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<document id="A8578B55A05451D5FD76F23B06591D6C" ID-CLB-Dataset="202083" ID-DOI="10.11646/zootaxa.5211.1.1" ID-GBIF-Dataset="bb97b9dd-2acc-4244-8f30-72bef07b9ed9" ID-ISSN="1175-5326" ID-Zenodo-Dep="7345007" ID-ZooBank="5754769C-B747-4714-BDD9-7D5509D48BEB" IM.materialsCitations_approvedBy="felipe" IM.metadata_approvedBy="felipe" IM.tables_requiresApprovalFor="existingObjects,plazi" IM.taxonomicNames_approvedBy="felipe" checkinTime="1669101340847" checkinUser="plazi" docAuthor="López-García, Margarita M. &amp; Deloya, Cuauhtémoc" docDate="2022" docId="A209493BFD61FFBEAFA61D99FEC0BF43" docLanguage="en" docName="zootaxa.5211.1.1.pdf" docOrigin="Zootaxa 5211 (1)" docSource="http://dx.doi.org/10.11646/zootaxa.5211.1.1" docStyle="DocumentStyle:5EBBA59367AD13919D70D935FA04F6A3.14:Zootaxa.2013-.monograph" docStyleId="5EBBA59367AD13919D70D935FA04F6A3" docStyleName="Zootaxa.2013-.monograph" docStyleVersion="14" docTitle="Tomarus laticaudus Lopez-Garcia &amp; Deloya 2019" docType="treatment" docVersion="10" lastPageNumber="64" masterDocId="5E303143FD5FFF81AF311956FFBEBC61" masterDocTitle="Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision" masterLastPageNumber="119" masterPageNumber="1" pageNumber="63" updateTime="1699396337306" updateUser="plazi">
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<mods:title id="979F4B603ED5DC8DAD3666D300564DCD">Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision</mods:title>
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<taxonomicName id="EDA083AEFD61FFBFAFA61D99FCBFB889" authority="Lopez-Garcia &amp; Deloya, 2019" authorityName="Lopez-Garcia &amp; Deloya" authorityYear="2019" box="[151,769,1229,1257]" class="Insecta" family="Dynastidae" genus="Tomarus" kingdom="Animalia" order="Coleoptera" pageId="62" pageNumber="63" phylum="Arthropoda" rank="species" species="laticaudus">
<emphasis id="18D4243FFD61FFBFAFA61D99FCBFB889" bold="true" box="[151,769,1229,1257]" pageId="62" pageNumber="63">
<emphasis id="18D4243FFD61FFBFAFA61D99FE39B888" bold="true" box="[151,391,1230,1257]" italics="true" pageId="62" pageNumber="63">Tomarus laticaudus</emphasis>
<bibRefCitation id="4E3185DCFD61FFBFAEBF1D9BFCBFB889" author="Lopez-Garcia, M. M. &amp; Deloya, C." box="[398,769,1229,1256]" pageId="62" pageNumber="63" pagination="127 - 141" refId="ref91328" refString="Lopez-Garcia, M. M. &amp; Deloya, C. (2019) Five new species of the dynastine genus Tomarus (Coleoptera: Scarabaeidae), with an illustrated key to species. The Coleopterists Bulletin, 73, 127 - 141. https: // doi. org / 10.1649 / 0010 - 065 X- 73.1.127" type="journal article" year="2019">López-García &amp; Deloya, 2019</bibRefCitation>
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<paragraph id="2A1FF82DFD61FFBFAFA61DA4FD85B96D" blockId="62.[151,769,1229,1292]" box="[151,571,1266,1292]" pageId="62" pageNumber="63">
(
<figureCitation id="B29BE4A8FD61FFBFAFAE1DA4FEAFB96D" box="[159,273,1266,1292]" captionStart="FIGURE 13" captionStartId="89.[151,250,1041,1065]" captionTargetBox="[154,1432,183,1016]" captionTargetId="figure-17@89.[151,1436,181,1017]" captionTargetPageId="89" captionText="FIGURE 13. Female genital plates. A) Euligyrus ebenus, B) Proculigyrus cicatricosus, C) Ligyrus (Anagrylius) moroni, D) L. (Ligyrodes) sallaei, E) L. (Ligyrus) allonasutus, F) L. (Ligyrus) bidentulus, G) L. (Ligyrus) neglectus, H) Tomarus laticaudus, I) T. maternus, J) T. rostratus, K) T. subtropicus. Abbreviations: SCx: Subcoxite, Cx: Coxite." figureDoi="http://doi.org/10.5281/zenodo.7345057" httpUri="https://zenodo.org/record/7345057/files/figure.png" pageId="62" pageNumber="63">Figs. 13H</figureCitation>
,
<figureCitation id="B29BE4A8FD61FFBFAE2C1DA4FEF1B96D" box="[285,335,1266,1292]" captionStart="FIGURE 16" captionStartId="91.[151,250,1633,1657]" captionTargetBox="[151,1426,189,1602]" captionTargetId="figure-17@91.[151,1436,181,1610]" captionTargetPageId="91" captionText="FIGURE 16. Parameres in frontal view. A) Tomarus adoceteus, B) T. amazonicus, C) T. bituberculatus, D) T. colombianus, E) T. discrepans, F) T. gyas, G) T. laevicollis, H) T. laticaudus, I) T. maimon, J) T. maracaiboensis, K) T. maternus, L) T. pilcopataensis, M) T. pullus, N) T. pumilus, O) T. roigjunenti, P) T. rostratus, Q) T. selanderi, R) T. subtropicus." figureDoi="http://doi.org/10.5281/zenodo.7345065" httpUri="https://zenodo.org/record/7345065/files/figure.png" pageId="62" pageNumber="63">16H</figureCitation>
