral above, median below, with baso-dorsal shield arrowed (original illustration). 44. Psolodesmus mandarinus, dorsal view of caudal gills, with baso-dorsal shield of median gill arrowed (redrawn from Matsuki &amp; Lien 1978). 45. Mnais andersoni, left lateral caudal gill with upper margin of median gill visible; baso-dorsal shield of median gill arrowed (original illustration). 46. Mnais gregoryi: caudal gills; left lateral above, median below, with baso-dorsal shield arrowed (drawn from Yang et al. 2021). 47. Caliphaea angka: caudal gills; left lateral above, median below (drawn from Yang et al. 2021). 48. Matrona basilaris: caudal gills; left lateral above, median below (drawn from Wang et al. 2017). 49. Vestalaria venusta: caudal gills; left lateral above, median below (drawn from Wang et al. 2017). 50. Vestalis amoena group: caudal gills; left lateral above, median below (redrawn from Lieftinck 1965). 51. Vestalis melania: caudal gills; left lateral above, median below (redrawn from Guadalquiver et al. 2022)." figureDoi="http://doi.org/10.5281/zenodo.13618337" httpUri="https://zenodo.org/record/13618337/files/figure.png" pageId="19" pageNumber="228">Fig. 45</figureCitation>
) or broadly sickle-shaped, about one third length of remainder of body; postocular lobes shallow in outline; one strong setae on lobes of prementum (
<figureCitation id="83E12A3EE026D741FBCEE9DEFBAFF9CF" box="[1056,1125,1534,1556]" captionStart="FIGURES 27–34. 27" captionStartId="7.[151,264,1845,1869]" captionTargetBox="[252,1326,159,1834]" captionTargetId="figure-15@7.[251,1336,153,1839]" captionTargetPageId="7" captionText="FIGURES 27–34. 27. Prementum of Calopterygidae showing measurements used in family key (based on Calopteryx japonica, redrawn from Miyakawa 1983). 28. Neurobasis chinensis, prementum and left labial palp (redrawn from Lieftinck 1965). 29. Matronoides cyaneipennis, prementum and labial palps (ventral view). 30. Matrona basilaris, prementum (dorsal view, drawn from Wang et al. 2017). 31. Atrocalopteryx atrata, prementum (dorsal view, redrawn from Miyakawa 1983). 32. Mnais tenuis, prementum (dorsal view, redrawn from Matsuki &amp; Lien 1977). 33. Echo uniformis, prementum and left labial palp dorsal view (redrawn from Lieftinck 1965). 34. Psolodesmus mandarinus, prementum (dorsal view, redrawn from Matsuki &amp; Lien 1978)." figureDoi="http://doi.org/10.5281/zenodo.13618333" httpUri="https://zenodo.org/record/13618333/files/figure.png" pageId="19" pageNumber="228">Fig. 32</figureCitation>
)....................
