taxonID	type	description	language	source
707787C6597E624DFF0A228132C69F98.taxon	description	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; 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.	en	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): 439-482, DOI: 10.11646/zootaxa.3722.4.2
707787C6597E624DFF0A228132C69F98.taxon	materials_examined	Type species: Gromphas dichroa Blanchard, 1846; present designation.	en	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): 439-482, DOI: 10.11646/zootaxa.3722.4.2
707787C6597E624DFF0A228132C69F98.taxon	etymology	Etymology: from Greek, an old sow (feminine) (Brullé 1837, Brown 1956). Redescription: Head: Clypeus with apex truncate (Fig. 26) or with two small lobes; if lobed, then margin between apical lobes and genae rounded (Figs. 24 – 25) or with another pair of lobes (Figs. 27 – 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), or only slightly arched upward and emarginate (Figs. 31 – 32) or apically truncate (Fig. 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); area lateral to antero-median angle sloped and granulate. Legs: Protibiae with four rounded teeth, proximalmost less developed but always present (Figs. 21 – 23). Dorsal surface of protibia with longitudinal carina extended from base to apex and with thin branches into three apical teeth (Fig. 21). Ventral longitudinal carina of protibia simple in both sexes (Figs. 22, 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, 45) or only slightly bent apically (Figs. 22, 46). Inner apical angle of protibia with a tuft of long setae sexually dimorphic (Figs. 23) or not (Fig. 22); in males of G. amazonica, inner apical angle with a long non-articulate spur (Fig. 21 b, white arrow); in males of the other species, inner apical angle only with a tiny and almost imperceptible tubercle (Fig. 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) or without (Figs. 22, 23 b, 46) a spiniform projection in apex. Mesotibiae with three slightly pointed teeth on apical margin. Metatibiae very broad and robust (Fig. 41) or very thin (Fig. 42), with four slightly pointed teeth on apical margin and apical spur clearly curved (Fig. 39) or straight (Fig. 40) apically. Mesotarsi and metatarsi with five tarsomeres progressively shorter from basal outward, outer margin with rows of rigid setae (Figs. 47 – 48); 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) or basal margin completely absent (Fig. 36). Abdomen with six visible sternites (i. e., six ventrites); sixth ventrite strongly narrowed at middle in males (Fig. 34 a) and very wide at middle in females (Fig. 34 b). Genitalia: Aedeagus: Parameres coated by a wide area of short setae and white tegument (Figs. 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 c, 54 c, 55 c, 56 c). Internal sac as in Fig. 62; single medial sclerite with variable curvature, with (Fig. 60) 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).	en	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): 439-482, DOI: 10.11646/zootaxa.3722.4.2
707787C6597E624DFF0A228132C69F98.taxon	description	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]). 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.	en	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): 439-482, DOI: 10.11646/zootaxa.3722.4.2
707787C6597E624DFF0A228132C69F98.taxon	discussion	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	en	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): 439-482, DOI: 10.11646/zootaxa.3722.4.2
707787C65973624BFF0A250036379D89.taxon	description	Gromphas aeruginosa: Harold 1859: 199; Harold 1869 a: 1016, 1869 b: 62; Waterhouse 1891: 60; Heyne & Taschenberg 1908: 64, plate 9 Fig. 42; Ohaus 1909: 135; Gillet 1911: 80; d’Olsoufieff 1924: 20, 58, 138, plate II Fig. 3, plate IX; Pessoa & Lane 1941: 478; Blackwelder 1944: 208; Barattini & Saenz 1960: 23, 1964: 177; Halffter & Matthews 1966: 131; Vulcano & Pereira 1967: 566; Halffter & Edmonds 1982: 86; Scherer 1983: 298; Kirk 1992: 49; Amézquita et al. 1999: 119; Escobar 2000: 206; Medina & Lopera-Toro 2000: 303 Fig. 3 c, 305 Fig. 4 c; Vaz-de-Mello 2000: 193; Medina et al. 2001: 138; Noriega 2002: 72, 2004: 40; Philips et al. 2004 b: 59 Fig. 33, 62; Quinteros et al. 2006: 78; Monaghan et al. 2007: 681; Hamel-Leigue et al. 2009: 49, 2013: 281; Medina & Pulido 2009: 61; González-Hernández & Navarrete-Heredia 2011: 479; Figueroa et al. 2012: 1; Korasaki et al. 2012: 427; Cupello 2013: 15. Gromphas lacordairei Blanchard, 1846: 181; Harold 1859: 199; Harold, 1869 a: 1016, 1869 b: 62; Gillet, 1911: 80; d’Olsoufieff, 1924: 138; Blackwelder, 1944: 208; Barattini & Saenz 1960: 23, 1964: 177; Hamel-Leigue et al. 2009: 61.	en	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): 439-482, DOI: 10.11646/zootaxa.3722.4.2
707787C65973624BFF0A250036379D89.taxon	materials_examined	Type specimen: — Onitis aeruginosus Perty, 1830: Lectotype: designated by Scherer (1983), undetermined sex, (“ Brasilien ”, “ 3 - 8 typi ”, “ Type von Gromphas aeruginosa Perty ”, “ alte Sammlung ”, “ Holotypus Onitis aeruginosus Perty. det. Dr. G. Scherer, 1981 ”, “ Gromphas aeruginosus (Perty) det. G. Scherer, 1981 ”), ZSMC (examined by photos [Figs. 6 a – e]). Paralectotypes: 7 specimens, ZSMC (not examined). — Gromphas lacordairei Blanchard, 1846: Not located at MNHN; unknown to us. Type locality: — Onitis aeruginosus: “ Habitat in mediterraneis Prov. S. Pauli et Minarum ” (Perty 1830; but see comments below). — Gromphas lacordairei Blanchard: “ Province de Santa-Cruz de la Sierra (Bolivia) ” (Blanchard 1846). Redescription: Color: Anterior region of head black, remainder of dorsum with bright and metallic reflections of green, yellow, copper, and burgundy; some of these colors predominating or, more commonly, a mixture of all. Pygidium colored as elytra. Metasternum bright and of uniform color. Head: Clypeus with two lobes in the apex (in worn specimens lobes absent and apex round or truncate); margin between these lobes and genae round (Fig. 24). Margin of genae extended beyond clypeal margin (Fig. 24). Clypeus with smooth tegument adjacent to eyes; granulation gradually denser and squamous toward outer margin (Fig. 24). Cephalic projection ranging from a low and transverse elevation arched upward to a horn flattened antero-posteriorly with apex distinctly bifid (apex truncate or rounded in worn specimens); horn granulate on both faces (Figs. 29 a – e). Frons smooth adjacent to eyes and granulate in center (Fig. 24). Thorax: Lateral region of pronotum with dense squamous granulation (Fig. 16); granulation gradually sparser toward posteromedian region (Figs. 1 – 5). Longitudinal midline with smooth and glossy tegument present at center of pronotum (clearly in Fig. 3). Posterior pronotal fossae always present; granulation absent in fossae. Posterior margin of pronotum rounded. Anterior region of pronotum with ornamentation ranging from two shallow impressions in anterior margin (Fig. 5) to prominence with two tapered projections apically obtuse or slightly acute; usually, projections largely divergent and directed towards sides of body; lateral margin of prominence curved (Fig. 1) or, less usually, straight (Fig. 2); separation between projections always very wide; anterior to prominence, a pair of concavities with smooth tegument (Fig. 16). Mesosternum with dense pilosity (Fig. 52). Metasternum with fine and sparse punctation in the center, almost imperceptible in worn specimens. Anteromedian angle of metasternum flat and with rounded apex (Fig. 52), area in front of angle with evident setae (Figs. 16, 52). Legs: Protibiae very broad in both sexes; in ventral view, longitudinal carina simple in both sexes (Fig. 22). Protibial spur with apex slightly bent laterally and downward (Figs. 22, 46); inner apical protibial angle with a tuft of long setae in both sexes (Fig. 22). Apical protarsomere tapered and only slightly elongate at apex (Figs. 22, 46). Metatibiae very broad and robust (Fig. 41). Mesotarsi and metatarsi not particularly enlarged and with apical tarsomeres slightly curved apically (Fig. 48). Metatibial spur with apex distinctly curved (Fig. 39). Elytra: Striae very fine and, especially those more medial, carinulate from base to no more than midlength of of elytra, where they become simple (Fig. 37); interstriae flat. Sutural margin only sparsely punctate (Figs. 1 – 5). Abdomen: Pygidium flat and not margined basally (Fig. 36), sometimes with a pair of weak and almost imperceptible impressions on sides. Groove of propygidium extending to base of pygidium or penetrating to its basal third. Abdominal sternites sparsely punctate. Aedeagus: Phallobase, in ventral view, with a very narrow apical membranous area (Fig. 53 c). Medial sclerite strongly curved downward (Fig. 57). Measurements: Males (65 specimens): TL: AV: 15.44; MX: 20.3; MN: 11.6; SD: 1.87. PL: AV: 12.42; MX: 15.9; MN: 9.6; SD: 1.32. PW: AV: 9.13; MX: 11.7; MN: 6.8; SD: 1.17. PP: AV: 2.79; MX: 4.7; MN: 0.9; SD: 0.84. Females (50 specimens): TL: AV: 15.71; MX: 20.3; MN: 11.9; SD: 1.87. PL: AV: 12.59; MX: 15.3; MN: 9.7; SD: 1.36. PW: AV: 9.29; MX: 12.4; MN: 6.9; SD: 1.2. PP: AV: 2.89; MX: 5; M: 1; SD: 0.95. Total (115 specimens): TL: AV: 15.56; SD: 1.87. PL: AV: 12.49; SD: 1.34. PW: AV: 9.19; SD: 1.18. PP: AV: 2.83; SD: 0.89. Intraspecific variation and taxonomic discussion: A wide variation in the degree of development of the ornamentation is observed in this species. Small specimens have the cephalic projection and especially the pronotal prominence weakly developed (in smaller specimens the pronotal prominence is completely absent and the two anterior concavities are reduced to a pair of shallow impressions [Fig. 5]), while large specimens have them well developed, with the width of pronotal prominence up to 5 mm (Figs. 1 – 2). Between these two extremes, an entire gradual variation is present, with most specimens in an intermediate position (Figs. 3 – 4). Previous authors mistakenly believed that this difference in development was related to sexual dimorphism. Perty (1830) stated that males had an armed head and thorax, while females had the head transversely carinate and thorax unarmed; and Blanchard (1846) thought that the cephalic horns of females of G. lacordairei Blanchard would be brief and rudimentary. d’Olsoufieff (1924) was the first to recognize correctly that the females of G. aeruginosa could have the ornamentation as developed as males; even so, in some recent works, such as the cladistic analysis of Philips et al. (2004 b), the ornamentation is still incorrectly regarded as sexually dimorphic. We observed that the average width of the pronotal prominence does not differ significantly between the sexes, and the same is true with the cephalic projection. Having already recognized this, Barattini & Saenz (1960, 1964) pointed out the presence of a wide allometry in this species. As demonstrated for a wide variety of other dung beetles, the ornamentation of G. aeruginosa is