,
<figureCitation id="B29BE4A8FD61FFBFAE6A1DA4FE30B96D" box="[347,398,1266,1292]" captionStart="FIGURE 19" captionStartId="94.[151,250,1155,1179]" captionTargetBox="[157,1432,188,1128]" captionTargetId="figure-24@94.[152,1436,181,1131]" captionTargetPageId="94" captionText="FIGURE 19. Parameres in lateral view. A) Tomarus adoceteus, B) T. amazonicus, C) T. bituberculatus, D) T. colombianus, E) T. discrepans, F) T. gyas, G) T. laevicollis, H) T. laticaudus, I) T. maimon." figureDoi="http://doi.org/10.5281/zenodo.7345079" httpUri="https://zenodo.org/record/7345079/files/figure.png" pageId="62" pageNumber="63">19H</figureCitation>
,
<figureCitation id="B29BE4A8FD61FFBFAEA81DA4FE75B96D" box="[409,459,1266,1292]" captionStart="FIGURE 23" captionStartId="98.[151,250,1728,1752]" captionTargetBox="[161,1421,189,1693]" captionTargetId="figure-24@98.[151,1436,181,1704]" captionTargetPageId="98" captionText="FIGURE 23. Internal sac. A) Tomarus amazonicus, B) T. discrepans, C) T. laticaudus, D) T. maimon. Abbreviations: AL: Accessory lamellae, CL: Copulatory lamella, LSB: Lamellar spiny belt, Ra: Raspula, SD: Sclerotized diverticulum." figureDoi="http://doi.org/10.5281/zenodo.7345091" httpUri="https://zenodo.org/record/7345091/files/figure.png" pageId="62" pageNumber="63">23C</figureCitation>
,
<figureCitation id="B29BE4A8FD61FFBFAEE71DA4FDB4B96D" box="[470,522,1266,1292]" captionStart="FIGURE 27" captionStartId="102.[151,250,1868,1892]" captionTargetBox="[158,1409,184,1839]" captionTargetId="figure-24@102.[151,1436,181,1844]" captionTargetPageId="102" captionText="FIGURE 27. Habitus of the species of Tomarus: A) Tomarus adoceteus, B) T. amazonicus, C) T. bituberculatus, D) T. colombianus, E) T. discrepans, F) T. gyas, G) T. laevicollis, H) T. laticaudus, I) T. maimon. Scale bar: 5 mm." figureDoi="http://doi.org/10.5281/zenodo.7345103" httpUri="https://zenodo.org/record/7345103/files/figure.png" pageId="62" pageNumber="63">27H</figureCitation>
;
<figureCitation id="B29BE4A8FD61FFBFAD241DA4FD8CB96D" box="[533,562,1266,1292]" captionStart="FIGURE 44" captionStartId="112.[151,250,1998,2022]" captionTargetBox="[224,1372,1135,1965]" captionTargetId="figure-38@112.[215,1372,1135,1973]" captionTargetPageId="112" captionText="FIGURE 44. Distributional records of Tomarus laevicollis, T. laticaudus, T. maracaiboensis, and T. colombianus." figureDoi="http://doi.org/10.5281/zenodo.7345164" httpUri="https://zenodo.org/record/7345164/files/figure.png" pageId="62" pageNumber="63">44</figureCitation>
)
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<paragraph id="2A1FF82DFD61FFBFAFA61C61FC08B92E" blockId="62.[151,1437,1335,1635]" box="[151,950,1335,1359]" pageId="62" pageNumber="63">
<treatmentCitationGroup id="0AB0DF03FD61FFBFAFA61C61FC08B92E" box="[151,950,1335,1359]" pageId="62" pageNumber="63">
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<taxonomicName id="EDA083AEFD61FFBFAFA61C61FD73B92E" authority="Lopez-Garcia &amp; Deloya, 2019: 129" authorityName="Lopez-Garcia &amp; Deloya" authorityPageNumber="129" authorityYear="2019" box="[151,717,1335,1359]" class="Insecta" family="Dynastidae" genus="Tomarus" kingdom="Animalia" order="Coleoptera" pageId="62" pageNumber="63" phylum="Arthropoda" rank="species" species="laticaudus">
<emphasis id="18D4243FFD61FFBFAFA61C61FEE1B92E" box="[151,351,1335,1359]" italics="true" pageId="62" pageNumber="63">Tomarus laticaudus</emphasis>
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<bibRefCitation id="4E3185DCFD61FFBFAE551C61FD73B92E" author="Lopez-Garcia, M. M. &amp; Deloya, C." box="[356,717,1335,1359]" pageId="62" pageNumber="63" pagination="127 - 141" refId="ref91328" refString="Lopez-Garcia, M. M. &amp; Deloya, C. (2019) Five new species of the dynastine genus Tomarus (Coleoptera: Scarabaeidae), with an illustrated key to species. The Coleopterists Bulletin, 73, 127 - 141. https: // doi. org / 10.1649 / 0010 - 065 X- 73.1.127" type="journal article" year="2019">López-García &amp; Deloya, 2019: 129</bibRefCitation>
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.
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combination.
