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<mods:title id="42A8EFF9BBAE0F30E62901EDB7DC1AE4">Taxonomic revision of the South American dung beetle genus Gromphas Brullé, 1837 (Coleoptera: Scarabaeidae: Scarabaeinae: Phanaeini: Gromphadina)</mods:title>
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<mods:namePart id="9DD9CA196E3BB1ACB33EB1B16F7A1B2F">Vaz-De-Mello, Fernando Z.</mods:namePart>
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<subSubSection id="B0C4655B5975624BFF0A24FD35129A55" pageId="15" pageNumber="454" type="nomenclature">
<paragraph id="F86136D05975624BFF0A24FD35D29A3F" blockId="15.[151,648,1015,1075]" box="[151,648,1015,1042]" pageId="15" pageNumber="454">
<heading id="A32981BC5975624BFF0A24FD35D29A3F" bold="true" box="[151,648,1015,1042]" fontSize="11" level="1" pageId="15" pageNumber="454" reason="1">
<emphasis id="CAAAEAC25975624BFF0A24FD35D29A3F" bold="true" box="[151,648,1015,1042]" pageId="15" pageNumber="454">
2. 
<taxonomicName id="3FDE4D535975624BFF2924F235D29A3F" ID-CoL="6L4GV" authority="Waterhouse, 1891" authorityName="Waterhouse" authorityYear="1891" box="[180,648,1015,1042]" class="Insecta" family="Scarabaeidae" genus="Gromphas" kingdom="Animalia" order="Coleoptera" pageId="15" pageNumber="454" phylum="Arthropoda" rank="species" species="lemoinei">
<emphasis id="CAAAEAC25975624BFF2924F236F89A3C" bold="true" box="[180,418,1015,1041]" italics="true" pageId="15" pageNumber="454">Gromphas lemoinei</emphasis>
Waterhouse, 1891
</taxonomicName>
</emphasis>
</heading>
</paragraph>
<paragraph id="F86136D05975624BFF0A231F36849A1E" blockId="15.[151,648,1015,1075]" box="[151,478,1050,1075]" pageId="15" pageNumber="454">
<figureCitation id="60E52A555975624BFF0A231F36549A1E" box="[151,270,1050,1075]" captionStart-0="FIGURES 7 – 10" captionStart-1="FIGURES 11 – 12. 11" captionStartId-0="16.[151,264,1939,1961]" captionStartId-1="17.[151,264,1901,1923]" captionTargetBox-0="[158,1436,199,1910]" captionTargetBox-1="[161,1428,193,1879]" captionTargetId-0="figure@16.[151,1436,193,1918]" captionTargetId-1="figure@17.[151,1436,193,1880]" captionTargetPageId-0="16" captionTargetPageId-1="17" captionText-0="FIGURES 7 – 10. Gromphas lemoinei Waterhouse, 1891. 7) Major specimen. 8 – 10) Series of intermediate and minor specimens. Scale = 5 mm." captionText-1="FIGURES 11 – 12. 11: Holotype of Gromphas lemoinei. 11 a) Dorsal view. 11 b) Lateral view. 11 c) Labels. 12: Lectotype of G. amazonica from Tefé (former Ega). 12 a) Dorsal view. 12 b) Labels." httpUri-0="https://zenodo.org/record/247629/files/figure.png" httpUri-1="https://zenodo.org/record/247630/files/figure.png" pageId="15" pageNumber="454">Figs. 7–11</figureCitation>
, 17, 25, 30, 58, 66
</paragraph>
<paragraph id="F86136D05975624BFF0A236735129A55" blockId="15.[151,1436,1122,1206]" box="[151,584,1122,1144]" pageId="15" pageNumber="454">
<taxonomicName id="3FDE4D535975624BFF0A2367351E9A55" authority="Waterhouse, 1891: 60" authorityName="Waterhouse" authorityPageNumber="60" authorityYear="1891" box="[151,580,1122,1144]" class="Insecta" family="Scarabaeidae" genus="Gromphas" kingdom="Animalia" order="Coleoptera" pageId="15" pageNumber="454" phylum="Arthropoda" rank="species" species="lemoinei">
<emphasis id="CAAAEAC25975624BFF0A236736049A55" box="[151,350,1122,1144]" italics="true" pageId="15" pageNumber="454">Gromphas lemoinei</emphasis>
Waterhouse, 1891: 60
</taxonomicName>
.
</paragraph>
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<subSubSection id="B0C4655B5975624BFF0A238436869A9B" pageId="15" pageNumber="454" type="description">
<paragraph id="F86136D05975624BFF0A238436869A9B" blockId="15.[151,1436,1122,1206]" pageId="15" pageNumber="454">
<taxonomicName id="3FDE4D535975624BFF0A238436239ABA" box="[151,377,1153,1175]" class="Insecta" family="Scarabaeidae" genus="Gromphas" kingdom="Animalia" order="Coleoptera" pageId="15" pageNumber="454" phylum="Arthropoda" rank="species" species="aeruginosa">
<emphasis id="CAAAEAC25975624BFF0A238436239ABA" box="[151,377,1153,1175]" italics="true" pageId="15" pageNumber="454">Gromphas aeruginosa</emphasis>
</taxonomicName>
<taxonomicName id="3FDE4D535975624BFE1D238735DF9ABA" authority="Waterhouse" authorityName="Waterhouse" box="[384,645,1153,1175]" pageId="15" pageNumber="454" rank="variety" variety="lemoinei">
var. 
<emphasis id="CAAAEAC25975624BFE31238435599ABA" box="[428,515,1153,1175]" italics="true" pageId="15" pageNumber="454">lemoinei</emphasis>
Waterhouse
</taxonomicName>
: Gillet 1911: 80; d’Olsoufieff 1924: 138; Blackwelder 1944: 208; Barattini &amp; Saenz 1960: 23, 1964: 177.
</paragraph>
</subSubSection>
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<paragraph id="F86136D05975624BFF0A23E734259B6E" blockId="15.[151,1436,1250,2031]" pageId="15" pageNumber="454">
<emphasis id="CAAAEAC25975624BFF0A23E736049AD6" bold="true" box="[151,350,1250,1275]" pageId="15" pageNumber="454">
<typeStatus id="276588725975624BFF0A23E737889AD6" box="[151,210,1250,1275]" pageId="15" pageNumber="454">Type</typeStatus>
specimen:
</emphasis>
<typeStatus id="276588725975624BFEEF23E736809AD6" box="[370,474,1250,1275]" pageId="15" pageNumber="454" type="holotype">Holotype</typeStatus>
: undetermined sex (“ 
<typeStatus id="276588725975624BFD6F23E6346B9AD6" box="[754,817,1251,1275]" pageId="15" pageNumber="454">Type</typeStatus>
”, “Caracas”, “ 
<taxonomicName id="3FDE4D535975624BFC7323E7321F9AD6" authority="Waterh." authorityName="Waterh." box="[1006,1349,1250,1275]" class="Insecta" family="Scarabaeidae" genus="Gromphas" kingdom="Animalia" order="Coleoptera" pageId="15" pageNumber="454" phylum="Arthropoda" rank="species" species="lemoinei">
<emphasis id="CAAAEAC25975624BFC7323E733829AD6" box="[1006,1240,1250,1275]" italics="true" pageId="15" pageNumber="454">Gromphas lemoinei</emphasis>
Waterh.
</taxonomicName>
<typeStatus id="276588725975624BFAC523E632D69AD6" box="[1368,1420,1251,1275]" pageId="15" pageNumber="454">type</typeStatus>
”, “ 
<emphasis id="CAAAEAC25975624BFF3E220336C49B32" box="[163,414,1286,1311]" italics="true" pageId="15" pageNumber="454">
Onthophoida 
<taxonomicName id="3FDE4D535975624BFEA3220336C49B32" authority="Reiche" authorityName="Reiche" box="[318,414,1286,1311]" class="Insecta" family="Scarabaeidae" genus="Gromphas" kingdom="Animalia" order="Coleoptera" pageId="15" pageNumber="454" phylum="Arthropoda" rank="species" species="lemoinei">lemoinei</taxonomicName>
</emphasis>
Reiche” “ 
<taxonomicName id="3FDE4D535975624BFD892203343F9B32" authority="Perty" authorityName="Perty" box="[532,869,1286,1311]" class="Insecta" family="Scarabaeidae" genus="Gromphas" kingdom="Animalia" order="Coleoptera" pageId="15" pageNumber="454" phylum="Arthropoda" rank="species" species="aeruginosa">
<emphasis id="CAAAEAC25975624BFD89220334559B33" box="[532,783,1286,1311]" italics="true" pageId="15" pageNumber="454">Gromphas aeruginosa</emphasis>
(Perty)
</taxonomicName>
614: 99”, “ 
<taxonomicName id="3FDE4D535975624BFC74220332319B32" authority="Perty, 1830" authorityName="Perty" authorityYear="1830" box="[1001,1387,1286,1311]" class="Insecta" family="Scarabaeidae" genus="Gromphas" kingdom="Animalia" order="Coleoptera" pageId="15" pageNumber="454" phylum="Arthropoda" rank="species" species="aeruginosa">
<emphasis id="CAAAEAC25975624BFC74220333BE9B33" box="[1001,1252,1286,1311]" italics="true" pageId="15" pageNumber="454">Gromphas aeruginosa</emphasis>
Perty, 1830
</taxonomicName>
det. CPDT Gillet, 2007”), BMNH (examined by photos [
<figureCitation id="60E52A555975624BFD7E222F34649B6E" box="[739,830,1322,1347]" captionStart="FIGURES 11 – 12. 11" captionStartId="17.[151,264,1901,1923]" captionTargetBox="[161,1428,193,1879]" captionTargetId="figure@17.[151,1436,193,1880]" captionTargetPageId="17" captionText="FIGURES 11 – 12. 11: Holotype of Gromphas lemoinei. 11 a) Dorsal view. 11 b) Lateral view. 11 c) Labels. 12: Lectotype of G. amazonica from Tefé (former Ega). 12 a) Dorsal view. 12 b) Labels." httpUri="https://zenodo.org/record/247630/files/figure.png" pageId="15" pageNumber="454">Figs. 11</figureCitation>
a–c]).
</paragraph>
<paragraph id="F86136D05975624BFF5A224B34139B4A" blockId="15.[151,1436,1250,2031]" box="[199,841,1358,1383]" pageId="15" pageNumber="454">
<emphasis id="CAAAEAC25975624BFF5A224B36339B4A" bold="true" box="[199,361,1358,1383]" pageId="15" pageNumber="454">
<typeStatus id="276588725975624BFF5A224B36589B4A" box="[199,258,1358,1383]" pageId="15" pageNumber="454">Type</typeStatus>
locality:
</emphasis>
“Caracas, La Guayra” (Waterhouse 1891).
</paragraph>
<paragraph id="F86136D05975624BFF5A227734BF9BFE" blockId="15.[151,1436,1250,2031]" pageId="15" pageNumber="454">
<emphasis id="CAAAEAC25975624BFF5A2277362C9BA6" bold="true" box="[199,374,1394,1419]" pageId="15" pageNumber="454">Redescription:</emphasis>
Color: Anterior region of head black; remainder of dorsum with bright and metallic reflections of green, copper, and burgundy, with some of these colors predominating or, more commonly, a mixture of all. Pygidium colored as elytra. Metasternum bright and with uniform coloring.
</paragraph>
<paragraph id="F86136D05975624BFF5A22DB35659886" blockId="15.[151,1436,1250,2031]" pageId="15" pageNumber="454">
Head: Clypeus with two lobes at apex (in worn specimens lobes absent and apex rounded or truncate); margin between these lobes and genae rounded (
<figureCitation id="60E52A555975624BFDE8210735969836" box="[629,716,1538,1563]" captionStart="FIGURES 24 – 28" captionStartId="29.[151,264,1615,1637]" captionTargetBox="[151,1436,200,1593]" captionTargetId="figure@29.[151,1436,193,1593]" captionTargetPageId="29" captionText="FIGURES 24 – 28. Dorsal view of head. 24) Gromphas aeruginosa (only one pair of clypeal lobes and tegument smooth adjacent to eyes). 25) G. lemoinei (same). 26) G. dichroa (clypeus truncate and strong granulation around eyes). 27) G. amazonica (margin of clypeus with two pairs of lobes; strong granulation around eyes). 28) G. inermis (same)." httpUri="https://zenodo.org/record/247633/files/figure.png" pageId="15" pageNumber="454">Fig. 25</figureCitation>
). Margin of genae extended beyond clypeal margin (
<figureCitation id="60E52A555975624BFAA5210732D59836" box="[1336,1423,1538,1563]" captionStart="FIGURES 24 – 28" captionStartId="29.[151,264,1615,1637]" captionTargetBox="[151,1436,200,1593]" captionTargetId="figure@29.[151,1436,193,1593]" captionTargetPageId="29" captionText="FIGURES 24 – 28. Dorsal view of head. 24) Gromphas aeruginosa (only one pair of clypeal lobes and tegument smooth adjacent to eyes). 25) G. lemoinei (same). 26) G. dichroa (clypeus truncate and strong granulation around eyes). 27) G. amazonica (margin of clypeus with two pairs of lobes; strong granulation around eyes). 28) G. inermis (same)." httpUri="https://zenodo.org/record/247633/files/figure.png" pageId="15" pageNumber="454">Fig. 25</figureCitation>
). Clypeus with smooth tegument adjacent to eyes; granulation gradually denser and squamous toward the outer margin (
<figureCitation id="60E52A555975624BFF67214F360E984E" box="[250,340,1610,1635]" captionStart="FIGURES 24 – 28" captionStartId="29.[151,264,1615,1637]" captionTargetBox="[151,1436,200,1593]" captionTargetId="figure@29.[151,1436,193,1593]" captionTargetPageId="29" captionText="FIGURES 24 – 28. Dorsal view of head. 24) Gromphas aeruginosa (only one pair of clypeal lobes and tegument smooth adjacent to eyes). 25) G. lemoinei (same). 26) G. dichroa (clypeus truncate and strong granulation around eyes). 27) G. amazonica (margin of clypeus with two pairs of lobes; strong granulation around eyes). 28) G. inermis (same)." httpUri="https://zenodo.org/record/247633/files/figure.png" pageId="15" pageNumber="454">Fig. 25</figureCitation>
). Cephalic projection ranging from a low and transverse elevation arched upward to anteroposteriorly flat horn with acuminate apex; horn granulate on both faces (
<figureCitation id="60E52A555975624BFC41216B336698AA" box="[988,1084,1646,1671]" captionStart="FIGURES 29 – 33" captionStartId="30.[151,264,1916,1938]" captionTargetBox="[195,1380,179,1890]" captionTargetId="figure@30.[195,1380,179,1891]" captionTargetPageId="30" captionText="FIGURES 29 – 33. Frontal view of cephalic projections. 29: Gromphas aeruginosa. 29 a) Major specimen with cephalic projection extremely broad. 29 b) Major specimen with apical notch. 29 c) Worn major specimen. 29 d) Medium specimen. 29 e) Minor specimen. 30: G. lemoinei. 30 a) Major specimen. 30 b) Medium specimen. 30 c) Minor specimen. 31) G. amazonica. 32) G. inermis. 33) G. dichroa." httpUri="https://zenodo.org/record/247634/files/figure.png" pageId="15" pageNumber="454">Figs. 30</figureCitation>
a–c). Frons smooth adjacent to eyes and granulate in center (
<figureCitation id="60E52A555975624BFE42219735699886" box="[479,563,1682,1707]" captionStart="FIGURES 24 – 28" captionStartId="29.[151,264,1615,1637]" captionTargetBox="[151,1436,200,1593]" captionTargetId="figure@29.[151,1436,193,1593]" captionTargetPageId="29" captionText="FIGURES 24 – 28. Dorsal view of head. 24) Gromphas aeruginosa (only one pair of clypeal lobes and tegument smooth adjacent to eyes). 25) G. lemoinei (same). 26) G. dichroa (clypeus truncate and strong granulation around eyes). 27) G. amazonica (margin of clypeus with two pairs of lobes; strong granulation around eyes). 28) G. inermis (same)." httpUri="https://zenodo.org/record/247633/files/figure.png" pageId="15" pageNumber="454">Fig. 25</figureCitation>
).
