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<document id="D8B7A0023C77EDDFF0F843A65F9CAF49" ID-DOI="10.5281/zenodo.5311784" ID-ISSN="0374-1036" ID-Zenodo-Dep="5311784" ID-ZooBank="15CF0A9E-44F3-492B-88F7-A7922EF9F73A" IM.materialsCitations_approvedBy="felipe" IM.metadata_approvedBy="felipe" IM.tables_requiresApprovalFor="existingObjects,plazi" IM.taxonomicNames_approvedBy="felipe" checkinTime="1630118666848" checkinUser="marcus" docAuthor="Angus, Robert B., Jia, Fenglong, Chen, Zhen-ning, Zhang, Ying, Vondráček, Dominik, Fikáček, Martin &amp; Sw, London" docDate="2016" docId="377A879F8D65284BFE555E31719BFB1F" docLanguage="en" docName="ActaEntMusNatPra.2016.56.1.109-148.pdf" docOrigin="Acta Entomologica Musei Nationalis Pragae 56 (1)" docStyle="DocumentStyle:E9BE312A6EDF9B84FC5211AAA80D80BF.10:ActaEntMusNatPra.2005-2016.journal_article" docStyleId="E9BE312A6EDF9B84FC5211AAA80D80BF" docStyleName="ActaEntMusNatPra.2005-2016.journal_article" docStyleVersion="10" docTitle="Helophorus (Lihelophorus) lamicola Zaitzev 1908" docType="treatment" docVersion="3" lastPageNumber="138" masterDocId="CB43FFE78D792856FFE05B4D7368FFAC" masterDocTitle="Taxonomy, larval morphology and cytogenetics of Lihelophorus, the Tibetan endemic subgenus of Helophorus (Coleoptera: Hydrophiloidea)" masterLastPageNumber="148" masterPageNumber="109" pageNumber="137" updateTime="1699158837459" updateUser="plazi" zenodo-license-document="CC-BY-4.0">
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<mods:title id="788D67D419E01479C94C745D38E92CA7">Taxonomy, larval morphology and cytogenetics of Lihelophorus, the Tibetan endemic subgenus of Helophorus (Coleoptera: Hydrophiloidea)</mods:title>
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<mods:namePart id="DE81B55EB0BDEFE1EDB1DAA28BD9E997">Angus, Robert B.</mods:namePart>
<mods:affiliation id="EB5127DBCBAEECD46A78C9F5D9138EF6">Division of Life Sciences (Insects), Natural History Museum, Cromwell Road, London SW 7 5 BD, UK; e-mail: r. angus @ royalholloway. ac. uk</mods:affiliation>
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<mods:namePart id="43BB8B71821D25FB1954231756DEFBBB">Jia, Fenglong</mods:namePart>
<mods:affiliation id="F48EF5CF0BB397395E48962B38EFD9FD">Museum of Biology, Sun Yat-sen University, Guangzhou, China; e-mail: lssjϐl @ mail. sysu. edu. cn</mods:affiliation>
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<mods:namePart id="450F4399BA46927CD23DE0087582B805">Chen, Zhen-ning</mods:namePart>
<mods:affiliation id="F368AE040B6D4B9FF0AD01C60C5FECF6">Biology and Geography School, Qinghai Normal University, Wusi West Road 38 #, 810000, Xining, Qinghai Province, China; e-mail: 149470880 @ qq. com</mods:affiliation>
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<mods:namePart id="6BCFD173E30775791A479FEA0EABFAE2">Zhang, Ying</mods:namePart>
<mods:affiliation id="C8E1ACE221A1C9263640659AD73B29AB">Computer Network Information Center, Chinese Academy of Sciences, Beijing, 100190, China; e-mail: 344280438 @ qq. com</mods:affiliation>
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<mods:namePart id="0939882ADE91AD3358C14D1214452559">Vondráček, Dominik</mods:namePart>
<mods:affiliation id="BEB28BA271CBE57B08D350F1A5BC818B">Charles University in Prague, Faculty of Science, Department of Zoology, Viničná 7, CZ- 128 43 Prague 2, Czech Republic; e-mail: dominik. vondracek @ gmail. com &amp; Department of Entomology, National Museum, Cirkusová 1740, CZ- 193 00 Praha 9 - Horní Počernice, Czech Republic; e-mail: mfikacek @ gmail. com</mods:affiliation>
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<emphasis id="8DA7EA9B8D65284AFE555E31715FFA3F" bold="true" box="[437,567,1404,1427]" italics="true" pageId="28" pageNumber="137">Helophorus</emphasis>
(
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) 
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<emphasis id="8DA7EA9B8D65284AFF775EE472EAFA10" bold="true" box="[151,386,1449,1468]" pageId="28" pageNumber="137">Larval material examined</emphasis>
(
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1 first instar larva (
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, DNA voucher 
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: 
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:
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, 
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, roadside pool, 
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, 
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&amp; 
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lgt
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.; 
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4 first instar larvae (BMNH): same label data. All larvae were reared from the single egg case (deposited in BMNH) laid by a field-collected female, association with adults from the same locality was confirmed by the 
<emphasis id="8DA7EA9B8D65284AFBDE5EB4770FF9A0" box="[1086,1127,1529,1548]" italics="true" pageId="28" pageNumber="137">cox1</emphasis>
sequence data
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.
