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<document id="DFC4C3DD06346442231FA187C0167862" ID-DOI="10.11646/zootaxa.4048.4.1" ID-GBIF-Dataset="bbb51c92-e184-489d-aba8-18d1dc9a25cd" ID-ISSN="1175-5326" ID-Zenodo-Dep="234023" ID-ZooBank="71E2B8DA-825E-429F-BAA0-F582702B4A80" IM.materialsCitations_approvedBy="felipe" IM.metadata_approvedBy="felipe" IM.tables_requiresApprovalFor="existingObjects,plazi" IM.taxonomicNames_approvedBy="felipe" checkinTime="1461348015003" checkinUser="plazi" docAuthor="López-García, Margarita M., Gasca-Álvarez, Héctor J. &amp; Amat-García, Germán" docDate="2015" docId="990DA529FF84FF9BFF4646B11580FE4A" docLanguage="en" docName="zt04048p492.pdf" docOrigin="Zootaxa 4048 (4)" docStyle="DocumentStyle:647186512141C8FC8976D5BCC54AEB7D.9:Zootaxa.2013-.journal_article" docStyleId="647186512141C8FC8976D5BCC54AEB7D" docStyleName="Zootaxa.2013-.journal_article" docStyleVersion="9" docTitle="Tomarus bituberculatus" docType="treatment" docVersion="11" lastPageNumber="479" masterDocId="6534DD51FF9EFF87FFD1425F1659FFE6" masterDocTitle="The scarab beetle tribe Pentodontini (Coleoptera: Scarabaeidae: Dynastinae) of Colombia: taxonomy, natural history, and distribution" masterLastPageNumber="492" masterPageNumber="451" pageNumber="477" updateTime="1698621022009" updateUser="plazi">
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<mods:title id="D100EC110DDD891478F5E306F94472F2">The scarab beetle tribe Pentodontini (Coleoptera: Scarabaeidae: Dynastinae) of Colombia: taxonomy, natural history, and distribution</mods:title>
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<paragraph id="111B143FFF84FF9DFF4646B11553FAEE" blockId="26.[151,778,1262,1322]" box="[151,778,1262,1288]" pageId="26" pageNumber="477">
<heading id="4A53A353FF84FF9DFF4646B11553FAEE" bold="true" box="[151,778,1262,1288]" fontSize="11" level="1" pageId="26" pageNumber="477" reason="1">
<emphasis id="23D0C82DFF84FF9DFF4646B11553FAEE" bold="true" box="[151,778,1262,1288]" pageId="26" pageNumber="477">
<taxonomicName id="D6A46FBCFF84FF9DFF4646B117EEFAEE" box="[151,439,1262,1288]" class="Insecta" family="Dynastidae" genus="Tomarus" kingdom="Animalia" order="Coleoptera" pageId="26" pageNumber="477" phylum="Arthropoda" rank="species" species="bituberculatus">
<emphasis id="23D0C82DFF84FF9DFF4646B117EEFAEE" bold="true" box="[151,439,1262,1288]" italics="true" pageId="26" pageNumber="477">Tomarus bituberculatus</emphasis>
</taxonomicName>
(Palisot de
<bibRefCitation id="753569CEFF84FF9DFD9946B1155BFAEE" author="Beauvois" box="[584,770,1262,1288]" pageId="26" pageNumber="477" refString="Palisot de Beauvois, A. M. F. J. (1811) Insectes Recueillis en Afrique et em Amerique, dans les Royaumes d´Oware et de Benin, a Saint-Dominigue et dans les Etats-Unis, pendant les Anees 1786 - 1797. Levrault, Schoell, et Cie, Paris, France, Livraison 7, pp. 101 - 120." type="journal article" year="1811">Beauvois, 1811</bibRefCitation>
)
</emphasis>
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</paragraph>
<paragraph id="111B143FFF84FF9DFF46474E1765FACC" blockId="26.[151,778,1262,1322]" box="[151,316,1297,1322]" pageId="26" pageNumber="477">
(
<figureCitation id="899F08BAFF84FF9DFF4E474E176AFACC" box="[159,307,1297,1322]" captionStart="FIGURES 94 101" captionStartId="27.[151,264,1159,1182]" captionTargetBox="[162,1433,376,1132]" captionTargetId="figure@27.[151,1436,373,1138]" captionTargetPageId="27" captionText="FIGURES 94 101. Tomarus bituberculatus. 94) habitus in dorsal view, 95 96) variations in shape pronotal fovea, 97) habitus in lateral view, 98) parameres in frontal view, 99) parameres in lateral view, 100) spiculum gastrale, 101) locality records in Colombia." httpUri="https://zenodo.org/record/234036/files/figure.png" pageId="26" pageNumber="477">Figs. 94101</figureCitation>
)
</paragraph>
</subSubSection>
<subSubSection id="59BE47B4FF84FF9DFF4647061785FA0C" pageId="26" pageNumber="477" type="reference_group">
<paragraph id="111B143FFF84FF9DFF46470617A2FA4A" blockId="26.[151,747,1369,1514]" pageId="26" pageNumber="477">
<treatmentCitationGroup id="31B43311FF84FF9DFF46470617A2FA4A" pageId="26" pageNumber="477">
<taxonomicName id="D6A46FBCFF84FF9DFF46470617A9FA89" ID-CoL="6Y84T" authority="Palisot" authorityName="Palisot" box="[151,496,1369,1391]" class="Insecta" family="Scarabaeidae" genus="Scarabaeus" kingdom="Animalia" order="Coleoptera" pageId="26" pageNumber="477" phylum="Arthropoda" rank="species" species="bituberculatus">
<emphasis id="23D0C82DFF84FF9DFF46470617FDFA89" box="[151,420,1369,1391]" italics="true" pageId="26" pageNumber="477">Scarabaeus bituberculatus</emphasis>
Palisot
</taxonomicName>
de
<treatmentCitation id="9005322EFF84FF9DFDC5470614BEFA89" author="Beauvois" box="[532,743,1369,1391]" page="103" pageId="26" pageNumber="477" year="1811">
<bibRefCitation id="753569CEFF84FF9DFDC5470614ECFA89" author="Beauvois" box="[532,693,1369,1391]" pageId="26" pageNumber="477" refString="Palisot de Beauvois, A. M. F. J. (1811) Insectes Recueillis en Afrique et em Amerique, dans les Royaumes d´Oware et de Benin, a Saint-Dominigue et dans les Etats-Unis, pendant les Anees 1786 - 1797. Levrault, Schoell, et Cie, Paris, France, Livraison 7, pp. 101 - 120." type="journal article" year="1811">Beauvois, 1811</bibRefCitation>
: 103
</treatmentCitation>
.