</paragraph>
<paragraph id="F86136D05975624BFF5A21B3333A9972" blockId="15.[151,1436,1250,2031]" pageId="15" pageNumber="454">
Thorax: Lateral region of pronotum with dense squamous granulation (Fig. 17); granulation always dense, without gradual decrease toward posteromedian region; posterior region with sparse granulation abruptly delimited (
<figureCitation id="60E52A555975624BFF0221FB367B993A" box="[159,289,1790,1815]" captionStart="FIGURES 7 – 10" captionStartId="16.[151,264,1939,1961]" captionTargetBox="[158,1436,199,1910]" captionTargetId="figure@16.[151,1436,193,1918]" captionTargetPageId="16" captionText="FIGURES 7 – 10. Gromphas lemoinei Waterhouse, 1891. 7) Major specimen. 8 – 10) Series of intermediate and minor specimens. Scale = 5 mm." httpUri="https://zenodo.org/record/247629/files/figure.png" pageId="15" pageNumber="454">Figs. 7–10</figureCitation>
). Longitudinal midline with smooth and glossy tegument present at center of pronotum (easily observable in Fig. 3). Posterior pronotal fossae absent or greatly reduced only in major specimens; in other specimens, fossae present without granulation. Posterior margin of pronotum rounded.
</paragraph>
<paragraph id="F86136D05975624BFF5A206F355B99C2" blockId="15.[151,1436,1250,2031]" pageId="15" pageNumber="454">
Anterior region of pronotum with ornamentation ranging from two shallow impressions in anterior margin (
<figureCitation id="60E52A555975624BFF02208B3640998A" box="[159,282,1934,1959]" captionStart="FIGURES 7 – 10" captionStartId="16.[151,264,1939,1961]" captionTargetBox="[158,1436,199,1910]" captionTargetId="figure@16.[151,1436,193,1918]" captionTargetPageId="16" captionText="FIGURES 7 – 10. Gromphas lemoinei Waterhouse, 1891. 7) Major specimen. 8 – 10) Series of intermediate and minor specimens. Scale = 5 mm." httpUri="https://zenodo.org/record/247629/files/figure.png" pageId="15" pageNumber="454">Figs. 9–10</figureCitation>
) to a prominence with two wide, triangular projections; lateral margin of prominence always straight (
<figureCitation id="60E52A555975624BFF0220B7365199E6" box="[159,267,1970,1995]" captionStart="FIGURES 7 – 10" captionStartId="16.[151,264,1939,1961]" captionTargetBox="[158,1436,199,1910]" captionTargetId="figure@16.[151,1436,193,1918]" captionTargetPageId="16" captionText="FIGURES 7 – 10. Gromphas lemoinei Waterhouse, 1891. 7) Major specimen. 8 – 10) Series of intermediate and minor specimens. Scale = 5 mm." httpUri="https://zenodo.org/record/247629/files/figure.png" pageId="15" pageNumber="454">Figs. 7–8</figureCitation>
); separation between projections always very narrow; anterior to prominence a with a pair of concavities with smooth tegument (Fig. 17).
</paragraph>
<caption id="ACA16658596A6254FF0A209636FA99E5" httpUri="https://zenodo.org/record/247629/files/figure.png" pageId="16" pageNumber="455" targetBox="[158,1436,199,1910]" targetPageId="16">
<paragraph id="F86136D0596A6254FF0A209636FA99E5" blockId="16.[151,1435,1939,1992]" pageId="16" pageNumber="455">
<emphasis id="CAAAEAC2596A6254FF0A209634729984" bold="true" box="[151,808,1939,1961]" pageId="16" pageNumber="455">
FIGURES 7–10. 
<taxonomicName id="3FDE4D53596A6254FEC42096345B9984" authority="Waterhouse, 1891" authorityName="Waterhouse" authorityYear="1891" box="[345,769,1939,1961]" class="Insecta" family="Scarabaeidae" genus="Gromphas" kingdom="Animalia" order="Coleoptera" pageId="16" pageNumber="455" phylum="Arthropoda" rank="species" species="lemoinei">
<emphasis id="CAAAEAC2596A6254FEC4209635779984" bold="true" box="[345,557,1939,1961]" italics="true" pageId="16" pageNumber="455">Gromphas lemoinei</emphasis>
Waterhouse, 1891
</taxonomicName>
. 7)
</emphasis>
Major specimen. 
<emphasis id="CAAAEAC2596A6254FC6C209633719984" bold="true" box="[1009,1067,1939,1961]" pageId="16" pageNumber="455">8–10)</emphasis>
Series of intermediate and minor specimens. Scale = 5 mm.
</paragraph>
</caption>
<caption id="ACA16658596B6255FF0A20683408998F" httpUri="https://zenodo.org/record/247630/files/figure.png" pageId="17" pageNumber="456" targetBox="[161,1428,193,1879]" targetPageId="17">
<paragraph id="F86136D0596B6255FF0A20683408998F" blockId="17.[151,1433,1901,1954]" pageId="17" pageNumber="456">
<emphasis id="CAAAEAC2596B6255FF0A2068345299AE" bold="true" box="[151,776,1901,1923]" pageId="17" pageNumber="456">
FIGURES 11–12. 11: Holotype of 
<taxonomicName id="3FDE4D53596B6255FD9C2068359599AE" box="[513,719,1901,1923]" class="Insecta" family="Scarabaeidae" genus="Gromphas" kingdom="Animalia" order="Coleoptera" pageId="17" pageNumber="456" phylum="Arthropoda" rank="species" species="lemoinei">
<emphasis id="CAAAEAC2596B6255FD9C2068359599AE" bold="true" box="[513,719,1901,1923]" italics="true" pageId="17" pageNumber="456">Gromphas lemoinei</emphasis>
</taxonomicName>
. 11a)
</emphasis>
Dorsal view. 
<emphasis id="CAAAEAC2596B6255FC092068349A99AE" bold="true" box="[916,960,1901,1923]" pageId="17" pageNumber="456">11b)</emphasis>
Lateral view. 
<emphasis id="CAAAEAC2596B6255FBCF2068332699AE" bold="true" box="[1106,1148,1901,1923]" pageId="17" pageNumber="456">11c)</emphasis>
Labels. 
<emphasis id="CAAAEAC2596B6255FB4E2068351F998F" bold="true" pageId="17" pageNumber="456">
12: Lectotype of 
<taxonomicName id="3FDE4D53596B6255FA1B20683652998F" class="Insecta" family="Scarabaeidae" genus="Gromphas" kingdom="Animalia" order="Coleoptera" pageId="17" pageNumber="456" phylum="Arthropoda" rank="species" species="amazonica">
<emphasis id="CAAAEAC2596B6255FA1B20683652998F" bold="true" italics="true" pageId="17" pageNumber="456">G. amazonica</emphasis>
</taxonomicName>
from Tefé (former Ega). 12a)
</emphasis>
Dorsal view. 
<emphasis id="CAAAEAC2596B6255FD4C2089345B998F" bold="true" box="[721,769,1932,1954]" pageId="17" pageNumber="456">12b)</emphasis>
Labels.
</paragraph>
</caption>
<paragraph id="F86136D059686256FF5A279D320B9EF8" blockId="18.[151,1436,151,2013]" pageId="18" pageNumber="457">
Mesosternum with dense pilosity (
<figureCitation id="60E52A5559686256FDD7279235C59E9D" box="[586,671,151,176]" captionStart="FIGURES 49 – 52" captionStartId="32.[151,264,1223,1245]" captionTargetBox="[151,1436,193,1201]" captionTargetId="figure@32.[151,1436,193,1202]" captionTargetPageId="32" captionText="FIGURES 49 – 52. Anteromedian angle of metasternum. 49) Gromphas dichroa. 50) G. inermis. 51) G. amazonica. 52) G. aeruginosa (arrow indicates median carina of mesosternum)." httpUri="https://zenodo.org/record/247636/files/figure.png" pageId="18" pageNumber="457">Fig. 52</figureCitation>
). Metasternum very densely punctate in center. Anteromedian angle of metasternum flat and with rounded apex (
<figureCitation id="60E52A5559686256FD1527B935819EF8" box="[648,731,188,213]" captionStart="FIGURES 49 – 52" captionStartId="32.[151,264,1223,1245]" captionTargetBox="[151,1436,193,1201]" captionTargetId="figure@32.[151,1436,193,1202]" captionTargetPageId="32" captionText="FIGURES 49 – 52. Anteromedian angle of metasternum. 49) Gromphas dichroa. 50) G. inermis. 51) G. amazonica. 52) G. aeruginosa (arrow indicates median carina of mesosternum)." httpUri="https://zenodo.org/record/247636/files/figure.png" pageId="18" pageNumber="457">Fig. 52</figureCitation>
), area in front of angle with evident setae (Figs. 16, 52).
