Coprophanaeus (Metallophanaeus) saphirinus ( Sturm, 1826 )

Cupello, Mario & Vaz-De-Mello, Fernando Z., 2014, Revalidation of the Brazilian Atlantic Forest dung beetle species Coprophanaeus (Metallophanaeus) machadoi (Pereira & d’Andretta, 1955) (Coleoptera: Scarabaeidae: Scarabaeinae: Phanaeini) based on morphological and distributional evidence, Zootaxa 3869 (4), pp. 435-451 : 443-449

Coprophanaeus (Metallophanaeus) saphirinus ( Sturm, 1826)

Figs. 10–11 , 13–23 .

Copris saphirina Sturm, 1826: 65 , 125, tab. II, fig. 15a, b.

Phanaeus sapphirinus: Dejean, 1833: 140 ; Dejean, 1836: 155; Nevinson, 1892: 7, 10; Gillet, 1911: 86.

Phanaeus saphirinus: Klug, 1841: 210 ; Sturm, 1843: 106; Harold, 1869a: 1019; Harold, 1869b: 65; Harold, 1875: 42 (misidentification of C. machadoi ); Waterhouse, 1891: 129; Ohaus, 1909: 28 –29; Kolbe, 1909: 529; Bruch, 1911: 190; d’Olsoufieff, 1924: 9, 15; Arnaud, 2002: 55; Almeida & Mise, 2009: 238; Edmonds & Zidek, 2010: 29.

Phaenaeus [sic] saphirinus: Moura et al., 1997: 271 –272.

Phanaeus (Metallophanaeus) saphirinus: d’Olsoufieff, 1924: 22 , 28, 63, 72, 74, 136, 144, pl. IV, fig. 15; Pessoa, 1934: 284 –285, 293, fig. 8a,b; Pessoa & Lane, 1941: 478; Pereira, 1949: 218 –222, fig. 1; Pereira & d’Andretta, 1955: 260; Martínez, 1959: 102; Halffter & Matthews, 1966: 30, 132; Edmonds, 1967: 97 –104 (probably in reference to C. machadoi ; see above).

Metallophanaeus sapphirinus: Blackwelder, 1944: 209 .

Phanaeus (Metallophanaeus) saphyrinus [sic]: Redtenbacher, 1868: 56; Lange, 1947: 313.

Coprophanaeus (Metallophanaeus) saphirinus: Edmonds, 1972: 842 –843, 874; Edmonds & Halffter, 1978: 322; Halffter & Edmonds, 1982: 89; Vaz-de-Mello, 2000: 192; Arnaud, 2002: 55; Cáceres & Monteiro-Filho, 2006: 1198 –1201; Almeida & Mise, 2009: 232, 238, fig. 34; Hernández & Vaz-de-Mello, 2009: 608, 611; Edmonds & Zidek, 2010: 2 –5, 22–24, 29–31, 33; Hernández et al., 2011: 5, 7–8; Silva et al., 2011: 334, 339, 342; Vaz-de-Mello et al., 2011: 54, fig. 55; Silva & Di Mare, 2012: 199–200, 202; Silva et al., 2012: 433–434, 438; Culot et al., 2013: 85, 87; Korasaki et al., 2013: 396; Silva et al., 2013: 685; Silva & Bogoni, 2014: 340 –341.

Coprophanaeus (Metallophanaeus) saphirinus saphirinus: Arnaud, 2002: 55 .

Phanaeus chabrillacii Thomson, 1857: 117 .

Phanaeus chabrillaci: Harold, 1869a: 1017 ; Harold, 1869b: 65; Edmonds & Zidek, 2010: 29 –28.

Phanaeus saphirinus var. chabrillaci: Gillet , 191: 86.

Phanaeus (Metallophanaeus) saphirinus var. chabrillaci: d’Olsoufieff, 1924: 28 , 75, 144; Pessoa, 1934: 293; Pessoa & Lane, 1941: 478.

Metallophanaeus sapphirinus [sic] var. chabrillaci: Blackwelder, 1944: 209 .

Phanaeus (Metallophanaeus) saphirinus chabrillacei: Pereira, 1949: 222 ; Pereira & d’Andretta, 1955: 260.

Coprophanaeus (Metallophanaeus) saphirinus var. chabrillacei : Vaz-de-Mello, 2000: 192.

Coprophanaeus (Metallophanaeus) saphirinus chabrillacii: Arnaud, 2002: 56 .

