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<mods:title id="98902F2C90A1043F623928A27AB78675">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="69DFBB099C34C942C5C268DDEB81E060">Cupello, Mario</mods:namePart>
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<mods:namePart id="1A18F02E33CF3CAB3D837060506A7649">Vaz-De-Mello, Fernando Z.</mods:namePart>
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<subSubSection id="B0C4655B596E6250FF0A26D934EF9CB7" pageId="20" pageNumber="459" type="nomenclature">
<paragraph id="F86136D0596E6250FF0A26D935369FDB" blockId="20.[151,703,476,535]" box="[151,620,476,502]" pageId="20" pageNumber="459">
<heading id="A32981BC596E6250FF0A26D935369FDB" bold="true" box="[151,620,476,502]" fontSize="11" level="1" pageId="20" pageNumber="459" reason="1">
<emphasis id="CAAAEAC2596E6250FF0A26D935369FDB" bold="true" box="[151,620,476,502]" pageId="20" pageNumber="459">
3. 
<taxonomicName id="3FDE4D53596E6250FF2926D935369FDB" ID-CoL="6KSGZ" authority="Blanchard, 1846" authorityName="Blanchard" authorityYear="1846" box="[180,620,476,502]" class="Insecta" family="Scarabaeidae" genus="Gromphas" kingdom="Animalia" order="Coleoptera" pageId="20" pageNumber="459" phylum="Arthropoda" rank="species" species="dichroa">
<emphasis id="CAAAEAC2596E6250FF2926D936CD9FDB" bold="true" box="[180,407,476,502]" italics="true" pageId="20" pageNumber="459">Gromphas dichroa</emphasis>
Blanchard, 1846
</taxonomicName>
</emphasis>
</heading>
</paragraph>
<paragraph id="F86136D0596E6250FF0A26FB35E59C3A" blockId="20.[151,703,476,535]" box="[151,703,510,535]" pageId="20" pageNumber="459">
<figureCitation id="60E52A55596E6250FF0A26FB37A99C3A" box="[151,243,510,535]" captionStart="FIGURES 13 – 15. 13" captionStartId="22.[151,264,1861,1883]" captionTargetBox="[155,1436,198,1838]" captionTargetId="figure@22.[151,1436,193,1840]" captionTargetPageId="22" captionText="FIGURES 13 – 15. 13: Gromphas amazonica Bates, 1870. 13 a) Dark blue specimen. 13 b) Reddish-brown specimen. 13 c) Dark green specimen. 14: Gromphas dichroa Blanchard, 1846. 14 a) Red-blue specimen. 14 b) Intermediate yellow-blue specimen. 14 c) Green specimen. 15: Gromphas inermis Harold, 1869. 15 a) Coppery specimen. 15 b) Dark green specimen. 15 c) Black specimen. Scale = 5 mm." httpUri="https://zenodo.org/record/247631/files/figure.png" pageId="20" pageNumber="459">Figs. 14</figureCitation>
, 19, 26, 33, 36, 42, 44, 47, 49, 54, 61, 66
</paragraph>
<paragraph id="F86136D0596E6250FF0A254334EF9CB7" blockId="20.[151,1436,582,696]" pageId="20" pageNumber="459">
<taxonomicName id="3FDE4D53596E6250FF0A254335649C71" authority="Blanchard, 1846: 182" authorityName="Blanchard" authorityPageNumber="182" authorityYear="1846" box="[151,574,582,604]" class="Insecta" family="Scarabaeidae" genus="Gromphas" kingdom="Animalia" order="Coleoptera" pageId="20" pageNumber="459" phylum="Arthropoda" rank="species" species="dichroa">
<emphasis id="CAAAEAC2596E6250FF0A2543360C9C71" box="[151,342,582,604]" italics="true" pageId="20" pageNumber="459">Gromphas dichroa</emphasis>
Blanchard, 1846: 182
</taxonomicName>
; Lacordaire 1856: 100; Harold 1869b: 62; Burmeister 1874: 130; Tremoleras 1910: 25; Gillet 1911: 80; d’Olsoufieff 1924: 21, 60, 139, plate IX; Pessoa &amp; Lane 1941: 473; Blackwelder 1944: 208; Martínez 1959: 95; Barattini &amp; Saenz, 1960: 22, 1964: 173; Figueroa 
<emphasis id="CAAAEAC2596E6250FCB42580340C9CB7" box="[809,854,644,666]" italics="true" pageId="20" pageNumber="459">et al</emphasis>
. 2012: 2.
</paragraph>
</subSubSection>
<subSubSection id="B0C4655B596E6250FF0A25A735779C95" box="[151,557,674,696]" pageId="20" pageNumber="459" type="description">
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<taxonomicName id="3FDE4D53596E6250FF0A25A735739C95" authority="Harold, 1869a: 1016" authorityName="Harold" authorityPageNumber="1016" authorityYear="1869" box="[151,553,674,696]" class="Insecta" family="Scarabaeidae" genus="Bolbites" kingdom="Animalia" order="Coleoptera" pageId="20" pageNumber="459" phylum="Arthropoda" rank="species" species="dichrous">
<emphasis id="CAAAEAC2596E6250FF0A25A736129C95" box="[151,328,674,696]" italics="true" pageId="20" pageNumber="459">Bolbites dichrous</emphasis>
: Harold, 1869a: 1016
</taxonomicName>
.
</paragraph>
</subSubSection>
<subSubSection id="B0C4655B596E6251FF0A25E0347B9F6D" lastPageId="21" lastPageNumber="460" pageId="20" pageNumber="459" type="materials_examined">
<paragraph id="F86136D0596E6250FF0A25E034D89CD0" blockId="20.[151,1437,741,2026]" box="[151,898,741,766]" pageId="20" pageNumber="459">
<emphasis id="CAAAEAC2596E6250FF0A25E0360A9CD3" bold="true" box="[151,336,741,766]" pageId="20" pageNumber="459">
<typeStatus id="27658872596E6250FF0A25E037889CD3" box="[151,210,741,766]" pageId="20" pageNumber="459">Type</typeStatus>
specimen:
</emphasis>
<typeStatus id="27658872596E6250FECA25E0369E9CD3" box="[343,452,741,766]" pageId="20" pageNumber="459" type="holotype">Holotype</typeStatus>
: Not located at MNHN; unknown to us.
</paragraph>
<paragraph id="F86136D0596E6250FF5A240C34D49D0F" blockId="20.[151,1437,741,2026]" box="[199,910,777,802]" pageId="20" pageNumber="459">
<emphasis id="CAAAEAC2596E6250FF5A240C36339D0F" bold="true" box="[199,361,777,802]" pageId="20" pageNumber="459">
<typeStatus id="27658872596E6250FF5A240C36589D0F" box="[199,258,777,802]" pageId="20" pageNumber="459">Type</typeStatus>
locality:
</emphasis>
“Montevideo, près de la mer” (Blanchard 1846).
</paragraph>
<paragraph id="F86136D0596E6250FF5A242832009DFB" blockId="20.[151,1437,741,2026]" pageId="20" pageNumber="459">
<emphasis id="CAAAEAC2596E6250FF5A2428362C9D6B" bold="true" box="[199,374,813,838]" pageId="20" pageNumber="459">Redescription:</emphasis>
Color: Two general patterns with intermediates (
<figureCitation id="60E52A55596E6250FC05242834B39D6B" box="[920,1001,813,838]" captionStart="FIGURES 13 – 15. 13" captionStartId="22.[151,264,1861,1883]" captionTargetBox="[155,1436,198,1838]" captionTargetId="figure@22.[151,1436,193,1840]" captionTargetPageId="22" captionText="FIGURES 13 – 15. 13: Gromphas amazonica Bates, 1870. 13 a) Dark blue specimen. 13 b) Reddish-brown specimen. 13 c) Dark green specimen. 14: Gromphas dichroa Blanchard, 1846. 14 a) Red-blue specimen. 14 b) Intermediate yellow-blue specimen. 14 c) Green specimen. 15: Gromphas inermis Harold, 1869. 15 a) Coppery specimen. 15 b) Dark green specimen. 15 c) Black specimen. Scale = 5 mm." httpUri="https://zenodo.org/record/247631/files/figure.png" pageId="20" pageNumber="459">Fig. 14</figureCitation>
b): 1) Anterior region of clypeus black; remainder of dorsum of head with strong metallic sheen of red, green, and yellow. Pronotum metallic blue on sides and along posterior margin and dark red in center; a narrow strip of metallic yellow and green surrounds entire red area. Metasternum, elytra, and pygidium blue or blue with green reflections (
<figureCitation id="60E52A55596E6250FB9F249C33059D9F" box="[1026,1119,921,946]" captionStart="FIGURES 13 – 15. 13" captionStartId="22.[151,264,1861,1883]" captionTargetBox="[155,1436,198,1838]" captionTargetId="figure@22.[151,1436,193,1840]" captionTargetPageId="22" captionText="FIGURES 13 – 15. 13: Gromphas amazonica Bates, 1870. 13 a) Dark blue specimen. 13 b) Reddish-brown specimen. 13 c) Dark green specimen. 14: Gromphas dichroa Blanchard, 1846. 14 a) Red-blue specimen. 14 b) Intermediate yellow-blue specimen. 14 c) Green specimen. 15: Gromphas inermis Harold, 1869. 15 a) Coppery specimen. 15 b) Dark green specimen. 15 c) Black specimen. Scale = 5 mm." httpUri="https://zenodo.org/record/247631/files/figure.png" pageId="20" pageNumber="459">Figs. 14</figureCitation>
a, 19). 2) Anterior region of clypeus black; remainder of dorsum, metasternum and pygidium predominantly bright light green (
<figureCitation id="60E52A55596E6250FB7024B832659DFB" box="[1261,1343,957,982]" captionStart="FIGURES 13 – 15. 13" captionStartId="22.[151,264,1861,1883]" captionTargetBox="[155,1436,198,1838]" captionTargetId="figure@22.[151,1436,193,1840]" captionTargetPageId="22" captionText="FIGURES 13 – 15. 13: Gromphas amazonica Bates, 1870. 13 a) Dark blue specimen. 13 b) Reddish-brown specimen. 13 c) Dark green specimen. 14: Gromphas dichroa Blanchard, 1846. 14 a) Red-blue specimen. 14 b) Intermediate yellow-blue specimen. 14 c) Green specimen. 15: Gromphas inermis Harold, 1869. 15 a) Coppery specimen. 15 b) Dark green specimen. 15 c) Black specimen. Scale = 5 mm." httpUri="https://zenodo.org/record/247631/files/figure.png" pageId="20" pageNumber="459">Fig. 14</figureCitation>
c).