</paragraph>
<paragraph id="F86136D059686256FF5A27E533BC9FA5" blockId="18.[151,1436,151,2013]" pageId="18" pageNumber="457">
Legs: Protibiae very broad in both sexes; in ventral view, longitudinal carina simple in both sexes (
<figureCitation id="60E52A5559686256FAA427DA32D59ED5" box="[1337,1423,223,248]" captionStart="FIGURES 21 – 23" captionStartId="28.[151,264,1921,1943]" captionTargetBox="[248,1348,181,1909]" captionTargetId="figure@28.[248,1349,181,1909]" captionTargetPageId="28" captionText="FIGURES 21 – 23. Protibiae. 21: Dorsal view of protibia of Gromphas amazonica. 21 a) Female. 21 b) Male (arrow indicates non-articulating spur). 22) Ventral view of female protibia of G. aeruginosa. 23: Ventral view of protibia of G. inermis. 23 a) Female. 23 b) Male (white arrow indicates tubercle of inner apical angle and blue arrow indicates row of tubercles on ventral carina of protibia)." httpUri="https://zenodo.org/record/247632/files/figure.png" pageId="18" pageNumber="457">Fig. 22</figureCitation>
). Protibial spur with apex slightly bent laterally and downward (
<figureCitation id="60E52A5559686256FCCB260134EE9F31" box="[854,948,260,285]" captionStart="FIGURES 21 – 23" captionStartId="28.[151,264,1921,1943]" captionTargetBox="[248,1348,181,1909]" captionTargetId="figure@28.[248,1349,181,1909]" captionTargetPageId="28" captionText="FIGURES 21 – 23. Protibiae. 21: Dorsal view of protibia of Gromphas amazonica. 21 a) Female. 21 b) Male (arrow indicates non-articulating spur). 22) Ventral view of female protibia of G. aeruginosa. 23: Ventral view of protibia of G. inermis. 23 a) Female. 23 b) Male (white arrow indicates tubercle of inner apical angle and blue arrow indicates row of tubercles on ventral carina of protibia)." httpUri="https://zenodo.org/record/247632/files/figure.png" pageId="18" pageNumber="457">Figs. 22</figureCitation>
, 
<figureCitation id="60E52A5559686256FC22260134859F30" box="[959,991,260,285]" captionStart="FIGURES 34 – 48. 34" captionStartId="31.[151,264,1827,1849]" captionTargetBox="[241,1350,179,1803]" captionTargetId="figure@31.[241,1350,179,1804]" captionTargetPageId="31" captionText="FIGURES 34 – 48. 34: Sixth visible abdominal sternite of Gromphas aeruginosa. 34 a) Male. 34 b) Female. 35) Margined pygidium of G. dichroa. 36) Non-margined pygidium of G. aeruginosa. 37) Elytral striae of G. aeruginosa. 38) Elytral striae of G. amazonica. 39) Curved metatibial spur of G. aeruginosa. 40) Straight metatibial spur of G. inermis. 41) Robust metatibia of G. aeruginosa. 42) Thin metatibia of G. d i c h ro a. 43) Posterior pronotal fossae present in some G. i n e rm i s. 44) Antennal club of G. dichroa. 45) Protarsus and expanded protibial spur of G. amazonica. 46) Protarsus and straight protibial spur of G. aeruginosa. 47) Metatarsus of G. dichroa with apical tarsomere strongly curved. 48) Metatarsus of G. aeruginosa with apical tarsomere only slighty curved at apex." httpUri="https://zenodo.org/record/247635/files/figure.png" pageId="18" pageNumber="457">46</figureCitation>
); inner apical protibial angle with a tuft of long setae in both sexes (
<figureCitation id="60E52A5559686256FE522622357E9F6D" box="[463,548,295,320]" captionStart="FIGURES 21 – 23" captionStartId="28.[151,264,1921,1943]" captionTargetBox="[248,1348,181,1909]" captionTargetId="figure@28.[248,1349,181,1909]" captionTargetPageId="28" captionText="FIGURES 21 – 23. Protibiae. 21: Dorsal view of protibia of Gromphas amazonica. 21 a) Female. 21 b) Male (arrow indicates non-articulating spur). 22) Ventral view of female protibia of G. aeruginosa. 23: Ventral view of protibia of G. inermis. 23 a) Female. 23 b) Male (white arrow indicates tubercle of inner apical angle and blue arrow indicates row of tubercles on ventral carina of protibia)." httpUri="https://zenodo.org/record/247632/files/figure.png" pageId="18" pageNumber="457">Fig. 22</figureCitation>
). Apical protarsomere tapered and only slightly elongate at apex (
<figureCitation id="60E52A5559686256FA9B262232399F6D" box="[1286,1379,295,320]" captionStart="FIGURES 21 – 23" captionStartId="28.[151,264,1921,1943]" captionTargetBox="[248,1348,181,1909]" captionTargetId="figure@28.[248,1349,181,1909]" captionTargetPageId="28" captionText="FIGURES 21 – 23. Protibiae. 21: Dorsal view of protibia of Gromphas amazonica. 21 a) Female. 21 b) Male (arrow indicates non-articulating spur). 22) Ventral view of female protibia of G. aeruginosa. 23: Ventral view of protibia of G. inermis. 23 a) Female. 23 b) Male (white arrow indicates tubercle of inner apical angle and blue arrow indicates row of tubercles on ventral carina of protibia)." httpUri="https://zenodo.org/record/247632/files/figure.png" pageId="18" pageNumber="457">Figs. 22</figureCitation>
, 
<figureCitation id="60E52A5559686256FAED262232CA9F6D" box="[1392,1424,295,320]" captionStart="FIGURES 34 – 48. 34" captionStartId="31.[151,264,1827,1849]" captionTargetBox="[241,1350,179,1803]" captionTargetId="figure@31.[241,1350,179,1804]" captionTargetPageId="31" captionText="FIGURES 34 – 48. 34: Sixth visible abdominal sternite of Gromphas aeruginosa. 34 a) Male. 34 b) Female. 35) Margined pygidium of G. dichroa. 36) Non-margined pygidium of G. aeruginosa. 37) Elytral striae of G. aeruginosa. 38) Elytral striae of G. amazonica. 39) Curved metatibial spur of G. aeruginosa. 40) Straight metatibial spur of G. inermis. 41) Robust metatibia of G. aeruginosa. 42) Thin metatibia of G. d i c h ro a. 43) Posterior pronotal fossae present in some G. i n e rm i s. 44) Antennal club of G. dichroa. 45) Protarsus and expanded protibial spur of G. amazonica. 46) Protarsus and straight protibial spur of G. aeruginosa. 47) Metatarsus of G. dichroa with apical tarsomere strongly curved. 48) Metatarsus of G. aeruginosa with apical tarsomere only slighty curved at apex." httpUri="https://zenodo.org/record/247635/files/figure.png" pageId="18" pageNumber="457">46</figureCitation>
). Metatibiae very broad and robust (
<figureCitation id="60E52A5559686256FDAD264935D29F49" box="[560,648,332,357]" captionStart="FIGURES 34 – 48. 34" captionStartId="31.[151,264,1827,1849]" captionTargetBox="[241,1350,179,1803]" captionTargetId="figure@31.[241,1350,179,1804]" captionTargetPageId="31" captionText="FIGURES 34 – 48. 34: Sixth visible abdominal sternite of Gromphas aeruginosa. 34 a) Male. 34 b) Female. 35) Margined pygidium of G. dichroa. 36) Non-margined pygidium of G. aeruginosa. 37) Elytral striae of G. aeruginosa. 38) Elytral striae of G. amazonica. 39) Curved metatibial spur of G. aeruginosa. 40) Straight metatibial spur of G. inermis. 41) Robust metatibia of G. aeruginosa. 42) Thin metatibia of G. d i c h ro a. 43) Posterior pronotal fossae present in some G. i n e rm i s. 44) Antennal club of G. dichroa. 45) Protarsus and expanded protibial spur of G. amazonica. 46) Protarsus and straight protibial spur of G. aeruginosa. 47) Metatarsus of G. dichroa with apical tarsomere strongly curved. 48) Metatarsus of G. aeruginosa with apical tarsomere only slighty curved at apex." httpUri="https://zenodo.org/record/247635/files/figure.png" pageId="18" pageNumber="457">Fig. 41</figureCitation>
). Mesotarsi and metatarsi not particularly enlarged and with apical tarsomeres slightly curved apically (
<figureCitation id="60E52A5559686256FDB0266A35D89FA5" box="[557,642,367,392]" captionStart="FIGURES 34 – 48. 34" captionStartId="31.[151,264,1827,1849]" captionTargetBox="[241,1350,179,1803]" captionTargetId="figure@31.[241,1350,179,1804]" captionTargetPageId="31" captionText="FIGURES 34 – 48. 34: Sixth visible abdominal sternite of Gromphas aeruginosa. 34 a) Male. 34 b) Female. 35) Margined pygidium of G. dichroa. 36) Non-margined pygidium of G. aeruginosa. 37) Elytral striae of G. aeruginosa. 38) Elytral striae of G. amazonica. 39) Curved metatibial spur of G. aeruginosa. 40) Straight metatibial spur of G. inermis. 41) Robust metatibia of G. aeruginosa. 42) Thin metatibia of G. d i c h ro a. 43) Posterior pronotal fossae present in some G. i n e rm i s. 44) Antennal club of G. dichroa. 45) Protarsus and expanded protibial spur of G. amazonica. 46) Protarsus and straight protibial spur of G. aeruginosa. 47) Metatarsus of G. dichroa with apical tarsomere strongly curved. 48) Metatarsus of G. aeruginosa with apical tarsomere only slighty curved at apex." httpUri="https://zenodo.org/record/247635/files/figure.png" pageId="18" pageNumber="457">Fig. 48</figureCitation>
). Metatibial spur with apex distinctly curved (
<figureCitation id="60E52A5559686256FB18266A33809FA5" box="[1157,1242,367,392]" captionStart="FIGURES 34 – 48. 34" captionStartId="31.[151,264,1827,1849]" captionTargetBox="[241,1350,179,1803]" captionTargetId="figure@31.[241,1350,179,1804]" captionTargetPageId="31" captionText="FIGURES 34 – 48. 34: Sixth visible abdominal sternite of Gromphas aeruginosa. 34 a) Male. 34 b) Female. 35) Margined pygidium of G. dichroa. 36) Non-margined pygidium of G. aeruginosa. 37) Elytral striae of G. aeruginosa. 38) Elytral striae of G. amazonica. 39) Curved metatibial spur of G. aeruginosa. 40) Straight metatibial spur of G. inermis. 41) Robust metatibia of G. aeruginosa. 42) Thin metatibia of G. d i c h ro a. 43) Posterior pronotal fossae present in some G. i n e rm i s. 44) Antennal club of G. dichroa. 45) Protarsus and expanded protibial spur of G. amazonica. 46) Protarsus and straight protibial spur of G. aeruginosa. 47) Metatarsus of G. dichroa with apical tarsomere strongly curved. 48) Metatarsus of G. aeruginosa with apical tarsomere only slighty curved at apex." httpUri="https://zenodo.org/record/247635/files/figure.png" pageId="18" pageNumber="457">Fig. 39</figureCitation>
).
</paragraph>
<paragraph id="F86136D059686256FF5A269134E49FD9" blockId="18.[151,1436,151,2013]" pageId="18" pageNumber="457">
Elytra: Striae very fine and, especially those more medial, carinulate from base to no more than midlength of elytra, where they become simple (
<figureCitation id="60E52A5559686256FDB026B235D99FFD" box="[557,643,439,464]" captionStart="FIGURES 34 – 48. 34" captionStartId="31.[151,264,1827,1849]" captionTargetBox="[241,1350,179,1803]" captionTargetId="figure@31.[241,1350,179,1804]" captionTargetPageId="31" captionText="FIGURES 34 – 48. 34: Sixth visible abdominal sternite of Gromphas aeruginosa. 34 a) Male. 34 b) Female. 35) Margined pygidium of G. dichroa. 36) Non-margined pygidium of G. aeruginosa. 37) Elytral striae of G. aeruginosa. 38) Elytral striae of G. amazonica. 39) Curved metatibial spur of G. aeruginosa. 40) Straight metatibial spur of G. inermis. 41) Robust metatibia of G. aeruginosa. 42) Thin metatibia of G. d i c h ro a. 43) Posterior pronotal fossae present in some G. i n e rm i s. 44) Antennal club of G. dichroa. 45) Protarsus and expanded protibial spur of G. amazonica. 46) Protarsus and straight protibial spur of G. aeruginosa. 47) Metatarsus of G. dichroa with apical tarsomere strongly curved. 48) Metatarsus of G. aeruginosa with apical tarsomere only slighty curved at apex." httpUri="https://zenodo.org/record/247635/files/figure.png" pageId="18" pageNumber="457">Fig. 37</figureCitation>
); interstriae, especially mesial, slightly convex, producing a grooved appearance of striae to the naked eyes. Sutural margin densely punctate.
</paragraph>
<paragraph id="F86136D059686256FF5A26FA35999C11" blockId="18.[151,1436,151,2013]" pageId="18" pageNumber="457">
Abdomen: Pygidium flat, not margined basally (
<figureCitation id="60E52A5559686256FC9126FA343D9C35" box="[780,871,511,536]" captionStart="FIGURES 34 – 48. 34" captionStartId="31.[151,264,1827,1849]" captionTargetBox="[241,1350,179,1803]" captionTargetId="figure@31.[241,1350,179,1804]" captionTargetPageId="31" captionText="FIGURES 34 – 48. 34: Sixth visible abdominal sternite of Gromphas aeruginosa. 34 a) Male. 34 b) Female. 35) Margined pygidium of G. dichroa. 36) Non-margined pygidium of G. aeruginosa. 37) Elytral striae of G. aeruginosa. 38) Elytral striae of G. amazonica. 39) Curved metatibial spur of G. aeruginosa. 40) Straight metatibial spur of G. inermis. 41) Robust metatibia of G. aeruginosa. 42) Thin metatibia of G. d i c h ro a. 43) Posterior pronotal fossae present in some G. i n e rm i s. 44) Antennal club of G. dichroa. 45) Protarsus and expanded protibial spur of G. amazonica. 46) Protarsus and straight protibial spur of G. aeruginosa. 47) Metatarsus of G. dichroa with apical tarsomere strongly curved. 48) Metatarsus of G. aeruginosa with apical tarsomere only slighty curved at apex." httpUri="https://zenodo.org/record/247635/files/figure.png" pageId="18" pageNumber="457">Fig. 36</figureCitation>
). Groove of propygidium extending to base of pygidium. Abdominal sternites sparsely punctate.
</paragraph>
<paragraph id="F86136D059686256FF5A254235E39CA8" blockId="18.[151,1436,151,2013]" pageId="18" pageNumber="457">
Aedeagus: Phallobase in ventral view with a very narrow apical membranous area (
<figureCitation id="60E52A5559686256FBE02542338B9C4D" box="[1149,1233,583,608]" captionStart="FIGURES 53 – 56" captionStartId="33.[151,264,1243,1265]" captionTargetBox="[159,1430,195,1222]" captionTargetId="figure@33.[156,1431,193,1222]" captionTargetPageId="33" captionText="FIGURES 53 – 56. Schematic drawings of aedeagi (a—dorsal view, b—lateral view, c—ventral view; dashed lines delimit ventral groove). 53) Gromphas aeruginosa. 54) G. dichroa. 55) G. inermis. 56) G. amazonica." httpUri="https://zenodo.org/record/247637/files/figure.png" pageId="18" pageNumber="457">Fig. 53</figureCitation>
c). Medial sclerite wide, flat and strongly curved laterally (
<figureCitation id="60E52A5559686256FDC5256935F69CA8" box="[600,684,620,645]" captionStart="FIGURES 57 – 62" captionStartId="34.[151,264,1134,1156]" captionTargetBox="[433,1142,199,1103]" captionTargetId="figure@34.[428,1160,193,1113]" captionTargetPageId="34" captionText="FIGURES 57 – 62. Schematic drawings. 57 – 61: Medial sclerite of internal sac. 57) Gromphas aeruginosa. 58) G. lemoinei. 59) G. inermis. 60) G. amazonica. 61) G. dichroa. 62) Internal sac of G. aeruginosa (TE = temones; MS = medial sclerite; AS = apical sclerites)." httpUri="https://zenodo.org/record/247638/files/figure.png" pageId="18" pageNumber="457">Fig. 58</figureCitation>
).
</paragraph>
<paragraph id="F86136D059686256FF5A258A32CF9D15" blockId="18.[151,1436,151,2013]" pageId="18" pageNumber="457">
<emphasis id="CAAAEAC259686256FF5A258A35989C85" bold="true" box="[199,706,655,680]" pageId="18" pageNumber="457">
Measurements: Males (
<specimenCount id="EED8FD5959686256FE41258A35259C85" box="[476,639,655,680]" pageId="18" pageNumber="457" type="generic">10 specimens</specimenCount>
): TL:
</emphasis>
AV: 16; 
<collectingCountry id="80C9764059686256FCB92595340F9C8A" box="[804,853,656,679]" name="Mexico" pageId="18" pageNumber="457">MX</collectingCountry>
: 20.1; MN: 12.7; SD: 2.04. 
<emphasis id="CAAAEAC259686256FB17258A33ED9C85" bold="true" box="[1162,1207,655,680]" pageId="18" pageNumber="457">PL:</emphasis>
AV: 13.07; 
<collectingCountry id="80C9764059686256FAA0259532359C8A" box="[1341,1391,656,679]" name="Mexico" pageId="18" pageNumber="457">MX</collectingCountry>
: 17; MN: 10.3; SD: 1.9. 
<emphasis id="CAAAEAC259686256FE1E25B136E29CE0" bold="true" box="[387,440,692,717]" pageId="18" pageNumber="457">PW:</emphasis>
AV: 9.4; 
<collectingCountry id="80C9764059686256FDB125B035079CE1" box="[556,605,693,716]" name="Mexico" pageId="18" pageNumber="457">MX</collectingCountry>
: 11.7; MN: 7.1; SD: 1.29. 