Type specimen: Copris saphirina Sturm, 1826 : Holotype: male (“ Brasil. B. Copris saphirinus Mihi. ” “ Type von Copris saphirinus Sturm ”, “Coll. Sturm”, “ Metallophanaeus saphirinus Sturm ”), ZSMC (examined by photos ( Fig. 15 )). Phanaeus chabrillaci Thomson, 1857 : Holotype: male, (“ Type ”, “Coll. J. Thomson”, “cf. Arch. Ent. I., 1857, p. 117”, “det. J. Thomson Phanaeus chabrillaci Th. ”, “ Ph. (Metal.) saphirinus var. chabrillaci A. Janssens rev., 1932”), ISNB (examined by photos (Fig. 16)).

Type locality: Copris saphirina : “Aus Brasilien ” ( Sturm, 1826).— Phanaeus chabrillaci : “ Brésil ” ( Thomson, 1857).

Measurements: Males (67 specimens): TL: AV: 18.93; MX: 23; MN: 15; SD: 2.16. PW: AV: 12.07; MX: 15; MN: 10; SD: 1.63. Females (66 specimens): TL: AV: 19.11; MX: 24; MN: 14; SD: 2.42. PW: AV: 12.27; MX: 19; MN: 9; SD: 1.98. Total (133 specimens): TL: AV: 19.02; SD: 2.29. PW: AV: 12.09; SD: 1.62.

Intraspecific variation and taxonomic discussion: Edmonds & Zidek (2010) stated that C. saphirinus of both sexes have paraocular areas (genae) with “ smooth, highly shining ” tegument. However, specimens observed in this work present a different condition: males follow Edmonds & Zidek’s description and have genae mostly smooth and shining, with rugosity restricted at most only adjacent to outer margin; females, on the other hand, possess coarsely rugose genae, in the same manner as the rest of the head. Small males generally possess genae that are more rugose than large males, but never as strongly as in females.

Dorsal coloration in C. saphirinus is highly varied, ranging from dark metallic blue, violet and green to entirely red, yellowish and greenish red, and dark red ( Figs. 17–23 ); ventral coloration, however, is more uniform and presents green and blue coloration and is never red ( Figs. 5 b, 15b, 16c–d). The most common dorsal form is predominantly blue, but specimens of all colors apparently coexist at least in the southern latitude or higher altitude populations. For example, the series collected in São Francisco de Paula (Rio Grande do Sul) is composed of blue, violet, green and red specimens. Similarly, specimens collected in Serra da Mantiqueira Mountains (localities as Itatiaia (Rio de Janeiro) and Passa Quatro and Virgínia (Minas Gerais)) since the beginning of the 20th century show that in this area all color variations coexist. Vaz-de-Mello (cited as personal communication in Arnaud (2002) and Edmonds & Zidek (2010)) hypothesized that red specimens tend to occur more frequently in higher elevations than does the blue form, but our present observations contradict this: red specimens occur in localities at sea level in southern locations (e.g., Porto Alegre, Rio Grande do Sul) as well as at high elevations above 1,500 m. (e.g., Campos do Jordão, São Paulo). In Virgínia (Minas Gerais), for instance, both blue and red specimens were collected together at 1,500 m., and our highest altitude record belongs, in fact, to a blue specimen (Campos do Jordão, 1,700 m.). Therefore, as these forms are apparently sympatric and there is no other characteristic differentiating them, we agree with Harold’s (1869b) and Edmonds & Zidek’s (2010) conclusions and considered color as a matter of intraspecific variation in C. saphirinus and thus Phanaeus chabrillaci as Thomson, 1857, name proposed to distinguish red specimens in a separate taxon, its junior synonym. But it is true and noteworthy that some of the localities examined by us with a large number of collected specimens, particularly São Bento do Sul (Santa Catarina), did not present red specimens, but only blue and green ones, and that, in southeastern Brazil, no red individual has been collected at lower altitudes.

Bionomics: Martínez (1959) and Halffter & Mathews (1966) erroneously regarded C. saphirinus to be strictly coprophagous. As stated by Edmonds (1972) and Edmonds & Zidek (2010), this species also presents necrophagous habits and is found in carcasses. Silva et al. (2012) and Silva & Di Mare (2012), after having collected almost 600 specimens with baited pitfall traps, observed that C. saphirinus exhibited a special attraction to rotting chicken organs and fish and thus considered this species as preferentially necrophagous, although it was also attracted to rotting banana and to human excrement. Specimens examined by us with collecting data are those collected with human feces by Mario Cupello in Parque Nacional do Itatiaia (Rio de Janeiro), M. I. M. Hernandez in Serra do Japi (São Paulo), E. Bovy in Parque Estadual da Serra do Mar (São Paulo), and G. Viegas in Rolante (Rio Grande do Sul); with tapir dung by E. Bovy in São Miguel Arcanjo (São Paulo); in a toad carcass by J. F. Zikán in Virgínia (Minas Gerais); and with carrion by M. Rocha in Porto Belo (Santa Catarina) and François Génier & Sérgio Ide in Estação Biológica da Boracéia (São Paulo). Ohaus (1909) reported this species from cattle and mule dung, Cáceres & Monteiro-Filho (2006) observed it consuming dung of a big-eared opossum ( Didelphis aurita Wied-Neuwied, 1826 ) and burying a carcass of a black-footed pygmy rice rat ( Oligoryzomys nigripes (Olfers, 1818)) , and Silva & Bogoni (2014) collected four specimens of C. saphirinus using rotten chicken eggs as bait.

