taxonID	type	description	language	source
3A7F7B2C7F778871FF34FAB32400EA32.taxon	diagnosis	Diagnosis. (based on Edmonds 1972, q. v.) Clypeus deeply, narrowly emarginated medially, bearing two prominent, acute median teeth united on lower side by U-shaped ridge; clypeal margin lateral to teeth often angulate. Eyes usually large (except Metallophanaeus), width of upper portion exceeding one-fourth of interocular distance, width of lower portion exceeding twice that of oculogular space. Circumnotal carina entire or broken behind eyes, but pronotal margin never excised to receive parietal lobes of head. Pronotum, at least anterolaterally, granulorugose, rugosities often coalescing into transverse ridging or scale-like sculpturing. Protibiae very strongly, acutely quadridentate. Protarsi absent in male, absent in female except some Megaphanaeus. Color most often dark, somber, with limited metallic highlights, seldom wholly metallic. Sexual dimorphism usually pronounced, but in different ways among subgenera and species groups (see below).	en	Edmonds, W. D., Zidek, J. (2010): A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 2010 (129): 1-111, DOI: 10.5281/zenodo.5352924
3A7F7B2C7F778871FF34FAB32400EA32.taxon	discussion	Comments. Members of this genus are easily distinguished from other Phanaeini by the combination of strong dentition of the tibiae and clypeus. The only other phanaeines likely to be confused with certain Coprophanaeus are those few robust species of Dendropaemon Perty, which, among other differences, possess only two or three meso- and metatarsal segments. Olsoufieff’s choice of the name for this group recognized “ … leur ressemblance avec les veritable Copris. ” As in most phanaeine groups, the taxonomy of Coprophanaeus is male-dominated. Nevertheless, the variation in the form and degree of sexual dimorphism in the genus is striking and much more complicated than in any other phanaeine group. The particulars of each case are treated in the appropriate taxonomic section below, but several cases suggest some interesting possibilities concerning the evolution of sexual dimorphism in this genus. In C. bellicosus, unlike other Megaphanaeus species, the male and female are quite different, especially in the form of the head. The male bears a long, tapering horn, whereas the female bears a trituberculate carina with enlarged central tubercle, similar to the saphirinus group of Metallophanaeus. Several other features suggest that C. bellicosus is an annectant between the two subgenera. Arnaud (2002 c), also struck by the uniqueness of this species, removed C. bellicosus from Megaphanaeus and placed it in Coprophanaeus s. str. (see Comments under C. bellicosus). Rowland and Emlen (2009) have provided new insights on the relationship between male horn polymorphism and reproductive strategy as exhibited primarily by the phanaeine dung beetles.	en	Edmonds, W. D., Zidek, J. (2010): A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 2010 (129): 1-111, DOI: 10.5281/zenodo.5352924
3A7F7B2C7F70886CFF34F8932296E952.taxon	diagnosis	Diagnosis. General – Width of lower portion of eye about twice that of oculogular space. Paraocular areas (genae) distinctly carinate lateral to eyes. Occipital areas of parietals with angulate prominence (Fig. 11, asterisk). Circumnotal ridge entire, not effaced behind eyes (as in Fig. 7). Pronotum completely sculptured, lacking smooth areas (except C. bellicosus); anterior surfaces transversely ridged, posterior surfaces granulorugose, becoming strongly granulate posteromedially (except C. bellicosus); basal fossae small, punctiform. Pronotum with thick, rounded ridge (Fig. 37, arrow) on each side extending ventrally from base of median prominence (reaching lateral fossa in C. bellicosus, Fig. 32, arrow). Striae (Fig. 13 - 16) conspicuous, always carinulate, carinulae straight or undulate. Hind wing not notched basally (as in Fig. 10). Posterior surface of each protibial tooth with basal, elongate, brush-like clump of densely packed, short setae (Fig. 5, arrow). Abdominal sterna clearly punctate along entire width; puncturing finer and sparser medially. Sexual dimorphism usually subtle, female secondary sexual features male-like. Male – Head bearing massive, posteriorly curved horn. Pronotum deeply concave; concavity transverse, posterior (dorsal) margin developed as massive prominence of varying shape. Parameres with dorsally directed apical, sometimes attenuated processes either rounded or acute in profile. Female – Protarsi present (except C. bonariensis). Head bearing massive, posteriorly curved horn like that of male (except C. bellicosus). Pronotum transversely concave; posterior (dorsal) margin of concavity developed as massive, saddle-shaped prominence usually broader than equivalent prominence in male (except C. bellicosus).	en	Edmonds, W. D., Zidek, J. (2010): A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 2010 (129): 1-111, DOI: 10.5281/zenodo.5352924
3A7F7B2C7F70886CFF34F8932296E952.taxon	distribution	Distribution. South America east of the Andes from the Amazon basin to northern Argentina comprising the Amazonian, Chacoan and Paranaian subregions of the Neotropical region.	en	Edmonds, W. D., Zidek, J. (2010): A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 2010 (129): 1-111, DOI: 10.5281/zenodo.5352924
3A7F7B2C7F70886CFF34F8932296E952.taxon	discussion	Comments. Megaphanaeus is used here exactly as conceived by Olsoufieff and followed by Blackwelder (1944) and Edmonds (1972). It includes the largest known phanaeines, C. ensifer and C. lancifer, some individuals of which can exceed 50 mm in length. The other two included species, C. bellicosus and C. bonariensis, are also large but can be equaled or exceeded in size by certain Sulcophanaeus, Diabroctis Gistel and members of the jasius species group of Coprophanaeus s. str. All four species are very well known and the absence of known types for three of the four is not a barrier to consistent use of the species names. Martínez’s (1944) review of the subgenus included six species, two of which Martínez and Pereira (1967) later synonymized. Our phylogenetic view of the subgenus has the species pair C. ensifer – C. bonariensis more closely related to C. lancifer than to C. bellicosus. The latter species is a taxonomic isolate that we place in a monobasic species group in apposition to the lancifer species group. Arnaud (2002 c) moved C. bellicosus to Coprophanaeus s. str. (see Comments below under C. bellicosus). The similar development and expression of secondary sexual features of the head and pronotum between the sexes is unique among “ armed ” phanaeines. The usual situation in Megaphanaeus might be called “ male dominant ”, because the female (except of C. bellicosus) has acquired a decidedly masculine form: a massive head horn and broad and massive saddle-shaped pronotal prominence. Large individuals clearly differ in details of pronotal shape, but medium-sized and small individuals are virtually identical. In two species, C. lancifer and C. ensifer, the female has retained protarsi, but in C. bonariensis, which lacks protarsi in both sexes, sexing these individuals requires dissection. In contrast, the females of C. bellicosus are easily recognizable by the tridentate cephalic process and presence of protarsi. Where present, female protarsi are very susceptible to loss, leaving only a very small empty socket at the base of the tibial spur. Determining sex in this subgenus using presence or absence of female protarsi therefore requires careful examination. The distribution of Megaphanaeus occupies much of the South American portion of the Neotropical region. The exclusive Amazonian representative is C. lancifer. Coprophanaeus bellicosus inhabits the Atlantic coastal forests (Paranaian subregion) of Brazil, while the C. ensifer – C. bonariensis pair splits the wide, xeric central swath of the continent (Chacoan subregion) from the cerrados to the Chaco thorn forests. The ranges of three of the four species (all but C. bellicosus) converge to within very short distances in the biodiversity hotspot, Parque Nacional Noel Kempff Mercado in northeastern Bolivia; but as far as we know, no two are ecologically sympatric (collected together in the same habitat). Besides the characters mentioned in the key, there are differences among the parameres of these species that were first pointed out by Lane and Carmargo-Andrade (1935). These are not treated in the species diagnoses below but are illustrated.	en	Edmonds, W. D., Zidek, J. (2010): A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 2010 (129): 1-111, DOI: 10.5281/zenodo.5352924
3A7F7B2C7F6D886EFF34F9D222B8ECB2.taxon	description	Fig. 16, 27 - 34	en	Edmonds, W. D., Zidek, J. (2010): A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 2010 (129): 1-111, DOI: 10.5281/zenodo.5352924
3A7F7B2C7F6D886EFF34F9D222B8ECB2.taxon	diagnosis	Diagnosis. General – Pronotum (Fig. 30 - 31) punctate posteromedially, not granulate. Posterolateral pronotal ridge long, extending to lateral pronotal fossa (Fig. 32, arrow). Metasternal shield largely smooth (as in Fig. 18); anterior portion of metasternum smooth. Elytral striae (Fig. 16) narrow, sides carinulate, straight, converging apically; interstriae 1 - 4 smooth, 5 - 7 finely rugose. Dorsal color black or dark brown, sometimes dark blue or bluish violet. Length 28 - 38 mm. Male (Fig. 31 - 33) – Pronotal concavity broad, shallow, lacking lateral protuberances. Dorsal prominence with narrow, median, scoop-shaped element. Apical process of parameres rounded in profile. Female (Fig. 29 - 30) – Cephalic process tridentate, middle tubercle much larger than laterals. Pronotal prominence a transverse, weakly bitumid, narrow process near posterior margin; pronotal concavity very broad, shallow, similar to that of male. Protarsi present. Specimens examined – 150.	en	Edmonds, W. D., Zidek, J. (2010): A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 2010 (129): 1-111, DOI: 10.5281/zenodo.5352924
3A7F7B2C7F6D886EFF34F9D222B8ECB2.taxon	distribution	Distribution. Atlantic coast forests of Brazil (Paranaian subregion) (Fig. 28). Collection Records. BRAZIL: Bahia – Entre Rios (Jan); Encruzilhada (Dec). Espiritu Santo – Sooretama (Jun, Nov); Tijuco Prêto (Dec); Fazenda Lagoa do Macuco [Municipio Linhares], 19 o 03 ’ 50 ” S 39 o 58 ’ 43 ” W (Jan); Timbuhy. Minas Gerais – Viçosa [Mata do Paraiso], 20 o 48 ’ 18 ” S 42 o 51 ’ 20 ” W, 750 m (Feb, Nov). Rio Grande do Sul – São Alberto (Jan). Rio de Janeiro – Nova Friburgo (Jan, Oct); 17 km E Nova Friburgo, 22 o 23 ’ 04 ” S 42 o 33 ’ 30 ” W, 750 m (Jan). Santa Catarina – Joinville (Nov-Dec); Tijuca Forest (Jan). São Paulo – São João dos Campos (Feb); Nova Cantareira (Apr); 50 km SE Mogi das Cruzes [Serra do Mar Biological Station “ Boraceia ”], 800 - 900 m (Apr); Tremembé (Dec).	en	Edmonds, W. D., Zidek, J. (2010): A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 2010 (129): 1-111, DOI: 10.5281/zenodo.5352924
3A7F7B2C7F6D886EFF34F9D222B8ECB2.taxon	discussion	Comments. Arnaud (2002 c) transferred this species to the subgenus Coprophanaeus s. str. stating only that, while it was isolated from other members of that subgenus, it could not “ … en aucune manière, si ce n’est sa taille, être associé au groupe lancifer. ” We disagree and maintain its placement in Megaphanaeus because of the suite of characters it shares with C. lancifer, C. ensifer and C. bonariensis (see key to subgenera). But we do agree that it is a taxonomic isolate within the subgenus with, in our view, a closer affinity with the saphirinus group of Metallophanaeus than with any constituent of Coprophanaeus s. str. In this context, the affinity is suggested by secondary sexual characteristics of both sexes and the texture of the elytra. Interestingly, Nevinson (1892) in his “ Suggested order of the species, and the groups into which they more-or-less naturally fall, ” proposed C. bellicosus as an isolated annectant between the lancifer and saphirinus groups. To Martínez (1944) the females of C. bellicosus “ … se asemejan a ciertos [two male symbols] del subgenero Coprophanaeus del grupo jasius. ” Coprophanaeus bellicosus is a denizen of the Atlantic coast forests of Brazil, and its range, along with that of C. cerberus, has undoubtedly contracted in recent decades because of widespread agriculturization and urbanization of Brazil’s eastern seaboard.	en	Edmonds, W. D., Zidek, J. (2010): A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 2010 (129): 1-111, DOI: 10.5281/zenodo.5352924
3A7F7B2C7F6F8868FF34FA922335EEB2.taxon	description	Fig. 13, 17 - 18, 23 - 26, 28, 35 - 39	en	Edmonds, W. D., Zidek, J. (2010): A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 2010 (129): 1-111, DOI: 10.5281/zenodo.5352924
3A7F7B2C7F6F8868FF34FA922335EEB2.taxon	materials_examined	Type. S. lancifer – unknown to us; P. heros – unknown to us; P. miles – unknown to us; S. satelles – unknown to us; P. septentrionalis – holotype female, Museu de Zoologia, Universidade de São Paulo, São Paulo.	en	Edmonds, W. D., Zidek, J. (2010): A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 2010 (129): 1-111, DOI: 10.5281/zenodo.5352924
3A7F7B2C7F6F8868FF34FA922335EEB2.taxon	diagnosis	Diagnosis. General – Pronotum (Fig. 25 - 26, 35, 38) strongly granulate posteromedially. Posterolateral pronotal ridge (Fig. 37, arrow) short, not extending to lateral pronotal fossa. Metasternal shield (Fig. 18) smooth centrally, punctate peripherally; anterior portion of metasternum completely smooth, shiny medially, finely granulate laterally. Basal two-thirds of elytral striae 2 - 4 moderately undulate, width at widest points about one-half of that at narrowings; carinulate margins usually tuberculate at narrowings Fig. 13). Interstriae granulorugose, granulation, if distinct, confined to apical one-third of interstriae 2 - 6. Dorsal color usually dark blue, rarely green or bluish green. Length 28 - 50 mm. Male (Fig. 23 - 24, 37 - 39) – Pronotal concavity flanked on each side beneath central prominence by sharp, oblique carina (Fig. 23, arrow). Dorsal prominence wide, width of saddle usually much greater than length. Apical process of parameres acute in profile. Female (Fig. 25 - 26, 35 - 36) – Protarsi present. Pronotal concavity lacking oblique carinae. Specimens examined – 351.	en	Edmonds, W. D., Zidek, J. (2010): A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 2010 (129): 1-111, DOI: 10.5281/zenodo.5352924
3A7F7B2C7F6F8868FF34FA922335EEB2.taxon	distribution	Distribution. Amazonian subregion of the Neotropical region (Fig. 28). Collection Records. BOLIVIA: Beni – Magdalena (Apr). Pando – Cobija [Reserva San Sebastian Tahuamanu], 11 o 24 ’ 27 ” S 69 o 01 ’ 04 ” W (Dec); Guayaramerin (Dec); near Guayamerin, 11 o 50 ’ S 65 o 22 ’ W, 120 m (Feb, Dec); 2 km E Fortaleza, 9 o 47 ’ S 65 o 30 ’ W, 120 m; Río Negro, 9 o 52 ’ S 65 o 42 ’ W, 120 m (Feb); 20 km SW Villa Bella, 10 o 22 ’ S 65 o 22 ’ W, 120 m (Feb); Tahuamanu, 11 o 24 ’ 27 ” S 69 o 01 ’ 04 ” W, 280 m; Malecón, 11 o 57 ’ S 68 o 48 ’ W, 190 m; Florida, 12 o 18 ’ S 68 o 40 ’ W, 190 m (Nov); Río Negro, 9 o 52 ” S 65 o 42 ’ W, 120 m (Feb); near Bella Vista, 10 o 22 ’ S 65 o 22 ’ W, 120 m; Santa Rosa, 12 o 00 ’ S 68 o 52 ’ W, 180 m (Oct). BRAZIL: Amazonas – 30 km N Manaus [Reserva Forestal Ducke] (Mar- May, Aug); Manaus (Jan, Mar-Apr, Jul, Dec); 60 km N Manaus [Fazenda Esteio] (Jun); Manacapurú (Mar); São Paulo de Olivença (May, Jul, Dec); Tefé (Aug). Mato Grosso – Rosário Oeste (Jul); Nobres (Jan, Jul). Pará – Belêm [Agua Preta] (Jan-Feb); Tucuruí; Obidos (Mar-May, Jul, Nov-Dec); Santarém (Jan, May); Taperinha; Monte Dourado [Apui], 00 o 46 ’ S 52 o 35.5 ’ W (Jan); Monte Dourado [Tingilingi], 00 o 57 ’ S 52 o 45.5 ’ W (Apr). Rondônia – 9 km NE Cacaulândia (Feb, Nov) GUYANA: East Berbice-Corentyne – Bartica (May). Upper Demerara-Berbice – Ituni (Jul). PERU: Madre de Díos – Río Madre de Díos, 12 o 34 ’ 10.0 ” S 70 o 06 ’ 01.4 ” W, 290 m (Apr); Río Tambopata, 12 o 38 ’ 59 ” S 69 o 06 ’ 24 ” W, 230 m (Sep); Río Palma Real Grande, 12 o 32 ’ 20 ” S 68 o 51 ’ 40 ” W, 220 m; 30 km SW Puerto Maldonato [Río Tambopata Reserve], 12 o 50 ’ S 69 o 20 ’ W. 290 m. SURINAME: Commewijne – Akintosoela, CELOS Camp, 39 km SE Suriname river bridge, road to Redi Dodi, 40 m, 5 ° 16 ’ 17 ” N 54 ° 55 ’ 15 ” W (Jul). Sipaliwini – Lely Plateau, 46 ’ 13 " N 54 o 44 ’ 18 " W, 650 m (Oct); Oelemarie, ~ 3 o 6 ’ 0 ' ’ N 54 o 32 ’ 00 ' ’ W (Feb, Aug, Oct); Palumeu, ~ 3 o 21 ’ 30.3 ' ’ N 55 o 26 ’ 38.2 ' ’ W (Jun): Kwamalasumutu (Jul). VENEZUELA: Amazonas – Río Mavaca camp, 2 o 02 ’ N 65 o 06 ’ W. Bolívar – Las Trincheras [Río Cauca] (Aug); Puerto Cabello (Río Cauca] (Aug).	en	Edmonds, W. D., Zidek, J. (2010): A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 2010 (129): 1-111, DOI: 10.5281/zenodo.5352924
3A7F7B2C7F6F8868FF34FA922335EEB2.taxon	discussion	Comments. This is a common species occurring sometimes is large numbers throughout much of the Amazon basin. Among all Coprophanaeus species, C. lancifer is the best choice as indicator species of Amazonia and is an important component of many local dung beetle communities (see, for example, Gardner et al. 2008, and Quintero and Halffter 2009). We have seen no specimens from eastern Ecuador and Colombia, but its absence there is unlikely. The color of this species is usually a consistent dark metallic blue; Trond Larsen has collected bright green examples in southeastern Peru. Besides the commonly used baits of dung and carrion, this species is also attracted to decomposing millipedes (Conrad Gillett, pers. comm.) Théry et al. (2008) interpreted the crepuscular flight activity of this species as a mechanism that improves visual communication among conspecifics. At mid-range (> 0.5 m) in the low-intensity dusk lighting, the dark blue, metallic (structural) body coloration is seen at higher contrast against ambient background and active individuals become more visible to each other; at shorter range (<0.5 m), enhancement of contrast between the black head horn and metallic pronotum probably functions in species recognition. Except for the presence of protarsi, medium-sized and small females closely resemble equal-sized males. Pessôa’s (1934) description of C. septentrionalis was based on five females; Lane and Camargo- Andrade (1935) re-examined the type series and found that it included one male. The pronotal prominence of large females (Fig. 25 - 26) is subject to considerable variation in width, but not as extreme as in C. ensifer (q. v.). This species is the first Coprophanaeus depicted in print, as far as we know. Voet’s (1766) illustrations (pl. 23, fig. 1 - 2 and 38) are unmistakably of C. lancifer. His fig. 1 and 2 are cited by Linné (1767) in his description of Scarabaeus lancifer and can be regarded as at least part of the type series. Voet’s fig. 38 is of interest because, as Olsoufieff (1924) pointed out, it depicts the body of a male C. lancifer “ … mais avec une tête (?) d’un autre Scarabaéide. ” In the text caption Voet stated that [specimen 38] “ Forgan haec femina est No. 1 vel 2 ” [“ Could be the female of either N. 1 or 2 ”]. Herbst’s (1789) illustration (pl. VIII, fig. 6, under the name Hamadrias) is a copy of Voet’s “ hybrid; ” but his pl. XV, fig. 1 depicts a male C. lancifer placed correctly under the name “ Scarab [aeus] lancifer. ”	en	Edmonds, W. D., Zidek, J. (2010): A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 2010 (129): 1-111, DOI: 10.5281/zenodo.5352924
3A7F7B2C7F6A886BFF34FF72224AEF52.taxon	description	Fig. 15, 28, 40 - 44	en	Edmonds, W. D., Zidek, J. (2010): A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 2010 (129): 1-111, DOI: 10.5281/zenodo.5352924
3A7F7B2C7F6A886BFF34FF72224AEF52.taxon	materials_examined	Type. P. bonariensis – unknown to us; P. vicinus s. st r. holotype male, Museo Argentino de Ciencias Naturales “ Bernadino Rivadavia, ” Buenos Aires; P. vicinus argentinus, holotype male, Museo Argentino de Ciencias Naturales “ Bernadino Rivadavia, ” Buenos Aires.	en	Edmonds, W. D., Zidek, J. (2010): A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 2010 (129): 1-111, DOI: 10.5281/zenodo.5352924
3A7F7B2C7F6A886BFF34FF72224AEF52.taxon	diagnosis	Diagnosis. General – Pronotum strongly granulate posteromedially. Posterolateral pronotal ridge (Fig. 40, cf. Fig. 37) short, not extending to lateral pronotal fossa. Metasternal shield completely coarsely punctured (as in Fig. 19); anterior portion of metasternum finely granulate laterally, smooth medially. Carinulate margins of striae 2 - 4 gently undulate such that width of stria at narrowings is at least threefourths of that at widest points (Fig. 15); carinulae simple, rarely tuberculate. Interstriae completely covered by large, round granules. Dorsal color usually shade of green or yellowish green. Length 21 - 35 mm. Male (Fig. 42 - 44) – Pronotal concavity with dorsally inclined spiniform process on each side below prominence (Fig. 43). Dorsal prominence narrow, scoop-shaped, width of saddle less than length (Fig. 42). Apical process of parameres acute in profile. Female (Fig. 40 - 41) – Pronotal concavity lacking lateral spines. Protarsi lacking. Specimens examined – 222.	en	Edmonds, W. D., Zidek, J. (2010): A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 2010 (129): 1-111, DOI: 10.5281/zenodo.5352924
3A7F7B2C7F6A886BFF34FF72224AEF52.taxon	distribution	Distribution. Cerrado and Chaco provinces of Chacoan subregion (Fig. 28). Collection Records. ARGENTINA: Catamarca – Catamarca (Feb). Chaco – Miraflores (Apr). Córdoba – San Javier, Las Rosas (Jan); Quilino (Dec). Corrientes – Bella Vista (Feb); Ituzaingo (Jan, Nov); Ituzaingo [Alto Paraná] (Sep). Formosa – Gran Guardia (Jan). Jujuy – 6 km W Yuto (Feb); El Quemado (Apr). Misiones – Ayolas (Oct). Santiago del Estero – Termas de Río Hondo (Dec); Santiago del Estero (Feb); Río Salado; Campo Gallo (Mar). Tucuman – Trancas (Jan); Benjamin Araoz (Mar); Gobernador Garmendia (Jan). BOLIVIA: Beni – Magdalena (Feb). Santa Cruz – San Matias, 16 o 34 ’ S 59 o 09 ’ W; Santa Cruz de la Sierra; Ichila, 400 m; Charagua, 20 o 14 ’ 11 ” S 63 o 13 ’ 52 ” W, 960 m (Apr); Urubigua, 20 o 14 ’ 11 ” S 63 o 13 ’ 52 ” W; Buena Vista (Jan); La Peña (Nov); Cabezas (Jan); Santiago de Chiquitos, 18 o 20 ’ S 59 o 36 ’ W (Feb); Santa Elena El Pantanal, 17 o 57 ’ 53 ” S 59 o 01 ’ 33 ” W (Apr); 1.6 km ESE Santiago de Chiquitos, 18 o 20.103 ’ S 59 o 35.007 ’ W, 622 m (Nov); 20 km NE Santiago de Chiquitos (Tucavaca River), 18 o 14.590 ’ S 59 o 27.907 ’ W, 215 m (Nov). Tarija – between Caiza and Creveaux, 21 o 50 ’ 19 ” S 63 o 24 ’ 58 ” W, 490 m (Jan). BRAZIL: Mato Grosso – 5 km S Cuiabá [Várzea Grande] (Dec); Rosario Oeste (Feb). PARAGUAY: Boquerón – Estación La Patria [trans-Chaco highway] (Mar, Dec); Nueva Asunción, 20 o 48 ’ S 61 o 55 ’ W; Mariscal Estigarribia (Feb, May). Central – Aregua (Dec); Altos; Colonia Nueva Italia (Feb); Asunción (Nov); Puerto Villeta (Jan). Concepción – Horqueta (Nov). Cordillera – Altos (Apr); Compañia Narajo (Nov). Misiones – Ayolas (Oct); Paraguari – Sapucay (Jan- Apr, Oct-Dec). Presidente Hays – Laguna Capitán (Jan).	en	Edmonds, W. D., Zidek, J. (2010): A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 2010 (129): 1-111, DOI: 10.5281/zenodo.5352924
3A7F7B2C7F6A886BFF34FF72224AEF52.taxon	discussion	Comments. This species largely replaces C. ensifer in the chiquitano, cerrado and chacoan habitats of eastern Bolivia and surrounding portions of Argentina, Paraguay and Brazil. We here correct the record concerning female protarsi in C. bonariensis; contrary to Edmonds (1972), they are absent in this species. Consequently, medium-sized females and smaller males are virtually indistinguishable externally.	en	Edmonds, W. D., Zidek, J. (2010): A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 2010 (129): 1-111, DOI: 10.5281/zenodo.5352924
3A7F7B2C7F6B8865FF34F90024B6EF72.taxon	description	Fig. 5, 11, 14, 19 - 22, 28, 45 - 49	en	Edmonds, W. D., Zidek, J. (2010): A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 2010 (129): 1-111, DOI: 10.5281/zenodo.5352924
3A7F7B2C7F6B8865FF34F90024B6EF72.taxon	materials_examined	Type. C. ensifer – unknown to us; P. ajax – unknown to us; P. ducalis – unknown to us.	en	Edmonds, W. D., Zidek, J. (2010): A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 2010 (129): 1-111, DOI: 10.5281/zenodo.5352924