<emphasis id="CAAAEAC259686256FC0425B1349E9CE0" bold="true" box="[921,964,692,717]" pageId="18" pageNumber="457">PP:</emphasis>
AV: 2.51; 
<collectingCountry id="80C9764059686256FBDB25B0332D9CE1" box="[1094,1143,693,716]" name="Mexico" pageId="18" pageNumber="457">MX</collectingCountry>
: 3.8; MN: 1.4; SD: 0.40. 
<emphasis id="CAAAEAC259686256FF0A25D236BB9CDD" bold="true" box="[151,481,727,752]" pageId="18" pageNumber="457">
Females (
<specimenCount id="EED8FD5959686256FE9425D236C49CDD" box="[265,414,727,752]" pageId="18" pageNumber="457" type="generic">4 specimens</specimenCount>
): TL:
</emphasis>
AV: 17.3; 
<collectingCountry id="80C9764059686256FDC725DD35D19CC2" box="[602,651,728,751]" name="Mexico" pageId="18" pageNumber="457">MX</collectingCountry>
: 19.2; MN: 13.9; SD: 2.09. 
<emphasis id="CAAAEAC259686256FC5925D234AB9CDD" bold="true" box="[964,1009,727,752]" pageId="18" pageNumber="457">PL:</emphasis>
AV: 13.73; 
<collectingCountry id="80C9764059686256FBE425DD33F09CC2" box="[1145,1194,728,751]" name="Mexico" pageId="18" pageNumber="457">MX</collectingCountry>
: 14.8; MN: 11.2; SD: 0.9. 
<emphasis id="CAAAEAC259686256FF5025F936589D38" bold="true" box="[205,258,764,789]" pageId="18" pageNumber="457">PW:</emphasis>
AV: 9.78; 
<collectingCountry id="80C9764059686256FE1125F836E79D39" box="[396,445,765,788]" name="Mexico" pageId="18" pageNumber="457">MX</collectingCountry>
: 10.4; MN: 8.3; SD: 0.63. 
<emphasis id="CAAAEAC259686256FC9425F9346E9D38" bold="true" box="[777,820,764,789]" pageId="18" pageNumber="457">PP:</emphasis>
AV: 2.63; 
<collectingCountry id="80C9764059686256FC2325F834B59D39" box="[958,1007,765,788]" name="Mexico" pageId="18" pageNumber="457">MX</collectingCountry>
: 3.1; MN: 2.1; SD: 0.46. 
<emphasis id="CAAAEAC259686256FAB125F936039D15" bold="true" pageId="18" pageNumber="457">
Total (
<specimenCount id="EED8FD5959686256FA1D25F9364C9D15" pageId="18" pageNumber="457" type="generic">14 specimens</specimenCount>
): TL:
</emphasis>
AV: 16.38; SD: 2.06. 
<emphasis id="CAAAEAC259686256FDCD241A35279D15" bold="true" box="[592,637,799,824]" pageId="18" pageNumber="457">PL:</emphasis>
AV: 13.25; SD: 1.68. 
<emphasis id="CAAAEAC259686256FCE8241A34F09D15" bold="true" box="[885,938,799,824]" pageId="18" pageNumber="457">PW:</emphasis>
AV: 9.5; SD: 1.14. 
<emphasis id="CAAAEAC259686256FB1B241A33EB9D15" bold="true" box="[1158,1201,799,824]" pageId="18" pageNumber="457">PP:</emphasis>
AV: 2.54; SD: 0.62.
</paragraph>
<paragraph id="F86136D059686256FF5A244133AE9BB0" blockId="18.[151,1436,151,2013]" pageId="18" pageNumber="457">
<emphasis id="CAAAEAC259686256FF5A2441344F9D70" bold="true" box="[199,789,836,861]" pageId="18" pageNumber="457">Intraspecific variation and taxonomic discussion:</emphasis>
The closest relative of 
<taxonomicName id="3FDE4D5359686256FB80244033C69D70" box="[1053,1180,836,861]" class="Insecta" family="Scarabaeidae" genus="Gromphas" kingdom="Animalia" order="Coleoptera" pageId="18" pageNumber="457" phylum="Arthropoda" rank="species" species="lemoinei">
<emphasis id="CAAAEAC259686256FB80244033C69D70" box="[1053,1180,836,861]" italics="true" pageId="18" pageNumber="457">G. lemoinei</emphasis>
</taxonomicName>
is 
<taxonomicName id="3FDE4D5359686256FB23244032069D71" box="[1214,1372,837,860]" class="Insecta" family="Scarabaeidae" genus="Gromphas" kingdom="Animalia" order="Coleoptera" pageId="18" pageNumber="457" phylum="Arthropoda" rank="species" species="aeruginosa">
<emphasis id="CAAAEAC259686256FB23244032069D71" box="[1214,1372,837,860]" italics="true" pageId="18" pageNumber="457">G. aeruginosa</emphasis>
</taxonomicName>
, with which it was synonymized for more than a century. The description given by Waterhouse (1891), although short and based only on a single specimen, is very accurate and uses basically the same morphological features used by us to distinguish these two species; it is surprising, therefore, that this synonymy has remained for so long. Gillet (1911) was the first author to mention Waterhouse’s species as “ 
<taxonomicName id="3FDE4D5359686256FCF524D134A09DC0" box="[872,1018,980,1005]" pageId="18" pageNumber="457" rank="variety" variety="lemoinei">
var. 
<emphasis id="CAAAEAC259686256FC0724D134A09DC0" box="[922,1018,980,1005]" italics="true" pageId="18" pageNumber="457">lemoinei</emphasis>
</taxonomicName>
” of 
<taxonomicName id="3FDE4D5359686256FBB024D033909DC1" box="[1069,1226,981,1004]" class="Insecta" family="Scarabaeidae" genus="Gromphas" kingdom="Animalia" order="Coleoptera" pageId="18" pageNumber="457" phylum="Arthropoda" rank="species" species="aeruginosa">
<emphasis id="CAAAEAC259686256FBB024D033909DC1" box="[1069,1226,981,1004]" italics="true" pageId="18" pageNumber="457">G. aeruginosa</emphasis>
</taxonomicName>
, not making clear, however, whether or not he considered them synonymous; d’Olsoufieff (1924) was the first to state unequivocally this synonymy. Nevertheless, neither of these two authors nor even Barattini &amp; Saénz (1960, 1964) discussed or justified this synonymy or the condition of variety of 
<taxonomicName id="3FDE4D5359686256FD65234534229A75" box="[760,888,1087,1112]" class="Insecta" family="Scarabaeidae" genus="Gromphas" kingdom="Animalia" order="Coleoptera" pageId="18" pageNumber="457" phylum="Arthropoda" rank="species" species="lemoinei">
<emphasis id="CAAAEAC259686256FD65234534229A75" box="[760,888,1087,1112]" italics="true" pageId="18" pageNumber="457">G. lemoinei</emphasis>
</taxonomicName>
, and it is very likely that they have not seen the 
<typeStatus id="2765887259686256FF0A2360379D9A50" box="[151,199,1125,1149]" pageId="18" pageNumber="457">type</typeStatus>
specimens. All this confusion probably occurred simply because of the superficial similarity that these two species show at the first glance; as they share only between themselves the following characteristics: genae and frons smooth adjacent to eyes (
<figureCitation id="60E52A5559686256FE6823A935259AE8" box="[501,639,1196,1221]" captionStart="FIGURES 24 – 28" captionStartId="29.[151,264,1615,1637]" captionTargetBox="[151,1436,200,1593]" captionTargetId="figure@29.[151,1436,193,1593]" captionTargetPageId="29" captionText="FIGURES 24 – 28. Dorsal view of head. 24) Gromphas aeruginosa (only one pair of clypeal lobes and tegument smooth adjacent to eyes). 25) G. lemoinei (same). 26) G. dichroa (clypeus truncate and strong granulation around eyes). 27) G. amazonica (margin of clypeus with two pairs of lobes; strong granulation around eyes). 28) G. inermis (same)." httpUri="https://zenodo.org/record/247633/files/figure.png" pageId="18" pageNumber="457">Figs. 24–25</figureCitation>
); cephalic projection very high (
<figureCitation id="60E52A5559686256FC7623A9332E9AE9" box="[1003,1140,1196,1221]" captionStart="FIGURES 29 – 33" captionStartId="30.[151,264,1916,1938]" captionTargetBox="[195,1380,179,1890]" captionTargetId="figure@30.[195,1380,179,1891]" captionTargetPageId="30" captionText="FIGURES 29 – 33. Frontal view of cephalic projections. 29: Gromphas aeruginosa. 29 a) Major specimen with cephalic projection extremely broad. 29 b) Major specimen with apical notch. 29 c) Worn major specimen. 29 d) Medium specimen. 29 e) Minor specimen. 30: G. lemoinei. 30 a) Major specimen. 30 b) Medium specimen. 30 c) Minor specimen. 31) G. amazonica. 32) G. inermis. 33) G. dichroa." httpUri="https://zenodo.org/record/247634/files/figure.png" pageId="18" pageNumber="457">Figs. 29–30</figureCitation>
); pronotum with a smooth midlongitudinal line; presence of pronotal prominence (Figs. 1–5, 7–10); flattened anteromedian angle of metasternum (
<figureCitation id="60E52A5559686256FEA523F136D79B20" box="[312,397,1268,1293]" captionStart="FIGURES 49 – 52" captionStartId="32.[151,264,1223,1245]" captionTargetBox="[151,1436,193,1201]" captionTargetId="figure@32.[151,1436,193,1202]" captionTargetPageId="32" captionText="FIGURES 49 – 52. Anteromedian angle of metasternum. 49) Gromphas dichroa. 50) G. inermis. 51) G. amazonica. 52) G. aeruginosa (arrow indicates median carina of mesosternum)." httpUri="https://zenodo.org/record/247636/files/figure.png" pageId="18" pageNumber="457">Fig. 52</figureCitation>
); tibiae very broad in both sexes (
<figureCitation id="60E52A5559686256FC9223F1343E9B21" box="[783,868,1268,1293]" captionStart="FIGURES 21 – 23" captionStartId="28.[151,264,1921,1943]" captionTargetBox="[248,1348,181,1909]" captionTargetId="figure@28.[248,1349,181,1909]" captionTargetPageId="28" captionText="FIGURES 21 – 23. Protibiae. 21: Dorsal view of protibia of Gromphas amazonica. 21 a) Female. 21 b) Male (arrow indicates non-articulating spur). 22) Ventral view of female protibia of G. aeruginosa. 23: Ventral view of protibia of G. inermis. 23 a) Female. 23 b) Male (white arrow indicates tubercle of inner apical angle and blue arrow indicates row of tubercles on ventral carina of protibia)." httpUri="https://zenodo.org/record/247632/files/figure.png" pageId="18" pageNumber="457">Fig. 22</figureCitation>
); protibia with ventral carina simple in both sexes and with apical tuft of setae without apparent sexual dimorphism (
<figureCitation id="60E52A5559686256FC12221234BF9B1D" box="[911,997,1303,1328]" captionStart="FIGURES 21 – 23" captionStartId="28.[151,264,1921,1943]" captionTargetBox="[248,1348,181,1909]" captionTargetId="figure@28.[248,1349,181,1909]" captionTargetPageId="28" captionText="FIGURES 21 – 23. Protibiae. 21: Dorsal view of protibia of Gromphas amazonica. 21 a) Female. 21 b) Male (arrow indicates non-articulating spur). 22) Ventral view of female protibia of G. aeruginosa. 23: Ventral view of protibia of G. inermis. 23 a) Female. 23 b) Male (white arrow indicates tubercle of inner apical angle and blue arrow indicates row of tubercles on ventral carina of protibia)." httpUri="https://zenodo.org/record/247632/files/figure.png" pageId="18" pageNumber="457">Fig. 22</figureCitation>
); protibial spur only slightly curved at apex (
<figureCitation id="60E52A5559686256FF46223936639B79" box="[219,313,1340,1365]" captionStart="FIGURES 21 – 23" captionStartId="28.[151,264,1921,1943]" captionTargetBox="[248,1348,181,1909]" captionTargetId="figure@28.[248,1349,181,1909]" captionTargetPageId="28" captionText="FIGURES 21 – 23. Protibiae. 21: Dorsal view of protibia of Gromphas amazonica. 21 a) Female. 21 b) Male (arrow indicates non-articulating spur). 22) Ventral view of female protibia of G. aeruginosa. 23: Ventral view of protibia of G. inermis. 23 a) Female. 23 b) Male (white arrow indicates tubercle of inner apical angle and blue arrow indicates row of tubercles on ventral carina of protibia)." httpUri="https://zenodo.org/record/247632/files/figure.png" pageId="18" pageNumber="457">Figs. 22</figureCitation>
, 
<figureCitation id="60E52A5559686256FED92239363E9B78" box="[324,356,1340,1365]" captionStart="FIGURES 34 – 48. 34" captionStartId="31.[151,264,1827,1849]" captionTargetBox="[241,1350,179,1803]" captionTargetId="figure@31.[241,1350,179,1804]" captionTargetPageId="31" captionText="FIGURES 34 – 48. 34: Sixth visible abdominal sternite of Gromphas aeruginosa. 34 a) Male. 34 b) Female. 35) Margined pygidium of G. dichroa. 36) Non-margined pygidium of G. aeruginosa. 37) Elytral striae of G. aeruginosa. 38) Elytral striae of G. amazonica. 39) Curved metatibial spur of G. aeruginosa. 40) Straight metatibial spur of G. inermis. 41) Robust metatibia of G. aeruginosa. 42) Thin metatibia of G. d i c h ro a. 43) Posterior pronotal fossae present in some G. i n e rm i s. 44) Antennal club of G. dichroa. 45) Protarsus and expanded protibial spur of G. amazonica. 46) Protarsus and straight protibial spur of G. aeruginosa. 47) Metatarsus of G. dichroa with apical tarsomere strongly curved. 48) Metatarsus of G. aeruginosa with apical tarsomere only slighty curved at apex." httpUri="https://zenodo.org/record/247635/files/figure.png" pageId="18" pageNumber="457">46</figureCitation>
); elytral striae carinulae extending at most only to midlenght of elytron (
<figureCitation id="60E52A5559686256FB10223933BA9B79" box="[1165,1248,1340,1365]" captionStart="FIGURES 34 – 48. 34" captionStartId="31.[151,264,1827,1849]" captionTargetBox="[241,1350,179,1803]" captionTargetId="figure@31.[241,1350,179,1804]" captionTargetPageId="31" captionText="FIGURES 34 – 48. 34: Sixth visible abdominal sternite of Gromphas aeruginosa. 34 a) Male. 34 b) Female. 35) Margined pygidium of G. dichroa. 36) Non-margined pygidium of G. aeruginosa. 37) Elytral striae of G. aeruginosa. 38) Elytral striae of G. amazonica. 39) Curved metatibial spur of G. aeruginosa. 40) Straight metatibial spur of G. inermis. 41) Robust metatibia of G. aeruginosa. 42) Thin metatibia of G. d i c h ro a. 43) Posterior pronotal fossae present in some G. i n e rm i s. 44) Antennal club of G. dichroa. 45) Protarsus and expanded protibial spur of G. amazonica. 46) Protarsus and straight protibial spur of G. aeruginosa. 47) Metatarsus of G. dichroa with apical tarsomere strongly curved. 48) Metatarsus of G. aeruginosa with apical tarsomere only slighty curved at apex." httpUri="https://zenodo.org/record/247635/files/figure.png" pageId="18" pageNumber="457">Fig. 37</figureCitation>
); and phallobase with a narrow apical membranous area (
<figureCitation id="60E52A5559686256FDC6225A35F59B55" box="[603,687,1375,1400]" captionStart="FIGURES 53 – 56" captionStartId="33.[151,264,1243,1265]" captionTargetBox="[159,1430,195,1222]" captionTargetId="figure@33.[156,1431,193,1222]" captionTargetPageId="33" captionText="FIGURES 53 – 56. Schematic drawings of aedeagi (a—dorsal view, b—lateral view, c—ventral view; dashed lines delimit ventral groove). 53) Gromphas aeruginosa. 54) G. dichroa. 55) G. inermis. 56) G. amazonica." httpUri="https://zenodo.org/record/247637/files/figure.png" pageId="18" pageNumber="457">Fig. 53</figureCitation>
c). Despite this extensive list of similarities, several morphological and biogeographical differences convincingly demonstrate the validity of each of these two species.