As for C. machadoi , C. saphirinus has been recorded for all months from October to May, suggesting that adults of this species are inactive during the coldest period of the year.

Geographic distribution: Chacoan subregion: Parana dominion: Atlantic, Parana Forest, and Araucaria Forest provinces. BRAZIL: Minas Gerais: Aiuruoca, Camanducaia, Conceição dos Ouros, Ingaí (Reserva Biológica Unilavras/Boqueirão), Passa Quatro, Poço de Caldas, Varginha, Virgínia. Rio de Janeiro: Itatiaia (Parque Nacional do Itatiaia). São Paulo: Bananal (Serra da Bocaina), Barueri, Campos do Jordão, Cubatão, Jundiaí, Leme, Mogi das Cruzes, Parque Estadual da Serra do Mar (Núcleo Picinguaba, Núcleo Santa Virgínia), Parque Estadual Xixová-Japuí, Pindamonhangaba, Ribeirão Preto, Salesópolis (Estação Biológica de Boracéia), Rio Grande da Serra, São Bernardo do Campo, São Miguel Arcanjo (Parque Estadual Carlos Botelho), São Paulo, Serra da Cantareira, Serra do Japi, Tremembé. Paraná: Balsa Nova (Serra de São Luiz do Purunã), Curitiba, Foz do Iguaçu, Guarapuava, Lapa, Londrina, Marechal Cândido Rondon, Matelândia, Ponta Grossa, Rio Negro. Santa Catarina: Anitápolis, Antônio Carlos, Blumenau, Bombinhas (Área de Relevante Interesse Ecológico ( ARIE) Costeira de Zimbros), Campos Novos, Canoinhas, Corupá, Florianópolis, Governador Celso Ramos, Ipumirim, Itapema, Joinville, Lages, Mafra, Monte Castelo, Nova Veneza, Porto Belo (Área de Proteção Ambiental (APA) do Araça), Rio das Antas, Santo Amaro da Imperatriz, São Bento do Sul, São Bonifácio, São Domingos, São Francisco do Sul, São Pedro de Alcântara, Seara, Timbó. Rio Grande do Sul: Barão de Cotegipe, Bento Gonçalves, Caxias do Sul, Cruz Alta, Ijuí, Porto Alegre, Rolante, Santa Maria, Santo Augusto, São Borja ( Edmonds & Zidek, 2010), São Francisco de Paula, Silveira Martins, Triunfo. PARAGUAY: Cordillera. Guairá: Colonia Independencia, Natalicio Talavera, Paso Yobai, Villarrica. Paraguarí: Paraguarí. Caazapá: Parque Nacional Caaguazú. Itapúa. ARGENTINA: Misiones: Aristóbulo del Valle, Cerro Azul, Dos de Mayo, Garuhapé, Loreto, Parque Nacional Iguazu, Posadas, San Ignacio, San Pedro, San Vicente, Santa María.

We consider the reports of C. saphirinus for the Paraguayan department of Alto Paraguay, for the Argentinean provinces of Tucuman and Chaco ( Edmonds & Zidek, 2010), and for Uruguay ( d’Olsoufieff, 1924), as well as the specimen examined by us with label indicating precedence from Porto Murtinho (Mato Grosso do Sul, Brazil), as very doubtful, and therefore were excluded from the distribution presented above. The two individuals examined in the MZSP with report from Pará (Itaituba and Óbidos) are certainly mislabeled.