<taxonomicName id="D6A46FBCFF84FF9DFF46472714C2FA68" authority="Burmeister, 1847: 100" authorityName="Burmeister" authorityPageNumber="100" authorityYear="1847" box="[151,667,1400,1422]" class="Insecta" family="Dynastidae" genus="Heteronychus" kingdom="Animalia" order="Coleoptera" pageId="26" pageNumber="477" phylum="Arthropoda" rank="species" species="scarabaeinus">
<emphasis id="23D0C82DFF84FF9DFF46472717E9FA68" box="[151,432,1400,1422]" italics="true" pageId="26" pageNumber="477">Heteronychus scarabaeinus</emphasis>
<treatmentCitation id="9005322EFF84FF9DFE67472714C2FA68" author="Burmeister" box="[438,667,1400,1422]" page="100" pageId="26" pageNumber="477" year="1847">
<bibRefCitation id="753569CEFF84FF9DFE6747271431FA68" author="Burmeister" box="[438,616,1400,1422]" pageId="26" pageNumber="477" refString="Burmeister, H. (1847) Handbuch der Entomologie. Coleoptera Lamellicornia, Xylophila et Pectinicornia, Band 5. Enslin, Berlin, Germany." type="book" year="1847">Burmeister, 1847</bibRefCitation>
: 100
</treatmentCitation>
</taxonomicName>
.
<taxonomicName id="D6A46FBCFF84FF9DFF4647C717AEFA4A" authority="Bates, 1888: 315" authorityName="Bates" authorityPageNumber="315" authorityYear="1888" box="[151,503,1430,1453]" class="Insecta" family="Scarabaeidae" genus="Ligyrus" higherTaxonomySource="CoL" kingdom="Animalia" order="Coleoptera" pageId="26" pageNumber="477" phylum="Arthropoda" rank="species" species="latifovea">
<emphasis id="23D0C82DFF84FF9DFF4647C7171DFA4A" box="[151,324,1430,1453]" italics="true" pageId="26" pageNumber="477">Ligyrus latifovea</emphasis>
<treatmentCitation id="9005322EFF84FF9DFE9847C817AEFA4A" author="Bates" box="[329,503,1430,1452]" page="315" pageId="26" pageNumber="477" year="1888">
<bibRefCitation id="753569CEFF84FF9DFE9847C8179DFA4A" author="Bates" box="[329,452,1430,1452]" pageId="26" pageNumber="477" refString="Bates, H. W. (1888) Pectinicornia and Lamellicornia, Family Dynastinae. In: Godman, F. D. &amp; Salvin, O. (Eds), Biologia Centrali-Americana, Insecta, Coleoptera, Vol. II, Part 2, 296 - 342." type="book chapter" year="1888">Bates, 1888</bibRefCitation>
: 315
</treatmentCitation>
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.
</treatmentCitationGroup>
</paragraph>
<paragraph id="111B143FFF84FF9DFF4647E9145CFA2D" blockId="26.[151,747,1369,1514]" box="[151,517,1461,1483]" pageId="26" pageNumber="477">
<treatmentCitationGroup id="31B43311FF84FF9DFF4647E9145CFA2D" box="[151,517,1461,1483]" pageId="26" pageNumber="477">
<taxonomicName id="D6A46FBCFF84FF9DFF4647E91458FA2D" ID-CoL="6PZ9Q" authority="Arrow, 1913: 466" authorityName="Arrow" authorityPageNumber="466" authorityYear="1913" box="[151,513,1461,1483]" class="Insecta" family="Scarabaeidae" genus="Ligyrus" higherTaxonomySource="CoL" kingdom="Animalia" order="Coleoptera" pageId="26" pageNumber="477" phylum="Arthropoda" rank="species" species="maximus">
<emphasis id="23D0C82DFF84FF9DFF4647E9171CFA2D" box="[151,325,1462,1483]" italics="true" pageId="26" pageNumber="477">Ligyrus maximus</emphasis>
<treatmentCitation id="9005322EFF84FF9DFE9A47E91458FA2D" author="Arrow" box="[331,513,1461,1483]" page="466" pageId="26" pageNumber="477" year="1913">
<bibRefCitation id="753569CEFF84FF9DFE9A47E91796FA2D" author="Arrow" box="[331,463,1461,1483]" pageId="26" pageNumber="477" refString="Arrow, G. J. (1913) Some new species of Lamellicorn Beetles from Brazil. The Annals and Magazine of natural History, including Zoology, Botany and Geology. London, series 8, volume 11, 456 - 466." type="journal article" year="1913">Arrow, 1913</bibRefCitation>
: 466
</treatmentCitation>
</taxonomicName>
.