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(USNM)
<materialsCitation id="9AC8F270FD61FFBFAB821C61FE9FB9ED" ID-GBIF-Occurrence="3986696676" collectionCode="USNM" collectorName="Two" pageId="62" pageNumber="63" specimenCount="2" specimenCount-female="1" typeStatus="holotype">
“FF // CorazalCZ / Pan IV2711 / AugBusck //
<taxonomicName id="EDA083AEFD61FFBFAE981C00FDD0B90C" authorityName="Lopez-Garcia &amp; Deloya" authorityYear="2019" box="[425,622,1365,1389]" class="Insecta" family="Dynastidae" genus="Tomarus" kingdom="Animalia" order="Coleoptera" pageId="62" pageNumber="63" phylum="Arthropoda" rank="species" species="laticaudus">Tomarus laticaudus</taxonomicName>
/ López-García &amp; Deloya /
<typeStatus id="F51B468FFD61FFBFACB61C03FBACB90C" box="[903,1042,1365,1389]" pageId="62" pageNumber="63" type="holotype">HOLOTYPE</typeStatus>
”.
<collectorName id="87559DFBFD61FFBFAB151C00FBEFB90C" box="[1060,1105,1366,1389]" pageId="62" pageNumber="63">Two</collectorName>
male and
<specimenCount id="3CA633A4FD61FFBFAB891C03FA8BB90C" box="[1208,1333,1365,1389]" pageId="62" pageNumber="63" type="female">three female</specimenCount>
<typeStatus id="F51B468FFD61FFBFAA0A1C00FA22B90F" box="[1339,1436,1366,1390]" pageId="62" pageNumber="63" type="paratype">paratypes</typeStatus>
(
<collectionCode id="4CB160E8FD61FFBFAFFF1C22FEA4B9ED" box="[206,282,1396,1420]" country="USA" lsid="urn:lsid:biocol.org:col:34871" name="Smithsonian Institution, National Museum of Natural History" pageId="62" pageNumber="63" type="Museum">USNM</collectionCode>
)
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<materialsCitation id="9AC8F270FD61FFBFAE181C22FE3EB9CA" ID-GBIF-Occurrence="3986697010" collectionCode="USNM" collectorName="Male" pageId="62" pageNumber="63" specimenCount="2" typeStatus="paratype">
“CorazalCZ / Pan IV 27 11 / AugBusck //
<taxonomicName id="EDA083AEFD61FFBFADD21C22FC15B9ED" authorityName="Lopez-Garcia &amp; Deloya" authorityYear="2019" box="[739,939,1396,1420]" class="Insecta" family="Dynastidae" genus="Tomarus" kingdom="Animalia" order="Coleoptera" pageId="62" pageNumber="63" phylum="Arthropoda" rank="species" species="laticaudus">Tomarus laticaudus</taxonomicName>
/ López-García &amp; Deloya /
<typeStatus id="F51B468FFD61FFBFABE01C22FAEBB9ED" box="[1233,1365,1396,1420]" pageId="62" pageNumber="63" type="paratype">PARATYPE</typeStatus>
”.
<collectorName id="87559DFBFD61FFBFAA591C22FA22B9ED" box="[1384,1436,1396,1420]" pageId="62" pageNumber="63">Male</collectorName>
<typeStatus id="F51B468FFD61FFBFAFF61CC5FEA0B9CA" box="[199,286,1427,1451]" pageId="62" pageNumber="63" type="paratype">paratype</typeStatus>
(
<collectionCode id="4CB160E8FD61FFBFAE1D1CC5FEC6B9CA" box="[300,376,1427,1451]" country="USA" lsid="urn:lsid:biocol.org:col:34871" name="Smithsonian Institution, National Museum of Natural History" pageId="62" pageNumber="63" type="Museum">USNM</collectionCode>
)
</materialsCitation>
<materialsCitation id="9AC8F270FD61FFBFAEB61CC5FD96B9A8" ID-GBIF-Occurrence="3986696467" collectionCode="USNM" collectorName="A. Busck" pageId="62" pageNumber="63" specimenCount="1" typeStatus="paratype">
“Corazal / CZ / Pn / iv.7.12 /
<collectorName id="87559DFBFD61FFBFAD851CC5FCB2B9CA" box="[692,780,1427,1451]" pageId="62" pageNumber="63">A.Busck</collectorName>
/ light //
<taxonomicName id="EDA083AEFD61FFBFAC5D1CC5FB8CB9CA" authorityName="Lopez-Garcia &amp; Deloya" authorityYear="2019" box="[876,1074,1427,1451]" class="Insecta" family="Dynastidae" genus="Tomarus" kingdom="Animalia" order="Coleoptera" pageId="62" pageNumber="63" phylum="Arthropoda" rank="species" species="laticaudus">Tomarus laticaudus</taxonomicName>
/ López-García &amp; Deloya / PARATYPE”. Female
<typeStatus id="F51B468FFD61FFBFAE411CE4FE79B9AB" box="[368,455,1458,1482]" pageId="62" pageNumber="63" type="paratype">paratype</typeStatus>
(
<collectionCode id="4CB160E8FD61FFBFAEE41CE7FD9FB9A8" box="[469,545,1457,1481]" country="USA" lsid="urn:lsid:biocol.org:col:34871" name="Smithsonian Institution, National Museum of Natural History" pageId="62" pageNumber="63" type="Museum">USNM</collectionCode>
)
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<materialsCitation id="9AC8F270FD61FFBFAD1F1CE7FD13B989" ID-GBIF-Occurrence="3986696457" collectingDate="1971-05-03" collectionCode="UNSM" collectorName="W. E. Bivin" pageId="62" pageNumber="63" specimenCount="2" typeStatus="paratype">
“Diablo Hts. / CZ / Pn /
<date id="5E1EDEEDFD61FFBFAC161CE7FCC3B9A8" box="[807,893,1457,1481]" pageId="62" pageNumber="63" value="1971-05-03">