Material examined: 252 males and 198 females. NO DATA — 3 males ( MNHN), 1 male and 1 female ( MNRJ; female of the red form), 1 male ( BMNH), and 1 male ( OUMNH). ARGENTINA: MISIONES: Alto Paraná, Puerto Aguirre, 1933-34, K. J. Hayward col.— 1 female ( BMNH); Arroyo Cuñápirú, 02.III.1996, without collector— 1 male and 1 female ( CEMT; male of the red form); “Papiri”, 22.III.1991, without collector— 2 males ( CEMT; red form); Parque Nacional Iguazu, XII.2004 without collector— 1 female ( CPFA, red form); Posadas, 22.XI.1995, without collector— 1 male ( CEMT); Puerto Azul (=Cerro Azul?), 10.XI.2002, R. Foerster col.— 2 males and 2 females ( CPFA, all red form); San Ignacio, Tijucuare, without date and collector— 1 female ( MNHN); San Pedro, XI.2012, P. Wagner col— 2 females ( CEMT; red form); San Vicente, Salto La Golondrina, 500 m., 27º7'19''S 54º30'43''W, XI.2006, E. Abadie col.— 1 male ( CEMT; red form). BRAZIL: no more data— 2 males and 1 female ( MNHN; female of the red form) and 4 males and 1 female ( CPFA, 2 males and female of red form). PARÁ [mislabeling]: Itaituba, Santarenzinho, Rio Tapajós, III.1961, Dirings col.— 1 male ( MZSP); Óbidos, XII.1931, B. Pohl col.— 1 male and 1 female ( MZSP). MATO GROSSO DO SUL [mislabeling?]: Porto Murtinho, XII.1929, Spitz col.— 1 female ( MZSP). MINAS GERAIS: Aiuruoca, VI.2009, P. H. Bernardo col.— 1 female ( CEMT; red form); Aiuruoca, Parque Estadual da Serra do Papagaio, 1,700m, X.1993, A. Machado col. – 1 male ( CEMT; red form); Camanducaia, Monte Verde, 13.XI.1970, J. Halik col.— 1 female ( MZSP); Conceição dos Ouros, I.2003, G. P. Almeida col.— 2 males ( CEMT); Fazenda dos Campos, 20.XII.1920, J. F. Zikán col.— 1 female ( FIOC); Ingaí, Reserva Biológica Unilavras/Boqueirão, II.2003, Fernando Z. Vaz-de-Mello col.— 1 female ( CEMT); Passa Quatro, without date, J. F. Zikán col.— 1 male ( FIOC); Passa Quatro, 1915, Jaeger col.— 1 female ( MNRJ; red form); Passa Quatro, 02.XII.1915, Zikán col.— 3 males ( MNRJ); Passa Quatro, 915 m., 14.XII.1922, J. F. Zikán col.— 1 male ( FIOC); Passa Quatro, Fazenda dos Campos, XI.1917, J. F. Zikán col.— 1 male and 1 female ( MNRJ); Passa Quatro, Faz. dos Campos, 17.XII.1918, J. F. Zikán col.— 1 male ( MNRJ); Passa Quatro, Faz. dos Campos, 01.XI.1920, J. F. Zikán col.— 1 female ( MNRJ); Passa Quatro, Faz. dos Campos, 09.XI.1920, J. F. Zikán col.— 1 male ( MNRJ; red form); Varginha, without date, Zoroastro col.— 1 female ( FIOC); Virgínia, 1,500 m., 08.X.1918, J. F. Zikán col.— 1 female ( FIOC); Virgínia, 1,500 m., 1920, J. Zikán col.— 1 female ( FIOC); Virgínia, 1,500 m., 06.I.1922, J. F. Zikán col.— 1 female ( FIOC); Virgínia, Fazenda dos Campos, 1,500 m., 25.X.1915, J. F. Zikán col.— 1 male ( FIOC; in a toad carcass); Virgínia, Faz. dos Campos, 1,500 m., 15.XII.1915, J. F. Zikán col.— 1 male ( FIOC); Virgínia, Faz. dos Campos, 1,500 m., 07-23.XI.1917, J. Zikán col. – 1 male ( MNRJ) and 1 female ( FIOC); Virgínia, Faz. dos Campos, 1,500 m., 25.I.1919, J. Zikán col.— 1 male ( MNRJ); Virgínia, Faz. dos Campos, 1,500 m., 22.X.1920, J. F. Zikán col.— 1 female ( FIOC); Virgínia, Faz. dos Campos, 1,500 m., 16.XI.1920, J. F. Zikán col.— 1 male ( DZUP; red form); 1 male ( FIOC; red form), 2 males and 1 female ( MNRJ), and 1 male ( MZSP; red form); Virgínia, Faz. dos Campos, 1,500 m, 25-30.XI.1920, J. F. Zikán col.— 2 males and 2 females ( FIOC);, Virgínia, Faz. dos Campos, 1,500 m., 28.XI.1921, J. F. Zikán col.— 1 male ( FIOC; red form); Virgínia, Faz. dos Campos, 1,500 m., 08.XII.1921, J. F. Zikán col.— 1 male ( CEMT; red form); Virgínia, Faz. dos Campos, 1,500 m., 08.XI.1926, J. F. Zikán col.— 1 female ( FIOC). PARANÁ: Balsa Nova, Serra de São Luiz de Purunã, Fazenda Monjolo, 1,116 m., 25º45'S 49º70'W, I.2007, P. Lowenberg Neto col.— 3 males and 1 female ( CEMT); Curitiba, without date and collector— 1 female ( MNHN); Curitiba, XII.1940, Claretiano col.