<taxonomicName id="DCDA4D38E026D741FAB0E9DEFA57F9CF" authorityName="Selys" authorityYear="1853" box="[1374,1437,1534,1555]" class="Insecta" family="Calopterygidae" genus="Mnais" kingdom="Animalia" order="Odonata" pageId="19" pageNumber="228" phylum="Arthropoda" rank="genus">
<emphasis id="29AEEAA9E026D741FAB0E9DEFA57F9CF" bold="true" box="[1374,1437,1534,1555]" italics="true" pageId="19" pageNumber="228">Mnais</emphasis>
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<paragraph id="1B6536BBE026D741FF79EA3CFA56F9ED" blockId="19.[151,1437,361,1762]" box="[151,1436,1564,1586]" pageId="19" pageNumber="228">
<keyLead id="A02B958EE026D741FF79EA3CFA56F9ED" box="[151,1436,1564,1586]" pageId="19" pageNumber="228">9’ Lateral gills bearing denticles on central outer rib and sometimes on margins..................................... 10</keyLead>
</paragraph>
</keyStep>
<keyStep id="A02E2E1EE026D741FF79EA19FA56F93E" pageId="19" pageNumber="228">
<paragraph id="1B6536BBE026D741FF79EA19FA56F956" blockId="19.[151,1437,361,1762]" pageId="19" pageNumber="228">
<keyLead id="A02B958EE026D741FF79EA19FA56F956" pageId="19" pageNumber="228">
10 (9’) Postocular lobe shallow in outline with small tubercle (
<figureCitation id="83E12A3EE026D741FD14EA19FC8AF993" box="[762,832,1593,1615]" captionStart="FIGURES 52–56. 52" captionStartId="10.[151,264,1941,1965]" captionTargetBox="[191,1388,163,1929]" captionTargetId="figure-21@10.[190,1397,151,1933]" captionTargetPageId="10" captionText="FIGURES 52–56. 52. Caliphaea angka, habitus (based on Yang et al. 2021). 53. Vestalis gracilis, habitus (redrawn from Rattanachan et al. 2022). 54. Neurobasis chinensis, habitus (original drawing). 55. Matrona basilaris, habitus (drawn from Wang et al. 2017). 56. Echo uniformis, habitus (redrawn from Lieftinck 1965)." figureDoi="http://doi.org/10.5281/zenodo.13618339" httpUri="https://zenodo.org/record/13618339/files/figure.png" pageId="19" pageNumber="228">Fig. 56</figureCitation>
) (but see note on Fraser 1924); lateral caudal gills about three times as long as broad with denticles along dorsal and ventral margins and small tubercles on outer central rib (
<figureCitation id="83E12A3EE026D741FAE3EA76FA9FF9B7" box="[1293,1365,1622,1644]" captionStart="FIGURES 42–51. 42" captionStartId="9.[151,264,1592,1616]" captionTargetBox="[165,1413,192,1557]" captionTargetId="figure-15@9.[151,1435,181,1568]" captionTargetPageId="9" captionText="FIGURES 42–51. 42. Archineura incarnata: caudal gills; left lateral above, median below (drawn from Yang et al. 2022). 43. Echo modesta: caudal gills; left lateral above, median below, with baso-dorsal shield arrowed (original illustration). 44. Psolodesmus mandarinus, dorsal view of caudal gills, with baso-dorsal shield of median gill arrowed (redrawn from Matsuki &amp; Lien 1978). 45. Mnais andersoni, left lateral caudal gill with upper margin of median gill visible; baso-dorsal shield of median gill arrowed (original illustration). 46. Mnais gregoryi: caudal gills; left lateral above, median below, with baso-dorsal shield arrowed (drawn from Yang et al. 2021). 47. Caliphaea angka: caudal gills; left lateral above, median below (drawn from Yang et al. 2021). 48. Matrona basilaris: caudal gills; left lateral above, median below (drawn from Wang et al. 2017). 49. Vestalaria venusta: caudal gills; left lateral above, median below (drawn from Wang et al. 2017). 50. Vestalis amoena group: caudal gills; left lateral above, median below (redrawn from Lieftinck 1965). 51. Vestalis melania: caudal gills; left lateral above, median below (redrawn from Guadalquiver et al. 2022)." figureDoi="http://doi.org/10.5281/zenodo.13618337" httpUri="https://zenodo.org/record/13618337/files/figure.png" pageId="19" pageNumber="228">Fig. 43</figureCitation>
).
