taxonID	type	description	language	source
03B6F8059638FFADB93CF9C2191962FA.taxon	diagnosis	Diagnosis of the Phanaeus chalcomelas species group Anteromedial margin of clypeus bidentate (Fig. 1 A). Head of females with transverse carina, straight (Fig. 1 A, D) or weakly trituberculate (in P. chalcomelas [Fig. 19 G – H]). Pronotum bearing contrasted black elevations, variously and irregularly shaped and frequently partly anastomosed on disc regions (Fig. 1 F), more abundant and anastomosed in females than males. Pronotal disc of major males triangularly flattened in dorsal view (Fig. 1 F), with a pair of anterior tubercles (Figs 8 D, 16 – 17). Pronotum of females with three (Fig. 1 D) or four (in P. chalcomelas [Fig. 19 G – H]) tubercles on anteromedial portion, sometimes poorly distinguishable from the irregular black elevations if not by their regular position. Posteromedial pronotal fossae present or absent (in P. cambeforti). Elytral striae punctate, basally impressed as distinct fossae (Fig. 1 H). Front tibiae tridentate, with faint, almost imperceptible fourth tooth. Dorsum dull, colored black, brown, green or blue, sometimes with red areas. Metallic sheen restricted to the following areas: around the eyes, pronotal margins, anterior margins of elytra, ventral portion of femora, hypomeron, and the pair of anteromedial pronotal tubercles in males. Widely distributed in the Amazon and along the Eastern slopes of Andes from Bolivia to Venezuela (Figs 20 – 21).	en	Bordin, Bruna Ribeiro, Arias-Buriticá, Jorge, Maldaner, María Eduarda, Vaz-De-Mello, Fernando Z. (2026): Taxonomic revision of the Phanaeus chalcomelas species group and a description of the new Phanaeus achilles species group (Coleoptera: Scarabaeidae: Scarabaeinae: Phanaeini). European Journal of Taxonomy 1080: 1-65, DOI: 10.5852/ejt.2026.1080.3310, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3310/14554
03B6F8059634FFA9B943F88F1DB16397.taxon	description	Figs 1 A, D, 2 G – H, 3 C, 4 A, 5, 13 A, 14 A, D, G, J, 16 C, 17 C, 18 C, 19 C	en	Bordin, Bruna Ribeiro, Arias-Buriticá, Jorge, Maldaner, María Eduarda, Vaz-De-Mello, Fernando Z. (2026): Taxonomic revision of the Phanaeus chalcomelas species group and a description of the new Phanaeus achilles species group (Coleoptera: Scarabaeidae: Scarabaeinae: Phanaeini). European Journal of Taxonomy 1080: 1-65, DOI: 10.5852/ejt.2026.1080.3310, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3310/14554
03B6F8059634FFA9B943F88F1DB16397.taxon	diagnosis	Diagnosis Dorsal color ranging from olive green to reddish brown. Ventral color dark green or black with green sheen (Fig. 5 C). Pronotal disc with faint punctation (× 40 magnification), absent in some specimens (Fig. 5 A – B, D). Posteromedial fossae of pronotum absent or shallow in a few large specimens, mainly in females (Fig. 4 A). Pronotum, in frontal view, with the transverse carina just posterior and adjacent to the anterior medial margin appearing straight (Figs 3 C, 16 C, 19 C). Pronotum of males with few black elevations, these rarely anastomosed (Fig. 5 A). Width of the pronotal disc in major males rarely exceeds the greatest width of elytra. Tip of anteromedial prominence of the metaventrite flat in lateral view (Fig. 2 H). Length 11 – 18.7 mm. Pygidium mostly brown (Fig. 18 C), rarely green. Endophallites: ME in dorsal view, elongate and blade-like, with a broad, rounded basal portion tapering gradually into a narrow, curved apex; left margin slightly concave, right margin nearly straight; left margin with a small fold near basal portion, not very pronounced compared to the other species of the group with a similar structure (Fig. 14 D). AE with a straight base, not sharp; right margin (dorsal view) almost straight, with a small tooth-like projection in the middle, more visible in lateral view (Fig. 14 G). SRP circular, as long as wide (Fig. 14 J).	en	Bordin, Bruna Ribeiro, Arias-Buriticá, Jorge, Maldaner, María Eduarda, Vaz-De-Mello, Fernando Z. (2026): Taxonomic revision of the Phanaeus chalcomelas species group and a description of the new Phanaeus achilles species group (Coleoptera: Scarabaeidae: Scarabaeinae: Phanaeini). European Journal of Taxonomy 1080: 1-65, DOI: 10.5852/ejt.2026.1080.3310, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3310/14554
03B6F8059634FFA9B943F88F1DB16397.taxon	materials_examined	Type material Holotype FRENCH GUIANA • ♂; “ Saül; Guyane française, IV- 1977, M. DURANTON leg. ”; MNHN (Arnaud 1982 b: 122: fig. 6. Paratypes FRENCH GUIANA • 1 ♂, 1 ♀ (examined from photographs provided in Arnaud 2002: 87). Non-type material examined (193 ♂♂, 203 ♀♀) BOLIVIA – La Paz • 3 ♂♂, 2 ♀♀; Iturralde, Rio Nuanu; 12 ° 33 ′ 23 ″ S, 67 ° 43 ′ 44 ″ W; 21 – 23 Sep. 2002; W. Aliaga leg.; MHNNKM • 2 ♂♂, 4 ♀♀; same data as for preceding; 12 ° 33 ′ 9 ″ S, 67 ° 42 ′ 32 ″ W; 22 – 24 Sep. 2002; MHNNKM. – Pando • 1 ♂, 2 ♀♀; Nicolás Suárez, Rutina; 25 May 2009; O. Terrán et al. leg.; MHNNKM • 1 ♂; same data as for preceding; Sep. 1998; MHNNKM. BRAZIL – Acre • 3 ♂♂, 5 ♀♀; Bujari, FES Antimary; 09 º 25 ′ 41 ″ S, 68 º 17 ′ 7 ″ W; 15 Mar. 2017; B. S. Bitencourt leg.; pitfall human dung; CEMT • 1 ♀; Mâncio Lima; 7 ° 28 ′ 58 ″ S, 72 ° 54 ′ 11 ″ W; 5 Dec. 2018; Luiz H. M. B. leg.; CEMT • 1 ♂; Mâncio Lima, Parque Nacional da Serra do Divisor; 7 ° 24 ′ 58 ″ S, 73 ° 37 ′ 56 ″ W; 10 – 14 Dec. 2019; R. A. Azevedo leg.; flight interception trap; CEMT • 1 ♂, 1 ♀; same data as for preceding; 7 ° 26 ′ 46 ″ S, 73 ° 39 ′ 28 ″ W; 7 – 11 Dec. 2019; CEMT • 1 ♂; same data as for preceding; 7 ° 26 ′ 46 ″ S, 73 ° 39 ′ 27 ″ W; pitfall human dung; CEMT • 2 ♂♂, 3 ♀♀; same data as for preceding; 7 ° 27 ′ 04 ″ S, 73 ° 38 ′ 57 ″ W; 6 – 14 Dec. 2019; flight interception trap; CEMT • 1 ♂; same data as for preceding; pitfall human dung; CEMT • 1 ♀; same data as for preceding; 7 ° 28 ′ 59 ″ S, 72 ° 54 ′ 11 ″ W; CEMT • 2 ♂♂, 3 ♀♀; Rio Branco, Fazenda Catuaba; Feb. 1997; F. Z. Vaz-de-Mello leg.; pitfall human dung; CEMT • 1 ♂, 1 ♀; Senador Guiomard; 10 ° 04 ′ S, 67 ° 37 ′ W; 14 Apr. 2017; B. S. Bitencourt; pitfall human dung; CEMT • 1 ♂, same data as for preceding; 20 Apr. 2018; CEMT. – Amapá • 1 ♂, 2 ♀♀; Serra do Navio, Cava Urucum-Amapari; 0 ° 53 ′ 06 ″ N, 51 ° 52 ′ 53 ″ W; Dec. 2000; R. Ribon leg.; CEMT. – Amazonas • 4 ♂♂, 3 ♀♀; Barcelos, Parque Nacional do Jaú; 2 ° 00 ′ 24 ″ S, 61 ° 33 ′ 1 ″ W; 10 – 14 Nov. 2021; R. A. Azevedo leg.; flight interception trap; CEMT • 2 ♀♀; same data as for preceding; 2 ° 00 ′ 26 ″ S, 61 ° 32 ′ 49 ″ W; 14 – 16 Nov. 2021; pitfall human dung; CEMT • 1 ♂, same data as for preceding; 2 ° 00 ′ 25 ″ S, 61 ° 32 ′ 55 ″ W; CEMT • 2 ♀♀; same data as for preceding; 2 ° 00 ′ 23 ″ S, 61 ° 32 ′ 01 ″ W; CEMT • 2 ♂♂, 4 ♀♀; same data as for preceding; 2 ° 00 ′ 22 ″ S, 61 ° 32 ′ 03 ″ W; CEMT • 2 ♀♀; same data as for preceding; 2 ° 00 ′ 21 ″ S, 61 ° 32 ′ 06 ″ W; CEMT • 1 ♂, same data as for preceding; 2 ° 00 ′ 20 ″ S, 61 ° 33 ′ 10 ″ W; CEMT • 2 ♂♂; same data as for preceding; 2 ° 00 ′ 19 ″ S, 61 ° 33 ′ 13 ″ W; CEMT • 2 ♂♂, 1 ♀; same data as for preceding; 2 ° 00 ′ 18 ″ S, 61 ° 33 ′ 16 ″ W; CEMT • 2 ♂♂, 2 ♀♀; same data as for preceding; 1 ° 50 ′ 54 ″ S, 61 ° 34 ′ 01 ″ W; CEMT • 3 ♀♀; same data as for preceding; 1 ° 50 ′ 35 ″ S, 61 ° 34 ′ 07 ″ W; flight interception trap; CEMT • 1 ♀; same data as for preceding; 2 ° 02 ′ 55 ″ S, 61 ° 32 ′ 52 ″ W; 13 – 17 Nov. 2021; CEMT • 1 ♀; Careiro, BR- 319; 4 ° 07 ′ 58 ″ S, 60 ° 44 ′ 09 ″ W; 2 Dec. 2015; D. Pires leg.; pitfall human dung; CEMT • 1 ♀; same data as for