</treatmentCitationGroup>
</paragraph>
<paragraph id="111B143FFF84FF9DFF46478A1785FA0C" blockId="26.[151,747,1369,1514]" box="[151,476,1492,1514]" pageId="26" pageNumber="477">
<treatmentCitationGroup id="31B43311FF84FF9DFF46478A1785FA0C" box="[151,476,1492,1514]" pageId="26" pageNumber="477">
<taxonomicName id="D6A46FBCFF84FF9DFF46478A1781FA0C" ID-CoL="6PYXQ" authority="Arrow, 1914: 273" authorityName="Arrow" authorityPageNumber="273" authorityYear="1914" box="[151,472,1492,1514]" class="Insecta" family="Scarabaeidae" genus="Ligyrus" higherTaxonomySource="CoL" kingdom="Animalia" order="Coleoptera" pageId="26" pageNumber="477" phylum="Arthropoda" rank="species" species="latus">
<emphasis id="23D0C82DFF84FF9DFF46478A1745FA0C" box="[151,284,1492,1514]" italics="true" pageId="26" pageNumber="477">Ligyrus latus</emphasis>
<treatmentCitation id="9005322EFF84FF9DFEF3478B1781FA0C" author="Arrow" box="[290,472,1492,1514]" page="273" pageId="26" pageNumber="477" year="1914">
<bibRefCitation id="753569CEFF84FF9DFEF3478B17FCFA0C" author="Arrow" box="[290,421,1492,1514]" pageId="26" pageNumber="477" refString="Arrow, G. J. (1914) Some further notes on lamellicorn beetles of the subfamily Dynastinae. Annals and Magazine of Natural History, series 8, volume 14, 257 - 276, 360." type="journal article" year="1914">Arrow, 1914</bibRefCitation>
: 273
</treatmentCitation>
</taxonomicName>
.
</treatmentCitationGroup>
</paragraph>
</subSubSection>
<subSubSection id="59BE47B4FF84FF9CFF464449170CFEA6" lastPageId="27" lastPageNumber="478" pageId="26" pageNumber="477" type="description">
<paragraph id="111B143FFF84FF9CFF464449170CFEA6" blockId="26.[151,1437,1558,2015]" lastBlockId="27.[151,1436,151,320]" lastPageId="27" lastPageNumber="478" pageId="26" pageNumber="477">
<emphasis id="23D0C82DFF84FF9DFF4644491770F9C9" bold="true" box="[151,297,1558,1583]" pageId="26" pageNumber="477">Description.</emphasis>
Habitus as in
<figureCitation id="899F08BAFF84FF9DFE1744491441F9C9" box="[454,536,1558,1583]" captionStart="FIGURES 94 101" captionStartId="27.[151,264,1159,1182]" captionTargetBox="[162,1433,376,1132]" captionTargetId="figure@27.[151,1436,373,1138]" captionTargetPageId="27" captionText="FIGURES 94 101. Tomarus bituberculatus. 94) habitus in dorsal view, 95 96) variations in shape pronotal fovea, 97) habitus in lateral view, 98) parameres in frontal view, 99) parameres in lateral view, 100) spiculum gastrale, 101) locality records in Colombia." httpUri="https://zenodo.org/record/234036/files/figure.png" pageId="26" pageNumber="477">Fig. 94</figureCitation>
. Length 26.0
<quantity id="D65CB9DAFF84FF9DFD6844491542F9C8" box="[697,795,1558,1583]" metricMagnitude="-2" metricUnit="m" metricValue="2.85" pageId="26" pageNumber="477" unit="mm" value="28.5">28.5 mm</quantity>
(♂), 25.030.0 mm (♀). Width 13.015.0 mm (♂),
<quantity id="D65CB9DAFF84FF9DFA8C444916A4F9B4" metricMagnitude="-2" metricUnit="m" metricValue="1.4" metricValueMax="1.45" metricValueMin="1.35" pageId="26" pageNumber="477" unit="mm" value="14.0" valueMax="14.5" valueMin="13.5">13.5 14.5 mm</quantity>
(♀). Color black to dark brown, venter lighter.
<emphasis id="23D0C82DFF84FF9DFCF544651535F9B5" bold="true" box="[804,876,1594,1619]" italics="true" pageId="26" pageNumber="477">Head:</emphasis>
Frons flat; surface strongly rugose and grooved; base nearly smooth, with sparse, deep punctures. Frontoclypeal suture only visible on the sides, interrupted by 2 frontal tubercles. Tubercles transverse and triangular, separated by about 4.55.0 times a tubercle diameter (
<figureCitation id="899F08BAFF84FF9DFABE44DD16EEF959" captionStart="FIGURES 94 101" captionStartId="27.[151,264,1159,1182]" captionTargetBox="[162,1433,376,1132]" captionTargetId="figure@27.[151,1436,373,1138]" captionTargetPageId="27" captionText="FIGURES 94 101. Tomarus bituberculatus. 94) habitus in dorsal view, 95 96) variations in shape pronotal fovea, 97) habitus in lateral view, 98) parameres in frontal view, 99) parameres in lateral view, 100) spiculum gastrale, 101) locality records in Colombia." httpUri="https://zenodo.org/record/234036/files/figure.png" pageId="26" pageNumber="477">Fig. 97</figureCitation>
). Clypeus strongly rugose and grooved; shape triangular, apex narrowed with 2 reflexed teeth. Interocular distance equals 3.53.7 times the transverse ocular diameter. Antennal subequal in length to antennomeres 27. Mandibles with 2 apical teeth and one basal rounded lobe.