</paragraph>
<paragraph id="F86136D0596E6250FF5A24E436E09A6F" blockId="20.[151,1437,741,2026]" pageId="20" pageNumber="459">
Head: Clypeus with truncate apex; genae with very dense and coarse granulation, including region adjacent to eyes (
<figureCitation id="60E52A55596E6250FF45230036779A33" box="[216,301,1029,1054]" 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="20" pageNumber="459">Fig. 26</figureCitation>
). Cephalic projection only slightly elevated and with truncate apex (
<figureCitation id="60E52A55596E6250FBB3230033DB9A33" box="[1070,1153,1029,1054]" 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="20" pageNumber="459">Fig. 33</figureCitation>
). Frons granulate at least adjacent to eyes (
<figureCitation id="60E52A55596E6250FEC4232C36F79A6F" box="[345,429,1065,1090]" 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="20" pageNumber="459">Fig. 26</figureCitation>
).
</paragraph>
<paragraph id="F86136D0596E6250FF5A234836E19AFF" blockId="20.[151,1437,741,2026]" pageId="20" pageNumber="459">Thorax: Pronotum globular; lateral region with very dense and coarse granulation (Fig. 19), density of granulation decreasing posteromedially; posteromedian region smooth or with strongly effaced granulation, except for a very narrow strip of dense granulation along posterior margin; posterior fossae present and granulate; posterior margin rounded.</paragraph>
<paragraph id="F86136D0596E6250FF5A23DB35869B13" blockId="20.[151,1437,741,2026]" pageId="20" pageNumber="459">
Mesosternum glabrous or with very sparse setae (
<figureCitation id="60E52A55596E6250FCA123D834C19ADB" box="[828,923,1245,1270]" 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="20" pageNumber="459">Fig. 49</figureCitation>
). Metasternum clearly punctate at center. Anteromedian angle strongly projecting and narrowed, with truncate apex (
<figureCitation id="60E52A55596E6250FC6A220433179B37" box="[1015,1101,1281,1306]" 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="20" pageNumber="459">Fig. 49</figureCitation>
); metasternal area in front of anteromedian angle entirely glabrous (Figs. 19, 49).
</paragraph>
<paragraph id="F86136D0596E6250FF5A224F33549BDF" blockId="20.[151,1437,741,2026]" pageId="20" pageNumber="459">
Legs: Protibiae very narrow in both sexes; in ventral view, longitudinal carina with a row of tubercles on basal half in males (Figs. 20d, 23b) and simple in females (
<figureCitation id="60E52A55596E6250FC99226834009BAB" box="[772,858,1389,1414]" 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="20" pageNumber="459">Fig. 23</figureCitation>
a). Protibial spur with apex strongly expanded and curved downward (
<figureCitation id="60E52A55596E6250FEEE229436889B87" box="[371,466,1425,1450]" 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="20" pageNumber="459">Figs. 23</figureCitation>
, 
<figureCitation id="60E52A55596E6250FE7D229436A59B87" box="[480,511,1425,1450]" 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="20" pageNumber="459">45</figureCitation>
); inner apical angle of protibia with tuft of setae longer in males than in females (
<figureCitation id="60E52A55596E6250FF0222B037A49BE3" box="[159,254,1461,1486]" 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="20" pageNumber="459">Figs. 23</figureCitation>
a–b). Metatibia very narrow (
<figureCitation id="60E52A55596E6250FDD222B035FF9BE3" box="[591,677,1461,1486]" 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="20" pageNumber="459">Fig. 42</figureCitation>
). Mesotarsi and metatarsi not enlarged and with apical tarsomeres strongly curved (
<figureCitation id="60E52A55596E6250FEC822DC36F39BDF" box="[341,425,1497,1522]" 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="20" pageNumber="459">Fig. 47</figureCitation>
). Metatibial spur with apex distinctly curved (
<figureCitation id="60E52A55596E6250FC3022DC33589BDF" box="[941,1026,1497,1522]" 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="20" pageNumber="459">Fig. 39</figureCitation>
).
</paragraph>
<paragraph id="F86136D0596E6250FF5A22F8356B9817" blockId="20.[151,1437,741,2026]" pageId="20" pageNumber="459">Elytra: Striae very fine, simple and effaced, almost imperceptible, especially eighth stria. Sutural margin smooth, distinctly depressed at base.</paragraph>
<paragraph id="F86136D0596E6250FF5A214033F498AF" blockId="20.[151,1437,741,2026]" pageId="20" pageNumber="459">
Abdomen: Pygidium with a pair of weak impressions on sides; basal margin complete and never interrupted by groove of propygidium (
<figureCitation id="60E52A55596E6250FE37216C36A498AF" box="[426,510,1641,1666]" 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="20" pageNumber="459">Fig. 35</figureCitation>
). Sixth sternite densely punctate, remaining sternites smooth.
</paragraph>
<paragraph id="F86136D0596E6250FF5A218836D898E7" blockId="20.[151,1437,741,2026]" pageId="20" pageNumber="459">
Aedeagus: Phallobase, in ventral view, with wide membranous apical area (
<figureCitation id="60E52A55596E6250FB8821883333988B" box="[1045,1129,1677,1702]" 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="20" pageNumber="459">Fig. 54</figureCitation>
c). Medial sclerite very thin and curved (
<figureCitation id="60E52A55596E6250FEBF21B4362F98E7" box="[290,373,1713,1738]" 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="20" pageNumber="459">Fig. 61</figureCitation>
).
</paragraph>
<paragraph id="F86136D0596E6250FF5A21D0327A9977" blockId="20.[151,1437,741,2026]" pageId="20" pageNumber="459">
<emphasis id="CAAAEAC2596E6250FF5A21D035EE98C3" bold="true" box="[199,692,1749,1774]" pageId="20" pageNumber="459">
Measurements: Males (
<specimenCount id="EED8FD59596E6250FE4021D0352B98C3" box="[477,625,1749,1774]" pageId="20" pageNumber="459" type="generic">2 specimens</specimenCount>
): TL:
</emphasis>
AV: 12.85; 
<collectingCountry id="80C97640596E6250FCA421D3343098C0" box="[825,874,1750,1773]" name="Mexico" pageId="20" pageNumber="459">MX</collectingCountry>
: 13; MN: 12.7; SD: 0.21. 
<emphasis id="CAAAEAC2596E6250FB1721D033ED98C3" bold="true" box="[1162,1207,1749,1774]" pageId="20" pageNumber="459">PL:</emphasis>
AV: 12.85; 
<collectingCountry id="80C97640596E6250FAA021D3323598C0" box="[1341,1391,1750,1773]" name="Mexico" pageId="20" pageNumber="459">MX</collectingCountry>
: 13; MN: 12.7; SD: 0.21. 
<emphasis id="CAAAEAC2596E6250FE1121FC369B993F" bold="true" box="[396,449,1785,1810]" pageId="20" pageNumber="459">PW:</emphasis>
AV: 7.6; 
<collectingCountry id="80C97640596E6250FDAF21FF3539993C" box="[562,611,1786,1809]" name="Mexico" pageId="20" pageNumber="459">MX</collectingCountry>
: 8.1; MN: 7.1; SD: 0.71. 
<emphasis id="CAAAEAC2596E6250FC1621FC3381993F" bold="true" box="[907,1243,1785,1810]" pageId="20" pageNumber="459">
Females (
<specimenCount id="EED8FD59596E6250FC6221FC33CC993F" box="[1023,1174,1785,1810]" pageId="20" pageNumber="459" type="generic">5 specimens</specimenCount>
): TL:
</emphasis>
AV: 14.84; 
<collectingCountry id="80C97640596E6250FAFA21FF32C2993C" box="[1383,1432,1786,1809]" name="Mexico" pageId="20" pageNumber="459">MX</collectingCountry>
: 16.6; MN: 12.5; SD: 1.54. 