3A7F7B2C7F6B8865FF34F90024B6EF72.taxon	diagnosis	Diagnosis. General – Pronotum strongly granulate posteromedially (Fig. 20 - 21, 46 - 47). Posterolateral pronotal ridge short, not extending to lateral pronotal fossa (Fig. 45, cf. Fig. 37). Metasternal shield completely, coarsely punctured (Fig. 19); anterior portion of metasternum completely finely granulate. Carinulate margins of striae 2 - 4 strongly undulate such that width of stria at narrowings is no more than one-half (usually much less) of that at widest points; carinulate margins strongly tuberculate at narrowings (Fig. 14). Interstriae completely covered by large, round granules. Dorsal color usually shade of green or bluish green, rarely blue. Length 30 - 56 mm. Male (Fig. 22, 47 - 49) – Pronotal concavity with dorsally inclined spiniform process on each side below prominence (Fig. 22). Dorsal prominence wide, width of saddle much greater than length. Apical process of parameres rounded in profile. Female (Fig. 20 - 21, 45 - 46) – Pronotal concavity lacking spiniform processes. Protarsi present. Specimens examined – 133.	en	Edmonds, W. D., Zidek, J. (2010): A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 2010 (129): 1-111, DOI: 10.5281/zenodo.5352924
3A7F7B2C7F6B8865FF34F90024B6EF72.taxon	distribution	Distribution. Caatinga and Cerrado provinces of Chacoan subregion (Fig. 28). Collection Records. ARGENTINA: Misiones – Loreto. BOLIVIA: Santa Cruz – Parque Nacional Noel Kempff Mercado [Los Fierros], 14 o 33 ’ 28 ” S 60 o 55 ’ 51 ” W and 15 o 15 ’ 6.3 ” S 61 o 14 ’ 41 ” W (Jan-Feb); Caparú, 14 o 50 ’ S 61 o 10 ’ W; Las Conchas, 17 o 34 ’ S 59 o 28 ’ W; La Guardia, 17 o 53 ’ S 63 o 20 ’ W; Basilio, 18 o 07 ’ S 63 o 12 ’ W; Estancia Perforación, 19 o 45 ’ S 62 o 00 ’ W; 20 km SW Perseverancia, 14 o 44 ’ S 62 o 47 ’ W, 230 m. BRAZIL: Alagoas – Maçeio (Jul). Bahia – Encruzilhada (Dec). Distrito Federal – Brasilia, 600 m (Nov-Dec). Goias – Bananeiras (Jan). Mato Grosso – Rio Verde (Nov); Fazenda São João, 14 o 14 ’ 10 ” S 56 o 08 ’ 11 ” W, 400 m (Jan); Arinos (Nov). Mato Grosso do Sul – Piraputanga (Mar); Selvíria (Jan). Minas Gerais – Lagoa Santa; Machacalis (Dec); Uberlândia (Dec). Paraíba – Mamanguape [Rebio Guaribas], 6 o 41 ’ S 35 o 10 ’ W; João Pessoa [Mata do Buraquinho] (May). Pernambuco – Recife [campus Universidade Federal de Pernambuco] (Aug). São Paulo – Assis (Dec); Descalvado (Mar); Colina (May); Itirapina (Feb); Ilha Solteira (Feb); São José do Rio Preto (Dec); Riberão Preto (Feb); Bálsamo (Nov); Rio Claro (Feb-Mar); Araraquara (Oct); Urupês (Nov); Planalto (Apr); Limeira (Feb); São José dos Campos (Oct-Nov); Rio Preto (Dec); Franca (Nov); Mogi Guaçu [Fazenda Campininas] (Jan); Campinas (Nov-Dec). PARAGUAY: Alto Paraná – Limoy (Sep-Oct). Anambay – Cerro Cora Feb, Dec). Caaguazú – Caaguazú (Nov-Dec). Concepción – Cororó (Nov). Guairá – Villarrica (Dec); Independencia; Melgarejo (Sep). Kanindeyu – Carapa (Apr).	en	Edmonds, W. D., Zidek, J. (2010): A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 2010 (129): 1-111, DOI: 10.5281/zenodo.5352924
3A7F7B2C7F6B8865FF34F90024B6EF72.taxon	discussion	Comments. Endres et al. (2005) studied the phenology of C. ensifer in Paraíba (Brazil) and found that adult abundance and activity are greatest during the April – September rainy season and that beetles showed no preference for carrion type used to bait traps. Otronen (1988) provided observations and interpretations on horn variation and combat in this species in both sexes. Fighting in C. ensifer occurs both intra- and intersexually. Males fight males to defend or gain resources and to win females; females fight females and males to defend their burrows. This species varies in color. The usual tone is dark metallic green, often with contrasting bluish reflections. Pessôa (1934) reported rare metallic blue individuals from Lussanvira, São Paulo, and bluishgreen individuals from Mato Grosso. A very dark bluish-green form occurs in the gallery forests of northern Mato Grosso (Querencia, Diamantino, Trivelato), where the species has evidently invaded a decidedly Amazonian habitat; the male of this form has a small median tooth between larger lateral ones born apically on the median projection of the pronotal prominence (Fernando Vaz-de-Mello, pers. comm.). The female pronotal prominence takes two forms. The summit of the more frequent one is broadly saddle-shaped (Fig. 20). In the less frequent condition the sides of the median saddle are closely appressed, producing a fluted salience (Fig. 21). Both forms occur together in the region of Brasilia.	en	Edmonds, W. D., Zidek, J. (2010): A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 2010 (129): 1-111, DOI: 10.5281/zenodo.5352924
3A7F7B2C7F658866FF34F9E42142E9B2.taxon	diagnosis	Diagnosis. General – Lower portion of eye small, width no more, and usually less, than twice that of oculogular space. Paraocular areas (genae) carinate or not lateral to eyes. Occipital areas of parietals more-or-less flattened, lacking angulate prominence (as in Fig. 12). Circumnotal ridge entire, not effaced behind eyes (as in Fig. 8). Pronotal sculpturing variable, usually granulorugose anterolaterally, punctate to some degree posteromedially; basal fossae variable. Pronotum lacking posterolateral ridge. Striae fine, simple or carinulate, anterior ends of 1 - 4 impressed to some degree. Hind wing not notched basally (Fig. 10). Posterior surface of protibial tooth with single basal row of long setae (as in Fig. 6). Abdominal sterna usually smooth medially. Sexual dimorphism marked. Male – Head bearing posteriorly curved horn. Pronotum variable, flattened or transversely concave anteromedially. Parameres variable. Female – Protarsi absent. Pronotum variable, usually with transverse process near anterior margin.	en	Edmonds, W. D., Zidek, J. (2010): A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 2010 (129): 1-111, DOI: 10.5281/zenodo.5352924
3A7F7B2C7F658866FF34F9E42142E9B2.taxon	distribution	Distribution. Eastern Brazil (east of Amazonia) southward to northeastern Argentina.	en	Edmonds, W. D., Zidek, J. (2010): A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 2010 (129): 1-111, DOI: 10.5281/zenodo.5352924
3A7F7B2C7F658866FF34F9E42142E9B2.taxon	discussion	Comments. This subgenus brings together eight mostly colorful species distributed between two different species groups recognized by Edmonds (1972) and Arnaud (2002 c). Only six of the eight are treated in the key and species accounts (see below). As far as we know, all Metallophanaeus species are diurnal and copronecrophagous. The subgenus as here constituted is undoubtedly paraphyletic. Affinities to Megaphanaeus (mainly via C. bellicosus, q. v.) are clearly suggested in the saphirinus group by, in particular, similarity in male secondary sexual characters (form of head and pronotum and apically hooked parameres). Affinities to the jasius group of Coprophanaeus s. st r. are equally demonstrable in the thalassinus group by male and female secondary sexual characters (male head horn and pronotal shape; swollen tips of parameres; pronotal ridge and oval concavity in female). A cogent argument can be made to isolate the two species groups of Metallophanaeus into separate subgenera. Indeed, Olsoufieff included only C. saphirinus and C. horus in this subgenus, which he defined mainly on the basis of the male pronotum; C. pertyi, C. punctatus (of which Olsoufieff had no male) and C. thalassinus he placed in Coprophanaeus s. str. We have combined the two groups here mainly because both lack (by symplesiomorphy, we presume) the notching of the hind wing (a synapomorphy) characteristic of the nominate subgenus. Our preference now is to maintain the current, probably artificial, inclusion of both groups in Metallophanaeus; any decision otherwise should await a more formal phylogenetic analysis of the entire Coprophanaeus lineage, including Dendropaemon. The salient differences between the two species groups recognized here reside in secondary sexual characters and in the shape of the anterior portion of the metasternum. Regarding the latter, the thalassinus group exhibits a tendency for the anterior metasternal angle (viewed in profile) to be enhanced to some degree – either squared or obviously prolonged anteriorly in contrast to the evenly curved profile seen in the saphirinus group. Accentuation of the anterior metasternal angle of the thalassinus group (supposedly most developed in C. vazdemeloi) correlates with a tendency for the profemora to be swollen posteriorly, and for the swollen posterior surface to be divided longitudinally into a flat, glabrous area and a rounded setose area behind it. Sexual dimorphism is striking but expressed differently in the two groups. In the saphirinus group, as in Phanaeus, Sulcophanaeus and Oxysternon, the male possesses a long, gradually tapering, cylindrical head horn and prominent posteromedian pronotal salience; the female, a trituberculate cephalic carina and a convex pronotum with subtle relief and (usually) a transverse crest near the anterior margin. In the thalassinus group, the male head horn is proportionately shorter with a distinctly swollen base and the pronotum bears a transverse ridge; the female has a trituberculate cephalic carina and the pronotum bears a strong anteromedian crest followed by an oval concavity (as in many species of Coprophanaeus s. str.). Protarsi are absent in both sexes of all Metallophanaeus species. We are not able to treat formally in this review two undoubtedly valid species belonging to the thalassinus group. One is C. vazdemeloi Arnaud (2002 a), known from the holotype male and one paratype female (BRAZIL: Piauí – São Raimundo Nonato, Parque Nacional Serra de Capivara [Jan]) housed in the private collection of Patrick Arnaud. We have been unable to examine these specimens, which are reported to be unique among the thalassinus group for the very strong prolongation of the anteromedian angle of the metasternum. Since we have no way to reliably assess other characters of this species and to formally compare it with other members of the species group, we are obliged to exclude it from formal consideration at this time. We refer the reader to the original description and to the photographs and comments in Arnaud (2002 c). The key and species treatments below also do not consider an eighth species from the Chapada do Parecis (northern Mato Grosso, Brazil) assignable to the thalassinus group and currently under study by Fernando Vaz-de-Mello and Patrick Arnaud. It will key out below to C. thalassinus, from which it differs by the following combination of characters: new species – a) posteromedian portion of pronotum granulorugose; b) paraocular areas flat; c) abdominal sterna distinctly punctured medially; d) elytra golden with green reflections basally and along elytral suture; pronotal disk bluish green, sides and head golden; antennal club golden; e) from Mato Grosso; f) all elytral interstriae more-or-less evenly convex; first interstria not more strongly raised than others. Coprophanaeus thalassinus differs from the new species as follows: a) posteromedian portion of pronotum with punctures grading to fine ridges to scalelike granules around basal fossae; b) posterior portion of paraocular areas convex; c) abdominal sterna smooth medially at least along midline; d) dorsum metallic green and dark blue; antennal club black; e) from Bahia; f) elytral interstriae weakly convex except first; first more strongly raised and shinier than others.	en	Edmonds, W. D., Zidek, J. (2010): A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 2010 (129): 1-111, DOI: 10.5281/zenodo.5352924
3A7F7B2C7F60885DFF34FA3222DEEB72.taxon	description	Fig. 10, 56, 60, 62 - 68	en	Edmonds, W. D., Zidek, J. (2010): A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 2010 (129): 1-111, DOI: 10.5281/zenodo.5352924
3A7F7B2C7F60885DFF34FA3222DEEB72.taxon	materials_examined	Type. P. horus – holotype male, The Natural History Museum, London (examined); P. sericeus – unknown to us.	en	Edmonds, W. D., Zidek, J. (2010): A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 2010 (129): 1-111, DOI: 10.5281/zenodo.5352924
3A7F7B2C7F60885DFF34FA3222DEEB72.taxon	diagnosis	Diagnosis. General – Posterior portion of paraocular areas (Fig. 60) strongly transversely raised in front of eyes (swelling sometimes ridge-like), coarsely punctured in front of swelling. Pronotum smooth posteromedially, otherwise weakly granulorugose; posterior angle normal, basal fossae large, conspicuous, incorporated into inner ends of sulcus paralleling posterior margin. Anteromedian angle of metasternum (seen in profile, as in Fig. 59) rounded, not salient; angle capped with elongate thickening (bead). Striae (x 5) simple, weakly impressed; bases of striae 1 - 4 impressed, not distinctly fossate. Elytral interstria usually distinctly convex, first, second and / or fourth often more strongly so. Ventral surface of protibia lateral to longitudinal carina paralleling inner margin entirely punctatorugose. Pygidium (Fig. 56) with broad basal groove. Length 15 - 22 mm. Color green or yellowish green with strong golden highlights. Male (Fig. 63 - 65) – Head with evenly tapering long horn whose base is not abruptly swollen. Pronotum with pair of closely set, apically convergent processes near posterior margin, separated by strong concavity; disk anterior to processes with pair of conspicuous round concavities. Pronotal disk punctate posteromedially, otherwise granulorugose. Apical processes of parameres angulate. Female (Fig. 67 - 68) – Pronotum with transverse, weakly margined ridge followed by midlongitudinal groove. Pronotum finely granulorugose on disk and sides, punctate posteromedially. Specimens examined – 31.	en	Edmonds, W. D., Zidek, J. (2010): A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 2010 (129): 1-111, DOI: 10.5281/zenodo.5352924
3A7F7B2C7F60885DFF34FA3222DEEB72.taxon	distribution	Distribution. Chaco and Cerrado provinces (Fig. 62). Collection Records. ARGENTINA: Buenos Aires – General Sarmiento (Jan); General Belgrano. Córdoba – Cruz Alta (Feb, Nov); Leones (Feb). Misiones – Loreto (Jan, Nov). Tucuman – Los Ralos. BRAZIL: Distrito Federal – Brasilia (Nov). Minas Gerais – 100 km SSE Lavras [Carrancas, Chapada das Perdízes] (Dec). Paraná – Mamborê (Feb). Rio Grande do Sul – Canôas (Oct); Colorado (Dec); Botucatu (Feb). PARAGUAY: Boquerón – Nueva Asunción (Jan, Apr, Dec). Caaguazú – Caaguazú (Dec). Presidente Hays – Laguna Capitan (Jan).	en	Edmonds, W. D., Zidek, J. (2010): A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 2010 (129): 1-111, DOI: 10.5281/zenodo.5352924
3A7F7B2C7F60885DFF34FA3222DEEB72.taxon	discussion	Comments. This is the only Metallophanaeus species whose female lacks an anteromedian transverse crest adjacent to anterior pronotal margin, a departure from “ normal ” female pronotal type also seen in C. cerberus (q. v.) References to C. horus by Pessôa (1934) and Pessôa and Lane (1941) are actually to C. pessoai (Pereira, 1949). Martínez (1959) reported C. horus to be strictly necrophagous. Felsche’s species (Phanaeus sericeus) was based on specimens from the southern part of the range, where individuals take on a more muted color tone than in the central Cerrado.	en	Edmonds, W. D., Zidek, J. (2010): A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 2010 (129): 1-111, DOI: 10.5281/zenodo.5352924
3A7F7B2C7F5C885FFF34FC322131EE72.taxon	description	Fig. 50, 59, 62, 69 - 74	en	Edmonds, W. D., Zidek, J. (2010): A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 2010 (129): 1-111, DOI: 10.5281/zenodo.5352924
3A7F7B2C7F5C885FFF34FC322131EE72.taxon	materials_examined	Type. P. saphirinus – holotype male, Zoologische Staatssammlung, Munich; P. chabrillaci – unknown to us; P. machadoi – holotype male, Museu de Zoologia, Universidade de São Paulo, São Paulo.	en	Edmonds, W. D., Zidek, J. (2010): A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 2010 (129): 1-111, DOI: 10.5281/zenodo.5352924
3A7F7B2C7F5C885FFF34FC322131EE72.taxon	diagnosis	Diagnosis. General – Paraocular areas (genae) mostly smooth, at most only weakly convex, not carinate lateral to eyes. Pronotum smooth posteromedially, otherwise weakly granulorugose; posterior angle flattened, slightly explanate, basal fossae and inner portions of sulcus paralleling posterior margin effaced. Anteromedian angle of metasternum (seen in profile, Fig. 59) rounded, not salient; angle capped with elongate thickening (bead) usually visible from below as broad V. Elytral interstriae flat; striae (x 5) simple, very fine, superficial, not at all impressed (x 5); bases of striae 1 - 4 distinctly fossate, fossae progressively larger laterally. Ventral surface of protibia lateral to longitudinal carina paralleling inner margin entirely punctatorugose (Fig. 50). Pygidium lacking basal groove. Length 12 - 22 mm. Color (Fig. 69 - 74) shining metallic blue, violet, red or rarely green. Male (Fig. 69 - 71) – Head with evenly tapering long horn, base not abruptly swollen. Pronotum with pair of closely set, apically convergent processes near posterior margin, separated by strong concavity; disk anterior to processes with deep, round concavity on each side, otherwise slightly convex, rarely with pair of closely set small acute tubercles about midway to anterior margin. Pronotal sculpturing limited to weak rugosity on lateral margins; otherwise surface nearly smooth. Apical processes of parameres rounded. Female (Fig. 73 - 74) – Pronotum with small, weakly trituberculate crest adjacent to anterior margin followed by weak concavity bounded posteriorly by pair of weak tumosities near middle of disk. Pronotum finely granulorugose on disk and sides, punctate posteromedially. Specimens examined – 381.	en	Edmonds, W. D., Zidek, J. (2010): A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 2010 (129): 1-111, DOI: 10.5281/zenodo.5352924
3A7F7B2C7F5C885FFF34FC322131EE72.taxon	distribution	Distribution. Paranaian subregion (Fig. 62). Collection Records. ARGENTINA: Chaco – Miraflores (Apr). Misiones – Posadas (Dec); Coñapirú (Dec); Garuhapé (Oct); Mocona (Dec); Dorado (Apr); Aristóbulo del Valle [Depto. Cainguas] (Dec); Dos de Mayo (Sep, Nov); Santa María; Parque Nacional Iguazú, 200 m (Jan-Feb, Dec); Cosapiel (Nov); Loreto (Dec). Tucumán – Tucumán (Feb). BRAZIL: Bahia – Entre Rios. Espiritu Santo – Timbuhy (Dec). Minas Gerais – Virginia, 1500 m (Nov); Poços do Caldas [Morro da Ferra] (Feb). Paraná – Londrina (Dec); Lapa (Mar); Curitiba (Feb, Apr, Oct – Dec); Bariqui (Apr). Rio de Janeiro – Itatiaia (Jan, Mar, Nov); Floresta da Tijuca (Oct, Dec); Ilha Grande (Apr); Petropolis (Nov-Dec); Nova Friburgo (Jan); 17 km E Nova Friburgo, 22 o 23 ’ 04 ” S 42 o 33 ’ 30 ” W, 750 m (Jan, Mar). Rio Grande do Sul – São Borja (Dec). Santa Catarina – Pinhal (Dec); Rio das Antas; Corupá (Jan, Oct-Nov); Nova Teutônia, 27 o 11 ’ S 52 o 23 ’ W (Sep-Nov, Jan); Rio Vermelho (Feb); São Francisco (Nov). São Paulo – 50 km SE Mogi das Cruzes [Serra do Mar Biological Station “ Boraceia ”], 800 - 900 m (Apr); São Bernardo (Jan); São Paulo (Mar); Cantareira (Dec); Campos do Jordão (Dec); Tremembé (Mar). PARAGUAY: Alto Paraguay – Bella Vista (Dec). Caazapá – Parque Nacional Caaguazú [San José Cristal] (Oct, Dec). Cordillera – Naranjo (Dec). Guairá – Paso Yobai (Sep); Villarrica (Nov); Colonia Independencia (Nov); Colonia Natalicio Talavera (Dec); Colonia Nueva Talavera; Yoveré [Cordillera Ybytyruzu] (Jan). Itapúa – Yatai [San Rafael Reserve], 26 o 38 ’ 13 ” S 55 o 39 ’ 50 ” W (Sep). Paraguarí – Paraguarí (Nov).	en	Edmonds, W. D., Zidek, J. (2010): A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 2010 (129): 1-111, DOI: 10.5281/zenodo.5352924
3A7F7B2C7F5C885FFF34FC322131EE72.taxon	discussion	Comments. Coprophanaeus saphirinus is one of the most common species of the genus and by far the most common in the subgenus. Its range is broad; and while it prefers forest habitat, it does venture into the more open Chaco formations of northeastern Argentina. The color of this species varies more than in any other member of the genus. Dark metallic blue predominates. A red form (Fig. 71), described as Phanaeus chabrillaci and sometimes regarded as a subspecies, occurs sporadically in the southern part of its extensive range; and a green form (Fig. 71) occurs most commonly in populations in the coastal forests of Rio de Janeiro. The bright colors of this species are associated with diurnal activity (Medina-Hernandez 2002). Concerning Phanaeus chabrillaci, we echo Harold’s (1869) sentiments, “ Das typische Stück [of C. chabrillaci] ist nichts als eine schön kupfrigrothe Varietät [of C. saphirinus] ”, and we agree with his synonymy. This morph of C. saphirinus occurs widely and in populations dominated by the bluish-violet form; it is not a distinct taxon in our opinion. Fernando Vaz-de-Mello (pers. comm.) has observed that the red form tends to occur more frequently in populations at somewhat higher elevations than does the blue form, and only in the southern portion of the species range. Thomson’s species was originally spelled “ chabrillacii ”, and in the literature it commonly appears so written. He named it for the collector of the type series, François Chabrillac. Harold (1869), followed by Blackwelder (1944), emended the name to “ chabrillaci ”. The elytral striae (more frequently the lateral ones) of this species are often micropunctate (x 10), but they are never carinulate. Pereira and d’Andretta’s (1955) description of C. machadoi was based on a single male with stronger strial and interstrial puncturing than typical C. saphirinus and with curious modifications of the pronotal disk. Fernando Vaz-de-Mello (pers. comm.) reports that coastal populations of this species (“ machadoi ”) in Rio de Janeiro, Espiritu Santo and some adjacent regions of Minas Gerais differ from populations in the main range of C. saphirinus. These populations, which include blue and green individuals, present a weak, midlongitudinal ridge on the anterior (declivitous) portion of the pronotum and stronger elytral striae that at times appear punctured (as in C. punctatus). Other large male variants exist; we have observed one with a pair of acute tubercles along the midline of the pronotal disk. Pereira (1949) reported observing dimorphic males of C. saphirinus. Besides those bearing a tapering head horn, he described small males with a bituberculate swelling of the kind also observed in the palaeno group of Phanaeus and in Oxysternon. In none of the smallest males observed by us have we seen a bituberculate swelling; the minimum condition has been only a small bump or an obsolete tubercle. Martínez (1959) cited C. saphirinus as strictly necrophagous, but it is regularly collected also from human feces.	en	Edmonds, W. D., Zidek, J. (2010): A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 2010 (129): 1-111, DOI: 10.5281/zenodo.5352924
3A7F7B2C7F5E8859FF34F93222D4ECF2.taxon	description	Fig. 54, 62, 75 - 79	en	Edmonds, W. D., Zidek, J. (2010): A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 2010 (129): 1-111, DOI: 10.5281/zenodo.5352924
3A7F7B2C7F5E8859FF34F93222D4ECF2.taxon	materials_examined	Type. Lectotype female (des. by Arnaud, 1982: 116), Muséum National d’Histoire Naturelle, Paris (examined).	en	Edmonds, W. D., Zidek, J. (2010): A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 2010 (129): 1-111, DOI: 10.5281/zenodo.5352924
3A7F7B2C7F5E8859FF34F93222D4ECF2.taxon	diagnosis	Diagnosis. General – Paraocular areas (genae) completely punctured, usually coarsely and densely so, at most only weakly convex, not carinate lateral to eyes. Pronotum smooth posteromedially, otherwise weakly granulorugose; posterior angle normal, basal fossae small, incorporated into inner ends of sulcus paralleling posterior margin. Anteromedian angle of metasternum (seen in profile, as in Fig. 59) rounded, not salient; angle capped with elongate thickening (bead). Elytral interstriae (Fig. 54) weakly convex; striae (x 5) weakly impressed, carinulate, appearing weakly punctate (x 5; see Comments below); bases of striae 1 - 4 impressed, not distinctly fossate. Ventral surface of protibia lateral to longitudinal carina paralleling inner margin entirely punctatorugose. Pygidium lacking basal groove. Length 13 - 17 mm. Elytra black, pronotum with green to yellowish-green areas (Fig. 75 - 78). Male (Fig. 77 - 79) – Head with evenly tapering long horn whose base is not abruptly swollen. Pronotum with pair of closely set, apically convergent processes near posterior margin, separated by strong concavity; disk anterior to processes lacking concavities. Pronotal disk punctate posteromedially, otherwise granulorugose. Apical processes of parameres rounded. Female (Fig. 75 - 76) – Pronotum with small, weakly trituberculate crest adjacent to anterior margin followed by weak concavity bounded posteriorly by pair of weak tumosities near middle of disk. Pronotum finely granulorugose on disk and sides, punctate posteromedially. Specimens examined – 6.	en	Edmonds, W. D., Zidek, J. (2010): A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 2010 (129): 1-111, DOI: 10.5281/zenodo.5352924
