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<document id="3C92BFAE5AC5E66F8BD793BA208AC5A6" ID-CLB-Dataset="121760" ID-DOI="10.1093/zoolinnean/zlab105" ID-GBIF-Dataset="83d946c2-fa80-4948-9160-16625c240886" ID-ISSN="0024-4082" ID-Zenodo-Dep="6994599" ID-ZooBank="C06A1F94-AF08-4A21-B1F3-A0865FB1A8DF" IM.bibliography_approvedBy="felipe" IM.materialsCitations_approvedBy="felipe" IM.metadata_approvedBy="felipe" IM.tables_requiresApprovalFor="existingObjects,plazi" IM.taxonomicNames_approvedBy="felipe" checkinTime="1660324512590" checkinUser="plazi" docAuthor="Sites, Robert W." docDate="2022" docId="E25E878FFF9CFFC0FC5481760482FAC9" docLanguage="en" docName="zlab105.pdf" docOrigin="Zoological Journal of the Linnean Society 195" docStyle="DocumentStyle:36B3BD6A90C22AB4F7F465C853188CC8.7:ZoolJLinnSoc.2017-2023.journal_article" docStyleId="36B3BD6A90C22AB4F7F465C853188CC8" docStyleName="ZoolJLinnSoc.2017-2023.journal_article" docStyleVersion="7" docTitle="Laccocorinae STAL 1876" docType="treatment" docVersion="7" lastPageNumber="1264" masterDocId="1E67FFF7FF8DFFD3FFC487270740FFAD" masterDocTitle="Phylogeny and revised classification of the saucer bugs (Hemiptera: Nepomorpha: Naucoridae)" masterLastPageNumber="1286" masterPageNumber="1245" pageNumber="1262" updateTime="1699366499196" updateUser="ExternalLinkService">
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<mods:title id="BAAB3082B9885634641B7FC0260AFEEB">Phylogeny and revised classification of the saucer bugs (Hemiptera: Nepomorpha: Naucoridae)</mods:title>
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<treatment id="E25E878FFF9CFFC0FC5481760482FAC9" ID-DOI="http://doi.org/10.5281/zenodo.6994701" ID-GBIF-Taxon="199315258" ID-Zenodo-Dep="6994701" LSID="urn:lsid:plazi:treatment:E25E878FFF9CFFC0FC5481760482FAC9" httpUri="http://treatment.plazi.org/id/E25E878FFF9CFFC0FC5481760482FAC9" lastPageId="19" lastPageNumber="1264" pageId="17" pageNumber="1262">
<subSubSection id="22ED6512FF9CFFC2FC548176020DF9C7" box="[912,1357,1617,1642]" pageId="17" pageNumber="1262" type="nomenclature">
<paragraph id="6A483699FF9CFFC2FC548176020DF9C7" blockId="17.[912,1357,1617,1642]" box="[912,1357,1617,1642]" pageId="17" pageNumber="1262">
SUBFAMILY
<taxonomicName id="ADF74D1AFF9CFFC2FBD38176020DF9C7" ID-CoL="JS3" authority="STAL, 1876" authorityName="STAL" authorityYear="1876" box="[1047,1357,1617,1642]" class="Insecta" family="Naucoridae" kingdom="Animalia" order="Hemiptera" pageId="17" pageNumber="1262" phylum="Arthropoda" rank="subFamily" status="stat. nov." subFamily="Laccocorinae">
LACCOCORINAE
<bibRefCitation id="0E664B68FF9CFFC2FB0A8176020DF9C7" author="Stal C" box="[1230,1357,1617,1642]" pageId="17" pageNumber="1262" pagination="1 - 167" refId="ref25070" refString="Stal C. 1876. Enumeratio Hemipterorum. Bidrag till en forteckning ofver alla hittills kanda. Hemiptera, jemte systematiska meddelanden. Kongliga Svenska VetenskapsAkademiens Handlingar 14: 1 - 167." type="journal article" year="1876">STÅL, 1876</bibRefCitation>
</taxonomicName>
</paragraph>
</subSubSection>
<subSubSection id="22ED6512FF9CFFC2FC43815A0216F93B" box="[903,1366,1661,1686]" pageId="17" pageNumber="1262" type="description">
<paragraph id="6A483699FF9CFFC2FC43815A0216F93B" blockId="17.[903,1366,1661,1686]" box="[903,1366,1661,1686]" pageId="17" pageNumber="1262">
<figureCitation id="F2CC2A1CFF9CFFC2FC43815A04A1F939" box="[903,993,1661,1686]" captionStart="Figure 3" captionStartId="8.[145,230,1468,1490]" captionTargetBox="[172,733,195,1428]" captionTargetId="figure-532@8.[172,733,195,1428]" captionTargetPageId="8" captionText="Figure 3. Mouthpart positions: recessed rostrum of Cheirochela thailandana (a); lateral views of Laccocoris spurcus (b) and Australambrysus crenulatus (c). White arrows indicate positions of labrum with respect to the functional anterior margin of the head." figureDoi="http://doi.org/10.5281/zenodo.6994613" httpUri="https://zenodo.org/record/6994613/files/figure.png" pageId="17" pageNumber="1262">FIGS 3B</figureCitation>
,
<figureCitation id="F2CC2A1CFF9CFFC2FC2A815A0358F93B" box="[1006,1048,1661,1686]" captionStart-0="Figure 4" captionStart-1="Figure 5" captionStart-2="Figure 6" captionStartId-0="9.[163,249,1379,1401]" captionStartId-1="10.[145,223,735,757]" captionStartId-2="10.[145,228,1329,1351]" captionTargetBox-0="[163,781,195,1339]" captionTargetBox-1="[145,1426,195,695]" captionTargetBox-2="[216,689,828,1289]" captionTargetId-0="figure-575@9.[163,781,195,1339]" captionTargetId-1="figure-384@10.[145,1426,195,695]" captionTargetId-2="figure-415@10.[216,689,828,1289]" captionTargetPageId-0="9" captionTargetPageId-1="10" captionTargetPageId-2="10" captionText-0="Figure 4. Foretibiae and foretarsi of Laccocorinae: Ctenipocoris asiaticus (a), Heleolaccocoris ovatus (b), Namtokocoris dalanta (c)." captionText-1="Figure 5. Heleocoris elongatus mesofemur (a), metafemur (b), Tsingala gen. nov. sp. C mesofemur (c), metafemur (d). White arrows indicate posteroventral spine rows, black arrows indicate anteroventral spine rows." captionText-2="Figure 6. Laccocoris spurcus showing the transverse sulcus at the base of the labrum, which is characteristic of most Laccocorini." figureDoi-0="http://doi.org/10.5281/zenodo.6994615" figureDoi-1="http://doi.org/10.5281/zenodo.6994617" figureDoi-2="http://doi.org/10.5281/zenodo.6994621" httpUri-0="https://zenodo.org/record/6994615/files/figure.png" httpUri-1="https://zenodo.org/record/6994617/files/figure.png" httpUri-2="https://zenodo.org/record/6994621/files/figure.png" pageId="17" pageNumber="1262">
4–
<geoCoordinate id="0FC3505EFF9CFFC2FBCF815A0358F93B" box="[1035,1048,1661,1686]" degrees="6, 10" direction="east" orientation="longitude" pageId="17" pageNumber="1262" precision="555" value="6.1">6</geoCoordinate>
</figureCitation>
,
<figureCitation id="F2CC2A1CFF9CFFC2FBE1815A0310F939" box="[1061,1104,1661,1686]" captionStart="Figure 10" captionStartId="14.[144,226,1765,1787]" captionTargetBox="[144,1424,676,1726]" captionTargetId="figure-25@14.[144,1424,676,1726]" captionTargetPageId="14" captionText="Figure 10. Habitus examples of each subfamily: Ambrysinae: Ambrysus signoreti (a), Cheirochelinae: Cheirochela thailandana (b), Cryphocricinae: Cryphocricos hungerfordi (c), Ilyocorinae: Ilyocoris cimicoides (d), Laccocorinae: Laccocoris limicola (e), Limnocorinae: Limnocoris inornatus (f), Macrocorinae subfam. nov.: Macrocoris flavicollis (g), Naucorinae: Naucoris scutellaris (Thailand) (h)." figureDoi="http://doi.org/10.5281/zenodo.6994631" httpUri="https://zenodo.org/record/6994631/files/figure.png" pageId="17" pageNumber="1262">10E</figureCitation>
,
<figureCitation id="F2CC2A1CFF9CFFC2FB99815A03C8F939" box="[1117,1160,1661,1686]" captionStart="Figure 11" captionStartId="15.[162,240,1540,1562]" captionTargetBox="[163,1443,194,1500]" captionTargetId="figure-103@15.[163,1443,194,1500]" captionTargetPageId="15" captionText="Figure 11. Examples of male genital capsule with pygophore, aedeagus and parameres (terga IX and X removed) for each subfamily: Ambrysinae: Ambrysus lunatus (a), Cheirochelinae: Coptocatus kinabalu (b), Cryphocricinae: Cryphocricos hungerfordi (c), Ilyocorinae:Ilyocoris cimicoides (d), Laccocorinae: Laccocoris spurcus (e), Limnocorinae: Limnocoris ovatulus (f), Macrocorinae subfam. nov.: Macrocoris flavicollis (g), Naucorinae: Afronaucoris gen. nov.: Afronaucoris madagascariensis (h)." figureDoi="http://doi.org/10.5281/zenodo.6994635" httpUri="https://zenodo.org/record/6994635/files/figure.png" pageId="17" pageNumber="1262">11E</figureCitation>