— 1 male ( MZSP); Curitiba, Bariguí, II.1941, Claretiano col.— 1 female ( MZSP); Curitiba, XI.1963, R. R. Teixeira col.— 2 females ( DZUP); Curitiba, 900 m., 12.XII.1968, Dep. Zoo. col.— 1 male ( DZUP); Foz do Iguaçu, XI.1984, A. Soares col.— 1 male ( CEMT); Guarapuava, XI.1960, I. Schneider col.— 1 female ( MNRJ; red form); Marechal Cândido Rondon, Porto Artaza, without date and collector— 1 male ( MZSP); Matelândia, XI.1961, A. Maller col.— 1 male ( MZSP; red form); Matelândia, I.1962, A. Maller col.— 1 female ( MNRJ; red form) and 1 male and 1 female ( MZSP; red form); Ponta Grossa, XII.1942, without collector— 1 male and 1 female ( DZUP); Rio Negro, Tijuco Preto, 810m, 26º45'S 39º45'53”W, V.2012, Randy Marcolino col.— 1 male and 1 female ( CEMT). RIO DE JANEIRO: Itatiaia, 08.II.1991, Celso Jr. col.— 1 male ( CEMT; red form); Itatiaia, XI.1991, Celso Godinho col.— 2 females ( CEMT); Itatiaia, 08.XI.1991, Celso Jr. col.— 1 male ( CPFA); Itatiaia, II.1992, Celso Godinho col.— 1 male ( CEMT); Itatiaia, IV.1992, Celso Godinho col.— 2 females ( CEMT; one of the red form); Itatiaia, 02.XI.1992, Celso Jr. col.— 1 male and 2 females ( CPFA, all red form); Itatiaia, III.1993, Celso Godinho col.— 4 males ( CEMT); Itatiaia, III.1995, Celso Godinho Jr.— 1 female ( CEMT; red form); Itatiaia, 850 m., 18.III.1934, J. F. Zikán col.— 1 female ( FIOC; red form); Itatiaia, Parque Nacional do Itatiaia, Casa do Pesquisador, 810 m., 22- 25.II.2013, Mario Cupello col.— 1 male and 2 females ( MNRJ). RIO GRANDE DO SUL: no more data— 3 males and 2 females ( MNHN) and 2 females ( BMNH); Barão de Cotegipe, 31.XII.1965, F. Giacomel col.— 1 male ( DZUP); Bento Gonçalves, IX.2001, C. Arioli col.— 4 males ( CEMT); Caxias do Sul, XI.1926, without collector— 2 females ( MNRJ); Ijuí, 19.IX.2012, J. Lima col.— 1 male ( CEMT; red form); Porto Alegre, without date and collector— 1 male ( MNRJ; red form); Rolante, 29º27'43''S 56º09'12''W, 19-22.X.2010, G. Viegas col.— 3 males and 1 female ( CEMT; one male of the red form); Santo Augusto, X.1958, O. Roppa col.— 3 males and 1 female ( MNRJ; red form); Santo Augusto, X.1966, without collector— 2 males ( CPFA, red form); São Francisco de Paula, 29º29'16”S 50º10'60''W, 20.II-19.III.2002, R. Ott col.— 3 males and 3 females ( CEMT); São Francisco de Paula, 13-14.I.2008, L. Audino col.— 3 males and 3 females ( CEMT; one female of the red form); São Francisco de Paula, 10-13.XII.2009, Paschoal C. Grossi col.— 1 male and 1 female ( CEMT); Triunfo, 17,IX- 17.X.1993, without collector— 1 male and 1 female ( CEMT). SANTA CATARINA: no more data— 2 males and 1 female ( MNHN), 1 female ( MNRJ), 1 male and 2 females ( CPFA), and 2 males and 2 females ( BMNH); without date, Friedenreich col.— 1 male ( MNRJ); Anitápolis, I.1928, B. Pohl col.— 2 females ( MZSP); Anitápolis, 27º54'S 49º07'W, 28.I.2013, T. R. O. Simões col.— 1 male and 1 female ( CEMT); Antônio Carlos, 27º25'S 48º51'W, 19.XII.2012, J. A. Bogoni col.— 1 male and 1 female ( CEMT); Blumenau, XII.1924, Luederwaldt col.— 1 male ( MZSP); Bombinhas, ARIE Costeira de Zimbros, 27ºS 48ºW, 14.X.2011, C. C. Oliveira col.— 1 female ( CEMT); Campos Novos, 27º23'S 51º12'W, II.2010, M. I. M. Hernández & G. Viegas cols.— 2 males ( CEMT); Canoinhas, Pinhal, XII.1951, A. Maller col.— 2 females ( MNRJ); Corupá, X (without year), Anton Maller col.— 3 males ( MNRJ); Corupá, XI (without year), Anton Maller col.— 2 females ( MNRJ); Corupá (Hansa Humboldt), X.1942, collector illegible— 1 male ( MZSP); Corupá, III.1952, A. Maller col.— 2 males and 1 female ( MNRJ); Corupá, XI.1952, A. Maller col.— 1 male ( MNRJ); Corupá, III.1953, A. Maller col.— 1 female ( MNRJ); Corupá, XII.1996, without collector— 3 males and 1 female ( CPFA); Corupá, II.1954, A. Maller col.— 1 female ( DZUP); Corupá, II.1992 without collector— 4 males and 2 females ( CPFA); Florianópolis, 27º31'S 48º30'W, 09-10.I.2013, P. G. Silva col.