3A7F7B2C7F5E8859FF34F93222D4ECF2.taxon	distribution	Distribution. Known only from isolated localities in northern Brazilian Atlantic Forest province (Fig. 62). Collection Records. BRAZIL: Espiritu Santo – Fazenda Lagoa do Macuco [Mun. Linhares], 19 o 03 ’ 50 ” S 39 o 58 ’ 43 ” W, 10 m (Jan). Pernambuco – Igarassú [Reserva Ecologica Charles Darwin] (Mar, May-Jul, Oct).	en	Edmonds, W. D., Zidek, J. (2010): A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 2010 (129): 1-111, DOI: 10.5281/zenodo.5352924
3A7F7B2C7F5E8859FF34F93222D4ECF2.taxon	discussion	Comments. Olsoufieff (1924) had only three females at hand when he described this species. But he was rightly suspicious of placing it in Coprophanaeus s. str., noting “ L’absence des tarses antérieurs et la forme de l’epistome les font placer ici [Coprophanaeus s. str.], ou les ranger au groupe du Ph. saphirinus, dont elles ont le caractère general. Ce n’est qu’en trouvant les [two male symbols] que l’on pourra résoudre cette question. ” Arnaud (2002 c) confirmed its placement in this subgenus. Olsoufieff’s name refers to apparent puncturing of the elytral striae. But strial puncturing in this species (x 10) is in fact vague and virtually effaced. The illusion of obvious puncturing is produced by serial widening of the striae to produce a beaded (“ moniliform ”) appearance. Each wide spot of the stria encloses a flat area with a micropuncture in the middle. Coprophanaeus punctatus is a scarce species of which we have seen, in addition to the type series and a single female from Espiritu Santo, only one recently collected series from the Reserva Ecologica Charles Darwin in Pernambuco. The only strongly developed male in the series (see Fig. 77 - 78) resembles that of C. saphirinus with the difference that the tooth-like posteromedian pronotal projections point upward rather than being inclined anteriorly.	en	Edmonds, W. D., Zidek, J. (2010): A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 2010 (129): 1-111, DOI: 10.5281/zenodo.5352924
3A7F7B2C7F58885AFF34FAB22341E9F2.taxon	description	Fig. 51, 57, 62, 80 - 84	en	Edmonds, W. D., Zidek, J. (2010): A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 2010 (129): 1-111, DOI: 10.5281/zenodo.5352924
3A7F7B2C7F58885AFF34FAB22341E9F2.taxon	materials_examined	Type. P. pertyi – lectotype male, Zoologische Staatssammlung, Munich (des. Arnaud 2002 a: 6); P. alvarengai – holotype male, Museo de Zoologia, Universidade de São Paulo, São Paulo.	en	Edmonds, W. D., Zidek, J. (2010): A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 2010 (129): 1-111, DOI: 10.5281/zenodo.5352924
3A7F7B2C7F58885AFF34FAB22341E9F2.taxon	diagnosis	Diagnosis. General – Paraocular areas (genae) nearly flat, strongly and densely punctate, with short carina lateral to eyes. Posteromedian portion of pronotum appearing smoother and shinier than remain- der of disk, simply punctate, less strongly and densely so near basal fossae; posterior angle normal, basal fossae small, incorporated into inner ends of distinct sulcus paralleling posterior margin. Salient anteromedian angle of metasternum (seen in profile, Fig. 57) rounded. Striae (x 5) weakly impressed, simple; bases of striae 1 - 4 impressed, not distinctly fossate. Elytral interstriae weakly convex. Ventral surface of protibia lateral to longitudinal carina mostly smooth, usually with single or double row of shallow, confluent punctures paralleling longitudinal carina (Fig. 51, arrow). Pygidium lacking basal groove. Length 14 - 20 mm. Elytra dull black; pronotum, head, pygidium and venter with strong bluishgreen metallic reflections. Male (Fig. 80 - 82) – Head with evenly tapering long horn, base of horn abruptly swollen. Pronotum of large male with deep anteromedian concavity at summit marked by a lobate tumosity. Tips of parameres only slightly swollen laterally, lacking distinct flanges. Female (Fig. 83 - 84) – Cephalic carina simple, scarcely elevated, at most only weakly trituberculate. Pronotum with deep, oval anteromedian concavity, bearing strongly raised, trituberculate, quadrate process adjacent to anterior margin. Specimens examined – 48.	en	Edmonds, W. D., Zidek, J. (2010): A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 2010 (129): 1-111, DOI: 10.5281/zenodo.5352924
3A7F7B2C7F58885AFF34FAB22341E9F2.taxon	distribution	Distribution. Caatinga province (Fig. 62). Collection Records. BRAZIL: Bahia – Cruz das Almas (Apr); Maracas (Feb); Feira de Santana (Jun). Ceará – Maranguape, Tabatinga, 38 o 43 ’ 48 " W; 04 o 00 ’ 42 " S, 126 (Mar); Maranguape, Cagado, 38 o 45 ’ 05 " W 04 o 01 ’ 57 " S, 114 m (Apr). Maranguape, Sao Benedito, Sanctuario da Nossa Senhora de Penha, 38 o 40 ’ 48 " W 03 o 52 ’ 29 " S, 96 m (Mar); Juazeiro do Norte. Minas Gerais – Aguas Vermelhas (Dec). Paraíba – Mamanguape (Jul); Soledade (Mar); São José dos Cordeiros [Fazenda Almas], 7 o 28 ’ S 36 o 53 ’ W, 650 m (Feb). Pernambuco – Brejo Novo (Mar); Garanhuns (Mar). Piauí – São Raimundo Nonato [Parque Nacional Serra da Capivara] (Jan). Rio Grande do Norte – Canguaretama (Aug); Natal (Jun, Nov).	en	Edmonds, W. D., Zidek, J. (2010): A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 2010 (129): 1-111, DOI: 10.5281/zenodo.5352924
3A7F7B2C7F58885AFF34FAB22341E9F2.taxon	discussion	Comments. Phanaeus pertyi was proposed as a new name for the taxon Felsche (1901) erroneously regarded as Phanaeus thalassinus Perty (Olsoufieff 1924) (see Comments under C. thalassinus below). Subsequently, Martínez and Pereira (1967) synonymized Phanaeus alvarengai Martínez and Pereira (1955) and P. pertyi. In accordance with Article 53.3 of the International Code of Zoological Nomenclature (ICZN 1999), Phanaeus alvarengai Martínez and Pereira is the senior primary homonym of Phanaeus alvarengai Arnaud (1984), currently still placed in the genus Phanaeus (Edmonds 1994; Arnaud 2002 c). The fact that Martínez and Pereira placed their taxon in the subgenus Coprophanaeus of Phanaeus does not, in accordance with Article 57.4 of the Code (ICZN 1999), shield Arnaud’s name from unavailability, and, therefore, the name alvarengai sensu Arnaud must be replaced. We have referred the issue to the species’ author, Patrick Arnaud, for rectification.	en	Edmonds, W. D., Zidek, J. (2010): A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 2010 (129): 1-111, DOI: 10.5281/zenodo.5352924
3A7F7B2C7F5B885AFF34FDB225C8EE12.taxon	description	Fig. 53, 55, 58, 85 - 89	en	Edmonds, W. D., Zidek, J. (2010): A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 2010 (129): 1-111, DOI: 10.5281/zenodo.5352924
3A7F7B2C7F5B885AFF34FDB225C8EE12.taxon	materials_examined	Type. Holotype male, Museu de Zoologia, Universidade de São Paulo, São Paulo.	en	Edmonds, W. D., Zidek, J. (2010): A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 2010 (129): 1-111, DOI: 10.5281/zenodo.5352924
3A7F7B2C7F5B885AFF34FDB225C8EE12.taxon	diagnosis	Diagnosis. General – Paraocular areas (genae) densely rugose, strongly raised in front of eye (Fig. 61), carina adjacent to eye largely obliterated by strong rugosity. Posteromedian portion of pronotum (Fig. 53) strongly and densely punctate, lacking distinct granules or ridges (sculpturing stronger in females); posterior angle normal, basal fossae small, incorporated into inner ends of distinct sulcus paralleling posterior margin. Salient anterior angle of metasternum acutely angled, sometimes almost conical (Fig. 58). Ventral surface of protibia lateral to longitudinal carina mostly smooth, usually with single or double row of shallow, confluent punctures paralleling longitudinal carina (as in Fig. 51, arrow). Striae (x 5) weakly impressed, simple; bases of striae 1 - 4 impressed, not distinctly fossate. Elytral interstriae flat, transversely wrinkled (Fig. 55). Pygidium with broad, shallow basal groove. Length 18 - 21 mm. Color bright metallic green, often with golden reflections. Male (Fig. 87 - 89) – Head with evenly tapering horn, base of horn abruptly swollen. Pronotum of large male with deep anteromedian concavity at summit bearing straight transverse ridge. Tips of parameres with strongly developed rounded, flat flanges. Female (Fig. 85 - 86) – Cephalic carina with prominent conical median tubercle. Pronotum with deep, oval anteromedian concavity, bearing strongly raised, trituberculate, quadrate process adjacent to anterior margin. Specimens examined – 8.	en	Edmonds, W. D., Zidek, J. (2010): A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 2010 (129): 1-111, DOI: 10.5281/zenodo.5352924
3A7F7B2C7F5B885AFF34FDB225C8EE12.taxon	distribution	Distribution. Chaco province (Fig. 62). Collection Records. ARGENTINA: Salta – Disto. San Martín (Mar). BOLIVIA: Santa Cruz – Cordillera Prov., Puesto Salas (Nov); Palmar de las Islas, 19 o 25 ’ S 60 o 32 ’ W, 270 m (Feb). PARAGUAY: Boquerón – Guachalla [Alto Río Pilcomayo] (Oct); Filadelfia (Colonia Fernheim). Chaco – Fortín Esmeralda.	en	Edmonds, W. D., Zidek, J. (2010): A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 2010 (129): 1-111, DOI: 10.5281/zenodo.5352924
3A7F7B2C7F5B885AFF34FDB225C8EE12.taxon	discussion	Comments. The brilliant coloration of this rare denizen of the Chaco is stunning. According to Pereira (1949), references by Pessôa (1935) and Pessôa and Lane (1941) to C. horus are actually to this species.	en	Edmonds, W. D., Zidek, J. (2010): A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 2010 (129): 1-111, DOI: 10.5281/zenodo.5352924
3A7F7B2C7F5B8854FF34F9122136EB72.taxon	description	Fig. 52, 62, 90 - 94	en	Edmonds, W. D., Zidek, J. (2010): A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 2010 (129): 1-111, DOI: 10.5281/zenodo.5352924
3A7F7B2C7F5B8854FF34F9122136EB72.taxon	materials_examined	Type. Holotype male, Zoologische Staatssammlung, Munich. Diagnosis. General – Paraocular areas (genae) densely rugose, at most only weakly raised in front of eye; carina adjacent to eye largely obliterated by strong rugosity. Posteromedian portion of pronotum, at least near fossae, granulate or granulorugose (Fig. 52); posterior angle normal, basal fossae small; basal pronotal sulcus usually obsolete medially, not reaching basal fossae. Salient anteromedian angle of metasternum (seen in profile) quadrate, not noticeably salient. Striae (x 5) weakly impressed, simple; bases of striae 1 - 4 impressed, not distinctly fossate. Elytral interstriae weakly convex. Ventral surface of protibia lateral to longitudinal carina mostly smooth, usually with single or double row of shallow, confluent punctures paralleling longitudinal carina (as in Fig. 51, arrow). Pygidium with broad, shallow basal groove. Length 13 - 19 mm. Color dark blue or green. Male – Head with evenly tapering horn, whose base is abruptly swollen. Pronotum of large male with deep anteromedian concavity at summit bearing transverse ridge. Tips of parameres bear rounded flanges with slightly upturned margins. Female – Cephalic carina of female with prominent conical median tubercle. Pronotum with deep, oval anteromedian concavity, bearing strongly raised, trituberculate, quadrate process adjacent to anterior margin. Specimens examined – 7.	en	Edmonds, W. D., Zidek, J. (2010): A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 2010 (129): 1-111, DOI: 10.5281/zenodo.5352924
3A7F7B2C7F5B8854FF34F9122136EB72.taxon	distribution	Distribution. Largely unknown (Fig. 62). Collection Records. BRAZIL: Bahia – 160 km W Barrieras, 11 o 51 ’ S 46 o 10 ’ W, 830 m (Jan).	en	Edmonds, W. D., Zidek, J. (2010): A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 2010 (129): 1-111, DOI: 10.5281/zenodo.5352924
3A7F7B2C7F5B8854FF34F9122136EB72.taxon	discussion	Comments. This is one of the two or three rarest species of the genus. We have seen only seven specimens and accept the distinct possibility that the above diagnosis may prove to be too restrictive. Martínez and Pereira (1967) provided a detailed description of the species and commentary on the prevailing confusion attending the name “ thalassinus ”. According to their analysis, Olsoufieff (1924) concluded that Felsche (1901) misidentified Perty’s species in his redescription of “ thalassinus ”, which he inadvertently based on a then undescribed species. To rectify the error, Olsoufieff redescribed Felsche’s “ thalassinus ” as Phanaeus pertyi, and then proceeded to invert characters of the two in the couplet of his key separating C. thalassinus and C. pertyi!	en	Edmonds, W. D., Zidek, J. (2010): A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 2010 (129): 1-111, DOI: 10.5281/zenodo.5352924
3A7F7B2C7F558854FF34FC3324DEEEF5.taxon	diagnosis	Diagnosis. General – Lower portion of eye large, width greater than twice (usually about three times or more) that of oculogular space. Paraocular areas (genae) not carinate lateral to eyes. Occipital areas of parietals more-or-less flattened, lacking angulate prominence (Fig. 12). Circumnotal ridge entire or broken behind eyes (Fig. 7 - 8). Pronotal sculpturing variable, usually granulorugose anterolaterally, punctate to some degree posteromedially; basal fossae variable. Pronotum lacking posterolateral ridge. Striae fine, simple or carinulate. Posterior edge of hind wing strongly notched basally (Fig. 9). Posterior surface of each protibial tooth with single basal row of long setae (Fig. 6). Abdominal sterna usually smooth medially. Sexual dimorphism marked. Male – Head bearing lamellate process of some sort or trituberculate carina, never a simple curved horn. Pronotum variable, flattened or transversely concave anteromedially, with strong median prominence. Parameres variable. Female – Head with trituberculate carina, sometimes strongly raised. Protarsi absent. Pronotum variable, usually with transverse process near anterior margin followed by oval concavity.	en	Edmonds, W. D., Zidek, J. (2010): A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 2010 (129): 1-111, DOI: 10.5281/zenodo.5352924
3A7F7B2C7F558854FF34FC3324DEEEF5.taxon	distribution	Distribution. Neotropical region.	en	Edmonds, W. D., Zidek, J. (2010): A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 2010 (129): 1-111, DOI: 10.5281/zenodo.5352924
3A7F7B2C7F558854FF34FC3324DEEEF5.taxon	discussion	Comments. This subgenus embraces 29 species distributed in four species groups separated in the key below. Edmonds (1972) divided the subgenus into two species groups (jasius and dardanus); only the jasius group is maintained here as defined then. Our organization of this subgenus is different from that presented by Arnaud (2002 c), but many of the subgroups of closely related species (Arnaud’s complexes) are the same. Keys to species appear under the treatments of each species group.	en	Edmonds, W. D., Zidek, J. (2010): A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 2010 (129): 1-111, DOI: 10.5281/zenodo.5352924
3A7F7B2C7F538852FF34FF72228AECB2.taxon	description	Fig. 7, 12, 97, 101, 108, 112 - 118	en	Edmonds, W. D., Zidek, J. (2010): A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 2010 (129): 1-111, DOI: 10.5281/zenodo.5352924
3A7F7B2C7F538852FF34FF72228AECB2.taxon	materials_examined	Type. Holotype male, Museu de Zoologia, Universidade de São Paulo, São Paulo.	en	Edmonds, W. D., Zidek, J. (2010): A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 2010 (129): 1-111, DOI: 10.5281/zenodo.5352924
3A7F7B2C7F538852FF34FF72228AECB2.taxon	diagnosis	Diagnosis. General – Dorsum except for irregular black areas, head and pronotum bright metallic green or yellowish green, elytra dark green to almost black. Clypeal margin rounded, not noticeably angulate or explanate lateral to median teeth (Fig. 114, 117). Pronotum with elongate triangular depression densely packed with coarse granules extending from posterior margin to middle of disk (Fig. 97); area adjacent to depression and extending along posterior pronotal margin somewhat swollen, smooth, punctured but sometimes only weakly so. Basal pronotal fossae effaced. Elytral striae distinctly carinulate (Fig. 101), even in specimens with dark elytra. Length 18 - 24 mm. Male (Fig. 97, 116 - 118) – Base of head horn weakly concave. Anteromedian pronotal concavity shallow, bordered posteriorly by weak transverse carina bearing rounded central tubercle; ends of transverse carina curving downward, lacking distinct tubercle. Female (Fig. 114 - 115) – Cephalic carina straight, median tubercle larger than laterals. Pronotal ridge not strongly elevated, narrowly bidentate medially. Specimens examined – 99.	en	Edmonds, W. D., Zidek, J. (2010): A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 2010 (129): 1-111, DOI: 10.5281/zenodo.5352924
3A7F7B2C7F538852FF34FF72228AECB2.taxon	distribution	Distribution. Chacoan subregion – southern Cerrado and Chaco provinces (Fig. 113). Collection Records. BOLIVIA: Beni – Magdalena (Dec). Santa Cruz – Parque Nacional Noel Kempff Mercado [Los Fierros], 14 o 33.5 ’ S 60 o 55.9 ’ W. BRAZIL: Bahia – 150 km W Barrieras (Jan); Encruzilhada (Dec). Distrito Federal – Estação Forestal Cabeça do Veado, 1100 m (Oct); Brasilia (Nov-Dec). Mato Grosso – Uirapurú [Chapada dos Parecis] (Dec). Mato Grosso do Sul – Selvíria (Jan, Mar, Oct). Minas Gerais – Cordisburgo [Fazenda Pontinha] (Dec); Turvolândia (Feb). Paraná – Vila Velha. São Paulo – Mogi Guaçu [Fazenda Campininas] (Jan); Jabaquara [Capital] (Dec); Novo Horizonte (Nov); Batatais (Mar, Dec); São Paulo [Ipiranga] (Mar); São José dos Campos (Nov). Goiás – Rio Verde; Vianopolis (Dec). PARAGUAY: Amambay – Cerro Corá (Feb, Dec). Caaguazú – Caaguazú. Concepción – Cororó (Nov); Concepción [Santa Morana], 23 o 24 ’ S 57 o 26 ’ W (Oct); Campo Zanja Morotí, 22 o 53 ’ S 57 o 23 ’ W (Nov). Presidente Hayes – Lolita (Yaragui) (Jan).	en	Edmonds, W. D., Zidek, J. (2010): A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 2010 (129): 1-111, DOI: 10.5281/zenodo.5352924
3A7F7B2C7F53884FFF34FA32244AEED4.taxon	description	Fig. 100, 113, 119 - 123	en	Edmonds, W. D., Zidek, J. (2010): A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 2010 (129): 1-111, DOI: 10.5281/zenodo.5352924
3A7F7B2C7F53884FFF34FA32244AEED4.taxon	materials_examined	Type. P. acrisius – neotype male (des. by Arnaud, 2002 b: 3), The Natural History Museum, London (examined); P. camargoi – holotype male, Museu de Zoologia, Universidade de São Paulo, São Paulo.	en	Edmonds, W. D., Zidek, J. (2010): A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 2010 (129): 1-111, DOI: 10.5281/zenodo.5352924
3A7F7B2C7F53884FFF34FA32244AEED4.taxon	diagnosis	Diagnosis. General – Posterior portion of head and pronotum bright metallic green or yellowish green; elytral interstriae dull bronze, striae and margins of elytra bright green. Clypeal margin rounded (Fig. 119, 122), not noticeably angulate or explanate lateral to median teeth. Pronotum (as in Fig. 97) with elongate triangular depression densely packed with coarse granules, extending from posterior margin to middle of disk; area adjacent to depression and extending along posterior pronotal margin somewhat swollen, densely and rather coarsely punctatorugose to granulorugose. Basal pronotal fossae distinct. Elytral striae at most only very weakly carinulate (Fig. 100). Length 22 - 30 mm. Male (Fig. 121 - 123) – Base of head horn flat. Anteromedian pronotal concavity shallow, bordered posteriorly by distinctly tridentate transverse carina. Female (Fig. 119 - 120) – Cephalic carina straight, median tubercle larger than laterals. Pronotal ridge elevated, trapezoidal, tridentate. Specimens examined – 35.	en	Edmonds, W. D., Zidek, J. (2010): A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 2010 (129): 1-111, DOI: 10.5281/zenodo.5352924
3A7F7B2C7F53884FFF34FA32244AEED4.taxon	distribution	Distribution. Chacoan subregion – northern Cerrado and Caatinga provinces (Fig. 113). Collection Records. BOLIVIA: Beni – Magdalena, 180 m (Dec). Santa Cruz – Parque Nacional Noel Kempff Mercado [Los Fierros], 14 o 33 ’ 28 ” S 60 o 55 ’ 51 ” W (Jan-Feb); Parque Nacional Noel Kempff Mercado [Huanchaca], 13 o 54 ’ 12.5 ” S 60 o 48 ’ 57.1 ” W (Oct). BRASIL: Bahia – Cruz das Almas (Apr, Jun). Maranhão – Santa Quiteria de Maranhão, Fazenda Rodiador, 42 o 40 ’ 46 " W 03 o 24 ’ 50 " S, 47 m (Feb, May). Mato Grosso – 14 km S Posto do Gil, 14 o 37 ’ 55 ” S 56 o 16 ’ 23 ” W, 240 m (Jan); 25 km N Posto do Gil, 14 o 22 ’ 29 ” S 56 o 07 ’ 28 ” W, 500 m (Jan); Fazenda São João, 14 o 23 ’ 38 ” S 56 o 08 ’ 50 ” W, 520 m (Jan); Serra do Tombador, 14 o 36 ’ 47 ” S 56 o 15 ’ 12 ” W, 450 m (Jan); Alto Rio Arinos, Diamantino (Dec). Paraiba – Joãozeirinho (Jun). Pernambuco – Recife [campus Universidade Federal de Recife]; Tejucupapo (Apr).	en	Edmonds, W. D., Zidek, J. (2010): A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 2010 (129): 1-111, DOI: 10.5281/zenodo.5352924
3A7F7B2C7F53884FFF34FA32244AEED4.taxon	discussion	Comments. Coprophanaeus acrisius is the northern partner of the C. spitzi – C. acrisius species pair. MacLeay proposed C. acrisius as variety of C. jasius, and we cite its original name above, in accordance with ICZN Article 45.6.4 (ICZN 1999), as Phanaeus jasius acrisius MacLeay. Fernando Vaz-de-Mello (pers. comm.) has also observed specimens from the coast of Maranhão.	en	Edmonds, W. D., Zidek, J. (2010): A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 2010 (129): 1-111, DOI: 10.5281/zenodo.5352924
3A7F7B2C7F4E884EFF34F955230AECD5.taxon	description	Fig. 9, 95, 104, 113, 124 - 128	en	Edmonds, W. D., Zidek, J. (2010): A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 2010 (129): 1-111, DOI: 10.5281/zenodo.5352924
3A7F7B2C7F4E884EFF34F955230AECD5.taxon	diagnosis	Diagnosis. General – Dorsum black or brown, lacking metallic coloration. Clypeal margin noticeably angulate or explanate lateral to median teeth (Fig. 124, 127). Pronotum weakly granulose anterolaterally, disk smooth except for elongate, anteriorly narrowed depression filled with coalesced coarse granules often mixed with punctures extending from posterior margin toward middle of disk (Fig. 95). Basal pronotal fossae effaced or only weakly indicated. Length 19 - 25 mm. Male (Fig. 95, 104, 126 - 128) – Middle process of head horn short, length about one-fourth that of horn base. Anteromedian pronotal concavity shallow, bordered posteriorly by weak transverse carina bearing rounded central tubercle; ends of transverse carina thickened, lacking distinct tubercle. Female (Fig. 124 - 125) – Cephalic carina straight, trituberculate, tubercles equal-sized and closely set, distance between lateral tubercles about one-half of interocular distance. Pronotum lacking anteromedian concavity, with simple transverse carina at upper edge of smooth, declivitous area reaching anterior margin; declivity bounded on each side by shallow depression extending toward eye. Specimens examined – 66.	en	Edmonds, W. D., Zidek, J. (2010): A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 2010 (129): 1-111, DOI: 10.5281/zenodo.5352924
3A7F7B2C7F4E884EFF34F955230AECD5.taxon	distribution	Distribution. Brazilian Atlantic Coast Forest province of Paranaian subregion (Fig. 113). Collection Records. BRAZIL: Espiritu Santo – Parque Estadual Pedra Azul, 1500 m (Feb-Mar). Minas Gerais – Poco de Caldas (Jan, Dec); Poços do Coicos (Dec). Rio de Janeiro – 17 km E Nova Friburgo, 22 o 23 ’ 04 ” S 42 o 33 ’ 30 ” W, 750 m (Jan); Mury, 8 km E. Nova Friburgo, 22 o 21 ’ 49 ” S 42 o 33 ’ 07 ” W, 1150 m (Jan); Itatiaia, 700 m (Feb, Nov); Tijuca Forest (Jan). São Paulo – Catareira (Mar); Piauiti; 50 km SE Mogi das Cruzes, Estacão Biológica Boraceia (Serra do Mar), 800 - 900 m (Oct, Dec); Bocaina (Apr).	en	Edmonds, W. D., Zidek, J. (2010): A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 2010 (129): 1-111, DOI: 10.5281/zenodo.5352924
