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<mods:title id="28EE664B3571FE77CF807204AE4F2382">Cladistic analysis reveals polyphyly of Tomarus (Coleoptera: Scarabaeidae: Dynastinae): new classification and taxonomic revision</mods:title>
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<taxonomicName id="EDA083AEFD65FFBBAFA619C1FC3EBCD0" authority="Lopez-Garcia &amp; Gasca-Alvarez, 2014" authorityName="Lopez-Garcia &amp; Gasca-Alvarez" authorityYear="2014" box="[151,896,150,181]" class="Insecta" family="Dynastidae" genus="Tomarus" kingdom="Animalia" order="Coleoptera" pageId="58" pageNumber="59" phylum="Arthropoda" rank="species" species="colombianus">
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<emphasis id="18D4243FFD65FFBBAFA619C1FE1ABCD3" bold="true" box="[151,420,151,178]" italics="true" pageId="58" pageNumber="59">Tomarus colombianus</emphasis>
<bibRefCitation id="4E3185DCFD65FFBBAE9A19C0FC3EBCD0" author="Lopez-Garcia, M. M. &amp; Gasca-Alvarez, H. J. &amp; Amat-Garcia, G." box="[427,896,150,181]" pageId="58" pageNumber="59" pagination="579 - 584" refId="ref91387" refString="Lopez-Garcia, M. M., Gasca-Alvarez, H. J. &amp; Amat-Garcia, G. (2014) A new species of Tomarus Erichson, 1847 (Coleoptera: Scarabaeoidea: Dynastinae: Pentodontini), with a key to the species in Colombia. Zootaxa, 3869 (5), 579 - 584. https: // doi. org / 10.11646 / zootaxa. 3869.5.8" type="journal article" year="2014">López-García &amp; Gasca-Álvarez, 2014</bibRefCitation>
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(
<figureCitation id="B29BE4A8FD65FFBBAFAE19EDFEBDBCB4" box="[159,259,187,213]" captionStart="FIGURE 9" captionStartId="87.[151,250,1074,1098]" captionTargetBox="[151,1436,181,1050]" captionTargetId="figure-17@87.[151,1436,181,1050]" captionTargetPageId="87" captionText="FIGURE 9. Pygidium of male. A) Euligyrus ebenus, B) Proculigyrus cicatricosus, C) Ligyrus (Anagrylius) cuniculus, D) L. (Ligyrus) fossor, E) L. (Ligyrus) paranaensis, F) Tomarus bituberculatus, G) T. colombianus, H) T. maracaiboensis, I) T. pilcopataensis, J) T. pullus, K) T. roigjunenti, L) T. rostratus." figureDoi="http://doi.org/10.5281/zenodo.7345041" httpUri="https://zenodo.org/record/7345041/files/figure.png" pageId="58" pageNumber="59">Figs. 9G</figureCitation>
,
<figureCitation id="B29BE4A8FD65FFBBAE3E19EDFE88BCB4" box="[271,310,187,213]" captionStart="FIGURE 11" captionStartId="88.[151,250,931,955]" captionTargetBox="[234,1354,186,899]" captionTargetId="figure-24@88.[231,1357,181,907]" captionTargetPageId="88" captionText="FIGURE 11. Protibia in frontal view.A) Euligyrus similis, B) Proculigyrus cicatricosus, C) Ligyrus (Anagrylius) moroni, D) L. (Ligyrodes) relictus, E) L. (Ligyrodes) sallaei, F) L. (Ligyrus) allonasutus, G) L. (Ligyrus) villosus, H) Tomarus bituberculatus, I) T. colombianus, J) T. pullus." figureDoi="http://doi.org/10.5281/zenodo.7345051" httpUri="https://zenodo.org/record/7345051/files/figure.png" pageId="58" pageNumber="59">11I</figureCitation>
,
<figureCitation id="B29BE4A8FD65FFBBAE7019EDFECABCB4" box="[321,372,187,213]" 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="58" pageNumber="59">16D</figureCitation>
,
<figureCitation id="B29BE4A8FD65FFBBAE4E19EDFE0CBCB4" box="[383,434,187,213]" 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="58" pageNumber="59">19D</figureCitation>
,
<figureCitation id="B29BE4A8FD65FFBBAE8F19EDFE4FBCB4" box="[446,497,187,213]" 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="58" pageNumber="59">27D</figureCitation>