—1 mal and 1 female ( CEMT); Florianópolis, 27º42'S 48º31'W, 01.II.2013, P. G. Silva col.— 1 male and 1 female ( CEMT); Florianópolis, Lagoa do Peri, 27º43'S 48º32'W, IX.2007, P. Condé & M. Hernández cols.— 1 male ( CEMT); Florianópolis, Lagoa do Peri, X.2007, P. Condé & M. Hernández cols.— 4 males and 1 female ( CEMT); Florianópolis, Lagoa do Peri, XI.2007, P. Condé & M. Hernández cols.— 1 male ( CEMT); Florianópolis, Lagoa do Peri, 27º49'S 45º92'W, I.2009, P. Condé & M. Hernández cols.— 1 female ( CEMT); Florianópolis, Vila Harmonia, XII.1914, Luederwaldt col.— 1 female ( MZSP); Governador Celso Ramos, 27º23'S 48º33'W, 27.I.2012, P. G. Silva col.— 1 male and 1 female ( CEMT); Governador Celso Ramos, 27º25'S 48º34'W, 24.I.2013, P. G. Silva col.— 1 male and 1 female ( CEMT); Ipumirim, 27º02'S 52º10'W, 25.XII.2012, J. A. Bogoni col.— 2 males ( CEMT); Itapema, 27º03'S 48º36'W, 17.I.2013, P. G. Silva col.— 1 male and 1 female ( CEMT); Joinville, without date, Dirings col.— 2 females ( MZSP); Joinville, 1919, Schmith col.— 2 males and 1 female ( MNRJ); Joinville, Rio Bracinho, without date, Dirings col.— 3 males and 2 females ( MZSP); Lages, without date and collector— 1 male ( MNHN; red form); Mafra, IV.1976, P. Arnaud col.— 3 males ( CPFA); Monte Castelo, 815m, 26º39'S 50º18'W, 24.XII.2012, A. L. Brandi col.— 1 female ( CEMT); Monte Castelo, 881m, 26º43'S 50º19'W, 07.I.2013, A. L. Brandi col.— 2 males and 1 female ( CEMT); Nova Veneza, 28º33'S 49º35'W, 20.III.2013, J. A. Bogoni col.— 1 male and 1 female ( CEMT); Porto Belo, APA do Araça, 27ºS 48ºW, 15.II.2011, M. Rocha col.— 1 male ( CEMT); Rio Vermelho, I.2003, without collector— 2 males ( CPFA); Rio Vermelho, II.1995, without collector— 1 male ( CPFA); Santo Amaro da Imperatriz, 27º44'S 48º48'W, XI.2009, C. B. Marcon & M. I. M. Hernández cols.— 3 males and 1 female ( CEMT); São Bento do Sul, without date, Dirings col.— 1 male and 1 female ( MZSP); São Bento do Sul, 1921, collector illegible— 1 male ( MNRJ); São Bento do Sul, I.1952, without collector— 1 male ( MNRJ); São Bento do Sul, IV.1949, Dirings col.— 1 male ( MZSP); São Bento do Sul, I.1950, Dirings col.— 1 male ( MZSP); São Bento do Sul, III.1952, without collector— 1 male ( MNRJ); São Bento do Sul, Rio Natal, III.1957, Dirings col.— 1 male and 1 female ( MZSP); São Bento do Sul, Rio Vermelho, without date, Dirings col.— 21 males and 16 females ( MZSP); São Bento do Sul, Rio Vermelho, XII.1948, Dirings col.— 2 males ( MZSP); São Bento do Sul, Rio Vermelho, III.1949, Dirings col.— 4 males ( MZSP); São Bento do Sul, Rio Vermelho, XI.1949, Dirings col.— 1 male ( MZSP); São Bento do Sul, Rio Vermelho, II.1957, A. Maller col.— 1 male ( MNRJ); São Bento do Sul, Rio Vermelho, I.1962, Dirings col.— 1 male ( MZSP); São Bento do Sul, Rio Vermelho, 850 m., IV.1974, Rank col.— 1 male ( DZUP); São Bento do Sul, Rio Vermelho, 850 m., V.1974, Rank col.— 2 females ( DZUP); São Bonifácio, 27º53'S 48º51'W, 15.XII.2012, J. A. Bogoni col.— 1 male and 1 female ( CEMT); São Domingos, 26º26'S 52º34'W, 28.XII.2011, A. Bugoni col.— 1 male and 1 female ( CEMT); São Pedro de Alcântara, 27º37'S 48º53'W, 27.II.2013, J. A. Bogoni col.— 1 female ( CEMT); Seara, Nova Teutônia (“Hansa”), without date, J. Clermont col.— 1 male and 2 females ( CPFA); Seara, Nova Teutônia (“Hansa Humboldt”), without date and collector— 5 males and 1 female ( MNHN); Seara, Nova Teutônia, I.1934, B. Pohl col.— 1 male and 1 female ( MZSP); Seara, Nova Teutônia, X.1934, B. Pohl col.— 1 male ( MZSP); Seara, Nova Teutônia, I.1935, B. Pohl col.— 1 male ( MZSP); Seara, Nova Teutônia, 31.X.1940, F. Plauman col.— 1 female ( MNRJ); Seara, Nova Teutônia, XII.1944, Dirings col.— 1 female ( MZSP); Seara, Nova Teutônia, VIII.1948, Dirings col.— 2 females ( MZSP); Seara, Nova Teutônia, III.1949, Dirings col.— 1 female ( MZSP); Seara, Nova Teutônia, 04.XI.1950, F.