3A7F7B2C7F4E884EFF34F955230AECD5.taxon	discussion	Comments. Olsoufieff’s (1924) erroneously synonymized P. perseus Harold with C. cerberus (see comments under C. corythus). The coarse sculpturing of the posteromedian groove of the pronotum links this species with C. spitzi and C. acrisius (cf. Fig. 95 and 97). The form of the female pronotum is unique to the genus and more reminiscent of females of Phanaeus mexicanus group than other Coprophanaeus. Sculpturing of head and pronotum is weakest of species in the jasius group.	en	Edmonds, W. D., Zidek, J. (2010): A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 2010 (129): 1-111, DOI: 10.5281/zenodo.5352924
3A7F7B2C7F4F8848FF34FB52224AEC32.taxon	description	Fig. 96, 103, 107, 109, 113, 129 - 133	en	Edmonds, W. D., Zidek, J. (2010): A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 2010 (129): 1-111, DOI: 10.5281/zenodo.5352924
3A7F7B2C7F4F8848FF34FB52224AEC32.taxon	materials_examined	Type. S. jasius – neotype male (des. by Arnaud 2002 a: 7), Muséum National d’Histoire Naturelle, Paris (examined by photo); P. satyrus – unknown to us.	en	Edmonds, W. D., Zidek, J. (2010): A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 2010 (129): 1-111, DOI: 10.5281/zenodo.5352924
3A7F7B2C7F4F8848FF34FB52224AEC32.taxon	diagnosis	Diagnosis. General – Metallic coloration green to yellowish green (seldom with red reflections) confined to posterior areas of head and pronotal sides and margins. Clypeal margin (Fig. 129, 132) clearly explanate / angulate lateral to median teeth. Posteromedian portion of pronotum (Fig. 96) smooth, bearing sparse, effaced puncturing (weakest in large males). Sulcus paralleling carinate posterior margin of pronotum smooth (Fig. 109), lacking conspicuous puncturing or asperities (x 25); carina entire. Posterior pronotal fossae with sparse puncturing and / or microgranules; area surrounding fossae virtually smooth. Length 18 - 27 mm. Male (Fig. 96, 103, 131 - 133) – Tip of head horn never reaching level of prothoracic carina; anterior surface of base usually coarsely punctate; central process (viewed laterally) inclined slightly anteriorly from base, its length less than one-half that of base. Pronotal carina (Fig. 96) weakly, broadly angulate medially, trituberculate, medial tooth only slightly larger than laterals. Parameres long, length at least three-fourths that of phallobase. Female (Fig. 107, 129 - 130) – Cephalic carina bowed or angulate medially, median tooth larger than laterals. Vertex coarsely, densely punctured. Pronotal ridge trapezoidal, summit weakly trituberculate, width at summit slightly less than interocular width. Anteromedian concavity shallow, posterior (upper) margin simple, not bitumose. Specimens examined – 274.	en	Edmonds, W. D., Zidek, J. (2010): A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 2010 (129): 1-111, DOI: 10.5281/zenodo.5352924
3A7F7B2C7F4F8848FF34FB52224AEC32.taxon	distribution	Distribution. Amazonian subregion (Fig. 113). Collection Records. BRAZIL: Amazonas – 26 km N Manaus [Reserva Forestal Ducke] (Oct-Dec); 60 km N. Manaus, Fazenda Esteio and Fazenda Dimona (Jan, Jun-Jul); Municipio Itacoatiara, Fazenda Aruana [AM Hwy 10, km 215] (Nov); Manacapurú (Mar). Pará – Belêm [Agua Preta] (Jan-Feb]; upper Rocana River. Rondônia – 9 km NE Cacaulândia (Feb). FRENCH GUIANA: 7 km N Saul, 3 km NW Les Eaux Claires, Mt. La Fumée, 490 m, 3 o 39 ’ 46 ” N 53 o 13 ’ 19 ” W (Jun); Saül, Mt. Galbao, 3 o 37 ’ 18 ” N 53 o 16 ’ 42 ” W 740 m (Jun); Nouragues Field Station, 4 o 05 ’ N 52 o 40 ’ W. GUYANA: Cuyuni-Mazaruni – Bartica District (May). Potaro- Siparuni – Iwokrama Forest Reserve, 4 o 40 ’ 19 ” N 58 o 41 ’ 04 ” W, 100 - 200 m (May-June). SURINAME: Sipaliwini – Central Suriname Nature Reserve, environs of Voltzberg research station, 4 o 40.90 ’ N 56 o 11.13 ' ’ W, 100 m (Mar); Central Suriname Nature Reserve, environs of Lolopasi field station, 4 o 42.91 ’ N 56 o 12.83 ' ’ W, 80 m (Mar); Central Suriname Nature Reserve, environs of Conservation International field station approx. 50 m (Mar); Oelemarie, ~ 3 o 6 ’ 0 ' ’ N 54 o 32 ’ 0 ' ’ W (Aug); Palumeu, ~ 3 o 21 ’ 30.3 ' ’ N 55 o 26 ’ 38.2 ' ’ W (Jun). VENEZUELA: Bolívar – Lago Guri Islands, 7 o 21 ” S 62 o 52 ’ W (Jun); Las Trincheras [Río Caura] (Aug); Puerto Cabello [Río Caura] (Aug); Jabillal [Río Caura – Orinoco] (Oct). Delta Amacuro – 11 km W Piacoa.	en	Edmonds, W. D., Zidek, J. (2010): A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 2010 (129): 1-111, DOI: 10.5281/zenodo.5352924
3A7F7B2C7F4F8848FF34FB52224AEC32.taxon	discussion	Comments. There has been much confusion attending the use of the name “ jasius ”; it has been regularly and erroneously applied to practically all members of the jasius group. Arnaud’s (2002 a) designation of a neotype resolved much of the identity problem of this, the type species of the genus. Coprophanaeus jasius is an Amazonian species commonly found in the same communities as its congeners C. parvulus, C. dardanus and C. lancifer. It is a copronecrophagous forest dweller. Its closest known relative is C. abas, originally described by MacLeay as a variety of C. jasius and first recognized as a separate species by Arnaud (2002 c). Of the members of the jasius group, only C. jasius and C. abas have a conspicuously smooth posteromedian portion of the pronotum. Pereira and Martínez (1956; see also 1960) synonymized C. cyanescens and C. camargoi with C. jasius, but neither is now considered in the context of Olivier’s taxon. We consider C. cyanescens (q. v.) a valid species and C. camargoi a synonym of C. acrisius (q. v.). Arnaud (2002 c) has a wider definition of C. jasius than our own; he cites the species from Colombia, Ecuador, Peru and Bolivia. While we have seen no specimens from any of these countries, its occurrence in Amazonian habitats in any or all of them is not out of the question and we so indicate in the distribution depicted in Fig. 113. Arnaud (1982) pointed out that the type of Phanaeus costatus is in fact a “ hybrid ” combining the forebody of a Coprophanaeus with the hindbody of Diabroctis mimas (Linné). He identified the forebody as C. jasius, which he synonymized with C. costatus. C. P. D. T. Gillett (pers. comm.) has kindly informed us that the forebody is of C. cyanescens, requiring that the synonymy be reassigned accordingly (see Comments under C. cyanescens).	en	Edmonds, W. D., Zidek, J. (2010): A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 2010 (129): 1-111, DOI: 10.5281/zenodo.5352924
3A7F7B2C7F49884AFF34FB122112EA92.taxon	description	Fig. 102, 105, 110, 113, 135 - 139	en	Edmonds, W. D., Zidek, J. (2010): A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 2010 (129): 1-111, DOI: 10.5281/zenodo.5352924
3A7F7B2C7F49884AFF34FB122112EA92.taxon	materials_examined	Type. P. abas – neotype male (des. by Arnaud 2002 b: 3), The Natural History Museum, London (examined); P. rex – holotype male, National Museum, Prague (examined).	en	Edmonds, W. D., Zidek, J. (2010): A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 2010 (129): 1-111, DOI: 10.5281/zenodo.5352924
3A7F7B2C7F49884AFF34FB122112EA92.taxon	diagnosis	Diagnosis. General – Metallic color green to yellowish green and confined to posterior areas of head and pronotal sides and margins. Clypeal margin clearly explanate / angulate lateral to median teeth (Fig. 135, 138). Posteromedian portion of pronotum smooth, bearing sparse, effaced puncturing (weakest in large males). Sulcus paralleling carinate posterior margin of pronotum (Fig. 110) with obvious sculpturing (x 25); carina entire. Posterior pronotal fossae with sparse puncturing and / or microgranules. Length 20 - 29 mm. Male (Fig. 102, 137 - 139) – Head horn long, tip reaching level of summit of pronotum; anterior surface of horn base usually only weakly punctured; central process (viewed laterally) inclined slightly posteriorly from base, its length at least twice that of base. Pronotal process strongly produced medially as lobate tooth, itself sometimes weakly bilobed, flanked on each side by small, acute tooth (process reduced to strongly angulate transverse carina in smallest individuals). Parameres long, length at least three-fourths that of phallobase. Female (Fig. 105, 135 - 136) – Cephalic carina bowed or angulate medially, median tooth larger than laterals. Vertex moderately punctured. Pronotal ridge (Fig. 105) strongly elevated, quadrate, summit strongly trituberculate, width at summit slightly more than interocular width. Anteromedian pronotal concavity oval, deep, posterior (upper) margin clearly bitumose (except in smallest individuals). Specimens examined – 57.	en	Edmonds, W. D., Zidek, J. (2010): A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 2010 (129): 1-111, DOI: 10.5281/zenodo.5352924
3A7F7B2C7F49884AFF34FB122112EA92.taxon	distribution	Distribution. Mesoamerica – Venezuelan Llanos province and adjacent Amazonia (Fig. 113). Collection Records. COLOMBIA: Guaviaré – San José [Finca Esmeralda], 2 o 33 ’ N 72 o 38 ’ W, 240 m (Oct). Caquetá – Puerto Solario, PNN La Serranía de Chiribiquete, 0 o 12 ’ 47.8 " N 72 o 25 ’ 25.4 " W, 250 m (Feb-Mar). TRINIDAD – TOBAGO: 11 km SE Arima [Arena Forest Reserve], 80 m (Jun-Jul); 13 km S Arima, 2 km N Talparo, 50 m (Jun-Jul). VENEZUELA: Amazonas – Gavilán (May). Araugua – 20 km NW Maracay [Rancho Grande], 1000 m (Jun). Barinas – Barinitas [Mpio. Libertador, Quebrada Miraflores], 650 m (Aug). Bolívar – Lago Guri Islands, 7 o 21 ’ N 62 o 52 ’ W (Jun); Río Caura, Las Trincheras (Aug); 85 km S El Dorado (Aug); 33 km S El Dorado, 220 m (Aug); Sipao [110 km E Caicara del Orinoco] (Jun-Aug); 26 km N Guasipati (Jun-Jul); Canaima [220 km SSE Cd. Bolívar] (Aug); Caura River, Jabillal, Orinoco, 200 m (Aug).	en	Edmonds, W. D., Zidek, J. (2010): A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 2010 (129): 1-111, DOI: 10.5281/zenodo.5352924
3A7F7B2C7F49884AFF34FB122112EA92.taxon	discussion	Comments. This species, along with C. acrisius, was proposed by MacLeay (1819) as a variety of C. jasius (q. v.); in accordance with Article 45.6.4 (ICZN 1999), its original name is Phanaeus jasius abas MacLeay. In smaller individuals the large median pronotal tab is replaced by a central tubercle and the overall appearance becomes more similar to C. jasius. We do not understand well the distribution of this species, which may later become clearer as more material is collected. Our information suggests that it occurs in the Venezuelan Llanos province (? mainly along gallery forests of the Orinoco basin) and adjacent Amazonian provinces. The general area, especially in the Venezuelan state of Bolívar, is also home to C. jasius and eastern populations of C. gamezi; but we do not know if they are ecologically sympatric in the region.	en	Edmonds, W. D., Zidek, J. (2010): A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 2010 (129): 1-111, DOI: 10.5281/zenodo.5352924
3A7F7B2C7F4B8844FF34FC9222EEE9F2.taxon	description	Fig. 99, 106, 111, 113, 140 - 144	en	Edmonds, W. D., Zidek, J. (2010): A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 2010 (129): 1-111, DOI: 10.5281/zenodo.5352924
3A7F7B2C7F4B8844FF34FC9222EEE9F2.taxon	materials_examined	Type. Holotype male, private collection of Patrick Arnaud, Saintry sur Seine, France (examined by photo).	en	Edmonds, W. D., Zidek, J. (2010): A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 2010 (129): 1-111, DOI: 10.5281/zenodo.5352924
3A7F7B2C7F4B8844FF34FC9222EEE9F2.taxon	diagnosis	Diagnosis. General – Metallic coloration dark green, blue or yellowish green, confined to posterior areas of head and pronotal sides and margins; elytra black or tinged with highlights of same tone as pronotum. Clypeal margin clearly explanate / angulate lateral to median teeth (Fig. 140, 143). Posteromedian portion of pronotum granulate, granules often coalescing as transverse, squamose ridging. Sulcus paralleling carinate posterior margin of pronotum (Fig. 111) distinctly sculptured (x 25); carina usually broken up by dense puncturing, often crenulate. Posterior pronotal fossae and surrounding area densely, finely granulate. Length 18 - 28 mm. Male (Fig. 142 - 144) – Tip of head horn almost reaching level of prothoracic carina; anterior surface of base usually punctate; length of central process equal to that of base. Pronotal prominence with strongly developed rounded, tab-like projection, itself often weakly bitumose; median process flanked on each side by small, acute tubercle. Length of parameres about one-half that of phallobase. Female (Fig. 99, 106, 140 - 141) – Cephalic carina bowed anteriorly, trituberculate, median tooth raised, much larger than lateral tubercles (Fig. 99). Vertex coarsely, densely punctured. Pronotal ridge quadrate, summit scarcely trituberculate, if at all (Fig. 106). Anteromedian concavity shallow, posterior (upper) margin simple, not bitumose. Specimens examined – 201.	en	Edmonds, W. D., Zidek, J. (2010): A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 2010 (129): 1-111, DOI: 10.5281/zenodo.5352924
3A7F7B2C7F4B8844FF34FC9222EEE9F2.taxon	distribution	Distribution. Mesoamerica – Maracaibo, Venezuelan Coast and Venezuelan Llanos provinces and adjacent Amazonia (Fig. 113). Collection Records. BRAZIL: Roraima – Serra da Moça (Jul). COLOMBIA: Bolívar – Mompox, 9 o 14 ’ N 74 o 25 ’ W (Jun); Zambrano, Hacienda Monterrey, 9 o 37 ’ 48 " N 74 o 54 ’ 44 " W, 135 m (Feb). Boyacá – El Mortiño (Aug). Cundinamarca – Tocaima (Mar). Magdalena – Onaca, 770 m (Dec); Bonda (Jun). GUYANA: Rupununi – Parara Ranch, 3 o 37.3 ’ N 59 o 44.2 ’ W (Apr). Vichada – Municipio Cumaribo, PNN El Tuparro, 5 o 21 ’ 03 " N 67 o 52 ’ 15 " W, 135 m (Feb); Puerto Correrio (Nov). VENEZUELA: Amazonas – Puerto Ayacucho (May); 32 km S Puerto Ayacucho [Cataniapo River] (May). Aragua – Maracay, 450 m (Nov). Barinas – Maravillas (Sep); Saboneta [Mpio. Alberto Torrealba], 200 m (Jun). Bolívar – 48 km ESE Cd. Bolívar [Rancho Santa Rita], 7 o 55 ’ N 63 o 10 ’ W (Jul); Suapuré River (Jul); Sipao, 110 km E Caicara del Orinoco (Jun, Aug); 70 km W. Caicara del Orinoco; 70 km W Cd. Bolívar, Puente Cocuizas (Jun-Aug); 35 km SW Puerto Ordaz (Jul-Aug); Guri (Jul); Caura River, Jabillal, Orinoco, 200 m (Aug); El Dorado [Minas de Payapal] (May). Carabobo – San Esteban [near Puerto Cabello] (Dec); Saman Mocho, Flor Amarillo (Feb, Jul). Delta Amacuro – 11 km W Piacoa (Jul). Guárico – Tucupido (Jul) Mérida – Mpio. Padre Noguera, Sector Cuenca del Río Capara, 7 o 50 ’ 32 ” N 71 o 30 ’ 34 ” W, 280 m (Apr). Sucre – Cumanacoa (Dec). Zulia – Santa Ana [Mpio. Colón Hacienda], 8 o 54 ’ 30 ” N 71 o 52 ’ 17 ” W, 175 m (Aug); El Caracolí [Mpio. Colón], 70 m (May); Mpio. Colón, Sector Bancada de Limones, Finca El Recreo (Apr).	en	Edmonds, W. D., Zidek, J. (2010): A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 2010 (129): 1-111, DOI: 10.5281/zenodo.5352924
3A7F7B2C7F4B8844FF34FC9222EEE9F2.taxon	discussion	Comments. This species most closely resembles C. cyanescens (q. v.) from which it differs, among other characters, by the coarse sculpturing of the posterior margin of the pronotum (Fig. 111) and female pronotal ridge (Fig. 99). Gámez (2004) and Gámez and Acconcia (2009) studied this species in the Maracaibo district of Mérida, where it prefers open habitats and readily invades induced pasture lands, and presented detailed information on its activity.	en	Edmonds, W. D., Zidek, J. (2010): A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 2010 (129): 1-111, DOI: 10.5281/zenodo.5352924
3A7F7B2C7F458846FF34FDB2233EED52.taxon	description	Fig. 98, 113, 145 - 150	en	Edmonds, W. D., Zidek, J. (2010): A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 2010 (129): 1-111, DOI: 10.5281/zenodo.5352924
3A7F7B2C7F458846FF34FDB2233EED52.taxon	materials_examined	Type. P. cyanescens – lectotype male (des. by Arnaud 1982 a: 116), Muséum National d’Histoire Naturelle, Paris (examined by photo); P. costatus - holotype, hybrid specimen (examined; see Comments), The Natural History Museum, London; P. worontzowi – holotype male, Museu de Zoologia, Universidade de São Paulo, São Paulo.	en	Edmonds, W. D., Zidek, J. (2010): A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 2010 (129): 1-111, DOI: 10.5281/zenodo.5352924
3A7F7B2C7F458846FF34FDB2233EED52.taxon	diagnosis	Diagnosis. General – Metallic coloration green or dark blue, or intermediate tones, confined to posterior areas of head and pronotal sides and margins; elytra black or tinged with highlights of same tone as pronotum. Clypeal margin clearly explanate / angulate lateral to median teeth (Fig. 45, 148). Posteromedian portion of pronotum, except around basal fossae, punctured or punctatorugose, sometimes densely so. Sulcus paralleling carinate posterior margin of pronotum weakly sculptured (x 25); carina entire, sometimes weakly punctate. Posterior pronotal fossae and surrounding area densely, finely granulate. Length 19 - 27 mm. Male (Fig. 147 - 150) – Tip of head horn almost reaching level of prothoracic carina; anterior surface of base usually punctate; length of central process equal to that of base. Pronotal prominence widely Vshaped, trituberculate, medial tooth distinctly larger than laterals, width less than distance separating outer margins of eyes. Length of parameres about one-half that of phallobase. Female (Fig. 145 - 146) – Cephalic carina weakly bowed anteriorly, trituberculate, median tooth slightly larger than laterals. Vertex coarsely, densely punctured. Pronotal ridge trapezoidal, summit weakly trituberculate. Anteromedian concavity shallow, posterior (upper) margin simple, not bitumose. Specimens examined – 416.	en	Edmonds, W. D., Zidek, J. (2010): A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 2010 (129): 1-111, DOI: 10.5281/zenodo.5352924
3A7F7B2C7F458846FF34FDB2233EED52.taxon	distribution	Distribution. Chacoan subregion (Fig. 113). Collection Records. ARGENTINA: Catamarca – La Viña (Jan). Chaco – Miraflores (Apr, Sep); Chaco National Park, 100 km NW Resistencia (Dec); Puerto Bermejo. Formosa – Gran Guardia (Nov). Jujuy – El Quemado (Apr); 12 km S. Ledesma [Río Ledesma], 500 m (Dec); Calilegua National Park [Mirador; El Cortaderal; Aguas Negras], 500 - 900 m (Dec); 6 km W Yuto (Feb). Misiones – Dos de Mayo (Feb); El Dorado (Nov); Iguazú National Park, 200 m (Dec); Garuhapé (Mar). Salta – Tablillas, 700 m (Feb); General Martín Miguel de Güemes (Feb); Pocitos [Distrito San Martín] (Jan, Nov); El Rey Nacional Park, 890 m (Nov-Dec); Rosario de Lerma, 1325 m (Mar). Tucuman – Famaillá (Jan). BOLIVIA: Chuquisaca – Rosario [Lago Rocagua] (Nov). Beni – Rurrenbaque (Jun); Magdalena, 180 m (Dec). Santa Cruz – Santa Cruz Botanic Garden, 17 o 47 ’ 02 ” S 63 o 03 ’ 47 ” W, 400 m (Nov); Buena Vista (Feb-Mar); El Refugio Reserve, 14 o 46 ’ S 61 o 02 ’ W; Santiago (Feb); Parque Nacional Noel Kempff Mercado [El Refugio], 14 o 45 ’ 58 ” S 61 o 01 ’ 59 ” W (Jan); Reserva Municipal Tucavaca, 18 o 12 ’ S 59 o 28 ’ W, 150 m (Apr); RPPN Caparú, 14 o 50 ’ S 61 o 10 ’ W, 180 m (Dec); 1.6 km ESE Santiago de Chiquitos, 18 o 20.103 ’ S 59 o 35.007 ’ W, 622 m (Nov); 20 km NE Santiago de Chiquitos (Tucavaca River), 18 o 14.590 ’ S 59 o 27.907 ’ W, 215 m (Nov). Tarija – between Caiga and Creveaux, 20 o 50 ’ 19 ” S 63 o 24 ’ 58 ” W, 490 m (Jan); between Yaguacua and Caiza, 21 o 50 ’ 52 ” S 63 o 36 ’ 26 ” W (Jan); Entre Ríos (Quebrada de Abra) (Feb); Yacuiba (May-Jun); TCO Yumao [Kaaguasu], 19 o 05 ’ S 63 o 35 ’ W, 500 m (Feb). BRAZIL: Bahia – Feira de Santana (Jun); Encruzilhada (Dec); Cruz das Almas (Apr). Ceará – Uirapuru, Serra de Baturité, 4 o 20 ’ S 38 o 56 ’ W (Jun); Maranguape, São Benedito, Sanc. Nossa Senhora de Penha, 38 o 40 ’ 48 " W 03 o 52 ’ 29 " S, 96 m (Mar). Maranguape, 3 o 54 ’ S 38 o 42 ’ W (Jun); Maranguape, São Benedito, Sanctuario da Nossa Senhora de Penha, 38 o 40 ’ 53 " W 03 o 52 ’ 18 " S, 160 m (Mar- Apr); Maranguape, Tabatinga, 38 o 43 ’ 48 " W 04 o 00 ’ 42 " S, 126 m (Mar); Serra de Baturité, Uirapuru, 38 o 54 ’ 22 " W 04 o 17 ’ 27 " S, 620 m (Mar, May); Maranguape, Piraponga, 38 o 41 ’ 59 " W 03 o 53 ’ 26 " S, 200 m (Feb- Mar). Distrito Federal – 15 km N Brasilia, 1250 m (Mar). Espíritu Santo – Timbuhy (Dec); Linhares (Mar, Nov, Jan). Maranhão – Codó (Feb); Santa Quiteria de Maranhão, Fazenda Rodiador, 42 o 40 ’ 46 " W 03 o 24 ’ 50 " S, 47 m (Feb, May). Mato Grosso – 13.4 km S Posto do Gil, 14 o 37 ’ 55 ” S 56 o 16 ’ 23 ” W, 240 m (Jan); 20.2 km S Posto do Gil, 14 o 40 ’ 58 ” S 56 o 17 ’ 57 ” W, 240 m (Jan); Fazenda São João, 14 o 14 ’ 10 ” S 56 o 08 ’ 11 ” W, 400 m.; Vale de Solidão, 14 o 22 ’ S 56 o 07 ’ W (Mar). Mato Grosso do Sul – Paranaíba (Mar); Selvíria (Oct); Campo Grande (Oct, Dec). Minas Gerais – Sete Lagoas (Dec); Ipatinga (Nov); Sabará; Rio das Velhas. Pará – Belêm (Feb). Paraná – Londrina [Mata Godoy] (Jan-Mar, Dec). Paraíba – Mamanguape [Rebio Guaribas], 6 o 41 ’ S 35 o 10 ’ W (Apr); João Pessoa [Mata do Buraquinho] (Jun); Fazenda Pacatuba (Jul). Pernambuco – Recife. Rio de Janeiro – Rio Bonito (Apr); Pocos de Caldas (Dec). Rio Grande do Norte – Lajes. São Paulo – Piracicaba (Oct); Indiana (Feb); Ilha Solteira (Mar); Campinas (Jan); Batatais (Dec); Anhembí [Fazenda Barriero Rico] (Nov). PARAGUAY: Alto Paraguay – Puerto Bertoni; Primavera (Nov). Alto Paraná – Estación Dimas (Jan-Feb). Amambay – Cerro Cora (Feb, Dec); Pedro Juan Caballero (Nov). Caaguazú – Caaguazú. Canindeyú – Reserva Nacional Bosque Mbaracayú [Jejuí-Mí] (Jan, Nov). Central – Asunción; Capiatá (Oct). Concepción – Concepción [Santa Morana], 23 o 24 ’ S 57 o 26 ’ W (Nov); Horqueta (Dec); Cororó (Nov); Campo Zanja Morotí, 22 o 53 ’ S 57 o 23 ’ W (Mar, Nov-Dec). Cordillera – Altos (Mar); Naranjo (Oct, Dec). Guairá – Melgarejo (Apr, Sep); Colonia Sudetia (Jan); Yaparucu (Nov); Villarrica (Oct). Itapúa – Encarnación; San Rafael Reserve, 26 o 31 ’ 24 ” S 55 o 48 ’ 18 ” W, 90 m (Nov). Misiones – Santa Rosa (Oct). Paraguari – Sapucay (Jan-Apr, Oct-Dec). San Pedro – Yatevó.	en	Edmonds, W. D., Zidek, J. (2010): A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 2010 (129): 1-111, DOI: 10.5281/zenodo.5352924
3A7F7B2C7F458846FF34FDB2233EED52.taxon	discussion	Comments. This is a widespread species that, along with C. ensifer and C. bonariensis, is an indicator of the Chacoan subregion. Metallic coloration can be blue or green, and there seems to be no geographic pattern associated with color variability. Fernando Vaz-de-Mello (pers. comm.) reports that in the Atlantic coast forest of Brazil the blue form frequents forest habitats while the green form predominates in pastures and other open habitats. Conrad Gillett (pers. comm.) reports that in Ceará it occurs commonly in Atlantic forest fragments and less commonly in caatinga; specimens from both habitats there are bluish green. Rare individuals can present a crenulate posterior pronotal margin similar to that of C. gamezi. We have not seen C. rigoutorum, but our interpretations of Arnaud’s descriptions (2002 a, 2002 c) place it near, or perhaps in synonymy with C. cyanescens. Arnaud (2002 c: 36) listed P. worontzowi as a synonym of C. jasius; our interpretation of the latter results in the new synonymy of P. worontzowi and C. cyanescens. Edmonds’ (1967) reference to Phanaeus jasius undoubtedly applies to C. cyanescens. We here reassign the synonymy of Phanaeus costatus to C. cyanescens, which Arnaud (1982 a) synonymized with C. jasius (see Comments under C. jasius).	en	Edmonds, W. D., Zidek, J. (2010): A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 2010 (129): 1-111, DOI: 10.5281/zenodo.5352924
3A7F7B2C7F43883EFF34F9122152EAF2.taxon	description	Fig. 162 - 163, 171 - 172, 179 - 184	en	Edmonds, W. D., Zidek, J. (2010): A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 2010 (129): 1-111, DOI: 10.5281/zenodo.5352924
3A7F7B2C7F43883EFF34F9122152EAF2.taxon	materials_examined	Type. C. morenoi – holotype male, Muséum National d’Histoire Naturelle, Paris (examined by photo); C. kohlmanni – holotype male, Instituto Nacional de Biodiversidad, Santo Domingo de Heredia.	en	Edmonds, W. D., Zidek, J. (2010): A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 2010 (129): 1-111, DOI: 10.5281/zenodo.5352924