<collectingDate id="4E5A2705FD61FFBFAC161CE7FCC3B9A8" box="[807,893,1457,1481]" pageId="62" pageNumber="63" value="1971-05-03">3.v.1971</collectingDate>
</date>
/
<collectorName id="87559DFBFD61FFBFACA01CE4FBBCB9A8" box="[913,1026,1457,1481]" pageId="62" pageNumber="63">W.E. Bivin</collectorName>
//
<taxonomicName id="EDA083AEFD61FFBFAB2C1CE4FB5DB9A8" authorityName="Lopez-Garcia &amp; Deloya" authorityYear="2019" box="[1053,1251,1457,1481]" class="Insecta" family="Dynastidae" genus="Tomarus" kingdom="Animalia" order="Coleoptera" pageId="62" pageNumber="63" phylum="Arthropoda" rank="species" species="laticaudus">Tomarus laticaudus</taxonomicName>
/ López-García &amp; Deloya /
<typeStatus id="F51B468FFD61FFBFAE151C86FE16B989" box="[292,424,1488,1512]" pageId="62" pageNumber="63" type="paratype">PARATYPE</typeStatus>
”. Male
<typeStatus id="F51B468FFD61FFBFAEC41C86FDF2B989" box="[501,588,1488,1512]" pageId="62" pageNumber="63" type="paratype">paratype</typeStatus>
(
<collectionCode id="4CB160E8FD61FFBFAD681C86FD1BB989" box="[601,677,1488,1512]" country="USA" lsid="urn:lsid:biocol.org:col:34868" name="University of Nebraska State Museum" pageId="62" pageNumber="63">UNSM</collectionCode>
)
</materialsCitation>
<materialsCitation id="9AC8F270FD61FFBFAD821C86FBA2B989" ID-GBIF-Occurrence="3986696305" box="[691,1052,1488,1512]" collectionCode="USNM" pageId="62" pageNumber="63" specimenCount="1" specimenCount-female="1" typeStatus="paratype">
and
<specimenCount id="3CA633A4FD61FFBFADEF1C86FCEAB989" box="[734,852,1488,1512]" pageId="62" pageNumber="63" type="female">four female</specimenCount>
(
<collectionCode id="4CB160E8FD61FFBFAC501C86FC13B989" box="[865,941,1488,1512]" country="USA" lsid="urn:lsid:biocol.org:col:34871" name="Smithsonian Institution, National Museum of Natural History" pageId="62" pageNumber="63" type="Museum">USNM</collectionCode>
)
<typeStatus id="F51B468FFD61FFBFAC8A1C86FBA2B989" box="[955,1052,1488,1512]" pageId="62" pageNumber="63" type="paratype">paratypes</typeStatus>
</materialsCitation>
<materialsCitation id="9AC8F270FD61FFBFAB131C86FA22BA66" ID-GBIF-Occurrence="3986696597" collectionCode="UNSM, USNM" collectorName="Four" pageId="62" pageNumber="63" specimenCount="2" typeStatus="paratype">
“Arc-light / Ancon, C.Z. / Pan. 12. V. II / AH Jennings //
<taxonomicName id="EDA083AEFD61FFBFAEB81CB9FDF1BA66" authorityName="Lopez-Garcia &amp; Deloya" authorityYear="2019" box="[393,591,1519,1543]" class="Insecta" family="Dynastidae" genus="Tomarus" kingdom="Animalia" order="Coleoptera" pageId="62" pageNumber="63" phylum="Arthropoda" rank="species" species="laticaudus">Tomarus laticaudus</taxonomicName>
/ López-García &amp; Deloya /
<typeStatus id="F51B468FFD61FFBFAC5C1CB9FC4FBA66" box="[877,1009,1519,1543]" pageId="62" pageNumber="63" type="paratype">PARATYPE</typeStatus>
”.
<collectorName id="87559DFBFD61FFBFAB331CB9FB8CBA67" box="[1026,1074,1519,1542]" pageId="62" pageNumber="63">Four</collectorName>
female
<typeStatus id="F51B468FFD61FFBFABB51CB9FB5BBA66" box="[1156,1253,1519,1543]" pageId="62" pageNumber="63" type="paratype">paratypes</typeStatus>
(
<collectionCode id="4CB160E8FD61FFBFABC21CB9FAFEBA66" box="[1267,1344,1519,1543]" country="USA" lsid="urn:lsid:biocol.org:col:34868" name="University of Nebraska State Museum" pageId="62" pageNumber="63">UNSM</collectionCode>
,
<collectionCode id="4CB160E8FD61FFBFAA7A1CB9FA28BA66" box="[1355,1430,1519,1543]" country="USA" lsid="urn:lsid:biocol.org:col:34871" name="Smithsonian Institution, National Museum of Natural History" pageId="62" pageNumber="63" type="Museum">USNM</collectionCode>
)
</materialsCitation>
<materialsCitation id="9AC8F270FD61FFBFAFF61F5BFE3EBA25" ID-GBIF-Occurrence="3986696744" collectingDate="1911-04" collectionCode="UNSM" collectorName="Female" pageId="62" pageNumber="63" specimenCount="2" typeStatus="paratype">
“arc-light globe / Ancon, CZPan /
<date id="5E1EDEEDFD61FFBFAD151F58FD38BA44" box="[548,646,1549,1574]" pageId="62" pageNumber="63" value="1911-04">
<collectingDate id="4E5A2705FD61FFBFAD151F58FD38BA44" box="[548,646,1549,1574]" pageId="62" pageNumber="63" value="1911-04">Apr. 1911</collectingDate>
</date>
Kraft //
<taxonomicName id="EDA083AEFD61FFBFADE91F58FC20BA44" authorityName="Lopez-Garcia &amp; Deloya" authorityYear="2019" box="[728,926,1549,1573]" class="Insecta" family="Dynastidae" genus="Tomarus" kingdom="Animalia" order="Coleoptera" pageId="62" pageNumber="63" phylum="Arthropoda" rank="species" species="laticaudus">Tomarus laticaudus</taxonomicName>
/ López-García &amp; Deloya /
<typeStatus id="F51B468FFD61FFBFAB8D1F5BFAFEBA44" box="[1212,1344,1549,1573]" pageId="62" pageNumber="63" type="paratype">PARATYPE</typeStatus>
”.