preceding; 4 ° 08 ′ 28 ″ S, 60 ° 43 ′ 57 ″ W; CEMT • 1 ♀; same data as for preceding; 4 ° 08 ′ 58 ″ S, 60 ° 43 ′ 44 ″ W; CEMT • 1 ♀; same data as for preceding; 4 ° 09 ′ 10 ″ S, 60 ° 43 ′ 08 ″ W; 3 Dec. 2015; CEMT • 1 ♂, 1 ♀; same data as for preceding; 4 ° 09 ′ 28 ″ S, 60 ° 43 ′ 32 ″ W; 2 Dec. 2015; CEMT • 1 ♂; Presidente Figueiredo; Oct. – Nov. 1996; E. Anderson leg.; CEMT • 1 ♂, 1 ♀; Silves, Fazenda Saracá; 2 ° 50 ′ 45 ″ S, 58 ° 35 ′ 07 ″ W; 5 – 12 May 2019; R. S. Moura leg.; CEMT. – Rondônia • 1 ♂; Cacaulândia, Fazenda Rancho Grande; Nov. 1996; K. Vulinec leg.; CEMT • 1 ♀; same data as for preceding; Feb. 1997; V. Z. Vaz-de-Mello leg.; CEMT • 1 ♀; same data as for preceding; Feb. 1998; CEMT • 1 ♂; Guajará-Mirim, Sítio Coletti; 10 ° 19 ′ 47 ″ S, 64 ° 45 ′ 12 ″ W; elev. 170 m; 28 Jan. 2010; pitfall human dung; F. Coletti leg.; CEMT • 1 ♀; same data as for preceding; carcass; CEMT • 1 ♀; Porto Velho, Abunã; 9 ° 38 ′ 09 ″ S, 65 ° 32 ′ 47 ″ W; 13 – 15 Jan. 2011; J. C. F. Falcão leg.; pitfall human dung; CEMT • 3 ♂♂, 2 ♀♀; Porto Velho, ESEC Cuniã; 8 ° 04 ′ 08 ″ S, 63 ° 28 ′ 06 ″ W; 11 – 15 Aug. 2012; M. A. P. A. Silveira leg.; pitfall human dung; CEMT • 4 ♀♀; same data as for preceding; 8 ° 04 ′ 11 ″ S, 63 ° 28 ′ 34 ″ W; elev. 83 m; 10 – 12 Nov. 2013; CEMT. COLOMBIA – Amazonas • 1 ♀; Leticia, Estación Biológica El Zafire; 4 ° 00 ′ 21 ″ S, 69 ° 53 ′ 55 ″ W; elev. 150 m; 2 – 4 Dec. 2007; L. Franco and S. Flórez; pitfall human dung; IAvH-E. – Caquetá • 1 ♂, 2 ♀♀; Puerto Solano, PNN Serranía de Chiribiquete, Puerto Abeja; 0 ° 04 ′ 27 ″ N, 72 ° 27 ′ 0.5 ″ W; elev. 200 m; Jul. 1999; M. Álvares and G. Mejía; IAvH-E • 1 ♂; Puerto Solano, PNN Serrania de Chiribiquete, Río Cunãre; 0 ° 20 ′ N, 72 ° 37 ′ 50 ″ W; elev. 250 m; 2 – 4 Nov. 2000; M. Ospina and E. González leg.; flight interception trap; UPTC-CE • 2 ♀♀; same data as for preceding; 0 ° 31 ′ 44 ″ N, 72 ° 37 ′ 50 ″ W; 22 – 24 Nov. 2000; UPTC-CE • 3 ♀♀; same data as for preceding; 0 ° 30 ′ N, 72 ° 37 ′ 50 ″ W; 2 – 4 Nov. 2000; pitfall human dung; IAvH-E • 3 ♀♀; same data as for preceding; 0 ° 32 ′ 4 ″ N, 72 ° 37 ′ 57.5 ″ W; 19 – 21 Nov. 2000; IAvH-E • 1 ♀; same data as for preceding; 15 Nov. 2000; flight interception trap; IAvH-E • 1 ♂; Puerto Solano, PNN Serranía de Chiribiquete, Río Cuñare - Amú; 0 ° 12 ′ 34.5 ″ N, 72 ° 24 ′ 55.1 ″ W; elev. 250 m; 14 Feb. 2001; E. González and M. Ospina leg.; flight interception trap; IAvH-E • 1 ♂; same data as for preceding; 15 – 17 Feb. 2001; pitfall human dung; IAvH-E • 1 ♀; same data as for preceding; 0 ° 12 ′ 42.8 ″ N, 72 ° 25 ′ 3.2 ″ W; 21 Feb. 2001; IAvH-E • 1 ♂; same data as for preceding; 0 ° 12 ′ 47.8 ″ N, 72 ° 25 ′ 25.4 ″ W; 1 – 3 Mar. 2001; IAvH-E • 1 ♀; same data as for preceding; 0 ° 13 ′ 3.4 ″ N, 72 ° 25 ′ 22.7 ″ W; 7 – 9 Mar. 2001; IAvH-E • 2 ♀♀; Puerto Solano, PNN Serranía de Chiribiquete, Río Mesay; 0 ° 15 ′ 38 ″ N, 72 ° 55 ′ 24 ″ W; elev. 250 m; 24 Jan. 2000; F. Quevedo leg.; pitfall human dung; IAvH-E • 1 ♀; same data as for preceding; 0 ° 14 ′ 24 ″ N, 72 ° 56 ′ 2 ″ W; 25 Jan. 2000; IAvH-E • 1 ♂, 1 ♀; same data as for preceding; 0 ° 14 ′ 54 ″ N, 72 ° 56 ′ 5 ″ W; 28 Jan. 2000; IAvH-E • 1 ♂; same data as for preceding; 0 ° 14 ′ 32 ″ N, 72 ° 56 ′ 15 ″ W; 4 Feb. 2000; IAvH-E • 1 ♂, 1 ♀; Puerto Solano, PNN Serranía de Chiribiquete, Río Sararamano; 0 ° 10 ′ 47 ″ N, 72 ° 37 ′ 24 ″ W; elev. 250 m; 4 Apr. 2000; E. González leg.; flight interception trap; IAvH-E • 1 ♂, 1 ♀; same data as for preceding; 0 ° 09 ′ 26 ″ N, 72 ° 37 ′ 45 ″ W; 11 Apr. 2000; pitfall human dung; IAvH-E • 1 ♂; same data as for preceding; 0 ° 08 ′ 26 ″ N, 72 ° 37 ′ 45 ″ W; 11 Nov. 2000; flight interception trap; UPTC-CE. – Cundinamarca • 1 ♀; Medina, alto del Río Gazaunta; 4 ° 35 ′ N, 73 ° 23 ′ W; elev. 550 m; Mar. 1997; F. Escobar leg.; pitfall trap; UPTC-CE • 1 ♂; same data as for preceding; IAvH-E • 4 ♀♀; Medina, Miralindo; 4 ° 35 ′ 33 ″ N, 73 ° 27 ′ 17 ″ W; elev. 550 m; Feb. – Mar. 1997; F. Escobar leg.; pitfall human dung; IAvH-E. – Guaviare • 1 ♀; Cerro Cocuy and Cerro Moyano; 2 ° 10 ′ 02 ″ N, 71 ° 10 ′ 54 ″ W; elev. 200 m; 30 – 31 Jan. 1996; F. Escobar leg.; pitfall human dung; CEMT • 1 ♂; same data as for preceding; IAvH-E • 2 ♂♂, 4 ♀♀; same data as for preceding; 2 ° 10 ′ 35 ″ N, 71 ° 15 ′ 58 ″ W; elev. 300 m; Feb. 1996; IAvH-E. – Meta • 1 ♀; Villavicencio, El Carmen; 4 ° 08 ′ 38 ″ N, 73 ° 39 ′ 57 ″ W; elev. 748 m; 23 Apr. 2019; D. C. Martínez leg.; carcass; CEMT • 1 ♂, 1 ♀; Villavicencio, Bosque Bavaria; 4 ° 09 ′ S, 73 ° 38 ′ W; elev. 550 m; J. Noriega leg.; pitfall human dung; CEMT • 3 ♂♂; Villavicencio; 29 Feb. – 5 Mar. 1972; S. Peck and J. Peck leg.; TAMU • 1 ♀; 30 km Villavicencio; 1 – 4 Mar. 1972; S. Peck and J. Peck leg.; dung; TAMU • 1 ♀; 16 km of Villavicencio; 2 – 4 Mar. 1972; S. Peck leg.; dung; TAMU • 1 ♀; 500 m of Villavicencio; 29 Feb. – 5 Mar. 1972; Peck and J. Peck leg.; bait traps; TAMU. – Putumayo • 1 ♀; Puerto Leguizamo; 00 ° 04 ′ 20 ″ S, 74 ° 59 ′ 19 ″ W; 17 Feb. 2008; L. Jiménez leg.; CCBUMAG. ECUADOR – Orellana • 1 ♀; Francisco de Orellana; 4 May 2000; A. Dávalos leg.; pitfall carcass; CEMT. – Napo • 1 ♂, 2 ♀♀; Estación Científica Yasuni Puce, Río Tiputini; 0 ° 40 ′ S, 76 ° 24 ′ W; Jul. 1999; A. Trischechkin leg.; flight interception trap; CEMT. – Zamora-Chinchipe • 120 ♂♂, 88 ♀♀; Padmi, Estación Experimental El Padmi UNL, BN Ladera; 3 ° 43 ′ 58 ″ S, 78 ° 37 ′ 09 ″ W; elev. 897 – 970 m; Dec. 2021 – Jan. 2022; J. Jiménez leg.; pitfall human dung; LOUNAZ • 2 ♂♂, 1 ♀; same data as for preceding; CEMT. FRENCH GUIANA – Cayenne • 1 ♀; Régina, Réserve Naturelle Nationale des Nouragues; 1996; CEMT • 1 ♀; same data as for preceding; 10 Apr. 1997; CEMT • 1 ♂; same data as for preceding; 31 Mar. 2002; F. Feer leg.; CEMT • 1 ♀; same data as for preceding; 21 Mar. 2003; CEMT. – Saint- Laurent-du-Maroni • 1 ♀; Bas Maroni; TAMU. PERU – Junín • 1 ♂, 1 ♀; Satipo, Río Venado; elev. 1137 m; May 2005; A. Santibáñez leg.; TAMU. – Lima • 1 ♀; 11 Oct. 1970 (data error); CEMT. – Loreto • 1 ♂, 1 ♀; Río Pucacuro; 2 ° 05 ′ 55 ″ S, 75 ° 40 ′ 39 ″ W; elev. 208 m; 26 Nov. 2007; C. Moreno leg.; pitfall human dung; CEMT. – Madre de Dios • 1 ♂, 1 ♀; CICRA Field Station, 12 ° 33 ′ 07 ″ S, 70 ° 06 ′ 34 ″ W; elev. 295 m; 7 – 9 Jun. 2011; Chaboo team leg.; flight interception trap; MUSM • 2 ♂♂, 1 ♀; Pantiacolla Lodge, Alto Madre de Dios River; 12 ° 39 ′ 10 ″ S, 71 ° 15 ′ 28 ″ W; elev. 500 m; 23 – 26 Oct. 2000; R. Brooks leg.; flight interception trap; CEMT • 1 ♂, 2 ♀♀; PNN Manu, Estación Biológica Pantiacolla; 12 ° 38 ′ 48 ″ S, 71 ° 14 ′ 36 ″ W; 2010; S. Pérez leg.; UPTC-CE. – San Martín • 2 ♂♂, 2 ♀♀; Juanjuí; 7 Aug. 2005; P. Melendes leg.; TAMU. – Ucayali • 1 ♀; Padre Abad, Boquerón del Padre Abad; elev. 500 – 550 m; 16 Sep. 1962; J. Schunke L. leg.; carcass; TAMU. Type locality French Guiana, Saül.	en	Bordin, Bruna Ribeiro, Arias-Buriticá, Jorge, Maldaner, María Eduarda, Vaz-De-Mello, Fernando Z. (2026): Taxonomic revision of the Phanaeus chalcomelas species group and a description of the new Phanaeus achilles species group (Coleoptera: Scarabaeidae: Scarabaeinae: Phanaeini). European Journal of Taxonomy 1080: 1-65, DOI: 10.5852/ejt.2026.1080.3310, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3310/14554
