Gromphas Brullé, 1837, Brulle, 1837
Cupello, Mario & Vaz-De-Mello, Fernando Z., 2013, Taxonomic revision of the South American dung beetle genus Gromphas Brullé, 1837 (Coleoptera: Scarabaeidae: Scarabaeinae: Phanaeini: Gromphadina), Zootaxa 3722 (4), pp. 439-482: 443-448
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|Gromphas Brullé, 1837|
Copris (Gromphas) Brullé, 1837: 283 , 298, 304.
Gromphas: Blanchard 1846: 100 ; Erichson 1847: 760; Lacordaire 1856: 95, 100; Harold 1868: 81; Harold 1869 a: 1016; Bates 1870: 175; Lacordaire & Chapuis 1876: 276, 365; Karsch 1887: 1; Kolbe 1905: 550; Packard 1909: 101; Gillet 1911: 80; Fabre 1919: 244, 247, 256; d’Olsoufieff 1924: 9, 12, 17, 20, 58, plate IX; Luederwaldt 1929: 614; Pessoa 1934: 282; Pessoa & Lane 1941: 470; Blackwelder 1944: 208; Barattini & Saenz 1960: 1; Barattini & Saenz 1964: 173; Halffter & Matthews 1966: 17, 30, 122, 131, 133, 194, 209, 257; Vulcano & Pereira 1967: 565; Edmonds 1972: 816; Halffter & Edmonds 1982: 52, 59, 65, 85, 86, 137; Zunino 1983: 22; 1985 a: 21; 1985 b: 104; Martínez 1989: 50, 67; Hanski & Cambefort 1991: 468; Walsh & Gandolfo 1996: 587 Fig. 7 View FIGURES 7 – 10 ; Medina & Lopera-Toro 2000: 300; Vaz-de-Mello 2000: 186; Davis et al. 2002: 1240; Philips et al. 2004 a: 55–56; 2004 b: 43, 51–54, 58; Ocampo & Hawks 2006: 558; Noriega et al. 2008: 131; Scholtz et al. 2009: 243, 246–249, 284; Gillet et al. 2010: 4; Noriega et al. 2010: 454; Bouchard et al. 2011: 245; Vaz-de-Mello et al. 2011: 5, 9, 17, 24, 31, 39, 44; Figueroa et al. 2012: 1; Gatty et al. 2012: 1; Korasaki et al. 2012: 427; Cupello 2013: 15–16.
Type species: Gromphas dichroa Blanchard, 1846 ; present designation.
Etymology: from Greek, an old sow (feminine) (Brullé 1837, Brown 1956).
Redescription: Head: Clypeus with apex truncate ( Fig. 26 View FIGURES 24 – 28 ) or with two small lobes; if lobed, then margin between apical lobes and genae rounded ( Figs. 24–25 View FIGURES 24 – 28 ) or with another pair of lobes ( Figs. 27–28 View FIGURES 24 – 28 ). Junction between clypeus and genae notched; margin of clypeus and genae folded up or not. Genae with granular tegument; with metallic reflections or not. Cephalic projection a high antero-posteriorly compressed horn ( Figs. 29–30 View FIGURES 29 – 33 ), or only slightly arched upward and emarginate ( Figs. 31–32 View FIGURES 29 – 33 ) or apically truncate ( Fig. 33 View FIGURES 29 – 33 ). Dorsal portion of eyes smaller than ventral. Clypeal process a transverse ridge with rounded margin.
Thorax: Pronotum, at its greatest width, greater than elytra; anterior margin not interrupted behind eyes; tegument granulate on sides and granulate or smooth in posteromedian region; with metallic reflections or not. Posterior pronotal fossae present or not. Pronotal prominence present or not.
Propleura not excavated anteriorly. Mesepisternum, metepisternum and outer sides of metasternum with glabrous and granulate tegument (Figs. 16–20). Metasternum wide, densely punctate or not; antero-median angle flat or convex and very projected ( Figs. 49–52 View FIGURES 49 – 52 ); area lateral to antero-median angle sloped and granulate.