</paragraph>
<paragraph id="F86136D059686256FF5A22AC34649890" blockId="18.[151,1436,151,2013]" pageId="18" pageNumber="457">
Morrone (2001, 2006) recognized four subregions of the Neotropical region (Caribbean, Amazonian, Chacoan, and Parana), each them divided into several provinces. 
<taxonomicName id="3FDE4D5359686256FC8022C934A79BC8" box="[797,1021,1484,1509]" class="Insecta" family="Scarabaeidae" genus="Gromphas" kingdom="Animalia" order="Coleoptera" pageId="18" pageNumber="457" phylum="Arthropoda" rank="species" species="lemoinei">
<emphasis id="CAAAEAC259686256FC8022C934A79BC8" box="[797,1021,1484,1509]" italics="true" pageId="18" pageNumber="457">Gromphas lemoinei</emphasis>
</taxonomicName>
and 
<taxonomicName id="3FDE4D5359686256FBA722C833809BC9" box="[1082,1242,1485,1508]" class="Insecta" family="Scarabaeidae" genus="Gromphas" kingdom="Animalia" order="Coleoptera" pageId="18" pageNumber="457" phylum="Arthropoda" rank="species" species="aeruginosa">
<emphasis id="CAAAEAC259686256FBA722C833809BC9" box="[1082,1242,1485,1508]" italics="true" pageId="18" pageNumber="457">G. aeruginosa</emphasis>
</taxonomicName>
inhabit northern South 
<collectingCountry id="80C9764059686256FF7C22EA361E9825" box="[225,324,1519,1544]" name="United States of America" pageId="18" pageNumber="457">America</collectingCountry>
, but have widely separated distributions. The former species occurs in the Venezuelan Coast and Venezuelan Llanos provinces of the Caribbean subregion, while the latter species is widely distributed in eight provinces of the Amazonian subregion and in the north of Chaco province of Chacoan subregion. Nowhere are these two species found in sympatry; in fact, they are separated by large physiographic barriers such as the 
<collectingCountry id="80C9764059686256FAD6215932C69858" box="[1355,1436,1628,1653]" name="Guyana" pageId="18" pageNumber="457">Guiana</collectingCountry>
Shield and the Orinoco River. This geographical separation certainly has contributed to the emergence and maintenance of the morphological distinction between them.
</paragraph>
<paragraph id="F86136D059686257FF5A21C234AB9F12" blockId="18.[151,1436,151,2013]" lastBlockId="19.[151,1437,151,2013]" lastPageId="19" lastPageNumber="458" pageId="18" pageNumber="457">
The main distinction in the external morphology between 
<taxonomicName id="3FDE4D5359686256FCCB21CD34A998CD" box="[854,1011,1736,1760]" class="Insecta" family="Scarabaeidae" genus="Gromphas" kingdom="Animalia" order="Coleoptera" pageId="18" pageNumber="457" phylum="Arthropoda" rank="species" species="aeruginosa">
<emphasis id="CAAAEAC259686256FCCB21CD34A998CD" box="[854,1011,1736,1760]" italics="true" pageId="18" pageNumber="457">G. aeruginosa</emphasis>
</taxonomicName>
and 
<taxonomicName id="3FDE4D5359686256FBB621CD33F098CD" box="[1067,1194,1735,1760]" class="Insecta" family="Scarabaeidae" genus="Gromphas" kingdom="Animalia" order="Coleoptera" pageId="18" pageNumber="457" phylum="Arthropoda" rank="species" species="lemoinei">
<emphasis id="CAAAEAC259686256FBB621CD33F098CD" box="[1067,1194,1735,1760]" italics="true" pageId="18" pageNumber="457">G. lemoinei</emphasis>
</taxonomicName>
is in the shape of the pronotal prominence of well-developed specimens: each of the two projections of the prominence of 
<taxonomicName id="3FDE4D5359686256FB6221E832C69929" box="[1279,1436,1773,1796]" class="Insecta" family="Scarabaeidae" genus="Gromphas" kingdom="Animalia" order="Coleoptera" pageId="18" pageNumber="457" phylum="Arthropoda" rank="species" species="aeruginosa">
<emphasis id="CAAAEAC259686256FB6221E832C69929" box="[1279,1436,1773,1796]" italics="true" pageId="18" pageNumber="457">G. aeruginosa</emphasis>
</taxonomicName>
is clearly thin and tapered, while the projections are wide and triangular in 
<taxonomicName id="3FDE4D5359686256FC7A2015333C9905" box="[999,1126,1807,1832]" class="Insecta" family="Scarabaeidae" genus="Gromphas" kingdom="Animalia" order="Coleoptera" pageId="18" pageNumber="457" phylum="Arthropoda" rank="species" species="lemoinei">
<emphasis id="CAAAEAC259686256FC7A2015333C9905" box="[999,1126,1807,1832]" italics="true" pageId="18" pageNumber="457">G. lemoinei</emphasis>
</taxonomicName>
. In general, the projections are strongly divergent in 
<taxonomicName id="3FDE4D5359686256FE26203035019961" box="[443,603,1845,1868]" class="Insecta" family="Scarabaeidae" genus="Gromphas" kingdom="Animalia" order="Coleoptera" pageId="18" pageNumber="457" phylum="Arthropoda" rank="species" species="aeruginosa">
<emphasis id="CAAAEAC259686256FE26203035019961" box="[443,603,1845,1868]" italics="true" pageId="18" pageNumber="457">G. aeruginosa</emphasis>
</taxonomicName>
, and always subparallel in 
<taxonomicName id="3FDE4D5359686256FC07203033469960" box="[922,1052,1844,1869]" class="Insecta" family="Scarabaeidae" genus="Gromphas" kingdom="Animalia" order="Coleoptera" pageId="18" pageNumber="457" phylum="Arthropoda" rank="species" species="lemoinei">
<emphasis id="CAAAEAC259686256FC07203033469960" box="[922,1052,1844,1869]" italics="true" pageId="18" pageNumber="457">G. lemoinei</emphasis>
</taxonomicName>
; hence the lateral margins of the prominence are usually curved in 
<taxonomicName id="3FDE4D5359686256FD88205D35E9995D" box="[533,691,1880,1904]" class="Insecta" family="Scarabaeidae" genus="Gromphas" kingdom="Animalia" order="Coleoptera" pageId="18" pageNumber="457" phylum="Arthropoda" rank="species" species="aeruginosa">
<emphasis id="CAAAEAC259686256FD88205D35E9995D" box="[533,691,1880,1904]" italics="true" pageId="18" pageNumber="457">G. aeruginosa</emphasis>
</taxonomicName>
(Fig. 1), while in 
<taxonomicName id="3FDE4D5359686256FC1D205D34A5995D" box="[896,1023,1879,1904]" class="Insecta" family="Scarabaeidae" genus="Gromphas" kingdom="Animalia" order="Coleoptera" pageId="18" pageNumber="457" phylum="Arthropoda" rank="species" species="lemoinei">
<emphasis id="CAAAEAC259686256FC1D205D34A5995D" box="[896,1023,1879,1904]" italics="true" pageId="18" pageNumber="457">G. lemoinei</emphasis>
</taxonomicName>
they are always straight (
<figureCitation id="60E52A5559686256FABF205232D5995D" box="[1314,1423,1879,1904]" captionStart="FIGURES 7 – 10" captionStartId="16.[151,264,1939,1961]" captionTargetBox="[158,1436,199,1910]" captionTargetId="figure@16.[151,1436,193,1918]" captionTargetPageId="16" captionText="FIGURES 7 – 10. Gromphas lemoinei Waterhouse, 1891. 7) Major specimen. 8 – 10) Series of intermediate and minor specimens. Scale = 5 mm." httpUri="https://zenodo.org/record/247629/files/figure.png" pageId="18" pageNumber="457">Figs. 7–8</figureCitation>
). The space between the apices of the projections in 
<taxonomicName id="3FDE4D5359686256FD792078343F99B8" box="[740,869,1916,1941]" class="Insecta" family="Scarabaeidae" genus="Gromphas" kingdom="Animalia" order="Coleoptera" pageId="18" pageNumber="457" phylum="Arthropoda" rank="species" species="lemoinei">
<emphasis id="CAAAEAC259686256FD792078343F99B8" box="[740,869,1916,1941]" italics="true" pageId="18" pageNumber="457">G. lemoinei</emphasis>
</taxonomicName>
is much narrower than in 
<taxonomicName id="3FDE4D5359686256FB042078326299B9" box="[1177,1336,1917,1940]" class="Insecta" family="Scarabaeidae" genus="Gromphas" kingdom="Animalia" order="Coleoptera" pageId="18" pageNumber="457" phylum="Arthropoda" rank="species" species="aeruginosa">
<emphasis id="CAAAEAC259686256FB042078326299B9" box="[1177,1336,1917,1940]" italics="true" pageId="18" pageNumber="457">G. aeruginosa</emphasis>
</taxonomicName>
, and the total width of the pronotal prominence is proportionally greater in 
<taxonomicName id="3FDE4D5359686256FC1F20A433459995" box="[898,1055,1953,1976]" class="Insecta" family="Scarabaeidae" genus="Gromphas" kingdom="Animalia" order="Coleoptera" pageId="18" pageNumber="457" phylum="Arthropoda" rank="species" species="aeruginosa">
<emphasis id="CAAAEAC259686256FC1F20A433459995" box="[898,1055,1953,1976]" italics="true" pageId="18" pageNumber="457">G. aeruginosa</emphasis>
</taxonomicName>
than in 
<taxonomicName id="3FDE4D5359686256FBE720A433A39994" box="[1146,1273,1952,1977]" class="Insecta" family="Scarabaeidae" genus="Gromphas" kingdom="Animalia" order="Coleoptera" pageId="18" pageNumber="457" phylum="Arthropoda" rank="species" species="lemoinei">
<emphasis id="CAAAEAC259686256FBE720A433A39994" box="[1146,1273,1952,1977]" italics="true" pageId="18" pageNumber="457">G. lemoinei</emphasis>
</taxonomicName>
. For example, the largest specimen of 
<taxonomicName id="3FDE4D5359686256FE2820C0356399F0" box="[437,569,1988,2013]" class="Insecta" family="Scarabaeidae" genus="Gromphas" kingdom="Animalia" order="Coleoptera" pageId="18" pageNumber="457" phylum="Arthropoda" rank="species" species="lemoinei">
<emphasis id="CAAAEAC259686256FE2820C0356399F0" box="[437,569,1988,2013]" italics="true" pageId="18" pageNumber="457">G. lemoinei</emphasis>
</taxonomicName>
examined by us has a total length of 
<quantity id="3F269B3559686256FB9120C1332F99F1" box="[1036,1141,1988,2012]" metricMagnitude="-2" metricUnit="m" metricValue="2.01" pageId="18" pageNumber="457" unit="mm" value="20.1">20.1 mm</quantity>
and a width of pronotal prominence of 
<quantity id="3F269B3559696257FEDD279D36C39E82" box="[320,409,152,176]" metricMagnitude="-3" metricUnit="m" metricValue="3.8" pageId="19" pageNumber="458" unit="mm" value="3.8">3.8 mm</quantity>
; at the same time, the average width of the prominence in three specimens of 
<taxonomicName id="3FDE4D5359696257FB62279D32C69E9D" box="[1279,1436,152,176]" class="Insecta" family="Scarabaeidae" genus="Gromphas" kingdom="Animalia" order="Coleoptera" pageId="19" pageNumber="458" phylum="Arthropoda" rank="species" species="aeruginosa">
<emphasis id="CAAAEAC259696257FB62279D32C69E9D" box="[1279,1436,152,176]" italics="true" pageId="19" pageNumber="458">G. aeruginosa</emphasis>
</taxonomicName>
with total length between 
<quantity id="3F269B3559696257FE2527B935419EF9" box="[440,539,188,212]" metricMagnitude="-2" metricUnit="m" metricValue="1.98" pageId="19" pageNumber="458" unit="mm" value="19.8">19.8 mm</quantity>
and 
<quantity id="3F269B3559696257FDCD27B935EE9EF9" box="[592,692,188,212]" metricMagnitude="-2" metricUnit="m" metricValue="2.03" pageId="19" pageNumber="458" unit="mm" value="20.3">20.3 mm</quantity>
is 
<quantity id="3F269B3559696257FD4E27B934739EF9" box="[723,809,188,213]" metricMagnitude="-3" metricUnit="m" metricValue="4.6" pageId="19" pageNumber="458" unit="mm" value="4.6">4.6 mm</quantity>
(the total average in 
<taxonomicName id="3FDE4D5359696257FB8E27B833EA9EF9" box="[1043,1200,189,212]" class="Insecta" family="Scarabaeidae" genus="Gromphas" kingdom="Animalia" order="Coleoptera" pageId="19" pageNumber="458" phylum="Arthropoda" rank="species" species="aeruginosa">
<emphasis id="CAAAEAC259696257FB8E27B833EA9EF9" box="[1043,1200,189,212]" italics="true" pageId="19" pageNumber="458">G. aeruginosa</emphasis>
</taxonomicName>
is 
<quantity id="3F269B3559696257FB5227B932699EF9" box="[1231,1331,188,212]" metricMagnitude="-3" metricUnit="m" metricValue="2.83" pageId="19" pageNumber="458" unit="mm" value="2.83">2.83 mm</quantity>
and in 
<taxonomicName id="3FDE4D5359696257FA1927B837A29ED5" class="Insecta" family="Scarabaeidae" genus="Gromphas" kingdom="Animalia" order="Coleoptera" pageId="19" pageNumber="458" phylum="Arthropoda" rank="species" species="lemoinei">
<emphasis id="CAAAEAC259696257FA1927B837A29ED5" italics="true" pageId="19" pageNumber="458">G. lemoinei</emphasis>
</taxonomicName>
<quantity id="3F269B3559696257FF6327DA363C9ED5" box="[254,358,223,248]" metricMagnitude="-3" metricUnit="m" metricValue="2.54" pageId="19" pageNumber="458" unit="mm" value="2.54">2.54 mm</quantity>
). In a few specimens of 
<taxonomicName id="3FDE4D5359696257FDE527E5344F9ED5" box="[632,789,224,248]" class="Insecta" family="Scarabaeidae" genus="Gromphas" kingdom="Animalia" order="Coleoptera" pageId="19" pageNumber="458" phylum="Arthropoda" rank="species" species="aeruginosa">
<emphasis id="CAAAEAC259696257FDE527E5344F9ED5" box="[632,789,224,248]" italics="true" pageId="19" pageNumber="458">G. aeruginosa</emphasis>
</taxonomicName>
the lateral margins of the prominence are subparallel and not strongly divergent (Fig. 2); however, in such cases, the thin and tapered projections and especially the wide separation between the projections are clear characteristics of 
<taxonomicName id="3FDE4D5359696257FCD3262D34B09F6D" box="[846,1002,296,320]" class="Insecta" family="Scarabaeidae" genus="Gromphas" kingdom="Animalia" order="Coleoptera" pageId="19" pageNumber="458" phylum="Arthropoda" rank="species" species="aeruginosa">
<emphasis id="CAAAEAC259696257FCD3262D34B09F6D" box="[846,1002,296,320]" italics="true" pageId="19" pageNumber="458">G. aeruginosa</emphasis>
</taxonomicName>
.