3A7F7B2C7F43883EFF34F9122152EAF2.taxon	diagnosis	Diagnosis. General – Completely black, only very rarely with metallic coloration along anterior pronotal margin. Clypeal teeth normal, length about equal to width at base. Width of upper portion of eye slightly greater than one-fourth of interocular distance. Frons smooth, at most with faint puncturing behind transverse carina. Posterior portion of paraocular area smooth. Posteromedian portion of pronotum smooth, devoid of distinct sculpturing; basal pronotal fossae effaced. Anterior angle of metasternum bearing elevated cap (seen in profile, as in Fig. 168). Elytral interstriae flat. Pygidium with wide basal groove (Fig. 162 - 163). Length 15 - 23 mm. Male (Fig. 181 - 184) – Pronotal prominence bimorphic, either cleat-shaped (Fig. 183, upper view) or quadrilobate with lobes linked by thick carina (Fig. 183, lower view); prominence flanked by broad concavities; declivitous surface of pronotum beneath carina smooth, devoid of sculpturing. Parameres with wide subapical tooth (Fig. 184). Female (Fig. 179 - 180) – Length of frons about one and one-half times that of clypeus. Transverse pronotal carina bidentate medially, followed by transverse depression summit of which weakly bitumid. Specimens examined – 327.	en	Edmonds, W. D., Zidek, J. (2010): A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 2010 (129): 1-111, DOI: 10.5281/zenodo.5352924
3A7F7B2C7F43883EFF34F9122152EAF2.taxon	distribution	Distribution. Chocó and Eastern Central American provinces (Fig. 172). Collection Records. COLOMBIA: Chocó – Quibdó (Sep); El Amargal Biological Station, 5.6 oN 77.4 oW, 150 - 500 m (Aug); 30 km S Quibdó [Lloró], 5 o 30 ’ N 76 o 33.5 ’ W, 90 m (Feb); Pacurita, 5 o 41 ’ N 76 o 40 ’ W, 50 m (Nov); 20 km NE Quibdó [Tutumendó], 60 m (Nov). Valle de Cauca – 70 km E Buenaventura [Anchicaya Dam], 350 m (Jul). COSTA RICA: Alajuela – 3.5 km E Bijagua [Heliconias Lodge], 10 o 42 ’ 45.2 ” N 85 o 02 ’ 28.5 ” W, 730 m (May-Jun). Heredia – Reserva Biológica La Selva, 10 o 26 ’ N 86 o 59 W, 75 m (Oct). Limón – Sector Cerro Cocori, Finca de E. Rojas, 150 m (Oct). San José – Estación Bijagual (north of Bijagualito), 600 m (Jun). ECUADOR: Esmeraldas – Borbón, 1 o 04 ’ 44 ” N 78 o 58 ’ 48 ” W, 25 m (Aug); San Miguel, 00 o 40 ’ 18 ” N 79 o 53 ’ 00 ” W (Dec); 11 km SE San Lorenzo [La Chiquita Forest Station] (Jun); Punta Venado (Nov-Dec); Yalere (Nov); Playa de Oro (Feb-Mar, Oct); La Concordia (Nov); Palma Real (Mar). Los Ríos – Río Palenque Research Station, 0 o 35 ’ S 79 o 22 ’ W, 200 m (Jan-Feb, Jun-Jul). Pichincha – Quevedo (Apr-May); 113 km NW Quito on Puerto Quito road, 800 m (Aug); i 1 km E Tinalandia, 600 m (Jul). NICARAGUA: Jinotega – 32 km NW Jinotega [Finca El Jaguar], 13 o 14 ’ 28 ” N 86 o 03 ’ 16 ” W, 1340 m (Dec). Matagalpa – Selva Negra, 13 o 00 ’ 01 ” N 85 o 54 ’ 32 ” W, 1350 m (May). Río San Juan – 8 km S El Castillo [Refugio Bartola], 10 o 56.6 ’ N 84 o 20.4 ’ W, 30 m (Mar, May). PANAMA: Colón – 16 km SE Colón [Santa Rita Ridge], 270 m (Jun). Darien – Estación Ambiental Cana, 7 o 45.32 ’ N 77 o 41.07 W, 600 - 750 m (Jun). Panamá – Capira [Cerro Campana], 8 o 44 ’ N 79 o 57 ’ W, 790 m (Jun, Dec); Chepo-Carti Road, 400 m (Jun-Aug); Cerro Azul, 650 m (May, Dec); 7.5 - 9.6 km N El Llano, 475 m (Jan); 12 km N El Llano on road to Carti, 250 m (May).	en	Edmonds, W. D., Zidek, J. (2010): A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 2010 (129): 1-111, DOI: 10.5281/zenodo.5352924
3A7F7B2C7F43883EFF34F9122152EAF2.taxon	discussion	Comments. The occurrence of bimorphic major males in this species is unique to the genus and, although in different descriptive terms and under the name “ ohausi ”, it was first described by Howden and Young (1981). The unusual form (Fig. 183, lower view) is so far known only from certain populations in the environs of El Llano (Panamá province), where it occurs with the typical cleat-shaped form (Fig. 183, upper view). Because the quadrilobate condition of the male pronotum occurs in individuals otherwise “ major ” in size and other respects, we consider it a developmental option to the cleat-shaped prominence in typical males rather than a step in the gradient from “ major ” to “ minor ” individuals. The middle of basal margin of pygidium is sometimes weakly angulate and tooth-like, at least partially interrupting basal groove (Fig. 162). This was a key character in the original description that has revealed itself to be highly variable. The toothed condition is more frequent in Ecuadorian, Colombian and eastern Panamanian populations, but it becomes very rare beyond the Darien province. For most of this study we regarded C. kohlmanni as a species distinct from C. morenoi; the primary criteria for doing so were its somewhat larger size and presumed isolated distribution. It turns out, however, that the distribution of C. morenoi is continuous from northwestern Ecuador through Panama along Caribbean coast into Nicaragua, and the species is subject to local variations along the way, the most striking of which is the unusual male type from western Panama. Costa Rican populations formerly assigned to C. kohlmanni are part of a continuum and tend to be somewhat larger than in other populations.	en	Edmonds, W. D., Zidek, J. (2010): A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 2010 (129): 1-111, DOI: 10.5281/zenodo.5352924
3A7F7B2C7F3F883EFF34FCB2237EEF52.taxon	description	Fig. 165, 172, 185 - 188	en	Edmonds, W. D., Zidek, J. (2010): A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 2010 (129): 1-111, DOI: 10.5281/zenodo.5352924
3A7F7B2C7F3F883EFF34FCB2237EEF52.taxon	materials_examined	Type. Holotype male, Canadian Museum of Nature, Ottawa (examined).	en	Edmonds, W. D., Zidek, J. (2010): A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 2010 (129): 1-111, DOI: 10.5281/zenodo.5352924
3A7F7B2C7F3F883EFF34FCB2237EEF52.taxon	diagnosis	Diagnosis. General – Black except for red to reddish-green metallic coloring on pronotum and pygidium. Clypeal teeth normal, length about equal to width at base (Fig. 161). Length of frons about equal to that of clypeus. Width of upper portion of eye one-fourth to one-fifth of interocular distance. Frons smooth, at most with faint puncturing behind transverse carina. Paraocular area completely sculptured, sculpturing weakening closer to eye. Posteromedian portion of pronotum only weakly punctured, becoming smooth near posterior margin; basal pronotal fossae distinct. Anterior angle of metasternum with rounded cap (seen in profile, as in Fig. 167). Elytral interstriae convex (Fig. 165). Pygidium with wide basal groove. Length 13 - 16 mm. Male (Fig. 185 - 187) – Tubercles of cephalic carina small, equal-sized. Pronotal prominence of large male cleat-shaped, width greater than that of emarginate, anteromedian portion of pronotal margin. Parameres lacking subapical tooth (Fig. 187). Female (Fig. 188) – Transverse pronotal carina bidentate medially, followed by transverse depression with summit very weakly bitumid. Specimens examined – 14.	en	Edmonds, W. D., Zidek, J. (2010): A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 2010 (129): 1-111, DOI: 10.5281/zenodo.5352924
3A7F7B2C7F3F883EFF34FCB2237EEF52.taxon	distribution	Distribution. Western Isthmus province of Costa Rica (Fig. 172). Collection Records. COSTA RICA: Puntarenas – Osa Peninsula [Río Piro La Joya], 8 o 23 ’ 56 ” N 83 o 19 ’ 51 ” W, 350 m (Aug-Sep); Rincón de Osa, 8 o 41.14 ’ N 83 o 31.11 ’ W, 150 m (Jun); Osa Peninsula [Rancho Quemado], 200 m (Jul, Dec); Estación Agujas, Sendero Zamia, 300 m (Apr, Oct); Estación Quebrada Bonita [Reserva Biológica Carara], 50 m (Sep); Estación Los Patos, 200 m (Sep).	en	Edmonds, W. D., Zidek, J. (2010): A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 2010 (129): 1-111, DOI: 10.5281/zenodo.5352924
3A7F7B2C7F3F883EFF34FCB2237EEF52.taxon	discussion	Comments. The distribution of C. solisi lies in the Pacific rainforest of the lower elevations of Puntarenas province of Costa Rica. This area includes the Osa Peninsula and extends northwest along the coast to the Tárcoles River, a landmark limit to Pacific coast species distributions discussed by Kohlmann and Wilkinson (2007). It is partially sympatric with C. pecki (q. v.).	en	Edmonds, W. D., Zidek, J. (2010): A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 2010 (129): 1-111, DOI: 10.5281/zenodo.5352924
3A7F7B2C7F3F883AFF34F8F22251EA52.taxon	description	Fig. 160, 172, 189 - 194	en	Edmonds, W. D., Zidek, J. (2010): A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 2010 (129): 1-111, DOI: 10.5281/zenodo.5352924
3A7F7B2C7F3F883AFF34F8F22251EA52.taxon	materials_examined	Type. Lectotype male (des. by Arnaud 2002 a: 9), The Natural History Museum, London (examined).	en	Edmonds, W. D., Zidek, J. (2010): A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 2010 (129): 1-111, DOI: 10.5281/zenodo.5352924
3A7F7B2C7F3F883AFF34F8F22251EA52.taxon	diagnosis	Diagnosis. General – Black except for green to yellowish-green metallic coloring on pronotum and pygidium. Clypeal teeth (subject to wear) elongate, length about one and half width at base (Fig. 160). Length of frons about equal to that of clypeus. Eyes large, width of upper portion about one-third to onefourth of interocular distance. Anterior one-half of frons punctate, smooth posteriorly. Paraocular area completely sculptured, sculpturing weakening closer to eye. Posteromedian portion of pronotum strongly punctured, punctures asperate, becoming weaker and sparser near posterior margin; basal pronotal fossae distinct. Anterior angle of metasternum with rounded cap (seen in profile, as in Fig. 167). Elytral interstriae at most only weakly convex (as in Fig. 166). Pygidium with wide basal groove (as in Fig. 163). Length 13 - 25 mm. Male (Fig. 191 - 194) – Tubercles of cephalic carina acute, equal-sized, somewhat elongate. Pronotal prominence of large male (Fig. 193) cleat-shaped, lateral angles reduced such that width is less than that of emarginate, anteromedian portion of pronotal margin. Parameres lacking subapical tooth (Fig. 194). Female (Fig. 189 - 190) – Transverse pronotal carina bidentate medially, followed by transverse depression with summit weakly bitumid. Specimens examined – 48.	en	Edmonds, W. D., Zidek, J. (2010): A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 2010 (129): 1-111, DOI: 10.5281/zenodo.5352924
3A7F7B2C7F3F883AFF34F8F22251EA52.taxon	distribution	Distribution. Extreme southern portion of the Pacific Coast province (Fig. 172). Collection Records. COSTA RICA: Guanacaste – Liberia (Aug); 34 km N Liberia (Hacienda Poco Sol) (May-Jul); Rincón de la Vieja National Park, 775 m (Jun); Estación Santa Rosa [Parque Nacional Santa Rosa], 10 o 50 ’ 23.0 ” N 85 o 37 ’ 07.0 ” W, 290 m (Jun-Jul);. San José – Santo Domingo (Aug). EL SAL- VADOR: San Salvador – San Salvador (Jul); Santa Tecla (Nueva San Salvador), 900 m (Sep); Lago Ilopango (Aug). HONDURAS: Cortés – Parque Nacional Cerro Azul-Meambar, 14 o 52.4 ’ N 87 o 54.7 ’ W, 800 m (May). Francisco Morazán – El Zamorano [Escuela Agrícola Panamericana], 800 m (Jul); Tatumbla, 1400 m (Oct). Olancho – 14 km N La Unión [Parque Nacional La Muralla], 1500 m (Aug).	en	Edmonds, W. D., Zidek, J. (2010): A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 2010 (129): 1-111, DOI: 10.5281/zenodo.5352924
3A7F7B2C7F3F883AFF34F8F22251EA52.taxon	discussion	Comments. Pereira (1953) regarded this species as a synonym of C. pluto. This very close relative of C. pluto is a denizen endemic to dry forests of the Pacific coast of Costa Rica, Honduras and El Salvador, where it occurs with Phanaeus eximius Bates and P. demon var. excelsus Bates. We have seen specimens labeled Tuxtla Gutierrez, Chiapas (530 m); while its occurrence there is a reasonable possibility, the record needs confirmation.	en	Edmonds, W. D., Zidek, J. (2010): A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 2010 (129): 1-111, DOI: 10.5281/zenodo.5352924
3A7F7B2C7F3B8835FF34FDF22320EE52.taxon	description	Fig. 151 - 153, 161, 166 - 167, 169 - 170, 172, 195 - 199	en	Edmonds, W. D., Zidek, J. (2010): A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 2010 (129): 1-111, DOI: 10.5281/zenodo.5352924
3A7F7B2C7F3B8835FF34FDF22320EE52.taxon	materials_examined	Type. P. pluto – lectotype male (des. by Arnaud 1982 a: 115), Muséum National d’Histoire Naturelle, Paris (examined by photo; see Comments); P. morio – holotype male, Museum of Comparative Zoology, Harvard University, Cambridge (examined by photo). C. nogueirai – holotype male, private collection of Patrick Arnaud, Saintry sur Seine, France.	en	Edmonds, W. D., Zidek, J. (2010): A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 2010 (129): 1-111, DOI: 10.5281/zenodo.5352924
3A7F7B2C7F3B8835FF34FDF22320EE52.taxon	diagnosis	Diagnosis. General – Black except for green to yellowish-green metallic coloring on pronotum and pygidium. Clypeal teeth normal, length about equal to width at base (Fig. 161). Length of frons about equal to that of clypeus. Eyes normal, width of upper portion one-fourth to one-fifth of interocular distance. Frons smooth, at most with faint puncturing behind transverse carina. Paraocular area completely sculptured, sculpturing weakening closer to eye. Posteromedian portion of pronotum distinctly punctured, punctures reaching posterior margin; basal pronotal fossae distinct. Anterior angle of metasternum with rounded cap (seen in profile, Fig. 167). Elytral interstriae at most only weakly convex (Fig. 166). Pygidium with wide basal groove. Length 14 - 25 mm. Male (Fig. 151 - 153, 169 - 170, 197 - 199) – Lateral tubercles of cephalic carina small, acute; middle tubercle variable: a) same size and shape as laterals (Fig. 151), b) elongate, about twice as high as laterals (Fig. 152), or c) lengthened, broadened and truncate or weakly emarginate apically (Fig. 153). Pronotal prominence of large male cleat-shaped (Fig. 197 - 198), width greater than that of emarginate anteromedian portion of pronotal margin. Parameres lacking subapical tooth (Fig. 169, 199). Female – Transverse pronotal carina bidentate medially, followed by transverse depression with summit weakly bitumid. Specimens examined – 377.	en	Edmonds, W. D., Zidek, J. (2010): A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 2010 (129): 1-111, DOI: 10.5281/zenodo.5352924
3A7F7B2C7F3B8835FF34FDF22320EE52.taxon	distribution	Distribution. All Mesoamerican provinces of Mexico (Fig. 172). Collection Records. GUATEMALA: Suchitepéquez – Variedades, 300 m. MEXICO: Aguascalientes – Auguascalientes (Sep). Chiapas – 19 km N Mapastepec, 550 m (Aug); Tapachula; Rosario Izapa (May). Colima – Manzanillo (Aug); Plantanarilla, 1200 m (Jul); 1 km S El Limón, 870 m (Jul). Guerrero – 7 km N Taxco (Jun); 45 km NE Ixtapa (Aug). Hidalgo – Laguna Azteca (Sep). Jalisco – El Tuito, 600 m (Jul); Guadalajara (Jul); Estación Biológica Chamela; Sierra de Manantlán, La Calera, 760 m (Oct); Sierra de Quila, Tecolotlán, 1580 m (Jul); Toanya, 950 m (Jul); Sierra de Manantlán, El Tigre, 700 m (Jul); Ejutla (la Estancia), 960 m (Oct); Autlán, 875 m (Jun); Puerto Los Mazos, 1350 m (Jul); Mezquitlán, 1000 m (Jul); El Parotal, 456 m (Sep); Zenzontla, 1000 m (Jun); ~ 5 km WNW Talpa de Allende, 20 o 23.893 ’ N 104 o 51.980 ’ W, 1263 m (Jul). Mexico – Valle de Bravo (Sep); 10 km SW Valle de Bravo (Jun). Michoacan – road between La Huacana and Ario de Rosales (Jul). Morelos – Cuernavaca; Tlaltizapan (Jul); Acamilpa (Jul). Nayarit – Tepic (Jul); Mesa de Nayar (Jul); Jesus María. Nuevo Leon – El Diente (Oct). Oaxaca – near Monte Albán, 17 o 03 ’ 34.0 ” N 96 o 45 ’ 53.4 ” W, 1690 m (Jul). Queretaro – 2 km S Valle Verde, 21.501 oN 99.177 oW, 1300 m (Aug). Sinaloa – 13 km E Villa Unión, 100 m (Jul). San Luis Potosí – 6.5 km E Cd. Maiz (Aug); 80 km NNW Cd. Valles (El Salto) (Jul). Sonora – 29 km NW Yécora, 28 o 23 ’ 34.7 ” N 109 o 05 ’ 30.8 ” W, 1350 m (Jul); 32 km NW Yécora, 28 o 24 ’ 03.6 ” N 109 o 06 ’ 24.4 ” W, 1190 m (Jul); Alamos, 400 m (Sep); 75 km W Yécora, 28 o 29 ’ 09 ” N 109 o 21 ’ 32 ” W, 935 m (Aug); 140 km W Yécora, 28 o 36 ’ 12 ” N 109 o 48 ’ 06 ” W, 550 m (Aug);. Tamaulipas – 75 km E Cd. Victoria [Villa de Casas] (Jul); Gómez Farías [Estación Biológica Los Cedros], 350 m (Jul); 4.5 km W Gómez Farías [Altas Cimas], 900 m (Jul); Villa Aldama (Aug); Villa Allende, 90 m (Aug). Veracruz – Palma Sola (Jun); 11 km W Palma Sola, 19 o 46 ’ N 96 o 25 ’ W, 120 m (Aug- Sep); Río Playa; Córdoba (Jul); Actopan (Aug); Cerro Sonpaso, 690 m (Aug); Apazapan, 19 o 19 ’ 18 ” N 96 o 42 ’ 39 ” W, 280 m (Aug); 14 km WSW Papantla [Plan de Hidalgo] (May); Alto Lucero, 690 m (Sep). UNITED STATES: Arizona – Santa Cruz Co., Nogales (Sep). Texas – Cameron Co. - Sabal Palm Grove Sanctuary (Oct); 12.5 km N junction hwy. 106 and FM 2925; Laguna Atascosa National Wildlife Refuge (Nov); Harlingen (Nov). Hidalgo Co. – Edinburg.	en	Edmonds, W. D., Zidek, J. (2010): A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 2010 (129): 1-111, DOI: 10.5281/zenodo.5352924
3A7F7B2C7F3B8835FF34FDF22320EE52.taxon	discussion	Comments. The male cephalic carina of C. pluto is trimorphic. In the “ major ” state (Fig. 153) the middle tubercle is greatly thickened, much higher than the laterals, and apically truncate or weakly excised. This form is characteristic of very large males along the Gulf coast of Veracruz, Tamaulipas and into southern Texas, where it occurs along with the “ minor ” form. In the “ minor ” state (Fig. 151) the three tubercles are equal-sized. The minor state (described as the subspecies “ nogueirai ” by Arnaud 2002 b; see below) is found in Gulf Coast populations and is the only known condition in populations along the Pacific coast. In the third state (Fig. 152), occurring infrequently in isolated populations in southern Mexico, the middle tubercle is acute and about twice the length of the laterals. This is the only species of the genus known to reach the United States, where it has been collected in extreme southern Texas and Arizona. Riley and Wolfe (2003) reported C. pluto from Cameron, Starr and Willacy counties, which, along with Hidalgo County, comprise the southern tip of Texas. Certain individuals from southern Texas exhibit the dark blue metallic color seen also in C. gilli, which they can resemble at first glance. Robinson (1948) reported C. pluto from Arizona. We regard the lectotype in the Paris museum as the primary type of this species on the assumption that Harold’s many phanaeine types remained there in the nineteenth century in the wake of the dispute between the Paris and London museums. However, there is a specimen in the London museum labeled as holotype; in this case none of the labels is attributable to Harold himself. Both specimens are, to our eyes, assignable to C. pluto. Arnaud (2002 b) based C. pluto nogueirai (original name “ noguerai ” emended to “ nogueirai ” by Zidek 2005) on the structure of the “ minor ” male cephalic carina. For much of this study we were inclined to recognize Arnaud’s taxon as valid. As far as we know, large males of “ nogueirai ” from southern Mexico and the Pacific coast always have an acute middle tubercle; that is, they are always “ minor. ” We have found, however, that Gulf coast populations often also include, along with “ major ” males, individuals identical to C. nogueirai. Our conclusion is that the only defining character of C. p. nogueirai – a male cephalic carina with three acute, usually equal-sized tubercles (Fig. 151) – is an expression of the normal intrapopulational variation of the male of C. pluto that has become fixed in southern and western populations of the species. The case of “ nogueirai ” poses interesting questions about male variation and the genetic status of species “ en statu nascendi ” that would be well worth an intensive population genetics study.	en	Edmonds, W. D., Zidek, J. (2010): A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 2010 (129): 1-111, DOI: 10.5281/zenodo.5352924
3A7F7B2C7F348837FF34F9F22338EC72.taxon	description	Fig. 172, 200 - 204	en	Edmonds, W. D., Zidek, J. (2010): A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 2010 (129): 1-111, DOI: 10.5281/zenodo.5352924
3A7F7B2C7F348837FF34F9F22338EC72.taxon	materials_examined	Type. P. conocephalus – holotype female, Muséum National d’Histoire Naturelle, Paris (examined by photo); P. roubali – holotype, National Museum, Prague (examined); C. edmondsi – holotype male, Canadian Museum of Nature, Ottawa (examined).	en	Edmonds, W. D., Zidek, J. (2010): A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 2010 (129): 1-111, DOI: 10.5281/zenodo.5352924
3A7F7B2C7F348837FF34F9F22338EC72.taxon	diagnosis	Diagnosis. General – Black except for dark blue metallic coloring on pronotum and pygidium. Clypeal teeth normal, length about equal to width at base. Eyes normal, width of upper portion one-fourth to onefifth of interocular distance. Frons smooth, at most with faint puncturing behind transverse carina (stronger puncturing in female). Paraocular area completely sculptured, sculpturing weakening closer to eye. Posterior portion of pronotum virtually smooth, if puncturing visible (x 10) then minute and sparse; basal pronotal fossae distinct. Anterior angle of metasternum with rounded cap (seen in profile, as in Fig. 167). Elytral interstriae at most only weakly convex, seldom shinier midlongitudinally. Pygidium usually lacking distinct basal grove, rarely with narrow basal groove. Length 15 - 20 mm. Male (Fig. 202 - 204) – Length of frons about one and half times that of clypeus. Teeth of cephalic carina closely set, distance between lateral teeth usually about one-half that of interocular distance. Pronotum lacking distinct fovea above lateral fossa; sometimes with prominent tubercle in front of lateral fossa (Fig. 202). Pronotal prominence of large male narrow, bitumid, tumescences connected by ridge. Parameres at most with indistinct subapical tooth. Female (Fig. 200 - 201) – Teeth of cephalic carina closely set, distance between lateral teeth about onehalf or less of interocular distance. Lengths of frons and clypeus about equal. Transverse pronotal carina bidentate medially, followed by very weak depression. Specimens examined – 17.	en	Edmonds, W. D., Zidek, J. (2010): A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 2010 (129): 1-111, DOI: 10.5281/zenodo.5352924
3A7F7B2C7F348837FF34F9F22338EC72.taxon	distribution	Distribution. Cauca province (Fig. 172). Collection Records. COLOMBIA: Nariño – Río Ñambi, 1 o 17 ’ N 78 o 15 ’ W, 1350 m (May). Valle de Cauca – Morro Frio, 1800 m (Mar); Q. Cristalina, AM Calima (Jul) ECUADOR: Cañar – Javin, 850 - 1400 m (Feb). Carchi, Parroquia El Goaltal, Hacienda San Francisco, 1200 m (Sep). Guayas – Balzar (Feb). Pichincha – Chiriboga, 1500 m (Feb); Chiriboga Road, 9.8 km NE Hwy. 30, 00 o 13 ’ 14 ” S 78 o 53 ’ 20 ” W, 1300 m (Mar).	en	Edmonds, W. D., Zidek, J. (2010): A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 2010 (129): 1-111, DOI: 10.5281/zenodo.5352924
3A7F7B2C7F348837FF34F9F22338EC72.taxon	discussion	Comments. There is no good reason in our opinion to maintain C. edmondsi as a distinct taxon in view of its close morphological and geographical relationship with C. conocephalus. The distribution of this species appears to parallel that of C. morenoi in Colombia and Ecuador at higher elevations (> 1000 m) along the western slopes of the cordillera (Note: The Ecuadorian portion of the range is not depicted in Fig. 172.) Along with C. chiriquensis and C. gilli, it forms a cluster differing from the pluto subgroup by its bitumid, rather than cleat-shaped, pronotal prominence.	en	Edmonds, W. D., Zidek, J. (2010): A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 2010 (129): 1-111, DOI: 10.5281/zenodo.5352924
3A7F7B2C7F368831FF34FB3220A8ECD2.taxon	description	Fig. 154 - 157, 164, 172, 205 - 209	en	Edmonds, W. D., Zidek, J. (2010): A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 2010 (129): 1-111, DOI: 10.5281/zenodo.5352924
3A7F7B2C7F368831FF34FB3220A8ECD2.taxon	materials_examined	Type. P. chiriquensis – female lectotype (des. by Arnaud, 1982: 116), Muséum National d’Histoire Naturelle, Paris (examined by photo); C. uhleri – holotype male, National Museum, Prague (examined).	en	Edmonds, W. D., Zidek, J. (2010): A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 2010 (129): 1-111, DOI: 10.5281/zenodo.5352924