<collectorName id="87559DFBFD61FFBFAA631F5BFA23BA44" box="[1362,1437,1549,1573]" pageId="62" pageNumber="63">Female</collectorName>
<typeStatus id="F51B468FFD61FFBFAFF61F7AFEA0BA25" box="[199,286,1580,1604]" pageId="62" pageNumber="63" type="paratype">paratype</typeStatus>
(
<collectionCode id="4CB160E8FD61FFBFAE1C1F7AFEC7BA25" box="[301,377,1580,1604]" country="USA" lsid="urn:lsid:biocol.org:col:34868" name="University of Nebraska State Museum" pageId="62" pageNumber="63">UNSM</collectionCode>
)
</materialsCitation>
<materialsCitation id="9AC8F270FD61FFBFAEB61F7AFDB4BA03" ID-GBIF-Occurrence="3986696641" country="Panama" location="Corozal" pageId="62" pageNumber="63" specimenCount="1" typeStatus="paratype">
“Paraiso CZ / iv.24.11 / August Busck //
<taxonomicName id="EDA083AEFD61FFBFAC1A1F7AFC4CBA25" authorityName="Lopez-Garcia &amp; Deloya" authorityYear="2019" box="[811,1010,1580,1604]" class="Insecta" family="Dynastidae" genus="Tomarus" kingdom="Animalia" order="Coleoptera" pageId="62" pageNumber="63" phylum="Arthropoda" rank="species" species="laticaudus">Tomarus laticaudus</taxonomicName>
/ López-García &amp; Deloya /
<typeStatus id="F51B468FFD61FFBFAA221F7AFA2ABA25" box="[1299,1428,1580,1604]" pageId="62" pageNumber="63" type="paratype">PARATYPE</typeStatus>
” Type locality:
<location id="2F7FAEF6FD61FFBFAE681F1DFE12BA02" LSID="urn:lsid:plazi:treatment:A209493BFD61FFBEAFA61D99FEC0BF43:2F7FAEF6FD61FFBFAE681F1DFE12BA02" box="[345,428,1611,1635]" country="Panama" name="Corozal" pageId="62" pageNumber="63">Corozal</location>
,
<collectingCountry id="52B7B8BDFD61FFBFAE871F1DFDB4BA03" box="[438,522,1611,1634]" name="Panama" pageId="62" pageNumber="63">Panama</collectingCountry>
</materialsCitation>
.
</paragraph>
</subSubSection>
<subSubSection id="62BAABA6FD61FFBEAFA61FC7FB77BDD3" lastPageId="63" lastPageNumber="64" pageId="62" pageNumber="63" type="description">
<paragraph id="2A1FF82DFD61FFBEAFA61FC7FB77BDD3" blockId="62.[151,1437,1681,2031]" lastBlockId="63.[151,1437,151,802]" lastPageId="63" lastPageNumber="64" pageId="62" pageNumber="63">
<emphasis id="18D4243FFD61FFBFAFA61FC7FE97BACA" bold="true" box="[151,297,1681,1707]" pageId="62" pageNumber="63">Description.</emphasis>
Habitus as in
<figureCitation id="B29BE4A8FD61FFBFAEFA1FC7FD8DBACA" box="[459,563,1681,1707]" captionStart="FIGURE 27" captionStartId="102.[151,250,1868,1892]" captionTargetBox="[158,1409,184,1839]" captionTargetId="figure-24@102.[151,1436,181,1844]" captionTargetPageId="102" captionText="FIGURE 27. Habitus of the species of Tomarus: A) Tomarus adoceteus, B) T. amazonicus, C) T. bituberculatus, D) T. colombianus, E) T. discrepans, F) T. gyas, G) T. laevicollis, H) T. laticaudus, I) T. maimon. Scale bar: 5 mm." figureDoi="http://doi.org/10.5281/zenodo.7345103" httpUri="https://zenodo.org/record/7345103/files/figure.png" pageId="62" pageNumber="63">Fig. 27H</figureCitation>
. Length
<quantity id="ED5855C8FD61FFBFADA41FC7FC85BACA" box="[661,827,1681,1707]" metricMagnitude="-2" metricUnit="m" metricValue="2.16" metricValueMax="2.44" metricValueMin="1.88" pageId="62" pageNumber="63" unit="mm" value="21.6" valueMax="24.4" valueMin="18.8">18.8–24.4 mm</quantity>
; humeral width
<quantity id="ED5855C8FD61FFBFACC31FC7FB35BACA" box="[1010,1163,1681,1707]" metricMagnitude="-2" metricUnit="m" metricValue="1.105" metricValueMax="1.22" metricValueMin="0.99" pageId="62" pageNumber="63" unit="mm" value="11.05" valueMax="12.2" valueMin="9.9">9.9–12.2 mm</quantity>
. Color reddish brown to black.