Plauman col.— 1 female ( MNRJ); Seara, Nova Teutônia, X.1951, F. Plauman col.— 1 male and 1 female ( MNRJ); Seara, Nova Teutônia, XII.1964, F. Plaumann col.— 1 male ( MZSP); Seara, Nova Teutônia, X.1965, F. Plaumann col.— 1 male ( MZSP); Seara, Nova Teutônia, 27º11’S 52º23’W, II.1980, Fritz Plaumann col.— 1 male ( DZUP); Timbó, IV.1956, Dirings col.— 1 female ( MZSP); Timbó, X.1961, Dirings col.— 1 male ( MZSP). SÃO PAULO: no more data— 1 male and 1 female ( BMNH); Bananal, Serra da Bocaina, I.1937, D. Mendes col.— 1 female ( MNRJ); Barueri, 16-23.II.1955, K. Lenko col.— 2 males ( DZUP); Barueri, 28.II.1956, K. Lenko col.— 1 male ( DZUP); Barueri, 04.II.1968, K. Lenko col.— 1 female ( MZSP); Campos do Jordão, without date and collector— 1 female ( MZSP); Campos do Jordão, 20.XI.1952, P. Pereira col.— 1 female ( MZSP; red form); Campos do Jordão, XII.1955, J. Lane col.— 1 male ( MZSP); Campos do Jordão, 09.V.1964, H. Ebert col.— 1 female ( FIOC); Campos do Jordão, I.2004, G. P. Almeida Neto col.— 8 females ( CEMT); Campos do Jordão, 1,600 m., 25.IV.1955, E. C. Freire col.— 1 male ( MNRJ; red form); Campos do Jordão, 1,600 m., 29.IV.1955, E. C. Freire col.— 1 female ( MNRJ); Campos do Jordão, 1,700 m., 24.I.1966, H. Ebert col.— 1 male ( FIOC); Cubatão, XI.2004, A. A. da Silva col.— 1 male ( MZSP); Cubatão, 14-19.II.2005, A. A. da Silva col.— 2 males ( MZSP); Estação Biológica da Boracéia, 800- 900m., 28-30.IV.1997, F. Génier & S. Ide cols.— 1 female ( MZSP); Jundiaí, without date and collector— 1 male ( MZSP); Leme, 1930, Riditer col.— 1 female ( MZSP); Mogi das Cruzes, 1932, Elisa & Vianna cols.— 1 female ( MZSP); Parque Estadual da Serra do Mar, Núcleo Caraguatatuba, 23º35'27''S 45º25'19''W, 13.I.2012, E. Bovy col.— 1 male ( CEMT); Parque Estadual da Serra do Mar, Núcleo Picinguaba, 23º20'11''S 44º50'03''W, 14.I.2012, E. Bovy col.— 4 males and 13 females ( CEMT); Parque Estadual da Serra do Mar, Núcleo Santa Virgínia, Sede Vargem Grande, 23º26'33''S 45º14'19''W, 17.I.2012, E. Bovy col.— 17 males and 15 females ( CEMT); Parque Estadual Xixová-Japuí, 28.X-17.XI.1999, “Grupo” col.— 1 female ( CEMT); Parque Estadual Xixová-Japuí, 21.II.2000, “Grupo” col.— 1 male; Pindamonhangaba, Eugênio Lefevre, 21.II.1963, Exp. Dep. Zoo. col.— 1 male ( MZSP); Ribeirão Preto, 1906, without collector— 1 female ( MZSP); Rio Grande da Serra, II.1963, without collector— 1 male ( MZSP); Salesópolis, Estação Biológica da Boracéia, 21.I.1966, Reichardt col.— 1 female ( MZSP); Salesópolis, Est. Biol. Boracéia, 01-28.XI.2001, S. A. Casari & I. B. Francini cols.— 1 male ( MZSP); São Bernardo do Campo, X.1933, without collector— 1 male ( MNRJ); São Miguel Arcanjo, Parque Estadual Carlos Botelho, 762 m., 24º04'01''S 47º59'40''W, 25.XI.2011, E. Bovy col.— 1 male ( CEMT); São Paulo, without date and collector— 1 male ( MZSP); São Paulo, Ipiranga, 1901, Lima col.— 1 female ( MZSP); São Paulo, Ipiranga, 1902, without collector— 1 male ( MZSP); São Paulo, Jabaquara, 08.I.1934, without collector— 1 male ( MNRJ); São Paulo, Santana, 04.XII.1965, J. Halik col.— 1 male and 1 female ( MZSP); São Paulo, Santo Amaro, 1933, without collector— 1 male ( MNRJ); Serra da Bocaina, IV.1924, Luederwaldt & R. Spitz cols.— 2 males and 1 female ( MZSP); Serra da Bocaina, 27.X.1963, H. S. Lopes col.— 1 female ( FIOC); Serra da Cantareira, IV.1931, R. Spitz col.— 1 female ( MZSP); Serra da Cantareira, I.1966, J. Halik col.— 1 male and 1 female ( MZSP); Serra do Japi, 1,050m., 23º14'S 46º56'W, 1998, M. I. M. Hernández col.— 1 male ( CEMT).