positively allometric in relation to body length and pronotal width, but lacks the dimorphic condition found in some other genera, such as Onthophagus and Oxysternon, and in other beetles (e. g., Eberhard & Gutiérrez 1991; Edmonds & Zidek 2004; Emlen 1997 a, b; Kawano 2006). Sexual dimorphism of G. aeruginosa thus appears to be restricted to that observed for the genus in general (i. e., protarsus present only in females and sixth visible abdominal sternite narrowed in the middle in males and of even width in females). Barattini & Saenz (1960, 1964) observed a sexual difference in the apex of clypeus; according to them, only the females have the clypeus upturned. It is true that many well-preserved specimens of both sexes have the margin of the clypeus (and of genae, to a lesser degree) slightly folded, but we did not observe any sexual difference. In addition to the degree of development, the shape of the ornamentation also varies. Specimens of G. aeruginosa may possess the projections of pronotal prominence with apices acuminate or rounded. Most specimens have the projections widely divergent, but in a few specimens the projections are subparallel and the pronotal prominence is “ U ” shaped (Fig. 2). Still, all this variation occurs within a very limited range, and a cohesive unit is clearly visible. The species closest to G. aeruginosa, G. lemoinei, has a different shape of this prominence (see discussion of G. lemoinei). Unrelated to the development, the carinulae of the elytral striae also vary widely. In most specimens only the two most medial striae have evident carinulae, and they never extend beyond the apical half of the elytra (Fig. 37). However, many specimens have the second stria entirely simple and the first very weakly carinulate, while in others the carinulae are evident even in the third and fourth striae. Harold (1869 b), after observing the type specimens of G. lacordairei Blanchard then deposited at the Jardin des Plantes (current MNHN), confirmed the synonymy between G. aeruginosa and G. lacordairei Blanchard that he proposed in 1859. Although we have not examined the type specimens of G. lacordairei Blanchard (see comments below), this seems to be a correct position, since the description of Blanchard is very clear and consistent with G. aeruginosa. Moreover, we examined more than 20 Bolivian specimens of G. aeruginosa and did not find any notable difference from the rest of the species. The other species present in Bolivia, G. inermis, clearly does not fit the description given by Blanchard (e. g., “ Le prothorax est trés-épais, élevé antérieurement et muni dans cette partie de deux pointes divergentes ”). The only other species that would be consistent with this description, G. lemoinei, does not occur in Bolivia or in the rest of the Amazon subregion. Also, the name Onitis aeruginosus Perty was at no time cited by Blanchard, suggesting that he was unaware of the description of Perty’s species.	en	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): 439-482, DOI: 10.11646/zootaxa.3722.4.2
707787C65973624BFF0A250036379D89.taxon	discussion	Comments: The type series of G. aeruginosa was collected during the expedition of Johann Baptist von Spix and Karl Friedrich Philip von Martius through Brazil (1817 – 1820), who, from Rio de Janeiro, toured the southeast, northeast and north of that country (Papavero 1971). Despite being cited as the type locality by Perty, the former provinces and now states of São Paulo and Minas Gerais in southeastern Brazil are certainly not part of the distribution of this strictly Amazonian species. None of paralectotypes or the lectotype of G. aeruginosa possess labels indicating something more precise than Brazil (Fig. 6 e). Although the material collected in southeastern Brazil and in the Amazon were sent to Bavaria at different times (Papavero 1971), it is possible that they have been mixed up, which would have led to Perty’s mistake (also revealing, Phanaeus chalcomelas, another Amazon species collected by Spix and Martius and originally described in Onitis by Perty (1830), also has the wrong type locality of São Paulo and Minas Gerais). Many type specimens described by Perty were lost during World War I and II, but most of the type specimens of Scarabaeidae survived, including eight syntypes of G. aeruginosa (Scherer 1983). According to Scherer (1983), the original labels written in pencil by Perty were removed from these specimens and replaced by other labels by subsequent curators. Designated by Scherer (1983), the lectotype of G. aeruginosa (Fig. 6) has seven labels, none written in pencil (Fig. 6 e). Based on information indicated by him, the orange label written “ type von Gromphas aeruginosa Perty ” was prepared by Hans Kulzer, former preparator of the Zoologische Staatssammlung München; and the circular green label indicates that the material came from South America. The red-border label written by Scherer states “ holotypus ”, although the specimen has been designated by Scherer himself (Scherer 1983) as a “ lectoholotypus ”, equivalent to lectotype (Evenhuis 2008). It is clear that Perty (1830) based his description on more than one specimen because he made comparisons between what he considered to be males and females. We examined only photographs of the lectotype, and therefore could not to confirm its sex. Scherer said it was a male, but it is likely he believed that the differences in the development of ornamentation were sexually dimorphic. The confusion in sex determination probably occurred also with the paralectotypes, of which, according to Scherer, there were three males and four females. We initially doubted the commonly cited date of publication of Onitis aeruginosus, but 1830 seems to be correct. Blackwelder (1957) and Scherer (1983) cited this year as the publication of pages 1 – 60 of the “ Insecta Brasiliensis ” part of “ Delectus Animalium Articulatorum ”; Onitis aeruginosus was described on page 39. Also published in 1830, the eighth plate has the figure “ 8. Onitis aeruginosus ”, the first published illustration of a specimen of Gromphas known to us. (The date of publication of G. lacordairei Blanchard is also discussed in the comments on G. dichroa). The type of G. lacordairei Blanchard was not found by us. Based on what Blanchard (1846) wrote, we assume that is a series of syntypes. This assumption is based on the length provided for G. lacordairei Blanchard; he did not mention an exact measurement, but a range (“ long. 15 à 30 millim ”). Two junior primary homonyms of Onitis aeruginosus Perty were proposed in the 19 th century: O. aeruginosus Gistel, 1831 and O. aeruginosus Klug, 1855. The first name was proposed for a Brazilian species, but has not been cited (as valid or invalid) by any other author since; as its description is too vague and no type specimens are known to exist, Cupello (2013) regarded it as a nomen dubium and asked the International Commission on Zoological Nomenclature (ICZN) to suppress the name O. aeruginosus Gistel for the purposes of the Principle of Priority and of the Principle of Homonymy. The Klug’s name, however, is consistently and largely applied to an Afrotropical species of Onitis, and it is not considered as congeneric of Perty’s species since the latter was transferred to Gromphas by Harold (1859). Therefore, in order to preserve stability, Cupello (2013) asked the ICZN to rule that the name O. aeruginosus Klug is not invalid by reason to being a junior primary homonym of O. aeruginosus Perty, and thus remains as the valid name of this African dung beetle species. The reported occurrences of G. aeruginosa in Paraguay (Heyne & Taschemberg 1908) and in the Guianas (Vulcano & Pereira 1967) are probably incorrect. The presence in the Colombian departments of Meta and Casanare, as pointed out by Medina & Pulido (2009), is also unlikely; it is possible that the specimens studied by them are misidentified G. lemoinei (in fact, we examined several specimens of G. lemoinei collected in Meta). The specimens from the MZSP collection recorded for Santa Catarina, Brazil, certainly are mislabeled. These specimens were found in a drawer among specimens of G. inermis that had the same labels, correct for that species. Bionomics: Gromphas aeruginosa seems to be a common species in artificial open environments and in natural Amazonian savannas (Amezquita et al. 1999, Kirk 1992, Ohaus 1909). Specimens are found in pastures feeding on cattle and equine dung, and are also collected in traps baited with human feces (Hamel-Leigue et al. 2009). Regarding the activity time, Kirk (1992) and Hamel-Leigue et al. (2009) observed that G. aeruginosa is diurnal. However, specimens collected in Cruzeiro do Sul (Acre, Brazil) by R. L. Gonçalves have labels stating that they have been caught in a light trap. This species is also common at lights in towns of Acre and Rondonia in Brazil (FZVM, personal observation). The altitudinal range recorded for this species is 190 – 500 m in Bolivia (Hamel- Leigue et al. 2009) and 200 – 350 m in Colombia (Amezquita et al. 1999, Noriega 2002). Based on the specimen labels examined in this present work, we can say that the adults of G. aeruginosa are collected throughout the year. Geographic distribution: Amazonian subregion: Napo, Imeri, Varzea, Ucayali, Madeira, Tapajós-Xingú, Pantanal and Yungas. Chacoan subregion: Chaco. COLOMBIA: Huila: Gigante. Amazonas: Letícia. BRAZIL: Amazonas: Benjamin Constant, Borba, Eirunepé, Fonte Boa, Guajará, Itacoatiara, Manacapuru, Manaus, Parintins, São Gabriel da Cachoeira, São Paulo de Olivença, Tabatinga, Tefé. Pará: Altamira, Juruti, Óbidos, Santarém. Acre: Bujari, Cruzeiro do Sul, Feijó, Rio Branco. Rondônia: Porto Velho. ECUADOR: Napo. Orellana: Francisco Orellana (Chiruisla Station). PERU: Loreto: Maynas (Iquitos). San Martin: San Martin (El Porvernir). Ucayali: Coronel Portillo (Iparía; Callería: Pucallpa; Yarinacocha. Huánuco: Leoncio Prado (Tingo María). Madre de Dios: Tahuamanu (Iñapari); Tambopata (Puerto Maldonado: Lago Sandoval). Cuzco: Quispicanchi (Camanti: Quince Mil). BOLIVIA: Beni: (El Trapiche: Estación Biológica del Beni). Santa Cruz: Agustín Saavedra, Andrés Ibáñez (El Torno: La Angostura; Palmasola; Potrerillos del Guenda), Cordillera, Florida (Pampa Grande: Mataral), Ichilo (Buena Vista), José Miguel de Velasco (San Miguel de Velasco), Ñuflo de Chávez (Los Troncos), Obispo Santistevan (Montero; Saavedra). Cochabamba: Chapare. (Fig. 66).	en	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): 439-482, DOI: 10.11646/zootaxa.3722.4.2