<emphasis id="CAAAEAC2596E6250FE53201836A1991B" bold="true" box="[462,507,1821,1846]" pageId="20" pageNumber="459">PL:</emphasis>
AV: 12.44; 
<collectingCountry id="80C97640596E6250FD15201B35E39918" box="[648,697,1822,1845]" name="Mexico" pageId="20" pageNumber="459">MX</collectingCountry>
: 13.3; MN: 11.8; SD: 0.74. 
<emphasis id="CAAAEAC2596E6250FC612018336B991B" bold="true" box="[1020,1073,1821,1846]" pageId="20" pageNumber="459">PW:</emphasis>
AV: 8.8; 
<collectingCountry id="80C97640596E6250FB3F201B33899918" box="[1186,1235,1822,1845]" name="Mexico" pageId="20" pageNumber="459">MX</collectingCountry>
: 9.7; MN: 8; SD: 0.79. 
<emphasis id="CAAAEAC2596E6250FF4B204436A09977" bold="true" box="[214,506,1857,1882]" pageId="20" pageNumber="459">
Total (
<specimenCount id="EED8FD59596E6250FEBF204436ED9977" box="[290,439,1857,1882]" pageId="20" pageNumber="459" type="generic">7 specimens</specimenCount>
): TL:
</emphasis>
AV: 14.27; SD: 1.59. 
<emphasis id="CAAAEAC2596E6250FD6F204434449977" bold="true" box="[754,798,1857,1882]" pageId="20" pageNumber="459">PL:</emphasis>
AV: 11.94; SD: 1.1. 
<emphasis id="CAAAEAC2596E6250FB95204433669977" bold="true" box="[1032,1084,1857,1882]" pageId="20" pageNumber="459">PW:</emphasis>
AV: 8.46; SD: 0.92.
</paragraph>
<paragraph id="F86136D0596E6251FF5A2060347B9F6D" blockId="20.[151,1437,741,2026]" lastBlockId="21.[151,1436,151,1977]" lastPageId="21" lastPageNumber="460" pageId="20" pageNumber="459">
<emphasis id="CAAAEAC2596E6250FF5A206034429953" bold="true" box="[199,792,1893,1918]" pageId="20" pageNumber="459">Intraspecific variation and taxonomic discussion:</emphasis>
The most striking intraspecific variation in 
<emphasis id="CAAAEAC2596E6250FA96206332249950" box="[1291,1406,1893,1918]" italics="true" pageId="20" pageNumber="459">G. d i ch ro a</emphasis>
is the color: two distinct patterns are observable, one where green is predominant (
<figureCitation id="60E52A55596E6250FBB8208C332D998F" box="[1061,1143,1929,1954]" captionStart="FIGURES 13 – 15. 13" captionStartId="22.[151,264,1861,1883]" captionTargetBox="[155,1436,198,1838]" captionTargetId="figure@22.[151,1436,193,1840]" captionTargetPageId="22" captionText="FIGURES 13 – 15. 13: Gromphas amazonica Bates, 1870. 13 a) Dark blue specimen. 13 b) Reddish-brown specimen. 13 c) Dark green specimen. 14: Gromphas dichroa Blanchard, 1846. 14 a) Red-blue specimen. 14 b) Intermediate yellow-blue specimen. 14 c) Green specimen. 15: Gromphas inermis Harold, 1869. 15 a) Coppery specimen. 15 b) Dark green specimen. 15 c) Black specimen. Scale = 5 mm." httpUri="https://zenodo.org/record/247631/files/figure.png" pageId="20" pageNumber="459">Fig. 14</figureCitation>
c) and the other with dark blue elytra and reddish-blue pronotum (
<figureCitation id="60E52A55596E6250FDC420A835F699EB" box="[601,684,1965,1990]" captionStart="FIGURES 13 – 15. 13" captionStartId="22.[151,264,1861,1883]" captionTargetBox="[155,1436,198,1838]" captionTargetId="figure@22.[151,1436,193,1840]" captionTargetPageId="22" captionText="FIGURES 13 – 15. 13: Gromphas amazonica Bates, 1870. 13 a) Dark blue specimen. 13 b) Reddish-brown specimen. 13 c) Dark green specimen. 14: Gromphas dichroa Blanchard, 1846. 14 a) Red-blue specimen. 14 b) Intermediate yellow-blue specimen. 14 c) Green specimen. 15: Gromphas inermis Harold, 1869. 15 a) Coppery specimen. 15 b) Dark green specimen. 15 c) Black specimen. Scale = 5 mm." httpUri="https://zenodo.org/record/247631/files/figure.png" pageId="20" pageNumber="459">Fig. 14</figureCitation>
a). However, the specimen from Porto Alegre (Rio Grande do Sul, 
<collectingCountry id="80C97640596E6250FF0A20D4378799C7" box="[151,221,2001,2026]" name="Brazil" pageId="20" pageNumber="459">Brazil</collectingCountry>
) has an intermediate coloration: the elytra are light blue and the pronotum and head are green with a large iridescent yellowish-red spot (
<figureCitation id="60E52A55596F6251FE6A279235179E9D" box="[503,589,151,176]" captionStart="FIGURES 13 – 15. 13" captionStartId="22.[151,264,1861,1883]" captionTargetBox="[155,1436,198,1838]" captionTargetId="figure@22.[151,1436,193,1840]" captionTargetPageId="22" captionText="FIGURES 13 – 15. 13: Gromphas amazonica Bates, 1870. 13 a) Dark blue specimen. 13 b) Reddish-brown specimen. 13 c) Dark green specimen. 14: Gromphas dichroa Blanchard, 1846. 14 a) Red-blue specimen. 14 b) Intermediate yellow-blue specimen. 14 c) Green specimen. 15: Gromphas inermis Harold, 1869. 15 a) Coppery specimen. 15 b) Dark green specimen. 15 c) Black specimen. Scale = 5 mm." httpUri="https://zenodo.org/record/247631/files/figure.png" pageId="21" pageNumber="460">Fig. 14</figureCitation>
b). The green female collected in 
<collectingCountry id="80C97640596F6251FC42279D33339E9D" box="[991,1129,152,176]" name="Sao Tome and Principe" pageId="21" pageNumber="460">Santo Tomé</collectingCountry>
(Corrientes, 
<collectingCountry id="80C97640596F6251FA982792322C9E9D" box="[1285,1398,151,176]" name="Argentina" pageId="21" pageNumber="460">Argentina</collectingCountry>
) is labeled “ 
<taxonomicName id="3FDE4D53596F6251FF6127B9368A9EF8" box="[252,464,188,213]" class="Insecta" family="Scarabaeidae" genus="Gromphas" kingdom="Animalia" order="Coleoptera" pageId="21" pageNumber="460" phylum="Arthropoda" rank="species" species="dichroa">
<emphasis id="CAAAEAC2596F6251FF6127B9368A9EF8" box="[252,464,188,213]" italics="true" pageId="21" pageNumber="460">Gromphas dichroa</emphasis>
</taxonomicName>
<taxonomicName id="3FDE4D53596F6251FE4527B835039EF8" box="[472,601,188,213]" pageId="21" pageNumber="460" rank="variety" variety="fulgida">
var. 
<emphasis id="CAAAEAC2596F6251FD9427B935039EF8" box="[521,601,188,213]" italics="true" pageId="21" pageNumber="460">fulgida</emphasis>
</taxonomicName>
nov. A. Martínez det. 1947”, a name never published. There is no reason for the recognition of a separate taxon for specimens of this color pattern, considering that this characteristic is not absolute, as is well demonstrated by the specimen from Porto Alegre, and there is no morphological or distributional difference from the reddish-blue specimens.