<emphasis id="23D0C82DFF84FF9DFCE744B015EAF8E0" bold="true" box="[822,947,1775,1798]" italics="true" pageId="26" pageNumber="477">Pronotum:</emphasis>
Surface with shallow and small punctures, denser and deeper on anterior and posterior angles. Apical tubercle prominent, apex strongly acute and slightly recurved (
<figureCitation id="899F08BAFF84FF9DFEC345691728F8A9" box="[274,369,1846,1871]" captionStart="FIGURES 94 101" captionStartId="27.[151,264,1159,1182]" captionTargetBox="[162,1433,376,1132]" captionTargetId="figure@27.[151,1436,373,1138]" captionTargetPageId="27" captionText="FIGURES 94 101. Tomarus bituberculatus. 94) habitus in dorsal view, 95 96) variations in shape pronotal fovea, 97) habitus in lateral view, 98) parameres in frontal view, 99) parameres in lateral view, 100) spiculum gastrale, 101) locality records in Colombia." httpUri="https://zenodo.org/record/234036/files/figure.png" pageId="26" pageNumber="477">Fig. 96</figureCitation>
). Subapical fovea transversally oval, deep, rugose and finely punctate (
<figureCitation id="899F08BAFF84FF9DFB23456913C9F8A9" box="[1266,1424,1846,1871]" captionStart="FIGURES 94 101" captionStartId="27.[151,264,1159,1182]" captionTargetBox="[162,1433,376,1132]" captionTargetId="figure@27.[151,1436,373,1138]" captionTargetPageId="27" captionText="FIGURES 94 101. Tomarus bituberculatus. 94) habitus in dorsal view, 95 96) variations in shape pronotal fovea, 97) habitus in lateral view, 98) parameres in frontal view, 99) parameres in lateral view, 100) spiculum gastrale, 101) locality records in Colombia." httpUri="https://zenodo.org/record/234036/files/figure.png" pageId="26" pageNumber="477">Figs. 95, 97</figureCitation>
).
<taxonomicName id="D6A46FBCFF84FF9DFF4645051728F895" authority="Surface" authorityName="Surface" box="[151,369,1882,1907]" class="Trilobita" family="Styginidae" genus="Scutellum" higherTaxonomySource="GBIF" kingdom="Animalia" order="Corynexochida" pageId="26" pageNumber="477" phylum="Arthropoda" rank="genus">
<emphasis id="23D0C82DFF84FF9DFF464505174CF895" bold="true" box="[151,277,1882,1907]" italics="true" pageId="26" pageNumber="477">Scutellum:</emphasis>
Surface
</taxonomicName>
with small to minute, sparse punctures.
<emphasis id="23D0C82DFF84FF9DFC8D450515F4F895" bold="true" box="[860,941,1882,1907]" italics="true" pageId="26" pageNumber="477">Elytra:</emphasis>
Surface with deep, large punctures, some smaller between intervals; 3 pairs of double rows of punctures; elytral suture present.
<emphasis id="23D0C82DFF84FF9DFB4C4521134CF871" bold="true" box="[1181,1301,1918,1943]" italics="true" pageId="26" pageNumber="477">
<taxonomicName id="D6A46FBCFF84FF9DFB4C45211354F871" box="[1181,1293,1918,1943]" family="Trichomycteridae" genus="Pygidium" higherTaxonomySource="CoL" kingdom="Animalia" order="Siluriformes" pageId="26" pageNumber="477" phylum="Chordata" rank="genus">Pygidium</taxonomicName>
:
</emphasis>
Basal third strongly punctate and rugose. Disc with sparser punctures. Surface weakly convex in males, nearly flat in females.
<emphasis id="23D0C82DFF84FF9DFF2D45981762F838" bold="true" box="[252,315,1991,2014]" italics="true" pageId="26" pageNumber="477">Legs:</emphasis>
Protibia tridentate, teeth spaced by a subequal distance. Male protarsus simple, not enlarged. Apex of metatibia strongly crenulate; apical margin with 1517 spinules. Apex of first metatarsomere subtruncate.
<emphasis id="23D0C82DFF85FF9CFEF742E31723FF35" bold="true" box="[294,378,188,211]" italics="true" pageId="27" pageNumber="478">Venter:</emphasis>
Prosternal process long, extending beyond procoxae; apex flat, longitudinally oval, posterior half bordered by long yellow setae.