707787C65973624BFF0A250036379D89.taxon	materials_examined	Material examined: 320 males and 271 females (381 with undetermined sex). BOLlVIA: without date and collector — 1 male (BMNH). COCHABAMBA: Chapare, without date and collector — 1 male (MZSP); Chapare, without date, Zischka col. — 1 female (MZSP); Chapare, 400 m., without date, Zischka col. — 1 male (CMNC). SANTA CRUZ: Flórida, Pampa Grande, Mataral, 08. XII. 1994, M. Quiroz col. — 1 male (CMNC); Cordillera, Rio Parapetí, I. 1964, Martínez col. — 1 male (CMNC); José Miguel de Velasco, San Miguel de Velasco, 26. XII. 2009, Vidaurre et al. cols. — 1 male (CEMT); Obispo Santisteva, Cuatro Ojos, without date and collector — 1 male and 1 female (MZSP); Obispo Santisteva, Montero, Rio Piraí, 23. III. 1987, F. D. B. col. — 1 female (CMNC); Obispo Santisteva, Rio Piraí, 08. I. 1994, S. Graz col. — 1 male and 1 female (CMNC); Obispo Santisteva, Saavedra, CIMCA (Centro Integral de Manejo del Comportamiento Animal), 10. IX. 1987, C. J. Pruett col. — 1 male (CMNC); Obispo Santisteva, Saavedra, CIMCA, 09. IX. 1988, C. J. Pruett col. — 1 male (CMNC); Gutierrez, Nueva Moka, III. 1956, A. Martínez col. — 1 female (MZSP); Ichilo, Buena Vista, 700 pés (518 m.), X. 1922, J. Steinbach co. l — 1 male (BMNH); Ichilo, Buena Vista, I. 1950, A. Martínez col. — 1 male and 1 female (MZSP); Ichilo, Buena Vista, II. 1950, without collector — 3 males (MLPA); Sara, Nueva Moka, I. 1963, A. Martínez col. — 1 male and 1 female (MZSP). BRAZIL: ACRE: Bujari, 20. VI. 1995, L. M. de Andrade col. — 1 female (CEMT); Bujari, 09. VII. 1995, L. M. de Andrade col. — 1 female (CEMT); Cruzeiro do Sul, VI. 1952, J. Correa col. — 2 males and 6 females (MNRJ); Cruzeiro do Sul, IX. 1952, J. Correia col. — 2 males and 5 females (MNRJ) and 2 males and 2 females (MZSP); Cruzeiro do Sul, IX. 1956, R. L. Gonçalves col. — 4 males and 5 females (MNRJ); Cruzeiro do Sul, Fazenda Morada Nova, 07 º 37 ’ 57 ” S / 72 º 43 ’ 43 ” W, 25 - 26. XI. 1996, C. S. Motta et al. col. — 4 males (INPA); Feijó, XI. 1956, Dirings col. — 22 males and 23 females; Feijó, XII. 1956, Werner col. — 1 male and 2 females (MZSP); Iquiri, VIII. 1951, without collector — 1 male (MZSP); Rio Branco, 30. V. 1988, Craveiro col. — 1 male (CEMT); Rio Branco, 11. VIII. 1994, M. L. Guerreiro col. — 1 female (CEMT); Rio Branco, 19. VIII. 1994, N. Batista col. — 1 male (CEMT); Rio Branco, 14. IV. 1995, M. F. S. de Oliveira & A. M. de Almeida — 1 female (CEMT); Rio Branco, 10. VI. 1995, S. M. N. Galdino col. — 1 female (CEMT); Rio Branco, 15. VI. 1995, S. M. F. Almeida col. — 1 male and 1 female (CEMT); Rio Branco, Campus Universitário, 15. III. 1982, William col. — 1 with undetermined sex (DZRJ). AMAZONAS: Benjamin Constant, III-IV. 1942, A. Parko col. — 3 males and 1 female (MNRJ); Benjamin Constant, XI. 1960, collector illegible — 1 female (MNRJ); Benjamin Constant, XII. 1960, L. G. Pereira col. — 2 males and 2 females (MNRJ); Benjamin Constant, 0 9. III. 2004, P. H. Silva col. — 1 female (CEMT); Benjamin Constant, 21 º 25 ’ 45 ’’ S 44 º 36 ’ 21 ’’ W, IX. 2008, V. Korasaki col. — 1 male (CEMT); Benjamin Constant, Rio Javari, IV. 1942, B. Pohl col. — 2 males and 2 females; Benjamin Constant, Rio Javari, IV-VI. 1942, A. Parko col. — 1 male and 1 female (MZSP); Benjamin Constant, Rio Javari, VII. 1942, A. Parko col. — 1 female (MNRJ); Benjamin Constant, Rio Javari, VIII. 1960, Dirings col. — 2 males and 2 females; Benjamin Constant, Rio Javari, I. 1961, Dirings col. — 3 males and 1 female (MZSP); Benjamin Constant, Rio Javari, III. 1961, Dirings col. — 1 male (MZSP); Benjamin Constant, Rio Javari, I-VIII. 1961, Dirings col. — 136 with undetermined sex (MZSP); Benjamin Constant, Rio Javari, V. 1961, Dirings col. — 1 female (MZSP); Benjamin Benjamin Constant, Rio Javari, VIII. 1961, Dirings col. — 13 males and 5 females (MZSP); Benjamin Constant, Rio Javari, XII. 1961, Dirings col. — 4 males and 5 females (MZSP); Benjamin Constant, Rio Javari, X. 1962, Dirings col. — 5 males and 6 females (MZSP); Benjamin Constant, Rio Javari, XI. 1962, Dirings col. — 1 female (MZSP); Benjamin Constant, Rio Javari, IX-XI. 1962, Dirings col. — 210 with undetermined sex (MZSP); Benjamin Constant, Rio Javari, IV. 1963, Dirings col. — 3 females (MZSP); Benjamin Constant, Rio Javari, X. 1963, Dirings col. — 3 females (MZSP); Benjamin Constant, Rio Javari, Alto Amazonas, without date, Dirings col. — 2 male and 1 female (MZSP); Borba, III. 1943, A. Parko col. — 4 males and 3 females (MNRJ) and 1 male (CEMT); Borba, Mata do Borba, II. 1943, A. Parko col. — 2 males (MZSP) and 2 females (MNRJ); Codajás, 10. X. 1935, without collector — 2 males and 1 female (MNRJ); Eirunepé (João Pessoa), Rio Juruá, 10. IX. 1936, Zellibor-Hauff col. — 1 female (MNRJ) and 1 male and 2 females (MZSP); Fonte Boa, X. 1975, F. M. Oliveira col. — 3 males and 2 females (MNRJ); Guajará, VII. 1943, A. Parko col. — 1 male (MZSP); Lago Amanã, XI. 1983, R. C. Best col. — 1 male and 1 female (INPA); Itacoatiara, XI. 1961, Dirings col. — 1 female (MZSP); Manacapuru, Costa do Pesqueiro II, 03 º 23 ’ 35 ” S / 60 º 36 ’ 57 ” W, 16. IX. 2006, F. F. Xavier col. — 1 male (INPA); Manaus, without date and collector — 3 males and 2 females (MZSP); Manaus, VII. 1927, J. F. Zikán col. — 1 female (MNRJ); Manaus, VIII. 1927, J. F. Zikán col. — 2 males (MZSP); Manaus, 1936, A. Ducke col. — 1 female (MNRJ); Manaus, INPA / Aleixo, 06. IV. 1990, J. A. O. Guerra col. — 2 males (INPA); Manaus, Rio Negro, VII (without year), J. F. Zikán col. — 1 male (FIOC); Manaus, Rio Negro, VIII. 1927, J. F. Zikán col. — 1 male and 1 female (MNRJ); Parintins, 1921, Garbe col. — 1 males (MZSP); São Gabriel da Cachoeira, Rio Negro, without date and collector — 1 female (MNRJ); São Paulo de Olivença, Rio Solimões, without date, Dirings col. — 1 male (MZSP); Tabatinga, XII. 1956, Pereira col. — 2 males and 1 female (MNRJ); Tefé, without date, Dirings col. — 9 males and 12 females (MZSP); Tefé, without date, J. F. Zikán col. — 1 female (CEMT); Tefé, VIII (without year), J. F. Zikán, — 1 male (MZSP); Tefé, VIII. 1921, J. F. Zikán col. — 2 males (FIOC) and 1 male and 1 female (MNRJ); Tefé, IX. 1921, without collector — 1 male (MZSP); Tefé, X. 1958, Dirings col. — 14 with undetermined sex (MZSP); Tefé, VIII. 1960, Dirings col. — 15 with undetermined sex (MZSP); Tefé, XII. 1961, Dirings col. — 3 males and 1 female (MZSP); Tefé, III. 1962, Dirings col. — 8 males and 10 female (MZSP); Tefé, VIII. 1991, O. Roppa & P. Magno cols. — 128 males and 87 females (MNRJ). PARÁ: Altamira, Rio Xingu, without date, J. F. Zikán col. — 1 male and 1 female (FIOC); Juruti, without date and collector — 1 male and 3 females (CEMT); Óbidos, XII. 1953, J. Brazilino col. — 1 male (MNRJ); Santarém, 27 - 28. XII. 1967, Exp. Perm. Amaz. col. — 1 male (MZSP). RONDÔNIA: Porto Velho, Rio Madeira, X. 1932, B. Pohl col. — 2 males and 2 females (MZSP). SANTA CATARINA: Rio Vermelho, XII. 1961, Dirings col. — 11 males and 7 females (MZSP); Rio Vermelho, IV. 1963, Dirings col. — 3 males and 4 females (MZSP). COLOMBIA: 1905, Fry col. — 1 male and 1 female (BMNH); 1910, Fry col. — 1 male (BMNH). AMAZONAS: Letícia, 700 ft., 08 - 09. VIII. 1970, H. & A. Howden cols. — 1 male and 1 female (CMNC); Letícia, 700 ft., 19 - 25. II. 1972, H. & A. Howden cols. — 1 male (CMNC); Letícia, 700 ft., 23. II- 02. III. 1974, H. & A. Howden cols. — 1 male and 1 female (CMNC). CALQUETÁ: Vegas Del Calquetá, without date, O. Rojas col. — 1 female (MIZA). HUILA: Gigante, VII. 1979, O. Rojas col. — 6 males and 9 females (MIZA); Gigante, 4. VI. 1981, O. Rojas col. — 1 male (MIZA); Gigante, parte alta da cordilheira, without date, O. Rojas col. — 1 female (MIZA); Gigante, parte alta da cordilheira, 1981, O. Rojas col. — 1 female (MIZA); Gigante, parte alta da cordilheira, 06 - 07. I. 1982, O. Rojas col. — 1 male (MIZA); Gigante, sopé da cordilheira, whitout date, O. Rojas col. — 1 male (MIZA). ECUADOR: NAPO: Rio Coca, IX. 2000, D. J. Inward & K. A. Jackson cols. — 1 male (BMNH). PERU: UCAYALI: Coronel Portillo, Pucallpa, XII. 1978, without collector — 1 female (CEMT); Coronel Portillo, Pucallpa, X-XI. 1992, L. E. Peña col. — 1 male and 2 females (CEMT); Coronel Portillo, Pucallpa, Rio Ucayalli, 200 m., IX. 1947, without collector — 1 male (MZSP); Coronel Portillo, Yarinacocha, 150 m., 8 - 15. VII. 2001, D. Curoe col. — 4 males and 4 females (CEMT); Coronel Portillo, Yarinacocha, Rio Ucayali, 11. X. 1946, J. Schunke col. — 1 female (MNRJ) and 2 females (MZSP).	en	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): 439-482, DOI: 10.11646/zootaxa.3722.4.2
707787C659756250FF0A24FD323A9FA5.taxon	description	Gromphas aeruginosa var. lemoinei Waterhouse: Gillet 1911: 80; d’Olsoufieff 1924: 138; Blackwelder 1944: 208; Barattini & Saenz 1960: 23, 1964: 177.	en	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): 439-482, DOI: 10.11646/zootaxa.3722.4.2
707787C659756250FF0A24FD323A9FA5.taxon	materials_examined	Type specimen: Holotype: undetermined sex (“ Type ”, “ Caracas ”, “ Gromphas lemoinei Waterh. type ”, “ Onthophoida lemoinei Reiche ” “ Gromphas aeruginosa (Perty) 614: 99 ”, “ Gromphas aeruginosa Perty, 1830 det. CPDT Gillet, 2007 ”), BMNH (examined by photos [Figs. 11 a – c]). Type locality: “ Caracas, La Guayra ” (Waterhouse 1891). Redescription: Color: Anterior region of head black; remainder of dorsum with bright and metallic reflections of green, copper, and burgundy, with some of these colors predominating or, more commonly, a mixture of all. Pygidium colored as elytra. Metasternum bright and with uniform coloring. Head: Clypeus with two lobes at apex (in worn specimens lobes absent and apex rounded or truncate); margin between these lobes and genae rounded (Fig. 25). Margin of genae extended beyond clypeal margin (Fig. 25). Clypeus with smooth tegument adjacent to eyes; granulation gradually denser and squamous toward the outer margin (Fig. 25). Cephalic projection ranging from a low and transverse elevation arched upward to anteroposteriorly flat horn with acuminate apex; horn granulate on both faces (Figs. 30 a – c). Frons smooth adjacent to eyes and granulate in center (Fig. 25). Thorax: Lateral region of pronotum with dense squamous granulation (Fig. 17); granulation always dense, without gradual decrease toward posteromedian region; posterior region with sparse granulation abruptly delimited (Figs. 7 – 10). Longitudinal midline with smooth and glossy tegument present at center of pronotum (easily observable in Fig. 3). Posterior pronotal fossae absent or greatly reduced only in major specimens; in other specimens, fossae present without granulation. Posterior margin of pronotum rounded. Anterior region of pronotum with ornamentation ranging from two shallow impressions in anterior margin (Figs. 9 – 10) to a prominence with two wide, triangular projections; lateral margin of prominence always straight (Figs. 7 – 8); separation between projections always very narrow; anterior to prominence a with a pair of concavities with smooth tegument (Fig. 17). Mesosternum with dense pilosity (Fig. 52). Metasternum very densely punctate in center. Anteromedian angle of metasternum flat and with rounded apex (Fig. 52), area in front of angle with evident setae (Figs. 16, 52). Legs: Protibiae very broad in both sexes; in ventral view, longitudinal carina simple in both sexes (Fig. 22). Protibial spur with apex slightly bent laterally