<taxonomicName id="DCDA4D38E026D741FA8EEA76FA5EF9B0" box="[1376,1428,1622,1644]" class="Insecta" family="Calopterygidae" genus="Echo" kingdom="Animalia" order="Odonata" pageId="19" pageNumber="228" phylum="Arthropoda" rank="genus">
<emphasis id="29AEEAA9E026D741FA8EEA76FA5EF9B0" bold="true" box="[1376,1428,1622,1644]" italics="true" pageId="19" pageNumber="228">Echo</emphasis>
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[Sumatra, Peninsular
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, eastern Himalayas,
<collectingCountry id="63CD762BE026D741FBF8EA54FBBFF955" box="[1046,1141,1652,1673]" name="Myanmar" pageId="19" pageNumber="228">Myanmar</collectingCountry>
,
<collectingCountry id="63CD762BE026D741FB90EA54FB19F956" box="[1150,1235,1652,1674]" name="Thailand" pageId="19" pageNumber="228">Thailand</collectingCountry>
and southern
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]
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</paragraph>
<paragraph id="1B6536BBE026D741FF79EAB1FA56F93E" blockId="19.[151,1437,361,1762]" pageId="19" pageNumber="228">
<keyLead id="A02B958EE026D741FF79EAB1FA56F93E" pageId="19" pageNumber="228">
10’ Postocular lobe well rounded in outline with small tubercle; lateral gills ca 0.33 times length of rest of body only outer central rib of lateral gills with very strong denticles; margins smooth (
<figureCitation id="83E12A3EE026D741FCD4EA8EFC4AF91F" box="[826,896,1710,1732]" captionStart="FIGURES 42–51. 42" captionStartId="9.[151,264,1592,1616]" captionTargetBox="[165,1413,192,1557]" captionTargetId="figure-15@9.[151,1435,181,1568]" captionTargetPageId="9" captionText="FIGURES 42–51. 42. Archineura incarnata: caudal gills; left lateral above, median below (drawn from Yang et al. 2022). 43. Echo modesta: caudal gills; left lateral above, median below, with baso-dorsal shield arrowed (original illustration). 44. Psolodesmus mandarinus, dorsal view of caudal gills, with baso-dorsal shield of median gill arrowed (redrawn from Matsuki &amp; Lien 1978). 45. Mnais andersoni, left lateral caudal gill with upper margin of median gill visible; baso-dorsal shield of median gill arrowed (original illustration). 46. Mnais gregoryi: caudal gills; left lateral above, median below, with baso-dorsal shield arrowed (drawn from Yang et al. 2021). 47. Caliphaea angka: caudal gills; left lateral above, median below (drawn from Yang et al. 2021). 48. Matrona basilaris: caudal gills; left lateral above, median below (drawn from Wang et al. 2017). 49. Vestalaria venusta: caudal gills; left lateral above, median below (drawn from Wang et al. 2017). 50. Vestalis amoena group: caudal gills; left lateral above, median below (redrawn from Lieftinck 1965). 51. Vestalis melania: caudal gills; left lateral above, median below (redrawn from Guadalquiver et al. 2022)." figureDoi="http://doi.org/10.5281/zenodo.13618337" httpUri="https://zenodo.org/record/13618337/files/figure.png" pageId="19" pageNumber="228">Fig. 44</figureCitation>
).................................
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<emphasis id="29AEEAA9E026D741FAF2EA8EFA56F918" bold="true" box="[1308,1436,1710,1732]" italics="true" pageId="19" pageNumber="228">Psolodesmus</emphasis>
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[
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and nearby islands]
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</treatment>
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122
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<mods:namePart id="E7306F50E744D7C2E6DDABA78C27478B">Orr, Albert G. W.</mods:namePart>