;
<figureCitation id="B29BE4A8FD65FFBBAECC19EDFDA4BCB4" box="[509,538,187,213]" 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="58" pageNumber="59">44</figureCitation>
)
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<taxonomicName id="EDA083AEFD65FFBBAFA61856FCB2BD79" authority="Lopez-Garcia &amp; Gasca-Alvarez, 2014" authorityName="Lopez-Garcia &amp; Gasca-Alvarez" authorityYear="2014" box="[151,780,256,282]" class="Insecta" family="Dynastidae" genus="Tomarus" kingdom="Animalia" order="Coleoptera" pageId="58" pageNumber="59" phylum="Arthropoda" rank="species" species="colombianus">
<emphasis id="18D4243FFD65FFBBAFA61856FEC7BD79" box="[151,377,256,280]" italics="true" pageId="58" pageNumber="59">Tomarus colombianus</emphasis>
<bibRefCitation id="4E3185DCFD65FFBBAEB21856FCB2BD79" author="Lopez-Garcia, M. M. &amp; Gasca-Alvarez, H. J. &amp; Amat-Garcia, G." box="[387,780,256,282]" pageId="58" pageNumber="59" pagination="579 - 584" refId="ref91387" refString="Lopez-Garcia, M. M., Gasca-Alvarez, H. J. &amp; Amat-Garcia, G. (2014) A new species of Tomarus Erichson, 1847 (Coleoptera: Scarabaeoidea: Dynastinae: Pentodontini), with a key to the species in Colombia. Zootaxa, 3869 (5), 579 - 584. https: // doi. org / 10.11646 / zootaxa. 3869.5.8" type="journal article" year="2014">
López-García &amp; Gasca-Álvarez,
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in
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López-García
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. 2014: 579
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. Original combination. Male
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(CIUQ) “Colombia—Valle / La Bocana-B/ventura / Altura:
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s.n.m / Fecha: 16/04/04 / Método: Directo / Col: Serna F. //
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/ López-García &amp; / Gasca-Álvarez / HOLOTIPO // CIUQ 1505”. Female
<typeStatus id="F51B468FFD65FFBBABC0186BFAF6BD34" box="[1265,1352,317,341]" pageId="58" pageNumber="59" type="paratype">paratype</typeStatus>
(CIUQ) “
<collectingCountry id="52B7B8BDFD65FFBBAFE1180AFEE3BD15" box="[208,349,348,372]" name="Colombia" pageId="58" pageNumber="59">COLOMBIA</collectingCountry>
:
<collectingRegion id="E86436CFFD65FFBBAE59180AFDFEBD15" box="[360,576,348,372]" country="Colombia" name="Valle del Cauca" pageId="58" pageNumber="59">Dep. Valle del Cauca</collectingRegion>
/ Buenaventura / La Bocana /
<date id="5E1EDEEDFD65FFBBAC4D180DFC59BD12" box="[892,999,347,371]" pageId="58" pageNumber="59" value="2004-04-15">15.iv.2004</date>
/ Light trap /
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altitud / Rivas L. Coll //
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<emphasis id="18D4243FFD65FFBBAFF6182DFE18BDF3" box="[199,422,378,402]" italics="true" pageId="58" pageNumber="59">Tomarus colombianus</emphasis>
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/ López-García &amp; / Gasca-Álvarez / PARATIPO // CIUQ 1504”. Female
<typeStatus id="F51B468FFD65FFBBABA0182DFB56BDF2" box="[1169,1256,379,403]" pageId="58" pageNumber="59" type="paratype">paratype</typeStatus>
(CIUQ) “Colombia- Valle
<collectingRegion id="E86436CFFD65FFBBAE0018CFFECEBDD0" box="[305,368,409,433]" country="Colombia" name="Cauca" pageId="58" pageNumber="59">Cauca</collectingRegion>
/ B/ventura-Bocana / Altura:
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.