3A7F7B2C7F368831FF34FB3220A8ECD2.taxon	diagnosis	Diagnosis. General – Black except for dark blue metallic coloring on pronotum and pygidium. Clypeal teeth normal, length about equal to width at base. Eyes normal, width of upper portion one-fourth to onefifth of interocular distance. Frons with weak, sparse puncturing behind transverse carina. Paraocular area completely sculptured, sculpturing weakening closer to eye. Posterior portion of pronotum usually smooth, at most with very weak, sparse puncturing; basal pronotal fossae distinct. Anterior angle of metasternum with rounded cap (seen in profile, as in Fig. 167). Elytral interstriae distinctly convex, usually shinier midlongitudinally (Fig. 205, 208). Pygidium usually lacking basal groove, rarely narrow groove present. Length 15 - 20 mm. Male (Fig. 154 - 157, 205 - 207) – Length of frons about one and half times that of clypeus. Teeth of cephalic carina closely set, distance between lateral teeth usually about one-half that of interocular distance. Pronotum with large, oval fovea mesial to lateral fossa (as in Fig. 210, arrow). Pronotal prominence of large male bitumid (Fig. 154 - 157); tumescences widely separated, usually connected by ridge, each with flattened, triangular area on dorsal surface. Parameres with acute subapical tooth (Fig. 207, 212). Female (Fig. 208 - 209) – Distance between lateral teeth of cephalic carina clearly greater than one-half of interocular distance. Lengths of frons and clypeus about equal. Transverse pronotal carina bidentate medially, followed by transverse depression. Specimens examined – 130.	en	Edmonds, W. D., Zidek, J. (2010): A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 2010 (129): 1-111, DOI: 10.5281/zenodo.5352924
3A7F7B2C7F368831FF34FB3220A8ECD2.taxon	distribution	Distribution. Central cordillera of Costa Rica between Eastern Central America and Western Isthmus provinces (Fig. 172). Collection Records. COSTA RICA: Alajuela – 3.5 km E Bijagua [Heliconas Lodge], 10 o 42 ’ 45.2 ” N 85 o 02 ’ 28.5 ” W (May-Jun); Parque Nacional Guanacaste [Estación San Ramón], 620 m (Jun); Reserva Biológica Alberto Brenes, Río San Lorencito, 850 m (Jun). Cartago – Río Grande de Orosi, 1150 - 1600 m (Dec); 19.3 km NE San José, 1010 m (May). Guanacaste – 9 km S Santa Cecilia [Estación Pitilla], 700 m (Jun-Jul); Tierras Morenas, 700 m (Mar); Rincón de la Vieja National Park, 775 m (Jun). Heredia – 16 km SSE La Virgen, 10 o 16 ’ N 84 o 05 ’ W, 1100 m (Mar). Puntarenas – San Vito [OTS Las Cruces Field Station], 1200 m (May-Jul); 6 km S San Vito, 1000 m (Apr-May); Monteverde, 1400 m (May-Aug); 1 km NE Sitio Portones [Zona Protectora Las Tablas], 1530 m (May); 1 km SW Cerro Biolley, Buenos Aires [Estación Altamira], 9 o 01 ’ 58 ” N 83 o 00 ’ 39 ” W, 1300 - 1450 m (Feb, Mar, Jul, Nov); 1.4 km NE La Tigra [Avenida El Pizote], 1300 m (Jul); Estación La Casona, Monte Verde, 10 o 18.5 ’ N 84 o 47.8 ’ W, 1520 m (Jun). PANAMA: Chiriquí – 4 km N Santa Clara [Hartmann’s Finca], 8 o 51 ’ N 82 o 46 ’ W, 1500 m (Jun-Aug); La Fortuna Dam, 1000 m (Jul); 15 km NE Gualaca [Cerro Hornito], 1200 m (Jul-Sep); 33.1 km S Chiriquí Grande, 800 m (Dec); Boquete (Jun). Veraguas – 8 km W Santa Fe [Cerro Tute], 8 o 30 ’ 26 ” N 81 o 06 ’ 49 ” W, 900 m.	en	Edmonds, W. D., Zidek, J. (2010): A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 2010 (129): 1-111, DOI: 10.5281/zenodo.5352924
3A7F7B2C7F368831FF34FB3220A8ECD2.taxon	discussion	Comments. Coprophanaeus uhleri is based on smaller specimens that fit well within the observed variation of C. chiriquensis. Pereira and Martínez (1956; see also 1960) erroneously synonymized this species with C. corythus. The posterior portion of the pronotum (as well as the frons) is smooth and usually devoid of any distinct punctures; rare individuals (usually large females) can show distinct, sparse puncturing (x 10) (see C. gilli). Coprophanaeus chiriquensis has been collected along with C. pecki in the Chiriquí region of Panama; but otherwise its distribution follows the cloud forests of the central Costa Rican cordillera that separates the Western Isthmus province from Eastern Central America province.	en	Edmonds, W. D., Zidek, J. (2010): A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 2010 (129): 1-111, DOI: 10.5281/zenodo.5352924
3A7F7B2C7F308830FF34FB522223ECB2.taxon	description	Fig. 158 - 159, 172, 210 - 212	en	Edmonds, W. D., Zidek, J. (2010): A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 2010 (129): 1-111, DOI: 10.5281/zenodo.5352924
3A7F7B2C7F308830FF34FB522223ECB2.taxon	materials_examined	Type. C. gilli – holotype male, Canadian Museum of Nature, Ottawa (examined); C. henryi – holotype male, National Museum, Prague (examined).	en	Edmonds, W. D., Zidek, J. (2010): A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 2010 (129): 1-111, DOI: 10.5281/zenodo.5352924
3A7F7B2C7F308830FF34FB522223ECB2.taxon	diagnosis	Diagnosis. General – Black except for dark blue metallic coloring on pronotum and pygidium (rarely also on elytra). Clypeal teeth normal, length about equal to width at base. Eyes normal, width of upper portion one-fourth to one-fifth of interocular distance. Frons weakly to moderately strongly punctured behind transverse carina. Paraocular area completely sculptured, sculpturing weakening closer to eye. Posteromedian portion of pronotum distinctly punctured, punctures progressively weaker and sparser approaching posterior margin; basal pronotal fossae distinct. Anterior angle of metasternum with rounded cap (seen in profile, as in Fig. 167). Elytral interstriae distinctly convex, usually shinier midlongitudinally (as in Fig. 205, 208). Pygidium usually lacking basal groove, rarely narrow groove present. Length 15 - 26 mm. Male (Fig. 158 - 159, 210 - 212) – Length of frons about one and half times that of clypeus. Teeth of cephalic carina closely set, distance between lateral teeth usually about one-half that of interocular distance. Pronotum with oval fovea above lateral fossa (Fig. 210, arrow). Pronotal prominence of large male thickened transverse carina with pair of closely set median tumescences, sometimes scarcely broken medially. Parameres with acute subapical tooth (Fig. 212, upper view). Female – Distance between lateral teeth of cephalic carina clearly greater than one-half of interocular distance. Lengths of frons and clypeus about equal. Transverse pronotal carina bidentate medially, followed by transverse depression. Specimens examined – 102.	en	Edmonds, W. D., Zidek, J. (2010): A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 2010 (129): 1-111, DOI: 10.5281/zenodo.5352924
3A7F7B2C7F308830FF34FB522223ECB2.taxon	distribution	Distribution. Gulf of Mexico and Eastern Central America provinces (Fig. 172). Collection Records. BELIZE: Toledo – Chiquibul National Park (Doyle’s Delight), 16 29 ’ 23 ” N 89 02 ’ 45 ” W, 950 m (Aug). HONDURAS: Cortes – Parque Nacional Cusuco, Santo Tomás, 700 - 800 m (Aug). Olancho – Parque Nacional La Muralla, 1800 m (Jun). GUATEMALA: Alta Verapaz – Finca Seacté Cobán (Apr). Izabal – 8 km N Las Escobas [Cerro San Gil], 800 m (Jun). Zacapa – 3 km S La Unión, 1400 m (Jun). MEXICO: Chiapas – 10.5 km W El Bosque, 1480 m (Aug); Santa Rosa (Aug); Bonampak, 16 o 44 ’ N 91 o 05 ’ W, 300 m (Sep). Hidalgo – Tlanchinol (Jun, Aug); Otongo, 1120 m (Jul). Oaxaca – Huautla de Jiménez (Nov); Los Metates (Sep). Puebla – Calapan 19 o 55 ’ 45.92 ” S 97 o 23 ’ 07.65 ” W, 1250 m (Jul). Veracruz – 33 km NE Catemaco, 160 m (Jul); Cuautlapan (Aug-Sep); Pipiapan [Parque de la Fauna y Flora Silvestre Tropical], 600 m (Nov); Dos Amates [Catemaco] (Jul, Sep); Sontecomapan [Estación Biológica Tropical “ Los Tuxtlas ”], 400 m (Jun, Oct); Ruiz Cortinas [Mpio. San Andrés Tuxtla], 1020 m (Apr). Cerro El Gallo [Mpio. Catemaco], 810 m (Apr); Guadalupe Victoria [Mpio. Tatahuicapan], 750 m (Jun); San Martín Pajapan [Mpio. Mecayapan], 720 m (Feb); Cuautlapan [Cerro Chicahuaxtla] (Aug); Ruiz Cortines (Apr).	en	Edmonds, W. D., Zidek, J. (2010): A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 2010 (129): 1-111, DOI: 10.5281/zenodo.5352924
3A7F7B2C7F308830FF34FB522223ECB2.taxon	discussion	Comments. This species is very closely related to C. chiriquensis, and, given the long history of intense collecting in Mexico and Central America, it is surprising that it went undiscovered until the late twentieth century. It differs from C. chiriquensis in geographic distribution, the shape of prominence of male pronotum, a somewhat larger average size, and usually in distinct (x 10) puncturing of the frons and posterior portion of the pronotum. Coprophanaeus henryi is based on smaller specimens that fit well within the variation of C. gilli. Among the best data we have on the local ecology of this species are those provided by Matthias Rös (pers. comm.). Careful sampling in the sierras of northern Puebla has revealed a rather broad habitat, from well conserved mesophilous forest to secondary growth, acahual (scrub), coffee plots and other highly disturbed areas between 1000 and 1300 m. Sharing this ecological spectrum there with C. gilli is another neotropical immigrant, C. corythus.	en	Edmonds, W. D., Zidek, J. (2010): A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 2010 (129): 1-111, DOI: 10.5281/zenodo.5352924
3A7F7B2C7F2F8828FF34FE5225C4ED72.taxon	description	Fig. 6, 213 - 216, 237, 239 - 243	en	Edmonds, W. D., Zidek, J. (2010): A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 2010 (129): 1-111, DOI: 10.5281/zenodo.5352924
3A7F7B2C7F2F8828FF34FE5225C4ED72.taxon	materials_examined	Type. P. dardanus – unknown to us (see Comments); P. bitias – unknown to us; P. jasion – holotype male, Staatliches Museum für Tierkunde, Dresden; P. arrowi – lectotype male (des. by Arnaud 1982: 116), Muséum National d’Histoire Naturelle, Paris (examined by photo).	en	Edmonds, W. D., Zidek, J. (2010): A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 2010 (129): 1-111, DOI: 10.5281/zenodo.5352924
3A7F7B2C7F2F8828FF34FE5225C4ED72.taxon	diagnosis	Diagnosis. General – Dorsum dark, sombre, metallic reflections restricted to pronotum and head (Fig. 239 - 242). Clypeal margin distinctly angulate adjacent to median teeth. Length of clypeus about equal to that of frons. Pronotal disk lacking midlongitudinal sulcus, posterior portion smooth to very weakly punctured. Cap of anterior metasternal angle usually simple, sometimes slightly forked; anterior surface of metasternum smooth above anterior margin of disk. Elytral striae not distinctly carinulate. Length 14 - 29 mm. Male – Head horn (Fig. 213 - 215) erect, plate-like, gently curved posteriorly; width approaching interocular distance, sides parallel to slightly divergent; apex bidentate, teeth elongate, widely separated by deep, rounded emargination. Pronotal prominence (Fig. 241 - 242) cleat-like, strongly bilobate, lobes directed anterolaterally and flanked by deep, oval concavities, lateral margin of each concavity with weak tumosity above lateral fossa; width of prominence slightly greater than distance between outer margins of eyes. Apical processes of parameres strong, hook-like. Female – Cephalic carina (Fig. 216) widely bidentate, apical margin between teeth widely raised, weakly angulate. Transverse crest of pronotum rounded, distinctly bidentate medially, followed by weak depression with feebly bitumose summit. Specimens examined – 535.	en	Edmonds, W. D., Zidek, J. (2010): A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 2010 (129): 1-111, DOI: 10.5281/zenodo.5352924
3A7F7B2C7F2F8828FF34FE5225C4ED72.taxon	distribution	Distribution. Northern Amazonian subregion and coastal Brazil (Fig. 237). Collection Records. BRAZIL: Amapá – Serra do Navio [Cava Urucum-Amapari], 00 o 53 ’ 06 ” N 51 o 52 ’ 53 ” W (Jan, Sep). Amazonas – 60 km N Manaus [Fazenda Esteio] Bahia – Porto Seguro (Jan). Ceará – Maranguape, Piraponga, 38 o 41 ’ 59 " W 03 o 53 ’ 26 " S, 200 m; Baturite, Uirapuru, 38 o 54 ’ 22 " W 04 o 17 ’ 27 " S, 620 m (Mar). Goiás – Jaraguá. Pará – Redenção, 7 o 46 ’ S 51 o 58 ’ W (Oct); Belêm (Jan-Mar); Alidos (Mar); Monte Dourado, 0 o 42 ’ S 52 o 38 ’ W, 100 m (Mar-Apr); 45 km E Canindé (Dec). Paraíba – João Pessoa [Mata do Buraquinho] (Aug); Aréia [Mata do Pau Ferro], 6 o 58 ’ S 35 o 42 ’ W (Apr). Paraná – Londrina (Apr). Pernambuco – Cabo [Reserva Ecologica Gurjaú] (May, Oct). Rio de Janeiro – Itatiaia, 700 m (Oct, Dec); Rio de Janeiro [Parque Lage] (Feb); Botafogo (Mar, Nov); Floresta da Tijuca (Dec); Petropolis (Sep, Dec). Roraima – Caracaraí (Jul). Santa Catarina – Corupá (Oct-Nov); Joinville (Apr, Jun). CO- LOMBIA: Guaviaré – San José [Finca La Esmeralda], 2 o 33 ’ N 72 o 38 ’ W, 240 m (Oct). FRENCH GUIANA: 8.4 km SSE Roura, 4 o 40 ’ 41 ” N 52 o 13 ’ 25 ” W, 200 m (May-Jun); 7 km N Saül [Les Eaux Claires], 3 o 39 ’ 46 ” N 53 o 13 ’ 19 ” W (May-Jun); Wanaboo [Marowijne River], 4 o 43 ’ 35 ” N 54 o 26 ’ 36 ” W, 40 m (Jun); 9.7 km NW – 4.3 km SE Patawa [Kaw Mountain], 4 o 32 ’ 40.1 ” N 52 o 09 ’ 08.1 ” W (Dec); Mana, 5 o 39.2 ’ N 53 o 49.9 ’ W (Jul); 100 km S Cayenne [Nouragues Research Station], 4 o 05 ’ N 52 o 40 ’ W. GUYANA: Mazaruni-Potaro – Takutu Mts., 6 o 15 ’ N 59 o 05 ’ W (Dec). Potaro-Siparuni – Iwokrama Forest Reserve, 4 o 40 ’ 19 ” N 58 o 41 ’ 04 ” W, 100 - 200 m (Jun). SURINAME: Brokopondo – Stoneiland, ~ approx. 4 o 59 ’ N 55 o 09 ’ W, 30 m (Apr-May); Rosebel gold mining area, ~ 5 o 07 ' N; 55 o 17 W (May). Para – ~ 20 km SSE Joden Savanne, 5 o 16 ’ 17 ” N 54 o 55 ’ 15 ” W, 40 m (Jun-Jul). Paramaribo – Pallisadenweg (Jul). Sipaliwini – Central Suriname Nature Reserve, environs of Voltzberg research station, 4 o 40.90 ” N 56 o 11.13 ' ’ W, 100 m (Mar); Central Suriname Nature Reserve, environs of Lolopasi field station, 4 o 42.91 ’ N 56 o 12.83 ’ W, 80 m (Mar); Central Suriname Nature Reserve, environs of Conservation International field station, 50 m (Mar); Central Suriname Nature Reserve, Raleighvallen, Fungu Island (= Foengoe Island), ~ 4 o 33 ’ 30 " N 56 o 12 ’ 21 " W (Jul); Bakhuis mountains, ~ 4 o 27 ’ 13 " N 56 o 57 ’ 39 " W (Nov); Palumeu, ~ 3 o 20 ’ 46.3 ' ’ N 55 o 26 ’ 30.8 ' ’ W (Jun). Wanica – Lelydorp (Feb); Zanderij, Hannover (Jun). TRINIDAD-TOBAGO: Port-of-Spain [Capuro Valley]; 16 km N Arima, 650 m (Jun); 8 km N Arima, 260 m; Maracas Valley, 600 m; 11 km SE Arima [Arena Forest Reserve], 80 m (Jun). VENEZUELA: Amazonas – Puerto Ayacucho (May); El Infierno (Jun); Atabapo (Jun); Camani (Jun-Jul). Aragua – Maracay [Rancho Grande] (Jun). Bolívar – 50 km SE El Manteco [Río El Supamo] (Dec); Paují, 1100 m; 10 km E San Francisco Yuruani, 1300 m (Jul); 20 km S Luepa, 1500 m (Jul); 10 km E Kavanayen, 1500 m (Jun); 22 km SE El Dorado (Jul); 10 km S El Dorado, 200 m (Jul-Aug); 100 km S El Dorado, 350 m (Jul-Aug); 6 km S San Isidro (Jun-Jul); Parupa, 1500 m (Jun-Jul). Miranda – 33 km N Altagracia de la Montaña [Guatopo National Park, La Crucita], 400 m (Jun). Monagas – 27 km SW Caripe, 300 m (Jun); Caripe, 700 m (Jul).	en	Edmonds, W. D., Zidek, J. (2010): A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 2010 (129): 1-111, DOI: 10.5281/zenodo.5352924
3A7F7B2C7F2F8828FF34FE5225C4ED72.taxon	discussion	Comments. We assume that the type, or a candidate for the lectotype of this species, will be found in the MacLeay Collection at the University of Sydney. We were unable to secure specimens from Sydney for this study. However, there is little room for doubt about the identity of C. dardanus even from the brief description of MacLeay (1819: 126 - 127): “ P. nigro-viridis, capitis cornubus duobus brevibus basi connatis, thorace antice carinâ bidentatâ instructo … Mas. Thorax fossulâ utrinque sub carinae dentibus unidentatâ … ” (Free translation: Black with green, head horn with two teeth broadly connected at the base; thorax with bidentate process … Male: Lateral concavity beneath median process with single tooth … ”). His diagnosis highlights the bidentate structure of the male head horn (Fig. 213 - 214), which is unique to the genus and seen otherwise in phanaeines only in Diabroctis. The length of the teeth varies, but usually amounts to at least the height of the base. In large individuals the upper margin of the base, between the lateral teeth, is slightly raised medially. In very small males (Fig. 215) the horn is reduced to a weakly bidentate swelling reminiscent of the small males of Oxysternon and certain Phanaeus. The cephalic carina of the female (Fig. 216), even in small individuals, is also unique and highly diagnostic. It bears a small denticle at each end, separated by a broad, rounded or slightly angulate elevation. We regard C. dardanus as a fundamentally Amazonian species, but it has managed to spread southward via the Atlantic coast of Brazil into portions of the Paranaian subregion as far south as Santa Catarina as well as into the central highlands (via gallery forests?). Remarkably, as far as we can see, the vast occupation of its combined range has not resulted in any detectable geographic variation. We have not seen Arnaud’s (2002 a) species, C. terrali, which was based on a single male from Mato Grosso and considered a close ally of C. dardanus. The single reported specimen (holotype) is part of a private collection that was not available for study. The shape of the cephalic carina of the holotype is strikingly different from that expected for C. dardanus, and the possibility that the shape is a teratologic form or a distinct morph in otherwise normal populations needs to be ruled out in any assessment of its validity.	en	Edmonds, W. D., Zidek, J. (2010): A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 2010 (129): 1-111, DOI: 10.5281/zenodo.5352924
3A7F7B2C7F29882AFF34FA32248CEAD2.taxon	description	Fig. 8, 227, 234, 236 - 237, 244 - 248	en	Edmonds, W. D., Zidek, J. (2010): A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 2010 (129): 1-111, DOI: 10.5281/zenodo.5352924
3A7F7B2C7F29882AFF34FA32248CEAD2.taxon	materials_examined	Type. Holotype male, private collection of Patrick Arnaud, Saintry sur Seine, France (examined by photo).	en	Edmonds, W. D., Zidek, J. (2010): A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 2010 (129): 1-111, DOI: 10.5281/zenodo.5352924
3A7F7B2C7F29882AFF34FA32248CEAD2.taxon	diagnosis	Diagnosis. General – Dorsum (Fig. 244 - 247) with widespread metallic green reflections. Clypeal margin at most only weakly angulate adjacent to median teeth. Length of clypeus greater than that of frons, especially in male. Pronotal disk with midlongitudinal sulcus expanding posteriorly into triangular depression embracing basal fossae (Fig. 247); sulcus and triangular depression metallic green. Cap of anterior metasternal angle usually simple, sometimes bifurcate; anterior surface of metasternum smooth above anterior margin of disk. Elytral striae distinctly carinulate (Fig. 234). Length 18 - 29 mm. Male – Head horn (Fig. 227) erect, plate-like, set very near eyes; width approaching interocular distance, sides more-or-less parallel; apex tridentate, emarginations separating teeth deep, rounded. Pronotal prominence (Fig. 247) strongly bilobate, lobes directed anteriorly and flanked by shallow concavities; width of prominence clearly less than distance between outer margins of eyes. Apical processes of parameres tooth-like. Female – Transverse crest of pronotum rounded, at most only very weakly bidentate; crest followed by weak concavity with feebly bitumose summit. Specimens examined – 62.	en	Edmonds, W. D., Zidek, J. (2010): A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 2010 (129): 1-111, DOI: 10.5281/zenodo.5352924
3A7F7B2C7F29882AFF34FA32248CEAD2.taxon	distribution	Distribution. Chaco and Cerrado provinces (Fig. 237). Collection Records. BOLIVIA: Beni – Estación Paraparu (May); Reyes (Dec); Magdalena (Dec); Estación Biológica del Beni [Bosque El Trapiche], 14 o 52 ’ 10 ” S 66 o 19 ’ 58 ” W, 195 m; Río Ichiguita, 15 o 8 ’ S 56 o 18 ’ W, 155 m. Chuquisaca – Rosario [Lago Rocagua] (Nov). La Paz – Ixiamas (Dec). Santa Cruz – RPPN Caparú, 14 o 47 ’ S 61 o 10 ’ W, 180 m; General Saavedra (May); Parque Nacional Noel Kempff Mercado [Los Fierros], 14 o 36 ’ 53 ” S 60 o 51 ’ 01 ” W and 14 o 34 ’ 45.6 ” S 60 o 54 ’ 30.3 ” W, 200 m (Jan-Feb); Concepción (Jan). BRAZIL: Goias – Leopoldo de Bulhões (Dec). Mato Grosso – Rio Taguarassu (Nov). Minas Gerais – Serra do Sipo, Vao da Lagoa (Dec); Serra Caraça, 138 m (Nov); Juiz de Fora (Nov). Paraná – Curitiba (Feb); Vilha Velha (Feb). São Paulo – São Paulo (Dec); Tremembé (Mar); São Miguel (Jan); Santo Amaro.	en	Edmonds, W. D., Zidek, J. (2010): A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 2010 (129): 1-111, DOI: 10.5281/zenodo.5352924
3A7F7B2C7F29882AFF34FA32248CEAD2.taxon	discussion	Comments. Because of the close similarity in size and coloring, a casual assessment may confuse this species with C. spitzi and C. acrisius. In the latter pair the pronotal sculpturing is much coarser and denser, especially posteromedially; moreover, the anterior part of the circumnotal ridge is not broken behind the eyes as it is in C. magnoi. Arnaud (2002 b) described this taxon as a subspecies of C. milon. Because the distributions of the two appear to be mutually exclusive and they are consistently distinct morphologically, we have chosen to regard C. magnoi and C. milon as separate species.	en	Edmonds, W. D., Zidek, J. (2010): A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 2010 (129): 1-111, DOI: 10.5281/zenodo.5352924
3A7F7B2C7F2B8827FF34FD5224F4EA92.taxon	description	Fig. 225 - 226, 233, 237, 249 - 253	en	Edmonds, W. D., Zidek, J. (2010): A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 2010 (129): 1-111, DOI: 10.5281/zenodo.5352924
3A7F7B2C7F2B8827FF34FD5224F4EA92.taxon	materials_examined	Type. P. milon – lectotype male (des. by Arnaud 1982 a: 114), Muséum National d’Histoire Naturelle, Paris (examined by photo); P. obscurus – holotype male, Zoologische Staatssammlung, Munich.	en	Edmonds, W. D., Zidek, J. (2010): A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 2010 (129): 1-111, DOI: 10.5281/zenodo.5352924
3A7F7B2C7F2B8827FF34FD5224F4EA92.taxon	diagnosis	Diagnosis. General – Dorsum (Fig. 249 - 252) dark, sombre, metallic reflections dark blue or bluish green. Clypeal margin at most only weakly angulate adjacent to median teeth. Length of clypeus greater than that of frons, especially in male. Pronotal disk (Fig. 252) with midlongitudinal sulcus expanding posteriorly into triangular depression embracing basal fossae; sulcus and triangular depression often with metallic color. Cap of anterior metasternal angle usually simple, sometimes bifurcate; anterior surface of metasternum smooth above anterior margin of disk. Elytral striae not distinctly carinulate (Fig. 233). Length 18 - 25 mm. Male – Head horn (Fig. 225 - 226) erect, plate-like, set very near eyes; width clearly less than interocular distance, sides convergent to divergent; apex tridentate, emarginations separating teeth shallow. Pronotal prominence (Fig. 252) strongly bilobate, lobes directed anterolaterally and flanked by deep, oval concavities; width of prominence approaching distance between outer margins of eyes. Apical processes of parameres hook-like. Female – Transverse crest of pronotum rounded, at most only very weakly bidentate; crest followed by weak concavity, summit weakly bitumose. Specimens examined – 138.	en	Edmonds, W. D., Zidek, J. (2010): A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 2010 (129): 1-111, DOI: 10.5281/zenodo.5352924
3A7F7B2C7F2B8827FF34FD5224F4EA92.taxon	distribution	Distribution. Chaco and Pampa provinces (Fig. 237). Collection Records. ARGENTINA: Buenos Aires – Mercedes (Mar); San Isidro (Jan); General Sarmiento San Miguel (Jan); Buenos Aires (Feb, Oct); San Fernando (Dec); Zelaya (Feb). Chaco – Cruce Loma Plata (Jan); Miraflores (Apr). Córdoba – Río Primero (Feb); Cruz Alta (Feb). Corrientes – Monte Caseros (Sep). Entre Ríos – Liebig (Dec); Pronunciamiento (Feb-Mar, Oct). Formosa – Formosa (Nov); Gran Guardia (Oct). Misiones – Dos de Mayo (Feb); Puerto Iguazú (Nov). Santiago del Estero – Santiago del Estero (Mar). BRAZIL: Mato Grosso do Sul – Corumbá [Pantanal do Miranda] (Jan). Paraná – Rolândia (Nov). Rio Grande do Sul – Pelotas (Dec). Santa Catarina – São Francisco (Nov). PARAGUAY: Boquerón – Mariscal Estgarribia (Feb, May). Presidente Hayes – Cruce Loma Plata (Feb); Estación Yui (Dec); Laguna Capitan (Jan); Lolita (Yaragui) (Jan). URUGUAY: Canelones – Las Piedras (Jan); Ruta 107, km 43 Canelones-Chico (Jan). Lavalleja – Sierra de Minas. Montevideo – La Teja (Dec). Río Negro – Haedo (Feb).	en	Edmonds, W. D., Zidek, J. (2010): A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 2010 (129): 1-111, DOI: 10.5281/zenodo.5352924