</paragraph>
<paragraph id="F86136D059696257FF5A2649321F9C4D" blockId="19.[151,1437,151,2013]" pageId="19" pageNumber="458">
Characteristics of the tegument also separate these two species. The punctation in the center of metasternum and along the sutural margin of the elytra is very sparse in 
<taxonomicName id="3FDE4D5359696257FCB52675349E9FA5" box="[808,964,368,392]" class="Insecta" family="Scarabaeidae" genus="Gromphas" kingdom="Animalia" order="Coleoptera" pageId="19" pageNumber="458" phylum="Arthropoda" rank="species" species="aeruginosa">
<emphasis id="CAAAEAC259696257FCB52675349E9FA5" box="[808,964,368,392]" italics="true" pageId="19" pageNumber="458">G. aeruginosa</emphasis>
</taxonomicName>
and seen only in unwarn specimens; in 
<taxonomicName id="3FDE4D5359696257FA19267537AD9F80" class="Insecta" family="Scarabaeidae" genus="Gromphas" kingdom="Animalia" order="Coleoptera" pageId="19" pageNumber="458" phylum="Arthropoda" rank="species" species="lemoinei">
<emphasis id="CAAAEAC259696257FA19267537AD9F80" italics="true" pageId="19" pageNumber="458">G. lemoinei</emphasis>
</taxonomicName>
, however, this punctation is much denser and clearly evident. Regarding the pronotal granulation, in 
<taxonomicName id="3FDE4D5359696257FA192690364F9FFD" class="Insecta" family="Scarabaeidae" genus="Gromphas" kingdom="Animalia" order="Coleoptera" pageId="19" pageNumber="458" phylum="Arthropoda" rank="species" species="aeruginosa">
<emphasis id="CAAAEAC259696257FA192690364F9FFD" italics="true" pageId="19" pageNumber="458">G. aeruginosa</emphasis>
</taxonomicName>
, the squamous granules of the lateral region become progressively finer and sparser toward the center of the pronotum; in 
<taxonomicName id="3FDE4D5359696257FE1526D835519FD8" box="[392,523,476,501]" class="Insecta" family="Scarabaeidae" genus="Gromphas" kingdom="Animalia" order="Coleoptera" pageId="19" pageNumber="458" phylum="Arthropoda" rank="species" species="lemoinei">
<emphasis id="CAAAEAC259696257FE1526D835519FD8" box="[392,523,476,501]" italics="true" pageId="19" pageNumber="458">G. lemoinei</emphasis>
</taxonomicName>
, on the other hand, the density of granulation is always very high across the pronotum, except within the smooth anterior concavities and along the midline. The presence of posterior pronotal fossae also distinguish these two species: in 
<taxonomicName id="3FDE4D5359696257FD17252034539C10" box="[650,777,548,573]" class="Insecta" family="Scarabaeidae" genus="Gromphas" kingdom="Animalia" order="Coleoptera" pageId="19" pageNumber="458" phylum="Arthropoda" rank="species" species="lemoinei">
<emphasis id="CAAAEAC259696257FD17252034539C10" box="[650,777,548,573]" italics="true" pageId="19" pageNumber="458">G. lemoinei</emphasis>
</taxonomicName>
, larger specimens have no trace of these fossae, which are apparent in smaller specimens; in 
<taxonomicName id="3FDE4D5359696257FD89254D35EB9C4D" box="[532,689,584,608]" class="Insecta" family="Scarabaeidae" genus="Gromphas" kingdom="Animalia" order="Coleoptera" pageId="19" pageNumber="458" phylum="Arthropoda" rank="species" species="aeruginosa">
<emphasis id="CAAAEAC259696257FD89254D35EB9C4D" box="[532,689,584,608]" italics="true" pageId="19" pageNumber="458">G. aeruginosa</emphasis>
</taxonomicName>
, the fossae are always present regardless of specimen size.
</paragraph>
<paragraph id="F86136D059696257FF5A256936AC9CC2" blockId="19.[151,1437,151,2013]" pageId="19" pageNumber="458">
The elytral striae are also distinct: in 
<taxonomicName id="3FDE4D5359696257FDF1256834509CA9" box="[620,778,621,644]" class="Insecta" family="Scarabaeidae" genus="Gromphas" kingdom="Animalia" order="Coleoptera" pageId="19" pageNumber="458" phylum="Arthropoda" rank="species" species="aeruginosa">
<emphasis id="CAAAEAC259696257FDF1256834509CA9" box="[620,778,621,644]" italics="true" pageId="19" pageNumber="458">G. aeruginosa</emphasis>
</taxonomicName>
, the interstriae are flat and in the same plane as the striae, whereas in 
<taxonomicName id="3FDE4D5359696257FE88259536CF9C85" box="[277,405,655,680]" class="Insecta" family="Scarabaeidae" genus="Gromphas" kingdom="Animalia" order="Coleoptera" pageId="19" pageNumber="458" phylum="Arthropoda" rank="species" species="lemoinei">
<emphasis id="CAAAEAC259696257FE88259536CF9C85" box="[277,405,655,680]" italics="true" pageId="19" pageNumber="458">G. lemoinei</emphasis>
</taxonomicName>
the interstriae, especially the more mesial ones, are slightly convex at the base and the striae appear furrowed to the naked eye. This furrowed appearance gives the impression of a more elongate elytra in 
<taxonomicName id="3FDE4D5359696257FA1925B037A29CDD" class="Insecta" family="Scarabaeidae" genus="Gromphas" kingdom="Animalia" order="Coleoptera" pageId="19" pageNumber="458" phylum="Arthropoda" rank="species" species="lemoinei">
<emphasis id="CAAAEAC259696257FA1925B037A29CDD" italics="true" pageId="19" pageNumber="458">G. lemoinei</emphasis>
</taxonomicName>
than in 
<taxonomicName id="3FDE4D5359696257FECF25DD36B59CDD" box="[338,495,728,752]" class="Insecta" family="Scarabaeidae" genus="Gromphas" kingdom="Animalia" order="Coleoptera" pageId="19" pageNumber="458" phylum="Arthropoda" rank="species" species="aeruginosa">
<emphasis id="CAAAEAC259696257FECF25DD36B59CDD" box="[338,495,728,752]" italics="true" pageId="19" pageNumber="458">G. aeruginosa</emphasis>
</taxonomicName>
.
</paragraph>
<paragraph id="F86136D059696257FF5A25F832339DE5" blockId="19.[151,1437,151,2013]" pageId="19" pageNumber="458">
More developed specimens of 
<taxonomicName id="3FDE4D5359696257FDB725F835909D39" box="[554,714,765,788]" class="Insecta" family="Scarabaeidae" genus="Gromphas" kingdom="Animalia" order="Coleoptera" pageId="19" pageNumber="458" phylum="Arthropoda" rank="species" species="aeruginosa">
<emphasis id="CAAAEAC259696257FDB725F835909D39" box="[554,714,765,788]" italics="true" pageId="19" pageNumber="458">G. aeruginosa</emphasis>
</taxonomicName>
have the cephalic projection distinctly notched and bifid (
<figureCitation id="60E52A5559696257FAF225F937EE9D15" captionStart="FIGURES 29 – 33" captionStartId="30.[151,264,1916,1938]" captionTargetBox="[195,1380,179,1890]" captionTargetId="figure@30.[195,1380,179,1891]" captionTargetPageId="30" captionText="FIGURES 29 – 33. Frontal view of cephalic projections. 29: Gromphas aeruginosa. 29 a) Major specimen with cephalic projection extremely broad. 29 b) Major specimen with apical notch. 29 c) Worn major specimen. 29 d) Medium specimen. 29 e) Minor specimen. 30: G. lemoinei. 30 a) Major specimen. 30 b) Medium specimen. 30 c) Minor specimen. 31) G. amazonica. 32) G. inermis. 33) G. dichroa." httpUri="https://zenodo.org/record/247634/files/figure.png" pageId="19" pageNumber="458">Fig. 29</figureCitation>
b) (round or truncate in worn specimens [
<figureCitation id="60E52A5559696257FD38241A345A9D15" box="[677,768,799,824]" captionStart="FIGURES 29 – 33" captionStartId="30.[151,264,1916,1938]" captionTargetBox="[195,1380,179,1890]" captionTargetId="figure@30.[195,1380,179,1891]" captionTargetPageId="30" captionText="FIGURES 29 – 33. Frontal view of cephalic projections. 29: Gromphas aeruginosa. 29 a) Major specimen with cephalic projection extremely broad. 29 b) Major specimen with apical notch. 29 c) Worn major specimen. 29 d) Medium specimen. 29 e) Minor specimen. 30: G. lemoinei. 30 a) Major specimen. 30 b) Medium specimen. 30 c) Minor specimen. 31) G. amazonica. 32) G. inermis. 33) G. dichroa." httpUri="https://zenodo.org/record/247634/files/figure.png" pageId="19" pageNumber="458">Fig. 29</figureCitation>
c]). In large 
<taxonomicName id="3FDE4D5359696257FC09242533429D15" box="[916,1048,799,824]" class="Insecta" family="Scarabaeidae" genus="Gromphas" kingdom="Animalia" order="Coleoptera" pageId="19" pageNumber="458" phylum="Arthropoda" rank="species" species="lemoinei">
<emphasis id="CAAAEAC259696257FC09242533429D15" box="[916,1048,799,824]" italics="true" pageId="19" pageNumber="458">G. lemoinei</emphasis>
</taxonomicName>
, on the other hand, the cephalic projection is always acuminate (
<figureCitation id="60E52A5559696257FD9C2441350C9D71" box="[513,598,836,861]" captionStart="FIGURES 29 – 33" captionStartId="30.[151,264,1916,1938]" captionTargetBox="[195,1380,179,1890]" captionTargetId="figure@30.[195,1380,179,1891]" captionTargetPageId="30" captionText="FIGURES 29 – 33. Frontal view of cephalic projections. 29: Gromphas aeruginosa. 29 a) Major specimen with cephalic projection extremely broad. 29 b) Major specimen with apical notch. 29 c) Worn major specimen. 29 d) Medium specimen. 29 e) Minor specimen. 30: G. lemoinei. 30 a) Major specimen. 30 b) Medium specimen. 30 c) Minor specimen. 31) G. amazonica. 32) G. inermis. 33) G. dichroa." httpUri="https://zenodo.org/record/247634/files/figure.png" pageId="19" pageNumber="458">Fig. 30</figureCitation>
). While one could argue that the lack of notch in specimens of 
<taxonomicName id="3FDE4D5359696257FA81244032C69D70" box="[1308,1436,836,861]" class="Insecta" family="Scarabaeidae" genus="Gromphas" kingdom="Animalia" order="Coleoptera" pageId="19" pageNumber="458" phylum="Arthropoda" rank="species" species="lemoinei">
<emphasis id="CAAAEAC259696257FA81244032C69D70" box="[1308,1436,836,861]" italics="true" pageId="19" pageNumber="458">G. lemoinei</emphasis>
</taxonomicName>
has resulted from wear, even the best preserved specimens of 
<taxonomicName id="3FDE4D5359696257FCD1246D34919DAD" box="[844,971,871,896]" class="Insecta" family="Scarabaeidae" genus="Gromphas" kingdom="Animalia" order="Coleoptera" pageId="19" pageNumber="458" phylum="Arthropoda" rank="species" species="lemoinei">
<emphasis id="CAAAEAC259696257FCD1246D34919DAD" box="[844,971,871,896]" italics="true" pageId="19" pageNumber="458">G. lemoinei</emphasis>
</taxonomicName>
have no trace of an incision and no worn 
<taxonomicName id="3FDE4D5359696257FF0A2488366F9D89" box="[151,309,909,932]" class="Insecta" family="Scarabaeidae" genus="Gromphas" kingdom="Animalia" order="Coleoptera" pageId="19" pageNumber="458" phylum="Arthropoda" rank="species" species="aeruginosa">
<emphasis id="CAAAEAC259696257FF0A2488366F9D89" box="[151,309,909,932]" italics="true" pageId="19" pageNumber="458">G. aeruginosa</emphasis>
</taxonomicName>
has the projection acuminate, thus ruling out this possibility. A very large Bolivian specimen of 
<taxonomicName id="3FDE4D5359696257FA192488364F9DE5" class="Insecta" family="Scarabaeidae" genus="Gromphas" kingdom="Animalia" order="Coleoptera" pageId="19" pageNumber="458" phylum="Arthropoda" rank="species" species="aeruginosa">
<emphasis id="CAAAEAC259696257FA192488364F9DE5" italics="true" pageId="19" pageNumber="458">G. aeruginosa</emphasis>
</taxonomicName>
shows the cephalic projection extremely broad in a way not seen in any other specimen (
<figureCitation id="60E52A5559696257FB6124AA32159DE5" box="[1276,1359,943,968]" captionStart="FIGURES 29 – 33" captionStartId="30.[151,264,1916,1938]" captionTargetBox="[195,1380,179,1890]" captionTargetId="figure@30.[195,1380,179,1891]" captionTargetPageId="30" captionText="FIGURES 29 – 33. Frontal view of cephalic projections. 29: Gromphas aeruginosa. 29 a) Major specimen with cephalic projection extremely broad. 29 b) Major specimen with apical notch. 29 c) Worn major specimen. 29 d) Medium specimen. 29 e) Minor specimen. 30: G. lemoinei. 30 a) Major specimen. 30 b) Medium specimen. 30 c) Minor specimen. 31) G. amazonica. 32) G. inermis. 33) G. dichroa." httpUri="https://zenodo.org/record/247634/files/figure.png" pageId="19" pageNumber="458">Fig. 29</figureCitation>
a).