</paragraph>
</subSubSection>
<subSubSection id="B0C4655B596F6251FF5A264932749B21" pageId="21" pageNumber="460" type="description">
<paragraph id="F86136D0596F6251FF5A264934B89D38" blockId="21.[151,1436,151,1977]" pageId="21" pageNumber="460">
<taxonomicName id="3FDE4D53596F6251FF5A264936C39F48" box="[199,409,332,357]" class="Insecta" family="Scarabaeidae" genus="Gromphas" kingdom="Animalia" order="Coleoptera" pageId="21" pageNumber="460" phylum="Arthropoda" rank="species" species="dichroa">
<emphasis id="CAAAEAC2596F6251FF5A264936C39F48" box="[199,409,332,357]" italics="true" pageId="21" pageNumber="460">Gromphas dichroa</emphasis>
</taxonomicName>
has the highest number of unique characters in the genus: (a) truncate apex of clypeus (
<figureCitation id="60E52A55596F6251FAF3264937ED9FA5" 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="21" pageNumber="460">Fig. 26</figureCitation>
); (b) clypeus not folded upwards to any degree (in 
<taxonomicName id="3FDE4D53596F6251FD6A267534319FA5" box="[759,875,368,392]" class="Insecta" family="Scarabaeidae" genus="Gromphas" kingdom="Animalia" order="Coleoptera" pageId="21" pageNumber="460" phylum="Arthropoda" rank="species" species="inermis">
<emphasis id="CAAAEAC2596F6251FD6A267534319FA5" box="[759,875,368,392]" italics="true" pageId="21" pageNumber="460">G. inermis</emphasis>
</taxonomicName>
and 
<taxonomicName id="3FDE4D53596F6251FC3F267533609FA5" box="[930,1082,368,392]" class="Insecta" family="Scarabaeidae" genus="Gromphas" kingdom="Animalia" order="Coleoptera" pageId="21" pageNumber="460" phylum="Arthropoda" rank="species" species="amazonica">
<emphasis id="CAAAEAC2596F6251FC3F267533609FA5" box="[930,1082,368,392]" italics="true" pageId="21" pageNumber="460">G. amazonica</emphasis>
</taxonomicName>
, the clypeus is strongly folded, while in 
<taxonomicName id="3FDE4D53596F6251FF66269036C39F81" box="[251,409,405,428]" class="Insecta" family="Scarabaeidae" genus="Gromphas" kingdom="Animalia" order="Coleoptera" pageId="21" pageNumber="460" phylum="Arthropoda" rank="species" species="aeruginosa">
<emphasis id="CAAAEAC2596F6251FF66269036C39F81" box="[251,409,405,428]" italics="true" pageId="21" pageNumber="460">G. aeruginosa</emphasis>
</taxonomicName>
and 
<taxonomicName id="3FDE4D53596F6251FE4F2690350B9F80" box="[466,593,404,429]" class="Insecta" family="Scarabaeidae" genus="Gromphas" kingdom="Animalia" order="Coleoptera" pageId="21" pageNumber="460" phylum="Arthropoda" rank="species" species="lemoinei">
<emphasis id="CAAAEAC2596F6251FE4F2690350B9F80" box="[466,593,404,429]" italics="true" pageId="21" pageNumber="460">G. lemoinei</emphasis>
</taxonomicName>
it is only slightly); (c) truncate cephalic projection (
<figureCitation id="60E52A55596F6251FB38269133A39F81" box="[1189,1273,404,429]" 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="21" pageNumber="460">Fig. 33</figureCitation>
); (d) posterior margin of pronotum with narrow margin of granulation, including posterior fossae; (e) mesosternum glabrous or with only very short and sparse pubescence, metasternum glabrous anteriorly to the anteromedian angle (Figs. 19, 49); (f) metatibia very narrow (
<figureCitation id="60E52A55596F6251FE6426FA35149C35" box="[505,590,511,536]" 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="21" pageNumber="460">Fig. 42</figureCitation>
); (g) apical mesotarsomeres and metatarsomeres strongly curved (
<figureCitation id="60E52A55596F6251FAA426FA32D49C35" box="[1337,1422,511,536]" 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="21" pageNumber="460">Fig. 47</figureCitation>
); (h) elytral suture depressed at base (
<figureCitation id="60E52A55596F6251FDDC252135FA9C11" box="[577,672,548,573]" captionStart="FIGURES 13 – 15. 13" captionStartId="22.[151,264,1861,1883]" captionTargetBox="[155,1436,198,1838]" captionTargetId="figure@22.[151,1436,193,1840]" captionTargetPageId="22" captionText="FIGURES 13 – 15. 13: Gromphas amazonica Bates, 1870. 13 a) Dark blue specimen. 13 b) Reddish-brown specimen. 13 c) Dark green specimen. 14: Gromphas dichroa Blanchard, 1846. 14 a) Red-blue specimen. 14 b) Intermediate yellow-blue specimen. 14 c) Green specimen. 15: Gromphas inermis Harold, 1869. 15 a) Coppery specimen. 15 b) Dark green specimen. 15 c) Black specimen. Scale = 5 mm." httpUri="https://zenodo.org/record/247631/files/figure.png" pageId="21" pageNumber="460">Figs. 14</figureCitation>
a–c); and (i) base of pygidium completely margined (
<figureCitation id="60E52A55596F6251FA88252132369C10" box="[1301,1388,548,573]" 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="21" pageNumber="460">Fig. 35</figureCitation>
). In addition, (j) the anteromedian angle of metasternum of 
<taxonomicName id="3FDE4D53596F6251FC99254D34239C4D" box="[772,889,583,608]" class="Insecta" family="Scarabaeidae" genus="Gromphas" kingdom="Animalia" order="Coleoptera" pageId="21" pageNumber="460" phylum="Arthropoda" rank="species" species="dichroa">
<emphasis id="CAAAEAC2596F6251FC99254D34239C4D" box="[772,889,583,608]" italics="true" pageId="21" pageNumber="460">G. dichroa</emphasis>
</taxonomicName>
differs from the other species: it is convex, as in 
<taxonomicName id="3FDE4D53596F6251FF0A256836759CA9" box="[151,303,621,644]" class="Insecta" family="Scarabaeidae" genus="Gromphas" kingdom="Animalia" order="Coleoptera" pageId="21" pageNumber="460" phylum="Arthropoda" rank="species" species="amazonica">
<emphasis id="CAAAEAC2596F6251FF0A256836759CA9" box="[151,303,621,644]" italics="true" pageId="21" pageNumber="460">G. amazonica</emphasis>
</taxonomicName>
and 
<taxonomicName id="3FDE4D53596F6251FEF92568368D9CA9" box="[356,471,621,644]" class="Insecta" family="Scarabaeidae" genus="Gromphas" kingdom="Animalia" order="Coleoptera" pageId="21" pageNumber="460" phylum="Arthropoda" rank="species" species="inermis">
<emphasis id="CAAAEAC2596F6251FEF92568368D9CA9" box="[356,471,621,644]" italics="true" pageId="21" pageNumber="460">G. inermis</emphasis>
</taxonomicName>
(
<figureCitation id="60E52A55596F6251FE7B256935349CA8" box="[486,622,620,645]" 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="21" pageNumber="460">Figs. 50–51</figureCitation>
), but is still much more projected and has a truncate apex (
<figureCitation id="60E52A55596F6251FA9D2569320E9CA9" box="[1280,1364,620,645]" 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="21" pageNumber="460">Fig. 49</figureCitation>
); in 
<taxonomicName id="3FDE4D53596F6251FA19256837A29C85" class="Insecta" family="Scarabaeidae" genus="Gromphas" kingdom="Animalia" order="Coleoptera" pageId="21" pageNumber="460" phylum="Arthropoda" rank="species" species="lemoinei">
<emphasis id="CAAAEAC2596F6251FA19256837A29C85" italics="true" pageId="21" pageNumber="460">G. lemoinei</emphasis>
</taxonomicName>
and 
<taxonomicName id="3FDE4D53596F6251FEB2259536969C85" box="[303,460,656,680]" class="Insecta" family="Scarabaeidae" genus="Gromphas" kingdom="Animalia" order="Coleoptera" pageId="21" pageNumber="460" phylum="Arthropoda" rank="species" species="aeruginosa">
<emphasis id="CAAAEAC2596F6251FEB2259536969C85" box="[303,460,656,680]" italics="true" pageId="21" pageNumber="460">G. aeruginosa</emphasis>
</taxonomicName>
, the angle is always very flat (
<figureCitation id="60E52A55596F6251FCBB258A34219C85" box="[806,891,655,680]" 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="21" pageNumber="460">Fig. 52</figureCitation>
). The smooth sutural margin of elytra (k) is also unique to 
<taxonomicName id="3FDE4D53596F6251FE9525B036279CE0" box="[264,381,692,717]" class="Insecta" family="Scarabaeidae" genus="Gromphas" kingdom="Animalia" order="Coleoptera" pageId="21" pageNumber="460" phylum="Arthropoda" rank="species" species="dichroa">
<emphasis id="CAAAEAC2596F6251FE9525B036279CE0" box="[264,381,692,717]" italics="true" pageId="21" pageNumber="460">G. dichroa</emphasis>
</taxonomicName>
(in 
<taxonomicName id="3FDE4D53596F6251FE3125B035139CE1" box="[428,585,693,716]" class="Insecta" family="Scarabaeidae" genus="Gromphas" kingdom="Animalia" order="Coleoptera" pageId="21" pageNumber="460" phylum="Arthropoda" rank="species" species="aeruginosa">
<emphasis id="CAAAEAC2596F6251FE3125B035139CE1" box="[428,585,693,716]" italics="true" pageId="21" pageNumber="460">G. aeruginosa</emphasis>
</taxonomicName>
the punctation is very sparse, but still present). The color pattern described above for this species (l) is also unique among 
<taxonomicName id="3FDE4D53596F6251FD2A25D234769CDD" box="[695,812,727,752]" class="Insecta" family="Scarabaeidae" genus="Gromphas" kingdom="Animalia" order="Coleoptera" pageId="21" pageNumber="460" phylum="Arthropoda" rank="genus">
<emphasis id="CAAAEAC2596F6251FD2A25D234769CDD" box="[695,812,727,752]" italics="true" pageId="21" pageNumber="460">Gromphas</emphasis>
</taxonomicName>
. Finally, the general shape of the medial sclerite of 
<taxonomicName id="3FDE4D53596F6251FA1925DD37B79D38" class="Insecta" family="Scarabaeidae" genus="Gromphas" kingdom="Animalia" order="Coleoptera" pageId="21" pageNumber="460" phylum="Arthropoda" rank="species" species="dichroa">
<emphasis id="CAAAEAC2596F6251FA1925DD37B79D38" italics="true" pageId="21" pageNumber="460">G. dichroa</emphasis>
</taxonomicName>
(m) is distinct from that of all other species of 
<taxonomicName id="3FDE4D53596F6251FD6025F934289D38" box="[765,882,764,789]" class="Insecta" family="Scarabaeidae" genus="Gromphas" kingdom="Animalia" order="Coleoptera" pageId="21" pageNumber="460" phylum="Arthropoda" rank="genus">
<emphasis id="CAAAEAC2596F6251FD6025F934289D38" box="[765,882,764,789]" italics="true" pageId="21" pageNumber="460">Gromphas</emphasis>
</taxonomicName>
(
<figureCitation id="60E52A55596F6251FC1C25F9348F9D38" box="[897,981,764,789]" 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="21" pageNumber="460">Fig. 61</figureCitation>
).