Legs: Protibiae with four rounded teeth, proximalmost less developed but always present ( Figs. 21–23 View FIGURES 21 – 23 ). Dorsal surface of protibia with longitudinal carina extended from base to apex and with thin branches into three apical teeth ( Fig. 21 View FIGURES 21 – 23 ). Ventral longitudinal carina of protibia simple in both sexes ( Figs. 22, 23 View FIGURES 21 – 23 a) or with a row of tubercles in its basal half in males (Figs. 20 d, 23 b). Protibial spur strongly expanded ( Figs. 21, 23 View FIGURES 21 – 23 , 45 View FIGURES 34 – 48. 34 ) or only slightly bent apically ( Figs. 22 View FIGURES 21 – 23 , 46 View FIGURES 34 – 48. 34 ). Inner apical angle of protibia with a tuft of long setae sexually dimorphic ( Figs. 23 View FIGURES 21 – 23 ) or not ( Fig. 22 View FIGURES 21 – 23 ); in males of G. amazonica , inner apical angle with a long non-articulate spur ( Fig. 21 View FIGURES 21 – 23 b, white arrow); in males of the other species, inner apical angle only with a tiny and almost imperceptible tubercle ( Fig. 23 View FIGURES 21 – 23 b, white arrow). Protarsus absent in males and present with four tarsomeres in females, its length equals one third to one half of protibial spur total length; apical protarsomere with ( Fig. 45 View FIGURES 34 – 48. 34 ) or without ( Figs. 22, 23 View FIGURES 21 – 23 b, 46) a spiniform projection in apex.
Mesotibiae with three slightly pointed teeth on apical margin. Metatibiae very broad and robust ( Fig. 41 View FIGURES 34 – 48. 34 ) or very thin ( Fig. 42 View FIGURES 34 – 48. 34 ), with four slightly pointed teeth on apical margin and apical spur clearly curved ( Fig. 39 View FIGURES 34 – 48. 34 ) or straight ( Fig. 40 View FIGURES 34 – 48. 34 ) apically. Mesotarsi and metatarsi with five tarsomeres progressively shorter from basal outward, outer margin with rows of rigid setae ( Figs. 47–48 View FIGURES 34 – 48. 34 ); apical mesotarsomeres and metatarsomeres spiniform.
Elytra: Outer margins of elytra, seen from above, rounded. Each elytron with 10 very fine and superficial, simple or carinulate striae; eighth stria present only on the apical half of elytra; ninth and tenth striae fused and distinct only at the apex. Sutural margin with glossy sheen strongly constrasting with remaining elytral disc of silky appearance; tegument smooth or punctate. Humeral angle with black glossy sheen. Epipleura strongly curved and wide from base of elytron to metacoxa; remainder length horizontal and narrowed (Figs. 16–20).
Abdomen: Pygidium flattened; basal margin complete and never interrupted by groove of propygidium ( Fig. 35 View FIGURES 34 – 48. 34 ) or basal margin completely absent ( Fig. 36 View FIGURES 34 – 48. 34 ). Abdomen with six visible sternites (i.e., six ventrites); sixth ventrite strongly narrowed at middle in males ( Fig. 34 View FIGURES 34 – 48. 34 a) and very wide at middle in females ( Fig. 34 View FIGURES 34 – 48. 34 b).
Genitalia: Aedeagus: Parameres coated by a wide area of short setae and white tegument ( Figs. 53–56 View FIGURES 53 – 56 ). Phallobase with membranous apex of variable width; in ventral view, basal margin with median incision followed by a midlongitudinal shallow groove (groove represented by dashed lines in Figs. 53 View FIGURES 53 – 56 c, 54 c, 55 c, 56 c). Internal sac as in Fig. 62 View FIGURES 57 – 62 ; single medial sclerite with variable curvature, with ( Fig. 60 View FIGURES 57 – 62 ) or without a spiniform projection at one end. Spermatheca: C-shaped; distal branch with pointed end and thinner than basal branch; inner cuticle smooth and without ornamentation; spermathecal duct very long. Spermathecal gland very large and well developed.
Geographic distribution: South America east of the Andes, from the Venezuelan coast south to the Argentine province of Buenos Aires; absent in the large areas of Guiana Shield and Caatinga in northeastern Brazil ( Fig. 66 View FIGURE 66 ).