03B6F8059634FFA9B943F88F1DB16397.taxon	distribution	Distribution Widely distributed in the Amazon, from the slopes of the Colombian Andes southwards to Bolivia and northeast wards all the way to French Guiana (Fig. 20) between 80 to 438 m a. s. l.	en	Bordin, Bruna Ribeiro, Arias-Buriticá, Jorge, Maldaner, María Eduarda, Vaz-De-Mello, Fernando Z. (2026): Taxonomic revision of the Phanaeus chalcomelas species group and a description of the new Phanaeus achilles species group (Coleoptera: Scarabaeidae: Scarabaeinae: Phanaeini). European Journal of Taxonomy 1080: 1-65, DOI: 10.5852/ejt.2026.1080.3310, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3310/14554
03B6F8059634FFA9B943F88F1DB16397.taxon	discussion	Taxonomic discussion The smallest species within the P. chalcomelas species group, specifically the males. Major males are very rare and seem never to show strongly developed secondary sex characteristics as males of the other species in the group. We observed that the width of the pronotal disc in males exceeded the width of the elytra only in a population collected in Ecuador (Estación Experimental El Padmi Universidad Nacional de Loja, Loja) (Fig. 5 D).	en	Bordin, Bruna Ribeiro, Arias-Buriticá, Jorge, Maldaner, María Eduarda, Vaz-De-Mello, Fernando Z. (2026): Taxonomic revision of the Phanaeus chalcomelas species group and a description of the new Phanaeus achilles species group (Coleoptera: Scarabaeidae: Scarabaeinae: Phanaeini). European Journal of Taxonomy 1080: 1-65, DOI: 10.5852/ejt.2026.1080.3310, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3310/14554
03B6F8059632FFB2B938FE861F616783.taxon	description	Figs 1 F, 2 A, 6 – 7, 16 G – H, 17 G – H, 18 G – H, 19 G – H, 28 A	en	Bordin, Bruna Ribeiro, Arias-Buriticá, Jorge, Maldaner, María Eduarda, Vaz-De-Mello, Fernando Z. (2026): Taxonomic revision of the Phanaeus chalcomelas species group and a description of the new Phanaeus achilles species group (Coleoptera: Scarabaeidae: Scarabaeinae: Phanaeini). European Journal of Taxonomy 1080: 1-65, DOI: 10.5852/ejt.2026.1080.3310, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3310/14554
03B6F8059632FFB2B938FE861F616783.taxon	diagnosis	Diagnosis Dorsum olive green, olive brown or reddish brown (Fig. 6). Ventral color dark green. Anteromedial prominence of metaventrite with regular callosity forming arrowhead pattern (Fig. 2 A). Major males with cephalic horn abruptly narrowed basally to mid length (Fig. 16 G – H). Pronotum of females with a shallow anteromedial concavity surrounded by four tubercles, anterior tubercle large and distinct, the other three smaller (middle posterior tubercle always posterior to lateral ones) and may blend with the black elevations (Figs 6 B, D, 19 G – H). Aedeagus: parameres gradually narrowing apically, exhibiting a conical shape (Fig. 7 A). Phallobase without sulcus or incisions (Fig. 7 B). Endophallus with six endophallites (Fig. 7 C – E). Endophallites: A + SA joined by a basal membrane (Fig. 7 C – E). Axial endophallite V-shaped, with one branch apically expanded and rounded, and the other narrow and unexpanded; subaxial endophallite broad. FLP narrow and curved, like an arch (Fig. 7 C). AE with sharp base and straight apical margin (Fig. 7 H). SRP vertically elongated, much larger than other species of P. chalcomelas group (Fig. 7 I). ME bears three main projections, roughly resembling a Y-shape with asymmetrical and smoothly angled arms (dorsal view); small, slightly more than half a millimeter (Fig. 7 F – G).	en	Bordin, Bruna Ribeiro, Arias-Buriticá, Jorge, Maldaner, María Eduarda, Vaz-De-Mello, Fernando Z. (2026): Taxonomic revision of the Phanaeus chalcomelas species group and a description of the new Phanaeus achilles species group (Coleoptera: Scarabaeidae: Scarabaeinae: Phanaeini). European Journal of Taxonomy 1080: 1-65, DOI: 10.5852/ejt.2026.1080.3310, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3310/14554
03B6F8059632FFB2B938FE861F616783.taxon	materials_examined	Type material Holotype of Phanaeus chalcomelas (studied from photographs) BRAZIL • ♀; Perty, 1830: 40; ZSM. The Amazon rainforest, somewhere visited by Spix and Martius along the Amazon River, the Japura, the lower Madeira, or the Rio Negro (see Papavero 1971 for a map with these places indicated) (see discussion below) Paratypes of Phanaeus chalcomelas grossii BOLIVIA – Chapare • 1 ♂, 1 ♀; (examined from photographs provided in Arnaud 2002: 84). – Cochabamba • 1 ♂; “ 67.5 km E. Villa Tunari, Est. Biol. Valle Sajta, Univ. San Simon 300 m, 17 ° 06 ′ 19 ″ S 64 ° 46 ′ 57 ″ W; 7 – 9. II. 1999, F. Génier, lowland rain for., ex. dung tp. 3, 99 - 045 ”; MHNNKM. Non-type material examined (447 ♂♂, 568 ♀♀) BOLIVIA – Cochabamba • 1 ♂; PN Carrasco, Río Sajta; Feb. 1993; L. Peña leg.; CEMT • 1 ♀; same data as for preceding; Feb. 1994; CEMT • 1 ♂, 1 ♀; PN Carrasco, Valle Sajta; 17 ° 06 ′ 19 ″ S, 64 ° 46 ′ 57 ″ W; elev. 300 m; 9 – 13 Feb. 1994; F. Genier leg.; MHNNKM • 3 ♂♂, 3 ♀♀; Palmar; 26 Dec. 2009; T. Vidaurre leg.; MHNNKM • 1 ♂, 2 ♀♀; Chapare, Villa Tunari; 26 – 29 Oct. 2004; J. Ledezma and I. García leg.; MHNNKM • 2 ♂♂; same data as for preceding; Feb. 1984; P. Arnaud leg.; CEMT. – La Paz • 21 ♂♂, 12 ♀♀; Iturralde, Río Nuanu; 21 – 23 Sep. 2002; W. Aliaga leg.; MHNNKM. – Santa Cruz • 1 ♂; Andrés Ibáñez, Espejillos; 27 Apr. 2007; M. Amaya leg.; MHNNKM • 3 ♂♂, 1 ♀; Andrés Ibáñez, Jardín Botánico; 17 ° 46 ′ S, 63 ° 03 ′ W; elev. 398 m; 30 Apr. 2006; T. Vidaurre et al. leg.; MHNNKM • 14 ♂♂, 9 ♀♀; same data as for preceding; 16 – 19 Jul. 2006; MHNNKM • 3 ♂♂, 3 ♀♀; Santa Cruz de la Sierra, Jardín Botánico; 17 ° 45 ′ 51 ″ S, 63 ° 39 ′ 30 ″ W; 10 Nov. 2006; Scarabnet exped. leg.; CEMT • 1 ♂; same data as for preceding; 17 ° 47 ′ 02 ″ S, 63 ° 03 ′ 47 ″ W; elev. 400 m; 7 – 8 Nov. 2006; W. D. Edmonds leg.; pitfall human dung; TAMU • 2 ♀♀; Caballero, Karawasi; 22 Nov. 1998; MHNNKM • 2 ♂♂, 2 ♀♀; Santa Cruz de la Sierra, 4 – 6 km SSE of Buena Vista, F & F Hotel; elev. 420 – 450 m; 2 – 12 Feb. 2000; J. E. Wappes leg.; pitfall dung / carrion; TAMU • 1 ♂, 2 ♀♀; Ichilo, Buena Vista; elev. 400 m; F. Steinbach leg.; TAMU • 10 ♂♂, 7 ♀♀; Ichilo, Buena Vista; 12 – 13 Feb. 2010; T. Vidaurre leg.; MHNNKM • 1 ♂, 1 ♀; same data as for preceding; 2 – 9 Nov. 2004; P. H. Sullivan leg.; MHNNKM • 2 ♂♂, 3 ♀♀; same data as for preceding; 2 – 12 Feb. 2000; J. E. Wappes leg.; MHNNKM • 2 ♂♂, 2 ♀♀; same data as for preceding; 2 – 13 Feb. 2000; M. C. Thomas leg.; MHNNKM • 1 ♀; same data as for preceding; 23 – 25 Apr. 1997; S. Amezquita leg.; MHNNKM • 1 ♂, 1 ♀; same data as for preceding; 27 – 30 Nov. 1997; S. Amezquita leg.; MHNNKM • 2 ♂♂, 3 ♀♀; Ichilo, Campamento Mataracú; 11 – 17 Oct. 1997; MHNNKM • 2 ♀♀; same data as for preceding; Oct. 1997; RAP- 97 exped. leg.; MHNNKM • 2 ♂♂, 2 ♀♀; same data as for preceding; 17 – 20 Sep. 1997; S. Aysama leg.; MHNNKM • 1 ♂, 1 ♀; same data as for preceding; 3 Mar. 1997; J. Ledezma leg.; MHNNKM • 1 ♂; Ichilo, Caranda; 6 – 7 Nov. 2004; F. Aguanta leg.; MHNNKM • 1 ♂, 1 ♀; Ichilo, Chore; 17 ° 01 ′ S, 64 ° 11 ′ W; elev. 253 m; 9 – 10 May 1997; F. Copa leg.; MHNNKM • 4 ♂♂, 8 ♀♀; Ichilo, Yapacani (BEEM); 8 – 12 Aug. 2006; I. García et al. leg.; MHNNKM • 1 ♂; Ñuflo de Chavez, San Javier; 29 Nov. 1995; I. García leg.; MHNNKM • 2 ♂♂; San Ignacio de Velasco, Las Gamas; 24 – 27 Feb. 1997; I. García leg.; MHNNKM • 1 ♂, 1 ♀; same data as for preceding; 21 Nov. 1997; MHNNKM • 3 ♂♂, 3 ♀♀; San Ignacio de Velasco, Los Fierros; 25 Jan. 1997; S. Spector et al. leg.; MHNNKM • 1 ♂; San Ignacio de Velasco, RPPN Caparú, 14 ° 48 ′ S, 61 ° 10 ′ W; elev. 150 m; 13 Dec. 2005; C. Hamel leg.; MHNNKM • 1 ♂; San Ignacio de Velasco, RPPN Caparú, Lagunitas; 20 Feb. 2007; A. Alcoba and M. Saldias leg.; MHNNKM • 1 ♂, 1 ♀; same data as for preceding; 11 Jan. 2008; R. Palomino