</paragraph>
<paragraph id="BF6C36898D64284BFF915B9170EFFEDF" blockId="29.[113,1090,220,1203]" pageId="29" pageNumber="138">
<emphasis id="8DA7EA9B8D64284BFF915B9173BBFF5F" bold="true" box="[113,211,220,243]" pageId="29" pageNumber="138">Egg case</emphasis>
(
<figureCitation id="27E82A0C8D64284BFF015B917246FF5F" box="[225,302,220,243]" captionStart="Fig" captionStartId="19.[113,144,1464,1483]" captionTargetBox="[114,1086,453,1447]" captionTargetId="figure-13@19.[113,1087,443,1448]" captionTargetPageId="19" captionText="Fig. 8. Immature stages of Lihelophorus. a–d – head and prothorax of first instar larvae: a–b – H. (Lihelophorus) lamicola Zaitzev, 1908 (a – dorsal; b – ventral); c–d – H. (Lihelophorus) yangae sp. nov. (c – dorsal; d – ventral). e–f – egg cases: e – H. (Lihelophorus) lamicola Zaitzev, 1908; f – H. (Lihelophorus) yangae sp. nov.; g–j – abdomen with spiracles containing air bubbles: g–h – H. (Lihelophorus) lamicola Zaitzev, 1908 (g – tracheal gills and its tracheal system; h – detail of tracheal branching around abdominal spiracle); i–j – H. (Lihelophorus) yangae sp. nov. (i – detail of tracheal system around abdominal spiracles; j – tracheal system of abdomen). Abbreviations: sp – spiracle; trg – tracheal gill." figureDoi="http://doi.org/10.5281/zenodo.5311800" httpUri="https://zenodo.org/record/5311800/files/figure.png" pageId="29" pageNumber="138">Fig. 8e</figureCitation>
). Consisting of an egg-bag surmounted by a thin trailing mast which takes the form of a long narrow tube, thus corresponding to 
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<specimenCount id="A9D5FD008D64284BFD095BB1704BFEBF" box="[745,803,252,275]" pageId="29" pageNumber="138" type="egg">4 egg</specimenCount>
case of 
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<collectingRegion id="7D17F86B8D64284BFC985BB170D6FEBD" box="[888,958,252,275]" country="United Kingdom" name="Angus" pageId="29" pageNumber="138">ANGUS</collectingRegion>
(1992)
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. The egg case was placed in the sand at the water’s edge in the aquarium housing the beetles, and has sand grains adhering to the egg-bag. A similar egg case was observed in the ground at the water’s edge in the Zuimatan locality where the beetles were collected
</paragraph>
</subSubSection>
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<emphasis id="8DA7EA9B8D64284BFF915A3172DFFE3F" bold="true" box="[113,439,380,403]" pageId="29" pageNumber="138">Larva. General morphology</emphasis>
(only characters different from 
<taxonomicName id="78D34D0A8D64284BFCFB5A3170E4FE3F" authorityName="Angus &amp; Jia &amp; Chen &amp; Zhang &amp; Vondráček &amp; Fikáček &amp; Sw" authorityYear="2016" box="[795,908,380,403]" class="Insecta" family="Helophoridae" genus="Helophorus" kingdom="Animalia" order="Coleoptera" pageId="29" pageNumber="138" phylum="Arthropoda" rank="species" species="yangae">
<emphasis id="8DA7EA9B8D64284BFCFB5A3170E4FE3F" box="[795,908,380,403]" italics="true" pageId="29" pageNumber="138">H. yangae</emphasis>
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are mentioned). Body (
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) elongate, slender, almost parallel-sided. Body length ca. 
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. 