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s.n.m / Fecha: Mayo/04 / Método: Directo-Bosque / Col: Mendoza L. //
<taxonomicName id="EDA083AEFD65FFBBAFEC18EEFE00BDB1" authorityName="Lopez-Garcia &amp; Gasca-Alvarez" authorityYear="2014" box="[221,446,440,464]" class="Insecta" family="Dynastidae" genus="Tomarus" kingdom="Animalia" order="Coleoptera" pageId="58" pageNumber="59" phylum="Arthropoda" rank="species" species="colombianus">
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/ López-García &amp; / Gasca-Álvarez / PARATIPO // CIUQ 1507”. Male parat–pe (ICN) “Colombia—Valle / La Bocana-B/ventura / Altura:
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s.n.m / Fecha: 17/04/04 / Método: Directo / Col: Serna F.” Female
<typeStatus id="F51B468FFD65FFBBAA741881FA22BD8E" box="[1349,1436,471,495]" pageId="58" pageNumber="59" type="paratype">paratype</typeStatus>
(ICN) “
<collectingRegion id="E86436CFFD65FFBBAE2518A3FEF1BE6C" box="[276,335,501,525]" country="Colombia" name="Huila" pageId="58" pageNumber="59">Huila</collectingRegion>
/ Neiva /
<date id="5E1EDEEDFD65FFBBAE8618A3FDA6BE6C" box="[439,536,501,525]" pageId="58" pageNumber="59" value="1974-03-05">5.iii.1974</date>
/ Bagos A. I. //
<taxonomicName id="EDA083AEFD65FFBBADF118A0FC1EBE6C" authorityName="Lopez-Garcia &amp; Gasca-Alvarez" authorityYear="2014" box="[704,928,501,525]" class="Insecta" family="Dynastidae" genus="Tomarus" kingdom="Animalia" order="Coleoptera" pageId="58" pageNumber="59" phylum="Arthropoda" rank="species" species="colombianus">
<emphasis id="18D4243FFD65FFBBADF118A0FC1EBE6C" box="[704,928,501,525]" italics="true" pageId="58" pageNumber="59">Tomarus colombianus</emphasis>
</taxonomicName>
/ López-García &amp; / Gasca-Álvarez / PARATIPO. Type locality: La Bocana,
<collectingRegion id="E86436CFFD65FFBBAEE31B42FDCABE4D" box="[466,628,532,556]" country="Colombia" name="Valle del Cauca" pageId="58" pageNumber="59">Valle del Cauca</collectingRegion>
, Colombia.
</paragraph>
</subSubSection>
<subSubSection id="62BAABA6FD65FFBBAFA61B02FC82B8CF" pageId="58" pageNumber="59" type="description">
<paragraph id="2A1FF82DFD65FFBBAFA61B02FC82B8CF" blockId="58.[151,1437,596,1559]" pageId="58" pageNumber="59">
<emphasis id="18D4243FFD65FFBBAFA61B02FE97BE0F" bold="true" box="[151,297,596,622]" pageId="58" pageNumber="59">Description.</emphasis>
Habitus as in
<figureCitation id="B29BE4A8FD65FFBBAEF31B02FD96BE0E" box="[450,552,596,623]" 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="58" pageNumber="59">Fig. 27D</figureCitation>
. Length 18.0–21.0 mm; humeral width
<quantity id="ED5855C8FD65FFBBACEB1B02FBCFBE0E" box="[986,1137,596,623]" metricMagnitude="-3" metricUnit="m" metricValue="9.5" metricValueMax="10.5" metricValueMin="8.5" pageId="58" pageNumber="59" unit="mm" value="9.5" valueMax="10.5" valueMin="8.5">8.5–10.5 mm</quantity>
. Color dark reddish brown.
<emphasis id="18D4243FFD65FFBBAFA61B2FFF5EBEF2" bold="true" box="[151,224,633,659]" pageId="58" pageNumber="59">Head:</emphasis>
Frons and clypeus coarsely and densely rugose. Frontoclypeal region with 2 conical tubercles separated by about 6 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 3.2 times an eye width.