Taxonomic discussion: As stated previously, the genus closest to Gromphas is certainly Oruscatus . They are distinguished primarily by the shape of the outer margin of elytra (rounded in Gromphas and parallel in Oruscatus ), the presence of the metepisternal tab (absent in Gromphas and present, though very weakly indicated, in Oruscatus ), the shape of epipleura (curved and wide at the base of elytra in Gromphas [Figs. 16–20] and entirely horizontal and narrow in Oruscatus ) and various characteristics of sexual dimorphism: protarsus absent in both sexes of Oruscatus and present with four tarsomeres in females of Gromphas ; prolegs longer in males than in females of Oruscatus and without this sexual dimorphism in Gromphas ; and the antero-median angle of metasternum sexually dimorphic in Oruscatus (strongly produced and lobate in males and simply acuminated in females) and not in Gromphas (although the interspecific variation of this character [ Figs. 49–52 View FIGURES 49 – 52 ]). In addition, in no species of Gromphas is the antero-median angle of metasternum so developed and sharp as in Oruscatus .
The pygidium of four species of Gromphas is not margined basally (or, in some specimens, the basal margin is marked only by a very thin and almost imperceptible line), whereas only G. dichroa has the basal margin complete and never interrupted by the groove of propygidium. In Oruscatus , the basal margin is always complete as G. dichroa . Consequently, step 54 of the key presented by Vaz-de-Mello et al. (2011) should be corrected by deleting the difference between these genera regarding the basal margin of pygidium or by adding a note excepting G. dichroa .
Taxonomic history and nomenclatural discussion: Gromphas has a long and confusing taxonomic history. Traditionally, the genus is taken as described by Brullé in 1834 and having the following species and their synonyms (dates are as have been considered until now): G. aeruginosa (Perty, 1830) (= G. lacordairei Blanchard, 1843 ; G. lemoinei Waterhouse, 1891 ); G. amazonica Bates, 1870 ; G. dichroa Blanchard, 1843 and G. lacordairei Brullé, 1834 (= G. inermis Harold, 1869 ). Nevertheless, when we reviewed the literature, we found several inconsistencies in this panorama; many of the errors refer to the dates of publication of taxa, but the most sensitive involve the authorship and the availability of some names. In the following, we review the historical development of the taxonomy of Gromphas and its species, discuss its nomenclatural problems and finally propose a satisfactory solution to them in accordance with the current edition of the International Code of Zoological Nomenclature (International Commission on Zoological Nomenclature 1999).
The first author who cited the name “ Gromphas ” was Dejean (1833), who proposed this new genus name for a single unpublished species, “ Lacordairei Dej.” from Buenos Aires, Argentina. However, Dejean provided no descriptions of his new taxa, limiting only to name them. The dates of publication of Dejean’s catalogs were discussed by Madge (1988) and here we use the dates proposed by him.
The first description of Gromphas was finally given by Brullé (1837, see comments on the date of this publication below), who regarded the taxon as one of the seven subgenera of Copris . Although he said that this subgenus possess only one species (“ Ce sous-genre se compose d’une seule espèce ...”), Brullé did not explicitly mention its name; he also vaguely ascribed the authorship of Gromphas to Dejean (“ Gromphas Dej. ”) and said that the subgenus was still unpublished (perhaps referring to the fact that Dejean failed to describe it).
Blanchard (1846, see the discussion of this date in the comments on G. dichroa ) treated Gromphas as a genus and recognized that Brullé, despite having described the taxon, did not include any species in it (“ Le genre Gromphas a été caractérisé par M. Brullé, Histoire des Insectes, tome 6, p. 304; mais cet entomologiste n’en décrit aucune espèce ”). Blanchard thus described two new species, the first included and described in Gromphas : G. dichroa from Montevideo, Uruguay and G. lacordairei from Santa Cruz de la Sierra, Bolivia.
Lacordaire (1856) redescribed the genus in more detail and stated that the species collected by him in Buenos Aires, cited as “ G. lacordairei ”, and G. dichroa could be synonyms (“ Je crois que ces deux espèces n’en font qu’une ”). Five pages later, he said that two species described by Perty (1830) from Brazil, Onitis aeruginosus and O. chalcomelas , should form a new genus related to Gromphas .
Harold (1859) did not follow Lacordaire and transferred O. chalcomelas and O. aeruginosus to different genera: the former to Phanaeus and the latter to Gromphas . Moreover, Harold also said that Gromphas aeruginosa ( O. aeruginosus ) would be a synonym of G. lacordairei , a species regarded by him as having been collected by Lacordaire, named by Dejean and described by Blanchard; so there would be only one “ G. lacordairei ” for him. Ten years later, however, Harold (1869 a) recognized two distinct names “ G. lacordairei ”: one described by Blanchard and considered a junior synonym of G. aeruginosa , and the other named by Dejean and synonymized with the still unpublished “ G. inermis ”. In the same year, Harold (1869 b) finally described G. inermis as an alternative name of the never described “ G. lacordairei Dejean ”. Having now examined the type specimens of Blanchard, Harold (1869 b) also concluded that the synonymy between G. aeruginosa and G. lacordairei Blanchard was correct.