<emphasis id="8DA7EA9B8D64284BFC235AD17096FE1F" bold="true" box="[963,1022,412,435]" italics="true" pageId="29" pageNumber="138">Head</emphasis>
width 
<quantity id="782B9B6C8D64284BFF915AF17265FE7F" box="[113,269,444,467]" metricMagnitude="-4" metricUnit="m" metricValue="4.45" metricValueMax="4.6" metricValueMin="4.3" pageId="29" pageNumber="138" unit="mm" value="0.445" valueMax="0.46" valueMin="0.43">0.43–0.46 mm</quantity>
(n = 5). Parietale (
<figureCitation id="27E82A0C8D64284BFE3B5AF1713DFE7F" box="[475,597,444,467]" captionStart="Fig" captionStartId="24.[151,182,1519,1538]" captionTargetBox="[153,1089,219,1436]" captionTargetId="figure-10@24.[151,1089,219,1436]" captionTargetPageId="24" captionText="Fig. 12. Labroclypeus and epistome of larvae of Lihelophorus and Helophorus s.str. a–b – H. (H.) liguricus Angus, 1970 (a – dorsal view; b – ventral view); c–d – H. (Lihelophorus) yangae sp. nov. (c – dorsal; d – ventral); e–f – H. (Lihelophorus) lamicola Zaitzev, 1908 (e – dorsal; f – ventral)." figureDoi="http://doi.org/10.5281/zenodo.5311810" httpUri="https://zenodo.org/record/5311810/files/figure.png" pageId="29" pageNumber="138">Figs 12a–b</figureCitation>
) with numerous cuticular spines dorsally and laterally, ventral face smooth. Frontoclypeus (
<figureCitation id="27E82A0C8D64284BFDAE5A9171CCFE5F" box="[590,676,476,499]" captionStart="Fig" captionStartId="24.[151,182,1519,1538]" captionTargetBox="[153,1089,219,1436]" captionTargetId="figure-10@24.[151,1089,219,1436]" captionTargetPageId="24" captionText="Fig. 12. Labroclypeus and epistome of larvae of Lihelophorus and Helophorus s.str. a–b – H. (H.) liguricus Angus, 1970 (a – dorsal view; b – ventral view); c–d – H. (Lihelophorus) yangae sp. nov. (c – dorsal; d – ventral); e–f – H. (Lihelophorus) lamicola Zaitzev, 1908 (e – dorsal; f – ventral)." figureDoi="http://doi.org/10.5281/zenodo.5311810" httpUri="https://zenodo.org/record/5311810/files/figure.png" pageId="29" pageNumber="138">Fig. 12e</figureCitation>
) with large bulged asymmetrical nasale bearing single median tooth, and large slightly asymmetrical epistomal lobes overlapping nasale. Gular suture absent, posterior tentorial pits minute, situated anteromesally (
<figureCitation id="27E82A0C8D64284BFC2A5951775DFD9F" box="[970,1077,540,563]" captionStart="Fig" captionStartId="19.[113,144,1464,1483]" captionTargetBox="[114,1086,453,1447]" captionTargetId="figure-13@19.[113,1087,443,1448]" captionTargetPageId="19" captionText="Fig. 8. Immature stages of Lihelophorus. a–d – head and prothorax of first instar larvae: a–b – H. (Lihelophorus) lamicola Zaitzev, 1908 (a – dorsal; b – ventral); c–d – H. (Lihelophorus) yangae sp. nov. (c – dorsal; d – ventral). e–f – egg cases: e – H. (Lihelophorus) lamicola Zaitzev, 1908; f – H. (Lihelophorus) yangae sp. nov.; g–j – abdomen with spiracles containing air bubbles: g–h – H. (Lihelophorus) lamicola Zaitzev, 1908 (g – tracheal gills and its tracheal system; h – detail of tracheal branching around abdominal spiracle); i–j – H. (Lihelophorus) yangae sp. nov. (i – detail of tracheal system around abdominal spiracles; j – tracheal system of abdomen). Abbreviations: sp – spiracle; trg – tracheal gill." figureDoi="http://doi.org/10.5281/zenodo.5311800" httpUri="https://zenodo.org/record/5311800/files/figure.png" pageId="29" pageNumber="138">Figs 8a–b</figureCitation>
). 