,
<figureCitation id="F2CC2A1CFF9CFFC2FB51815A03F1F93B" box="[1173,1201,1661,1686]" captionStart="Figure 12" captionStartId="17.[163,243,870,892]" captionTargetBox="[163,1443,195,830]" captionTargetId="figure-534@17.[163,1443,195,830]" captionTargetPageId="17" captionText="Figure 12. Mesotibia and mesotarsus of male and female Heleolaccocoris ovatus showing a sexually dimorphic ventral tomentose pad of hairs." figureDoi="http://doi.org/10.5281/zenodo.6994637" httpUri="https://zenodo.org/record/6994637/files/figure.png" pageId="17" pageNumber="1262">12</figureCitation>
,
<figureCitation id="F2CC2A1CFF9CFFC2FB7A815A039AF93B" box="[1214,1242,1661,1686]" captionStart="Figure 14" captionStartId="19.[163,244,765,787]" captionTargetBox="[163,1442,195,726]" captionTargetId="figure-559@19.[163,1442,195,726]" captionTargetPageId="19" captionText="Figure 14. Revised generic representation of the former genus Heleocoris. Dorsal habitus of Heleocoris vicinus (a), Laccocoris minisculus (b), Heleolaccocoris ovatus (c), Tsingala humeralis (d)." figureDoi="http://doi.org/10.5281/zenodo.6994645" httpUri="https://zenodo.org/record/6994645/files/figure.png" pageId="17" pageNumber="1262">14</figureCitation>
,
<figureCitation id="F2CC2A1CFF9CFFC2FB2C815A0244F93B" box="[1256,1284,1661,1686]" captionStart="Figure 15" captionStartId="20.[145,226,1055,1077]" captionTargetBox="[145,1425,195,1015]" captionTargetId="figure-409@20.[145,1425,195,1015]" captionTargetPageId="20" captionText="Figure 15. Revised generic representation of the former genus Laccocoris. Dorsal habitus of Laccocoris spurcus (a), Heleolaccocoris horvathi (b)." figureDoi="http://doi.org/10.5281/zenodo.6994647" httpUri="https://zenodo.org/record/6994647/files/figure.png" pageId="17" pageNumber="1262">15</figureCitation>
,
<figureCitation id="F2CC2A1CFF9CFFC2FAD5815A026DF93B" box="[1297,1325,1661,1686]" captionStart="Figure 20" captionStartId="28.[146,226,1107,1129]" captionTargetBox="[145,1425,195,1067]" captionTargetId="figure-347@28.[145,1425,195,1067]" captionTargetPageId="28" captionText="Figure 20. Tsingala humeralis male: dorsal habitus (a), genital capsule with terga IX and X removed (b), left and right parameres (c). Sizes are not proportional among figures." figureDoi="http://doi.org/10.5281/zenodo.6994662" httpUri="https://zenodo.org/record/6994662/files/figure.png" pageId="17" pageNumber="1262">20</figureCitation>
,
<figureCitation id="F2CC2A1CFF9CFFC2FAFE815A0216F93B" box="[1338,1366,1661,1686]" captionStart="Figure 21" captionStartId="29.[164,242,1876,1898]" captionTargetBox="[163,1443,921,1837]" captionTargetId="figure-364@29.[163,1443,921,1837]" captionTargetPageId="29" captionText="Figure 21. Heleolaccocoris horvathi male: dorsal habitus (a), genital capsule with terga IX and X removed (b)." figureDoi="http://doi.org/10.5281/zenodo.6994664" httpUri="https://zenodo.org/record/6994664/files/figure.png" pageId="17" pageNumber="1262">21</figureCitation>
</paragraph>
</subSubSection>
<subSubSection id="22ED6512FF9CFFC2FCFF8194026CF965" box="[827,1324,1714,1736]" pageId="17" pageNumber="1262" type="type_taxon">
<paragraph id="6A483699FF9CFFC2FCFF8194026CF965" blockId="17.[827,1324,1714,1736]" box="[827,1324,1714,1736]" pageId="17" pageNumber="1262">
<emphasis id="5883EA8BFF9CFFC2FCFF8194048BF965" box="[827,971,1715,1736]" italics="true" pageId="17" pageNumber="1262">
<typeStatus id="B54C883BFF9CFFC2FCFF81940430F965" box="[827,880,1715,1736]" pageId="17" pageNumber="1262">Type</typeStatus>
species:
</emphasis>
<taxonomicName id="ADF74D1AFF9CFFC2FC1381940265F965" ID-CoL="3RLYX" authority="Stal, 1856" authorityName="Stal" authorityYear="1856" box="[983,1317,1714,1736]" class="Insecta" family="Naucoridae" genus="Laccocoris" kingdom="Animalia" order="Hemiptera" pageId="17" pageNumber="1262" phylum="Arthropoda" rank="species" species="spurcus">
<emphasis id="5883EA8BFF9CFFC2FC13819403ECF965" box="[983,1196,1715,1736]" italics="true" pageId="17" pageNumber="1262">Laccocoris spurcus</emphasis>
<bibRefCitation id="0E664B68FF9CFFC2FB7781950265F965" author="Stal C" box="[1203,1317,1714,1736]" pageId="17" pageNumber="1262" pagination="193 - 199" refId="ref24930" refString="Stal C. 1856. Hemiptera samlade af Victorin i Caplandet. Ofversigt af Kongliga Svenska Vetenskaps-Akademiens Forhandlingar 13: 193 - 199." type="journal article" year="1856">Stål, 1856</bibRefCitation>
</taxonomicName>
.
</paragraph>
</subSubSection>
<subSubSection id="22ED6512FF9CFFC0FCFF81D5056CF929" lastPageId="19" lastPageNumber="1264" pageId="17" pageNumber="1262" type="discussion">
<paragraph id="6A483699FF9CFFC1FCFF81D506D0FA25" blockId="17.[827,1443,1777,1891]" lastBlockId="18.[145,761,1241,1416]" lastPageId="18" lastPageNumber="1263" pageId="17" pageNumber="1262">
<emphasis id="5883EA8BFF9CFFC2FCFF81D50356F8AB" box="[827,1046,1777,1799]" italics="true" pageId="17" pageNumber="1262">Taxonomic history:</emphasis>
<bibRefCitation id="0E664B68FF9CFFC2FBE781D603E3F8AA" author="Stal C" box="[1059,1187,1777,1799]" pageId="17" pageNumber="1262" pagination="1 - 167" refId="ref25070" refString="Stal C. 1876. Enumeratio Hemipterorum. Bidrag till en forteckning ofver alla hittills kanda. Hemiptera, jemte systematiska meddelanden. Kongliga Svenska VetenskapsAkademiens Handlingar 14: 1 - 167." type="journal article" year="1876">Stål (1876)</bibRefCitation>
proposed the division Laccocoraria to contain two genera,
<taxonomicName id="ADF74D1AFF9CFFC2FB378037022DF888" authorityName="Stal" authorityYear="1876" box="[1267,1389,1808,1829]" class="Insecta" family="Naucoridae" genus="Heleocoris" kingdom="Animalia" order="Hemiptera" pageId="17" pageNumber="1262" phylum="Arthropoda" rank="genus">
<emphasis id="5883EA8BFF9CFFC2FB378037022DF888" box="[1267,1389,1808,1829]" italics="true" pageId="17" pageNumber="1262">Heleocoris</emphasis>
</taxonomicName>
and
<taxonomicName id="ADF74D1AFF9CFFC2FCFF800804FAF8E9" authorityName="Stal" authorityYear="1856" box="[827,954,1839,1860]" class="Insecta" family="Naucoridae" genus="Laccocoris" kingdom="Animalia" order="Hemiptera" pageId="17" pageNumber="1262" phylum="Arthropoda" rank="genus">
<emphasis id="5883EA8BFF9CFFC2FCFF800804FAF8E9" box="[827,954,1839,1860]" italics="true" pageId="17" pageNumber="1262">Laccocoris</emphasis>
</taxonomicName>
, that he described.
<bibRefCitation id="0E664B68FF9CFFC2FB70800802E3F8E8" author="Montandon AL" box="[1204,1443,1839,1861]" pageId="17" pageNumber="1262" pagination="435 - 454" refId="ref23009" refString="Montandon AL. 1897 b. Hemiptera Cryptocerata. Fam. Naucoridae. Sous fam. Laccocorinae. Verhandlungen der Kaiserlich-Koniglichen Zoologisch-Botanischen Gesellschaft in Wien 47: 435 - 454." type="journal article" year="1897">Montandon (1897b)</bibRefCitation>
e l e v a t e d L a c c o c o r a r i a t o s u b f a m i l y s t a t u s a s
<taxonomicName id="ADF74D1AFF9FFFC1FF5583FE0673FB42" authorityName="STAL" authorityYear="1876" box="[145,307,1241,1263]" class="Insecta" family="Naucoridae" kingdom="Animalia" order="Hemiptera" pageId="18" pageNumber="1285" phylum="Arthropoda" rank="subFamily" subFamily="Laccocorinae">Laccocorinae</taxonomicName>
.