</paragraph>
<paragraph id="F86136D0596F6251FF5A241A35659A8D" blockId="21.[151,1436,151,1977]" pageId="21" pageNumber="460">
<taxonomicName id="3FDE4D53596F6251FF5A241A36C39D15" box="[199,409,799,824]" class="Insecta" family="Scarabaeidae" genus="Gromphas" kingdom="Animalia" order="Coleoptera" pageId="21" pageNumber="460" phylum="Arthropoda" rank="species" species="dichroa">
<emphasis id="CAAAEAC2596F6251FF5A241A36C39D15" box="[199,409,799,824]" italics="true" pageId="21" pageNumber="460">Gromphas dichroa</emphasis>
</taxonomicName>
shares with 
<taxonomicName id="3FDE4D53596F6251FDBB242535FF9D15" box="[550,677,799,824]" class="Insecta" family="Scarabaeidae" genus="Gromphas" kingdom="Animalia" order="Coleoptera" pageId="21" pageNumber="460" phylum="Arthropoda" rank="species" species="lemoinei">
<emphasis id="CAAAEAC2596F6251FDBB242535FF9D15" box="[550,677,799,824]" italics="true" pageId="21" pageNumber="460">G. lemoinei</emphasis>
</taxonomicName>
and 
<taxonomicName id="3FDE4D53596F6251FD41242534239D15" box="[732,889,800,824]" class="Insecta" family="Scarabaeidae" genus="Gromphas" kingdom="Animalia" order="Coleoptera" pageId="21" pageNumber="460" phylum="Arthropoda" rank="species" species="aeruginosa">
<emphasis id="CAAAEAC2596F6251FD41242534239D15" box="[732,889,800,824]" italics="true" pageId="21" pageNumber="460">G. aeruginosa</emphasis>
</taxonomicName>
a clypeus with rounded sides (
<figureCitation id="60E52A55596F6251FB4E241A327C9D15" box="[1235,1318,799,824]" 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="21" pageNumber="460">Fig. 26</figureCitation>
), and only with 
<taxonomicName id="3FDE4D53596F6251FF4D244036379D71" box="[208,365,837,860]" class="Insecta" family="Scarabaeidae" genus="Gromphas" kingdom="Animalia" order="Coleoptera" pageId="21" pageNumber="460" phylum="Arthropoda" rank="species" species="aeruginosa">
<emphasis id="CAAAEAC2596F6251FF4D244036379D71" box="[208,365,837,860]" italics="true" pageId="21" pageNumber="460">G. aeruginosa</emphasis>
</taxonomicName>
the presence of posterior pronotal fossae in specimens of any size (in 
<taxonomicName id="3FDE4D53596F6251FB1A2440325C9D70" box="[1159,1286,836,861]" class="Insecta" family="Scarabaeidae" genus="Gromphas" kingdom="Animalia" order="Coleoptera" pageId="21" pageNumber="460" phylum="Arthropoda" rank="species" species="lemoinei">
<emphasis id="CAAAEAC2596F6251FB1A2440325C9D70" box="[1159,1286,836,861]" italics="true" pageId="21" pageNumber="460">G. lemoinei</emphasis>
</taxonomicName>
, these fossae are absent or greatly reduced in larger specimens) and the two tenuous impressions on the sides of pygidium (this feature is very subtle and often absent in 
<taxonomicName id="3FDE4D53596F6251FD16248834779D89" box="[651,813,909,932]" class="Insecta" family="Scarabaeidae" genus="Gromphas" kingdom="Animalia" order="Coleoptera" pageId="21" pageNumber="460" phylum="Arthropoda" rank="species" species="aeruginosa">
<emphasis id="CAAAEAC2596F6251FD16248834779D89" box="[651,813,909,932]" italics="true" pageId="21" pageNumber="460">G. aeruginosa</emphasis>
</taxonomicName>
). On the other hand, it resembles 
<taxonomicName id="3FDE4D53596F6251FB512488321E9D89" box="[1228,1348,909,932]" class="Insecta" family="Scarabaeidae" genus="Gromphas" kingdom="Animalia" order="Coleoptera" pageId="21" pageNumber="460" phylum="Arthropoda" rank="species" species="inermis">
<emphasis id="CAAAEAC2596F6251FB512488321E9D89" box="[1228,1348,909,932]" italics="true" pageId="21" pageNumber="460">G. inermis</emphasis>
</taxonomicName>
and 
<taxonomicName id="3FDE4D53596F6251FA192488364B9DE5" class="Insecta" family="Scarabaeidae" genus="Gromphas" kingdom="Animalia" order="Coleoptera" pageId="21" pageNumber="460" phylum="Arthropoda" rank="species" species="amazonica">
<emphasis id="CAAAEAC2596F6251FA192488364B9DE5" italics="true" pageId="21" pageNumber="460">G. amazonica</emphasis>
</taxonomicName>
by the absence of any kind of pronotal prominence; by the protibial spur expanded at the apex (
<figureCitation id="60E52A55596F6251FAA724AA32CD9DE5" box="[1338,1431,943,968]" 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="21" pageNumber="460">Figs. 21</figureCitation>
, 
<figureCitation id="60E52A55596F6251FF0A24D137ED9DC0" box="[151,183,980,1005]" 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="21" pageNumber="460">45</figureCitation>
); and by the sexual dimorphism in the tuft of setae at the inner apical angle of protibiae (distinctly more developed in males than in females [
<figureCitation id="60E52A55596F6251FDAC24F235D59A3D" box="[561,655,1015,1040]" 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="21" pageNumber="460">Figs. 23</figureCitation>
a–b]) and in the ventral carina of protibiae (with a row of tubercles in males [Figs. 20d, 23b] and simple in females [
<figureCitation id="60E52A55596F6251FD06231935AB9A19" box="[667,753,1052,1077]" 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="21" pageNumber="460">Fig. 23</figureCitation>
a]). Only with 
<taxonomicName id="3FDE4D53596F6251FC0F2318335E9A19" box="[914,1028,1053,1076]" class="Insecta" family="Scarabaeidae" genus="Gromphas" kingdom="Animalia" order="Coleoptera" pageId="21" pageNumber="460" phylum="Arthropoda" rank="species" species="inermis">
<emphasis id="CAAAEAC2596F6251FC0F2318335E9A19" box="[914,1028,1053,1076]" italics="true" pageId="21" pageNumber="460">G. inermis</emphasis>
</taxonomicName>
does 
<taxonomicName id="3FDE4D53596F6251FBD8231833E09A18" box="[1093,1210,1052,1077]" class="Insecta" family="Scarabaeidae" genus="Gromphas" kingdom="Animalia" order="Coleoptera" pageId="21" pageNumber="460" phylum="Arthropoda" rank="species" species="dichroa">
<emphasis id="CAAAEAC2596F6251FBD8231833E09A18" box="[1093,1210,1052,1077]" italics="true" pageId="21" pageNumber="460">G. dichroa</emphasis>
</taxonomicName>
share simple elytral striae (in 
<taxonomicName id="3FDE4D53596F6251FE9C2345362F9A75" box="[257,373,1087,1112]" class="Insecta" family="Scarabaeidae" genus="Gromphas" kingdom="Animalia" order="Coleoptera" pageId="21" pageNumber="460" phylum="Arthropoda" rank="species" species="dichroa">
<emphasis id="CAAAEAC2596F6251FE9C2345362F9A75" box="[257,373,1087,1112]" italics="true" pageId="21" pageNumber="460">G. dichroa</emphasis>
</taxonomicName>
, they are very indistinct and almost imperceptible, especially the eighth and ninth striae, while in 
<taxonomicName id="3FDE4D53596F6251FF2B236036719A51" box="[182,299,1125,1148]" class="Insecta" family="Scarabaeidae" genus="Gromphas" kingdom="Animalia" order="Coleoptera" pageId="21" pageNumber="460" phylum="Arthropoda" rank="species" species="inermis">
<emphasis id="CAAAEAC2596F6251FF2B236036719A51" box="[182,299,1125,1148]" italics="true" pageId="21" pageNumber="460">G. inermis</emphasis>
</taxonomicName>
all striae are easily observed); and only with 
<taxonomicName id="3FDE4D53596F6251FCA2236034809A51" box="[831,986,1125,1148]" class="Insecta" family="Scarabaeidae" genus="Gromphas" kingdom="Animalia" order="Coleoptera" pageId="21" pageNumber="460" phylum="Arthropoda" rank="species" species="amazonica">
<emphasis id="CAAAEAC2596F6251FCA2236034809A51" box="[831,986,1125,1148]" italics="true" pageId="21" pageNumber="460">G. amazonica</emphasis>
</taxonomicName>
does it share a smooth posteriomedian region of the pronotum (
<figureCitation id="60E52A55596F6251FE342382356B9A8D" box="[425,561,1159,1184]" captionStart="FIGURES 13 – 15. 13" captionStartId="22.[151,264,1861,1883]" captionTargetBox="[155,1436,198,1838]" captionTargetId="figure@22.[151,1436,193,1840]" captionTargetPageId="22" captionText="FIGURES 13 – 15. 13: Gromphas amazonica Bates, 1870. 13 a) Dark blue specimen. 13 b) Reddish-brown specimen. 13 c) Dark green specimen. 14: Gromphas dichroa Blanchard, 1846. 14 a) Red-blue specimen. 14 b) Intermediate yellow-blue specimen. 14 c) Green specimen. 15: Gromphas inermis Harold, 1869. 15 a) Coppery specimen. 15 b) Dark green specimen. 15 c) Black specimen. Scale = 5 mm." httpUri="https://zenodo.org/record/247631/files/figure.png" pageId="21" pageNumber="460">Figs. 13, 14</figureCitation>
).