and downward (Figs. 22, 46); inner apical protibial angle with a tuft of long setae in both sexes (Fig. 22). Apical protarsomere tapered and only slightly elongate at apex (Figs. 22, 46). Metatibiae very broad and robust (Fig. 41). Mesotarsi and metatarsi not particularly enlarged and with apical tarsomeres slightly curved apically (Fig. 48). Metatibial spur with apex distinctly curved (Fig. 39). Elytra: Striae very fine and, especially those more medial, carinulate from base to no more than midlength of elytra, where they become simple (Fig. 37); interstriae, especially mesial, slightly convex, producing a grooved appearance of striae to the naked eyes. Sutural margin densely punctate. Abdomen: Pygidium flat, not margined basally (Fig. 36). Groove of propygidium extending to base of pygidium. Abdominal sternites sparsely punctate. Aedeagus: Phallobase in ventral view with a very narrow apical membranous area (Fig. 53 c). Medial sclerite wide, flat and strongly curved laterally (Fig. 58). Measurements: Males (10 specimens): TL: AV: 16; MX: 20.1; MN: 12.7; SD: 2.04. PL: AV: 13.07; MX: 17; MN: 10.3; SD: 1.9. PW: AV: 9.4; MX: 11.7; MN: 7.1; SD: 1.29. PP: AV: 2.51; MX: 3.8; MN: 1.4; SD: 0.40. Females (4 specimens): TL: AV: 17.3; MX: 19.2; MN: 13.9; SD: 2.09. PL: AV: 13.73; MX: 14.8; MN: 11.2; SD: 0.9. PW: AV: 9.78; MX: 10.4; MN: 8.3; SD: 0.63. PP: AV: 2.63; MX: 3.1; MN: 2.1; SD: 0.46. Total (14 specimens): TL: AV: 16.38; SD: 2.06. PL: AV: 13.25; SD: 1.68. PW: AV: 9.5; SD: 1.14. PP: AV: 2.54; SD: 0.62. Intraspecific variation and taxonomic discussion: The closest relative of G. lemoinei is G. aeruginosa, with which it was synonymized for more than a century. The description given by Waterhouse (1891), although short and based only on a single specimen, is very accurate and uses basically the same morphological features used by us to distinguish these two species; it is surprising, therefore, that this synonymy has remained for so long. Gillet (1911) was the first author to mention Waterhouse’s species as “ var. lemoinei ” of G. aeruginosa, not making clear, however, whether or not he considered them synonymous; d’Olsoufieff (1924) was the first to state unequivocally this synonymy. Nevertheless, neither of these two authors nor even Barattini & Saénz (1960, 1964) discussed or justified this synonymy or the condition of variety of G. lemoinei, and it is very likely that they have not seen the type specimens. All this confusion probably occurred simply because of the superficial similarity that these two species show at the first glance; as they share only between themselves the following characteristics: genae and frons smooth adjacent to eyes (Figs. 24 – 25); cephalic projection very high (Figs. 29 – 30); pronotum with a smooth midlongitudinal line; presence of pronotal prominence (Figs. 1 – 5, 7 – 10); flattened anteromedian angle of metasternum (Fig. 52); tibiae very broad in both sexes (Fig. 22); protibia with ventral carina simple in both sexes and with apical tuft of setae without apparent sexual dimorphism (Fig. 22); protibial spur only slightly curved at apex (Figs. 22, 46); elytral striae carinulae extending at most only to midlenght of elytron (Fig. 37); and phallobase with a narrow apical membranous area (Fig. 53 c). Despite this extensive list of similarities, several morphological and biogeographical differences convincingly demonstrate the validity of each of these two species. Morrone (2001, 2006) recognized four subregions of the Neotropical region (Caribbean, Amazonian, Chacoan, and Parana), each them divided into several provinces. Gromphas lemoinei and G. aeruginosa inhabit northern South America, but have widely separated distributions. The former species occurs in the Venezuelan Coast and Venezuelan Llanos provinces of the Caribbean subregion, while the latter species is widely distributed in eight provinces of the Amazonian subregion and in the north of Chaco province of Chacoan subregion. Nowhere are these two species found in sympatry; in fact, they are separated by large physiographic barriers such as the Guiana Shield and the Orinoco River. This geographical separation certainly has contributed to the emergence and maintenance of the morphological distinction between them. The main distinction in the external morphology between G. aeruginosa and G. lemoinei is in the shape of the pronotal prominence of well-developed specimens: each of the two projections of the prominence of G. aeruginosa is clearly thin and tapered, while the projections are wide and triangular in G. lemoinei. In general, the projections are strongly divergent in G. aeruginosa, and always subparallel in G. lemoinei; hence the lateral margins of the prominence are usually curved in G. aeruginosa (Fig. 1), while in G. lemoinei they are always straight (Figs. 7 – 8). The space between the apices of the projections in G. lemoinei is much narrower than in G. aeruginosa, and the total width of the pronotal prominence is proportionally greater in G. aeruginosa than in G. lemoinei. For example, the largest specimen of G. lemoinei examined by us has a total length of 20.1 mm and a width of pronotal prominence of 3.8 mm; at the same time, the average width of the prominence in three specimens of G. aeruginosa with total length between 19.8 mm and 20.3 mm is 4.6 mm (the total average in G. aeruginosa is 2.83 mm and in G. lemoinei 2.54 mm). In a few specimens of G. aeruginosa the lateral margins of the prominence are subparallel and not strongly divergent (Fig. 2); however, in such cases, the thin and tapered projections and especially the wide separation between the projections are clear characteristics of G. aeruginosa. Characteristics of the tegument also separate these two species. The punctation in the center of metasternum and along the sutural margin of the elytra is very sparse in G. aeruginosa and seen only in unwarn specimens; in G. lemoinei, however, this punctation is much denser and clearly evident. Regarding the pronotal granulation, in G. aeruginosa, the squamous granules of the lateral region become progressively finer and sparser toward the center of the pronotum; in G. lemoinei, on the other hand, the density of granulation is always very high across the pronotum, except within the smooth anterior concavities and along the midline. The presence of posterior pronotal fossae also distinguish these two species: in G. lemoinei, larger specimens have no trace of these fossae, which are apparent in smaller specimens; in G. aeruginosa, the fossae are always present regardless of specimen size. The elytral striae are also distinct: in G. aeruginosa, the interstriae are flat and in the same plane as the striae, whereas in G. lemoinei the interstriae, especially the more mesial ones, are slightly convex at the base and the striae appear furrowed to the naked eye. This furrowed appearance gives the impression of a more elongate elytra in G. lemoinei than in G. aeruginosa. More developed specimens of G. aeruginosa have the cephalic projection distinctly notched and bifid (Fig. 29 b) (round or truncate in worn specimens [Fig. 29 c]). In large G. lemoinei, on the other hand, the cephalic projection is always acuminate (Fig. 30). While one could argue that the lack of notch in specimens of G. lemoinei has resulted from wear, even the best preserved specimens of G. lemoinei have no trace of an incision and no worn G. aeruginosa has the projection acuminate, thus ruling out this possibility. A very large Bolivian specimen of G. aeruginosa shows the cephalic projection extremely broad in a way not seen in any other specimen (Fig. 29 a). Finally, there is a difference between these two species also in the medial sclerite. While in G. aeruginosa the curvature is mainly downwards (Fig. 57), in G. lemoinei this sclerite is flatter, wider and very much more strongly curved to the side (Fig. 58). However, these two forms are clearly more similar to each other than to the other species of Gromphas.	en	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): 439-482, DOI: 10.11646/zootaxa.3722.4.2
707787C659756250FF0A24FD323A9FA5.taxon	discussion	Comments: The presence of G. lemoinei in San Fernando de Apure is inferred from the geographic distribution of G. aeruginosa presented by Barattini & Saénz (1960, 1964). These authors regarded the latter species (which does not occur in Venezuela) as a senior synonym of the former, but it is likely that these data refer to G. lemoinei. Bionomics: Noriega (2002) found five specimens of G. lemoinei (cited as G. aeruginosa) on the floodplain of Rio Duda, La Macarena, Meta, Colombia; the only other species of Scarabaeinae found by him in this place was Sulcophanaeus leander (Waterhouse, 1891). According Noriega (personal communication) these two species share the same activity time and the same food resource. As G. lemoinei is found only in savannas, open areas and river sandbanks and never enter in the forest, it is possible that this species use the rivers and their banks as corridors from the savannas of the Llanos Orientales to inside Amazon region in Colombia (Noriega, personal communication). Noriega also reported that all specimens collected by him were associated with cattle (Bos indicus Linnaeus, 1758), capybara (Hydrochoerus hydrochaeris (Linnaeus, 1766 )), or crocodile (Crocodylus intermedius Graves, 1819) dung. Based on the specimens examined in this study, G. lemoinei seems to be active throughout the year, as is G. aeruginosa. The recorded months are January, March, June, July, September, November, and December. Geographical distribution: Caribbean subregion: Venezuelan Coast and Venezuelan Llanos. COLOMBIA: Arauca: Arauca. Meta: La Macarena, Puerto Gaitan, San Martín, Villavicencio. Guaviare: San José del Guaviare. VENEZUELA: Distrito Capital: Caracas. Barinas: Andrés Eloy Blanco (El Cantón), Ezequiel Zamora (San Antonio de Pagen). Táchira: Libertador (San Joaquín de Navay), San Cristóbal, Santo Domingo. Apure: San Fernando de Apure. (Fig. 66).	en	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): 439-482, DOI: 10.11646/zootaxa.3722.4.2
707787C659756250FF0A24FD323A9FA5.taxon	materials_examined	Material examined: 17 males and 10 females. COLOMBIA: ARAUCA: Arauca, 120 m., IX. 2005, Jorge Ari Noriega col. — 1 male (CJAN). GUAVIARE: San Jose del Guaviare, sabana, IX. 2000, J. Noriega col. — 2 males (CJAN). META: La Macarena, CIEM, Playa del Rio Duda, 350 m, XII. 1995, Jorge A. Noriega col. — 2 females (CJAN); La Macarena, CIEM, I. 1998, J. Noriega col. — 1 male (CJAN); Puerto Gaitan, 150 m., X. 2001, Jorge Ari Noriega col. — 1 female (CJAN); San Martín, 350 m., VII. 2007, Jorge Ari Noriega col. — 1 male and 1 female (CJAN); Villavicencio, 13. XII. 1994, Oscar Gómez col. — 1 female (CEMT). VENEZUELA: BARINAS: Andrés Eloy Blanco, El Cantón, XI. 1983, C. Borges col. — 1 male (MIZA); Ezequiel Zamora, San Antonio de Pagen, VII. 1989, without collector — 1 female (MZSP) and 2 males (CEMT); San Antonio, VII. 2007, J. Noriega col. — 1 male and 1 female (CJAN). DISTRITO CAPITAL: Caracas, 1905, Fry col. — 1 male (BMNH). TÁCHIRA: Libertador, San Joaquín de Navay, 100 m, 24. III. 1989, D. Havranek col. — 1 male (CMNC); San Cristóbal, without date and collector — 2 male (MIZA); San Cristóbal, 17. III. 1980, without collector — 1 male (MIZA); Santo Domingo, Hacienda Santa Rosa, VI. 1992, J. Blanco col. — 1 male (MZSP) and 2 male and 2 female (CEMT).	en	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): 439-482, DOI: 10.11646/zootaxa.3722.4.2