<bibRefCitation id="0E664B68FF9FFFC1FE8583FE06A0FB43" author="Popov YA" box="[321,480,1241,1262]" pageId="18" pageNumber="1263" pagination="93 - 98" refId="ref23866" refString="Popov YA. 1970. Notes on the classification of the recent Naucoridae (Heteroptera: Nepomorpha). Bulletin of the Polish Academy of Sciences (Biological Sciences) 18: 93 - 98." type="journal article" year="1970">Popov (1970)</bibRefCitation>
considered this group as a tribe-level taxon,
<taxonomicName id="ADF74D1AFF9FFFC1FE5F83DF0563FAA3" authorityName="POPOV" authorityYear="1970" box="[411,547,1272,1294]" class="Insecta" family="Naucoridae" kingdom="Animalia" order="Hemiptera" pageId="18" pageNumber="1266" phylum="Arthropoda" rank="tribe" tribe="Laccocorini">Laccocorini</taxonomicName>
, within subfamily
<taxonomicName id="ADF74D1AFF9FFFC1FF5582310656FA81" authorityName="LEACH" authorityYear="1815" box="[145,278,1302,1324]" class="Insecta" family="Naucoridae" kingdom="Animalia" order="Hemiptera" pageId="18" pageNumber="1263" phylum="Arthropoda" rank="subFamily" subFamily="Naucorinae">Naucorinae</taxonomicName>
, but this classification scheme has not been followed by other specialists. Presently, ten genera are recognized in the subfamily, with no suprageneric organization as tribes.
</paragraph>
<caption id="3E886611FF9FFFC1FF5583060454FB00" ID-DOI="http://doi.org/10.5281/zenodo.6994641" ID-Zenodo-Dep="6994641" httpUri="https://zenodo.org/record/6994641/files/figure.png" pageId="18" pageNumber="1263" startId="18.[145,223,1057,1079]" targetBox="[149,1425,195,1017]" targetPageId="18">
<paragraph id="6A483699FF9FFFC1FF5583060454FB00" blockId="18.[145,1425,1057,1197]" pageId="18" pageNumber="1263">
<emphasis id="5883EA8BFF9FFFC1FF5583060645FB9B" bold="true" box="[145,261,1057,1079]" pageId="18" pageNumber="1263">Figure 13.</emphasis>
Detail of Most Probable Phylogeny dendogram of
<taxonomicName id="ADF74D1AFF9FFFC1FCCA830604D8FB9A" authorityName="STAL" authorityYear="1876" box="[782,920,1057,1079]" class="Insecta" family="Naucoridae" kingdom="Animalia" order="Hemiptera" pageId="18" pageNumber="1285" phylum="Arthropoda" rank="subFamily" subFamily="Laccocorinae">Laccocorinae</taxonomicName>
. The red-coloured clade includes the Madagascar genera
<taxonomicName id="ADF74D1AFF9FFFC1FF1983180613FBF8" authorityName="Montandon" authorityYear="1897" box="[221,339,1087,1109]" class="Insecta" family="Naucoridae" genus="Temnocoris" kingdom="Animalia" order="Hemiptera" pageId="18" pageNumber="1263" phylum="Arthropoda" rank="genus">
<emphasis id="5883EA8BFF9FFFC1FF1983180613FBF8" box="[221,339,1087,1109]" italics="true" pageId="18" pageNumber="1263">Temnocoris</emphasis>
</taxonomicName>
and
<taxonomicName id="ADF74D1AFF9FFFC1FE40831806A1FBF8" authority="Sites, 2022" authorityName="Sites" authorityYear="2022" box="[388,481,1087,1109]" class="Insecta" family="Naucoridae" genus="Tsingala" kingdom="Animalia" order="Hemiptera" pageId="18" pageNumber="1263" phylum="Arthropoda" rank="genus" status="gen. nov.">
<emphasis id="5883EA8BFF9FFFC1FE40831806A1FBF8" box="[388,481,1087,1109]" italics="true" pageId="18" pageNumber="1263">Tsingala</emphasis>
</taxonomicName>
<taxonomicNameLabel id="43B057F0FF9FFFC1FE218318057EFBF8" box="[485,574,1087,1109]" pageId="18" pageNumber="1263" rank="genus">gen. nov.</taxonomicNameLabel>
(formerly
<taxonomicName id="ADF74D1AFF9FFFC1FD6383180452FBF8" authorityName="Stal" authorityYear="1876" box="[679,786,1087,1109]" class="Insecta" family="Naucoridae" genus="Heleocoris" kingdom="Animalia" order="Hemiptera" pageId="18" pageNumber="1263" phylum="Arthropoda" rank="genus">
<emphasis id="5883EA8BFF9FFFC1FD6383180452FBF8" box="[679,786,1087,1109]" italics="true" pageId="18" pageNumber="1263">Heleocoris</emphasis>
</taxonomicName>
). The blue-coloured clade represents
<taxonomicName id="ADF74D1AFF9FFFC1FB4B8318026EFBF8" authority="Sites, 2022" authorityName="Sites" authorityYear="2022" box="[1167,1326,1087,1109]" class="Insecta" family="Naucoridae" genus="Heleolaccocoris" kingdom="Animalia" order="Hemiptera" pageId="18" pageNumber="1263" phylum="Arthropoda" rank="genus" status="gen. nov.">
<emphasis id="5883EA8BFF9FFFC1FB4B8318026EFBF8" box="[1167,1326,1087,1109]" italics="true" pageId="18" pageNumber="1263">Heleolaccocoris</emphasis>
</taxonomicName>
<taxonomicNameLabel id="43B057F0FF9FFFC1FAF7831802CEFBF8" box="[1331,1422,1087,1109]" pageId="18" pageNumber="1263" rank="genus">gen. nov.</taxonomicNameLabel>
, which comprises species of the former genera
<taxonomicName id="ADF74D1AFF9FFFC1FDAC837B0593FBDF" authorityName="Stal" authorityYear="1876" box="[616,723,1116,1138]" class="Insecta" family="Naucoridae" genus="Heleocoris" kingdom="Animalia" order="Hemiptera" pageId="18" pageNumber="1263" phylum="Arthropoda" rank="genus">
<emphasis id="5883EA8BFF9FFFC1FDAC837B0593FBDF" box="[616,723,1116,1138]" italics="true" pageId="18" pageNumber="1263">Heleocoris</emphasis>
</taxonomicName>
and
<taxonomicName id="ADF74D1AFF9FFFC1FCC1837B0432FBDF" authorityName="Stal" authorityYear="1856" box="[773,882,1116,1138]" class="Insecta" family="Naucoridae" genus="Laccocoris" kingdom="Animalia" order="Hemiptera" pageId="18" pageNumber="1263" phylum="Arthropoda" rank="genus">
<emphasis id="5883EA8BFF9FFFC1FCC1837B0432FBDF" box="[773,882,1116,1138]" italics="true" pageId="18" pageNumber="1263">Laccocoris</emphasis>
</taxonomicName>
. All species of
<taxonomicName id="ADF74D1AFF9FFFC1FBC2837B0323FBDF" authority="Sites, 2022" authorityName="Sites" authorityYear="2022" box="[1030,1123,1116,1138]" class="Insecta" family="Naucoridae" genus="Tsingala" kingdom="Animalia" order="Hemiptera" pageId="18" pageNumber="1263" phylum="Arthropoda" rank="genus" status="gen. nov.">
<emphasis id="5883EA8BFF9FFFC1FBC2837B0323FBDF" box="[1030,1123,1116,1138]" italics="true" pageId="18" pageNumber="1263">Tsingala</emphasis>
</taxonomicName>
<taxonomicNameLabel id="43B057F0FF9FFFC1FBAC837B0381FBDF" box="[1128,1217,1116,1138]" pageId="18" pageNumber="1263" rank="genus">gen. nov.</taxonomicNameLabel>
and
<taxonomicName id="ADF74D1AFF9FFFC1FB36837B02D1FBDF" authority="Sites, 2022" authorityName="Sites" authorityYear="2022" box="[1266,1425,1116,1138]" class="Insecta" family="Naucoridae" genus="Heleolaccocoris" kingdom="Animalia" order="Hemiptera" pageId="18" pageNumber="1263" phylum="Arthropoda" rank="genus" status="gen. nov.">
<emphasis id="5883EA8BFF9FFFC1FB36837B02D1FBDF" box="[1266,1425,1116,1138]" italics="true" pageId="18" pageNumber="1263">Heleolaccocoris</emphasis>
</taxonomicName>
<taxonomicNameLabel id="43B057F0FF9FFFC1FF55835E07AAFB22" box="[145,234,1145,1167]" pageId="18" pageNumber="1263" rank="genus">gen. nov.</taxonomicNameLabel>
are new combinations. Geographic associations for major clades within and among tribes are indicated. The red box in the inset shows the part of Figure 9 depicted in this figure.