</paragraph>
<paragraph id="F86136D0596F6251FF5A23A832749B21" blockId="21.[151,1436,151,1977]" pageId="21" pageNumber="460">
In Harold’s (1869a) catalog, 
<taxonomicName id="3FDE4D53596F6251FD8123A835CF9AE8" box="[540,661,1196,1221]" class="Insecta" family="Scarabaeidae" genus="Gromphas" kingdom="Animalia" order="Coleoptera" pageId="21" pageNumber="460" phylum="Arthropoda" rank="species" species="dichroa">
<emphasis id="CAAAEAC2596F6251FD8123A835CF9AE8" box="[540,661,1196,1221]" italics="true" pageId="21" pageNumber="460">G. dichroa</emphasis>
</taxonomicName>
was transferred to the genus 
<taxonomicName id="3FDE4D53596F6251FC6423A9330F9AE8" box="[1017,1109,1196,1221]" class="Insecta" family="Scarabaeidae" genus="Bolbites" kingdom="Animalia" order="Coleoptera" pageId="21" pageNumber="460" phylum="Arthropoda" rank="genus">
<emphasis id="CAAAEAC2596F6251FC6423A9330F9AE8" box="[1017,1109,1196,1221]" italics="true" pageId="21" pageNumber="460">Bolbites</emphasis>
</taxonomicName>
, but soon thereafter Harold (1869b) returned this species to its original genus. From all the characters discussed in the description of 
<taxonomicName id="3FDE4D53596F6251FABA23CA32C69AC5" box="[1319,1436,1231,1256]" class="Insecta" family="Scarabaeidae" genus="Gromphas" kingdom="Animalia" order="Coleoptera" pageId="21" pageNumber="460" phylum="Arthropoda" rank="genus">
<emphasis id="CAAAEAC2596F6251FABA23CA32C69AC5" box="[1319,1436,1231,1256]" italics="true" pageId="21" pageNumber="460">Gromphas</emphasis>
</taxonomicName>
and in the diagnosis of Gromphadina, it is clear that 
<emphasis id="CAAAEAC2596F6251FD7F23F0340E9B21" box="[738,852,1268,1293]" italics="true" pageId="21" pageNumber="460">G. di ch ro a</emphasis>
belongs in 
<taxonomicName id="3FDE4D53596F6251FC4523F133179B20" box="[984,1101,1268,1293]" class="Insecta" family="Scarabaeidae" genus="Gromphas" kingdom="Animalia" order="Coleoptera" pageId="21" pageNumber="460" phylum="Arthropoda" rank="genus">
<emphasis id="CAAAEAC2596F6251FC4523F133179B20" box="[984,1101,1268,1293]" italics="true" pageId="21" pageNumber="460">Gromphas</emphasis>
</taxonomicName>
and not in 
<taxonomicName id="3FDE4D53596F6251FB5123F132729B20" box="[1228,1320,1268,1293]" class="Insecta" family="Scarabaeidae" genus="Bolbites" kingdom="Animalia" order="Coleoptera" pageId="21" pageNumber="460" phylum="Arthropoda" rank="genus">
<emphasis id="CAAAEAC2596F6251FB5123F132729B20" box="[1228,1320,1268,1293]" italics="true" pageId="21" pageNumber="460">Bolbites</emphasis>
</taxonomicName>
.
</paragraph>
</subSubSection>
<subSubSection id="B0C4655B596F6253FF5A2212336B9FD8" lastPageId="23" lastPageNumber="462" pageId="21" pageNumber="460" type="discussion">
<paragraph id="F86136D0596F6251FF5A2212359D9BB0" blockId="21.[151,1436,151,1977]" pageId="21" pageNumber="460">
<emphasis id="CAAAEAC2596F6251FF5A221236129B1D" bold="true" box="[199,328,1303,1328]" pageId="21" pageNumber="460">Comments</emphasis>
: We have not been able to examine the 
<typeStatus id="27658872596F6251FC9B221234329B1D" box="[774,872,1303,1328]" pageId="21" pageNumber="460" type="holotype">holotype</typeStatus>
of 
<emphasis id="CAAAEAC2596F6251FC13221D335A9B1D" box="[910,1024,1303,1328]" italics="true" pageId="21" pageNumber="460">G. di ch ro a</emphasis>
. Blanchard's (1846) description was based on a single specimen that he believed to be a female (“ 
<emphasis id="CAAAEAC2596F6251FCA3223835029B55" italics="true" pageId="21" pageNumber="460">Nous n’en avons vu qu’une femelle, que M. d’Orbigny a trouvée à Montevideo, près de la mer</emphasis>
”). d’Olsoufieff (1924) studied the 
<typeStatus id="27658872596F6251FC7A225A33139B55" box="[999,1097,1375,1400]" pageId="21" pageNumber="460" type="holotype">holotype</typeStatus>
and said it was housed in the MNHN collection, but we have not found it there.
</paragraph>
<paragraph id="F86136D0596F6251FF5A22A2326D98B5" blockId="21.[151,1436,151,1977]" pageId="21" pageNumber="460">
<taxonomicName id="3FDE4D53596F6251FF5A22A236C39BED" box="[199,409,1447,1472]" class="Insecta" family="Scarabaeidae" genus="Gromphas" kingdom="Animalia" order="Coleoptera" pageId="21" pageNumber="460" phylum="Arthropoda" rank="species" species="dichroa">
<emphasis id="CAAAEAC2596F6251FF5A22A236C39BED" box="[199,409,1447,1472]" italics="true" pageId="21" pageNumber="460">Gromphas dichroa</emphasis>
</taxonomicName>
and 
<taxonomicName id="3FDE4D53596F6251FE4D22AD35BF9BED" authority="Blanchard" authorityName="Blanchard" box="[464,741,1447,1472]" class="Insecta" family="Scarabaeidae" genus="Gromphas" kingdom="Animalia" order="Coleoptera" pageId="21" pageNumber="460" phylum="Arthropoda" rank="species" species="lacordairei">
<emphasis id="CAAAEAC2596F6251FE4D22AD35319BED" box="[464,619,1447,1472]" italics="true" pageId="21" pageNumber="460">G. lacordairei</emphasis>
Blanchard
</taxonomicName>
were published respectively on pages 181 and 182 of volume 6, part 2, of 
<emphasis id="CAAAEAC2596F6251FEBC22C835999BC8" box="[289,707,1484,1509]" italics="true" pageId="21" pageNumber="460">Voyage dans l’Amérique méridionale</emphasis>
, series published between 
<date id="8C601010596F6251FC6C22C933C29BC9" box="[1009,1176,1484,1509]" pageId="21" pageNumber="460" value="1835" valueMax="1847">
1835 and 
<quantity id="3F269B35596F6251FBFD22C933C29BC9" box="[1120,1176,1484,1508]" metricMagnitude="1" metricUnit="m" metricValue="4.69138" pageId="21" pageNumber="460" unit="in" value="1847.0">1847</quantity>
</date>
in 90 livrasions, seven text volumes and two volumes of plates by Alcide d’Orbigny (Evenhuis 1997). In the sixth volume, Auguste Gaspard Brullé was responsible for the species published on pages 1–60, while Émile Blanchard was for those on pages 61–222. Thus, Blanchard should be regarded as the author of these two species of 
<taxonomicName id="3FDE4D53596F6251FB7B21323201987D" box="[1254,1371,1591,1616]" class="Insecta" family="Scarabaeidae" genus="Gromphas" kingdom="Animalia" order="Coleoptera" pageId="21" pageNumber="460" phylum="Arthropoda" rank="genus">
<emphasis id="CAAAEAC2596F6251FB7B21323201987D" box="[1254,1371,1591,1616]" italics="true" pageId="21" pageNumber="460">Gromphas</emphasis>
</taxonomicName>
. The publication date usually cited for these species, 
<emphasis id="CAAAEAC2596F6251FD27215835869859" box="[698,732,1629,1652]" italics="true" pageId="21" pageNumber="460">i.e.</emphasis>
, 1843, is incorrect. According Sherborn &amp; Woodward (1901) and Evenhuis (1997), livrasion 80, containing pages 169–184 of the sixth volume, was published in 1846.