</paragraph>
<paragraph id="F86136D059696257FF5A24D136D39A7A" blockId="19.[151,1437,151,2013]" pageId="19" pageNumber="458">
Finally, there is a difference between these two species also in the medial sclerite. While in 
<taxonomicName id="3FDE4D5359696257FB4924D032289DC1" box="[1236,1394,981,1004]" class="Insecta" family="Scarabaeidae" genus="Gromphas" kingdom="Animalia" order="Coleoptera" pageId="19" pageNumber="458" phylum="Arthropoda" rank="species" species="aeruginosa">
<emphasis id="CAAAEAC259696257FB4924D032289DC1" box="[1236,1394,981,1004]" italics="true" pageId="19" pageNumber="458">G. aeruginosa</emphasis>
</taxonomicName>
the curvature is mainly downwards (
<figureCitation id="60E52A5559696257FD9424F235069A3D" box="[521,604,1015,1040]" captionStart="FIGURES 57 – 62" captionStartId="34.[151,264,1134,1156]" captionTargetBox="[433,1142,199,1103]" captionTargetId="figure@34.[428,1160,193,1113]" captionTargetPageId="34" captionText="FIGURES 57 – 62. Schematic drawings. 57 – 61: Medial sclerite of internal sac. 57) Gromphas aeruginosa. 58) G. lemoinei. 59) G. inermis. 60) G. amazonica. 61) G. dichroa. 62) Internal sac of G. aeruginosa (TE = temones; MS = medial sclerite; AS = apical sclerites)." httpUri="https://zenodo.org/record/247638/files/figure.png" pageId="19" pageNumber="458">Fig. 57</figureCitation>
), in 
<taxonomicName id="3FDE4D5359696257FD1224FD34549A3D" box="[655,782,1015,1040]" class="Insecta" family="Scarabaeidae" genus="Gromphas" kingdom="Animalia" order="Coleoptera" pageId="19" pageNumber="458" phylum="Arthropoda" rank="species" species="lemoinei">
<emphasis id="CAAAEAC259696257FD1224FD34549A3D" box="[655,782,1015,1040]" italics="true" pageId="19" pageNumber="458">G. lemoinei</emphasis>
</taxonomicName>
this sclerite is flatter, wider and very much more strongly curved to the side (
<figureCitation id="60E52A5559696257FEE5231936949A18" box="[376,462,1052,1077]" captionStart="FIGURES 57 – 62" captionStartId="34.[151,264,1134,1156]" captionTargetBox="[433,1142,199,1103]" captionTargetId="figure@34.[428,1160,193,1113]" captionTargetPageId="34" captionText="FIGURES 57 – 62. Schematic drawings. 57 – 61: Medial sclerite of internal sac. 57) Gromphas aeruginosa. 58) G. lemoinei. 59) G. inermis. 60) G. amazonica. 61) G. dichroa. 62) Internal sac of G. aeruginosa (TE = temones; MS = medial sclerite; AS = apical sclerites)." httpUri="https://zenodo.org/record/247638/files/figure.png" pageId="19" pageNumber="458">Fig. 58</figureCitation>
). However, these two forms are clearly more similar to each other than to the other species of 
<taxonomicName id="3FDE4D5359696257FE90233A36D89A75" box="[269,386,1087,1112]" class="Insecta" family="Scarabaeidae" genus="Gromphas" kingdom="Animalia" order="Coleoptera" pageId="19" pageNumber="458" phylum="Arthropoda" rank="genus">
<emphasis id="CAAAEAC259696257FE90233A36D89A75" box="[269,386,1087,1112]" italics="true" pageId="19" pageNumber="458">Gromphas</emphasis>
</taxonomicName>
.
</paragraph>
</subSubSection>
<subSubSection id="B0C4655B59696257FF5A236136BC9905" pageId="19" pageNumber="458" type="discussion">
<paragraph id="F86136D059696257FF5A236136609ACA" blockId="19.[151,1437,151,2013]" pageId="19" pageNumber="458">
<emphasis id="CAAAEAC259696257FF5A2361360B9A50" bold="true" box="[199,337,1124,1149]" pageId="19" pageNumber="458">Comments:</emphasis>
The presence of 
<taxonomicName id="3FDE4D5359696257FDAE236035E09A50" box="[563,698,1124,1149]" class="Insecta" family="Scarabaeidae" genus="Gromphas" kingdom="Animalia" order="Coleoptera" pageId="19" pageNumber="458" phylum="Arthropoda" rank="species" species="lemoinei">
<emphasis id="CAAAEAC259696257FDAE236035E09A50" box="[563,698,1124,1149]" italics="true" pageId="19" pageNumber="458">G. lemoinei</emphasis>
</taxonomicName>
in San Fernando de Apure is inferred from the geographic distribution of 
<taxonomicName id="3FDE4D5359696257FEDA238D36B29A8D" box="[327,488,1160,1184]" class="Insecta" family="Scarabaeidae" genus="Gromphas" kingdom="Animalia" order="Coleoptera" pageId="19" pageNumber="458" phylum="Arthropoda" rank="species" species="aeruginosa">
<emphasis id="CAAAEAC259696257FEDA238D36B29A8D" box="[327,488,1160,1184]" italics="true" pageId="19" pageNumber="458">G. aeruginosa</emphasis>
</taxonomicName>
presented by Barattini &amp; Saénz (1960, 1964). These authors regarded the latter species (which does not occur in 
<collectingCountry id="80C9764059696257FD9123A935DB9AE8" box="[524,641,1196,1221]" name="Venezuela" pageId="19" pageNumber="458">Venezuela</collectingCountry>
) as a senior synonym of the former, but it is likely that these data refer to 
<taxonomicName id="3FDE4D5359696257FF2923D536689AC5" box="[180,306,1231,1256]" class="Insecta" family="Scarabaeidae" genus="Gromphas" kingdom="Animalia" order="Coleoptera" pageId="19" pageNumber="458" phylum="Arthropoda" rank="species" species="lemoinei">
<emphasis id="CAAAEAC259696257FF2923D536689AC5" box="[180,306,1231,1256]" italics="true" pageId="19" pageNumber="458">G. lemoinei</emphasis>
</taxonomicName>
.
</paragraph>
<paragraph id="F86136D059696257FF5A23F136269800" blockId="19.[151,1437,151,2013]" pageId="19" pageNumber="458">
<emphasis id="CAAAEAC259696257FF5A23F136179B20" bold="true" box="[199,333,1268,1293]" pageId="19" pageNumber="458">Bionomics:</emphasis>
Noriega (2002) found five specimens of 
<taxonomicName id="3FDE4D5359696257FCBD23F034C59B20" box="[800,927,1268,1293]" class="Insecta" family="Scarabaeidae" genus="Gromphas" kingdom="Animalia" order="Coleoptera" pageId="19" pageNumber="458" phylum="Arthropoda" rank="species" species="lemoinei">
<emphasis id="CAAAEAC259696257FCBD23F034C59B20" box="[800,927,1268,1293]" italics="true" pageId="19" pageNumber="458">G. lemoinei</emphasis>
</taxonomicName>
(cited as 
<taxonomicName id="3FDE4D5359696257FB9323F033F19B21" box="[1038,1195,1269,1292]" class="Insecta" family="Scarabaeidae" genus="Gromphas" kingdom="Animalia" order="Coleoptera" pageId="19" pageNumber="458" phylum="Arthropoda" rank="species" species="aeruginosa">
<emphasis id="CAAAEAC259696257FB9323F033F19B21" box="[1038,1195,1269,1292]" italics="true" pageId="19" pageNumber="458">G. aeruginosa</emphasis>
</taxonomicName>
) on the floodplain of Rio Duda, La Macarena, Meta, 
<collectingCountry id="80C9764059696257FD9C221235299B1D" box="[513,627,1303,1328]" name="Colombia" pageId="19" pageNumber="458">Colombia</collectingCountry>
; the only other species of 
<taxonomicName id="3FDE4D5359696257FC02221233689B1D" box="[927,1074,1303,1328]" class="Insecta" family="Scarabaeidae" kingdom="Animalia" order="Coleoptera" pageId="19" pageNumber="458" phylum="Arthropoda" rank="subFamily" subFamily="Scarabaeinae">Scarabaeinae</taxonomicName>
found by him in this place was 
<taxonomicName id="3FDE4D5359696257FF0A223935D99B79" authority="Waterhouse, 1891" authorityName="Waterhouse" authorityYear="1891" box="[151,643,1340,1365]" class="Insecta" family="Scarabaeidae" genus="Sulcophanaeus" kingdom="Animalia" order="Coleoptera" pageId="19" pageNumber="458" phylum="Arthropoda" rank="species" species="leander">
<emphasis id="CAAAEAC259696257FF0A223936C69B78" box="[151,412,1340,1365]" italics="true" pageId="19" pageNumber="458">Sulcophanaeus leander</emphasis>
(Waterhouse, 1891)
</taxonomicName>
. According Noriega (personal communication) these two species share the same activity time and the same food resource. As 
<taxonomicName id="3FDE4D5359696257FC9F226534DB9B55" box="[770,897,1375,1400]" class="Insecta" family="Scarabaeidae" genus="Gromphas" kingdom="Animalia" order="Coleoptera" pageId="19" pageNumber="458" phylum="Arthropoda" rank="species" species="lemoinei">
<emphasis id="CAAAEAC259696257FC9F226534DB9B55" box="[770,897,1375,1400]" italics="true" pageId="19" pageNumber="458">G. lemoinei</emphasis>
</taxonomicName>
is found only in savannas, open areas and river sandbanks and never enter in the forest, it is possible that this species use the rivers and their banks as corridors from the savannas of the Llanos Orientales to inside Amazon region in 
<collectingCountry id="80C9764059696257FBA222A233F79BED" box="[1087,1197,1447,1472]" name="Colombia" pageId="19" pageNumber="458">Colombia</collectingCountry>
(Noriega, personal communication). Noriega also reported that all specimens collected by him were associated with cattle (
<taxonomicName id="3FDE4D5359696257FA8622C836129825" authority="Linnaeus, 1758" authorityName="Linnaeus" authorityYear="1758" class="Mammalia" family="Bovidae" genus="Bos" kingdom="Animalia" order="Artiodactyla" pageId="19" pageNumber="458" phylum="Chordata" rank="species" species="indicus">
<emphasis id="CAAAEAC259696257FA8622C832C69BC8" box="[1307,1436,1484,1509]" italics="true" pageId="19" pageNumber="458">Bos indicus</emphasis>
Linnaeus, 1758
</taxonomicName>
), capybara (
<taxonomicName id="3FDE4D5359696257FE4B22EA34BD9825" authority="Linnaeus, 1766" authorityName="Linnaeus" authorityYear="1766" box="[470,999,1519,1544]" class="Mammalia" family="Caviidae" genus="Hydrochoerus" kingdom="Animalia" order="Rodentia" pageId="19" pageNumber="458" phylum="Chordata" rank="species" species="hydrochaeris">
<emphasis id="CAAAEAC259696257FE4B22EA344B9825" box="[470,785,1519,1544]" italics="true" pageId="19" pageNumber="458">Hydrochoerus hydrochaeris</emphasis>
(Linnaeus, 1766))
</taxonomicName>
, or crocodile (
<taxonomicName id="3FDE4D5359696257FB1222EA36779801" authority="Graves, 1819" authorityName="Graves" authorityYear="1819" class="Reptilia" family="Crocodylidae" genus="Crocodylus" kingdom="Animalia" order="Crocodylia" pageId="19" pageNumber="458" phylum="Chordata" rank="species" species="intermedius">
<emphasis id="CAAAEAC259696257FB1222EA32C69825" box="[1167,1436,1519,1544]" italics="true" pageId="19" pageNumber="458">Crocodylus intermedius</emphasis>
Graves, 1819
</taxonomicName>
) dung.