</paragraph>
</caption>
<paragraph id="6A483699FF9FFFC1FF558296057CF8B5" blockId="18.[145,761,1457,1908]" pageId="18" pageNumber="1263">
<emphasis id="5883EA8BFF9FFFC1FF5582960628FA6A" box="[145,360,1457,1479]" italics="true" pageId="18" pageNumber="1263">Revised taxonomy:</emphasis>
<taxonomicName id="ADF74D1AFF9FFFC1FEB18296054CFA6A" authorityName="STAL" authorityYear="1876" box="[373,524,1457,1479]" class="Insecta" family="Naucoridae" kingdom="Animalia" order="Hemiptera" pageId="18" pageNumber="1285" phylum="Arthropoda" rank="subFamily" subFamily="Laccocorinae">Laccocorinae</taxonomicName>
remains unchanged with respect to constituent species, although synonymies, new genera and tribes are proposed here. In my Most Probable Phylogeny dendrogram (
<figureCitation id="F2CC2A1CFF9FFFC1FD6D812A05AAF98F" box="[681,746,1549,1571]" captionStart="Figure 9" captionStartId="12.[145,223,1678,1700]" captionTargetBox="[308,1266,197,1636]" captionTargetId="figure-82@12.[302,1268,195,1638]" captionTargetPageId="12" captionText="Figure 9. Most Probable Phylogeny dendogram based on Bayesian inference with substitutions from maximum likelihood for Laccocorinae (except Ctenipocorini trib. nov. and Idiocarini trib. nov.). Closer views of Laccocorinae and Naucorinae individually can be seen in Figures 13 and 18, respectively." figureDoi="http://doi.org/10.5281/zenodo.6994629" httpUri="https://zenodo.org/record/6994629/files/figure.png" pageId="18" pageNumber="1263">Fig. 9</figureCitation>
), I substituted the ML topology of the clade sister to
<taxonomicName id="ADF74D1AFF9FFFC1FF55816C065EF9CD" authorityName="Montandon" authorityYear="1897" box="[145,286,1611,1632]" class="Insecta" family="Naucoridae" genus="Ctenipocoris" kingdom="Animalia" order="Hemiptera" pageId="18" pageNumber="1263" phylum="Arthropoda" rank="genus">
<emphasis id="5883EA8BFF9FFFC1FF55816C065EF9CD" box="[145,286,1611,1632]" italics="true" pageId="18" pageNumber="1263">Ctenipocoris</emphasis>
</taxonomicName>
to represent what I consider to be the most plausible evolutionary relationships in the subfamily.
<taxonomicName id="ADF74D1AFF9FFFC1FF5581A0066DF930" authorityName="STAL" authorityYear="1876" box="[145,301,1671,1693]" class="Insecta" family="Naucoridae" kingdom="Animalia" order="Hemiptera" pageId="18" pageNumber="1285" phylum="Arthropoda" rank="subFamily" subFamily="Laccocorinae">Laccocorinae</taxonomicName>
is represented by four distinct major clades, which are herein recognized as tribe-level taxa (
<figureCitation id="F2CC2A1CFF9FFFC1FF5D81E207A9F977" box="[153,233,1733,1755]" captionStart="Figure 13" captionStartId="18.[145,223,1057,1079]" captionTargetBox="[149,1425,195,1017]" captionTargetId="figure-432@18.[145,1425,195,1017]" captionTargetPageId="18" captionText="Figure 13. Detail of Most Probable Phylogeny dendogram of Laccocorinae.The red-coloured clade includes the Madagascar genera Temnocoris and Tsingala gen. nov.(formerly Heleocoris). The blue-coloured clade represents Heleolaccocoris gen.nov., which comprises species of the former genera Heleocoris and Laccocoris. All species of Tsingala gen. nov. and Heleolaccocoris gen. nov. are new combinations. Geographic associations for major clades within and among tribes are indicated. The red box in the inset shows the part of Figure 9 depicted in this figure." figureDoi="http://doi.org/10.5281/zenodo.6994641" httpUri="https://zenodo.org/record/6994641/files/figure.png" pageId="18" pageNumber="1263">Fig. 13</figureCitation>
). Two of the predominant genera (
<taxonomicName id="ADF74D1AFF9FFFC1FD4581E205B9F977" authorityName="Stal" authorityYear="1876" box="[641,761,1733,1754]" class="Insecta" family="Naucoridae" genus="Heleocoris" kingdom="Animalia" order="Hemiptera" pageId="18" pageNumber="1263" phylum="Arthropoda" rank="genus">
<emphasis id="5883EA8BFF9FFFC1FD4581E205B9F977" box="[641,761,1733,1754]" italics="true" pageId="18" pageNumber="1263">Heleocoris</emphasis>
</taxonomicName>
and
<taxonomicName id="ADF74D1AFF9FFFC1FF0781C3067AF954" authorityName="Stal" authorityYear="1856" box="[195,314,1764,1785]" class="Insecta" family="Naucoridae" genus="Laccocoris" kingdom="Animalia" order="Hemiptera" pageId="18" pageNumber="1263" phylum="Arthropoda" rank="genus">
<emphasis id="5883EA8BFF9FFFC1FF0781C3067AF954" box="[195,314,1764,1785]" italics="true" pageId="18" pageNumber="1263">Laccocoris</emphasis>
</taxonomicName>
) in the subfamily are polyphyletic and occur in multiple clades in two tribes.
</paragraph>
<paragraph id="6A483699FF9FFFC0FF6D800606C7FC09" blockId="18.[145,761,1457,1908]" lastBlockId="19.[163,780,880,1668]" lastPageId="19" lastPageNumber="1264" pageId="18" pageNumber="1263">
Species of
<taxonomicName id="ADF74D1AFF9FFFC1FEE4800606D6F89B" authorityName="Stal" authorityYear="1876" box="[288,406,1825,1846]" class="Insecta" family="Naucoridae" genus="Heleocoris" kingdom="Animalia" order="Hemiptera" pageId="18" pageNumber="1263" phylum="Arthropoda" rank="genus">
<emphasis id="5883EA8BFF9FFFC1FEE4800606D6F89B" box="[288,406,1825,1846]" italics="true" pageId="18" pageNumber="1263">Heleocoris</emphasis>
</taxonomicName>
occur in three disparate clades (
<figureCitation id="F2CC2A1CFF9FFFC1FF5D801807ACF8F8" box="[153,236,1855,1877]" captionStart="Figure 13" captionStartId="18.[145,223,1057,1079]" captionTargetBox="[149,1425,195,1017]" captionTargetId="figure-432@18.[145,1425,195,1017]" captionTargetPageId="18" captionText="Figure 13. Detail of Most Probable Phylogeny dendogram of Laccocorinae.The red-coloured clade includes the Madagascar genera Temnocoris and Tsingala gen. nov.(formerly Heleocoris). The blue-coloured clade represents Heleolaccocoris gen.nov., which comprises species of the former genera Heleocoris and Laccocoris. All species of Tsingala gen. nov. and Heleolaccocoris gen. nov. are new combinations. Geographic associations for major clades within and among tribes are indicated. The red box in the inset shows the part of Figure 9 depicted in this figure." figureDoi="http://doi.org/10.5281/zenodo.6994641" httpUri="https://zenodo.org/record/6994641/files/figure.png" pageId="18" pageNumber="1263">Figs 13</figureCitation>
,
<figureCitation id="F2CC2A1CFF9FFFC1FF3C80670654F8F8" box="[248,276,1856,1877]" captionStart="Figure 14" captionStartId="19.[163,244,765,787]" captionTargetBox="[163,1442,195,726]" captionTargetId="figure-559@19.[163,1442,195,726]" captionTargetPageId="19" captionText="Figure 14. Revised generic representation of the former genus Heleocoris. Dorsal habitus of Heleocoris vicinus (a), Laccocoris minisculus (b), Heleolaccocoris ovatus (c), Tsingala humeralis (d)." figureDoi="http://doi.org/10.5281/zenodo.6994645" httpUri="https://zenodo.org/record/6994645/files/figure.png" pageId="18" pageNumber="1263">14</figureCitation>
). The
<typeStatus id="B54C883BFF9FFFC1FE9E806706CCF8F8" box="[346,396,1856,1877]" pageId="18" pageNumber="1263">type</typeStatus>
species is
<taxonomicName id="ADF74D1AFF9FFFC1FDC08018060DF8DE" authority="(Spinola, 1837)" baseAuthorityName="Spinola" baseAuthorityYear="1837" class="Insecta" family="Naucoridae" genus="Heleocoris" kingdom="Animalia" order="Hemiptera" pageId="18" pageNumber="1263" phylum="Arthropoda" rank="species" species="obliquatus">
<emphasis id="5883EA8BFF9FFFC1FDC0801805B8F8F9" box="[516,760,1855,1876]" italics="true" pageId="18" pageNumber="1263">Heleocoris obliquatus</emphasis>
(
<bibRefCitation id="0E664B68FF9FFFC1FF5E80790603F8DE" author="Spinola M" box="[154,323,1886,1908]" pageId="18" pageNumber="1263" refId="ref24867" refString="Spinola M. 1837. Essai sur les genres d'insectes appartenants a l'ordre des Hemipteres, Lin. ou Rhyngotes, Fab. et a la section des Heteropteres, Dufour. Chez Yves Graviers, Genes, 383 pp." type="book" year="1837">Spinola, 1837</bibRefCitation>
)
</taxonomicName>
from
<collectingCountry id="12E07609FF9FFFC1FE5F807906BDF8D9" box="[411,509,1886,1908]" name="India" pageId="18" pageNumber="1263">Bombay</collectingCountry>
(
<collectingCountry id="12E07609FF9FFFC1FDD480790539F8D9" box="[528,633,1886,1908]" name="India" pageId="18" pageNumber="1263">Mumbai</collectingCountry>
,
<collectingCountry id="12E07609FF9FFFC1FD4D8079058BF8D9" box="[649,715,1886,1908]" name="India" pageId="18" pageNumber="1263">India</collectingCountry>
) as designated by
<bibRefCitation id="0E664B68FF9FFFC1FC1583FF0311FB42" author="Stal C" box="[977,1105,1240,1263]" pageId="18" pageNumber="1263" pagination="1 - 167" refId="ref25070" refString="Stal C. 1876. Enumeratio Hemipterorum. Bidrag till en forteckning ofver alla hittills kanda. Hemiptera, jemte systematiska meddelanden. Kongliga Svenska VetenskapsAkademiens Handlingar 14: 1 - 167." type="journal article" year="1876">Stål (1876)</bibRefCitation>
, who based his designation on a specimen in the collection of Signoret.