leg.; MHNNKM • 1 ♂, 1 ♀; same data as for preceding; 16 Mar. 2008; MHNNKM • 1 ♂; San Ignacio de Velasco, San Miguel; 26 Jan. 2009; T. Vidaurre leg.; MHNNKM • 1 ♀; P. N. Noel Kempf Mercado; 13 ° 54 ′ 59 ″ S, 60 ° 48 ′ 51 ″ W; elev. 500 m; 17 – 20 Nov. 1999; S. Spector and A. Ayzama leg.; CEMT. BRAZIL – Acre • 3 ♂♂, 4 ♀♀; Mâncio Lima, Parque Nacional da Serra do Divisor; 7 ° 27 ′ 4 ″ S, 73 ° 38 ′ 57 ″ W; 6 – 8 Dec. 2019; R. A. Azevedo leg.; pitfall human dung; CEMT • 1 ♂, 1 ♀; same data as for preceding; 2 – 14 Dec. 2019; flight interception trap; CEMT • 5 ♂♂, 6 ♀♀; same data as for preceding; 7 ° 24 ′ 58 ″ S, 73 ° 37 ′ 56 ″ W; CEMT • 1 ♂, 1 ♀; same data as for preceding; 7 ° 26 ′ 46 ″ S, 73 ° 39 ′ 27 ″ W; 7 – 9 Dec. 2019; pitfall human dung; CEMT • 5 ♂♂, 8 ♀♀; same data as for preceding; 7 ° 24 ′ 59 ″ S, 73 ° 37 ′ 56 ″ W; 6 – 8 Dec. 2019; pitfall human dung; CEMT • 1 ♀; Manoel Urbano, Parque Estadual Chandless; 9 ° 22 ′ 26 ″ S, 69 ° 55 ′ 20 ″ W; 11 Jul. 2013; T. F. Brito leg.; CEMT • 5 ♂♂, 1 ♀; Rio Branco, Fazenda Catuaba; Feb. 1997; F. Z. Vaz-de-Mello leg.; CEMT • 1 ♂, 2 ♀♀; Senador Guiomard; 10 ° 04 ′ S, 67 ° 37 ′ W; 14 – 20 Apr. 2017; B. S. Bitencourt leg.; pitfall human dung; CEMT • 1 ♀; Senador Guiomard, Reserva Catuaba; 10 ° 4 ′ 27 ″ S, 67 ° 36 ′ 14 ″ W; 16 – 21 Jul. 2019; Casagrande leg.; CEMT • 1 ♂; Xapuri, Reserva Chico Mendes; 10 ° 14 ′ S, 68 ° 42 ′ W; 19 Oct. 2008; J. Silveira leg.; pitfall human dung; CEMT • 1 ♀; same data as for preceding; 10 ° 15 ′ S, 68 ° 42 ′ W; CEMT • 1 ♂; same data as for preceding; 10 ° 31 ′ S, 68 ° 42 ′ W; CEMT. – Amapá • 1 ♂, 1 ♀; Serra do Navio, Cava Urucum - Amapari; 0 ° 53 ′ 6 ″ N, 51 ° 52 ′ 53 ″ W; Sep. 2000; R. Ribon leg.; CEMT. – Amazonas • 1 ♂; Benjamin Constant, Rio Javari; Jan. 1961.; CEMT • 1 ♂, 1 ♀; Benjamin Constant, Guanabara II; 10 Mar. 2004; P. H. Silva leg.; CEMT • 3 ♀♀; Borba, BR – 319, km 220; 4 ° 24 ′ 3 ″ S, 60 ° 56 ′ 10 ″ W; 29 Dec. 2015; D. Pires leg.; pitfall human dung; CEMT • 1 ♂; same data as for preceding; 4 ° 24 ′ 6 ″ S, 60 ° 55 ′ 22 ″ W; 30 Dec. 2015; CEMT • 1 ♀; same data as for preceding; 4 ° 22 ′ 55 ″ S, 60 ° 57 ′ 19 ″ W; 29 Dec. 2015; CEMT • 1 ♂; Carauari, Reserva de Desenvolvimento Sustentável de Uacari; 5 ° 46 ′ 06 ″ S, 67 ° 42 ′ 40 ″ W; Jul. 2009; Nicols et al. leg.; CEMT • 1 ♀; Careiro, BR – 319; 4 ° 8 ′ 9 ″ S, 60 ° 43 ′ 31 ″ W; 3 Dec. 2015; D. Pires leg.; pitfall human dung; CEMT • 3 ♀♀; same data as for preceding; 4 ° 7 ′ 39 ″ S, 60 ° 43 ′ 43 ″ W; CEMT • 6 ♂♂, 3 ♀♀; same data as for preceding; 4 ° 8 ′ 40 ″ S, 60 ° 43 ′ 20 ″ W; CEMT • 3 ♀♀; same data as for preceding; 4 ° 7 ′ 58 ″ S, 60 ° 43 ′ 9 ″ W; 2 Dec. 2015; CEMT • 1 ♂, 6 ♀♀; same data as for preceding; 4 ° 8 ′ 58 ″ S, 60 ° 43 ′ 44 ″ W; CEMT • 1 ♀; same data as for preceding; 3 ° 21 ′ 46 ″ S, 59 ° 51 ′ 10 ″ W; 9 Dec. 2015; CEMT • 2 ♀♀; same data as for preceding; 4 ° 8 ′ 28 ″ S, 60 ° 43 ′ 57 ″ W; 2 Dec. 2015; CEMT • 1 ♀; same data as for preceding; 4 ° 9 ′ 28 ″ S, 60 ° 43 ′ 32 ″ W; CEMT • 2 ♀♀; Coari, Lago Uauaçu; 4 ° 12 ′ 59 ″ S, 62 ° 27 ′ 10 ″ W; elev. 66 m; Jan. – Sep. 2018; J. B. R. Alencar leg.; pitfall human dung; CEMT • 1 ♀; same data as for preceding; 4 ° 13 ′ 7 ″ S, 62 ° 27 ′ 9 ″ W; elev. 63 m; CEMT • 1 ♀; same data as for preceding; 4 ° 15 ′ 45 ″ S, 62 ° 16 ′ 2 ″ W; elev. 55 m; 29 Oct. 2018; CEMT • 1 ♀; Itapiranga, Fazenda Itapiranga; 2 ° 39 ′ 56 ″ S, 58 ° 40 ′ 23 ″ W; 3 May 2019; R. S. Moura leg.; CEMT • 1 ♂, 2 ♀♀; Manaus, Reserva Florestal Adolpho Ducke; 18 Jul. 1978; B. C. Ratcliffe leg.; CEMT • 1 ♂, 1 ♀; same data as for preceding; 25 Oct. – 22 Nov. 1977; CEMT • 1 ♀; same data as for preceding; 8 Nov. 1978; CEMT • 4 ♂♂, 5 ♀♀; Manicoré, Reserva Extrativista Lago Capanã Grande, BR 319, km 45; 5 ° 54 ′ 33 ″ S, 62 ° 23 ′ 56 ″ W; 30 Oct. – 1 Nov. 2019; R. A. Azevedo leg.; pitfall human dung; CEMT • 2 ♀♀; same data as for preceding; 5 ° 1 ′ 5 ″ S, 62 ° 29 ′ 25 ″ W; 31 Oct. – 2 Nov. 2019; CEMT • 1 ♂, 3 ♀♀; same data as for preceding; 5 ° 58 ′ 24 ″ S, 62 ° 27 ′ 38 ″ W; CEMT • 4 ♂♂, 4 ♀♀; same data as for preceding; 5 ° 54 ′ 33 ″ S, 62 ° 23 ′ 56 ″ W; 22 – 26 Oct. 2019; flight interception trap; CEMT • 6 ♀♀; same data as for preceding; 26 – 30 Oct. 2019; CEMT • 2 ♂♂, 3 ♀♀; same data as for preceding; 5 ° 58 ′ 24 ″ S, 62 ° 27 ′ 38 ″ W; 23 – 27 Oct. 2019; CEMT • 3 ♀♀; same data as for preceding; 6 ° 1 ′ 5 ″ S, 62 ° 29 ′ 25 ″ W; CEMT • 2 ♂♂, 3 ♀♀; same data as for preceding; 27 – 31 Oct. 2019; CEMT • 1 ♂; same data as for preceding; 31 Oct. – 4 Nov. 2019; CEMT • 2 ♂♂, 1 ♀; Manicoré, BR 319, km 350; 5 ° 12 ′ 56 ″ S, 61 ° 50 ′ 22 ″ W; 30 Jul. – 2 Aug. 2007; H. Gasca leg.; CEMT • 1 ♂, 5 ♀♀; Presidente Figueiredo; Nov. 1997; E. Andresen leg.; CEMT • 1 ♀; same data as for preceding; Oct. – Nov. 1996; CEMT • 1 ♂; same data as for preceding; Alouatta dung; CEMT • 1 ♂; Silves; 2 ° 53 ′ 53 ″ S, 58 ° 54 ′ 10 ″ W; 4 – 11 May 2019; R. S. Moura leg.; CEMT. – Mato Grosso • 2 ♂♂; Alta Floresta; May 2008; E. Berenguer leg.; pitfall human dung; CEMT • 2 ♂♂, 2 ♀♀; same data as for preceding; 9 ° 56 ′ 52 ″ S, 56 ° 3 ′ 2 ″ W; CEMT • 2 ♂♂, 4 ♀♀; same data as for preceding; 9 ° 41 ′ 24 ″ S, 55 ° 56 ′ 28 ″ W; Jun. 2008; CEMT • 1 ♀; same data as for preceding; 9 ° 58 ′ 46 ″ S, 56 ° 5 ′ 31 ″ W; CEMT • 2 ♀♀; same data as for preceding; 9 ° 52 ′ 53 ″ S, 56 ° 6 ′ 13 ″ W; CEMT • 2 ♀♀; same data as for preceding; 9 ° 53 ′ 40 ″ S, 56 ° 16 ′ 34 ″ W; CEMT • 4 ♂♂, 8 ♀♀; same data as for preceding; 9 ° 49 ′ 44 ″ S, 56 ° 19 ′ 51 ″ W; CEMT • 4 ♀♀; Alta Floresta, Ceplac; 24 May 2009; V. Gonçalves leg.; CEMT • 1 ♂, 1 ♀; Alta Floresta, Parque Estadual do Cristalino; 9 ° 35 ′ 50 ″ S, 55 ° 53 ′ 40 ″ W; 11 – 15 Nov. 2019; R. A. Azevedo leg.; flight interception trap; CEMT • 1 ♂, 5 ♀♀; same data as for preceding; 9 ° 40 ′ 59 ″ S, 55 ° 54 ′ 58 ″ W; 12 – 16 Nov. 2019; CEMT • 1 ♂, 3 ♀♀; same data as for preceding; 20 – 24 Nov. 2019; CEMT • 1 ♀; same data as for preceding; 16 – 20 Nov. 2019; CEMT • 2 ♂♂; same data as for preceding; 9 ° 35 ′ 50 ″ S, 55 ° 53 ′ 40 ″ W; 15 – 17 Nov. 2019; CEMT • 2 ♀♀; same data as for preceding; pitfall human dung; CEMT • 3 ♀♀; same data as for preceding; 9 ° 32 ′ 45 ″ S, 55 ° 54 ′ 56 ″ W; CEMT • 3 ♀♀; same data as for preceding; 9 ° 40 ′ 59 ″ S, 55 ° 53 ′ 40 ″ W; CEMT • 2 ♂♂, 1 ♀; Araputanga, Fazenda Bandeirantes; 15 ° 23 ′ 26 ″ S, 58 ° 24 ′ 48 ″ W; elev. 296 m; 18 – 20 Jan. 2013; R. J Silva leg.; pitfall human dung; CEMT • 1 ♂, 2 ♀♀; same data as for preceding; 15 ° 23 ′ 6 ″ S, 58 ° 26 ′ 2 ″ W; elev. 308 m; 19 – 21 Jan. 2013; CEMT • 2 ♂♂; same data as for preceding; 15 ° 23 ′ 4 ″ S, 58 ° 26 ′ W; elev. 296 m; 21 – 23 Jan. 2013; CEMT • 1 ♂, 2 ♀♀; Aripuanã, Fazenda Águas Claras; 10 ° 12 ′ 9 ″ S, 52 ° 34 ′ 26 ″ W; elev. 275 m; 9 – 11 Oct. 2018; R. J Silva leg.; flight interception trap; CEMT • 1 ♀; Aripuanã, Serra do Expedito; 10 ° 4 ′ 22 ″ S, 59 ° 30 ′ 41 ″ W; 25 Jan. 2017; T. Semedo leg.; CEMT • 1 ♂; Brasnorte, Fazenda Estrela do Sul; 12 ° 27 ′ 30 ″ S, 58 ° 6 ′ 29 ″ W; elev. 336 m; 30 Jan. – 1 Feb. 2018; U. F. Santos leg.; pitfall human dung; CEMT • 1 ♀; same data as for preceding; Fazenda Agriter; 13 ° 2 ′ 37 ″ S, 57 ° 52 ′ 55 ″ W; elev. 434 m; CEMT • 1 ♀; Carlinda; 13 May 2008; C. R. Menezes leg.; CEMT • 1 ♀; Carlinda; 14 Oct. 2008; I. B. Banheza leg.; CEMT • 1 ♂; Carlinda; 15 Oct. 2008; B. R. Krause leg.; CEMT • 1 ♂; Carlinda; 16 Oct. 2008; I. B. Banheza leg.; CEMT • 1 ♀; same data as for preceding; L. A. Aguiar leg.; CEMT • 3 ♂♂, 10 ♀♀; Carlinda; 21 – 23 Oct. 2008; K. Segatto et al. leg.; CEMT • 1 ♂; Carlinda; 12 Oct. 2009; J. S. Costa leg.; CEMT • 1 ♀; Claudia, Fazenda Continental; 11 ° 25 ′ 41 ″ S, 55 ° 18 ′ 54 ″ W; 20 Oct. 2011; M. F. Souza leg.; pitfall human dung; CEMT • 1 ♀; Cotriguaçu, Fazenda São Nicolau; 9 ° 50 ′ 24 ″ S, 58 ° 15 ′ 10 ″ W; elev. 250 m; 5 Oct. 2009; F. Z. Vaz-de-Mello leg.; pitfall human dung; CEMT • 7 ♂♂, 10 ♀♀; same data as for preceding; 8 – 12 Oct. 2009; CEMT • 1 ♀; same data as for preceding; 9 ° 51 ′ 18 ″ S, 58 ° 13 ′ 20 ″ W; elev. 200 m; Oct. 2009; CEMT • 1 ♂, 6 ♀♀; same data as for preceding; 9 ° 50 ′ 19 ″ S, 58 ° 15 ′ 3 ″ W; 8 Oct. 2009; CEMT • 2 ♀♀; same data as for preceding; pitfall cow dung; CEMT • 7 ♂♂, 2 ♀♀; same data as for preceding; 9 ° 51 ′ 36 ″ S, 58 ° 12 ′ 53 ″ W; Oct. 2009; pitfall human dung; CEMT • 3 ♂♂, 3 ♀♀; same data as for preceding; 9 ° 50 ′ 53 ″ S, 58 ° 14 ′ 36 ″ W; 11 Dec. 2009; CEMT • 4 ♂♂; same data as for preceding; 9 ° 50 ′ 24 ″ S, 58 ° 15 ′ 10 ″ W; elev. 250 m; 13 Dec. 2009; CEMT • 1 ♂, 2 ♀♀; same data as for preceding; 9 ° 50 ′ 19 ″ S, 58 ° 15 ′ 3 ″ W; Dec. 2009; horse dung; CEMT • 6 ♂♂, 9 ♀♀; same data as for preceding; 11 – 13 Dec. 2009; pitfall human dung; CEMT • 1 ♀; same data as for preceding; 9 ° 49 ′ 9 ″ S, 58 ° 15 ′ 47 ″ W; 8 – 10 Dec. 2009; CEMT • 1 ♂, 1 ♀; same data as for preceding; 9 ° 51 ′ 36 ″ S, 58 ° 12 ′ 53 ″ W; 11 – 13 Dec. 2009; CEMT • 1 ♀; same data as for preceding; 9 ° 49 ′ 9 ″ S, 58 ° 15 ′ 30 ″ W; 24 – 31 Oct. 2014; M. Karam-Gemael leg.; CEMT • 2 ♀♀; same data as for preceding; 9 ° 50 ′ 19 ″ S, 58 ° 15 ′ 15 ″ W; 30 Oct. 2017; F. Z. Vaz-de-Mello leg.; matinha do Fernando; flight interception trap; CEMT • 3 ♂♂, 5 ♀♀; same data as for preceding; 3 Nov. 2017; CEMT • 3 ♂♂, 2 ♀♀; same data as for preceding; 9 ° 49 ′ 17 ″ S, 58 ° 15 ′ 32 ″ W; PPBio 1; 1 – 3 Nov. 2017; CEMT • 1 ♀; same data as for preceding; 9 ° 52 ′ 31 ″ S, 58 ° 16 ′ 21 ″ W; 2 Nov. 2017; R. Stofel leg.; trail of garrote; CEMT • 8 ♂♂, 4 ♀♀; same data as for preceding; 9 ° 51 ′ S, 58 ° 14 ′ W; 4 – 6 Nov. 2017; J. P. Santos leg.; pitfall human dung; CEMT • 1 ♂, 1 ♀; same data as for preceding; 9 ° 50 ′ 15 ″ S, 58 ° 15 ′ 14 ″ W; 23 – 29 Oct. 2018; V. Costa-Silva leg.; CEMT • 2 ♀♀; same data as for preceding; 9 ° 50 ′ 05 ″ S, 58 ° 16 ′ 04 ″ W; 18 – 20 Oct. 2021; J. Mariano et al. leg.; pitfall human dung; CEMT • 1 ♂, 1 ♀; same data as for preceding; 9 ° 50 ′ 07 ″ S, 58 ° 16 ′ W; CEMT • 2 ♀♀; same data as for preceding; 9 ° 50 ′ 10 ″ S, 58 ° 16 ′ W; CEMT • 3 ♂♂, 2 ♀♀; same data as for preceding; 9 ° 50 ′ 47 ″ S, 58 ° 14 ′ 09 ″ W; 19 – 21 Oct. 2021; CEMT • 1 ♂, 3 ♀♀; same data as for preceding; 9 ° 50 ′ 49 ″ S, 58 ° 14 ′ 04 ″ W; CEMT • 3 ♀♀; same data as for preceding; 9 ° 52 ′ 08 ″ S, 58 ° 15 ′ 19 ″ W; CEMT • 4 ♀♀; same data as for preceding; 9 ° 51 ′ 25 ″ S, 58 ° 15 ′ 16 ″ W; 27 – 29 Oct. 2021; CEMT • 6 ♂♂, 4 ♀♀; same data as for preceding; 9 ° 50 ′ S, 58 ° 15 ′ W; 28 – 30 Oct. 2021; CEMT • 7 ♂♂, 1 ♀; same data as for preceding; 9 ° 49 ′ 12 ″ S, 58 ° 15 ′ 30 ″ W; CEMT • 3 ♂♂, 2 ♀♀; same data as for preceding; 9 ° 49 ′ 47 ″ S, 58 ° 16 ′ 19 ″ W; 1 – 3 Nov. 2021; CEMT • 2 ♂♂, 2 ♀♀; same data as for preceding; 9 ° 49 ′ 47 ″ S, 58 ° 16 ′ 21 ″ W; CEMT • 4 ♂♂, 4 ♀♀; same data as for preceding; 9 ° 49 ′ 47 ″ S, 58 ° 16 ′ 22 ″ W; CEMT • 3 ♂♂; same data as for preceding; 9 ° 49 ′ 47 ″ S, 58 ° 16 ′ 24 ″ W; CEMT • 2 ♀♀; same data as for preceding; 9 ° 49 ′ 47 ″ S, 58 ° 16 ′ 26 ″ W; CEMT • 1 ♂, 2 ♀♀; same data as for preceding; 9 ° 50 ′ 10 ″ S, 58 ° 15 ′ 49 ″ W; CEMT • 1 ♂, 2 ♀♀; same data as for preceding; 9 ° 50 ′ 11 ″ S, 58 ° 15 ′ 50 ″ W; CEMT • 2 ♀♀; same data as for preceding; 9 ° 51 ′ 43 ″ S, 58 ° 13 ′ 10 ″ W; 2 – 4 Nov. 2021; CEMT • 1 ♂; same data as for preceding; 9 ° 51 ′ 42 ″ S, 58 ° 13 ′ 11 ″ W; CEMT • 1 ♂, 1 ♀; same data as for preceding; 9 ° 51 ′ 26 ″ S, 58 ° 14 ′ 24 ″ W; CEMT • 1 ♂; same data as for preceding; 9 ° 51 ′ 32 ″ S, 58 ° 14 ′ 23 ″ W; CEMT • 1 ♂; same data as for preceding; 9 ° 51 ′ 31 ″ S, 58 ° 14 ′ 23 ″ W; CEMT • 1 ♀; same data as for preceding; 9 ° 51 ′ 28 ″ S, 58 ° 14 ′ 23 ″ W; CEMT • 2 ♂♂, 5 ♀♀; same data as for preceding; 9 ° 49 ′ 43 ″ S, 58 ° 16 ′ 30 ″ W; 3 – 5 Nov. 2021; CEMT • 1 ♂, 1 ♀; same data as for preceding; 9 ° 49 ′ 43 ″ S, 58 ° 16 ′ 32 ″ W; CEMT • 1 ♂, 2 ♀♀; same data as for preceding; 9 ° 49 ′ 43 ″ S, 58 ° 16 ′ 34 ″ W; CEMT • 3 ♀♀; same data as for preceding; 9 ° 49 ′ 43 ″ S, 58 ° 16 ′ 35 ″ W; CEMT • 1 ♂, same data as for preceding; 9 ° 49 ′ 44 ″ S, 58 ° 16 ′ 29 ″ W; CEMT • 1 ♀; same data as for preceding; 9 ° 49 ′ 26 ″ S, 58 ° 15 ′ 49 ″ W; 4 – 6 Nov. 2021; CEMT • 1 ♀; same data as for preceding; 9 ° 49 ′ 48 ″ S, 58 ° 16 ′ 11 ″ W; CEMT • 1 ♀; same data as for preceding; 9 ° 49 ′ 49 ″ S, 58 ° 16 ′ 09 ″ W; CEMT • 1 ♂, same data as for preceding; 9 ° 49 ′ 50 ″ S, 58 ° 16 ′ 06 ″ W; CEMT • 2 ♂♂, 1 ♀; same data as for preceding; 9 ° 50 ′ 53 ″ S, 58 ° 15 ′ 19 ″ W; CEMT • 7 ♂♂, 4 ♀♀; same data as for preceding; 9 ° 50 ′ 54 ″ S, 58 ° 15 ′ 21 ″ W; CEMT • 2 ♀♀; same data as for preceding; 9 ° 50 ′ 55 ″ S, 58 ° 15 ′ 21 ″ W; CEMT • 1 ♀; same data as for preceding; 9 ° 50 ′ 56 ″ S, 58 ° 15 ′ 22 ″ W; CEMT • 1 ♀; same data as for preceding; 9 ° 50 ′ 58 ″ S, 58 ° 15 ′ 23 ″ W; CEMT • 1 ♂, 1 ♀; same data as for preceding; 9 ° 50 ′ 55 ″ S, 58 ° 14 ′ 10 ″ W; 5 – 7 Nov. 2021; CEMT • 2 ♀♀; same data as for preceding; 9 ° 50 ′ 56 ″ S, 58 ° 14 ′ 08 ″ W; CEMT • 3 ♂♂, 3 ♀♀; same data as for preceding; 9 ° 51 ′ 01 ″ S, 58 ° 14 ′ 10 ″ W; CEMT • 1 ♂, 3 ♀♀; same data as for preceding; 9 ° 51 ′ 02 ″ S, 58 ° 14 ′ 07 ″ W; CEMT • 3 ♂♂; same data as for preceding; 9 ° 51 ′ 03 ″ S, 58 ° 14 ′ 06 ″ W; CEMT • 1 ♂, 4 ♀♀; same data as for preceding; 9 ° 49 ′ 25 ″ S, 58 ° 16 ′ 17 ″ W; CEMT • 5 ♀♀; same data as for preceding; 9 ° 49 ′ 26 ″ S, 58 ° 16 ′ 38 ″ W; CEMT • 2 ♀♀; same data as for preceding; 9 ° 49 ′ 27 ″ S, 58 ° 16 ′ 38 ″ W; CEMT • 1 ♂; same data as for preceding; 9 ° 49 ′ 27 ″ S, 58 ° 16 ′ 14 ″ W; CEMT • 1 ♂; same data as for preceding; 9 ° 49 ′ 29 ″ S, 58 ° 16 ′ 39 ″ W; CEMT • 2 ♂♂, 6 ♀♀; same data as for preceding; 9 ° 49 ′ 30 ″ S, 58 ° 16 ′ 12 ″ W; CEMT • 2 ♂♂, 4 ♀♀; same data as for preceding; 9 ° 49 ′ 30 ″ S, 58 ° 16 ′ 39 ″ W; CEMT • 1 ♂, 3 ♀♀; same data as for preceding; 9 ° 49 ′ 32 ″ S, 58 ° 16 ′ 39 ″ W; CEMT • 3 ♂♂, 4 ♀♀; same data as for preceding; 9 ° 50 ′ 56 ″ S, 58 ° 14 ′ 06 ″ W; CEMT • 2 ♂♂, 1 ♀; same data as for preceding; 9 ° 50 ′ 57 ″ S, 58 ° 14 ′ 05 ″ W; CEMT • 1 ♂, 1 ♀; same data as for preceding; 9 ° 51 ′ 02 ″ S, 58 ° 14 ′ 09 ″ W; CEMT • 2 ♀♀; same data as for preceding; 9 ° 51 ′ 08 ″ S, 58 ° 16 ′ 20 ″ W; CEMT • 1 ♀; same data as for preceding; 9 ° 51 ′ 09 ″ S, 58 ° 16 ′ 21 ″ W; CEMT • 1 ♀; same data as for preceding; 9 ° 51 ′ 11 ″ S, 58 ° 16 ′ 22 ″ W; CEMT • 1 ♂, 1 ♀; same data as for preceding; 9 ° 51 ′ 43 ″ S, 58 ° 15 ′ 50 ″ W; CEMT • 1 ♂, 1 ♀; same data as for preceding; 9 ° 51 ′ 48 ″ S, 58 ° 15 ′ 50 ″ W; CEMT • 1 ♀; Diamantino; Feb. 2003; E. Furtado leg.; CEMT • 1 ♂; Diamantino; 16 Mar. 2018; L. G. Nunes and E. Carvalho leg.; CEMT • 1 ♂, 1 ♀; same data as for preceding; 13 ° 48 ′ 07 ″ S, 56 ° 41 ′ 32 ″ W; 17 Mar. 2018; pitfall human dung; CEMT • 1 ♀; same data as for preceding; 13 ° 48 ′ 02 ″ S, 56 ° 41 ′ 08 ″ W; pitfall human dung; CEMT • 1 ♂; same data as for preceding; Semidouro; 13 ° 48 ′ 51 ″ S, 56 ° 41 ′ 32 ″ W; pitfall human dung; CEMT • 3 ♂♂, 1 ♀; Indiavaí, Faz. Alto Jaurú; 15 ° 27 ′ 18 ″ S, 58 ° 33 ′ 12 ″ W; elev. 231 m; 14 – 16 Jan. 2013; R. J. Silva leg.; pitfall human and pig dung; CEMT • 2 ♂♂, 1 ♀; Juína, Fazenda Quiles; 29 Nov. 2013; R. J. Silva leg.; carcass; CEMT • 1 ♂; Nova Maringá, Faz. Nova Fronteira; 13 ° 15 ′ 02 ″ S, 57 ° 