<emphasis id="18D4243FFD61FFBFAFD51FE3FE93BAAE" bold="true" box="[228,301,1717,1743]" pageId="62" pageNumber="63">Head:</emphasis>
Frons coarsely rugose. Frontoclypeal region with 2 conical tubercles separated by about 7 tubercle diameters. Clypeus narrowed towards apex, base 3 times wider than apex. Clypeal teeth triangular, separated by less than a tooth diameter. Mandible with 2 apical teeth and a lateral, narrowly rounded tooth. Mentum abruptly constricted at apical 3rd. Galea of maxilla with teeth 5 and 6 well developed. Interocular distance 3.3 times an eye width.
<emphasis id="18D4243FFD61FFBFAE261E13FE27BB3E" bold="true" box="[279,409,1861,1887]" pageId="62" pageNumber="63">Pronotum:</emphasis>
Surface smooth on disc, with minute punctures on lateral margins. Apical tubercle small, rounded, not visible in lateral view. Fovea shallow, striate, and about 1/2 as wide as interocular distance.
<emphasis id="18D4243FFD61FFBFAA7B1E3FFF71BBC6" bold="true" pageId="62" pageNumber="63">Scutellum:</emphasis>
Surface without or with sparse, small punctures, not forming lines parallel to margins.
<emphasis id="18D4243FFD61FFBFAB9F1EDBFABDBBC6" bold="true" box="[1198,1283,1933,1959]" pageId="62" pageNumber="63">Elytra:</emphasis>
First interval punctate, punctures smaller than on other intervals, sutural stria complete. Inner surface of apex with rounded, large tubercles forming 14–16 parallel lines.
<emphasis id="18D4243FFD61FFBFAD7A1E83FD77BB8E" bold="true" box="[587,713,2005,2031]" pageId="62" pageNumber="63">Abdomen:</emphasis>
Apex of tergite IV with a triangular wide area with small, irregular tubercles forming 9–10 diagonal nearly parallel lines. Pygidial surface rugose on basal 4th and lateral angles, disc with minute punctures; apex regularly rounded in male, strongly dilated in female.
<emphasis id="18D4243FFD60FFBEAB5D19EAFB7BBCB7" bold="true" box="[1132,1221,188,214]" pageId="63" pageNumber="64">Venter:</emphasis>
Apex of prosternal process flat, transversely oval to rounded. Metasternum with minute setae.
<emphasis id="18D4243FFD60FFBEACD719B6FB99BC9B" bold="true" box="[998,1063,224,250]" pageId="63" pageNumber="64">Legs:</emphasis>
Protibia tridentate, with an additional small, basal denticle. Metatibia not narrowed before apex, sides nearly parallel. Apex of metatibia crenulate; with 10–12 spinules.
<emphasis id="18D4243FFD60FFBEAEA0187CFDF7BD25" bold="true" box="[401,585,298,324]" pageId="63" pageNumber="64">Male genitalia:</emphasis>
Spiculum gastrale without basal plates. Phallobase 1.2 times longer than parameres. Parameres with 2 pairs of dorso-lateral teeth; basal tooth long, wide, and upwardly arched, apical tooth short and joined to the basal tooth (
<figureCitation id="B29BE4A8FD60FFBEAD161825FD24BDEC" box="[551,666,371,397]" captionStart="FIGURE 16" captionStartId="91.[151,250,1633,1657]" captionTargetBox="[151,1426,189,1602]" captionTargetId="figure-17@91.[151,1436,181,1610]" captionTargetPageId="91" captionText="FIGURE 16. Parameres in frontal view. A) Tomarus adoceteus, B) T. amazonicus, C) T. bituberculatus, D) T. colombianus, E) T. discrepans, F) T. gyas, G) T. laevicollis, H) T. laticaudus, I) T. maimon, J) T. maracaiboensis, K) T. maternus, L) T. pilcopataensis, M) T. pullus, N) T. pumilus, O) T. roigjunenti, P) T. rostratus, Q) T. selanderi, R) T. subtropicus." figureDoi="http://doi.org/10.5281/zenodo.7345065" httpUri="https://zenodo.org/record/7345065/files/figure.png" pageId="63" pageNumber="64">Figs. 16H</figureCitation>
,
<figureCitation id="B29BE4A8FD60FFBEAD971822FD64BDEF" box="[678,730,372,398]" captionStart="FIGURE 19" captionStartId="94.[151,250,1155,1179]" captionTargetBox="[157,1432,188,1128]" captionTargetId="figure-24@94.[152,1436,181,1131]" captionTargetPageId="94" captionText="FIGURE 19. Parameres in lateral view. A) Tomarus adoceteus, B) T. amazonicus, C) T. bituberculatus, D) T. colombianus, E) T. discrepans, F) T. gyas, G) T. laevicollis, H) T. laticaudus, I) T. maimon." figureDoi="http://doi.org/10.5281/zenodo.7345079" httpUri="https://zenodo.org/record/7345079/files/figure.png" pageId="63" pageNumber="64">19H</figureCitation>
). Internal sac with copulatory lamella and short lamellar spiny belt; with a complex of 7 spine-like, accessory lamellae, with dense granules at base (
<figureCitation id="B29BE4A8FD60FFBEAB6718CEFB02BDD3" box="[1110,1212,408,434]" captionStart="FIGURE 23" captionStartId="98.[151,250,1728,1752]" captionTargetBox="[161,1421,189,1693]" captionTargetId="figure-24@98.[151,1436,181,1704]" captionTargetPageId="98" captionText="FIGURE 23. Internal sac. A) Tomarus amazonicus, B) T. discrepans, C) T. laticaudus, D) T. maimon. Abbreviations: AL: Accessory lamellae, CL: Copulatory lamella, LSB: Lamellar spiny belt, Ra: Raspula, SD: Sclerotized diverticulum." figureDoi="http://doi.org/10.5281/zenodo.7345091" httpUri="https://zenodo.org/record/7345091/files/figure.png" pageId="63" pageNumber="64">Fig. 23C</figureCitation>
).