<bibRefCitation id="0E664B68FF9FFFC1FAC983D0043FFA81" author="Montandon AL" pageId="18" pageNumber="1263" pagination="652 - 658" refId="ref23277" refString="Montandon AL. 1910 b. Especes nouvelles ou peu connues d'Hydrococrises de l'Inde. Bulletin de la Societe des Sciences de Bucarest, Roumanie 19: 652 - 658." type="journal article" year="1910">Montandon (1910b)</bibRefCitation>
questioned the existence of the
<typeStatus id="B54C883BFF9FFFC1FB2C8230025AFA81" box="[1256,1306,1303,1324]" pageId="18" pageNumber="1263">type</typeStatus>
specimen, which was based on a brief original description, and provided a redescription of the species based on a specimen in Distant’s collection from the Dawna Hills of eastern
<collectingCountry id="12E07609FF9FFFC1FC2982B60302FA0B" box="[1005,1090,1425,1446]" name="Myanmar" pageId="18" pageNumber="1263">Burma</collectingCountry>
. Given that most species in these groups are restricted to particular geographic regions, it is highly doubtful that the specimen upon which Montandon redescribed
<taxonomicName id="ADF74D1AFF9FFFC1FB5E82CA027BF9AF" baseAuthorityName="Spinola" baseAuthorityYear="1837" box="[1178,1339,1517,1538]" class="Insecta" family="Naucoridae" genus="Heleocoris" kingdom="Animalia" order="Hemiptera" pageId="18" pageNumber="1263" phylum="Arthropoda" rank="species" species="obliquatus">
<emphasis id="5883EA8BFF9FFFC1FB5E82CA027BF9AF" box="[1178,1339,1517,1538]" italics="true" pageId="18" pageNumber="1263">H. obliquatus</emphasis>
</taxonomicName>
was in fact that species. Further vitiating this identification is the figure in
<bibRefCitation id="0E664B68FF9FFFC1FC2A810D039EF992" author="Distant WL" box="[1006,1246,1578,1600]" pageId="18" pageNumber="1263" refId="ref21795" refString="Distant WL. 1910. The fauna of British India, including Ceylon and Burma. Rhynchota. Vol. V. Heteroptera: Appendix. London: Taylor &amp; Francis." type="book" year="1910">Distant (1910: 322)</bibRefCitation>
of a specimen from ‘Lower Burma’ identified by Montandon as
<taxonomicName id="ADF74D1AFF9FFFC1FCED814F0490F9D1" baseAuthorityName="Spinola" baseAuthorityYear="1837" box="[809,976,1639,1661]" class="Insecta" family="Naucoridae" genus="Heleocoris" kingdom="Animalia" order="Hemiptera" pageId="18" pageNumber="1263" phylum="Arthropoda" rank="species" species="obliquatus">
<emphasis id="5883EA8BFF9FFFC1FCED814F0490F9D1" box="[809,976,1639,1661]" italics="true" pageId="18" pageNumber="1263">H. obliquatus</emphasis>
</taxonomicName>
, which instead clearly is
<taxonomicName id="ADF74D1AFF9FFFC1FAD78140032EF931" authority="Montandon, 1897 b" authorityName="Montandon" authorityYear="1897" class="Insecta" family="Naucoridae" genus="Heleocoris" kingdom="Animalia" order="Hemiptera" pageId="18" pageNumber="1263" phylum="Arthropoda" rank="species" species="strabus">
<emphasis id="5883EA8BFF9FFFC1FAD7814004C3F936" italics="true" pageId="18" pageNumber="1263">Heleocoris strabus</emphasis>
<bibRefCitation id="0E664B68FF9FFFC1FC4881A1032EF931" author="Montandon AL" box="[908,1134,1670,1692]" pageId="18" pageNumber="1263" pagination="435 - 454" refId="ref23009" refString="Montandon AL. 1897 b. Hemiptera Cryptocerata. Fam. Naucoridae. Sous fam. Laccocorinae. Verhandlungen der Kaiserlich-Koniglichen Zoologisch-Botanischen Gesellschaft in Wien 47: 435 - 454." type="journal article" year="1897">Montandon, 1897b</bibRefCitation>
</taxonomicName>
. As such,
<taxonomicName id="ADF74D1AFF9FFFC1FB2881A002D1F936" baseAuthorityName="Spinola" baseAuthorityYear="1837" box="[1260,1425,1670,1692]" class="Insecta" family="Naucoridae" genus="Heleocoris" kingdom="Animalia" order="Hemiptera" pageId="18" pageNumber="1263" phylum="Arthropoda" rank="species" species="obliquatus">
<emphasis id="5883EA8BFF9FFFC1FB2881A002D1F936" box="[1260,1425,1670,1692]" italics="true" pageId="18" pageNumber="1263">H. obliquatus</emphasis>
</taxonomicName>
from
<collectingCountry id="12E07609FF9FFFC1FCB1818204A0F916" box="[885,992,1701,1723]" name="India" pageId="18" pageNumber="1263">Mumbai</collectingCountry>
remains enigmatic and nearly impossible to diagnose without an authoritatively identified specimen, and thus is a
<emphasis id="5883EA8BFF9FFFC1FB7181C402CBF95A" box="[1205,1419,1762,1784]" italics="true" pageId="18" pageNumber="1263">species inquirenda</emphasis>
. Nonetheless, because the
<typeStatus id="B54C883BFF9FFFC1FB99802603D0F8BB" box="[1117,1168,1793,1814]" pageId="18" pageNumber="1263">type</typeStatus>
locality was given as ‘Bombay’, I here consider the clade containing species of
<taxonomicName id="ADF74D1AFF9FFFC1FC87801904F9F8FE" authorityName="Stal" authorityYear="1876" box="[835,953,1854,1875]" class="Insecta" family="Naucoridae" genus="Heleocoris" kingdom="Animalia" order="Hemiptera" pageId="18" pageNumber="1263" phylum="Arthropoda" rank="genus">
<emphasis id="5883EA8BFF9FFFC1FC87801904F9F8FE" box="[835,953,1854,1875]" italics="true" pageId="18" pageNumber="1263">Heleocoris</emphasis>
</taxonomicName>
from central
<collectingCountry id="12E07609FF9FFFC1FB8B801903CCF8F9" box="[1103,1164,1854,1876]" name="India" pageId="18" pageNumber="1263">India</collectingCountry>
(all but the far eastern states) to be valid
<taxonomicName id="ADF74D1AFF9FFFC1FC38807A0332F8DF" authorityName="Stal" authorityYear="1876" box="[1020,1138,1885,1906]" class="Insecta" family="Naucoridae" genus="Heleocoris" kingdom="Animalia" order="Hemiptera" pageId="18" pageNumber="1263" phylum="Arthropoda" rank="genus">
<emphasis id="5883EA8BFF9FFFC1FC38807A0332F8DF" box="[1020,1138,1885,1906]" italics="true" pageId="18" pageNumber="1263">Heleocoris</emphasis>
</taxonomicName>
. Other clades containing
<taxonomicName id="ADF74D1AFF9EFFC0FF6784570659FC28" authorityName="Stal" authorityYear="1876" box="[163,281,880,901]" class="Insecta" family="Naucoridae" genus="Heleocoris" kingdom="Animalia" order="Hemiptera" pageId="19" pageNumber="1264" phylum="Arthropoda" rank="genus">
<emphasis id="5883EA8BFF9EFFC0FF6784570659FC28" box="[163,281,880,901]" italics="true" pageId="19" pageNumber="1264">Heleocoris</emphasis>
</taxonomicName>
from Africa and Indochina each must have the genus renamed.