<emphasis id="23D0C82DFF85FF9CFDF942BF14F7FF1E" bold="true" box="[552,686,223,248]" pageId="27" pageNumber="478">
<emphasis id="23D0C82DFF85FF9CFDF942BF14FDFF11" bold="true" box="[552,676,224,247]" italics="true" pageId="27" pageNumber="478">Parameres</emphasis>
:
</emphasis>
Base broad; lateral teeth small and acute; apex broad, with 2 small lobed projections on each apex (
<figureCitation id="899F08BAFF85FF9CFDD4435B14D6FEFA" box="[517,655,260,285]" captionStart="FIGURES 94 101" captionStartId="27.[151,264,1159,1182]" captionTargetBox="[162,1433,376,1132]" captionTargetId="figure@27.[151,1436,373,1138]" captionTargetPageId="27" captionText="FIGURES 94 101. Tomarus bituberculatus. 94) habitus in dorsal view, 95 96) variations in shape pronotal fovea, 97) habitus in lateral view, 98) parameres in frontal view, 99) parameres in lateral view, 100) spiculum gastrale, 101) locality records in Colombia." httpUri="https://zenodo.org/record/234036/files/figure.png" pageId="27" pageNumber="478">Figs. 9899</figureCitation>
).
<emphasis id="23D0C82DFF85FF9CFD73435C1524FEFA" bold="true" box="[674,893,259,284]" italics="true" pageId="27" pageNumber="478">Spiculum gastrale:</emphasis>
Base larger than lateral branches, apex strongly broad (
<figureCitation id="899F08BAFF85FF9CFF3743781711FEA6" box="[230,328,295,320]" captionStart="FIGURES 94 101" captionStartId="27.[151,264,1159,1182]" captionTargetBox="[162,1433,376,1132]" captionTargetId="figure@27.[151,1436,373,1138]" captionTargetPageId="27" captionText="FIGURES 94 101. Tomarus bituberculatus. 94) habitus in dorsal view, 95 96) variations in shape pronotal fovea, 97) habitus in lateral view, 98) parameres in frontal view, 99) parameres in lateral view, 100) spiculum gastrale, 101) locality records in Colombia." httpUri="https://zenodo.org/record/234036/files/figure.png" pageId="27" pageNumber="478">Fig. 100</figureCitation>
).
</paragraph>
</subSubSection>
<caption id="45DB44B7FF85FF9CFF4646D8175BFB3D" httpUri="https://zenodo.org/record/234036/files/figure.png" pageId="27" pageNumber="478" targetBox="[162,1433,376,1132]" targetPageId="27">
<paragraph id="111B143FFF85FF9CFF4646D8175BFB3D" blockId="27.[151,1436,1159,1243]" pageId="27" pageNumber="478">
<emphasis id="23D0C82DFF85FF9CFF4646D81738FB7A" bold="true" box="[151,353,1159,1182]" pageId="27" pageNumber="478">FIGURES 94101.</emphasis>
<taxonomicName id="D6A46FBCFF85FF9CFEB646D6140FFB7B" box="[359,598,1159,1182]" class="Insecta" family="Dynastidae" genus="Tomarus" kingdom="Animalia" order="Coleoptera" pageId="27" pageNumber="478" phylum="Arthropoda" rank="species" species="bituberculatus">
<emphasis id="23D0C82DFF85FF9CFEB646D6140FFB7B" box="[359,598,1159,1182]" italics="true" pageId="27" pageNumber="478">Tomarus bituberculatus</emphasis>
</taxonomicName>
. 94) habitus in dorsal view, 95
<emphasis id="23D0C82DFF85FF9CFC4146D715C4FB7B" bold="true" box="[912,925,1160,1181]" pageId="27" pageNumber="478"></emphasis>
96) variations in shape pronotal fovea, 97) habitus in lateral view, 98) parameres in frontal view, 99) parameres in lateral view, 100) spiculum gastrale, 101) locality records in Colombia.
</paragraph>
</caption>
<subSubSection id="59BE47B4FF85FF9BFF16475D124FFF32" lastPageId="28" lastPageNumber="479" pageId="27" pageNumber="478" type="diagnosis">
<paragraph id="111B143FFF85FF9CFF16475D1240FAD9" blockId="27.[151,1437,1282,1991]" pageId="27" pageNumber="478">
<emphasis id="23D0C82DFF85FF9CFF16475D1718FAFD" bold="true" box="[199,321,1282,1307]" pageId="27" pageNumber="478">Diagnosis.</emphasis>
This species is similar to
<taxonomicName id="D6A46FBCFF85FF9CFDB5475C14E9FAFC" box="[612,688,1283,1306]" class="Insecta" family="Dynastidae" genus="Tomarus" kingdom="Animalia" order="Coleoptera" pageId="27" pageNumber="478" phylum="Arthropoda" rank="species" species="gyas">
<emphasis id="23D0C82DFF85FF9CFDB5475C14E9FAFC" box="[612,688,1283,1306]" italics="true" pageId="27" pageNumber="478">T. gyas</emphasis>
</taxonomicName>
by the size of the pronotal tubercle and fovea. However, the fovea of
<taxonomicName id="D6A46FBCFF85FF9CFF644778172BFAD9" box="[181,370,1318,1343]" class="Insecta" family="Dynastidae" genus="Tomarus" kingdom="Animalia" order="Coleoptera" pageId="27" pageNumber="478" phylum="Arthropoda" rank="species" species="bituberculatus">
<emphasis id="23D0C82DFF85FF9CFF644778172BFAD9" box="[181,370,1318,1343]" italics="true" pageId="27" pageNumber="478">T. bituberculatus</emphasis>
</taxonomicName>
is oval and wider, and the protibia has no fourth basal tooth.