<emphasis id="18D4243FFD65FFBBAE4F1A5FFDBEBF42" bold="true" box="[382,512,777,803]" pageId="58" pageNumber="59">Pronotum:</emphasis>
Surface with minute punctures; denser and deeper on anterior and lateral angles, disc impunctate. Apical tubercle small, rounded, not visible in lateral view. Subapical fovea shallow, elongate, narrow (1/3 the interocular distance); equal in both sexes, surface rugopunctate.
<emphasis id="18D4243FFD65FFBBAB301A06FB3CBF0B" bold="true" box="[1025,1154,848,874]" pageId="58" pageNumber="59">Scutellum:</emphasis>
Surface with deep, large punctures forming 2 lines parallel to margins.
<emphasis id="18D4243FFD65FFBBADA61A23FD53BFEE" bold="true" box="[663,749,885,911]" pageId="58" pageNumber="59">Elytra:</emphasis>
Punctures on first interval as large as those on other intervals. Inner surface of apex with rounded, large tubercles forming 14–16 parallel lines.
<emphasis id="18D4243FFD65FFBBAB291ACEFB28BFD3" bold="true" box="[1048,1174,920,946]" pageId="58" pageNumber="59">Abdomen:</emphasis>
Apex of tergite IV with a triangular wide area with small, irregular tubercles forming 9–10 diagonal nearly parallel lines. Pygidium with dense rugosity on basal 3rd (
<figureCitation id="B29BE4A8FD65FFBBAEEC1AB6FD86BF9A" box="[477,568,992,1019]" captionStart="FIGURE 9" captionStartId="87.[151,250,1074,1098]" captionTargetBox="[151,1436,181,1050]" captionTargetId="figure-17@87.[151,1436,181,1050]" captionTargetPageId="87" captionText="FIGURE 9. Pygidium of male. A) Euligyrus ebenus, B) Proculigyrus cicatricosus, C) Ligyrus (Anagrylius) cuniculus, D) L. (Ligyrus) fossor, E) L. (Ligyrus) paranaensis, F) Tomarus bituberculatus, G) T. colombianus, H) T. maracaiboensis, I) T. pilcopataensis, J) T. pullus, K) T. roigjunenti, L) T. rostratus." figureDoi="http://doi.org/10.5281/zenodo.7345041" httpUri="https://zenodo.org/record/7345041/files/figure.png" pageId="58" pageNumber="59">Fig. 9G</figureCitation>
), apex rounded.
<emphasis id="18D4243FFD65FFBBADC51AB7FCF3BF9A" bold="true" box="[756,845,993,1019]" pageId="58" pageNumber="59">Venter:</emphasis>
Apex of prosternal process flat, transversely oval to rounded. Metasternum with minute setae on anterior angles.
<emphasis id="18D4243FFD65FFBBAC771D53FC39B87E" bold="true" box="[838,903,1029,1055]" pageId="58" pageNumber="59">Legs:</emphasis>
Protibia tridentate, with a basal, small denticle (
<figureCitation id="B29BE4A8FD65FFBBAFAE1D7EFF45B822" box="[159,251,1064,1091]" captionStart="FIGURE 11" captionStartId="88.[151,250,931,955]" captionTargetBox="[234,1354,186,899]" captionTargetId="figure-24@88.[231,1357,181,907]" captionTargetPageId="88" captionText="FIGURE 11. Protibia in frontal view.A) Euligyrus similis, B) Proculigyrus cicatricosus, C) Ligyrus (Anagrylius) moroni, D) L. (Ligyrodes) relictus, E) L. (Ligyrodes) sallaei, F) L. (Ligyrus) allonasutus, G) L. (Ligyrus) villosus, H) Tomarus bituberculatus, I) T. colombianus, J) T. pullus." figureDoi="http://doi.org/10.5281/zenodo.7345051" httpUri="https://zenodo.org/record/7345051/files/figure.png" pageId="58" pageNumber="59">Fig. 11I</figureCitation>
). Apex of metatibia crenulate, with 9–11 spinules.