</paragraph>
<paragraph id="F86136D0596F6251FF5A21A1341E9928" blockId="21.[151,1436,151,1977]" pageId="21" pageNumber="460">
Martinez (1959) reported 
<taxonomicName id="3FDE4D53596F6251FE7021A035399890" box="[493,611,1700,1725]" class="Insecta" family="Scarabaeidae" genus="Gromphas" kingdom="Animalia" order="Coleoptera" pageId="21" pageNumber="460" phylum="Arthropoda" rank="species" species="dichroa">
<emphasis id="CAAAEAC2596F6251FE7021A035399890" box="[493,611,1700,1725]" italics="true" pageId="21" pageNumber="460">G. dichroa</emphasis>
</taxonomicName>
from 
<collectingCountry id="80C97640596F6251FD3621A0344F9890" box="[683,789,1701,1725]" name="Paraguay" pageId="21" pageNumber="460">Paraguay</collectingCountry>
. Although we do not know specimens from this country, we examined specimens collected in two 
<collectingCountry id="80C97640596F6251FDEF21C235BB98CD" box="[626,737,1735,1760]" name="Argentina" pageId="21" pageNumber="460">Argentine</collectingCountry>
provinces bordering 
<collectingCountry id="80C97640596F6251FC4821CD336498CD" box="[981,1086,1736,1760]" name="Paraguay" pageId="21" pageNumber="460">Paraguay</collectingCountry>
, Misiones and Corrientes. It is thus possible that this species is actually present in 
<collectingCountry id="80C97640596F6251FD4B21E8341A9928" box="[726,832,1773,1797]" name="Paraguay" pageId="21" pageNumber="460">Paraguay</collectingCountry>
.
</paragraph>
<paragraph id="F86136D0596F6251FF5A20153317995D" blockId="21.[151,1436,151,1977]" pageId="21" pageNumber="460">
None of the females of 
<taxonomicName id="3FDE4D53596F6251FE512015351B9905" box="[460,577,1807,1832]" class="Insecta" family="Scarabaeidae" genus="Gromphas" kingdom="Animalia" order="Coleoptera" pageId="21" pageNumber="460" phylum="Arthropoda" rank="species" species="dichroa">
<emphasis id="CAAAEAC2596F6251FE512015351B9905" box="[460,577,1807,1832]" italics="true" pageId="21" pageNumber="460">G. dichroa</emphasis>
</taxonomicName>
observed in this study had intact protarsi, all were broken and had only three tarsomeres or less or had the apex of apical tarsomere broken off, which made it impossible to observe its actual morphology (
<emphasis id="CAAAEAC2596F6251FEAD205C3602995D" box="[304,344,1881,1904]" italics="true" pageId="21" pageNumber="460">e.g.</emphasis>
, if the spiniform projection seen in 
<taxonomicName id="3FDE4D53596F6251FD77205D34DB995D" box="[746,897,1880,1904]" class="Insecta" family="Scarabaeidae" genus="Gromphas" kingdom="Animalia" order="Coleoptera" pageId="21" pageNumber="460" phylum="Arthropoda" rank="species" species="amazonica">
<emphasis id="CAAAEAC2596F6251FD77205D34DB995D" box="[746,897,1880,1904]" italics="true" pageId="21" pageNumber="460">G. amazonica</emphasis>
</taxonomicName>
is present or not).
</paragraph>
<paragraph id="F86136D0596F6251FF5A207935C89994" blockId="21.[151,1436,151,1977]" pageId="21" pageNumber="460">
<emphasis id="CAAAEAC2596F6251FF5A2079361799B8" bold="true" box="[199,333,1916,1941]" pageId="21" pageNumber="460">Bionomics:</emphasis>
There are no bionomic studies published on 
<taxonomicName id="3FDE4D53596F6251FCC62078348899B8" box="[859,978,1916,1941]" class="Insecta" family="Scarabaeidae" genus="Gromphas" kingdom="Animalia" order="Coleoptera" pageId="21" pageNumber="460" phylum="Arthropoda" rank="species" species="dichroa">
<emphasis id="CAAAEAC2596F6251FCC62078348899B8" box="[859,978,1916,1941]" italics="true" pageId="21" pageNumber="460">G. dichroa</emphasis>
</taxonomicName>
. The specimens examined in this study were collected between October and January.
</paragraph>
<caption id="ACA16658596C6252FF0A20403551999A" httpUri="https://zenodo.org/record/247631/files/figure.png" pageId="22" pageNumber="461" targetBox="[155,1436,198,1838]" targetPageId="22">
<paragraph id="F86136D0596C6252FF0A20403551999A" blockId="22.[151,1437,1861,1975]" pageId="22" pageNumber="461">
<emphasis id="CAAAEAC2596C6252FF0A2040346D9976" bold="true" box="[151,823,1861,1883]" pageId="22" pageNumber="461">
FIGURES 13–15. 13: 
<taxonomicName id="3FDE4D53596C6252FE16204035A09976" authority="Bates, 1870" authorityName="Bates" authorityYear="1870" box="[395,762,1861,1883]" class="Insecta" family="Scarabaeidae" genus="Gromphas" kingdom="Animalia" order="Coleoptera" pageId="22" pageNumber="461" phylum="Arthropoda" rank="species" species="amazonica">
<emphasis id="CAAAEAC2596C6252FE16204035299976" bold="true" box="[395,627,1861,1883]" italics="true" pageId="22" pageNumber="461">Gromphas amazonica</emphasis>
Bates, 1870
</taxonomicName>
. 13a)
</emphasis>
Dark blue specimen. 
<emphasis id="CAAAEAC2596C6252FBBC2040330A9976" bold="true" box="[1057,1104,1861,1883]" pageId="22" pageNumber="461">13b)</emphasis>
Reddish-brown specimen. 
<emphasis id="CAAAEAC2596C6252FAF2204032C19976" bold="true" box="[1391,1435,1861,1883]" pageId="22" pageNumber="461">13c)</emphasis>
Dark green specimen. 
<emphasis id="CAAAEAC2596C6252FE1A206134219957" bold="true" box="[391,891,1892,1914]" pageId="22" pageNumber="461">
14: 
<taxonomicName id="3FDE4D53596C6252FE29206134649957" authority="Blanchard, 1846" authorityName="Blanchard" authorityYear="1846" box="[436,830,1892,1914]" class="Insecta" family="Scarabaeidae" genus="Gromphas" kingdom="Animalia" order="Coleoptera" pageId="22" pageNumber="461" phylum="Arthropoda" rank="species" species="dichroa">
<emphasis id="CAAAEAC2596C6252FE29206135269957" bold="true" box="[436,636,1892,1914]" italics="true" pageId="22" pageNumber="461">Gromphas dichroa</emphasis>
Blanchard, 1846
</taxonomicName>
. 14a)
</emphasis>
Red-blue specimen. 
<emphasis id="CAAAEAC2596C6252FBC3206133D79957" bold="true" box="[1118,1165,1892,1914]" pageId="22" pageNumber="461">14b)</emphasis>
Intermediate yellow-blue specimen. 
<emphasis id="CAAAEAC2596C6252FE982087366B99B5" bold="true" box="[261,305,1922,1944]" pageId="22" pageNumber="461">14c)</emphasis>
Green specimen. 
<emphasis id="CAAAEAC2596C6252FE70208734F499B5" bold="true" box="[493,942,1922,1944]" pageId="22" pageNumber="461">
15: 
<taxonomicName id="3FDE4D53596C6252FD8A2087342999B5" authority="Harold, 1869" authorityName="Harold" authorityYear="1869" box="[535,883,1922,1944]" class="Insecta" family="Scarabaeidae" genus="Gromphas" kingdom="Animalia" order="Coleoptera" pageId="22" pageNumber="461" phylum="Arthropoda" rank="species" species="inermis">
<emphasis id="CAAAEAC2596C6252FD8A2087358699B5" bold="true" box="[535,732,1922,1944]" italics="true" pageId="22" pageNumber="461">Gromphas inermis</emphasis>
Harold, 1869
</taxonomicName>
. 15a)
</emphasis>
Coppery specimen. 
<emphasis id="CAAAEAC2596C6252FB1E208733E999B5" bold="true" box="[1155,1203,1922,1944]" pageId="22" pageNumber="461">15b)</emphasis>
Dark green specimen. 
<emphasis id="CAAAEAC2596C6252FF0A20A43799999A" bold="true" box="[151,195,1953,1975]" pageId="22" pageNumber="461">15c)</emphasis>
Black specimen. Scale = 5 mm.