Burmeister (1874) did not agree with Harold’s (1869 b) conclusions and modified the nomenclature: he thought that the beetle collected by Lacordaire in Buenos Aires was named as “ G. lacordairii ” by Dejean and later described under this name by Brullé; in turn, Blanchard was confused and described under this same name a synonym of G. aeruginosa (thus a homonym of Dejean’s name). Hence, in the view of Burmeister, the name “ G. lacordairii Dejean ” was valid, and therefore Harold had modified the species name to G. inermis unjustly. Burmeister also described the sexual dimorphism of “ G. lacordairii Dejean ” and presented its distribution.
From then until the early 20 th century, authors were divided on adopting as the valid name G. inermis (e.g., Preudhomme-de-Borre 1886; Judulien 1899; Gahan & Arrow 1903; Heyne & Taschenberg 1908), “ G. lacordairei Dejean ” (e.g., Tremoleras 1910) or “ G. lacordairei Brullé ” (e.g., Bruch 1911). In the catalogs of Gillet (1911) and Blackwelder (1944) and in the revision of d’Olsoufieff (1924), “ G. lacordairei Brullé, 1834 ” was taken as valid and senior synonym of G. inermis , in which were followed by all subsequent authors until now.
Two more species of Gromphas were described in the 19 th century: G. amazonica Bates, 1870 from the Brazilian Amazon and G. lemoinei Waterhouse, 1891 from Venezuela. However, the latter species was soon thereafter treated in a dubious way as a variety of G. aeruginosa by Gillet (1911) and finally clearly synonymized with this species by d’Olsoufieff (1924), remaining so until now (see discussion of G. lemoinei ). Finally, d’Olsoufieff (1924) described a variety ( var. bipunctata ) of “ G. lacordairei Brullé ” (see discussion of G. inermis ).
The names associated with Gromphas that were published by Dejean (1833) are unavailable because they fail to comply with the provisions of Article 12 of the Code (ICNZ 1999), which requires, among other points, the description of new taxa, or, for the genus, at least the inclusion of a valid species name. “ Gromphas Dejean ” and “ G. lacordairei Dejean ” thus should be regarded as nomina nuda (see Bousquet & Bouchard [2013 a, b] for a discussion on the genus-group names published in the Dejean’s catalogs). Brullé (1837) was the first to properly describe Gromphas and therefore has its authorship. Despite failing to include any species name in Gromphas, Brullé consistently applied the Principle of Binomial Nomenclature throughout his work and therefore his description of Gromphas must be accepted as available (Article 11.4.1). However, because he did not cite the name “ G. lacordairei ” anywhere in his work, Brullé obviously cannot have the authorship of this name.
The first author to properly describe a species under the name Gromphas lacordairei was Blanchard (1846). We agree with the synonymy between G. aeruginosa (Perty) and G. lacordairei Blanchard first proposed by Harold (1859) (see also the discussion of G. aeruginosa ). Burmeister (1874) actually recognized the existence of G. lacordairei Blanchard and its synonymy with G. aeruginosa . Nevertheless, he also considered that the nomina nuda of Dejean would be valid and that “ G. lacordairii Dejean ” would be a senior synonym of G. inermis Harold and therefore valid. Since he described the species credited to Dejean and explicitly stated that considered G. lacordairei Blanchard and “ G. lacordairii Dejean ” distinguished names referring to different species (“ G. lacordairii Dejean ” expressly cited as a synonym of G. inermis ), Burmeister inadvertently gave availability to a new name G. lacordairii Burmeister, 1874 , a junior objective synonym of G. inermis . However, as Burmeister did not followed the original spelling of Dejean (“ lacordairei ”) and changed it to “ lacordairii ”, G. lacordairii and G. lacordairei Blanchard are not homonyms (see Article 58 of the Code). Thus, G. lacordairei Blanchard = G. aeruginosa (Perty) and G. l a co rd ai r i i Burmeister = G. inermis Harold. All post- 1874 citations of “ G. lacordairei Dejean ” and “ G. lacordairei Brullé ” refer, in fact, to G. lacordairii Burmeister.