3A7F7B2C7F2B8827FF34FD5224F4EA92.taxon	discussion	Comments. Barattini and Sáenz (1953) described the reproductive behavior of C. milon. The shape of the male horn varies in this species. Usually it is parallel-sided, but in certain large individuals the sides diverge apically. Coprophanaeus (C.) christophorowi (Olsoufieff, 1924) Fig. 238, 254 - 258	en	Edmonds, W. D., Zidek, J. (2010): A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 2010 (129): 1-111, DOI: 10.5281/zenodo.5352924
3A7F7B2C7F2B8827FF34FD5224F4EA92.taxon	materials_examined	Type. Holotype male, Muséum National d’Histoire Naturelle, Paris (examined by photo).	en	Edmonds, W. D., Zidek, J. (2010): A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 2010 (129): 1-111, DOI: 10.5281/zenodo.5352924
3A7F7B2C7F2B8827FF34FD5224F4EA92.taxon	diagnosis	Diagnosis. General – Largely weakly shining black; anterior pronotal margin with weak violet-green reflections (reddish sheen in Fig. 257 produced by teneral surface). Posterior (ventral) surface of protibia smooth except for single or double row of coarse punctures (as in Fig. 230, arrow) paralleling longitudinal carina. Cap of anterior angle of metasternum very weakly forked. Anterior surface of metasternum glabrous above anterior margin of metasternal disk, otherwise sparsely, weakly punctured. Basal pronotal fossae effaced. Length 17 - 23 mm. Male – Head horn (Fig. 256) attenuated, base very short, lateral tubercles very acute. Pronotal prominence (Fig. 256 - 257) topped by simple transverse carina, anterior surface almost vertical, with elongate lateral concavities. Apical processes of parameres large, hook-like (Fig. 258). Female – Cephalic carina weakly bowed anteriorly, trituberculate, middle tubercle somewhat larger than laterals; length of clypeus equal to or greater than that of frons. Pronotal ridge angulate medially, not distinctly tuberculate. Specimens examined – 18.	en	Edmonds, W. D., Zidek, J. (2010): A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 2010 (129): 1-111, DOI: 10.5281/zenodo.5352924
3A7F7B2C7F2B8827FF34FD5224F4EA92.taxon	distribution	Distribution. French Guiana (Fig. 238). Collection Records. FRENCH GUIANA: Saül, Mt. Galbao, 3 o 37 ’ 18 ” N 53 o 16 ’ 42 ” N, 740 m (Jun); 7 km N Saül, 11 km NW Les Eaux Claires [Rue de Belizon Trail] 3 o 39 ’ 46 ” N 53 o 13 ’ 19 ” W, 280 m; 18.4 km SSE Roura, 4 o 36 ’ 38 ” N 53 o 13 ’ 25 ” W, 240 m (May); Nouragues Field Station, 4 o 05 ’ N 52 o 40 ’ W (Apr).	en	Edmonds, W. D., Zidek, J. (2010): A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 2010 (129): 1-111, DOI: 10.5281/zenodo.5352924
3A7F7B2C7F2B8827FF34FD5224F4EA92.taxon	discussion	Comments. Coprophanaeus christophorowi is a taxonomic isolate within the telamon species subgroup; its highly localized distribution is similar to that of Oxysternon durantoni (Edmonds and Zidek 2004). The violet-green color and shape of the male pronotum are unique to the subgroup.	en	Edmonds, W. D., Zidek, J. (2010): A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 2010 (129): 1-111, DOI: 10.5281/zenodo.5352924
3A7F7B2C7F268827FF34FC92234DEF52.taxon	description	Fig. 224, 228, 230, 238, 259 - 263	en	Edmonds, W. D., Zidek, J. (2010): A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 2010 (129): 1-111, DOI: 10.5281/zenodo.5352924
3A7F7B2C7F268827FF34FC92234DEF52.taxon	materials_examined	Type. Holotype male, private collection of Patrick Arnaud, Saintry sur Seine, France.	en	Edmonds, W. D., Zidek, J. (2010): A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 2010 (129): 1-111, DOI: 10.5281/zenodo.5352924
3A7F7B2C7F268827FF34FC92234DEF52.taxon	diagnosis	Diagnosis. General – Dorsum black with weak green to greenish red reflections along margin of pronotum. Posterior (ventral) surface of protibia smooth except for row of coarse punctures paralleling longitudinal carina (Fig. 230, arrow). Cap of anterior angle of metasternum deeply bifurcated, arms of cap embracing small, round fossa (Fig. 224). Anterior surface of metasternum densely, coarsely punctured, punctures extending onto metasternal disk adjacent to mesocoxae (Fig. 228). Basal pronotal fossae small, distinct. Length 13 - 17 mm. Male – Head horn (Fig. 261) small, sides of base convergent, central process straight, length equal to that of base. Pronotal prominence (Fig. 261 - 262) a pair of rounded, blister-like tumosities separated by longitudinal depression. Apical processes of parameres tooth-like, attenuated, tips scarcely visible (viewed from side, Fig. 263), not appressed dorsally. Female – Cephalic carina trituberculate, weakly bowed anteriorly; length of clypeus greater than that of frons. Pronotal carina only weakly raised, feebly bidentate medially. Specimens examined – 13.	en	Edmonds, W. D., Zidek, J. (2010): A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 2010 (129): 1-111, DOI: 10.5281/zenodo.5352924
3A7F7B2C7F268827FF34FC92234DEF52.taxon	distribution	Distribution. Southern Amazonian subregion (Madeira and Tapajós-Xingu provinces) (Fig. 238). Collection Records. BRAZIL: Acre – Rio Branco, Fazenda Catuaba (Feb). Pará – Redenção, 7 o 46 ’ S 51 o 58 ’ W (Nov). Rondônia – 62 km S Ariquemes [Fazenda Rancho Grande], 10 o 32 ’ S 62 o 48 ’ W (Nov); 9 km NE Cacaulândia (Jan, Nov). PERU: Madre de Díos – Manu [Los Amigos Field Station], 12 o 34 ’ S 70 o 06 ’ W, 290 m (Apr).	en	Edmonds, W. D., Zidek, J. (2010): A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 2010 (129): 1-111, DOI: 10.5281/zenodo.5352924
3A7F7B2C7F268827FF34FC92234DEF52.taxon	discussion	Comments. This species is similar in size and shape to C. parvulus, from which it is most easily distinguished by the completely coarsely punctate anterior metasternal surface and shape of the parameres. We have seen specimens only from widely scattered localities in the southern portion of the Amazon basin. The form of the male head horn and pronotum of this species, as well as C. parvulus, is like that of the smallest males of C. telamon and C. corythus.	en	Edmonds, W. D., Zidek, J. (2010): A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 2010 (129): 1-111, DOI: 10.5281/zenodo.5352924
3A7F7B2C7F208821FF34FF72232AEED2.taxon	description	Fig. 229, 238, 264 - 266	en	Edmonds, W. D., Zidek, J. (2010): A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 2010 (129): 1-111, DOI: 10.5281/zenodo.5352924
3A7F7B2C7F208821FF34FF72232AEED2.taxon	materials_examined	Type. Lectotype male (des. by Arnaud, 1982 a: 116), Muséum National d’Histoire Naturelle, Paris (examined by photo).	en	Edmonds, W. D., Zidek, J. (2010): A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 2010 (129): 1-111, DOI: 10.5281/zenodo.5352924
3A7F7B2C7F208821FF34FF72232AEED2.taxon	diagnosis	Diagnosis. General – Dorsum black with weak green to greenish red reflections along margin of pronotum. Posterior (ventral) surface of protibia smooth except for row of coarse punctures paralleling longitudinal carina (as in Fig. 230, arrow). Cap of anterior angle of metasternum deeply bifurcated, arms of cap embracing small, round fossa. Anterior surface of metasternum glabrous above anterior margin of metasternal disk, otherwise sparsely, weakly punctured. Basal pronotal fossae effaced (Fig. 229). Length 13 - 17 mm. Male – Head horn small, sides of base convergent, central process straight, length equal to that of base. Pronotal prominence (Fig. 264) a pair of rounded, blister-like tumosities separated by longitudinal depression. Apical processes of parameres tooth-like, closely appressed dorsally and clearly visible laterally (Fig. 266). Female – Cephalic carina trituberculate, weakly bowed anteriorly; length of clypeus greater than that of frons. Pronotal carina only weakly raised, feebly bidentate medially. Specimen examined – 104.	en	Edmonds, W. D., Zidek, J. (2010): A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 2010 (129): 1-111, DOI: 10.5281/zenodo.5352924
3A7F7B2C7F208821FF34FF72232AEED2.taxon	distribution	Distribution. Northern Amazonian subregion (Guyana, Humid Guyana, Amapá and Roraima provinces) (Fig. 238). Collection Records. BRAZIL: Amazonas – Manaus (Oct); 26 km NE Manaus [Reserva Forestal Ducke] (Feb, Apr, Jul, Sep-Oct, Dec); 60 km N Manaus [Fazenda Esteio], 2 o 25 ’ S 59 o 50 ’ W (Jan, Jul). Pará – Tucurí (Dec); Monte Dourado, 1 o 01 ’ S 52 o 44 ’ W, 100 m. COLOMBIA: Amazonas – Leticia [Tarapacá] (Jul-Aug). FRENCH GUIANA: Crique Plomb (Sep); Belizon (May); Saül (Jul); 7 km NW Saül, Mt. La Fumée, 3 o 39 ’ 49 ” N 53 o 13 ’ 19 ” W, 490 m (Jun); 18.4 km SSE Roura, 4 o 36 ’ 38 ” N 53 o 13 ’ 25 ” W, 240 m; Nouveau Chantier; St. Jean du Maroni; Nouragues Field Station, 4 o 05 ’ N 52 o 40 ’ W (Mar-Apr, Oct-Nov). SURINAME: Paramaribo – Pallisadenweg (Jul). Sipaliwini – Central Suriname Nature Reserve, environs of Lolopasi field station, 4 o 42.91 ’ N 56 o 12.83 ' ’ W, 80 m (Mar); Nassau Mt., 4 o 48 ’ 36 ” N 54 o 31 ’ 16 ” W (Jun); Lely Plateau, 4 o 16 ’ N 54 o 44 ’ W, 640 m (Oct). VENEZUELA: Amazonas – El Infierno [Distr. Atures] (May, Jul). Bolívar – 20 km NW Guri (Jul-Aug); 8 km N Guri, 200 m (Jul-Aug); Lago Guri Lslands, 7 o 21 ’ N 62 o 52 ’ W, 270 m (Jun-Jul).	en	Edmonds, W. D., Zidek, J. (2010): A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 2010 (129): 1-111, DOI: 10.5281/zenodo.5352924
3A7F7B2C7F208821FF34FF72232AEED2.taxon	discussion	Comments. This is a widely distributed Amazonian species that often occurs in the same communities as C. jasius and C. lancifer. It is similar to C. degallieri (q. v.), but the distributions of the two species as now known are largely separate latitudinally similar to the case of Oxysternon festivum and O. macleayi (Edmonds and Zidek 2004). The form of the male head horn and pronotum of this species, as well as C. degallieri, is like that of the smallest males of C. telamon and C. corythus. After the distribution map (Fig. 238) was prepared, we had the opportunity to examine a single specimen of C. parvulus from northern Loreto, Peru (2 o 55 ’ 26 " S 74 o 49 ’ 08 " W; ~ 190 km WNW Iquitos), a considerable distance (roughly 850 km) west of the range indicated in Fig. 238. Its presence there is unsurprising, and we suspect that intervening collection sites will be identified in the future.	en	Edmonds, W. D., Zidek, J. (2010): A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 2010 (129): 1-111, DOI: 10.5281/zenodo.5352924
3A7F7B2C7F208823FF34F972240EEE72.taxon	description	Fig. 232, 235, 238, 267 - 272	en	Edmonds, W. D., Zidek, J. (2010): A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 2010 (129): 1-111, DOI: 10.5281/zenodo.5352924
3A7F7B2C7F208823FF34F972240EEE72.taxon	materials_examined	Type. Lectotype male, here designated, Staatliches Museum für Tierkunde, Dresden.	en	Edmonds, W. D., Zidek, J. (2010): A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 2010 (129): 1-111, DOI: 10.5281/zenodo.5352924
3A7F7B2C7F208823FF34F972240EEE72.taxon	diagnosis	Diagnosis. General – Metallic coloration (Fig. 267 - 271), where present, bright red (rarely with green highlights, Fig. 269). Clypeus strongly and abruptly angulate lateral to median teeth. Posterior (ventral) surface of protibia completely granulorugose (as in Fig. 231). Cap of anterior angle of metasternum deeply bifurcate. Anterior surface of metasternum completely densely punctured. Metasternal disk finely punctured, bearing compound row of larger punctures along anterior margin. Basal pronotal fossae distinct. Elytral interstriae 2 - 5 raised midlongitudinally (Fig. 232). Length 20 - 25 mm. Male – Sides of base of head horn usually slightly convergent, rarely slightly divergent; anterior surface weakly concave, lateral angles acutely tuberculate; central process about same length as base, almost upright and angularly produced on posterior surface (as in Fig. 220). Pronotal prominence (Fig. 267 - 269) with rounded lateral angles, cleat-shaped, flanked by deep oval concavities; width equal to distance between outer margins of eyes. Female – Cephalic carina trituberculate; length of clypeus greater than that of frons. Specimen examined – 87.	en	Edmonds, W. D., Zidek, J. (2010): A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 2010 (129): 1-111, DOI: 10.5281/zenodo.5352924
3A7F7B2C7F208823FF34F972240EEE72.taxon	distribution	Distribution. Yungas province in Bolivia and Peru (Fig. 238). Collection Records. BOLIVIA: Cochabamba – Prov. Chaparé, 17 o 10.47 ’ S 65 o 46.66 ’ W, 1550 m (Feb, Nov); Yungas del Palmar, 1200 m (Jan-Feb); Limbo, 2000 m (Jan); 109 km E Cochabamba, 17 o 08 ’ 52 ” S 65 o 42 ’ 54 ” W, 1400 m (Feb); 124 km E Cochabamba [at Río Espírito Santo], 17 o 03 ’ 45 ” S 68 o 38 ’ 38 ” W, 700 m (Feb); road from Cochabamba to Villa Tunari, 17 o 08 ’ 50 ” S 65 o 42 ’ 29 ” W, 1480 m (Feb); Cordillera Mostenes, Isiboro-Securé National Park [Laguna Carachupa], 16 o 13 ’ 58 ” S 66 o 24 ’ 54 ” W, 1310 m (Sep); Carmen Pampa, 16 o 37 ’ 06 ” S 66 o 28 ’ 50 ” W, 1820 m. Santa Cruz – near Bermejo [Refugio Los Volcanes], 18 o 06 ’ 50 ” S 63 o 36 ’ 15 ” W, 1080 m. PERU: Cusco – San Pedro, 13 o 03 ’ 15 ” S 71 o 32 ’ 54 ” W; Megatoni National Sanctuary, 12 o 11 ’ 14 ” S 72 o 28 ’ 14 ” W, 1350 - 1900 m (May); La Convención, Echarate, 12 o 40 ’ 14 " S 73 o 10 ’ 21 " W, 1722 m (Oct). Junin – Puerto Eden [Río Tambo valley], 1400 m (Mar-Apr); Calabaza, 2000 m (Sep); near Satipo, 1200 m (Dec).	en	Edmonds, W. D., Zidek, J. (2010): A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 2010 (129): 1-111, DOI: 10.5281/zenodo.5352924
3A7F7B2C7F208823FF34F972240EEE72.taxon	discussion	Comments. Olsoufieff (1924) was unable to see specimens of Felsche’s species, which he tentatively regarded as a variety of C. telamon. Janssens (1940) countered Olsoufieff “ … cette espèce est parfaitemente valable … ”, but was himself countered by Pereira and Martínez (1956; see also 1960), who combined it along with C. corythus as a subspecies of C. telamon. We agree with Janssens and with Arnaud (2002 c) that Felsche’s taxon merits species recognition. The form of the male horn and pronotum of C. ignecinctus is very similar to that of C. telamon (q. v.). The central projection of the horn is strongly bent anteriorly, and the angle of bend is angularly produced but never bituberculate. Felsche’s (1909) description of this species is brief and includes no mention of the number of specimens he had on hand. The Felsche collection holdings in Dresden include six C. ignecinctus, only two (a pair) of which bear labels specifying the type locality, Marcapata, Peru. These two we consider syntypes, and it is the male which we hereby designate the lectotype. It is a small, worn individual missing the protarsal spurs as well as the left mesotarsus and both metatarsi. It bears five labels: a) handwritten in black on white “ Marcapata, Peru ”; b) printed on green “ Coll. C. Felsche, Kauf 20, 1918 ”; c) handwritten “ Ph. ignecinctus [male symbol] [unintelligible] ”; d) printed on white “ Staatsl. Museum für Tierkunde Dresden ”; and e) our lectotype designation printed on white with red margin.	en	Edmonds, W. D., Zidek, J. (2010): A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 2010 (129): 1-111, DOI: 10.5281/zenodo.5352924
3A7F7B2C7F22881DFF34F9122382EDD2.taxon	description	Fig. 217, 220 - 222, 231, 238, 273 - 274	en	Edmonds, W. D., Zidek, J. (2010): A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 2010 (129): 1-111, DOI: 10.5281/zenodo.5352924
3A7F7B2C7F22881DFF34F9122382EDD2.taxon	materials_examined	Type. Holotype male, Museum für Naturkunde der Humboldt-Universität, Berlin. Diagnosis. General – Dorsum almost completely black, metallic coloration, where present, green or yellowish green. Clypeus strongly and abruptly angulate lateral to median teeth. Posterior (ventral) surface of protibia completely granulorugose (Fig. 231). Cap of anterior angle of metasternum simple, not bifurcate. Anterior surface of metasternum glabrous above anterior margin of metasternal disk, otherwise sparsely, weakly punctured. Metasternal disk finely punctured, lacking row of larger punctures along anterior margin. Basal pronotal fossae distinct. Elytral interstriae weakly and evenly convex. Length 14 - 26 mm (usually> 20 mm). Male – Base of head horn parallel-sided, anterior surface strongly concave, lateral angles acutely tuberculate; central process as long or longer than base, usually strongly inclined posteriorly and angularly bent in the middle (Fig. 221 - 222), bend often tuberculate on each side (Fig. 222, arrow). Pronotal prominence (Fig. 274) with acute lateral angles, cleat-shaped, flanked by deep oval concavities; width equal to distance between outer margins of eyes. Female – Cephalic carina trituberculate, weakly bowed anteriorly; length of clypeus greater than that of frons. Specimen examined – 531.	en	Edmonds, W. D., Zidek, J. (2010): A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 2010 (129): 1-111, DOI: 10.5281/zenodo.5352924
3A7F7B2C7F22881DFF34F9122382EDD2.taxon	distribution	Distribution. Western Amazonia from Bolivia to Venezuela (Pantanal, Ucayali, Napo, Venezuelan Llanos provinces) (Fig. 238). Collection Records. BOLIVIA: Beni – Villa El Carmen, 14 o 38 ’ 26 ” S 65 o 01 ’ 33 ” W (Jul); Paraparu, 13 o 49 ’ S 64 o 26 ’ W, 160 m; Río Negro, 13 o 43 ’ S 67 o 22 ’ W, 260 m. Cochabamba – Parque Departamental Altamachi, 16 o 02 ’ S 66 o 40 ’ W, 1150 m (Sep); Laguna Carachupa (Cordillera Mostenes), 16 o 14 ’ S 66 o 25 ’ W, 1250 m; Sacta, 17 o 04 ’ 18 ” S 64 o 46 ’ 00 ” W and 17 o 06 ’ 19 ” S 64 o 46 ’ 57 ” W, 250 m (Dec); 67.5 km E Villa Tunari [Estación Biológica Valle Sajta], 17 o 06 ’ 19 ” S 64 o 46 ’ 57 ” W, 300 m (Feb); General Román, 450 m (Feb); Yungas del Palmar, 1200 m; Chimoré, 250 m (Jan); Río Coni, 400 m; Villa Tunari (Nov); 117 km E. Cochabamba, at Lagunitas 17 o 06 ’ 22 ” S 65 o 40 ’ 57 ” W, 1000 m (Feb); 124 km E. Cochabamba, at Río Espirito Santo, 17 o 03 ’ 45 ” S 65 o 38 ’ 38 ” W, 700 m (Feb); Río Ipiri, 16 o 2 ’ S 66 o 40 ’ W, 1100 m, 16 o 3 ’ S 66 o 41 ’ W, 700 m, 16 o 3 ’ S 66 o 40 ’ W, 900 m; San Antonio; Valle Sacta, 17 o 07 ’ S 64 o 45 ’ W, 230 m (Mar); Sacta, 17 o 06 ’ 19 ” S 64 o 46 ’ 57 ” W, 250 m (Dec); San Francisco, 280 m (Nov). La Paz – Parque Nacional Madidi, 13 o 38 ’ S 68 o 44 ’ W, 260 m (Aug); Chalalán, 14 o 26 ’ S 67 o 55 ’ W, 360 m; Río Cocos, 14 o 4 ’ S 68 o 50 ’ W, 1000 m; Serranía del Tigre, 13 o 35 ’ S 68 o 38 ’ W, 700 m; Alto Madidi, 13 o 38 ’ S 68 o 44 ’ W, 260 m; San Miguel de Huachi [Boopi Playa] (Dec); Guanay [Uyapi] (Oct). Pando – 2 km E Fortaleza, 120 m; Guayamerín, 11 o 50 ’ S 65 o 22 ’ W, 120 m (Feb); 2.5 km SW Fortaleza del Abuna, 9 o 47 ’ S 65 o 30 ’ W (Feb). Santa Cruz – Lagunita Caparú, 14 o 48 ’ S 61 o 10 ’ W, 180 m; Parque Nacional Noel Kempff Mercado, 13 o 57 ’ 43 ” S 60 o 49 ’ 45 ” W, 14 o 33.5 ’ S 60 o 55.9 ’ W and 14 o 31 ’ 16 ” S 60 o 22 ’ 48 ” W (Jan); Los Volcanes, 17 o 57.5 ’ S 63 o 25 ’ W, 560 m (Feb); 4.5 km S Buena Vista [Hotel Flora y Fauna], 17 o 29.925 ’ S 63 o 39.128 ’ W, 440 m (Dec, Feb). BRAZIL: Acre – Sector Florestal Rio Branco [PZ-UFAC] (Feb); Palmar de las Islas, 19 o 25 ’ S 60 o 32 ’ W, 270 m (Feb). Mato Grosso – Mun. Diamantino, Fazenda São João, 14 o 23 ’ 49 ” S 56 o 09 ’ 30 ” W, 480 m (Jan). Rondônia – Porto Velho (Mar); 62 km S Ariquemes (Fazenda Rancho Grande), 10 o 32 ’ S 62 o 48 ’ W, 165 m (Sep-Oct); 9 km NE Cacaulândia (Feb); 378 km S Porto Velho [Vilha Rondônia] (Jan-Feb). COLOMBIA: Amazonas – Leticia, 200 m (Feb-Mar); Parque Nacional Amacayacu [Leticia], 70 m (Dec). Boyacá – El Mortiño (Aug). Boyacá – near Pajarito, 5 o 23 ' 39 " N 72 o 41 ’ 17 " W, 1000 m (Jun). Caquetá – Puerto Solario, PNN La Serranía de Chiribiquete, 0 o 12 ’ 47.8 " N 72 o 25 ’ 25.4 " W, 250 m (Feb- Mar); Gigante (Feb-Apr). Guainía – Caño Bocón, Santa Rosa, 3 o 04 ’ 46 " N 68 o 02 ’ 26 " W, 100 m (Nov.). Guaviare – San José [Finca La Esmeralda], 2 o 33 ’ N 72 o 38 ’ W, 240 m (Oct); RN Nukak Maku, 2 o 10 ’ 35 " N 71 o 10 ’ 58 " W. Meta – Puerto Gaitan, 150 m (Nov); Pedro de Arimena, 250 m (Nov); Centro Agroturistico CAFAM, 4 o 17.6 ’ N 72 o 32.8 ’ W (Nov); San Martín, 335 m (Apr); Villavicencio (Feb-Mar, Nov); Río Duda, PNN Tinigua, 2 o 40 ’ N 74 o 10 ’ W, 350 m (Jan); Vista Hermosa, 3 o 02 ’ 44 " N 73 o 35 ’ 42 " W, 200 m (Apr) Norte de Santander – 3 km N Chinácota, 1000 m (May); Sur [Bucaramanga] (Nov); ~ 40 km SSE Toledo, PNN Tamá, 7 o 07 ' N 72 o 13 ’ W, 1450 m (Sep). Putumayo – Mocoa (Mar). Vaupés – Caparú Biological Station, Río Apoporis, 1.1 oS 69.5 oW, 200 m (Nov-Dec). ECUADOR: Morona Santiago – Untsuants [Cordillera de Cutucú], 2 o 32 ’ 57 ” S 77 o 53 ’ 23 ” W, 600 m. Napo-Orellana – Yasuní Biological Station, 00 o 40 ’ S 77 o 24 ’ W, 215 m (Feb, Jun-Oct); Puerto Franciso de Orellana [aka Coca] (Jun); Ávila (Jul); Dureno [Río Aguarico] (Sep); Tena, 400 m (Feb, Jul); 3.3 km E Puerto Napo (Jul); 12 km WSW Tena, 600 m (Jul); 20 km S Tena (Jul); 21 km E Puerto Napo [Jatun Sacha Biological Station], 1 o 03 ’ 57.6 ” S 77 o 36 ’ 59.6 ” W (Feb-Mar, Jul); Lago Agrio, 200 m (Jun); Daimi (Sep); Talag Pimpilata, 750 m (Feb); Yampuna (Jan); Archidona (Sep); Tiputini Biological Station, 00 o 38 ’ S 76 o 09 ’ W, 220 m (Sep); Aliñahui, 24 km E Atahualpa, 1 o 02 ’ 56.8 ” S 77 o 36 ’ 06.5 ” W (Mar); 24.5 km E Ahuano, 1 o 00 ’ 08.0 ” S 77 o 27 ’ 13.8 ” W (Feb-Mar); 29 km E, 1.5 km N San Pedro de Arajuno, 00 o 58 ’ 48.2 ” S 77 o 25 ’ 25.2 ” W, 360 m (Dec); 0.6 km E Río Arajuno, 1 o 05 ’ 26.5 ” S 77 o 35 ’ 30.1 ” W, 380 m (Dec); Payamino Research Station, 00 29 ’ 36.01 ” S 77 17 ’ 29.15 ” W, 400 m (Jul-Aug). Pastaza – Llandia [17 km N Puyo], 1000 m (Jul); Puyo, 1 o 28 ’ 56 ” S 77 o 59 ’ 56 ” W, 940 m (Mar, Jun, Nov); 22 km SE Puyo (Jul). Sucumbíos – Limoncocha, 250 m (Jun); 2 km N Limoncocha, 250 m (Jun). Tungurahua – Baños (Jan); Río Negro, 1 o 24 ’ S 78 o 13 ’ W. PERU: Quiroz, Río Paucartambo. Cusco – Huayllcyumbre, 13 o 11 ’ 42.2 S 70 o 39 ’ 27.7 ” W, 650 m (Jun); Megantoni National Sanctuary, 12 o 09 ’ 44 ” S 72 o 34 ’ 28 ” W, 730 - 900 m (Apr); Saucaetambo, Cosnipata (Koshñipata) River (Mar). Huánuco – Tingo María, 700 m (Jul, Oct- Dec); Cuchras [Haullaga River Valley] (Jun). Junín – Calabaza (Sep); Satipo (Mar, May). Loreto – Campamento San Lorenzo, 2 o 18 ’ 44.85 ” S 75 o 51 ’ 46 ” W, 175 - 215 m (Jun); Pucallpa (Jul, Aug). Madre de Díos – Pantiacolla Lodge, 8 km NW El Mirador trail, upper Madre de Díos River, 12 o 39 ’ 22 ” S 71 o 13 ’ 55 ” W, 400 m (Oct); Cocha Cashu Biological Station, Manu National Park, 11 o 53 ’ 45 ” S 71 o 24 ’ 24 ” W, 350 m (Oct); Cocha Salvador Reserve Zone, Manu National Park, 12 o 00 ’ 13.5 ” S 71 o 31 ’ 36 ” W, 310 m; 15 km NE Puerto Maldonado [Reserva Cuzco Amazónica], 200 m, 12 o 33 ’ S 69 o 03 ’ W (Jun); Río Tambopata, Ccolpa de Guacamayos, 13 o 08.5 ’ S 69 o 36.4 ’ W, 300 m (Oct); Atalaya [Hotel Amazonia, Río Madre de Díos] (Jun); Río Patuyacu [La Viuda Camp], 12 o 52 ’ S 68 o 55 ’ 33 ” W, 400 m (Mar); Río Patuyacu [Oculto Camp], 12 o 39 ’ 00 ” S 65 o 55 ’ 33 ” W, 230 m (Sep); Río Madre de Díos [Los Dos Amigos camp], 12 o 34 ’ 10.0 ” S 70 o 06 ’ 01.4 ” W, 250 m (Apr). Pasco – Santa Rosa [Río Palcazu] (Sep); Paujil [Parque Nacional Yanachaga-Chemillén], 500 m (Sep). San Martín – San Jacinto (Dec); Moyobamba [Dist. Naranjos, San Augustín] (Jan, May-Jul); Mishqui-Yacu, Moyobamba, 1200 m (Jul). VENEZUELA: Amazonas – La Tuquerena Rubio (Apr). Aragua – Portochuelo Pass, Parque Nacional Henri Pittier, 1200 m (Jun); Estación Biológica Rancho Grande, Parque Nacional Henri Pittier, 1100 m (Feb, Jun, Aug, Dec); Cuyagua, 800 m (Oct). Bolívar – El Dorado [Minas de Paypal] (May); km 90 on road from El Dorado to Santa Elena (Feb). Falcón – Sector Cumbre de Urea-El Chorro [Mpio. Petit], 1200 m (Jul). Mérida – 20 km SE Azulita [La Carbonera Biological Reserve], 2300 m (Jun); Sector Cuenca del Río Capara [Mpio. Padre Noguera], 7 o 50 ’ 32 ” N 71 o 30 ’ 34 ” W. Táchira – Presa La Honda, 1100 m (Dec); Pregonero (Las Trampitas), 1240 m (Jul); La Fundación [Presa Las Cuevas], 600 m (Jul); San Cristóbal, 1200 m (Aug); 42 km SE San Cristóbal, 700 m; 20 km NE San Cristóbal, 1200 m (May); 10 km NE San Cristóbal (Parque Nacional Chorro del Indio), 7 o 43 ’ 46 ” N 72 o 12 ’ 17 ” W, 1300 m (May, Oct); Río Frio, 500 m (Aug); Santo Domingo, 300 m (Aug).	en	Edmonds, W. D., Zidek, J. (2010): A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 2010 (129): 1-111, DOI: 10.5281/zenodo.5352924