Figures

FIGURE 5. Holotype of Coprophanaeus machadoi. 5 a) Dorsal view. 5 b) Ventral view. 5 c) Labels.

FIGURE 15. Holotype of Coprophanaeus saphirinus (Sturm, 1826): 15 a) Dorsal view. 15 b) Ventral view. 15 c) Frontal view. 15 d) Lateral view. 15 e) Labels. 15 f) Sturm’s (1826) illustration of the holotype. 16: Holotype of Phanaeus chabrillaci Thomson, 1857: 16 a) Labels. 16 b) Dorsal view. 16 c) Ventral view. 16 d) Lateral view.

FIGURES 6 – 11. Male head and pronotum. 6 – 9) Coprophanaeus machadoi. 10 – 11) C. saphirinus. White arrow indicates the midlongitudinal prolongation of the anterior margin of pronotum, which is triangular and flat in C. saphirinus and long, filiform, and sometimes raised in a sharp ridge in C. machadoi.

FIGURE 12 – 13. Aedeagi. 12) Coprophanaeus machadoi: 12 a) Dorsal view. 12 b) Lateral view. 13) C. saphirinus: 13 a) Dorsal view. 13 b) Lateral view. Arrow indicates the posterior flap of the apical process, which is just slightly elevated in C. machadoi and is strongly elevated and curved in C. saphirinus.

FIGURE 14. Distribution map of Coprophanaeus machadoi and C. saphirinus. Coprophanaeus machadoi is distributed in eastern Minas Gerais, Espírito Santo, and Rio de Janeiro, while C. saphirinus is present from the Serra da Mantiqueira, in the border of Minas Gerais, Rio de Janeiro and São Paulo, to southern Brazil, Paraguay and Misiones (Argentina). Both species are restricted to the Brazilian Atlantic Forest in Parana Dominion of the Chacoan subregion.

FIGURES 17 – 23. Color variation in Coprophanaeus saphirinus. 17 – 18) Dark blue female and male with purple reflections. 19) Blue male with greenish reflections. 20) Predominantly green female with bluish reflections. 21) Red male with greenish and yellowish reflections. 22) Predominantly red female. 23) Dark red male.