04 ′ 45 ″ W; elev. 356 m; 19 – 21 Dec. 2017; U. F. Santos leg.; pitfall human dung; CEMT • 2 ♂♂, 1 ♀; Novo Mundo, Parque Estadual do Cristalino; 9 ° 28 ′ 01 ″ S, 55 ° 47 ′ 43 ″ W; May 2013; V. Magalhães leg.; pitfall; CEMT • 1 ♂, 1 ♀; same data as for preceding; 9 ° 28 ′ 06 ″ S, 55 ° 50 ′ 36 ″ W; CEMT • 1 ♂, 1 ♀; same data as for preceding; 9 ° 28 ′ 09 ″ S, 55 ° 48 ′ 15 ″ W; CEMT • 1 ♂; same data as for preceding; 9 ° 27 ′ 6 ″ S, 55 ° 50 ′ 02 ″ W; CEMT • 1 ♂, 1 ♀; Paranaíta; 9 ° 14 ′ 16 ″ S, 57 ° 0 ′ 14 ″ W; Dec. 2015; M. A. L. Bragança leg.; pitfall; CEMT • 1 ♂; same data as for preceding; 9 ° 15 ′ 39 ″ S, 56 ° 48 ′ 50 ″ W; CEMT • 1 ♂; 9 ° 15 ′ 56 ″ S, 56 ° 58 ′ 45 ″ W; CEMT • 1 ♀; same data as for preceding; 9 ° 15 ′ 40 ″ S, 56 ° 48 ′ 46 ″ W; CEMT • 2 ♂♂; Paranaíta, Rio Teles Pires; 24 Oct. 2009; H. A. B. Faria leg.; CEMT • 3 ♂♂; São José do Rio Claro; 13 ° 53 ′ 10 ″ S, 56 ° 40 ′ 42 ″ W; 30 Jun. 2018; L. G. Nunes and E. Carvalho leg.; pitfall human dung; CEMT • 1 ♀; Tangará da Serra, Fazenda Curitiba; 14 ° 22 ′ 18 ″ S, 57 ° 29 ′ 24 ″ W; elev. 343 m; 31 Mar. 2012; R. J. Silva leg.; pitfall cow dung; CEMT • 1 ♂, 1 ♀; Tangará da Serra, Fazenda Rosa Branca; 14 ° 34 ′ S, 57 ° 45 ′ 24 ″ W; elev. 468 m; 18 – 27 Feb. 2011; R. J. Silva leg.; pitfall human and pig dung; CEMT • 1 ♂, 3 ♀♀; Tangará da Serra, Fazenda Netolândia; 14 ° 42 ′ 18 ″ S, 57 ° 54 ′ 14 ″ W; elev. 310 m; 14 – 21 Dec. 2012; R. J. Silva leg.; flight interception trap; CEMT • 1 ♂, 1 ♀; same data as for preceding; 14 ° 41 ′ 05 ″ S, 57 ° 54 ′ 08 ″ W; elev. 263 m; 25 – 27 Mar. 2012; pitfall cow dung; CEMT • 1 ♂; same data as for preceding; 14 ° 39 ′ 56 ″ S, 57 ° 54 ′ 08 ″ W; elev. 304 m; 20 – 22 Mar. 2012; CEMT • 1 ♂; same data as for preceding; 14 ° 39 ′ 54 ″ S, 57 ° 55 ′ 05 ″ W; elev. 330 m; 12 – 14 Mar. 2012; pitfall human and pig dung; CEMT. – Pará • 1 ♂; Almeirim, Monte Dourado, 1 ° 1 ′ S, 52 ° 44 ′ W; elev. 150 m; Jul. 2004; T. A. Gardner leg.; pitfall dung; CEMT • 5 ♂♂, 1 ♀; Monte Dourado, Pacanari; 0 ° 38 ′ S, 52 ° 35 ′ W; Mar. 2006; Gardner and Hernandez leg.; dung trap; TAMU • 1 ♂, 1 ♀; Cachimbo; Oct. 1995; Pe. Pereira leg.; TAMU • 5 ♂♂, 2 ♀♀; 9 ° 12 ′ 21 ″ S, 56 ° 57 ′ 34 ″ W; Dec. 2015; M. A. L. Bragança leg.; pitfall; CEMT • 1 ♂; same data as for preceding; 9 ° 12 ′ 35 ″ S, 56 ° 57 ′ 39 ″ W; CEMT • 1 ♀; Jacareacanga, Rio Teles Pires; 9 ° 15 ′ 48 ″ S, 56 ° 46 ′ 42 ″ W; 9 Jun. 2009; M. F. Souza and M. Uehara-Prado leg.; pitfall human dung; CEMT • 1 ♂; same data as for preceding; 9 ° 19 ′ 3 ″ S, 56 ° 46 ′ 41 ″ W; 8 Jun. 2009; CEMT • 2 ♀♀; Novo Progresso, Fazenda Florentino; 7 ° 8 ′ 51 ″ S, 55 ° 23 ′ 38 ″ W; Jan. 2011; T. D. Pelissari leg.; pitfall human dung; CEMT • 1 ♀; same data as for preceding; 7 ° 6 ′ 54 ″ S, 55 ° 24 ′ 23 ″ W; CEMT • 1 ♂; same data as for preceding; 7 ° 6 ′ 21 ″ S, 55 ° 25 ′ 13 ″ W; Mar. 2011; CEMT • 1 ♀; same data as for preceding; 7 ° 54 ′ 19 ″ S, 55 ° 29 ′ 59 ″ W; 2 Jan. 2011; CEMT • 1 ♀; same data as for preceding; 7 ° 53 ′ 18 ″ S, 55 ° 30 ′ 26 ″ W; 12 Jan. 2011; CEMT • 3 ♂♂, 1 ♀; Redenção, Fazenda Marajoara; 7 ° 50 ′ S, 50 ° 16 ′ W; Oct. 1998; P. Y. Scheffer leg.; CEMT • 1 ♂, 1 ♀; Redenção, Pinkaiti - Aik; 7 ° 46 ′ S, 51 ° 58 ′ W; Oct. 1999; P. Y. Scheffer leg.; CEMT • 1 ♂, 1 ♀; Santarém, Reserva Tapajós; 2 ° 46 ′ S, 55 ° 40 ′ W; 25 Dec. 2008; R. Andrade leg.; pitfall human dung; CEMT • 1 ♂, 1 ♀; Tailândia, Empresa Agropalma; 9 – 15 Jul. 2016; F. Silva leg.; CEMT. – Rondônia • 1 ♂; Alto Paraíso; Jul. 2002; E. Furtado leg.; CEMT • 2 ♂♂, 1 ♀; same data as for preceding; 12 – 17 Aug. 2002; CEMT • 2 ♀♀; same data as for preceding; Sep. 2002; CEMT • 5 ♂♂, 2 ♀♀; same data as for preceding; W. Furtado leg.; CEMT • 1 ♂, 1 ♀; same data as for preceding; Oct. 2002; CEMT • 3 ♂♂, 2 ♀♀; same data as for preceding; Nov. 2002; CEMT • 1 ♀; same data as for preceding; 13 – 23 Apr. 2013; E. Furtado leg.; CEMT • 1 ♂, 1 ♀; 62 km SW of Ariquemes, Fazenda Rancho Grande; 18 – 29 Sep. 1996; D. G. Marqua leg.; TAMU • 1 ♀; 60 km SW of Ariquemes, Fazenda Rancho Grande; Feb. 1997; F. Z. Vaz-de-Mello leg.; CEMT • 1 ♂, 2 ♀♀; Cacaulândia, Fazenda Rancho Grande; Feb. 1997; F. Z. Vaz-de-Mello leg.; CEMT • 1 ♀; Guajará-Mirim, Fazenda Agropecuária AJB; elev. 180 m; 15 Jan. 2010; F. Coletti leg.; carcass; CEMT • 1 ♂; Guajará-Mirim, Aristeu; 10 ° 22 ′ 16 ″ S, 64 ° 44 ′ 47 ″ W; elev. 220 m; 27 Jan. 2010; F. Coletti leg.; carcass; CEMT • 1 ♂; same data as for preceding; Fandinho; 10 ° 44 ′ 57 ″ S, 64 ° 19 ′ W; elev. 138 m; 24 Aug. 2010; cow dung; CEMT • 1 ♀; Itapuã do Oeste, FloNa Jamari; 9 ° 11 ′ 34 ″ S, 63 ° 06 ′ 34 ″ W; 20 Feb. 2013; J. F. Cerveira leg.; Mazama americana dung; CEMT • 1 ♀; same data as for preceding; 9 ° 11 ′ 33 ″ S, 63 ° 06 ′ 47 ″ W; CEMT • 1 ♀; same data as for preceding; 9 ° 11 ′ 29 ″ S, 63 ° 06 ′ 48 ″ W; CEMT • 1 ♀; same data as for preceding; 9 ° 11 ′ 27 ″ S, 63 ° 06 ′ 36 ″ W; Mazama gouazoubira dung; CEMT • 1 ♂; same data as for preceding; 9 ° 11 ′ 32 ″ S, 63 ° 06 ′ 35 ″ W; CEMT • 1 ♀; same data as for preceding; 9 ° 05 ′ 39 ″ S, 63 ° 08 ′ 55 ″ W; CEMT • 1 ♀; same data as for preceding; 9 ° 11 ′ 24 ″ S, 63 ° 06 ′ 50 ″ W; Mazama nemorivaga dung; CEMT • 2 ♀♀; Pimenta Bueno; 11 ° 43 ′ 3.21 ″ S, 61 ° 28 ′ 19.8 ″ W; 27 – 29 Jul. 2015; elev. 249 m; D. C. Castro leg.; pitfall human dung; CEMT • 1 ♂; same data as for preceding; 11 ° 43 ′ 3.26 ″ S, 61 ° 28 ′ 20.41 ″ W; CEMT • 2 ♂♂; same data as for preceding; 11 ° 43 ′ 3.28 ″ S, 61 ° 28 ′ 21 ″ W; CEMT • 1 ♂; same data as for preceding; 11 ° 43 ′ 3.32 ″ S, 61 ° 28 ′ 21.64 ″ W; CEMT • 2 ♀♀; same data as for preceding; 11 ° 43 ′ 3.8 ″ S, 61 ° 28 ′ 21.5 ″ W; CEMT • 3 ♀♀; same data as for preceding; 11 ° 43 ′ 4.8 ″ S, 61 ° 28 ′ 18.4 ″ W; CEMT • 1 ♂; same data as for preceding; 11 ° 43 ′ 4.96 ″ S, 61 ° 28 ′ 21.64 ″ W; CEMT • 1 ♀; same data as for preceding; 11 ° 43 ′ 5.43 ″ S, 61 ° 28 ′ 18.39 ″ W; CEMT • 1 ♂, 1 ♀; same data as for preceding; 11 ° 43 ′ 3.17 ″ S, 61 ° 28 ′ 19.2 ″ W; 9 – 11 Dec. 2015; CEMT 2 • ♀♀; same data as for preceding; 11 ° 43 ′ 3.82 ″ S, 61 ° 28 ′ 20.32 ″ W; CEMT • 1 ♀; same data as for preceding; 11 ° 43 ′ 4.29 ″ S, 61 ° 28 ′ 18.39 ″ W; CEMT • 2 ♀♀; same data as for preceding; 11 ° 43 ′ 4.32 ″ S, 61 ° 28 ′ 19.08 ″ W; CEMT • 1 ♀; same data as for preceding; 11 ° 43 ′ 4.9 ″ S, 61 ° 28 ′ 21.35 ″ W; CEMT • 1 ♂; same data as for preceding; 11 ° 43 ′ 5.49 ″ S, 61 ° 28 ′ 20.26 ″ W; CEMT • 1 ♀; same data as for preceding; 11 ° 43 ′ 5.49 ″ S, 61 ° 28 ′ 21.71 ″ W; CEMT • 1 ♀; Porto Velho, Abunã; 9 ° 35 ′ 46 ″ S, 60 ° 40 ′ 19 ″ W; 17 – 19 May 2010; L. R. Silva and F. F. Albertoni leg.; pitfall human dung; CEMT • 1 ♂; same data as for preceding; 9 ° 36 ′ 38 ″ S, 65 ° 21 ′ 33 ″ W; elev. 200 m; 19 Sep. 2010; J. C. F. Falcão leg.; pitfall human dung; CEMT • 1 ♀; same data as for preceding; Jirau; 9 ° 35 ′ 48 ″ S, 65 ° 21 ′ 56 ″ W; elev. 250 m; J. C. F. Falcão and L. R. Silva leg.; flight interception trap; CEMT • 1 ♂, 2 ♀♀; Porto Velho, EsEc Cuniã; 8 ° 4 ′ 11 ″ S, 63 ° 28 ′ 34 ″ W; elev. 83 m; 10 – 12 Nov. 2013; M. A. P. A. Silveira leg.; pitfall human dung; CEMT • 1 ♂; same data as for preceding; 13 – 15 Aug. 2012; C. Moreno leg.; CEMT • 1 ♂, 1 ♀; Porto Velho, Mutum Paraná; 9 ° 35 ′ S, 63 ° 03 ′ W; 15 – 17 Sep. 2010; J. C. F. Falcão and L. R. Silva leg.; pitfall human dung; CEMT • 1 ♂, 2 ♀♀; same data as for preceding; 9 ° 35 ′ 46 ″ S, 63 ° 02 ′ 27 ″ W; Sep. 2012; R. V. Nunes leg.; CEMT • 2 ♀♀; Porto Velho, Nova Mutum Paraná; 9 ° 26 ′ 31 ″ S, 60 ° 12 ′ 44 ″ W; 27 – 28 Feb. 2010; L. R. Silva and R. L. R. Silva leg.; pitfall human