<emphasis id="18D4243FFD65FFBBAC1E1D7FFC5CB823" bold="true" box="[815,994,1064,1091]" pageId="58" pageNumber="59">Male genitalia:</emphasis>
Spiculum gastrale without basal plates. Phallobase 1.1 times longer than parameres. Parameres with 2 dorsal teeth on each side; basal tooth acute, longer than apical (
<figureCitation id="B29BE4A8FD65FFBBAE101D26FE2DB8EB" box="[289,403,1136,1162]" 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="58" pageNumber="59">Figs. 16D</figureCitation>
,
<figureCitation id="B29BE4A8FD65FFBBAEAE1D27FE6DB8EA" box="[415,467,1137,1163]" 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="58" pageNumber="59">19D</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.
</paragraph>
</subSubSection>
<subSubSection id="62BAABA6FD65FFBBAFF61DEEFD0AB902" pageId="58" pageNumber="59" type="diagnosis">
<paragraph id="2A1FF82DFD65FFBBAFF61DEEFD0AB902" blockId="58.[151,1437,596,1559]" pageId="58" pageNumber="59">
<emphasis id="18D4243FFD65FFBBAFF61DEEFEFCB8B3" bold="true" box="[199,322,1208,1234]" pageId="58" pageNumber="59">Diagnosis.</emphasis>
<taxonomicName id="EDA083AEFD65FFBBAE781DEFFDFFB8B3" authorityName="Lopez-Garcia &amp; Gasca-Alvarez" authorityYear="2014" box="[329,577,1208,1234]" class="Insecta" family="Dynastidae" genus="Tomarus" kingdom="Animalia" order="Coleoptera" pageId="58" pageNumber="59" phylum="Arthropoda" rank="species" species="colombianus">
<emphasis id="18D4243FFD65FFBBAE781DEFFDFFB8B3" box="[329,577,1208,1234]" italics="true" pageId="58" pageNumber="59">Tomarus colombianus</emphasis>
</taxonomicName>
can be differentiated by the following character combination: frontoclypeal tubercles conical; pronotal surface nearly smooth (
<figureCitation id="B29BE4A8FD65FFBBADFE1D8AFC86B896" box="[719,824,1244,1271]" 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="58" pageNumber="59">Fig. 27D</figureCitation>
); pronotal tubercle not visible in lateral view; pronotal fovea shallow and narrow (1/3 as wide as interocular distance); protibia tridentate, with an additional basal denticle (
<figureCitation id="B29BE4A8FD65FFBBAFAE1C72FF44B95E" box="[159,250,1316,1343]" captionStart="FIGURE 11" captionStartId="88.[151,250,931,955]" captionTargetBox="[234,1354,186,899]" captionTargetId="figure-24@88.[231,1357,181,907]" captionTargetPageId="88" captionText="FIGURE 11. Protibia in frontal view.A) Euligyrus similis, B) Proculigyrus cicatricosus, C) Ligyrus (Anagrylius) moroni, D) L. (Ligyrodes) relictus, E) L. (Ligyrodes) sallaei, F) L. (Ligyrus) allonasutus, G) L. (Ligyrus) villosus, H) Tomarus bituberculatus, I) T. colombianus, J) T. pullus." figureDoi="http://doi.org/10.5281/zenodo.7345051" httpUri="https://zenodo.org/record/7345051/files/figure.png" pageId="58" pageNumber="59">Fig. 11I</figureCitation>
); apex of metatibia with 9–11 spinules; parameres with 2 dorsal teeth on each side, main teeth long, widely separate from secondary teeth (
<figureCitation id="B29BE4A8FD65FFBBAEC71C1EFDD9B903" box="[502,615,1352,1378]" 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="58" pageNumber="59">Figs. 16D</figureCitation>
,
<figureCitation id="B29BE4A8FD65FFBBAD421C1FFD19B902" box="[627,679,1353,1379]" 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="58" pageNumber="59">19D</figureCitation>
).