707787C6596E6253FF0A26D936089CE1.taxon	description	Bolbites dichrous: Harold, 1869 a: 1016.	en	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): 439-482, DOI: 10.11646/zootaxa.3722.4.2
707787C6596E6253FF0A26D936089CE1.taxon	materials_examined	Type specimen: Holotype: Not located at MNHN; unknown to us. Type locality: “ Montevideo, près de la mer ” (Blanchard 1846). Redescription: Color: Two general patterns with intermediates (Fig. 14 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 (Figs. 14 a, 19). 2) Anterior region of clypeus black; remainder of dorsum, metasternum and pygidium predominantly bright light green (Fig. 14 c). Head: Clypeus with truncate apex; genae with very dense and coarse granulation, including region adjacent to eyes (Fig. 26). Cephalic projection only slightly elevated and with truncate apex (Fig. 33). Frons granulate at least adjacent to eyes (Fig. 26). 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. Mesosternum glabrous or with very sparse setae (Fig. 49). Metasternum clearly punctate at center. Anteromedian angle strongly projecting and narrowed, with truncate apex (Fig. 49); metasternal area in front of anteromedian angle entirely glabrous (Figs. 19, 49). Legs: Protibiae very narrow in both sexes; in ventral view, longitudinal carina with a row of tubercles on basal half in males (Figs. 20 d, 23 b) and simple in females (Fig. 23 a). Protibial spur with apex strongly expanded and curved downward (Figs. 23, 45); inner apical angle of protibia with tuft of setae longer in males than in females (Figs. 23 a – b). Metatibia very narrow (Fig. 42). Mesotarsi and metatarsi not enlarged and with apical tarsomeres strongly curved (Fig. 47). Metatibial spur with apex distinctly curved (Fig. 39). Elytra: Striae very fine, simple and effaced, almost imperceptible, especially eighth stria. Sutural margin smooth, distinctly depressed at base. Abdomen: Pygidium with a pair of weak impressions on sides; basal margin complete and never interrupted by groove of propygidium (Fig. 35). Sixth sternite densely punctate, remaining sternites smooth. Aedeagus: Phallobase, in ventral view, with wide membranous apical area (Fig. 54 c). Medial sclerite very thin and curved (Fig. 61). Measurements: Males (2 specimens): TL: AV: 12.85; MX: 13; MN: 12.7; SD: 0.21. PL: AV: 12.85; MX: 13; MN: 12.7; SD: 0.21. PW: AV: 7.6; MX: 8.1; MN: 7.1; SD: 0.71. Females (5 specimens): TL: AV: 14.84; MX: 16.6; MN: 12.5; SD: 1.54. PL: AV: 12.44; MX: 13.3; MN: 11.8; SD: 0.74. PW: AV: 8.8; MX: 9.7; MN: 8; SD: 0.79. Total (7 specimens): TL: AV: 14.27; SD: 1.59. PL: AV: 11.94; SD: 1.1. PW: AV: 8.46; SD: 0.92. Intraspecific variation and taxonomic discussion: The most striking intraspecific variation in G. d i ch ro a is the color: two distinct patterns are observable, one where green is predominant (Fig. 14 c) and the other with dark blue elytra and reddish-blue pronotum (Fig. 14 a). However, the specimen from Porto Alegre (Rio Grande do Sul, Brazil) has an intermediate coloration: the elytra are light blue and the pronotum and head are green with a large iridescent yellowish-red spot (Fig. 14 b). The green female collected in Santo Tomé (Corrientes, Argentina) is labeled “ Gromphas dichroa var. fulgida 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.	en	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): 439-482, DOI: 10.11646/zootaxa.3722.4.2
707787C6596E6253FF0A26D936089CE1.taxon	description	Gromphas dichroa has the highest number of unique characters in the genus: (a) truncate apex of clypeus (Fig. 26); (b) clypeus not folded upwards to any degree (in G. inermis and G. amazonica, the clypeus is strongly folded, while in G. aeruginosa and G. lemoinei it is only slightly); (c) truncate cephalic projection (Fig. 33); (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 (Fig. 42); (g) apical mesotarsomeres and metatarsomeres strongly curved (Fig. 47); (h) elytral suture depressed at base (Figs. 14 a – c); and (i) base of pygidium completely margined (Fig. 35). In addition, (j) the anteromedian angle of metasternum of G. dichroa differs from the other species: it is convex, as in G. amazonica and G. inermis (Figs. 50 – 51), but is still much more projected and has a truncate apex (Fig. 49); in G. lemoinei and G. aeruginosa, the angle is always very flat (Fig. 52). The smooth sutural margin of elytra (k) is also unique to G. dichroa (in G. aeruginosa the punctation is very sparse, but still present). The color pattern described above for this species (l) is also unique among Gromphas. Finally, the general shape of the medial sclerite of G. dichroa (m) is distinct from that of all other species of Gromphas (Fig. 61). Gromphas dichroa shares with G. lemoinei and G. aeruginosa a clypeus with rounded sides (Fig. 26), and only with G. aeruginosa the presence of posterior pronotal fossae in specimens of any size (in G. lemoinei, 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 G. aeruginosa). On the other hand, it resembles G. inermis and G. amazonica by the absence of any kind of pronotal prominence; by the protibial spur expanded at the apex (Figs. 21, 45); 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 [Figs. 23 a – b]) and in the ventral carina of protibiae (with a row of tubercles in males [Figs. 20 d, 23 b] and simple in females [Fig. 23 a]). Only with G. inermis does G. dichroa share simple elytral striae (in G. dichroa, they are very indistinct and almost imperceptible, especially the eighth and ninth striae, while in G. inermis all striae are easily observed); and only with G. amazonica does it share a smooth posteriomedian region of the pronotum (Figs. 13, 14). In Harold’s (1869 a) catalog, G. dichroa was transferred to the genus Bolbites, but soon thereafter Harold (1869 b) returned this species to its original genus. From all the characters discussed in the description of Gromphas and in the diagnosis of Gromphadina, it is clear that G. di ch ro a belongs in Gromphas and not in Bolbites.	en	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): 439-482, DOI: 10.11646/zootaxa.3722.4.2
707787C6596E6253FF0A26D936089CE1.taxon	discussion	Comments: We have not been able to examine the holotype of G. di ch ro a. Blanchard's (1846) description was based on a single specimen that he believed to be a female (“ Nous n’en avons vu qu’une femelle, que M. d’Orbigny a trouvée à Montevideo, près de la mer ”). d’Olsoufieff (1924) studied the holotype and said it was housed in the MNHN collection, but we have not found it there. Gromphas dichroa and G. lacordairei Blanchard were published respectively on pages 181 and 182 of volume 6, part 2, of Voyage dans l’Amérique méridionale, series published between 1835 and 1847 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 Gromphas. The publication date usually cited for these species, i. e., 1843, is incorrect. According Sherborn & Woodward (1901) and Evenhuis (1997), livrasion 80, containing pages 169 – 184 of the sixth volume, was published in 1846. Martinez (1959) reported G. dichroa from Paraguay. Although we do not know specimens from this country, we examined specimens collected in two Argentine provinces bordering Paraguay, Misiones and Corrientes. It is thus possible that this species is actually present in Paraguay. None of the females of G. dichroa 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 (e. g., if the spiniform projection seen in G. amazonica is present or not). Bionomics: There are no bionomic studies published on G. dichroa. The specimens examined in this study were collected between October and January. It is probable that G. dichroa 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 1922 and 1947. It is truly remarkable that no specimen has been found over the past 65 years since G. dichroa is distributed in one of the most intensely collected regions of South America. As nothing is known about the natural history of this species, it is difficult to surmise the reasons for the putative disappearance of G. dichroa, but it is possible that the intense human pressure in the Southern Cone (which includes southern Brazil, Paraguay, Uruguay, Argentina, and Chile) has acted negatively on what were already small populations. Geographic distribution: Chacoan subregion: Chaco and Pampa. BRAZIL: Rio Grande do Sul: Cerro Largo, Porto Alegre. PARAGUAY. ARGENTINA: Misiones: Concepción (Santa Maria). Corrientes: Santo Tomé. Buenos Aires. URUGUAY: Tacuarembó, Soriano, Flórida, Montevidéu. (Fig. 66).	en	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): 439-482, DOI: 10.11646/zootaxa.3722.4.2
707787C6596E6253FF0A26D936089CE1.taxon	materials_examined	Material examined: 3 males and 6 females. ARGENTINA: BUENOS AIRES: Anchorena, X. 1935, M. J. Viana col. — 1 male (CMNC). CORRIENTES: Santo Tomé, XII. 1922, without collector — 1 female (CMNC). MISIONES: Concepción, Santa Maria, XI. 1947, M. J. Viana col. — 1 male (MLPA). BRAZIL: RIO GRANDE DO SUL: Cerro Largo (former Serro Azul), I. 1931 (or 1933?), without collector — 1 male (CEMT); Porto Alegre, X. 1928, without collector — 1 female (MZSP). URUGUAY: without date and collector — 3 females (MZSP) and 1 female (CEMT).	en	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): 439-482, DOI: 10.11646/zootaxa.3722.4.2