</paragraph>
<paragraph id="111B143FFF85FF9CFF16471515F0F821" blockId="27.[151,1437,1282,1991]" pageId="27" pageNumber="478">
<emphasis id="23D0C82DFF85FF9CFF16471517CEFA85" bold="true" box="[199,407,1354,1379]" pageId="27" pageNumber="478">Locality records.</emphasis>
(
<figureCitation id="899F08BAFF85FF9CFE7D4715144BFA85" box="[428,530,1354,1379]" captionStart="FIGURES 94 101" captionStartId="27.[151,264,1159,1182]" captionTargetBox="[162,1433,376,1132]" captionTargetId="figure@27.[151,1436,373,1138]" captionTargetPageId="27" captionText="FIGURES 94 101. Tomarus bituberculatus. 94) habitus in dorsal view, 95 96) variations in shape pronotal fovea, 97) habitus in lateral view, 98) parameres in frontal view, 99) parameres in lateral view, 100) spiculum gastrale, 101) locality records in Colombia." httpUri="https://zenodo.org/record/234036/files/figure.png" pageId="27" pageNumber="478">Fig. 101</figureCitation>
)
<specimenCount id="07A2DFB6FF85FF9CFDF64715148FFA85" box="[551,726,1354,1379]" pageId="27" pageNumber="478" type="generic">216 specimens</specimenCount>
,
<specimenCount id="07A2DFB6FF85FF9CFD3647141570FA85" box="[743,809,1355,1379]" pageId="27" pageNumber="478" type="generic">104♂</specimenCount>
, 112♀. Specimens were seen from ANDES, BCRC, CEUA, CEUN, CIUQ, CTI, HJG, IAvH, ICN, LEUC, MEFLG, MLS, MPUJ, UNAB, UNSM, UPN.
<emphasis id="23D0C82DFF85FF9CFACE47311696FA4D" bold="true" pageId="27" pageNumber="478">Amazonas (17):</emphasis>
Leticia (15). Leticia,
<collectingCountry id="69B354AFFF85FF9CFE1447CD141DFA4D" box="[453,580,1426,1451]" name="Macedonia" pageId="27" pageNumber="478">Macedonia</collectingCountry>
, Resguardo indígena Ticuna (1). Tarapacá, Río Tacana (1).
<emphasis id="23D0C82DFF85FF9CFB3547CD13C5FA4D" bold="true" box="[1252,1436,1426,1451]" pageId="27" pageNumber="478">Antioquia (12):</emphasis>
Apartadó (2). El Carmen de Viboral (1). Medellín (4). Puerto Berrío (4).
<collectingCountry id="69B354AFFF85FF9CFC1D47E8123DFA29" box="[972,1124,1462,1487]" name="Saint Helena, Ascension and Tristan da Cunha" pageId="27" pageNumber="478">Santa Helena</collectingCountry>
, Vda. El Mazo (1).
<emphasis id="23D0C82DFF85FF9CFA9247E91699FA15" bold="true" pageId="27" pageNumber="478">Arauca (3):</emphasis>
Alred. Río Arauca (3).
<emphasis id="23D0C82DFF85FF9CFE0347851402FA15" bold="true" box="[466,603,1498,1523]" pageId="27" pageNumber="478">Boyacá (2):</emphasis>
Villa de Leyva, SFF Iguaque (2).
<emphasis id="23D0C82DFF85FF9CFC3547851230FA15" bold="true" box="[996,1129,1498,1523]" pageId="27" pageNumber="478">Caldas (6):</emphasis>
La Dorada (2). Manizales,
<collectingCountry id="69B354AFFF85FF9CFF4647A116BAF9F1" box="[151,227,1534,1559]" name="Cyprus" pageId="27" pageNumber="478">Chipre</collectingCountry>
(1). Manizales, Vda. Hoyo Frío, Reserva Planalto CENICAFÉ (1). Manizales, Vda. Maltería, Granja Tesorito (1).
<collectingCountry id="69B354AFFF85FF9CFEF9447D17C9F9DD" box="[296,400,1570,1595]" name="Palestine" pageId="27" pageNumber="478">Palestina</collectingCountry>
, Vda. Santágueda, Granja Montelindo, Valle de Santagueda (1).
<emphasis id="23D0C82DFF85FF9CFBB7447D12A5F9DD" bold="true" box="[1126,1276,1570,1595]" pageId="27" pageNumber="478">Caquetá (2):</emphasis>
Florencia (2).
<emphasis id="23D0C82DFF85FF9CFF464419171EF9B9" bold="true" box="[151,327,1606,1631]" pageId="27" pageNumber="478">Casanare (20):</emphasis>
Tauramena, Vda. Villa Rosa, Fca. Barcelona (2). Yopal, Agua Azul, Hacienda Los Rosales (2). W. Agua Azul 1 KM (2). Yopal (14).
<emphasis id="23D0C82DFF85FF9CFDF14435154DF965" bold="true" box="[544,788,1642,1667]" pageId="27" pageNumber="478">Cundinamarca (12):</emphasis>
Bogotá (3). Fusagasugá, Cuenca Río
<collectingCountry id="69B354AFFF85FF9CFB1744351348F965" box="[1222,1297,1642,1667]" name="Jordan" pageId="27" pageNumber="478">Jordán</collectingCountry>
(1). Guasca (4). La Calera, Vda. La Jagada (1). La Mesa, Vda. Acacías (2). Tocaima (1).