</paragraph>
</subSubSection>
<subSubSection id="62BAABA6FD60FFBEAFF618EBFEA8BE0B" pageId="63" pageNumber="64" type="diagnosis">
<paragraph id="2A1FF82DFD60FFBEAFF618EBFEA8BE0B" blockId="63.[151,1437,151,802]" pageId="63" pageNumber="64">
<emphasis id="18D4243FFD60FFBEAFF618EBFEFCBDB6" bold="true" box="[199,322,445,471]" pageId="63" pageNumber="64">Diagnosis.</emphasis>
<taxonomicName id="EDA083AEFD60FFBEAE7818E8FD98BDB6" authorityName="Lopez-Garcia &amp; Deloya" authorityYear="2019" box="[329,550,445,471]" class="Insecta" family="Dynastidae" genus="Tomarus" kingdom="Animalia" order="Coleoptera" pageId="63" pageNumber="64" phylum="Arthropoda" rank="species" species="laticaudus">
<emphasis id="18D4243FFD60FFBEAE7818E8FD98BDB6" box="[329,550,445,471]" italics="true" pageId="63" pageNumber="64">Tomarus laticaudus</emphasis>
</taxonomicName>
can be recognized by the nearly impunctate pronotum; small pronotal tubercle not visible in lateral view; pronotal fovea shallow and narrow (1/2 as wide as interocular distance); and protibia tridentate with an additional basal 4th tooth. It is similar to
<taxonomicName id="EDA083AEFD60FFBEAC0C1B51FC73BE40" baseAuthorityName="Bates" baseAuthorityYear="1888" box="[829,973,519,545]" class="Insecta" family="Dynastidae" genus="Tomarus" kingdom="Animalia" order="Coleoptera" pageId="63" pageNumber="64" phylum="Arthropoda" rank="species" species="laevicollis">
<emphasis id="18D4243FFD60FFBEAC0C1B51FC73BE40" box="[829,973,519,545]" italics="true" pageId="63" pageNumber="64">T. laevicollis</emphasis>
</taxonomicName>
but can be differentiated by the strongly dilated pygidial apex in female, and the parameres with the main teeth wider and joined to secondary teeth (
<figureCitation id="B29BE4A8FD60FFBEAA551B7DFF77BE0B" captionStart="FIGURE 16" captionStartId="91.[151,250,1633,1657]" captionTargetBox="[151,1426,189,1602]" captionTargetId="figure-17@91.[151,1436,181,1610]" captionTargetPageId="91" captionText="FIGURE 16. Parameres in frontal view. A) Tomarus adoceteus, B) T. amazonicus, C) T. bituberculatus, D) T. colombianus, E) T. discrepans, F) T. gyas, G) T. laevicollis, H) T. laticaudus, I) T. maimon, J) T. maracaiboensis, K) T. maternus, L) T. pilcopataensis, M) T. pullus, N) T. pumilus, O) T. roigjunenti, P) T. rostratus, Q) T. selanderi, R) T. subtropicus." figureDoi="http://doi.org/10.5281/zenodo.7345065" httpUri="https://zenodo.org/record/7345065/files/figure.png" pageId="63" pageNumber="64">Figs. 16H</figureCitation>
,
<figureCitation id="B29BE4A8FD60FFBEAFE41B06FEB7BE0B" box="[213,265,592,618]" captionStart="FIGURE 19" captionStartId="94.[151,250,1155,1179]" captionTargetBox="[157,1432,188,1128]" captionTargetId="figure-24@94.[152,1436,181,1131]" captionTargetPageId="94" captionText="FIGURE 19. Parameres in lateral view. A) Tomarus adoceteus, B) T. amazonicus, C) T. bituberculatus, D) T. colombianus, E) T. discrepans, F) T. gyas, G) T. laevicollis, H) T. laticaudus, I) T. maimon." figureDoi="http://doi.org/10.5281/zenodo.7345079" httpUri="https://zenodo.org/record/7345079/files/figure.png" pageId="63" pageNumber="64">19H</figureCitation>
).
</paragraph>
</subSubSection>
<subSubSection id="62BAABA6FD60FFBEAFF61B23FEC0BF43" pageId="63" pageNumber="64" type="distribution">
<paragraph id="2A1FF82DFD60FFBEAFF61B23FC91BEEE" blockId="63.[151,1437,151,802]" box="[199,815,629,656]" pageId="63" pageNumber="64">
<emphasis id="18D4243FFD60FFBEAFF61B23FEDFBEEE" bold="true" box="[199,353,629,655]" pageId="63" pageNumber="64">Distribution.</emphasis>
<collectingCountry id="52B7B8BDFD60FFBEAE561B20FE7EBEF1" box="[359,448,630,656]" name="Panama" pageId="63" pageNumber="64">Panama</collectingCountry>
(
<bibRefCitation id="4E3185DCFD60FFBEAEFF1B23FC9DBEEE" author="Lopez-Garcia, M. M. &amp; Deloya, C." box="[462,803,629,655]" pageId="63" pageNumber="64" pagination="127 - 141" refId="ref91328" refString="Lopez-Garcia, M. M. &amp; Deloya, C. (2019) Five new species of the dynastine genus Tomarus (Coleoptera: Scarabaeidae), with an illustrated key to species. The Coleopterists Bulletin, 73, 127 - 141. https: // doi. org / 10.1649 / 0010 - 065 X- 73.1.127" type="journal article" year="2019">López-García &amp; Deloya 2019</bibRefCitation>
).