<emphasis id="8DA7EA9B8D64284BFF915971738DFDFF" bold="true" box="[113,229,572,595]" italics="true" pageId="29" pageNumber="138">Mandibles</emphasis>
(
<figureCitation id="27E82A0C8D64284BFF1159717200FDFF" box="[241,360,572,595]" captionStart="Fig" captionStartId="25.[113,144,1437,1456]" captionTargetBox="[120,1085,227,1306]" captionTargetId="figure-13@25.[113,1091,219,1316]" captionTargetPageId="25" captionText="Fig. 13. Head appendages of larvae of Lihelophorus and Helophorus s.str. a–d – mandibles in dorsal view: a – H. (H.) liguricus Angus, 1970 (left mandible); b – H. (Lihelophorus) yangae sp. nov. (left mandible); c–d – H. (Lihelophorus) lamicola Zaitzev, 1908 (c – left mandible; d – right mandible). e–f – left antenna in dorsal view: e – H. (H.) liguricus Angus, 1970; f – H. (Lihelophorus) yangae sp. nov. g–j – labium: g–h – H. (Lihelophorus) yangae sp. nov. (g – ventral; h – dorsal); i–j – H. (Lihelophorus) lamicola Zaitzev, 1908 (i – ventral; j – dorsal).Abbreviations: mp – mandibular penicillus; rt – retinacular tuft." figureDoi="http://doi.org/10.5281/zenodo.5311814" httpUri="https://zenodo.org/record/5311814/files/figure.png" pageId="29" pageNumber="138">Figs 13c–d</figureCitation>
) slightly asymmetrical, with falcate apical portion. Retinaculum with two teeth; distal tooth large, bearing dense tuft of long cuticular spines; basal tooth small, with numerous cuticular spines in left mandible and few projections in right one. Basal inner face projecting into a small lobe, with a basal field of fine spine-like cuticular projections. 
<emphasis id="8DA7EA9B8D64284BFF9159F173AFFD7F" bold="true" box="[113,199,700,723]" italics="true" pageId="29" pageNumber="138">Maxilla</emphasis>
(
<figureCitation id="27E82A0C8D64284BFF3559F17223FD7F" box="[213,331,700,723]" captionStart="Fig" captionStartId="26.[151,182,1513,1532]" captionTargetBox="[157,1119,224,1369]" captionTargetId="figure-10@26.[151,1126,216,1372]" captionTargetPageId="26" captionText="Fig. 14. Head appendages of larvae of Lihelophorus and Helophorus s.str. a–b – labium of H. (H.) liguricus Angus, 1970 (a – ventral; b – dorsal). c–h – maxilla: a–d – H. (H.) liguricus (c – ventral; d – dorsal); e–f – H. (Lihelophorus) yangae sp. nov. (e – ventral; f – dorsal); g–h – H. (Lihelophorus) lamicola Zaitzev, 1908 (g – ventral; h – dorsal)." figureDoi="http://doi.org/10.5281/zenodo.5311816" httpUri="https://zenodo.org/record/5311816/files/figure.png" pageId="29" pageNumber="138">Figs 14e–f</figureCitation>
). Stipes ca. as long as palpifer, sinuate laterally, mesal face with a series of irregular cuticular spines, lateral face with small blunt cuticular spines; palpifer slightly longer than stipes. 
<emphasis id="8DA7EA9B8D64284BFEDE59B172FCFCBF" bold="true" box="[318,404,764,787]" italics="true" pageId="29" pageNumber="138">Labium</emphasis>
(
<figureCitation id="27E82A0C8D64284BFE4359B1717DFCBF" box="[419,533,764,787]" captionStart="Fig" captionStartId="25.[113,144,1437,1456]" captionTargetBox="[120,1085,227,1306]" captionTargetId="figure-13@25.[113,1091,219,1316]" captionTargetPageId="25" captionText="Fig. 13. Head appendages of larvae of Lihelophorus and Helophorus s.str. a–d – mandibles in dorsal view: a – H. (H.) liguricus Angus, 1970 (left mandible); b – H. (Lihelophorus) yangae sp. nov. (left mandible); c–d – H. (Lihelophorus) lamicola Zaitzev, 1908 (c – left mandible; d – right mandible). e–f – left antenna in dorsal view: e – H. (H.) liguricus Angus, 1970; f – H. (Lihelophorus) yangae sp. nov. g–j – labium: g–h – H. (Lihelophorus) yangae sp. nov. (g – ventral; h – dorsal); i–j – H. (Lihelophorus) lamicola Zaitzev, 1908 (i – ventral; j – dorsal).Abbreviations: mp – mandibular penicillus; rt – retinacular tuft." figureDoi="http://doi.org/10.5281/zenodo.5311814" httpUri="https://zenodo.org/record/5311814/files/figure.png" pageId="29" pageNumber="138">Figs 13i–j</figureCitation>
). Submentum completely fused with parietale, submental suture absent. Mentum without cuticular spines on dorsal surface. 