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<mods:date id="E5B162650003077A75AC54E4F759CA8E">2024</mods:date>
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<mods:number id="B0B879B6E05229EFE3392D2595E7117A">5497</mods:number>
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<subSubSection id="53C06530E03BD75CFF79EB5BFE88F84A" box="[151,322,1915,1942]" pageId="14" pageNumber="223" type="nomenclature">
<paragraph id="1B6536BBE03BD75CFF79EB5BFE88F84A" blockId="14.[151,1436,1915,2014]" box="[151,322,1915,1942]" pageId="14" pageNumber="223">
<heading id="402D81D7E03BD75CFF79EB5BFE88F84A" bold="true" box="[151,322,1915,1942]" fontSize="11" level="1" pageId="14" pageNumber="223" reason="1">
<taxonomicName id="DCDA4D38E03BD75CFF79EB5BFE88F84A" ID-CoL="35D7" authorityName="Dumont, Vanfleteren, De Jonckheere &amp; Weekers" authorityYear="2005" box="[151,322,1915,1942]" class="Insecta" family="Calopterygidae" genus="Atrocalopteryx" kingdom="Animalia" order="Odonata" pageId="14" pageNumber="223" phylum="Arthropoda" rank="genus">
<emphasis id="29AEEAA9E03BD75CFF79EB5BFE88F84A" bold="true" box="[151,322,1915,1942]" italics="true" pageId="14" pageNumber="223">Atrocalopteryx</emphasis>
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<subSubSection id="53C06530E03BD75DFF79EBBFFD6DFDC5" lastPageId="15" lastPageNumber="224" pageId="14" pageNumber="223" type="discussion">
<paragraph id="1B6536BBE03BD75DFF79EBBFFD6DFDC5" blockId="14.[151,1436,1915,2014]" lastBlockId="15.[151,1437,151,537]" lastPageId="15" lastPageNumber="224" pageId="14" pageNumber="223">
The genus ranges from
<collectingCountry id="63CD762BE03BD75CFE43EB80FE27F866" box="[429,493,1952,1978]" name="Japan" pageId="14" pageNumber="223">Japan</collectingCountry>
to
<collectingCountry id="63CD762BE03BD75CFDF6EBBFFD91F865" box="[536,603,1951,1977]" name="China" pageId="14" pageNumber="223">China</collectingCountry>
and northern Indochina and includes eight species. The larva is known for only one species,
<taxonomicName id="DCDA4D38E03BD75CFE60EBE3FD11F801" authority="(Selys)" baseAuthorityName="Selys" box="[398,731,1987,2013]" class="Insecta" family="Calopterygidae" genus="Atrocalopteryx" kingdom="Animalia" order="Odonata" pageId="14" pageNumber="223" phylum="Arthropoda" rank="species" species="atrata">
<emphasis id="29AEEAA9E03BD75CFE60EBE3FD48F801" box="[398,642,1987,2013]" italics="true" pageId="14" pageNumber="223">Atrocalopteryx atrata</emphasis>
(Selys)
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which has a wide range from
<collectingCountry id="63CD762BE03BD75CFBD5EBE4FBB1F802" box="[1083,1147,1988,2014]" name="Japan" pageId="14" pageNumber="223">Japan</collectingCountry>
to
<collectingCountry id="63CD762BE03BD75CFB4DEBE4FB2DF802" box="[1187,1255,1988,2014]" name="South Korea" pageId="14" pageNumber="223">Korea</collectingCountry>
and throughout most of
<collectingCountry id="63CD762BE03AD75DFF14ECB7FE8BFF6D" box="[250,321,151,177]" name="China" pageId="15" pageNumber="224">China</collectingCountry>
. It was first described by Asahina (1940) from an immature specimen, with the habitus, caudal gills and anterior mask depicted. Subsequently a description based on a F instar larva was provided by the same author (
<bibRefCitation id="7F4B4B4AE03AD75DFF00ECFFFE5AFF25" author="Asahina, S." box="[238,400,222,249]" pageId="15" pageNumber="224" pagination="52 - 55" refId="ref21577" refString="Asahina, S. (1956) An annotated list of Odonata of Japan with brief descriptions of their larva. Shin-konchu, 9 (12), 52 - 55." type="journal article" year="1956">Asahina 1956</bibRefCitation>