</paragraph>
<caption id="3E886611FF9EFFC0FF6785DA0480FC9C" ID-DOI="http://doi.org/10.5281/zenodo.6994645" ID-Zenodo-Dep="6994645" httpUri="https://zenodo.org/record/6994645/files/figure.png" pageId="19" pageNumber="1264" startId="19.[163,244,765,787]" targetBox="[163,1442,195,726]" targetPageId="19">
<paragraph id="6A483699FF9EFFC0FF6785DA0480FC9C" blockId="19.[163,1443,765,817]" pageId="19" pageNumber="1264">
<emphasis id="5883EA8BFF9EFFC0FF6785DA065EFCBF" bold="true" box="[163,286,765,787]" pageId="19" pageNumber="1264">Figure 14.</emphasis>
Revised generic representation of the former genus
<taxonomicName id="ADF74D1AFF9EFFC0FCAF85D9049BFCB9" authorityName="Stal" authorityYear="1876" box="[875,987,766,788]" class="Insecta" family="Naucoridae" genus="Heleocoris" kingdom="Animalia" order="Hemiptera" pageId="19" pageNumber="1264" phylum="Arthropoda" rank="genus">
<emphasis id="5883EA8BFF9EFFC0FCAF85D9049BFCB9" box="[875,987,766,788]" italics="true" pageId="19" pageNumber="1264">Heleocoris</emphasis>
</taxonomicName>
. Dorsal habitus of
<taxonomicName id="ADF74D1AFF9EFFC0FB7585D90237FCB9" authorityName="Montandon" authorityYear="1910" box="[1201,1399,766,788]" class="Insecta" family="Naucoridae" genus="Heleocoris" kingdom="Animalia" order="Hemiptera" pageId="19" pageNumber="1264" phylum="Arthropoda" rank="species" species="vicinus">
<emphasis id="5883EA8BFF9EFFC0FB7585D90237FCB9" box="[1201,1399,766,788]" italics="true" pageId="19" pageNumber="1264">Heleocoris vicinus</emphasis>
</taxonomicName>
(a),
<taxonomicName id="ADF74D1AFF9EFFC0FF67843C06C9FC9C" box="[163,393,795,817]" class="Insecta" family="Naucoridae" genus="Laccocoris" kingdom="Animalia" order="Hemiptera" pageId="19" pageNumber="1264" phylum="Arthropoda" rank="species" species="minisculus">
<emphasis id="5883EA8BFF9EFFC0FF67843C06C9FC9C" box="[163,393,795,817]" italics="true" pageId="19" pageNumber="1264">Laccocoris minisculus</emphasis>
</taxonomicName>
(b),
<taxonomicName id="ADF74D1AFF9EFFC0FE72843C05DFFC9C" authorityName="Sites" authorityYear="2022" baseAuthorityName="Montandon" baseAuthorityYear="1897" box="[438,671,795,817]" class="Insecta" family="Naucoridae" genus="Heleolaccocoris" kingdom="Animalia" order="Hemiptera" pageId="19" pageNumber="1264" phylum="Arthropoda" rank="species" species="ovatus">
<emphasis id="5883EA8BFF9EFFC0FE72843C05DFFC9C" box="[438,671,795,817]" italics="true" pageId="19" pageNumber="1264">Heleolaccocoris ovatus</emphasis>
</taxonomicName>
(c),
<taxonomicName id="ADF74D1AFF9EFFC0FD0E843C04D7FC9C" authorityName="Sites" authorityYear="2022" baseAuthorityName="Signoret" baseAuthorityYear="1860" box="[714,919,795,817]" class="Insecta" family="Naucoridae" genus="Tsingala" kingdom="Animalia" order="Hemiptera" pageId="19" pageNumber="1264" phylum="Arthropoda" rank="species" species="humeralis">
<emphasis id="5883EA8BFF9EFFC0FD0E843C04D7FC9C" box="[714,919,795,817]" italics="true" pageId="19" pageNumber="1264">Tsingala humeralis</emphasis>
</taxonomicName>
(d).
</paragraph>
</caption>
<paragraph id="6A483699FF9EFFC0FF7F848A056CF929" blockId="19.[163,780,880,1668]" pageId="19" pageNumber="1264">
Species of
<taxonomicName id="ADF74D1AFF9EFFC0FEF0848906EAFC6E" authorityName="Stal" authorityYear="1856" box="[308,426,942,963]" class="Insecta" family="Naucoridae" genus="Laccocoris" kingdom="Animalia" order="Hemiptera" pageId="19" pageNumber="1264" phylum="Arthropoda" rank="genus">
<emphasis id="5883EA8BFF9EFFC0FEF0848906EAFC6E" box="[308,426,942,963]" italics="true" pageId="19" pageNumber="1264">Laccocoris</emphasis>
</taxonomicName>
were recovered in two distinct clades within different tribes (
<figureCitation id="F2CC2A1CFF9EFFC0FDCF84EB0522FC4C" box="[523,610,972,994]" captionStart="Figure 13" captionStartId="18.[145,223,1057,1079]" captionTargetBox="[149,1425,195,1017]" captionTargetId="figure-432@18.[145,1425,195,1017]" captionTargetPageId="18" captionText="Figure 13. Detail of Most Probable Phylogeny dendogram of Laccocorinae.The red-coloured clade includes the Madagascar genera Temnocoris and Tsingala gen. nov.(formerly Heleocoris). The blue-coloured clade represents Heleolaccocoris gen.nov., which comprises species of the former genera Heleocoris and Laccocoris. All species of Tsingala gen. nov. and Heleolaccocoris gen. nov. are new combinations. Geographic associations for major clades within and among tribes are indicated. The red box in the inset shows the part of Figure 9 depicted in this figure." figureDoi="http://doi.org/10.5281/zenodo.6994641" httpUri="https://zenodo.org/record/6994641/files/figure.png" pageId="19" pageNumber="1264">Figs 13</figureCitation>
,
<figureCitation id="F2CC2A1CFF9EFFC0FDB584EB05CDFC4F" box="[625,653,972,994]" captionStart="Figure 15" captionStartId="20.[145,226,1055,1077]" captionTargetBox="[145,1425,195,1015]" captionTargetId="figure-409@20.[145,1425,195,1015]" captionTargetPageId="20" captionText="Figure 15. Revised generic representation of the former genus Laccocoris. Dorsal habitus of Laccocoris spurcus (a), Heleolaccocoris horvathi (b)." figureDoi="http://doi.org/10.5281/zenodo.6994647" httpUri="https://zenodo.org/record/6994647/files/figure.png" pageId="19" pageNumber="1264">15</figureCitation>
). The
<typeStatus id="B54C883BFF9EFFC0FD1C84EB044AFC4C" box="[728,778,972,993]" pageId="19" pageNumber="1264">type</typeStatus>
species is
<taxonomicName id="ADF74D1AFF9EFFC0FED384CC053AFBAD" authority="(Stal, 1856)" baseAuthorityName="Stal" baseAuthorityYear="1856" box="[279,634,1002,1024]" class="Insecta" family="Naucoridae" genus="Laccocoris" kingdom="Animalia" order="Hemiptera" pageId="19" pageNumber="1264" phylum="Arthropoda" rank="species" species="spurcus">
<emphasis id="5883EA8BFF9EFFC0FED384CC06AFFBAD" box="[279,495,1003,1024]" italics="true" pageId="19" pageNumber="1264">Laccocoris spurcus</emphasis>
(
<bibRefCitation id="0E664B68FF9EFFC0FE3A84CD0532FBAD" author="Stal C" box="[510,626,1002,1024]" pageId="19" pageNumber="1264" pagination="193 - 199" refId="ref24930" refString="Stal C. 1856. Hemiptera samlade af Victorin i Caplandet. Ofversigt af Kongliga Svenska Vetenskaps-Akademiens Forhandlingar 13: 193 - 199." type="journal article" year="1856">Stål, 1856</bibRefCitation>
)
</taxonomicName>
from Africa; thus, the species of
<taxonomicName id="ADF74D1AFF9EFFC0FE5D832D0556FBB2" authorityName="Stal" authorityYear="1856" box="[409,534,1034,1055]" class="Insecta" family="Naucoridae" genus="Laccocoris" kingdom="Animalia" order="Hemiptera" pageId="19" pageNumber="1264" phylum="Arthropoda" rank="genus">
<emphasis id="5883EA8BFF9EFFC0FE5D832D0556FBB2" box="[409,534,1034,1055]" italics="true" pageId="19" pageNumber="1264">Laccocoris</emphasis>
</taxonomicName>
from Africa remain unchanged. However, the species of
<taxonomicName id="ADF74D1AFF9EFFC0FD94830F058AFB90" authorityName="Stal" authorityYear="1856" box="[592,714,1064,1085]" class="Insecta" family="Naucoridae" genus="Laccocoris" kingdom="Animalia" order="Hemiptera" pageId="19" pageNumber="1264" phylum="Arthropoda" rank="genus">
<emphasis id="5883EA8BFF9EFFC0FD94830F058AFB90" box="[592,714,1064,1085]" italics="true" pageId="19" pageNumber="1264">Laccocoris</emphasis>
</taxonomicName>
from Sundaland and the
<collectingCountry id="12E07609FF9EFFC0FEB1836106B3FBF1" box="[373,499,1094,1116]" name="Philippines" pageId="19" pageNumber="1264">Philippines</collectingCountry>
are here placed in a newly erected genus,