<emphasis id="8DA7EA9B8D64284BFC97585170A0FC9F" bold="true" box="[887,968,796,819]" italics="true" pageId="29" pageNumber="138">Thorax</emphasis>
. Legs (
<figureCitation id="27E82A0C8D64284BFBF5585173F2FCFF" captionStart="Fig" captionStartId="27.[113,144,1395,1414]" captionTargetBox="[122,1075,219,1349]" captionTargetId="figure-13@27.[113,1088,216,1363]" captionTargetPageId="27" captionText="Fig. 15. Thorax and abdomen of Lihelophorus and Helophorus s.str. a–d – metathoracic legs (a – H. (H.) liguricus Angus, 1970, frontal view; b–c – H. (Lihelophorus) yangae sp. nov.(b – frontal; c – posterior); d – H. (Lihelophorus) lamicola Zaitzev, 1908). e–f – thorax and first abdominal segments of H. (Lihelophorus) yangae sp. nov. (e – lateral view; f – dorsal view). g–h – abdominal segments (dorsal and ventral views): g – H. (Lihelophorus) yangae sp. nov.; h – H. (Lihelophorus) lamicola Zaitzev, 1908. i–j – abdominal segment 9 and urogomphi: i – H. (H.) liguricus Angus, 1970; j – H. (Lihelophorus) yangae sp. nov.Abbreviations:abd1 – abdominal segment 1; msth – mesothorax; mtth – metathorax; pth – prothorax; sp – spiracle." figureDoi="http://doi.org/10.5281/zenodo.5311820" httpUri="https://zenodo.org/record/5311820/files/figure.png" pageId="29" pageNumber="138">Fig. 15d</figureCitation>
) very long, 5-segmented; trochanter ca. half as long as femur; tibiotarsus cylindrical, slightly longer than trochanter and femur combined. Claw ca. as long as tibiotarsus, slightly bent ventrad. 
<emphasis id="8DA7EA9B8D64284BFEE358317204FC3F" bold="true" box="[259,364,892,915]" italics="true" pageId="29" pageNumber="138">Abdomen</emphasis>
(
<quantity id="782B9B6C8D64284BFE9B583172FEFC3F" box="[379,406,892,915]" metricMagnitude="-3" metricUnit="kg" metricValue="8.0" pageId="29" pageNumber="138" unit="g" value="8.0">8g</quantity>
–h, 15h). Segments 1–8 each with a pair of long finger-like projections laterally below spiracles, bearing lateral sclerite on apex; each projection with trachea connected to tracheal branch connecting main tracheal trunk with spiracle lying above base of the particular projection.
</paragraph>
<paragraph id="BF6C36898D64284BFF9158B1719BFB1F" blockId="29.[113,1090,220,1203]" pageId="29" pageNumber="138">
<emphasis id="8DA7EA9B8D64284BFF9158B17228FBBF" bold="true" box="[113,320,1020,1043]" pageId="29" pageNumber="138">
Chaetotaxy. 
<emphasis id="8DA7EA9B8D64284BFF1F58B17228FBBF" bold="true" box="[255,320,1020,1043]" italics="true" pageId="29" pageNumber="138">Head.</emphasis>
</emphasis>
Frontal pores FR2 closer to each other than setae FR3; setae FR8 widely separated from each other; parietal setae PA7, PA12, and PA14 much longer than in 
<taxonomicName id="78D34D0A8D64284BFC335F517751FB9F" authorityName="Angus &amp; Jia &amp; Chen &amp; Zhang &amp; Vondráček &amp; Fikáček &amp; Sw" authorityYear="2016" box="[979,1081,1052,1075]" class="Insecta" family="Helophoridae" genus="Helophorus" kingdom="Animalia" order="Coleoptera" pageId="29" pageNumber="138" phylum="Arthropoda" rank="species" species="yangae">
<emphasis id="8DA7EA9B8D64284BFC335F517751FB9F" box="[979,1081,1052,1075]" italics="true" pageId="29" pageNumber="138">L. yangae</emphasis>
</taxonomicName>
. Labial setae LA3 very long. Maxillary setae 
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5 and 
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6 of the same length, both rather long; setae 
<collectingCountry id="C7C476198D64284BFF0E5F117274FBDF" box="[238,284,1116,1139]" name="Mexico" pageId="29" pageNumber="138">MX</collectingCountry>
8–11 very long, situated along the whole length of stipes; apical maxillary palpomere with multiple additional digitiform sensilla. 
<emphasis id="8DA7EA9B8D64284BFD365F317057FB3F" bold="true" box="[726,831,1148,1171]" italics="true" pageId="29" pageNumber="138">Abdomen</emphasis>
with lateral abdominal sclerite (situated on the top of tracheal gill) with three setae.
</paragraph>
</subSubSection>
</treatment>
</document>