), who figured the prementum and labial palps, caudal gills, head and antenna and a habitus photograph. Further detail was provided by
<bibRefCitation id="7F4B4B4AE03AD75DFD96ED23FCC5FEC1" author="Ishida, K." box="[632,783,259,285]" pageId="15" pageNumber="224" refId="ref23597" refString="Ishida, K. (1996) Monograph of Odonata larvae in Japan. Hokkaido University Press, Sapporo, 446 pp. [in Japanese]" type="book" year="1996">Ishida (1996)</bibRefCitation>
. In the northern part of its range it overlaps with
<taxonomicName id="DCDA4D38E03AD75DFACEED23FF21FE9D" pageId="15" pageNumber="224">
<emphasis id="29AEEAA9E03AD75DFACEED23FA56FEC1" box="[1312,1436,259,285]" italics="true" pageId="15" pageNumber="224">Calopteryx</emphasis>
species
</taxonomicName>
, a genus not present in the Oriental region, and
<bibRefCitation id="7F4B4B4AE03AD75DFCF4ED07FC78FE9D" author="Ishida, K." box="[794,946,295,321]" pageId="15" pageNumber="224" refId="ref23597" refString="Ishida, K. (1996) Monograph of Odonata larvae in Japan. Hokkaido University Press, Sapporo, 446 pp. [in Japanese]" type="book" year="1996">Ishida (1996)</bibRefCitation>
and
<bibRefCitation id="7F4B4B4AE03AD75DFC03ED07FB7DFE9D" author="Miyakawa, K." box="[1005,1207,295,321]" pageId="15" pageNumber="224" pagination="25 - 34" refId="ref24906" refString="Miyakawa, K. (1983) Description of the larva of Calopteryx japonica Selys, in comparison with C. virgo (L.) and C. atrata Selys larvae (Odonata, Calopterygidae). Proceedings of the Japanese Society of Systematic Zoology, 26, 25 - 34." type="journal article" year="1983">Miyakawa (1983)</bibRefCitation>
demonstrate how it may be separated from sympatric species of that genus. In Indochina and south-eastern
<collectingCountry id="63CD762BE03AD75DFB90ED6BFB0BFEB9" box="[1150,1217,331,357]" name="China" pageId="15" pageNumber="224">China</collectingCountry>
the genus overlaps broadly with species of
<taxonomicName id="DCDA4D38E03AD75DFE46ED4FFDC3FE54" authorityName="Selys" authorityYear="1853" box="[424,521,367,392]" class="Insecta" family="Calopterygidae" genus="Matrona" kingdom="Animalia" order="Odonata" pageId="15" pageNumber="224" phylum="Arthropoda" rank="genus">
<emphasis id="29AEEAA9E03AD75DFE46ED4FFDC3FE54" box="[424,521,367,392]" italics="true" pageId="15" pageNumber="224">Matrona</emphasis>
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with which
<taxonomicName id="DCDA4D38E03AD75DFD76ED4FFD36FE54" box="[664,764,367,393]" class="Insecta" family="Calopterygidae" genus="Atrocalopteryx" kingdom="Animalia" order="Odonata" pageId="15" pageNumber="224" phylum="Arthropoda" rank="species" species="atrata">
<emphasis id="29AEEAA9E03AD75DFD76ED4FFD36FE54" box="[664,764,367,393]" italics="true" pageId="15" pageNumber="224">A. atrata</emphasis>
</taxonomicName>
larvae share some similarities, hence it is not certain if the key provided separates all species, but with interspecific variation in adult
<taxonomicName id="DCDA4D38E03AD75DFC02EDB3FB59FE71" authorityName="Dumont, Vanfleteren, De Jonckheere &amp; Weekers" authorityYear="2005" box="[1004,1171,403,429]" class="Insecta" family="Calopterygidae" genus="Atrocalopteryx" kingdom="Animalia" order="Odonata" pageId="15" pageNumber="224" phylum="Arthropoda" rank="genus">
<emphasis id="29AEEAA9E03AD75DFC02EDB3FB59FE71" box="[1004,1171,403,429]" italics="true" pageId="15" pageNumber="224">Atrocalopteryx</emphasis>
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being modest, we may suppose that this is true also of larvae. The clearest character distinguishing