dung; CEMT. COLOMBIA – Amazonas • 1 ♂, 1 ♀; Leticia; elev. 213 m; 23 Feb. – 2 Mar. 1974; S. Peck leg.; TAMU • 1 ♀; Leticia, Estación Biológica El Zafire; 4 ° 00 ′ 21 ″ S, 69 ° 53 ′ 55 ″ W; elev. 150 m; 2 – 4 Dec. 2007; L. Franco and S. Flórez leg.; pitfall human dung; IAvH-E • 3 ♂♂, 1 ♀; same data as for preceding; 3 ° 59 ′ 15 ″ S, 69 ° 54 ′ 10 ″ W; elev. 164 m; 9 – 11 Dec. 2007; IAvH-E. – Caquetá • 2 ♀♀; Puerto Solano, PNN Serranía de Chiribiquete, Estación Puerto Abeja; 0 ° 04 ′ 27 ″ N, 72 ° 27 ′ 0.5 ″ W; elev. 200 m; Jul. 1999; M. Álvarez and G. Mejía leg.; IAvH-E • 1 ♀; Puerto Solano, PNN Serranía de Chiribiquete, Río Amú; 0 ° 12 ′ 42 ″ N, 72 ° 25 ′ 3 ″ W; elev. 250 m; 21 Mar. 2001; M. Ospina and E. González leg.; pitfall human dung; UPTC-CE • 1 ♂, 1 ♀; Puerto Solano, PNN Serranía de Chiribiquete, Río Cuñare; 0 ° 20 ′ N, 72 ° 37 ′ 50 ″ W; elev. 250 m; 2 – 4 Nov. 2000; M. Ospina and E. González leg.; pitfall human dung; UPTC-CE • 4 ♂♂, 1 ♀; same data as for preceding; IAvH-E • 2 ♂♂, 4 ♀♀; same data as for preceding; 0 ° 31 ′ 3 ″ N, 72 ° 38 ′ 27.7 ″ W; 10 – 12 Nov. 2000; IAvH-E • 1 ♂, 1 ♀; same data as for preceding; 0 ° 32 ′ 4 ″ N, 72 ° 37 ′ 57.5 ″ W; 19 – 21 Nov. 2000; IAvH-E • 1 ♂; same data as for preceding; Río Cuñare - Amú; 0 ° 12 ′ 34.5 ″ N, 72 ° 24 ′ 55.1 ″ W; 15 – 17 Feb. 2001; IAvH-E • 2 ♂♂, 2 ♀♀; same data as for preceding; 0 ° 12 ′ 42.8 ″ N, 72 ° 25 ′ 3.2 ″ W; 21 Feb. 2001; IAvH-E • 1 ♂, 1 ♀; same data as for preceding; 0 ° 12 ′ 47.8 ″ N, 72 ° 25 ′ 25.4 ″ W; 1 – 3 Mar. 2001; IAvH-E • 1 ♂, 3 ♀♀; Puerto Solano, PNN Serranía de Chiribiquete, Río Mesay; 0 ° 15 ′ 38 ″ N, 72 ° 55 ′ 24 ″ W; elev. 250 m; 24 Jan. 2000; F. Quevedo leg.; pitfall human dung; IAvH-E • 2 ♂♂, 1 ♀; Puerto Solano, PNN Serranía de Chiribiquete, Río Sararamano; 0 ° 09 ′ 26 ″ N, 72 ° 37 ′ 45 ″ W; elev. 250 m; 11 Apr. 2000; E. González leg.; pitfall human dung; UPTC-CE • 2 ♀♀; same data as for preceding; IAvH-E • 1 ♂, 2 ♀♀; same data as for preceding; flight interception trap; UPTC-CE • 2 ♀♀; same data as for preceding; 0 ° 10 ′ 02 ″ N, 72 ° 36 ′ 35 ″ W; 8 Apr. 2000; pitfall human dung; IAvH-E • 1 ♂, 2 ♀♀; same data as for preceding; 4 Apr. 2000; flight interception trap; IAvH-E • 1 ♂, 2 ♀♀; same data as for preceding; 0 ° 10 ′ 47 ″ N, 72 ° 37 ′ 24 ″ W; IAvH-E • ♀; same data as for preceding; pitfall human dung; IAvH-E • 1 ♂, 1 ♀; same data as for preceding; 6 Apr. 2000; IAvH-E • 2 ♀♀; same data as for preceding; 19 Apr. 2000; IAvH-E. – Meta • 10 ♂♂, 15 ♀♀; PNN Tinigua, CIEM, Río Duda; 2 ° 40 ′ N, 74 ° 10 ′ W; elev. 350 m; Jan. 1996; I. Jiménez leg.; pitfall human dung; IAvH-E • 3 ♂♂, 6 ♀♀; Puerto López, El Naranjal; 04 ° 05 ′ 14 ″ N, 72 ° 57 ′ 30 ″ W; elev. 220 m; Feb. 1997; F. Escobar leg.; IAvH-E • 1 ♂; Puerto López, Vereda Remolino; 4 ° 11 ′ 57.8 ″ N, 72 ° 33 ′ 36.5 ″ W; elev. 2015 m; 10 Nov. 2018; pitfall human dung; D. C. Martínez leg.; IAvH-E • 1 ♀; same data as for preceding; 4 ° 15 ′ 10.1 ″ N, 72 ° 34 ′ 10.8 ″ W; elev. 210 m; IAvH-E • 1 ♀; same data as for preceding; 4 ° 12 ′ 34.5 ″ N, 72 ° 34 ′ 54.9 ″ W; elev. 201 m; 11 Nov. 2018; IAvH-E • 1 ♂; same data as for preceding; 4 ° 13 ′ 30.9 ″ N, 72 ° 34 ′ 3.2 ″ W; elev. 193 m; IAvH-E • 1 ♀; same data as for preceding; 4 ° 15 ′ 3.7 ″ N, 72 ° 34 ′ 11.5 ″ W; elev. 207 m; IAvH-E • 1 ♂; same data as for preceding; 4 ° 11 ′ 57.2 ″ N, 72 ° 33 ′ 32.8 ″ W; elev. 214 m; 12 Nov. 2018; IAvH-E • 1 ♂, 2 ♀♀; San Martín, La laguna; 3 ° 42 ′ 27.09 ″ N, 73 ° 32 ′ 32.76 ″ W; elev. 337, 4 m; I. Lina leg.; pitfall human dung; IAvH-E • 3 ♂♂, 3 ♀♀; Villavicencio; 4 ° 03 ′ 07 ″ N, 74 ° 15 ′ 57 ″ S; elev. 350 m; Mar. 1998; A. Camacho leg.; pitfall human dung; UPTC-CE • 1 ♂, 1 ♀; same data as for preceding; IAvH-E • 1 ♂, 3 ♀♀; same data as for preceding; 04 ° 02 ′ N, 75 ° 78 ′ 15 ″ W; elev. 150 m; IAvH-E • 2 ♂♂, 1 ♀; same data as for preceding; 4 ° 02 ′ N, 73 ° 15 ′ W; elev. 200 m; 19 Oct. 2009; D. López leg.; UPTC-CE • 1 ♂, 1 ♀; 33 km E of Villavicencio; 2 – 4 Mar. 1972; S. Peck and J. Peck leg.; dung trap; TAMU. – Putumayo • 1 ♂, 1 ♀; Puerto Leguizamo; 00 ° 04 ′ 20 ″ S, 74 ° 59 ′ 19 ″ W; 17 Feb. 2008; L. Jiménez leg.; CCBUMAG • 1 ♂; same data as for preceding; 00 ° 05 ′ 03 ″ S, 74 ° 59 ′ 28 ″ W; 20 Feb. 2008; L. Jiménez leg.; CCBUMAG • 1 ♂; Santa Rosa, headwaters Río San Miguel; 24 – 25 Oct. 1970; Malkin and Burchard leg.; TAMU. – Vaupés • 2 ♂♂, 4 ♀♀; Reserva Natural Mosiro-Itajura (Caparú); 1 ° 04 ′ S, 69 ° 31 ′ W, elev. 100 m; Nov. 1995; D. Forero leg.; pitfall human dung; IavH-E. ECUADOR – Pastaza • 1 ♀; Ashuara, Río Macuma, 10 km from Rio Morona; 3 elev. 00 m; 7 – 16 Jul. 1971; B. Malkin leg.; TAMU. – Pichincha • 2 ♂♂, 1 ♀; Gualea; Jun. 1929; TAMU • 1 ♀; Quito; TAMU. PERU – Amazonas • 1 ♀; Condorcanqui, Santa Maria de Nieva; 4 ° 28 ′ 23 ″ S, 77 ° 53 ′ 02 ″ W; elev. 274 m; 12 – 13 Oct. 2019; F. Paz leg.; MUSM. – Cusco • 5 ♂♂, 3 ♀♀; La Convención, Megantoni; elev. 420 – 442 m; 13 Mar. 2022; S. Bejar and B. Cocamareshi leg.; MUSM • 1 ♂; same data as for preceding; 20 Jun. 2023; R. Curo leg.; MUSM. – Huánuco • 1 ♂; Leoncia Prado Tingo Maria, Monson Valley; 20 Nov. 1954; E. I. Schlinger and E. S. Ross leg.; TAMU. – Loreto • 1 ♀; Datem del Marañón; 4 ° 59 ′ 44 ″ S, 76 ° 18 ′ 25 ″ W; elev. 132 m; 3 Jul. 2012; C. Espinoza leg.; MUSM • 1 ♀; Loreto; 4 ° 38 ′ 23 ″ S, 76 ° 10 ′ W; elev. 114 m; 30 Jun. 2012; C. Espinoza leg.; MUSM • 1 ♂, 1 ♀; Río Ungumayo; elev. 137 m; 29 – 30 Nov. 2009; L. Sulca leg.; MUSM. – Madre de Dios • 5 ♂♂, 5 ♀♀; CICRA Field Station; 12 ° 33 ′ 09 ″ S, 70 ° 06 ′ 36 ″ W; elev. 295 m; 10 – 15 Jul. 2010; Chaboo team leg.; flight interception trap; MUSM • 2 ♂♂, 2 ♀♀; same data as for preceding; 12 ° 33 ′ 07 ″ S, 70 ° 06 ′ 34 ″ W; 9 – 13 Jun. 2011; MUSM • 1 ♀; Nueva Arequipa; elev. 231 m; 23 – 27 Sep. 2009; C. Peña and E. Martínez leg.; MUSM • 1 ♂, 1 ♀; PN Bahuaja-Sonene; 13 ° 11 ′ 38 ″ S, 70 ° 07 ′ 58 ″ W; elev. 347 m; 3 – 19 Jun. 2013; J. Grados leg.; MUSM • 2 ♀♀; PNN Manú, Estación Biológica Pantiacolla; 12 ° 38 ′ 48 ″ S, 71 ° 14 ′ 36 ″ W; 2010; S. Pérez leg.; UPTC-CE • 1 ♂, 1 ♀; Reserva Comunal Amarakaeri; naciente Qda. Pinquiri; 12 ° 55 ′ 43 ″ S, 70 ° 51 ′ 41 ″ W; elev. 421 m; 4 Jun. 2011; M. Vilchez leg.; MUSM • 1 ♂, 1 ♀; Rio Tambopata; Collpa de Guacamayos; 13 ° 8.5 ′ S, 69 ° 36.4 ′ W; elev. 300 m; Oct. 1995; F. Forsyth leg.; MUSM • 1 ♀; Río Tambopata; RR. NN. Explorer’s inn.; 12 ° 50 ′ 30 ″ S, 69 ° 17 ′ 31 ″ W; elev. 161 m; 15 – 18 May 2019; L. Figueroa and M. Alvarado leg.; MUSM.	en	Bordin, Bruna Ribeiro, Arias-Buriticá, Jorge, Maldaner, María Eduarda, Vaz-De-Mello, Fernando Z. (2026): Taxonomic revision of the Phanaeus chalcomelas species group and a description of the new Phanaeus achilles species group (Coleoptera: Scarabaeidae: Scarabaeinae: Phanaeini). European Journal of Taxonomy 1080: 1-65, DOI: 10.5852/ejt.2026.1080.3310, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3310/14554
03B6F8059632FFB2B938FE861F616783.taxon	distribution	Distribution Widely distributed in the Amazon, from Colombia and Bolivia to French Guiana and northeastern Pará, in Brazil (Fig. 21). In lowlands, below 500 m a. s. l.	en	Bordin, Bruna Ribeiro, Arias-Buriticá, Jorge, Maldaner, María Eduarda, Vaz-De-Mello, Fernando Z. (2026): Taxonomic revision of the Phanaeus chalcomelas species group and a description of the new Phanaeus achilles species group (Coleoptera: Scarabaeidae: Scarabaeinae: Phanaeini). European Journal of Taxonomy 1080: 1-65, DOI: 10.5852/ejt.2026.1080.3310, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3310/14554