707787C6596D625DFF0A242532569A19.taxon	materials_examined	Type locality: Tefé (former Ega), Amazonas, Brazil (type locality of the lectotype, according Article 76.2 of the Code [International Commission on Zoological Nomenclature 1999]). Redescription: Color: Dorsum and metasternum usually entirely shiny black, dark blue (Fig. 13 a), or dark green (Fig. 13 c), without metallic reflections; some specimens reddish-brown (Fig. 13 b). Head: Margin of clypeus with four lobes (Fig. 27) and folded upwardly, especially two medial lobes. Genae and frons completely granulate, including region adjacent to eyes (Fig. 27). Cephalic projection a raised carina with emarginate apex in major specimens (Fig. 31), and only slightly projecting with rounded apex in minor specimens. Thorax: Pronotum globular; lateral region with dense granulation (Fig. 18), density of granulation decreasing posteromedially; posteromedian region smooth or with strongly effaced granulation (Fig. 13); posterior fossae usually absent, apparent in very few specimens only as two very shallow and almost imperceptible impressions (Fig. 43). Posterior margin of pronotum rounded. Mesosternum with dense pilosity (Fig. 52). Metasternum with variable density of punctation. Anteromedian angle of metasternum convex and with globose apex (Fig. 51); area in front of anteromedian angle with evident setae (Figs. 18, 51). Legs: Protibia narrower in males than in females (Figs. 21 a – b); in ventral view, longitudinal carina with a row of tubercles in its basal half in males (Figs. 20 d, 23 b) and simple in females (Fig. 23 a). Protibial spur with apex strongly expanded and curved downward (Figs. 23, 45). In males, inner apical angle of protibia with a long nonarticulated spur bearing a row of setae on its dorsal surface (Fig. 21 b, white arrow); in females, inner apical angle only with a tuft of short setae (Fig. 21 a). Apex of apical protarsomere with a long spiniform projection (Fig. 45). Mesotarsi and metatarsi very broad and with apical tarsomeres only slightly curved apically (Fig. 48). Metatibiae very broad and robust (Fig. 41). Metatibial spur with apex distinctly curved (Fig. 39). Elytra: Striae very fine and, especially striae 1 – 4, distinctly carinulate from base to apical slope of elytra (Fig. 38). Sutural margin glossy and punctate; in some specimens basal third or basal half of sutural margin with sheen and punctation extending laterally into first or second interstria (easily seen in Fig. 15 b). Abdomen: Pygidium flat, not margined basally (Fig. 36). Groove of propygidium extending to base of pygidium. Abdominal sternites smooth or sparsely punctate. Aedeagus: Phallobase, in ventral view, with a very broad apical membranous area (Fig. 56 c). Medial sclerite only slightly curved, almost flat, with a narrow and spiniform projection at one end (Fig. 60). Measurements: Males (18 specimens): TL: AV: 15.59; MX: 18.1; MN: 13.6; SD: 1.32. PL: AV: 13.73; MX: 15.3; MN: 13.6; SD: 1.32. PW: AV: 9.38; MX: 10.4; MN: 7.7; SD: 0.72. Females (20 specimens): TL: AV: 14.72; MX: 17; MN: 12; SD: 1.32. PL: AV: 12.93; MX: 14.4; MN: 10.5; SD: 1.12. PW: AV: 8.86; MX: 10.1; MN: 7; SD: 0.83. Total (38 specimens): TL: AV: 15.13; SD: 1.38. PL: AV: 13.31; SD: 1.09. PW: AV: 9.11; SD: 0.82. Intraspecific variation and taxonomic discussion: Gromphas amazonica shows a wide variation in the punctation of the sutural margin of elytra and metasternum; some specimens show conspicuous punctation, while in others it is only slight. Variation is also seen in the spiniform projection of the female protarsus; most specimens have it well developed and evident (Fig. 45), while some others seem to have the projections worn and consequently shortened. Posterior pronotal fossae are usually absent in G. amazonica, but occasionally they are represented by a pair of small, almost imperceptible, punctiform pits (Fig. 43). We observed these fossae only in two specimens; a small male from Itacoatiara (Amazonas, Brazil) and a large female from Afuá (Pará, Brazil) (Fig. 13 c). Although the other specimens did not show any trace of these fossae, it is clear that this is no more than individual variation, the same occurring in G. inermis, also a species typically without posterior pronotal fossae (see more on the discussion of G. inermis). Although the non-articulated spur of the male protibiae is unique to G. amazonica (Fig. 21 b), its homology with the small tubercle found in the male protibiae of the other species of Gromphas (Fig. 23 b, white arrow) is likely (the tubercle, as the non-articulated spur, is absent in females). However, in no other species is this tubercle developed more than an almost imperceptible; in G. amazonica, the non-articulated spur is visible to the naked eye even in smaller specimens. Other unique characters of G. amazonica are (a) the elytral striae carinulate beyond midpoint of elytra (Fig. 38); (b) apical protarsomere with a long spiniform projection (Fig. 45); (c) mesotarsi and metatarsi very wide; (d) usual absence of metallic reflections; and (e) medial sclerite of internal sac with a spiniform projection (Fig. 60). The phallobase with membranous apical area expanded in ventral view and the medial sclerite only slightly curved are characters shared in some way with G. dichroa and G. inermis, respectively, although in G. amazonica both characters are much more pronounced. With G. aeruginosa and G. lemoinei, G. amazonica shares the elytral striae carinulate (in G. lemoinei and G. aeruginosa the carinulae are always absent in the apical half of elytra [Fig. 37], while in G. amazonica the carinulae reach the apical slope of elytra [Fig. 38]). Gromphas amazonica shares only with G. inermis the margin of the clypeus with four lobes (Fig. 27) and folded upward; the cephalic projection emarginate apically (Figs. 27, 31); and the usual absence of posterior pronotal fossae (and their occasional presence in some specimens [Fig. 43]). (See comparison with G. dichroa in the discussion of that species). We examined a single female of G. amazonica that bears the unexpected locality of Cáceres (Mato Grosso, Brazil) in the Alto Pantanal region, about 1,000 km south of Contamana, Peru, the second southernmost point known for this species. This specimen has distinctly carinulate elytral striae and apical protarsomere with a long spiniform projection, but, unlike other G. amazonica, it has a strong green color with metallic reflections covering the entire body, especially the head and pronotum. With only one specimen known, it is impossible to say whether this condition is typical of this population of G. amazonica or just an individual variation (or even that this is the result of mislabeling). If indeed the existence of this Patanal population is confirmed, it would be a case analogous to, for example, Sulcophanaeus faunus (Fabricius, 1775), a species widely distributed throughout Amazon Basin with small isolated populations in Paraguay and western São Paulo state (Edmonds 2000).	en	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): 439-482, DOI: 10.11646/zootaxa.3722.4.2
707787C6596D625DFF0A242532569A19.taxon	discussion	Comments: Bates (1870) based his description of G. amazonica on four specimens (“ I do not find any sexual difference in the four specimens I posses ”). Nonetheless, we were able to locate only three of the syntypes at the MNHN, all females; one of them (that from Tefé, former Ega) was designated by us as the lectotype of G. amazonica. Bionomics: Figueroa et al. (2012) were the first to publish data on the natural history of G. amazonica. They report that this species is collected in pitfall traps baited with human feces in forests habitats and, in one case, several specimens were collected in a secondary forest associated with mandioca cultivation (Manihot esculenta Crantz). The specimens that we examined from Afuá, Pará, were also collected on human feces (Paulo R. Magno, MNRJ, personal communication). The specimens examined in this study were collected in July, August, September, November, January, February, and March.	en	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): 439-482, DOI: 10.11646/zootaxa.3722.4.2
707787C6596D625DFF0A242532569A19.taxon	description	Geographic distribution: Amazonian subregion: Amapa, Varzea, Ucayali, Madeira, and Pantanal. COLOMBIA: Amazonas: Letícia. BRAZIL: Amapá: Macapá. Amazonas: Benjamin Constant, Carauari, Eirunepé, Itacoatiara, São Paulo de Olivença, Tefé. Pará: Afuá (Ilha do Pará), Porto de Moz (Tapara). Mato Grosso: Cáceres. PERU: Loreto: Mariscal Ramón Castilla (Pebas), Maynas (Iquitos); Ucayali (Contamana; Padre Márquez). San Martin: San Martin (El Porvenir). Ucayali: Coronel Portillo (Callería: Pucallpa; Yarinacocha). (Fig. 66).	en	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): 439-482, DOI: 10.11646/zootaxa.3722.4.2
707787C6596D625DFF0A242532569A19.taxon	materials_examined	Material examined: 22 males and 26 females. BRAZIL: AMAPÁ: Macapá, BR 156 km. 14, 26. XI. 1981, I. S. Gorayeb & Equipe cols. — 2 females (CEMT). AMAZONAS: Benjamin Constant, IX. 1953, I. C. Lima col. — 1 male (MNRJ); Carauari, 57 º 28 ’ 05 ” S 05 º 40 ’ 75 ” W, 85 m, 19. VII. 2009, L. Nichols, R. Braga & G. Schiffler cols. — 1 male and 1 female (CEMT); Eirunepé (former João Pessoa), Rio Juruá, 10. IX. 1936, Zellibor-Hauff col. — 2 males and 1 female (MNRJ), 1 male and 1 female (FIOC) and 1 female (MZSP); Eirunepé, Rio Juruá, IX. 1936, without collector — 1 male (MZSP); Itacoatiara, without date, Mann & Baker cols. — 1 male (BMNH); Tefé, VIII. 1991, O. Roppa & P. Magno cols. — 2 females (MNRJ). MATO GROSSO: Cáceres, 10. X. 2008, E. Silva col. — 1 females (CEMT). PARÁ: Afuá, Ilha do Pará, Igarapé Juntinta, XI. 1995, P. Magno & C. Júlio cols. — 10 males and 13 females (MNRJ); Porto de Moz, Tapara, I. 1923, J. F. Zikán col. — 1 male (FIOC); Porto de Moz, Tapara, II. 1923, J. F. Zikán col. — 1 female (CEMT). COLOMBIA: AMAZONAS: Letícia, 700 ft., 23. II- 02. III. 1974, without collector — 1 male (CMNC); Letícia, 700 ft., Isla Sta. Sophia, 23. II- 02. III. 1974, J. Peck col. — 1 female (CMNC); Parque Nacional Natural Amacayacu, 90 m, 23. IX. 1993, collector illegible — 1 female (CMNC). PERU: Ucayali, Coronel Portillo, Yarinacocha, 150 m, 08 - 15. VII. 2001, D. Curoe col. — 3 males and 1 female (CEMT).	en	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): 439-482, DOI: 10.11646/zootaxa.3722.4.2
707787C659636261FF0A238D330E9A75.taxon	description	Gromphas lacordairii Burmeister, 1874: 130 (cited as “ G. lacordairei Dejean ” or “ G. lacordairei Brullé ”); Lacordaire 1876: 10, plate 27 Fig. 4; Tremoleras 1910: 25; Bruch 1911: 189; Gillet 1911: 80; Fabre 1919: 244; d’Olsoufieff 1924: 20, 59, 139, plate II Fig. 2, plate IX; Manter 1928: 345; Pessoa & Lane 1941: 472; Blackwelder 1944: 208; Lange 1947: 313; Lima 1953: 62; Ruffinelli & Carbonell 1953: 25; Janssens 1954: plate II; Martínez 1959: 95; Barattini & Saenz 1960: 25, 1964: 179; Halffter & Matthews 1966: 131, 225; Halffter & Edmonds 1982: 84, 86; Martínez, 1989: 67; Flechmann et al. 1995: 267; Flechmann & Rodrigues 1995: 304; Louzada et al. 1996: 164; Monteresino et al. 1996: 109; Walsh & Gandolfo 1996: 582, 583 Fig. 1 E, 285 – 287; Walsh & Cordo 1997: 194; Martínez & Cruz 1999: 805; Koller et al. 1999: 405, 2007: 85; Vazde-Mello 2000: 193; Morelli et al. 2002: 54; Hamel-Leigue 2006: 6, 2009: 49, 2013: 281; Damborsky et al. 2008: 149; Sánchez & Genise 2008: 49; Vieira et al. 2008: 722; Almeida & Mise 2009: 238; Silva et al. 2009: 36; Audino et al. 2011: 125; González-Hernández & Navarrete-Heredia 2011: 479; Silva 2011: 552; Vaz-de-Mello et al. 2011: 59, Fig. 95; Figueroa et al. 2012: 2. Gromphas lacordairei bipunctata d’Olsoufieff, 1924: 59 (cited as “ var. bipunctata ”); Vaz-de-Mello 2000: 193. New synonymy	en	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): 439-482, DOI: 10.11646/zootaxa.3722.4.2