<taxonomicName id="DCDA4D38E03AD75DFC1FED97FB9FFE0C" box="[1009,1109,439,465]" class="Insecta" family="Calopterygidae" genus="Atrocalopteryx" kingdom="Animalia" order="Odonata" pageId="15" pageNumber="224" phylum="Arthropoda" rank="species" species="atrata">
<emphasis id="29AEEAA9E03AD75DFC1FED97FB9FFE0C" box="[1009,1109,439,465]" italics="true" pageId="15" pageNumber="224">A. atrata</emphasis>
</taxonomicName>
from
<taxonomicName id="DCDA4D38E03AD75DFB75ED97FB36FE0C" authorityName="Selys" authorityYear="1853" box="[1179,1276,439,464]" class="Insecta" family="Calopterygidae" genus="Matrona" kingdom="Animalia" order="Odonata" pageId="15" pageNumber="224" phylum="Arthropoda" rank="genus">
<emphasis id="29AEEAA9E03AD75DFB75ED97FB36FE0C" box="[1179,1276,439,464]" italics="true" pageId="15" pageNumber="224">Matrona</emphasis>
</taxonomicName>
is the form of the prementum (
<figureCitation id="83E12A3EE03AD75DFEA0EDFBFE6BFE29" box="[334,417,475,501]" captionStart="FIGURES 27–34. 27" captionStartId="7.[151,264,1845,1869]" captionTargetBox="[252,1326,159,1834]" captionTargetId="figure-15@7.[251,1336,153,1839]" captionTargetPageId="7" captionText="FIGURES 27–34. 27. Prementum of Calopterygidae showing measurements used in family key (based on Calopteryx japonica, redrawn from Miyakawa 1983). 28. Neurobasis chinensis, prementum and left labial palp (redrawn from Lieftinck 1965). 29. Matronoides cyaneipennis, prementum and labial palps (ventral view). 30. Matrona basilaris, prementum (dorsal view, drawn from Wang et al. 2017). 31. Atrocalopteryx atrata, prementum (dorsal view, redrawn from Miyakawa 1983). 32. Mnais tenuis, prementum (dorsal view, redrawn from Matsuki &amp; Lien 1977). 33. Echo uniformis, prementum and left labial palp dorsal view (redrawn from Lieftinck 1965). 34. Psolodesmus mandarinus, prementum (dorsal view, redrawn from Matsuki &amp; Lien 1978)." figureDoi="http://doi.org/10.5281/zenodo.13618333" httpUri="https://zenodo.org/record/13618333/files/figure.png" pageId="15" pageNumber="224">Fig. 31</figureCitation>
). The caudal gills of
<emphasis id="29AEEAA9E03AD75DFD67EDFBFD39FE28" box="[649,755,475,501]" italics="true" pageId="15" pageNumber="224">
<taxonomicName id="DCDA4D38E03AD75DFD67EDFBFD24FE28" box="[649,750,475,501]" class="Insecta" family="Calopterygidae" genus="Atrocalopteryx" kingdom="Animalia" order="Odonata" pageId="15" pageNumber="224" phylum="Arthropoda" rank="species" species="atrata">A. atrata</taxonomicName>
,
</emphasis>
well depicted by (
<bibRefCitation id="7F4B4B4AE03AD75DFC51EDFBFBB2FE29" author="Miyakawa, K." box="[959,1144,475,501]" pageId="15" pageNumber="224" pagination="25 - 34" refId="ref24906" refString="Miyakawa, K. (1983) Description of the larva of Calopteryx japonica Selys, in comparison with C. virgo (L.) and C. atrata Selys larvae (Odonata, Calopterygidae). Proceedings of the Japanese Society of Systematic Zoology, 26, 25 - 34." type="journal article" year="1983">Miyakawa 1983</bibRefCitation>
), are very similar in shape and proportions to those of known
<taxonomicName id="DCDA4D38E03AD75DFDF1EDDFFD6DFDC5" box="[543,679,511,537]" class="Insecta" family="Calopterygidae" genus="Matrona" kingdom="Animalia" order="Odonata" pageId="15" pageNumber="224" phylum="Arthropoda" rank="species" species="undetermined">
<emphasis id="29AEEAA9E03AD75DFDF1EDDFFD4AFDC4" box="[543,640,511,536]" italics="true" pageId="15" pageNumber="224">Matrona</emphasis>
sp.
</taxonomicName>
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