</paragraph>
</caption>
<paragraph id="F86136D0596D6253FF5A279D341C9FA5" blockId="23.[151,1437,151,717]" pageId="23" pageNumber="462">
It is probable that 
<taxonomicName id="3FDE4D53596D6253FE38279D35449E9D" box="[421,542,151,176]" class="Insecta" family="Scarabaeidae" genus="Gromphas" kingdom="Animalia" order="Coleoptera" pageId="23" pageNumber="462" phylum="Arthropoda" rank="species" species="dichroa">
<emphasis id="CAAAEAC2596D6253FE38279D35449E9D" box="[421,542,151,176]" italics="true" pageId="23" pageNumber="462">G. dichroa</emphasis>
</taxonomicName>
has always been a rare species, but it is interesting to note that all the six specimens known to us with collection date available were collected between the years 
<date id="8C601010596D6253FBE627B9327D9EF9" box="[1147,1319,188,213]" pageId="23" pageNumber="462" value="1922" valueMax="1947">1922 and 1947</date>
. It is truly remarkable that no specimen has been found over the past 65 years since 
<taxonomicName id="3FDE4D53596D6253FC7627E533389ED5" box="[1003,1122,223,248]" class="Insecta" family="Scarabaeidae" genus="Gromphas" kingdom="Animalia" order="Coleoptera" pageId="23" pageNumber="462" phylum="Arthropoda" rank="species" species="dichroa">
<emphasis id="CAAAEAC2596D6253FC7627E533389ED5" box="[1003,1122,223,248]" italics="true" pageId="23" pageNumber="462">G. dichroa</emphasis>
</taxonomicName>
is distributed in one of the most intensely collected regions of South 
<collectingCountry id="80C97640596D6253FDF12601358A9F30" box="[620,720,260,285]" name="United States of America" pageId="23" pageNumber="462">America</collectingCountry>
. As nothing is known about the natural history of this species, it is difficult to surmise the reasons for the putative disappearance of 
<taxonomicName id="3FDE4D53596D6253FC02262D334C9F6D" box="[927,1046,295,320]" class="Insecta" family="Scarabaeidae" genus="Gromphas" kingdom="Animalia" order="Coleoptera" pageId="23" pageNumber="462" phylum="Arthropoda" rank="species" species="dichroa">
<emphasis id="CAAAEAC2596D6253FC02262D334C9F6D" box="[927,1046,295,320]" italics="true" pageId="23" pageNumber="462">G. dichroa</emphasis>
</taxonomicName>
, but it is possible that the intense human pressure in the Southern Cone (which includes southern 
<collectingCountry id="80C97640596D6253FCF3264934EF9F48" box="[878,949,332,357]" name="Brazil" pageId="23" pageNumber="462">Brazil</collectingCountry>
, 
<collectingCountry id="80C97640596D6253FC5F264833719F48" box="[962,1067,333,357]" name="Paraguay" pageId="23" pageNumber="462">Paraguay</collectingCountry>
, 
<collectingCountry id="80C97640596D6253FBAA264833C09F48" box="[1079,1178,333,357]" name="Uruguay" pageId="23" pageNumber="462">Uruguay</collectingCountry>
, 
<collectingCountry id="80C97640596D6253FB3B264932439F48" box="[1190,1305,332,357]" name="Argentina" pageId="23" pageNumber="462">Argentina</collectingCountry>
, and 
<collectingCountry id="80C97640596D6253FACB264932CE9F48" box="[1366,1428,332,357]" name="Chile" pageId="23" pageNumber="462">Chile</collectingCountry>
) has acted negatively on what were already small populations.
</paragraph>
<paragraph id="F86136D0596D6253FF5A2691336B9FD8" blockId="23.[151,1437,151,717]" pageId="23" pageNumber="462">
<emphasis id="CAAAEAC2596D6253FF5A269136AA9F80" bold="true" box="[199,496,404,429]" pageId="23" pageNumber="462">Geographic distribution:</emphasis>
Chacoan subregion: Chaco and Pampa. 
<emphasis id="CAAAEAC2596D6253FC2E2691337D9F80" bold="true" box="[947,1063,404,429]" pageId="23" pageNumber="462">
<collectingCountry id="80C97640596D6253FC2E269133789F80" box="[947,1058,404,429]" name="Brazil" pageId="23" pageNumber="462">BRAZIL</collectingCountry>
:
</emphasis>
Rio Grande do Sul: Cerro Largo, Porto Alegre. 
<emphasis id="CAAAEAC2596D6253FEDE26B235FC9FFD" bold="true" box="[323,678,439,464]" pageId="23" pageNumber="462">
<collectingCountry id="80C97640596D6253FEDE26B236849FFD" box="[323,478,439,464]" name="Paraguay" pageId="23" pageNumber="462">PARAGUAY</collectingCountry>
. 
<collectingCountry id="80C97640596D6253FE6F26B235F89FFD" box="[498,674,439,464]" name="Argentina" pageId="23" pageNumber="462">ARGENTINA</collectingCountry>
:
</emphasis>
Misiones: Concepción (Santa Maria). Corrientes: 
<collectingCountry id="80C97640596D6253FA9A26BD32C29FFD" box="[1287,1432,440,464]" name="Sao Tome and Principe" pageId="23" pageNumber="462">Santo Tomé</collectingCountry>
. Buenos Aires. 
<emphasis id="CAAAEAC2596D6253FEA126D9368A9FD8" bold="true" box="[316,464,476,501]" pageId="23" pageNumber="462">
<collectingCountry id="80C97640596D6253FEA126D936909FD8" box="[316,458,476,501]" name="Uruguay" pageId="23" pageNumber="462">URUGUAY</collectingCountry>
:
</emphasis>
Tacuarembó, Soriano, Flórida, Montevidéu. (
<figureCitation id="60E52A55596D6253FC4D26D9337E9FD8" box="[976,1060,476,501]" 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="23" pageNumber="462">Fig. 66</figureCitation>
).
</paragraph>
</subSubSection>
<subSubSection id="B0C4655B596D6253FF5A26FA36089CE1" pageId="23" pageNumber="462" type="materials_examined">
<paragraph id="F86136D0596D6253FF5A26FA36089CE1" blockId="23.[151,1437,151,717]" pageId="23" pageNumber="462">
<emphasis id="CAAAEAC2596D6253FF5A26FA36EF9C35" bold="true" box="[199,437,511,536]" pageId="23" pageNumber="462">Material examined:</emphasis>
<specimenCount id="EED8FD59596D6253FE22250535429C35" box="[447,536,511,536]" pageId="23" pageNumber="462" type="male">3 males</specimenCount>
and 
<specimenCount id="EED8FD59596D6253FDC826FA35929C35" box="[597,712,511,536]" pageId="23" pageNumber="462" type="female">6 females</specimenCount>
. 
<emphasis id="CAAAEAC2596D6253FD4826FA34D09C35" bold="true" box="[725,906,511,536]" pageId="23" pageNumber="462">
<collectingCountry id="80C97640596D6253FD4826FA34DF9C35" box="[725,901,511,536]" name="Argentina" pageId="23" pageNumber="462">ARGENTINA</collectingCountry>
:
</emphasis>
BUENOS AIRES: Anchorena, 
<date id="8C601010596D6253FB6126FA320B9C35" box="[1276,1361,511,536]" pageId="23" pageNumber="462" value="1935-10">X.1935</date>
, M. J. Viana col.— 
<specimenCount id="EED8FD59596D6253FEB2252136219C10" box="[303,379,548,573]" pageId="23" pageNumber="462" type="male">1 male</specimenCount>
(CMNC). CORRIENTES: 
<collectingCountry id="80C97640596D6253FD5F2521340B9C11" box="[706,849,548,572]" name="Sao Tome and Principe" pageId="23" pageNumber="462">Santo Tomé</collectingCountry>
, 
<date id="8C601010596D6253FCFF252134919C11" box="[866,971,548,572]" pageId="23" pageNumber="462" value="1922-12">XII.1922</date>
, without collector— 
<specimenCount id="EED8FD59596D6253FB5F252132799C10" box="[1218,1315,548,573]" pageId="23" pageNumber="462" type="female">1 female</specimenCount>
(CMNC). MISIONES: Concepción, Santa Maria, 
<date id="8C601010596D6253FDCF254D35EB9C4D" box="[594,689,584,608]" pageId="23" pageNumber="462" value="1947-11">XI.1947</date>
, M. J. Viana col.— 
<specimenCount id="EED8FD59596D6253FC10254234899C4D" box="[909,979,583,608]" pageId="23" pageNumber="462" type="male">1 male</specimenCount>
(MLPA). 
<emphasis id="CAAAEAC2596D6253FBDB254233E09C4D" bold="true" box="[1094,1210,583,608]" pageId="23" pageNumber="462">
<collectingCountry id="80C97640596D6253FBDB254233EF9C4D" box="[1094,1205,583,608]" name="Brazil" pageId="23" pageNumber="462">BRAZIL</collectingCountry>
:
</emphasis>
RIO GRANDE DO SUL: Cerro Largo (former Serro Azul), 
<date id="8C601010596D6253FDEE256935E19CA9" box="[627,699,620,644]" pageId="23" pageNumber="462" value="1931-01">I.1931</date>
(or 1933?), without collector— 
<specimenCount id="EED8FD59596D6253FBAA256933DB9CA8" box="[1079,1153,620,645]" pageId="23" pageNumber="462" type="male">1 male</specimenCount>
(CEMT); Porto Alegre, 
<date id="8C601010596D6253FF0A259537B79C85" box="[151,237,656,680]" pageId="23" pageNumber="462" value="1928-10">X.1928</date>
, without collector— 
<specimenCount id="EED8FD59596D6253FE47258A356C9C85" box="[474,566,655,680]" pageId="23" pageNumber="462" type="female">1 female</specimenCount>
(MZSP). 
<emphasis id="CAAAEAC2596D6253FD35258A34669C85" bold="true" box="[680,828,655,680]" pageId="23" pageNumber="462">
<collectingCountry id="80C97640596D6253FD35258A346C9C85" box="[680,822,655,680]" name="Uruguay" pageId="23" pageNumber="462">URUGUAY</collectingCountry>
:
</emphasis>
without date and collector— 
<specimenCount id="EED8FD59596D6253FB15258A33A89C85" box="[1160,1266,655,680]" pageId="23" pageNumber="462" type="female">3 females</specimenCount>
(MZSP) and 
<specimenCount id="EED8FD59596D6253FA13259537B99CE0" pageId="23" pageNumber="462" type="female">1 female</specimenCount>
(CEMT).
</paragraph>
</subSubSection>
</treatment>
</document>