</paragraph>
</subSubSection>
<subSubSection id="62BAABA6FD65FFBBAFF61C3AFCE6BA76" pageId="58" pageNumber="59" type="distribution">
<paragraph id="2A1FF82DFD65FFBBAFF61C3AFCB1B9E6" blockId="58.[151,1437,596,1559]" box="[199,783,1388,1415]" pageId="58" pageNumber="59">
<emphasis id="18D4243FFD65FFBBAFF61C3AFEDFB9E7" bold="true" box="[199,353,1388,1414]" pageId="58" pageNumber="59">Distribution.</emphasis>
<collectingCountry id="52B7B8BDFD65FFBBAE561C3AFE6BB9E7" box="[359,469,1388,1414]" name="Colombia" pageId="58" pageNumber="59">Colombia</collectingCountry>
(
<bibRefCitation id="4E3185DCFD65FFBBAED51C3BFCBDB9E6" author="Lopez-Garcia, M. M. &amp; Gasca-Alvarez, H. J. &amp; Amat-Garcia, G." box="[484,771,1388,1415]" pageId="58" pageNumber="59" pagination="579 - 584" refId="ref91387" refString="Lopez-Garcia, M. M., Gasca-Alvarez, H. J. &amp; Amat-Garcia, G. (2014) A new species of Tomarus Erichson, 1847 (Coleoptera: Scarabaeoidea: Dynastinae: Pentodontini), with a key to the species in Colombia. Zootaxa, 3869 (5), 579 - 584. https: // doi. org / 10.11646 / zootaxa. 3869.5.8" type="journal article" year="2014">
López-García
<emphasis id="18D4243FFD65FFBBADB61C3BFD06B9E7" box="[647,696,1388,1414]" italics="true" pageId="58" pageNumber="59">et al</emphasis>
. 2014
</bibRefCitation>
).
</paragraph>
<paragraph id="2A1FF82DFD65FFBBAFF61CC6FDC2B9AE" blockId="58.[151,1437,596,1559]" pageId="58" pageNumber="59">
<emphasis id="18D4243FFD65FFBBAFF61CC6FDB8B9CB" bold="true" box="[199,518,1424,1451]" pageId="58" pageNumber="59">
Locality records (
<figureCitation id="B29BE4A8FD65FFBBAE911CC6FE47B9CB" box="[416,505,1424,1450]" 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="58" pageNumber="59">Fig. 44</figureCitation>
).
</emphasis>
Five specimens examined from ICN and CIUQ.
<emphasis id="18D4243FFD65FFBBAB7B1CC6FA23B9CB" bold="true" box="[1098,1437,1424,1450]" pageId="58" pageNumber="59">
<collectingCountry id="52B7B8BDFD65FFBBAB7B1CC6FB59B9CB" box="[1098,1255,1424,1450]" name="Colombia" pageId="58" pageNumber="59">COLOMBIA</collectingCountry>
(5).
<collectingRegion id="E86436CFFD65FFBBAA141CC6FAD6B9CB" box="[1317,1384,1424,1450]" country="Colombia" name="Huila" pageId="58" pageNumber="59">Huila</collectingRegion>
(1):
</emphasis>
Neiva.
<emphasis id="18D4243FFD65FFBBAFD91CE3FE6CB9AF" bold="true" box="[232,466,1460,1487]" pageId="58" pageNumber="59">
<collectingRegion id="E86436CFFD65FFBBAFD91CE3FE1FB9AF" box="[232,417,1460,1487]" country="Colombia" name="Valle del Cauca" pageId="58" pageNumber="59">Valle del Cauca</collectingRegion>
(4):
</emphasis>
Buenaventura.
</paragraph>
<paragraph id="2A1FF82DFD65FFBBAFF61C8FFCE6BA76" blockId="58.[151,1437,596,1559]" pageId="58" pageNumber="59">
<emphasis id="18D4243FFD65FFBBAFF61C8FFE38B993" bold="true" box="[199,390,1496,1523]" pageId="58" pageNumber="59">Natural history.</emphasis>
The few known specimens were found in tropical humid forest near a mangrove swamp and Guandal forest. They were collected by hand and at light traps.
</paragraph>
</subSubSection>
</treatment>
</document>
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