<emphasis id="23D0C82DFF85FF9CFC2144D112C9F941" bold="true" box="[1008,1168,1678,1703]" pageId="27" pageNumber="478">Guaviare (2):</emphasis>
Caño Mocuare (1). San José del Guaviare, Fca. La Floresta (1).
<emphasis id="23D0C82DFF85FF9CFD8644ED1492F92D" bold="true" box="[599,715,1714,1739]" pageId="27" pageNumber="478">Huila (2):</emphasis>
San Agustín (2).
<emphasis id="23D0C82DFF85FF9CFC4344ED1210F92D" bold="true" box="[914,1097,1714,1739]" pageId="27" pageNumber="478">Magdalena (1):</emphasis>
Santa Ana (1).
<emphasis id="23D0C82DFF85FF9CFB2B44ED132EF92D" bold="true" box="[1274,1399,1714,1739]" pageId="27" pageNumber="478">Meta (15):</emphasis>
La Macarena (1). Puerto López (4). Puerto López, Vda. Menegua (1). Puerto López, Colegio INSTIVAL (2). Puerto Gaitán (1). Puerto Gaitán, Altamira, Los Llaneros (2). Villavicencio (1); Correg. Remolinos, Cafam Llanos (3).
<emphasis id="23D0C82DFF85FF9CFF464541171CF8D1" bold="true" box="[151,325,1822,1847]" pageId="27" pageNumber="478">Putumayo (1):</emphasis>
Puerto Asís (1).
<emphasis id="23D0C82DFF85FF9CFDDD454114EDF8D1" bold="true" box="[524,692,1822,1847]" pageId="27" pageNumber="478">Risaralda (1):</emphasis>
Pereira (1).
<emphasis id="23D0C82DFF85FF9CFC97454115AFF8D1" bold="true" box="[838,1014,1822,1847]" pageId="27" pageNumber="478">Santander (5):</emphasis>
Carare-Opón (1). Cimitarra, Correg. Puerto Olaya, Central Termocentro Isagen (3). Vélez, Campo Capote (1).
<emphasis id="23D0C82DFF85FF9CFC0E451D122BF8BD" bold="true" box="[991,1138,1858,1883]" pageId="27" pageNumber="478">Tolima (46):</emphasis>
Honda (45). Valencia (1).
<emphasis id="23D0C82DFF85FF9CFF46453917CDF899" bold="true" box="[151,404,1894,1919]" pageId="27" pageNumber="478">Valle del Cauca (26):</emphasis>
Buga (1). Bugalagrande (1). Cali (1). Calipuerto (5). “Mares”, Cordillera Occidental (12). Palmira (5). Roldanillo (1).
<emphasis id="23D0C82DFF85FF9CFE0445D51430F845" bold="true" box="[469,617,1930,1955]" pageId="27" pageNumber="478">Vichada (6):</emphasis>
Puerto Carreño, Fca. El Toro (1). Puerto Carreño, Barrio El Centro (4). Santa Rosalia (1).
<emphasis id="23D0C82DFF85FF9CFEB445F11430F821" bold="true" box="[357,617,1966,1991]" pageId="27" pageNumber="478">Ambiguous data (36):</emphasis>
<collectingCountry id="69B354AFFF85FF9CFDA145F11487F821" box="[624,734,1966,1991]" name="Colombia" pageId="27" pageNumber="478">Colombia</collectingCountry>
(4). No data (32).
</paragraph>
<paragraph id="111B143FFF82FF9BFF1642C8124FFF32" blockId="28.[151,1436,151,429]" pageId="28" pageNumber="479">
<emphasis id="23D0C82DFF82FF9BFF1642C8178FFF56" bold="true" box="[199,470,151,176]" pageId="28" pageNumber="479">Temporal distribution.</emphasis>
January (6), February (6), March (21), April (18), May (8), June (6), July (13), August (5), September (10), October (14), November (18), December (4). No data (87).
</paragraph>
</subSubSection>
<subSubSection id="59BE47B4FF82FF9BFF1642801580FE4A" pageId="28" pageNumber="479" type="distribution">
<paragraph id="111B143FFF82FF9BFF16428017DAFE82" blockId="28.[151,1436,151,429]" pageId="28" pageNumber="479">
<emphasis id="23D0C82DFF82FF9BFF1642801739FF1E" bold="true" box="[199,352,223,248]" pageId="28" pageNumber="479">Distribution.</emphasis>
<collectingCountry id="69B354AFFF82FF9BFEA142801791FF1E" box="[368,456,223,248]" name="Mexico" pageId="28" pageNumber="479">Mexico</collectingCountry>
,
<collectingCountry id="69B354AFFF82FF9BFE0C42801400FF1E" box="[477,601,223,248]" name="Guatemala" pageId="28" pageNumber="479">Guatemala</collectingCountry>
,
<collectingCountry id="69B354AFFF82FF9BFDBC428014A2FF1E" box="[621,763,223,248]" name="El Salvador" pageId="28" pageNumber="479">El Salvador</collectingCountry>
,
<collectingCountry id="69B354AFFF82FF9BFCDE42801526FF1E" box="[783,895,223,248]" name="Honduras" pageId="28" pageNumber="479">Honduras</collectingCountry>
,
<collectingCountry id="69B354AFFF82FF9BFC4242801250FF1E" box="[915,1033,223,248]" name="Nicaragua" pageId="28" pageNumber="479">Nicaragua</collectingCountry>
,
<collectingCountry id="69B354AFFF82FF9BFBCC42BF12FAFF1E" box="[1053,1187,223,248]" name="Costa Rica" pageId="28" pageNumber="479">Costa Rica</collectingCountry>