707787C659636261FF0A238D330E9A75.taxon	materials_examined	Type specimens: — G. inermis Harold, 1869: Lectotype: here designated, male (handwritten “ Buenos Aires ”, Harold’s handwritten “ inermis Harold ”, printed on white paper bordered in black “ Ex musaeo E. Harold ”, LECTOTYPE ”, “ LECTOTYPE Gromphas inermis Har. Vaz-de-Mello, 2013 ”), MNHN, ex col. Oberthür (examined). Paralectotype: Not located at MNHN unknown to us. — G. lacordairei bipunctata d’Olsoufieff, 1924: Lectotype: here designated, male (bordered in black “ Brésil, Prov. Matto Grosso, P. Germain, 1886 ”, “ LECTOTYPE ”, “ LECTOTYPE Gromphas lacordairei var. bipunctata Ols. Vaz-de-Mello, 2013 ”), MNHN, ex col. Oberthür (examined). Paralectotype: Not located at MNHN; unknown to us. FIGURES 16 – 20. Lateral views. 16) Gromphas aeruginosa. 17) G. lemoinei. 18) G. amazonica. 19) G. dichroa. 20: G. inermis. 20 a) Major specimen (the white arrow indicates “ pronotal hump ” and blue arrow indicates elevation of sutural margin of elytra). 20 b) Medium-sized specimen. 20 c) Minor specimen. 20 d) Lateral view of row of tubercles on ventral carina of protibia (indicated by arrow). Scale = 5 mm. Type locality: G. inermis: Buenos Aires, Argentina (type locality of the lectotype, according Article 76.2 of the Code [International Commission on Zoological Nomenclature 1999]). G. lacordairei bipunctata: Mato Grosso, Brazil (type locality of the lectotype). Redescription: Color: Anterior region of clypeus black; remainder of dorsum with metallic reflections and color ranging from black (Fig. 15 c) to dark green (Fig. 15 b) or entirely copper with green reflections (Fig. 15 a). Metasternum with distinct color, usually bright dark blue, blue with green reflections, or shiny green. Pygidium green or blue. Head: Margin of clypeus with four lobes (Fig. 28); apical margin folded upwardly, especially two median lobes. Genae and frons completely granulate, including region adjacent to eyes (Fig. 28). Cephalic projection raised in a carina with emarginate apex in major specimens (Fig. 32), and only slightly projecting with rounded apex in minor specimens. Thorax: Pronotum globular, in larger specimens, with evident anterior swelling (“ pronotal hump ”, Fig. 20 a, white arrow); lateral region and center with dense granulation (Figs. 20 a – d); posterior region smooth (Figs. 15 a – c). Longitudinal midline of glossy and smooth tegument absent or, rarely, very tenuous; posterior fossae usually absent, represented in some specimens by two very shallow, faint impressions (Fig. 43). Posterior margin projected at middle (Fig. 15 a – c). Mesosternum with dense pilosity (Fig. 52). Metasternum with very dense punctation in center. Anteromedian angle of metasternum convex and with globose apex (Fig. 50); area in front of anteromedian angle with evident setae (Figs. 20, 50). Legs: Protibiae narrower in males than in females (Fig. 23); in ventral view, longitudinal carina with a row of tubercles on basal one-half in males (Figs. 20 d, 23 b) and simple in females (Fig. 23 a). Protibial spur with apex strongly expanded and curved downward (Figs. 21, 23, 45); inner apical angle of protibiae with a tuft of setae longer in males than in females (Fig. 23). Apical protarsomere short, without spiniform projection (Fig. 46). Mesotarsi and metatarsi not particularly enlarged and with apical tarsomeres only slightly curved apically (Fig. 48). Metatibiae very broad and robust (Fig. 41). Metatibial spur with apex straight (Fig. 40). Elytra: Striae very fine and simple, not carinulate. Sutural margin densely punctate and clearly raised in its apical half in major specimens (Fig. 20 a, blue arrow); in basal third or basal half, sheen and punctation of sutural margin extend onto first or second interstriae (easily seen in Fig. 15 b). Abdomen: Pygidium flat, not margined basally (Fig. 36). Groove of propygidium extending to base of pygidium. Abdominal sternites densely punctate. Aedeagus: Phallobase, in ventral view, with apical membranous area only slightly wider (Fig. 55 c). Medial sclerite only slightly curved (Fig. 59). Measurements: Males (43 specimens): TL: AV: 13.49; MX: 17.4: MN: 9.3; SD: 1.65. PL: AV: 11.78; MX: 14.6; MN: 8.4; SD: 1.41. PW: AV: 8.08; MX: 10.5; MN: 5.6; SD: 1.04. Females (44 specimens): TL: AV: 13.56; MX: 16.6; MN: 9.9; SD: 1.46. PL: AV: 11.79; MX: 13.8; MN: 8.8; SD: 1.21; PW: AV: 8.15; MX: 9.8; MN: 6; SD: 0.88. Total (87 specimens): TL: AV: 13.53; SD: 1.55. PL: AV: 11.77; SD: 1.3. PW: AV: 8.11; SD: 0.96. Intraspecific variation and taxonomic discussion: Much of the intraspecific variation of G. inermis is a function of size. Very small specimens have the cephalic projection almost entirely reduced, showing only a carina slightly arched upward; in major specimens, on the other hand, this projection rises beyond that, but never as in G. aeruginosa and G. lemoinei (Fig. 32). In minor specimens the “ pronotal hump ” is not evident (Fig. 20 c), while in the major specimens the anterior region of the pronotum is notably swollen (Fig. 20 a, white arrow). The elevation of the sutural margin of elytra is more obvious in larger specimens than in small ones (Fig. 20 a, blue arrow). Most specimens of G. inermis have no posterior pronotal fossae, as in G. amazonica. However, we examined 11 specimens of both sexes that have a pair of weak circular impressions in the posterior region of pronotum, but differ from the posterior fossae of G. aeruginosa, G. lemoinei, and G. dichroa in position and being much smaller (Fig. 43). These specimens have “ partial length ” between 8.4 and 11.3 mm and pronotal width between 5.6 and 7.8 mm, i. e., measurements below the average for G. inermis. Interestingly, other specimens in the same size range do not have any trace of these fossae. D’Olsoufieff (1924) was the first to observe the presence of fossae in this species, claiming to have seen them in two small specimens from Mato Grosso, Brazil. Despite not having found any other morphological distinction with the rest of the species, he proposed the “ var. bipunctata ” for these specimens based only on the presence of these fossae (“ Deux petits exemplaires de Matto-Grosso ont deux fossettes basales du pronotum petites mais bien marquées, ce que je n’ai pas observé chez les autres. Le matériel manquant d’un côté et n’ayant pas trouvé d’autres différences avec les lacordairei typiques, je ne les compte pas en espèce distincte (…). Je me borne, pour les exemplaires cités, à leur donner le nom var. bipunctata nov. ”). Following the Article 45.6.4 of the Code, as the name “ bipunctata ” was expressly described as a variety by d’Olsoufieff, it must be regarded as of subspecific category and thus available. Nevertheless, as we observed that these fossae are present throughout the distribution of G. inermis (Porto Murtinho, Selvíria, Seropédica, Curitiba, Timbó, Torres, Santiago del Estero, Córdoba and Bañado de Medina), are always restricted to small specimens, and by the absence of any other distinctive characters, we believe that it is only a matter of individual variability and therefore propose synonymy with the nominotypical subspecies. Similarly, two specimens of G. amazonica examined by us also have a pair of these fossae; however, one has above average size for that species, indicating that, at least in G. amazonica, they are not restricted to small specimens (but nonetheless simple individual variability). Sexual dimorphism in this species is easily observable, contrary to Barattini & Saénz (1964): “ la morphologie externe de cette espèce n’est pas affectée par le sexe, seuls les caractères sexuels primaires permettent de les différencier ”. Among other sexual characteristics of Gromphas, it is possible to determine the sex of G. inermis by examining the ventral carina of the protibia, which is simple in females (Fig. 23 a) and with a row of tubercles in males (Figs. 20 d, 23 b). Interestingly, Barattini & Saenz (1964) mentioned this characteristic, but did not recognize it as a male sexual feature. Only in very worn specimens these tubercles are difficult to see. Unique features of G. inermis are: (a) posterior margin of pronotum projected at middle (Figs. 15 a – c); (b) metatibial spur straight at apex (Fig. 40) (in all other species it is distinctly curved [Fig. 39]); (c) and sutural margin of elytra clearly raised in its apical half (Fig. 20 a, blue arrow). The pronotal hump is also limited to large specimens of G. inermis (Fig. 20 a, white arrow). As in G. aeruginosa and especially G. lemoinei, the pronotum of G. inermis has granulation evident at center (in G. amazonica and G. dichroa, granulation is absent). Gromphas inermis shares only with G. lemoinei a densely punctate sutural margin of elytra and metasternum (in G. amazonica the punctation is also dense, but never at the same density of these two species). See comparison with G. amazonica and G. dichroa in the discussion of these respective species.	en	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): 439-482, DOI: 10.11646/zootaxa.3722.4.2
707787C659636261FF0A238D330E9A75.taxon	discussion	Comments: Although we have found only one specimen that was undoubtedly part of the type series of G. inermis, we know that Harold (1869 b) based his description in more than one specimen because he gave a size range (“ long. 12 – 15 mill. ”), mentioned sexual dimorphism and cited more than one collection locality (“ St. Catharina, Buenos Aires, Corrientes ” and “ G. inermis findet sich in Montivideo und Buenos Aires, dann im südlichen Brasilien von Porto Allegro bis Santa Catarina ”). Thus, we designate this specimen as the lectotype of G. inermis; the paralectotypes are unknown to us. D’Olsoufieff (1924) made clear that he based his description of G. lacordairei var. bipunctata in two specimens from Mato Grosso, Brazil. However, we were able to locate only one of these specimens at MNHN. Here, we designate this specimen as the lectotype of G. lacordairei bipunctata; the paralectotype is unknown to us. Judulien (1899) briefly described the larva of G. inermis. His observation that it bears five antennomeres contradicts Edmonds & Halffter (1978), who observed that all known larvae of Scarabaeinae have four antennomeres. Bionomics: Brullé (1837) was the first author to publish information on the natural history of Gromphas, probably referring to G. inermis. According to him, Lacordaire noted that this species burrows beneath horse dung. Subsequent authors (Judulien 1899, Fabre 1919, Halffter & Matthews 1966, Halffter & Edmonds 1982, Sánchez & Genise 2008) studied more thoroughly the nesting behavior of this species. Gromphas inermis has been also reported as a consumer of human, equine, bovine, and ovine dung in pastures (Lange 1947, Flechtmann & Rodrigues 1995, Flechtmann et al. 1995, Walsh & Cordo 1997, Morelli et al. 2002, Hamel-Leigue et al. 2006, Silva et al. 2009), and one specimen was collected by Louzada et al. (1996) using rotten cow spleen as bait. Flechtmann et al. (1995) and Monteresino et al. (1996) reported that G. inermis is diurnal. This species was also recognized as an intermediate host of the acanthocephalan worm Macracanthorhynchus hirudinaceus (Pallas, 1781), a parasite of the domestic