<taxonomicName id="ADF74D1AFF9EFFC0FE9783420545FBD7" authority="Because" authorityName="Sites" authorityYear="2022" box="[339,517,1125,1146]" class="Insecta" family="Naucoridae" genus="Heleolaccocoris" kingdom="Animalia" order="Hemiptera" pageId="19" pageNumber="1264" phylum="Arthropoda" rank="genus" status="gen. nov.">
<emphasis id="5883EA8BFF9EFFC0FE9783420545FBD7" box="[339,517,1125,1146]" italics="true" pageId="19" pageNumber="1264">Heleolaccocoris</emphasis>
</taxonomicName>
<taxonomicNameLabel id="43B057F0FF9EFFC0FDCA83410534FBD6" box="[526,628,1126,1147]" pageId="19" pageNumber="1264" rank="genus">gen. nov.</taxonomicNameLabel>
Because the well-supported ML topology recovered
<taxonomicName id="ADF74D1AFF9EFFC0FD9A83A3044BFB34" authorityName="Sites" authorityYear="2022" box="[606,779,1156,1177]" class="Insecta" family="Naucoridae" genus="Heleolaccocoris" kingdom="Animalia" order="Hemiptera" pageId="19" pageNumber="1264" phylum="Arthropoda" rank="genus">
<emphasis id="5883EA8BFF9EFFC0FD9A83A3044BFB34" box="[606,779,1156,1177]" italics="true" pageId="19" pageNumber="1264">Heleolaccocoris</emphasis>
</taxonomicName>
as sister to three species of Indochinese
<taxonomicName id="ADF74D1AFF9EFFC0FDB2838505ACFB1A" authorityName="Stal" authorityYear="1876" box="[630,748,1186,1207]" class="Insecta" family="Naucoridae" genus="Heleocoris" kingdom="Animalia" order="Hemiptera" pageId="19" pageNumber="1264" phylum="Arthropoda" rank="genus">
<emphasis id="5883EA8BFF9EFFC0FDB2838505ACFB1A" box="[630,748,1186,1207]" italics="true" pageId="19" pageNumber="1264">Heleocoris</emphasis>
</taxonomicName>
in successively more inclusive clades, this necessitated either designation of additional genera to accommodate the clades of Indochinese
<taxonomicName id="ADF74D1AFF9EFFC0FE0B83D90505FABE" authorityName="Stal" authorityYear="1876" box="[463,581,1278,1299]" class="Insecta" family="Naucoridae" genus="Heleocoris" kingdom="Animalia" order="Hemiptera" pageId="19" pageNumber="1264" phylum="Arthropoda" rank="genus">
<emphasis id="5883EA8BFF9EFFC0FE0B83D90505FABE" box="[463,581,1278,1299]" italics="true" pageId="19" pageNumber="1264">Heleocoris</emphasis>
</taxonomicName>
, or placing them all in
<taxonomicName id="ADF74D1AFF9EFFC0FF27823A06CCFA9F" authorityName="Sites" authorityYear="2022" box="[227,396,1309,1330]" class="Insecta" family="Naucoridae" genus="Heleolaccocoris" kingdom="Animalia" order="Hemiptera" pageId="19" pageNumber="1264" phylum="Arthropoda" rank="genus">
<emphasis id="5883EA8BFF9EFFC0FF27823A06CCFA9F" box="[227,396,1309,1330]" italics="true" pageId="19" pageNumber="1264">Heleolaccocoris</emphasis>
</taxonomicName>
as I have done here (
<figureCitation id="F2CC2A1CFF9EFFC0FDBF823A0586FA9F" box="[635,710,1309,1331]" captionStart="Figure 13" captionStartId="18.[145,223,1057,1079]" captionTargetBox="[149,1425,195,1017]" captionTargetId="figure-432@18.[145,1425,195,1017]" captionTargetPageId="18" captionText="Figure 13. Detail of Most Probable Phylogeny dendogram of Laccocorinae.The red-coloured clade includes the Madagascar genera Temnocoris and Tsingala gen. nov.(formerly Heleocoris). The blue-coloured clade represents Heleolaccocoris gen.nov., which comprises species of the former genera Heleocoris and Laccocoris. All species of Tsingala gen. nov. and Heleolaccocoris gen. nov. are new combinations. Geographic associations for major clades within and among tribes are indicated. The red box in the inset shows the part of Figure 9 depicted in this figure." figureDoi="http://doi.org/10.5281/zenodo.6994641" httpUri="https://zenodo.org/record/6994641/files/figure.png" pageId="19" pageNumber="1264">Fig. 13</figureCitation>
: bluecoloured clades). The previously recognized distinction between the two genera (
<taxonomicName id="ADF74D1AFF9EFFC0FE15827D0509FAC2" authorityName="Stal" authorityYear="1876" box="[465,585,1370,1391]" class="Insecta" family="Naucoridae" genus="Heleocoris" kingdom="Animalia" order="Hemiptera" pageId="19" pageNumber="1264" phylum="Arthropoda" rank="genus">
<emphasis id="5883EA8BFF9EFFC0FE15827D0509FAC2" box="[465,585,1370,1391]" italics="true" pageId="19" pageNumber="1264">Heleocoris</emphasis>
</taxonomicName>
and
<taxonomicName id="ADF74D1AFF9EFFC0FD43827C0441FADD" authorityName="Stal" authorityYear="1856" box="[647,769,1371,1392]" class="Insecta" family="Naucoridae" genus="Laccocoris" kingdom="Animalia" order="Hemiptera" pageId="19" pageNumber="1264" phylum="Arthropoda" rank="genus">
<emphasis id="5883EA8BFF9EFFC0FD43827C0441FADD" box="[647,769,1371,1392]" italics="true" pageId="19" pageNumber="1264">Laccocoris</emphasis>
</taxonomicName>
) was based on the shape of the labrum as wider than long with a rounded apex or longer than wide with an acute apex. This has been generally considered a feature inconsistent with phylogeny by recent scientists who have studied the group. In a treatment of the
<taxonomicName id="ADF74D1AFF9EFFC0FD4F82D3044BF9A7" authorityName="LEACH" authorityYear="1815" box="[651,779,1524,1546]" class="Insecta" family="Naucoridae" kingdom="Animalia" order="Hemiptera" pageId="19" pageNumber="1264" phylum="Arthropoda" rank="family">Naucoridae</taxonomicName>
in
<collectingCountry id="12E07609FF9EFFC0FF0481350673F985" box="[192,307,1554,1576]" name="Singapore" pageId="19" pageNumber="1264">Singapore</collectingCountry>
,
<bibRefCitation id="0E664B68FF9EFFC0FEFB813505CBF98A" author="Polhemus DA &amp; Polhemus JT" box="[319,651,1554,1576]" pageId="19" pageNumber="1264" pagination="665 - 686" refId="ref23727" refString="Polhemus DA, Polhemus JT. 2013. Guide to the aquatic Heteroptera of Singapore and Peninsular Malaysia. XI. Infraorder Nepomorpha - families Naucoridae and Aphelocheiridae. Raffles Bulletin of Zoology 61: 665 - 686." type="journal article" year="2013">Polhemus &amp; Polhemus (2013)</bibRefCitation>
considered the generic limits of
<taxonomicName id="ADF74D1AFF9EFFC0FE5E81160552F9EB" authorityName="Stal" authorityYear="1876" box="[410,530,1585,1606]" class="Insecta" family="Naucoridae" genus="Heleocoris" kingdom="Animalia" order="Hemiptera" pageId="19" pageNumber="1264" phylum="Arthropoda" rank="genus">
<emphasis id="5883EA8BFF9EFFC0FE5E81160552F9EB" box="[410,530,1585,1606]" italics="true" pageId="19" pageNumber="1264">Heleocoris</emphasis>
</taxonomicName>
and
<taxonomicName id="ADF74D1AFF9EFFC0FD8B81150589F9EA" authorityName="Stal" authorityYear="1856" box="[591,713,1586,1607]" class="Insecta" family="Naucoridae" genus="Laccocoris" kingdom="Animalia" order="Hemiptera" pageId="19" pageNumber="1264" phylum="Arthropoda" rank="genus">
<emphasis id="5883EA8BFF9EFFC0FD8B81150589F9EA" box="[591,713,1586,1607]" italics="true" pageId="19" pageNumber="1264">Laccocoris</emphasis>
</taxonomicName>
to be indistinct and correctly predicted the possible synonymy of Asian
<taxonomicName id="ADF74D1AFF9EFFC0FEC681480635F929" authorityName="Stal" authorityYear="1856" box="[258,373,1647,1668]" class="Insecta" family="Naucoridae" genus="Laccocoris" kingdom="Animalia" order="Hemiptera" pageId="19" pageNumber="1264" phylum="Arthropoda" rank="genus">
<emphasis id="5883EA8BFF9EFFC0FEC681480635F929" box="[258,373,1647,1668]" italics="true" pageId="19" pageNumber="1264">Laccocoris</emphasis>
</taxonomicName>
with
<taxonomicName id="ADF74D1AFF9EFFC0FE7081490565F92E" authorityName="Stal" authorityYear="1876" box="[436,549,1646,1667]" class="Insecta" family="Naucoridae" genus="Heleocoris" kingdom="Animalia" order="Hemiptera" pageId="19" pageNumber="1264" phylum="Arthropoda" rank="genus">
<emphasis id="5883EA8BFF9EFFC0FE7081490565F92E" box="[436,549,1646,1667]" italics="true" pageId="19" pageNumber="1264">Heleocoris</emphasis>
</taxonomicName>
.