As Gromphas was described by Brullé (1837) without any included species, one of the two species described by Blanchard (1846) should be designated as the type species of Gromphas , since they were the first included in the genus (Article 67.2.2). Although Recommendation 69 A. 1 of the Code recommends choosing the most common species, that is G. lacordairei Blanchard , the potential confusion created by designating as the type a species synonymized with another older, leads us to prefer designating the other species described by Blanchard, the rare G. dichroa , as the type species of Gromphas .
After the revalidation of G. lemoinei and having solved the problem with the “ var. bipunctata ” (see discussion of G. inermis ), the current status of the nomenclature is as follows: Gromphas Brullé, 1837 : G. aeruginosa (Perty, 1830) (= G. l a co rd ai re i Blanchard, 1846); G. amazonica Bates, 1870 ; G. dichroa Blanchard, 1846 (type species); G. inermis Harold, 1869 (= G. lacordairii Burmeister, 1974 ; = G. lacordairei bipunctata d’Olsoufieff, 1924); and G. lemoinei Waterhouse, 1891 . Table 1 summarizes the current situation of names proposed for Gromphas cited in this taxonomic history.
Comments: The year of the original publication of Gromphas by Brullé (i.e., of the third treatise of Coleoptera and sixth volume of the series “ Histoire Naturelle des Insectes ”, by Audouin & Brullé) is controversial. Regarding the taxonomic history of Gromphas, Burmeister (1874) was the first author to cite 1834 as the year of publication of this genus. This date was not questioned until 2011, when Bouchard et al. (2011) cited 1837 as the year of publication of Gromphas . These authors did not cite the sixth volume of “ Histoire Naturelle des Insectes ” in their references, but dated, with some doubt, the publication of volumes 4 and 5, respectively, as 1834 and 1835. Figueroa et al. (2012) also agreed on 1837 as being the date of publication of Gromphas . Smith (2006) did not cite Gromphas directly, but dated the sixth volume of the “ Histoire Naturelle des Insectes ” in his references as 1837. In fact, the date printed on the title page of the original book is 1837, which indicates that the publication of volume 6, even if it was issued in separate fascicles, at least began in that year, which rules out 1834. In the absence of evidence to the contrary (Article 21.2 of ICZN), we agree with Bouchard et al. (2011) and Figueroa et al. (2012) and adopt 1837 as the date of publication of Gromphas .
Taxon name Availability Validity Remarks
1. Pronotum with midlongitudinal line of smooth and glossy tegument at center (clearly seen in Fig. 3). Pronotal prominence formed by a pair of horizontal projections preceded by pair of smooth concavities (Figs. 1 –4, 7– 9); in very small specimens, pronotal prominence absent and concavities reduced to two impressions in anterior margin of pronotum (Figs. 5, 10). Protibial spur only slightly curved apically ( Figs. 22 View FIGURES 21 – 23 , 46 View FIGURES 34 – 48. 34 ). Ventral carina of protibia simple in both sexes ( Fig. 22 View FIGURES 21 – 23 ). Genae and frons with smooth tegument adjacent to eyes; remaining tegument of genae and frons with squamose granulation ( Figs. 24–25 View FIGURES 24 – 28 )....... 2
- Pronotum with smooth or rudimentary granulation at center ( Figs. 13–14 View FIGURES 13 – 15. 13 ); or, if granulation dense, midlongitunal line absent or very tenuous ( Fig. 15 View FIGURES 13 – 15. 13 ). Pronotal prominence absent. Protibial spur strongly expanded apically ( Figs. 21, 23 View FIGURES 21 – 23 , 45 View FIGURES 34 – 48. 34 ). Ventral carina of protibia with a row of tubercles in males ( Figs. 22 View FIGURES 21 – 23 d, 23 b) and simple in females ( Fig. 23 View FIGURES 21 – 23 a). Genae and frons with strong granulation adjacent to eyes ( Figs. 26–28 View FIGURES 24 – 28 )............................................................. 3