,
<collectingCountry id="69B354AFFF82FF9BFB6642BF134AFF11" box="[1207,1299,224,247]" name="Panama" pageId="28" pageNumber="479">Panama</collectingCountry>
,
<collectingCountry id="69B354AFFF82FF9BFAF6428013C1FF1E" box="[1319,1432,223,248]" name="Colombia" pageId="28" pageNumber="479">Colombia</collectingCountry>
,
<collectingCountry id="69B354AFFF82FF9BFF46435B1754FEFB" box="[151,269,260,285]" name="Venezuela" pageId="28" pageNumber="479">Venezuela</collectingCountry>
,
<collectingCountry id="69B354AFFF82FF9BFEC8435B1722FEFB" box="[281,379,260,285]" name="Trinidad and Tobago" pageId="28" pageNumber="479">Trinidad</collectingCountry>
,
<collectingCountry id="69B354AFFF82FF9BFE59435B17BEFEFB" box="[392,487,260,285]" name="Ecuador" pageId="28" pageNumber="479">Ecuador</collectingCountry>
,
<collectingCountry id="69B354AFFF82FF9BFE25435A1473FEFA" box="[500,554,261,284]" name="Peru" pageId="28" pageNumber="479">Peru</collectingCountry>
, and
<collectingCountry id="69B354AFFF82FF9BFDB6435B14F2FEFB" box="[615,683,260,285]" name="Brazil" pageId="28" pageNumber="479">Brazil</collectingCountry>
(
<bibRefCitation id="753569CEFF82FF9BFD6D435B1502FEFB" author="Endrodi" box="[700,859,260,285]" pageId="28" pageNumber="479" refString="Endrodi, S. (1969) Monographie der Dynastinae. 4. Tribus: Pentodontini (Coleoptera, Lamellicornia). Entomologische Abhandlungen Museum fur Tierkunde, Dresden, 37, 1 - 145." type="journal article" year="1969">Endrödi 1969</bibRefCitation>
;
<bibRefCitation id="753569CEFF82FF9BFCB9435B1236FEFB" author="Ratcliffe" box="[872,1135,260,285]" pageId="28" pageNumber="479" refString="Ratcliffe, B. C. &amp; Cave, R. D. (2006) The Dynastinae Scarab Beetles of Honduras, Nicaragua and El Salvador. Bulletin of the University of Nebraska State Museum, 21, 1 - 424." type="journal article" year="2006">Ratcliffe &amp; Cave 2006</bibRefCitation>
). In
<collectingCountry id="69B354AFFF82FF9BFB72435B134DFEFB" box="[1187,1300,260,285]" name="Colombia" pageId="28" pageNumber="479">Colombia</collectingCountry>
, the species has been found also in Chocó (
<bibRefCitation id="753569CEFF82FF9BFDD0437814FAFEA6" author="Endrodi" box="[513,675,295,320]" pageId="28" pageNumber="479" refString="Endrodi, S. (1969) Monographie der Dynastinae. 4. Tribus: Pentodontini (Coleoptera, Lamellicornia). Entomologische Abhandlungen Museum fur Tierkunde, Dresden, 37, 1 - 145." type="journal article" year="1969">Endrödi 1969</bibRefCitation>
), Cauca (
<bibRefCitation id="753569CEFF82FF9BFCC04378121CFEA6" author="Pardo-Locarno" box="[785,1093,295,320]" pageId="28" pageNumber="479" refString="Pardo-Locarno, L. C., Montoya, J. &amp; Schoonhoven, A. (2005) Abundancia de chisas rizofagas (Coleoptera: Melolonthidae) en agroecosistemas de Caldono y Buenos Aires, Cauca, Colombia. Revista Colombiana de Entomologia, 29, 177 - 184." type="journal article" year="2005">
Pardo-Locarno
<emphasis id="23D0C82DFF82FF9BFC15437615A0FEA6" box="[964,1017,295,320]" italics="true" pageId="28" pageNumber="479">et al</emphasis>
. 2005
</bibRefCitation>
), Cesar, and Córdoba (Pardo- Locarno
<emphasis id="23D0C82DFF82FF9BFF2A43121772FE83" box="[251,299,332,357]" italics="true" pageId="28" pageNumber="479">et al</emphasis>
. 2012).
</paragraph>
<paragraph id="111B143FFF82FF9BFF1643301580FE4A" blockId="28.[151,1436,151,429]" pageId="28" pageNumber="479">
<emphasis id="23D0C82DFF82FF9BFF16433017D5FE6E" bold="true" box="[199,396,367,392]" pageId="28" pageNumber="479">Natural history.</emphasis>
Specimens were collected with light traps, including black light, in different habitats including premontane and gallery forests. They were taken at
<quantity id="D65CB9DAFF82FF9BFC9D43CB158CFE4A" box="[844,981,404,429]" metricMagnitude="3" metricUnit="m" metricValue="1.415" metricValueMax="2.8" metricValueMin="0.03" pageId="28" pageNumber="479" unit="m" value="1415.0" valueMax="2800.0" valueMin="30.0">
30
<emphasis id="23D0C82DFF82FF9BFCB943CB152FFE4B" bold="true" box="[872,886,404,429]" pageId="28" pageNumber="479"></emphasis>
2,800 m
</quantity>
.
</paragraph>
</subSubSection>
</treatment>
</document>