3A7F7B2C7F22881DFF34F9122382EDD2.taxon	discussion	Comments. This species is widely distributed at lower elevations (<1000 m) along the eastern slopes of the Andes. Its range comes close to that of C. corythus in northwestern Venezuela, where the dividing line between the two is the Mérida – Zulia cordillera. While its ecological distribution is normally in mesophilic forests, it can enter more xeric habitats (Vidaurre et al. 2008). The normal form of the male head horn is unique and closely approached only in C. ignecinctus. In some populations (e. g. from Bolivia’s Parque Nacional Noel Kempff Mercado) the central process is cylindrical, bent forward and lacks any hint of the posterior tubercles.	en	Edmonds, W. D., Zidek, J. (2010): A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 2010 (129): 1-111, DOI: 10.5281/zenodo.5352924
3A7F7B2C7F1C881EFF34FA5222A0ED72.taxon	description	Fig. 218 - 219, 223, 238, 275 - 279	en	Edmonds, W. D., Zidek, J. (2010): A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 2010 (129): 1-111, DOI: 10.5281/zenodo.5352924
3A7F7B2C7F1C881EFF34FA5222A0ED72.taxon	materials_examined	Type. P. corythus – lectotype male (des. by Arnaud 1982 a: 115), Muséum National d’Histoire Naturelle, Paris (examined by photo); P. perseus – lectotype male (des. by Arnaud 1982 a: 115), Muséum National d’Histoire Naturelle, Paris (examined by photo); C. nevinsoni -- holotype male, private collection of Patrick Arnaud, Saintry sur Seine, France.	en	Edmonds, W. D., Zidek, J. (2010): A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 2010 (129): 1-111, DOI: 10.5281/zenodo.5352924
3A7F7B2C7F1C881EFF34FA5222A0ED72.taxon	diagnosis	Diagnosis. General – Metallic coloration, where present, green or yellowish green. Clypeus strongly and abruptly angulate lateral to median teeth. Posterior (ventral) surface of protibia completely granulorugose (as in Fig. 231). Cap of anterior angle of metasternum simple, not bifurcate. Anterior surface of metasternum glabrous above anterior margin of metasternal disk, otherwise sparsely, weakly punctured. Metasternal disk finely punctured, lacking row of larger punctures along anterior margin. Basal pronotal fossae distinct. Elytral interstriae weakly and evenly convex. Length 16 - 26 mm (usually> 20 mm). Male – Base of head horn (Fig. 218 - 219) with convergent sides, anterior surface at most only weakly concave, lateral angles acutely tuberculate; central process conspicuously shorter than base, at most only weakly inclined posteriorly, evenly curved posteriorly and never tuberculate or angulate on posterior surface. Pronotal prominence (Fig. 275 - 276) bilobate, saddle-shaped, never with acute lateral lobes, flanked by shallow concavities; width less than distance between outer margins of eyes. Female – Cephalic carina trituberculate, weakly bowed anteriorly; length of clypeus greater than that of frons. Specimen examined – 563.	en	Edmonds, W. D., Zidek, J. (2010): A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 2010 (129): 1-111, DOI: 10.5281/zenodo.5352924
3A7F7B2C7F1C881EFF34FA5222A0ED72.taxon	distribution	Distribution. Middle America from Venezuela and Colombia along Caribbean coast to southeastern Mexico (Cauca, Chocó, Eastern Central America and Gulf of Mexico provinces) (Fig. 238). Collection Records. BELIZE: Cayo – Belmopan (Jul-Aug); Chiquibul Forest Reserve [Las Cuevas Research Station], 16 o 44 ’ N 88 o 59 ’ W, 550 m (Jun, Aug). Orange Walk – Lamanai (Feb); Río Bravo Preserve (Apr). Toledo – Punta Gorda (Apr, Sep-Oct); Blue Creek Village (Jun). COLOMBIA: Antioquia – Amalfi, 1100 m (Mar); San Luis [Río Claro], 1440 m (Mar). Bolívar – SFF Los Colorados, 9 o 51 ’ 33 " N 73 o 06 ’ 38 " W, 300 m (Aug); Mompox (Jun). Cesar – Valledupar; San Alberto Indupalma (Sep). Chocó – Pacurita, 5 o 41 ’ N 76 o 40 ’ W; Tutumendó; Quibdó; El Amargal Biological Station, 5.6 oN 77.4 oW, 150 - 500 m (Aug). Cundinamarca – Santardercito (Jul). Quindío – Buenavista, 4 o 23 ’ N 75 o 44 ’ W, 1200 m. Risaralda – Pereira, 1850 m (Apr). Santander – Vélez (Sep); Bucaramanga (Jul). Tolima – Mariquita, 650 m (Sep); Ibagué (Mar); Guyabal [Armero], 250 m (Oct). Valle de Cauca – Buenaventura [Bajo Calima], 50 m (Apr); Palmira (Feb). COSTA RICA: Cartago – Turrialba (May). Guanacaste – Parque Nacional Santa Rosa, Estación Santa Rosa, 10 o 50 ’ 21.4 ” N 85 o 37 ’ 05.8 ” W, 295 m (Jun-Jul, Nov); Canas [Taboga] (Jun-Jul). Heredia – Puerto Viejo [Finca La Selva] (Jun). Limón – La Selva (Jul, Apr); Pandora [Valle Río Estrella] (Feb); La Ceiba (Jun); Sector Cerro Cocori, Finca de E. Rojas, 150 m (Jun). Puntarenas – Parque Nacional Manuel Antonio (Nov-Dec); Estación Sirena, Parque Nacional Corcovado, 100 m (Mar, Dec); Estación Quebrada Bonita, Estación Biológica Carara, 50 m (Mar-Apr); Osa Peninsula, Río Piro La Joya, 8 o 23 ’ 56 ” N 83 o 19 ’ 51 ” W, 350 m (Aug-Sep). GUATEMALA: Petén – Uaxactún (Dec). Zacapa – La Unión, 850 m (Jul); 3.5 km SE La Unión, 1500 m (Jun). HONDURAS: Atlántida – Tela [Lancetilla Botanic Gardens], 15 o 46 ’ N 87 o 27 ’ W (May-Jun). Francisco Morazán – El Zamorano; Tegucigalpa (May). Gracias a Díos – Krausirpi, 15 o 03 ’ N 84 o 52 ’ W (May). La Paz – La Paz (May). MEXICO: Campeche – Escarcega (Jul). Chiapas – San Antonio (Aug); Bonampak, 16 o 42 ’ N 91 o 04 ’ W (Sep); Santa Rosa (Aug); Boca de Chijul (Dec); 1.2 km and 6.2 km NE Estación Biológica Chajul, 16 o 07 ’ 07.1 ” N 90 o 55 ’ 51.9 ” W and 16 o 09 ’ 10.0 ” N 90 o 54 ’ 17.1 ” W, 200 m (May); Laguna Bélgica, 16 km NW Ocozocoautla, 970 m (Jun); Palenque, 600 m (May-Jul); Rosario Izapa (May); Selva Lacandona, Laguna Miramar (Oct). Hidalgo – 6.5 km SW Chapulhuacan (Jul). Oaxaca – Distrito Yautepec, Juquila Mixes (Sep); 10 km S. Valle Nacional, 600 m (Jul); confluencia Ríos Sarabia y Coatzaocoalcos [Finca San Carlos (Mar); La Nueva Esperanza (Aug); Tuxtepec (Nov). Puebla – Calapan 19 o 55 ’ 45.92 ” S 97 o 23 ’ 07.65 ” W, 1250 m (Jul). Quintana Roo – 12 km N. Felipe Carrillo Puerto (Oct); Cedral (Mar); Tintal (Mar); X-Can (Jun-Jul, Oct); Nuevo X-Can (Oct); 68 km W Chetumal (Jul). Veracruz – Veracruz (Aug); Córdoba (Jul-Aug); Coatepec, 1200 m (May); Guadalupe Victoria, 750 m (Jun); Omealca (Oct); 33 km N Catemaco, 160 m (Jul-Aug); San Andrés Tuxtla (May- Jun); Fortín de la Flores, 900 m (Jun); Los Tuxtlas (Jul); Tezonapa (Sep); Palma Sola (Jun); 11 km W Palma Sola (Sep); Xalapa, 1350 m (Jul, Sep); Actopan, 240 m (Aug); Espinal, 900 m (Aug); Cerro Leon, 700 m (Nov); Sontecomapan (Estación Biológica Tropical Los Tuxtlas), 400 m (Jun, Aug, Oct). NICARA- GUA: Atlántico Norte – Waslala (Dec); Río San Juan – 8 km SE El Castillo [Refugio Bartola], 10 o 56.6 ’ N 84 o 20.4 ’ W (May). PANAMA: Colón – 16 km SE Colón, Santa Rita Ridge, 270 m (Jun); 14 km N junction Escobal Road and Piña Road, 20 m (Jun); Gatún Island (May-Jun, Dec); Corozal (May); Barro Colorado Island (Jul). Darien – Río Tacarcuna, 580 m (Jul); Estación Ambiental Cana, 7 o 45.32 ’ N 77 o 41.07 ’ W, 600 - 750 m (Jun). Panama – 11 - 15 km N El Llano, 350 m (May); Cerro Azul, 650 m (May); 31 km E Canita, 50 m (May); Lago Bayano, Isla Majé (Dec); Howard Air Force Base (Pacific entrance Panama Canal), 10 m (Jul-Aug); Cerro Jefe, 1000 m (May); Cerro Campana, 860 m (May). VENEZUELA: Mérida – Finca Campo Alegre [Mpio. Zea-Mérida], 8 o 28 ’ 56 ” N 71 o 44 ’ 38 ” W, 650 m (Sep).	en	Edmonds, W. D., Zidek, J. (2010): A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 2010 (129): 1-111, DOI: 10.5281/zenodo.5352924
3A7F7B2C7F1C881EFF34FA5222A0ED72.taxon	discussion	Comments. Pereira and Martínez (1956; see also 1960) regarded C. corythus (along with C. ignecinctus [q. v.]) as a subspecies of C. telamon. We here return to former usage (Gillett 1911; Olsoufieff 1924; and Blackwelder 1944, among others) and restore C. corythus to species rank. Pereira and Martínez (1956) erroneously synonymized C. chiriquensis with C. corythus. Coprophanaeus corythus is a common denizen of forest – pasture mosaic habitats of Chiapas (Arellano et al. 2008; Navarrete and Halffter 2008), more common in continuous forest than pastures. The phenology and daily activity of this species (cited as C. telamon nevinsoni) in the Maracaibo depression of extreme western Mérida, Venezuela, was reported by Gámez et al. (2006). There it occurs in both relictual forest patches and open cultivated areas, with highest frequency in forest habitat; it is collected all year with greatest abundance during rainfall peaks in April – May and October – November. The large male head horn is most variable in the South American populations of this species, where it can be intermediate between the shape in typical C. corythus and that of certain C. telamon (cf. Fig. 218 - 219, 275 - 276). Certain of these intermediates, which we regard as intraspecific variants, were named C. telamon nevinsoni by Arnaud and Gámez (2002). There seems to be no solid basis at present to maintain the taxonomic distinction. Olsoufieff (1924) erroneously synonymized Phanaeus perseus Harold with Phanaeus cerberus Harold. Olsoufieff, who examined Harold’s type, correctly pointed out that Harold did not notice that his specimen was damaged, “ … l’auteur n’a pas fait attention que l’epistome était presque symétriquement grignoté (cassé) des deux côtés des lobes aigus, ce qui faisant illusion de 4 dents séparées ”; but he mistakenly associated it with C. cerberus, the female of which is quite different. The holotype agrees fully with Harold’s description, including the misplaced reference to the broken clypeus, “ … das Kopfschild vorn mit 4 Zähnen … die seitlichen … zwischen diesen [die mittleren] und dem stumpfwinkeligen vorderen Wangeneck eine ziemlich tiefe Ausbuchtung. ” And it was the apparently strongly quadridentate clypeus that prompted him to recognize it as a new species, “ … so habe ich doch in Anbetracht der höchst eigenthümlichen Zahnung des Clypeus kein Bedenken getragen, dieselbe als neu zu beschreiben. ” Ironically, he unconsciously corrected his own error, declaring “ Sieht man von der verschiedenen Kopfbildung ab, so stimmt das Weibchen in allem übrigen vollkommen mit dem des Corythus überein. ” Harold’s type is a female C. corythus with a broken clypeus.	en	Edmonds, W. D., Zidek, J. (2010): A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 2010 (129): 1-111, DOI: 10.5281/zenodo.5352924
3A7F7B2C7F188814FF34F93225F1E912.taxon	description	Fig. 287 - 288, 291 - 296	en	Edmonds, W. D., Zidek, J. (2010): A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 2010 (129): 1-111, DOI: 10.5281/zenodo.5352924
3A7F7B2C7F188814FF34F93225F1E912.taxon	materials_examined	Type. Holotype male, private collection of Patrick Arnaud, Saintry sur Seine, France. Diagnosis. General – Dorsum largely black, metallic color (red, green or coppery green) restricted to posterior portion of head and anterior portion and margin of pronotum and pygidium; venter with subdued violet reflections. Frons short, length less than that of clypeus. Anterior (declivitous) face of metasternum with setigerous punctures concentrated medially. Length 11 - 15 mm. Male – Apical processes of parameres strongly elevated, clearly visible in profile. Pronotal prominence arcuate, lacking well-defined tubercles or angles, narrow, width less than distance between outer margins of eyes (in small males prominence reduced to arcuate ridge very near anterior margin). Female – Pronotal carina very near anterior pronotal margin, only weakly raised, lacking distinct median tubercles. Specimen examined – 34.	en	Edmonds, W. D., Zidek, J. (2010): A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 2010 (129): 1-111, DOI: 10.5281/zenodo.5352924
3A7F7B2C7F188814FF34F93225F1E912.taxon	distribution	Distribution. Upper Amazon River basin of Napo province. Collection Records. PERU: Loreto – Iquitos (Feb) (see Comments below).	en	Edmonds, W. D., Zidek, J. (2010): A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 2010 (129): 1-111, DOI: 10.5281/zenodo.5352924
3A7F7B2C7F188814FF34F93225F1E912.taxon	discussion	Comments. Thanks to Darren Mann (Oxford University Museum of Natural History) we examined and assigned to this species a long series from the type locality (Iquitos). A paratype male was cited by Arnaud (2002 a) as from “ Brasil: Para, S. Paulo de Olivença … ”. We regard the state record (Pará) as erroneous; São Paulo de Olivença lies in the state of Amazonas about 450 km downriver from Iquitos. Arnaud’s description was based on two males, neither regarded as well developed: “ Ce critère [description of male pronotum] est à prendre avec prudence car l’examen de peu de spécimens ne permet pas de juger si les exemplares sont de forme majeure. ” Our series of 34 specimens includes 15 males, none of which is more robustly developed than the holotype pictured by Arnaud (2002 c). Since we have examined neither the holotype nor the paratype, our identification of this series as C. callegarii must be regarded as provisional.	en	Edmonds, W. D., Zidek, J. (2010): A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 2010 (129): 1-111, DOI: 10.5281/zenodo.5352924
3A7F7B2C7F158814FF34FE122556EDD2.taxon	description	Fig. 291, 297 - 301	en	Edmonds, W. D., Zidek, J. (2010): A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 2010 (129): 1-111, DOI: 10.5281/zenodo.5352924
3A7F7B2C7F158814FF34FE122556EDD2.taxon	materials_examined	Type. Holotype male, Colección Boliviana de Fauna, La Paz (examined; on loan to Museo de Historia Natural Noel Kempff Mercado, Santa Cruz).	en	Edmonds, W. D., Zidek, J. (2010): A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 2010 (129): 1-111, DOI: 10.5281/zenodo.5352924
3A7F7B2C7F158814FF34FE122556EDD2.taxon	diagnosis	Diagnosis. General – Dorsum largely black, red metallic color restricted to posterior portion of head and anterior portion and margin of pronotum and pygidium; venter with scattered reflections of same tone as dorsum. Frons long, length at least one and half times that of clypeus. Anterior (declivitous) face of metasternum with setigerous punctures widely distributed over most of surface. Length 17 - 22 mm. Male – Apical processes of parameres attenuated, tips barely visible in profile. Pronotal prominence of larger individuals consisting of two conical bumps separated by small concavity. Female – Pronotal carina a distinctly raised, medially bidentate crest followed by very weak depression. Specimen examined – 23.	en	Edmonds, W. D., Zidek, J. (2010): A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 2010 (129): 1-111, DOI: 10.5281/zenodo.5352924
3A7F7B2C7F158814FF34FE122556EDD2.taxon	distribution	Distribution. Yungas province in Bolivia. Collection Records. BOLIVIA: Cochabamba -- Cordillera Mosetenes, Isiboro-Securé National Park, 16 14 ’ 10 ” S 66 24 ’ 46 ” W, 1350 m (Sep).	en	Edmonds, W. D., Zidek, J. (2010): A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 2010 (129): 1-111, DOI: 10.5281/zenodo.5352924
3A7F7B2C7F158814FF34FE122556EDD2.taxon	discussion	Comments. Coprophanaeus caroliae is known only from the low yungas of Cordillera Mosetenes, whose dung beetle community received close scrutiny from Hamel-Leigue et al. (2008). This species is similar to C. strandi (Balthasar, 1939) and apparently also to C. lecromi Arnaud, 2002 a, and it may later be judged a synonym of either or both. These two species were based on single male specimens. We have examined the holotype and only known specimen of Balthasar’s species deposited at the National Museum of Natural History in Prague, which we illustrate here for the record (Fig. 302). But we have not seen that of C. lecromi, which is housed in the private collection of its describer. Until more specimens (including females) and information on C. strandi and C. lecromi become available, a thorough comparison and analysis of the three taxa will not be possible (see Edmonds 2008, for further observations).	en	Edmonds, W. D., Zidek, J. (2010): A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 2010 (129): 1-111, DOI: 10.5281/zenodo.5352924
3A7F7B2C7F158816FF34FA522562E992.taxon	description	Fig. 289 - 291, 303 - 307	en	Edmonds, W. D., Zidek, J. (2010): A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 2010 (129): 1-111, DOI: 10.5281/zenodo.5352924
3A7F7B2C7F158816FF34FA522562E992.taxon	materials_examined	Type. Holotype male, private collection of Patrick Arnaud, Saintry sur Seine, France.	en	Edmonds, W. D., Zidek, J. (2010): A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 2010 (129): 1-111, DOI: 10.5281/zenodo.5352924
3A7F7B2C7F158816FF34FA522562E992.taxon	diagnosis	Diagnosis. General – Dorsum (Fig. 303 - 307) mostly black, golden or red coloration with green highlights present on head and anterior portion and margins of pronotum; venter with metallic highlights of same tone as dorsum. Anterior (declivitous) face of metasternum with setigerous punctures widely distributed over most of surface. Length 12 - 14 mm. Male – Pronotal prominence (Fig. 290) more-or-less trapezoidal, with acute lateral angles. Apical processes of parameres attenuated, scarcely visible in profile (Fig. 305). Female – Pronotal ridge curved, very slightly bidentate medially. Specimen examined – 9.	en	Edmonds, W. D., Zidek, J. (2010): A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 2010 (129): 1-111, DOI: 10.5281/zenodo.5352924
3A7F7B2C7F158816FF34FA522562E992.taxon	distribution	Distribution. Eastern Napo province. Collection Records. COLOMBIA: Amazonas – Leticia (Feb). Vaupés – Río Apaporis (Caparú Biological Station), 1.1 S 69.5 W, 200 m (Nov-Dec). PERU: Loreto – Quebrada Ubuya, 74 o 01 ’ 24 " W 6 o 57 ’ 19 " S, 202 m (Mar); Quebrada Punga, 74 o 00 ’ 59 " W 6 o 27 ’ 40 " S, 145 m (Feb).	en	Edmonds, W. D., Zidek, J. (2010): A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 2010 (129): 1-111, DOI: 10.5281/zenodo.5352924
3A7F7B2C7F158816FF34FA522562E992.taxon	discussion	Comments. Arnaud (2002 a, 2002 c) places C. larseni near C. suredai. Among the few specimens of C. suredai that we have seen are the Leticia paratypes in the Canadian Museum of Nature. The original description was based on 10 specimens from western Amazonia, including Lago Agrio (Ecuador) and “ Peru ”. The Canadian Museum also has two female specimens which resemble C. suredai, from Brazil: Amazonas – Estirão do Ecuador, Rio Javari (Feb); and Bolivia: Pando – 11 o 50 ’ S 65 o 22 ’ W (Feb). We could not locate Estirão do Ecuador, but it presumably lies along the Brazil – Peru border (formed by the Rio Javari) near Tabatinga (Brazil) and Leticia. The coordinates specified for the latter lie in the northern extremity of the department of El Beni (near but not in Pando), about 1000 km SSE of Leticia. Hamel et al. (2009) cite this species from 2 km E Fortaleza (Pando) and Guayaramerin (Beni) in Bolivia.	en	Edmonds, W. D., Zidek, J. (2010): A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 2010 (129): 1-111, DOI: 10.5281/zenodo.5352924
3A7F7B2C7F178810FF34FD9224A9EAB2.taxon	description	Fig. 280 - 286, 291, 308 - 313	en	Edmonds, W. D., Zidek, J. (2010): A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 2010 (129): 1-111, DOI: 10.5281/zenodo.5352924
3A7F7B2C7F178810FF34FD9224A9EAB2.taxon	materials_examined	Type. P. ohausi – lectotype male, here designated, Staatliches Museum für Tierkunde, Dresden (examined); C. florenti – holotype male, private collection of Patrick Arnaud, Saintry sur Seine, France.	en	Edmonds, W. D., Zidek, J. (2010): A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 2010 (129): 1-111, DOI: 10.5281/zenodo.5352924
3A7F7B2C7F178810FF34FD9224A9EAB2.taxon	diagnosis	Diagnosis. General – Dorsum largely black, metallic color (dark green or blue), if any, usually restricted to posterior portion of head and anterior portion and margin of pronotum and pygidium; sometimes present on elytra; venter with scattered reflections of same tone as dorsum. Frons long, length at least one and half times that of clypeus, often longer. Anterior (declivitous) face of metasternum with setigerous punctures widely distributed over most of surface. Length 16 - 21 mm. Male – Apical processes of parameres strongly elevated, clearly visible in profile. Pronotal prominence of larger individuals variable, usually distinctly bitumid, lobes rounded and usually finely carinate anteriorly (in small males prominence reduced to transverse carina above anterior margin). Female – Pronotal ridge distinctly raised, medially bidentate and followed by very weak depression. Specimen examined – 68.	en	Edmonds, W. D., Zidek, J. (2010): A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 2010 (129): 1-111, DOI: 10.5281/zenodo.5352924
3A7F7B2C7F178810FF34FD9224A9EAB2.taxon	distribution	Distribution. Yungas and western Napo provinces. Collection Records. COLOMBIA: Casanare – Aguazul, 375 m (Nov). Meta – Puerto Gaitan, 400 m (Oct). Santander – Veléz, 2400 m (Sep). ECUADOR: El Oro – Piñas, 3 42 ’ S 79 42 ’ W (Mar). Morona- Santiago – Angel Rouby (Cordillera Cutucú), 2 o 21.6 ’ S 78 o 4.0 ’ W (Feb); Cordillera Cutucú, 2 32 ’ 57 ” S 77 53 ’ 23 ” W. Napo-Orellana – Loreto road, 7.9 km E Narupa junction, 00 43 ’ 06.9 ” S 77 45 ’ 44.0 ” W, 1380 m (Mar); Río Hollin, 00 41.702 ’ S 77 43.842 ’ W, 1068 m (Dec); Aliñahui (Feb); Puerto Napo (Apr); Yasuní National Park, Yasuní Research Station, 00 o 38 ’ S 76 o 36 ’ W, 215 m; Limoncocha, 250 m (Jun); Puerto Francisco de Orellana (“ Coca ”) (May); km 11.1 road Sarayacu to Loreto, 1200 m (Jul). Pastaza – 22 km SE Puyo, 900 m (Jul); 17 km N Puyo [Llandia], 1000 m (Jul). Tungurahua – 6 km and 8 km E Río Negro, 1400 m (Jul). Zamora-Chinchipe – Bombuscaro (Parque Nacional Podocarpus), 1146 m, 4 o 06 ’ 09 ” S 78 o 57 ’ 46 ” W (May). PERU: Junin – Satipo, 1100 m (Mar). San Martín – Moyobamba (Jan); Alto Nieva (Aug).	en	Edmonds, W. D., Zidek, J. (2010): A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 2010 (129): 1-111, DOI: 10.5281/zenodo.5352924
3A7F7B2C7F178810FF34FD9224A9EAB2.taxon	discussion	Comments. Of the species comprising this group, C. ohausi exhibits the greatest variation in form of the well-developed male prothorax. The median prominence is bitumid, but its swellings are connected by a transverse carina that varies in shape. Figures 280 - 284 illustrate this variation in a single population from Satipo, Peru. In well-developed individuals of both sexes the frons widens anteriorly and produces the false impression that the transverse carina is markedly narrower than in other species. Howden and Young’s (1981) reference to C. ohausi is in fact to C. morenoi Arnaud (Howden and Gill 1987). The description of C. lichyi Arnaud suggests that it is closely related to, or perhaps synonymous with C. ohausi. The type material of C. florenti was not available to us for study. Our point of reference for synonymizing this name with C. ohausi is a large series of C. florenti in the Canadian Museum of Nature collection identified by Arnaud. The color and male pronotal structure cited by Arnaud in his description, as well as that in the aforementioned series, fit well within the range of variation of C. ohausi as here defined. The Staatliches Museum für Tierkunde collection in Dresden includes three specimens that we regard as the type series of this species. Two are labeled “ Macas ” [Ecuador] and the third “ Teremotillo ”. Felsche’s original description of Phanaeus ohausi cites two Ecuadorian localities for the material at hand: Teremotillo and Jivaria. Felsche stated that Ohaus gave him the specimen from Teremotillo – “ … das andere, das er mir gütigst überlassen, bei Teremotillo; … ”. We exclude the Macas specimens from the type series, leaving the Teremotillo specimen as putative type. We hereby designate this specimen the lectotype. We assume that the Jivaria specimen seen by Felsche is lost or elsewhere. The lectotype is a large male with dark blue dorsal metallic coloration (including on the elytra), somewhat worn and dirty, and missing the left mesotarsus and the right metatarsus. It bears nine labels: a) [white rectangle] “ Ecuad. Ostcordill. / Teremotillo / F. Ohs. 14.12.05 ”; b) [green rectangle] “ Col. C. Felsche / Kauf 20, 1918 ”; c) [green square] “ 1910 / 4 ”; d) [folded white] “ Ohausi ” handwritten in red ink; e) [oblong white] “ Ph. ohausi / {illegible} ” handwritten in black ink; f) [red] “ Typus ” printed, “ ohausi Felsche ” handwritten; g) white “ Staatsl. Museum für / Tierkunde ”; h) our lectotype label printed on white with red border.	en	Edmonds, W. D., Zidek, J. (2010): A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 2010 (129): 1-111, DOI: 10.5281/zenodo.5352924