</paragraph>
<paragraph id="F86136D059696257FF5A213232729858" blockId="19.[151,1437,151,2013]" pageId="19" pageNumber="458">
Based on the specimens examined in this study, 
<taxonomicName id="3FDE4D5359696257FD69213D342E987D" box="[756,884,1591,1616]" class="Insecta" family="Scarabaeidae" genus="Gromphas" kingdom="Animalia" order="Coleoptera" pageId="19" pageNumber="458" phylum="Arthropoda" rank="species" species="lemoinei">
<emphasis id="CAAAEAC259696257FD69213D342E987D" box="[756,884,1591,1616]" italics="true" pageId="19" pageNumber="458">G. lemoinei</emphasis>
</taxonomicName>
seems to be active throughout the year, as is 
<taxonomicName id="3FDE4D5359696257FA19213D364F9859" class="Insecta" family="Scarabaeidae" genus="Gromphas" kingdom="Animalia" order="Coleoptera" pageId="19" pageNumber="458" phylum="Arthropoda" rank="species" species="aeruginosa">
<emphasis id="CAAAEAC259696257FA19213D364F9859" italics="true" pageId="19" pageNumber="458">G. aeruginosa</emphasis>
</taxonomicName>
. The recorded months are January, March, June, July, September, November, and December.
</paragraph>
<paragraph id="F86136D059696257FF5A217A36BC9905" blockId="19.[151,1437,151,2013]" pageId="19" pageNumber="458">
<emphasis id="CAAAEAC259696257FF5A217A355298B5" bold="true" box="[199,520,1663,1688]" pageId="19" pageNumber="458">Geographical distribution:</emphasis>
Caribbean subregion: Venezuelan Coast and Venezuelan Llanos. 
<emphasis id="CAAAEAC259696257FB69217A32C698B5" bold="true" box="[1268,1436,1663,1688]" pageId="19" pageNumber="458">
<collectingCountry id="80C9764059696257FB69217A32CD98B5" box="[1268,1431,1663,1688]" name="Colombia" pageId="19" pageNumber="458">COLOMBIA</collectingCountry>
:
</emphasis>
Arauca: Arauca. Meta: La Macarena, Puerto Gaitan, San Martín, Villavicencio. Guaviare: San José del Guaviare. 
<emphasis id="CAAAEAC259696257FF0A21C2361598CD" bold="true" box="[151,335,1735,1760]" pageId="19" pageNumber="458">
<collectingCountry id="80C9764059696257FF0A21C2361098CD" box="[151,330,1735,1760]" name="Venezuela" pageId="19" pageNumber="458">VENEZUELA</collectingCountry>
:
</emphasis>
Distrito Capital: Caracas. Barinas: Andrés Eloy Blanco (El Cantón), Ezequiel Zamora (San Antonio de Pagen). Táchira: Libertador (San Joaquín de Navay), 
<collectingCountry id="80C9764059696257FC3D21E9331A9928" box="[928,1088,1772,1797]" name="Saint Kitts and Nevis" pageId="19" pageNumber="458">San Cristóbal</collectingCountry>
, Santo 
<collectingCountry id="80C9764059696257FB0621E932539928" box="[1179,1289,1772,1797]" name="Dominican Republic" pageId="19" pageNumber="458">Domingo</collectingCountry>
. Apure: San Fernando de Apure. (
<figureCitation id="60E52A5559696257FE18200A36809905" box="[389,474,1807,1832]" captionStart="FIGURE 66" captionStartId="35.[151,250,1617,1639]" captionTargetBox="[151,1436,183,1595]" captionTargetId="figure@35.[151,1436,181,1595]" captionTargetPageId="35" captionText="FIGURE 66. Distribution map of the five species of Gromphas." httpUri="https://zenodo.org/record/247640/files/figure.png" pageId="19" pageNumber="458">Fig. 66</figureCitation>
).
</paragraph>
</subSubSection>
<subSubSection id="B0C4655B59696250FF5A2031323A9FA5" lastPageId="20" lastPageNumber="459" pageId="19" pageNumber="458" type="materials_examined">
<paragraph id="F86136D059696250FF5A2031323A9FA5" blockId="19.[151,1437,151,2013]" lastBlockId="20.[151,1437,151,392]" lastPageId="20" lastPageNumber="459" pageId="19" pageNumber="458">
<emphasis id="CAAAEAC259696257FF5A203136E89960" bold="true" box="[199,434,1844,1869]" pageId="19" pageNumber="458">Material examined:</emphasis>
<specimenCount id="EED8FD5959696257FE24203135449960" box="[441,542,1844,1869]" pageId="19" pageNumber="458" type="male">17 males</specimenCount>
and 
<specimenCount id="EED8FD5959696257FDC92031358B9960" box="[596,721,1844,1869]" pageId="19" pageNumber="458" type="female">10 females</specimenCount>
. 
<emphasis id="CAAAEAC259696257FD46203134D99960" bold="true" box="[731,899,1844,1869]" pageId="19" pageNumber="458">
<collectingCountry id="80C9764059696257FD46203134249960" box="[731,894,1844,1869]" name="Colombia" pageId="19" pageNumber="458">COLOMBIA</collectingCountry>
:
</emphasis>
ARAUCA: Arauca, 
<specimenCount id="EED8FD5959696257FBF3203133E79961" box="[1134,1213,1844,1868]" pageId="19" pageNumber="458" type="generic">
<quantity id="3F269B3559696257FBF3203133E39961" box="[1134,1209,1844,1868]" metricMagnitude="2" metricUnit="m" metricValue="1.2" pageId="19" pageNumber="458" unit="m" value="120.0">120 m</quantity>
.
</specimenCount>
, 
<date id="8C60101059696257FB54203132739960" box="[1225,1321,1844,1869]" pageId="19" pageNumber="458" value="2005-09">IX.2005</date>
, Jorge Ari Noriega col.— 
<specimenCount id="EED8FD5959696257FED8205236D4995D" box="[325,398,1879,1904]" pageId="19" pageNumber="458" type="male">1 male</specimenCount>
(CJAN). GUAVIARE: San Jose del Guaviare, sabana, 
<date id="8C60101059696257FB8D205D3335995D" box="[1040,1135,1880,1904]" pageId="19" pageNumber="458" value="2000-09">IX.2000</date>
, J. Noriega col.— 
<specimenCount id="EED8FD5959696257FADE205232C6995D" box="[1347,1436,1879,1904]" pageId="19" pageNumber="458" type="male">2 males</specimenCount>
(CJAN). META: La Macarena, CIEM, Playa del Rio Duda, 
<quantity id="3F269B3559696257FCD1207934CD99B9" box="[844,919,1916,1941]" metricMagnitude="2" metricUnit="m" metricValue="3.5" pageId="19" pageNumber="458" unit="m" value="350.0">350 m</quantity>
, 
<date id="8C60101059696257FC382079335799B8" box="[933,1037,1916,1941]" pageId="19" pageNumber="458" value="1995-12">XII.1995</date>
, Jorge A. Noriega col.— 
<specimenCount id="EED8FD5959696257FAB3207932C699B8" box="[1326,1436,1916,1941]" pageId="19" pageNumber="458" type="female">2 females</specimenCount>
(CJAN); La Macarena, CIEM, 
<date id="8C60101059696257FE6920A535659995" box="[500,575,1952,1976]" pageId="19" pageNumber="458" value="1998-01">I.1998</date>
, J. Noriega col.— 
<specimenCount id="EED8FD5959696257FC9220A5340C9994" box="[783,854,1952,1977]" pageId="19" pageNumber="458" type="male">1 male</specimenCount>
(CJAN); Puerto Gaitan, 
<specimenCount id="EED8FD5959696257FBF320A533E49995" box="[1134,1214,1952,1977]" pageId="19" pageNumber="458" type="generic">
<quantity id="3F269B3559696257FBF320A533E09995" box="[1134,1210,1952,1977]" metricMagnitude="2" metricUnit="m" metricValue="1.5" pageId="19" pageNumber="458" unit="m" value="150.0">150 m</quantity>
.
</specimenCount>
, 
<date id="8C60101059696257FB4F20A532729995" box="[1234,1320,1952,1976]" pageId="19" pageNumber="458" value="2001-10">X.2001</date>
, Jorge Ari Noriega col.— 
<specimenCount id="EED8FD5959696257FEDB20C136FD99F0" box="[326,423,1988,2013]" pageId="19" pageNumber="458" type="female">1 female</specimenCount>
(CJAN); San Martín, 
<specimenCount id="EED8FD5959696257FD2F20C135A699F0" box="[690,764,1988,2013]" pageId="19" pageNumber="458" type="generic">
<quantity id="3F269B3559696257FD2F20C135A299F0" box="[690,760,1988,2013]" metricMagnitude="2" metricUnit="m" metricValue="3.5" pageId="19" pageNumber="458" unit="m" value="350.0">350m</quantity>
.
</specimenCount>
, 
<date id="8C60101059696257FC9120C1342F99F1" box="[780,885,1988,2012]" pageId="19" pageNumber="458" value="2007-07">VII.2007</date>
, Jorge Ari Noriega col.— 
<specimenCount id="EED8FD5959696257FB3120C133AD99F0" box="[1196,1271,1988,2013]" pageId="19" pageNumber="458" type="male">1 male</specimenCount>
and 
<specimenCount id="EED8FD5959696257FAAB20C132C699F0" box="[1334,1436,1988,2013]" pageId="19" pageNumber="458" type="female">1 female</specimenCount>
(CJAN); Villavicencio, 
<date id="8C601010596E6250FE39279D35759E9D" box="[420,559,152,176]" pageId="20" pageNumber="459" value="1994-12-13">13.XII.1994</date>
, Oscar Gómez col.— 
<specimenCount id="EED8FD59596E6250FCB7279234DD9E9D" box="[810,903,151,176]" pageId="20" pageNumber="459" type="female">1 female</specimenCount>
(CEMT). 
<emphasis id="CAAAEAC2596E6250FB9D279233ED9E9D" bold="true" box="[1024,1207,151,176]" pageId="20" pageNumber="459">
<collectingCountry id="80C97640596E6250FB9D279233E89E9D" box="[1024,1202,151,176]" name="Venezuela" pageId="20" pageNumber="459">VENEZUELA</collectingCountry>
:
</emphasis>
BARINAS: Andrés Eloy Blanco, El Cantón, 
<date id="8C601010596E6250FE2227B935449EF9" box="[447,542,188,212]" pageId="20" pageNumber="459" value="1983-11">XI.1983</date>
, C. Borges col.— 
<specimenCount id="EED8FD59596E6250FD6B27B9341B9EF8" box="[758,833,188,213]" pageId="20" pageNumber="459" type="male">1 male</specimenCount>
(MIZA); Ezequiel Zamora, San Antonio de Pagen, 
<date id="8C601010596E6250FF0A27E537A59ED5" box="[151,255,224,248]" pageId="20" pageNumber="459" value="1989-07">VII.1989</date>
, without collector— 
<specimenCount id="EED8FD59596E6250FE7327DA35109ED5" box="[494,586,223,248]" pageId="20" pageNumber="459" type="female">1 female</specimenCount>
(MZSP) and 
<specimenCount id="EED8FD59596E6250FD7527E534659ED5" box="[744,831,223,248]" pageId="20" pageNumber="459" type="male">2 males</specimenCount>
(CEMT); San Antonio, 
<date id="8C601010596E6250FBCB27E533E59ED5" box="[1110,1215,224,248]" pageId="20" pageNumber="459" value="2007-07">VII.2007</date>
, J. Noriega col.— 
<specimenCount id="EED8FD59596E6250FA0E27DA37979F30" pageId="20" pageNumber="459" type="male">1 male</specimenCount>
and 
<specimenCount id="EED8FD59596E6250FE8C260136229F30" box="[273,376,260,285]" pageId="20" pageNumber="459" type="female">1 female</specimenCount>
(CJAN). DISTRITO CAPITAL: Caracas, 1905, Fry col.— 
<specimenCount id="EED8FD59596E6250FBDF260133D49F30" box="[1090,1166,260,285]" pageId="20" pageNumber="459" type="male">1 male</specimenCount>
(BMNH). TÁCHIRA: Libertador, San Joaquín de Navay, 
<quantity id="3F269B35596E6250FD82262D35329F12" box="[543,616,296,320]" metricMagnitude="2" metricUnit="m" metricValue="1.0" pageId="20" pageNumber="459" unit="m" value="100.0">100 m</quantity>
, 
<date id="8C601010596E6250FDE9262D35AE9F6D" box="[628,756,296,320]" pageId="20" pageNumber="459" value="1989-03-24">24.III.1989</date>
, D. Havranek col.— 
<specimenCount id="EED8FD59596E6250FC422622337C9F6D" box="[991,1062,295,320]" pageId="20" pageNumber="459" type="male">1 male</specimenCount>
(CMNC); 
<collectingCountry id="80C97640596E6250FB3C262D32669F6D" box="[1185,1340,295,320]" name="Saint Kitts and Nevis" pageId="20" pageNumber="459">San Cristóbal</collectingCountry>
, without date and collector— 
<specimenCount id="EED8FD59596E6250FE16264936879F48" box="[395,477,332,357]" pageId="20" pageNumber="459" type="male">2 male</specimenCount>
(MIZA); 
<collectingCountry id="80C97640596E6250FDC02649345A9F48" box="[605,768,332,357]" name="Saint Kitts and Nevis" pageId="20" pageNumber="459">San Cristóbal</collectingCountry>
, 
<date id="8C601010596E6250FC8E264934C99F49" box="[787,915,332,356]" pageId="20" pageNumber="459" value="1980-03-17">17.III.1980</date>
, without collector— 
<specimenCount id="EED8FD59596E6250FB13264933869F48" box="[1166,1244,332,357]" pageId="20" pageNumber="459" type="male">1 male</specimenCount>
(MIZA); Santo 
<collectingCountry id="80C97640596E6250FF0A266A365E9FA5" box="[151,260,367,392]" name="Dominican Republic" pageId="20" pageNumber="459">Domingo</collectingCountry>
, Hacienda Santa Rosa, 
<date id="8C601010596E6250FD96267535309FA5" box="[523,618,368,392]" pageId="20" pageNumber="459" value="1992-06">VI.1992</date>
, J. Blanco col.— 
<specimenCount id="EED8FD59596E6250FCB3266A342F9FA5" box="[814,885,367,392]" pageId="20" pageNumber="459" type="male">1 male</specimenCount>
(MZSP) and 
<specimenCount id="EED8FD59596E6250FB93267533039FA5" box="[1038,1113,367,392]" pageId="20" pageNumber="459" type="male">2 male</specimenCount>
and 
<specimenCount id="EED8FD59596E6250FB0D267533AB9FA5" box="[1168,1265,367,392]" pageId="20" pageNumber="459" type="female">2 female</specimenCount>
(CEMT).
</paragraph>
</subSubSection>
</treatment>
</document>