</paragraph>
</subSubSection>
<subSubSection id="22ED6512FF9EFFC0FF6781940228FB34" pageId="19" pageNumber="1264" type="diagnosis">
<paragraph id="6A483699FF9EFFC0FF6781940228FB34" blockId="19.[163,780,1714,1890]" lastBlockId="19.[827,1443,880,1178]" pageId="19" pageNumber="1264">
<emphasis id="5883EA8BFF9EFFC0FF6781940663F965" box="[163,291,1715,1736]" italics="true" pageId="19" pageNumber="1264">Diagnosis:</emphasis>
This subfamily has a suite of unique diagnostic attributes, some of which exhibit variation among genera. The front of the head is folded posteroventrally such that the labrum is set back from the functional anterior margin of the head (
<figureCitation id="F2CC2A1CFF9EFFC0FD6F800A05BBF8EE" box="[683,763,1837,1859]" captionStart="Figure 3" captionStartId="8.[145,230,1468,1490]" captionTargetBox="[172,733,195,1428]" captionTargetId="figure-532@8.[172,733,195,1428]" captionTargetPageId="8" captionText="Figure 3. Mouthpart positions: recessed rostrum of Cheirochela thailandana (a); lateral views of Laccocoris spurcus (b) and Australambrysus crenulatus (c). White arrows indicate positions of labrum with respect to the functional anterior margin of the head." figureDoi="http://doi.org/10.5281/zenodo.6994613" httpUri="https://zenodo.org/record/6994613/files/figure.png" pageId="19" pageNumber="1264">Fig. 3b</figureCitation>
). Males of all genera have a dense tomentose pad ventrally over much of the mesotibia and tarsomeres 2 and 3 (
<figureCitation id="F2CC2A1CFF9EFFC0FC8784A904CEFC0E" box="[835,910,910,932]" captionStart="Figure 12" captionStartId="17.[163,243,870,892]" captionTargetBox="[163,1443,195,830]" captionTargetId="figure-534@17.[163,1443,195,830]" captionTargetPageId="17" captionText="Figure 12. Mesotibia and mesotarsus of male and female Heleolaccocoris ovatus showing a sexually dimorphic ventral tomentose pad of hairs." figureDoi="http://doi.org/10.5281/zenodo.6994637" httpUri="https://zenodo.org/record/6994637/files/figure.png" pageId="19" pageNumber="1264">Fig. 12</figureCitation>
), and to a lesser extent the protibia and tarsus, whereas in females the pads are greatly reduced. The prothoracic pretarsal claws are paired and articulated, except in
<taxonomicName id="ADF74D1AFF9EFFC0FC6884CD030BFC52" authorityName="Sites" authorityYear="2007" box="[940,1099,1002,1023]" class="Insecta" family="Naucoridae" genus="Namtokocoris" kingdom="Animalia" order="Hemiptera" pageId="19" pageNumber="1264" phylum="Arthropoda" rank="genus">
<emphasis id="5883EA8BFF9EFFC0FC6884CD030BFC52" box="[940,1099,1002,1023]" italics="true" pageId="19" pageNumber="1264">Namtokocoris</emphasis>
</taxonomicName>
in which they are single and fused with the tarsus, which is the condition found in all other subfamilies (
<figureCitation id="F2CC2A1CFF9EFFC0FBD4830F031AFB90" box="[1040,1114,1064,1086]" captionStart="Figure 4" captionStartId="9.[163,249,1379,1401]" captionTargetBox="[163,781,195,1339]" captionTargetId="figure-575@9.[163,781,195,1339]" captionTargetPageId="9" captionText="Figure 4. Foretibiae and foretarsi of Laccocorinae: Ctenipocoris asiaticus (a), Heleolaccocoris ovatus (b), Namtokocoris dalanta (c)." figureDoi="http://doi.org/10.5281/zenodo.6994615" httpUri="https://zenodo.org/record/6994615/files/figure.png" pageId="19" pageNumber="1264">Fig. 4c</figureCitation>
). Males have two-segmented front tarsi and females only one, except in
<taxonomicName id="ADF74D1AFF9EFFC0FADC836002E2FBF1" authorityName="Montandon" authorityYear="1897" box="[1304,1442,1095,1116]" class="Insecta" family="Naucoridae" genus="Ctenipocoris" kingdom="Animalia" order="Hemiptera" pageId="19" pageNumber="1264" phylum="Arthropoda" rank="genus">
<emphasis id="5883EA8BFF9EFFC0FADC836002E2FBF1" box="[1304,1442,1095,1116]" italics="true" pageId="19" pageNumber="1264">Ctenipocoris</emphasis>
</taxonomicName>
where both sexes have two, and in
<taxonomicName id="ADF74D1AFF9EFFC0FB158342022FFBD7" authorityName="Sites" authorityYear="2007" box="[1233,1391,1125,1146]" class="Insecta" family="Naucoridae" genus="Namtokocoris" kingdom="Animalia" order="Hemiptera" pageId="19" pageNumber="1264" phylum="Arthropoda" rank="genus">
<emphasis id="5883EA8BFF9EFFC0FB158342022FFBD7" box="[1233,1391,1125,1146]" italics="true" pageId="19" pageNumber="1264">Namtokocoris</emphasis>
</taxonomicName>
and
<taxonomicName id="ADF74D1AFF9EFFC0FCFF83A304A4FB34" authorityName="Sites &amp; Zettel" authorityYear="2011" box="[827,996,1156,1177]" class="Insecta" family="Naucoridae" genus="Pogonocaudina" kingdom="Animalia" order="Hemiptera" pageId="19" pageNumber="1264" phylum="Arthropoda" rank="genus">
<emphasis id="5883EA8BFF9EFFC0FCFF83A304A4FB34" box="[827,996,1156,1177]" italics="true" pageId="19" pageNumber="1264">Pogonocaudina</emphasis>
</taxonomicName>
where both sexes have one (
<figureCitation id="F2CC2A1CFF9EFFC0FAD983A3021AFB34" box="[1309,1370,1156,1178]" captionStart="Figure 4" captionStartId="9.[163,249,1379,1401]" captionTargetBox="[163,781,195,1339]" captionTargetId="figure-575@9.[163,781,195,1339]" captionTargetPageId="9" captionText="Figure 4. Foretibiae and foretarsi of Laccocorinae: Ctenipocoris asiaticus (a), Heleolaccocoris ovatus (b), Namtokocoris dalanta (c)." figureDoi="http://doi.org/10.5281/zenodo.6994615" httpUri="https://zenodo.org/record/6994615/files/figure.png" pageId="19" pageNumber="1264">Fig. 4</figureCitation>
).
</paragraph>
</subSubSection>
<subSubSection id="22ED6512FF9EFFC0FCFF83920482FAC9" pageId="19" pageNumber="1264" type="discussion">
<paragraph id="6A483699FF9EFFC0FCFF83920482FAC9" blockId="19.[827,1443,1205,1380]" pageId="19" pageNumber="1264">
<emphasis id="5883EA8BFF9EFFC0FCFF83920486FB67" box="[827,966,1205,1226]" italics="true" pageId="19" pageNumber="1264">Comments:</emphasis>
This subfamily is circumtropical in distribution and is most common in Africa and Asia. It is not known from
<collectingCountry id="12E07609FF9EFFC0FBD483D5033FFAA5" box="[1040,1151,1266,1288]" name="Australia" pageId="19" pageNumber="1264">Australia</collectingCountry>
. Although
<taxonomicName id="ADF74D1AFF9EFFC0FB3F83D402C9FAA5" authorityName="Montandon" authorityYear="1897" box="[1275,1417,1267,1288]" class="Insecta" family="Naucoridae" genus="Ctenipocoris" kingdom="Animalia" order="Hemiptera" pageId="19" pageNumber="1264" phylum="Arthropoda" rank="genus">
<emphasis id="5883EA8BFF9EFFC0FB3F83D402C9FAA5" box="[1275,1417,1267,1288]" italics="true" pageId="19" pageNumber="1264">Ctenipocoris</emphasis>
</taxonomicName>
is circumtropical in distribution, each of the other major clades in the subfamily has a distinct geographic association.
</paragraph>
</subSubSection>
</treatment>
</document>
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