pig, Sus scrofa Linnaeus, 1758 (Manter 1928, Lima 1953, Martínez 1959, Monteresino et al., 1996). Our data indicate that G. inermis is collected throughout the year. A curious pattern of edaphic preference was observed between G. inermis and Ontherus sulcator (Fabricius, 1775) in two studies. Flechtmann & Rodrigues (1995), studying the fauna of dung beetles of Jaraguá do Sul (Santa Catarina, Brazil), observed a clear differential distribution between these two species; while G. inermis was found only in flooded areas, O. sulcator was largely confined to sloping areas with no accumulation of water. Likewise, Sánchez & Genise (2008), working at Navarro (Buenos Aires, Argentina), noted that G. inermis was more abundant around a lake, while O. sulcator was predominant on higher ground. As pointed out by Flechtmann & Rodrigues (1995), it is likely that water saturation of the soil influences the spatial distribution of these species. Edaphic preference has been reported for other Phanaeini, in particular for North American species of Phanaeus (e. g., Fincher 1973, Blume & Aga 1978, Edmonds 1994, Rasmussen 1994). Geographic distribution: Amazon subregion: Pantanal. Chacoan subregion: Cerrado, Chaco, and Pampa. Parana subregion: Brazilian Atlantic Forest, Parana Forest, and Araucaria angustifolia Forest. BRAZIL: Bahia: Mucuri. Mato Grosso: Chapada dos Guimarães, Nobres, Poconé, Rosário Oeste, Santo Antônio de Leverger. Mato Grosso do Sul: Aquidauana, Campo Grande, Corumbá, Porto Murtinho, Selvíria. Minas Gerais: Pouso Alegre. Espírito Santo: Linhares, São Mateus (Ilha de Guriri). Rio de Janeiro: Armação de Búzios, Cabo Frio, Duque de Caxias, Rio de Janeiro, São Gonçalo, Seropédica. São Paulo: Araçatuba, Cássia dos Coqueiros, Indiana, Piracicaba, Ribeirão Preto, São José do Rio Preto. Paraná: Castro, Curitiba, Foz do Iguaçu, Ivaí, Lapa, Marechal Cândido Rondon, Ponta Grossa. Santa Catarina: Blumenau, Canoinhas, Corupá, Jaraguá do Sul, Joinville, Mafra, Rio Negrinho, Rodeio, São Bento do Sul, Seara, Timbó. Rio Grande do Sul: Aceguá, Bagé, Mostardas, Pelotas, Porto Alegre, Santo Augusto, Torres, Tramandaí. BOLIVIA: Beni. Santa Cruz: Chiquitos. Tarija (Tarija).	en	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): 439-482, DOI: 10.11646/zootaxa.3722.4.2
707787C659636261FF0A238D330E9A75.taxon	description	PARAGUAY: Amambay: Pedro Juan Caballero. San Pedro. Caaguazú: Caaguazú. Distrito Capital: Assunção. Paraguarí: Ybycuí. Guairá: Colonia Independencia, Villarrica. Caazapá: Caazapá. Itapúa: Coronel Bogado. ARGENTINA: Salta: Guachipas, Salta. Formosa: Guaycolec. Chaco. Santiago del Estero: Río Hondo (Termas do Río Hondo). Misiones. Corrientes: Ituzaingó, Santo Tomé. Santa Fé. Córdoba: Río Primero (La Para), San Justo (Miramar). Entre Ríos. Buenos Aires: Florencio Varela, La Plata, Navarro, Puán, San Isidro (Boulogne Sur Mer). Ciudad Autónoma de Buenos Aires. URUGUAY: Artigas. Rivera: Vichadero; Tacuarembó. Cerro Largo: Melo. Durazno. Flórida. Maldonado. Montevidéu. (Fig. 66).	en	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): 439-482, DOI: 10.11646/zootaxa.3722.4.2
707787C659636261FF0A238D330E9A75.taxon	materials_examined	Material examined: 244 males and 211 females (590 with undetermined sex). ARGENTINA: 1939, Parko col. — 1 female (MNRJ). BUENOS AIRES: La Plata, without date and collector — 3 males and 1 female (MLPA); Puán, F. Sola, I. 1959, A. Martínez col. — 2 males and 4 females (MZSP). CIUDAD AUTÓNOMA DE BUENOS AIRES: I. 1993, without collector — 1 male (CEMT). CHACO: without date and collector — 1 male (MZSP). CÓRDOBA: 16. III. 1939, without collector — 1 male (MNRJ); 04. IV. 1939, without collector — 1 male (MNRJ); Río Primero, La Para, X. 1986, Z. Monteresino col. — 1 male (CEMT). CORRIENTES: Santo Tomé, XII. 1925, without collector — 1 male (MLPA). FORMOSA: Guaycolec, II. 1949, A. Martínez col. — 2 males (MZSP). SALTA: Guachipas, I. 1951, A. Martínez col. — 1 female (MZSP). SANTIAGO DEL ESTERO: without date, Wagner col. — 4 males and 2 females (MLPA); Rio Salado, without date, Wagner col — 1 male (MLPA). BOLIVIA: TARIJA: Tarija, between Yaguacua-Caiza, 21 º 50 ’ 52 ’’ S 63 º 36 ’ 26 ’’ W, 620 m, 03. I. 2005, Mann, Hamel & Herzog cols. — 1 male and 1 female (CEMT). BRAZIL: BAHIA: Mucuri, I. 1999, G. L. D. Leite col. — 1 male (CEMT). ESPÍRITO SANTO: Linhares, X. 1995, T. M. Virgens col. — 1 female (CEMT). MATO GROSSO: Nobres, XI. 1985, without collector — 1 female (CEMT); Poconé, Fazenda Alvorada, 16 º 26 ’ 53 ” S 56 º 24 ’ 44 ” W, 21. XI. 2011, M. B. Pessoa col. — 1 male (CEMT); Rosário Oeste, without date and collector — 2 males and 2 females (MNRJ); Rosário Oeste, without date, Dirings col. — 1 male (MZSP); Santo Antonio do Leverger, Fazenda Vale Esperança, 20. X. 1990, Marinez Marques col. — 1 female (CEMT). MATO GROSSO DO SUL: Aquidauana, VII. 2011, C. M. A. Correa col. — 2 females (CEMT); Campo Grande, 1990 - 1992, I. Bianchin col. — 1 male (CEMT); Campo Grande, Jaraguá, XII. 1936, W. Zikán col. — 1 male and 1 female (MNRJ); Corumbá, BEP / UFMS, 19 º 33 ’ 53 ” S 57 º 00 ’ 39 ” W, 06. X. 2011, M. B. Pessoa col. — 6 males and 2 females (CEMT); Corumbá, Passo da Lontra, BEP / UFMS, without date, V. Lopes col. — 1 male (CEMT); Porto Murtinho, XII. 1929, without collector — 1 male (MZSP); Porto Murtinho, I. 1930, without collector — 1 male and 1 female (MZSP); Riacho do Herval, Rio Paraná, XII. 1951, B. Pohl col. — 4 males (MZSP); Selvíria, Fazenda UNESP, 21. XI. 1992, C. A. H. Flechtmann col. — 1 female (CEMT); Três Marias, margem esquerda do rio Sucuriú, Fazenda Caimã, X. 1966, F. Lane col. — 1 female (MZSP). MINAS GERAIS: Pouso Alegre, 27. XII. 1958, M. Vogas col. — 1 male (MNRJ). PARANÁ: Curitiba, III. 1898, without collector — 1 female (MZSP); Curitiba, 25. XII. 1936, C. Westerman col. — 3 males (MZSP); Curitiba, XI. 1941, B. Pohl col. — 2 males (MZSP); Curitiba, II. 1944, B. Pohl col. — 1 male (MZSP); Ivaí, Rio Ivaí, II (without year), W. Kosak col. — 1 female (MNRJ); Marechal Cândido Rondon, Porto Artaza, without date and collector — 1 female (MZSP); Parque Nacional do Iguaçu, XI. 1992, A. F. A. Luna Dias col. — 1 female (FIOC). RIO DE JANEIRO: Armação de Búzios, Praia Rasa, XII. 1995, L. H. Gil Azevedo col. — 1 sem sexo (DZRJ); Cabo Frio, VII. 1980, Jane M. Costa col. — 1 male (FIOC); Duque de Caxias, IX. 1990, F. Z. Vaz-de-Mello col. — 1 female (CEMT); Duque de Caxias, São Bento, VIII. 1960, P. A. Teles col. — 1 female (MNRJ); Rio de Janeiro, Copacabana, XI. 1990, F. Z. Vazde-Mello col. — 1 female (CEMT); Rio de Janeiro, Copacabana, XII. 1992, R. L. Vaz-de-Mello col. — 1 female (FIOC) and 1 female (CEMT); Rio de Janeiro, Copacabana, XII, 1993, F. Z. Vaz-de-Mello col. — 1 male and 1 female (CEMT); Rio de Janeiro, Copacabana, XII. 1994, R. L. Vaz-de-Mello col. — 1 female (FIOC) and 2 males (CEMT); Rio de Janeiro, Jacarepaguá, IX. 1990, F. Z. Vaz-de-Mello col. — 1 female (FIOC); Rio de Janeiro, Marapendi, XI. 1987, Hugo col. — 1 female (CEMT); São Gonçalo, X. 1982, Pêssoa col. — 1 sem sexo (DZRJ); Seropédica, Estrada Rio-São Paulo, km 47, 30. VII. 1951, J. F. Zikán Neto col. — 1 male (MNRJ); Seropédica, Universidade Federal Rural do Rio de Janeiro (UFRRJ), 21. X. 1990, A. Saraiva col. — 1 male and 1 female (FIOC); Seropédica, UFRRJ, 17. XI. 1990, without collector — 2 males (FIOC). RIO GRANDE DO SUL: III. 1915, without collector — 1 female (MZSP); Mostardas, I. 1945, Pe. Buck col. — 1 male (MZSP); Aceguá, Fazenda Centinela, 31 º 27 ’ 30 ” S 54 º 21 ’ 18 ” W, 08 - 14. XI. 2011, R. M. Moraes col. — 2 males and 5 females (CEMT); Bagé, Embrapa / CPPSUL, 06. XII. 2006, L. D. Audino col. — 1 female (CEMT); Bagé, Embrapa / CPPSUL, 17. I. 2007, L. D. Audino col. — 1 male (CEMT); Bagé, Fazenda Santo Antônio, 07 - 13. I. 2012, R. M. Moraes col. — 2 males and 2 females (CEMT); Pelotas, I. 1935, without collector — 1 male and 1 female (MNRJ); Pelotas, XII. 1934, without collector — 1 male and 1 female (MNRJ); Porto Alegre, without date and collector — 1 male (MNRJ); Porto Alegre, VI. 1927, without collector — 1 female (MZSP); Torres, 09. XII. 1964, Pe. P. Buck col. — 1 male (CEMT); Tramandaí, I. 1979, C. Coimbra Jr. col. — 1 male (MZSP). SANTA CATARINA: Blumenau, XI. 1924, without collector — 1 male (MZSP); Canoinhas, Pinhal, XI. 1951, A. Maller col. — 1 female (MNRJ); Canoinhas, Pinhal, XII. 1952, A. Maller col. — 2 females (MNRJ); Corupá, V (without year), Anton Maller col. — 1 male (MNRJ); Corupá, X (without year), Anton Maller col. — 1 male (MNRJ); Corupá, XII. 1953, A. Maller col. — 1 male (MNRJ); Joinville, VI. 1899, without collector — 1 female (MZSP); Mafra, without date and collector — 1 male (MNRJ); Rio Negrinho, XI. 1925, A. Maller col. — 1 male and 1 female (MNRJ); Rio Vermelho, without date, Dirings col. — 1 male and 1 female (MZSP); Rio Vermelho, IV. 1963, Dirings col. — 3 males and 1 female (MZSP); Rio Vermelho, III. 1964, Dirings col. — 1 female (MZSP); Rodeio, Rio Benedito, without date, Dirings col. — 6 males and 2 females (MZSP); Seara, Nova Teutônia, I. 1966, F. Plaumann col. — 1 male (MZSP); São Bento do Sul, I. 1948, Dirings col. — 1 male (MZSP); Timbó, II. 1952, Dirings col. — 42 with undetermined sex (MZSP); Timbó, V. 1956, Dirings col. — 22 males and 24 females (MZSP); Timbó, II. 1958, Dirings col. — 104 with undetermined sex (MZSP); Timbó, XI. 1965, Dirings col. — 161 with undetermined sex (MZSP); Timbó, XII. 1956, Dirings col. — 9 males and 11 females (MZSP); Timbó, V. 1962, Dirings col. — 37 males, 34 females and 190 with undetermined sex (MZSP); Timbó, III. 1965, Dirings col. — 78 males, 70 females and 92 outros with undetermined sex (MZSP). SÃO PAULO: Araçatuba, Anhangaí, XII. 1926, without collector — 1 male (MZSP); Cássia dos Coqueiros, II. 1955, M. P. Barreto col. — 2 males (MZSP); Indiana, 10. II. 1935, without collector — 1 female (MNRJ); Piracicaba, without date and collector — 1 male and 1 female (MZSP); Ribeirão Preto, XII. 1954, Barreto col. — 2 females (MZSP); Ribeirão Preto, VIII. 1955, E. X. Rabello col. — 1 espécime (MZSP); São José do Rio Preto, I. 1932, B. Pohl col. — 1 female (MZSP). PARAGUAY: AMAMBAY: Pedro Juan Caballero, XI. 1998, Maruíz Díaz col. — 2 males (CEMT). GUAIRÁ: Colonia Independencia, I. 1950, Foerster col. — 1 female (MNRJ); Colonia Independencia, III. 1950, Juan Foerster col. — 1 male (MNRJ). ITAPÚA: Coronel Bogado, I. 1944, Martínez col. — 1 female (FIOC). URUGUAY: without date and collector — 1 male (MZSP). ARTIGAS: Arroy Tres Cruces Grande, 18. I- 20. II. 1958, M. A. Monné col. — 1 female (MZSP). CERRO LARGO: Bañado de Medina, 13 - 20. XII. 2011, R. M. Moraes col. — 4 males and 4 females (CEMT); Melo, Fazenda La Invernada, 13 - 20. XII. 2011, R. M. Moraes col. — 1 male and 1 female (CEMT). MONTEVIDEO: Montevideo, without date and collector — 1 male (MZSP). RIVERA: Vichadero, Fazenda El Cerro, 21 - 27. XI. 2011, R. M. Moraes col. — 1 male and 1 female (CEMT).	en	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): 439-482, DOI: 10.11646/zootaxa.3722.4.2