2 (1). Pronotal prominence with narrow projections; projections strongly divergent and with lateral margins strongly curved outwards (Fig. 1) or rarely straight (Fig. 2); separation between these projections always very wide (Figs. 1–2). Sutural margin of elytra and metasternum sparsely punctured. Cephalic projection bifid in major specimens ( Fig. 29 View FIGURES 29 – 33 ) (in worn specimens truncate, never acuminate). Pronotal granulation gradually decreasing in density from sides to center. Posterior pronotal fossae always present (Figs. 1–5). Colombia, Brazil, Ecuador, Bolivia, and Peru (Amazon subregion and north of the province of Chaco in Chacoan subregion)................................................ Gromphas aeruginosa (Perty, 1830)
- Pronotal prominence with broad projections; lateral margins straight and parallel to each other; separation between projections very narrow ( Figs. 7–8 View FIGURES 7 – 10 ). Sutural margin of elytra and metasternum densely punctured. Cephalic projection distinctly acuminate in major specimens ( Fig. 30 View FIGURES 29 – 33 ). Pronotum densely granulated, except in posterior region and midlongitudinal line. Posterior pronotal fossae absent or greatly reduced in major specimens, and present in minor specimens. Colombia and Venezuela (Caribbean subregion).................................................... Gromphas lemoinei Waterhouse, 1891
3 (1). Pygidium margined basally ( Fig. 35 View FIGURES 34 – 48. 34 ). Apical mesotarsomeres and metatarsomeres strongly curved ( Fig. 47 View FIGURES 34 – 48. 34 ). Metatibiae very thin ( Fig. 42 View FIGURES 34 – 48. 34 ). Posterior pronotal fossae always present. Anteromedian angle of metasternum strongly projecting, narrow and truncate apically ( Fig. 49 View FIGURES 49 – 52 ); region in front of anteromedian angle glabrous (Fig. 19, 49). Clypeus truncate apically and not lobed ( Fig. 26 View FIGURES 24 – 28 ), never folded upward. Cephalic projection truncate apically ( Fig. 33 View FIGURES 29 – 33 ). Brazil (Rio Grande do Sul), Paraguay, Argentina, and Uruguay (Chacoan subregion)................................... Gromphas dichroa Blanchard, 1846
- Pygidium not margined basally ( Fig. 36 View FIGURES 34 – 48. 34 ). Apical mesotarsomeres and metatarsomeres only slightly curved apically ( Fig. 48 View FIGURES 34 – 48. 34 ). Metatibiae very broad and robust ( Fig. 41 View FIGURES 34 – 48. 34 ). Posterior pronotal fossae usually absent; if rarely present, then very faint ( Fig. 43 View FIGURES 34 – 48. 34 ). Anteromedian angle of metasternum wide and rounded apically, not so projected ( Figs. 50–51 View FIGURES 49 – 52 ); region in front of the antero-
median angle with evident pilosity (Figs. 18, 20, 50–51). Clypeus with four lobes ( Figs. 27–28 View FIGURES 24 – 28 ) and distinctly folded upward. Cephalic projection emarginate apically ( Figs. 32–33 View FIGURES 29 – 33 ) (in very small and worn specimens, apex rounded or truncate)....... 4 4 (3). Elytral striae carinulate from base to the apical slope of elytra ( Fig. 38 View FIGURES 34 – 48. 34 ). Posteromedian region of pronotum smooth or with strongly effaced granulation ( Fig. 13 View FIGURES 13 – 15. 13 ). Posterior margin of pronotum rounded ( Fig. 13 View FIGURES 13 – 15. 13 ). Metatibial spur strongly curved apically ( Fig. 39 View FIGURES 34 – 48. 34 ). Male protibia with a long non-articulated spur ( Fig. 21 View FIGURES 21 – 23 b; arrow). Colombia, Brazil, and Peru (Amazon subregion).................................................................... Gromphas amazonica Bates, 1870
- Elytral striae simple. Pronotum with strong granulation in center, only posterior region completely smooth ( Figs. 15 View FIGURES 13 – 15. 13 , 43 View FIGURES 34 – 48. 34 ). Posterior margin of pronotum projecting at middle ( Fig. 15 View FIGURES 13 – 15. 13 ). Metatibial spur straight apically ( Fig. 40 View FIGURES 34 – 48. 34 ). Protibiae of both sexes without long non-articulate spur ( Figs. 23 View FIGURES 21 – 23 a –b). Brazil, Bolivia, Paraguay, Argentina, and Uruguay (Chacoan and Parana subregions)...................................................................... Gromphas inermis Harold, 1869
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