</paragraph>
<paragraph id="2A1FF82DFD60FFBEAFF61BCCFC1ABEB9" blockId="63.[151,1437,151,802]" pageId="63" pageNumber="64">
<emphasis id="18D4243FFD60FFBEAFF61BCCFE47BED2" bold="true" box="[199,505,665,692]" pageId="63" pageNumber="64">
Locality records (
<figureCitation id="B29BE4A8FD60FFBEAEA61BCFFE52BED2" box="[407,492,665,691]" captionStart="FIGURE 44" captionStartId="112.[151,250,1998,2022]" captionTargetBox="[224,1372,1135,1965]" captionTargetId="figure-38@112.[215,1372,1135,1973]" captionTargetPageId="112" captionText="FIGURE 44. Distributional records of Tomarus laevicollis, T. laticaudus, T. maracaiboensis, and T. colombianus." figureDoi="http://doi.org/10.5281/zenodo.7345164" httpUri="https://zenodo.org/record/7345164/files/figure.png" pageId="63" pageNumber="64">Fig. 44</figureCitation>
).
</emphasis>
<materialsCitation id="9AC8F270FD60FFBEAECE1BCCFC21BEB9" ID-GBIF-Occurrence="3986696662" collectionCode="UNSM, USNM" country="Panama" location="Ancon" municipality="Corozal" pageId="63" pageNumber="64" specimenCount="1" stateProvince="Los Santos">
21 examined specimens from
<collectionCode id="4CB160E8FD60FFBEAC7D1BCCFC23BED5" box="[844,925,666,692]" country="USA" lsid="urn:lsid:biocol.org:col:34868" name="University of Nebraska State Museum" pageId="63" pageNumber="64">UNSM</collectionCode>
and
<collectionCode id="4CB160E8FD60FFBEACE31BCCFB99BED5" box="[978,1063,666,692]" country="USA" lsid="urn:lsid:biocol.org:col:34871" name="Smithsonian Institution, National Museum of Natural History" pageId="63" pageNumber="64" type="Museum">USNM</collectionCode>
.
<emphasis id="18D4243FFD60FFBEAB011BCCFA22BED2" bold="true" box="[1072,1436,665,692]" pageId="63" pageNumber="64">
<collectingCountry id="52B7B8BDFD60FFBEAB011BCCFB17BED5" box="[1072,1193,666,692]" name="Panama" pageId="63" pageNumber="64">PANAMA</collectingCountry>
(21).
<collectingRegion id="E86436CFFD60FFBEABDA1BCCFAD2BED5" box="[1259,1388,666,692]" country="Panama" name="Los Santos" pageId="63" pageNumber="64">Los Santos</collectingRegion>
(8):
</emphasis>
<collectingMunicipality id="CA7B6257FD60FFBEAFA61BE9FF4CBEB8" box="[151,242,703,729]" pageId="63" pageNumber="64">Corozal</collectingMunicipality>
, Paraíso.
<emphasis id="18D4243FFD60FFBEAE6F1BE8FE41BEB9" bold="true" box="[350,511,702,729]" pageId="63" pageNumber="64">
<collectingRegion id="E86436CFFD60FFBEAE6F1BE8FE7EBEB8" box="[350,448,702,729]" country="Panama" name="Panama" pageId="63" pageNumber="64">Panamá</collectingRegion>
(12):
</emphasis>
<location id="2F7FAEF6FD60FFBEAD371BE9FDEABEB8" LSID="urn:lsid:plazi:treatment:A209493BFD61FFBEAFA61D99FEC0BF43:2F7FAEF6FD60FFBEAD371BE9FDEABEB8" box="[518,596,703,729]" country="Panama" municipality="Corozal" name="Ancon" pageId="63" pageNumber="64" stateProvince="Los Santos">Ancon</location>
; Diablo Heights.
<emphasis id="18D4243FFD60FFBEAC291BE9FC21BEB9" bold="true" box="[792,927,702,729]" pageId="63" pageNumber="64">No data (1)</emphasis>
</materialsCitation>
<emphasis id="18D4243FFD60FFBEACAF1BE8FC1ABEB9" bold="true" box="[926,932,702,728]" pageId="63" pageNumber="64">.</emphasis>
</paragraph>
<paragraph id="2A1FF82DFD60FFBEAFF61BB2FEC0BF43" blockId="63.[151,1437,151,802]" pageId="63" pageNumber="64">
<emphasis id="18D4243FFD60FFBEAFF61BB2FE3BBE9C" bold="true" box="[199,389,739,766]" pageId="63" pageNumber="64">Natural history.</emphasis>
The species inhabits tropical moist forest areas between
<quantity id="ED5855C8FD60FFBEAB301BB2FBC2BE9F" box="[1025,1148,740,766]" metricMagnitude="1" metricUnit="m" metricValue="4.5" metricValueMax="8.0" metricValueMin="1.0" pageId="63" pageNumber="64" unit="m" value="45.0" valueMax="80.0" valueMin="10.0">
10 to
<specimenCount id="3CA633A4FD60FFBEAB711BB2FBC2BE9F" box="[1088,1148,740,766]" pageId="63" pageNumber="64" type="generic">80 m</specimenCount>
</quantity>
. It has been found mainly in April using lights.
</paragraph>
</subSubSection>
</treatment>
</document>
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