03B6F805962BFFBCB987FC441EAE6668.taxon	description	urn: lsid: zoobank. org: act: 16 D 03340 - F 1 FA- 4 E 1 D-B 16 F-FF 9 FEA 86 A 94 D Figs 2 D – E, 3 D, 4 B, 8, 13 C, 14 B, E, H, K, 16 B, 17 B, 18 B, 19 B	en	Bordin, Bruna Ribeiro, Arias-Buriticá, Jorge, Maldaner, María Eduarda, Vaz-De-Mello, Fernando Z. (2026): Taxonomic revision of the Phanaeus chalcomelas species group and a description of the new Phanaeus achilles species group (Coleoptera: Scarabaeidae: Scarabaeinae: Phanaeini). European Journal of Taxonomy 1080: 1-65, DOI: 10.5852/ejt.2026.1080.3310, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3310/14554
03B6F805962BFFBCB987FC441EAE6668.taxon	diagnosis	Diagnosis This new species is diagnosed by the following combination of character states: venter bright green color (Fig. 8 C); tip of anteromedial prominence of metaventrite convex in lateral view, usually with irregular and small calluses or carinulae (Fig. 2 D – E); pronotum with deep posteromedial fossae with punctations inside (Figs 4 B, 8 A – B); pronotal punctures strong, clearly visible over entire pronotal surface under 20 × magnification; carina adjacent to anteromedial pronotal margin of females appearing regularly arched in frontal view (Fig. 3 D); major males with elongate triangular horn and well-developed, pyramidal pronotal tubercles (Fig. 8 D).	en	Bordin, Bruna Ribeiro, Arias-Buriticá, Jorge, Maldaner, María Eduarda, Vaz-De-Mello, Fernando Z. (2026): Taxonomic revision of the Phanaeus chalcomelas species group and a description of the new Phanaeus achilles species group (Coleoptera: Scarabaeidae: Scarabaeinae: Phanaeini). European Journal of Taxonomy 1080: 1-65, DOI: 10.5852/ejt.2026.1080.3310, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3310/14554
03B6F805962BFFBCB987FC441EAE6668.taxon	etymology	Etymology The specific epithet refers to the type locality, Serra do Divisor National Park.	en	Bordin, Bruna Ribeiro, Arias-Buriticá, Jorge, Maldaner, María Eduarda, Vaz-De-Mello, Fernando Z. (2026): Taxonomic revision of the Phanaeus chalcomelas species group and a description of the new Phanaeus achilles species group (Coleoptera: Scarabaeidae: Scarabaeinae: Phanaeini). European Journal of Taxonomy 1080: 1-65, DOI: 10.5852/ejt.2026.1080.3310, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3310/14554
03B6F805962BFFBCB987FC441EAE6668.taxon	materials_examined	Type material (9 ♂♂, 10 ♀♀) Holotype BRAZIL • ♂; Acre, Mâncio Lima, Parque Nacional da Serra do Divisor; 7 ° 26 ′ 46 ″ S, 73 ° 39 ′ 27 ″ W; 7 – 9 Dec. 2019; R. A. Azevedo leg.; pitfall human dung; plotAP 06; CEMT. Paratypes BRAZIL – Acre • 6 ♂♂, 9 ♀♀; same data as for holotype; CEMT • 1 ♂, 1 ♀; same data as for holotype; TAMU • 1 ♂; same data as for holotype; 7 ° 26 ′ 46 ″ S, 73 ° 39 ′ 28 ″ W; 11 – 15 Dec. 2019; flight interception trap; CEMT. Type locality Brazil, Acre, Mâncio Lima, Serra do Divisor National Park.	en	Bordin, Bruna Ribeiro, Arias-Buriticá, Jorge, Maldaner, María Eduarda, Vaz-De-Mello, Fernando Z. (2026): Taxonomic revision of the Phanaeus chalcomelas species group and a description of the new Phanaeus achilles species group (Coleoptera: Scarabaeidae: Scarabaeinae: Phanaeini). European Journal of Taxonomy 1080: 1-65, DOI: 10.5852/ejt.2026.1080.3310, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3310/14554
03B6F805962BFFBCB987FC441EAE6668.taxon	description	Description Major male (holotype) MEASUREMENTS. Length 18.5 mm. Width 10.3 mm. HEAD. Clypeus bidentate with striated sculpture (Fig. 8 A). Anterior margin of clypeus and horn black (Fig. 8 A). Olive brown on paraocular area and posterior portion of clypeus (Figs 8 A, 16 B). Cephalic horn triangular, long, curved posteriorly over pronotum (Figs 8 A, 16 B, 17 B). Discrete sheen around the eyes and at apex of horn (Fig. 8 A). PRONOTUM. Olive brown with irregularly black elevations (Fig. 8 A). Punctures strong, clearly visible over entire pronotal surface under 20 × magnification. Disc triangular, flattened, with two well-developed, pyramidal anterior tubercles (Fig. 8 D). Carina adjacent to anteromedial pronotal margin elevated (Fig. 8 D). Lateral pronotal expansion of pronotum slightly curved posteriorly in dorsal view (Fig. 8 A). Posteromedial fossae semicircular with punctations inside (Figs 4 B, 8 A). Discrete sheen on entire pronotal margin (Fig. 8 A). ELYTRA. Striae punctate, basally impressed as distinct fossae; each strial puncture within a black macula (Fig. 8 A). Interstriae olive brown with scabriculous surface giving opaque appearance. Anterior margin with discrete sheen (Fig. 8 A). METAVENTRITE. Tip of anteromedial prominence of metaventrite convex in lateral view, with irregular carinulae (Fig. 2 D – E). Surface with large, deep, and dense punctures (Fig. 2 D). Black color in the central portion; green color near the insertion of the mesocoxae and metacoxae (Fig. 8 C). PYGIDIUM. Almost completely brown, except for green anterior portion (Fig. 18 B). Scabriculous surface with lateral waves and a smooth medial area, with strong punctures. ENDOPHALLITES. ME in dorsal view elongate and blade-like, with a broad, rounded basal portion tapering gradually into a narrow, curved apex; left margin slightly concave, right margin arched, convex; left margin with a pronounced fold, occupying the basal half portion (Fig. 14 E). AE with tapered base; right margin (dorsal view) sinuous, with a small tooth-like projection near the base, more visible in lateral view (Fig. 14 H). SRP wider than long (Fig. 14 K). Variation Dorsal color varies from reddish brown to olive green, with iridescence between green and brown in some specimens. Male sizes: length 16 – 18.5 mm, width 9 – 11.3 mm. Female sizes: length 15.4 – 17.6 mm, width 9.6 – 10.9 mm. Females: head with transversal carina (Figs 3 D, 8 B); pronotum with three tubercles arranged in V, two anterior rounded and one posterior elongated longitudinally (Figs 8 B, 19 B); carina adjacent to anteromedial pronotal margin appearing regularly arched in frontal view (Fig. 3 D); posteromedial pronotal fossae triangular (Fig. 8 B). Minor males: short horn or without horn; if hornless, only with a transverse carina on the head; triangular pronotal disc, but without lateral expansions and diminute pair of anterior pronotal tubercles.	en	Bordin, Bruna Ribeiro, Arias-Buriticá, Jorge, Maldaner, María Eduarda, Vaz-De-Mello, Fernando Z. (2026): Taxonomic revision of the Phanaeus chalcomelas species group and a description of the new Phanaeus achilles species group (Coleoptera: Scarabaeidae: Scarabaeinae: Phanaeini). European Journal of Taxonomy 1080: 1-65, DOI: 10.5852/ejt.2026.1080.3310, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3310/14554
03B6F805962BFFBCB987FC441EAE6668.taxon	distribution	Distribution Brazil, Acre, Serra do Divisor National Park (Fig. 20). As shown for another dung beetle originally described exclusively from the Brazilian side of the Divisor Range (Arias-Buriticá et al. 2023, 2024), this species is also likely present on the Peruvian side and perhaps in other areas further within this country.	en	Bordin, Bruna Ribeiro, Arias-Buriticá, Jorge, Maldaner, María Eduarda, Vaz-De-Mello, Fernando Z. (2026): Taxonomic revision of the Phanaeus chalcomelas species group and a description of the new Phanaeus achilles species group (Coleoptera: Scarabaeidae: Scarabaeinae: Phanaeini). European Journal of Taxonomy 1080: 1-65, DOI: 10.5852/ejt.2026.1080.3310, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3310/14554
03B6F805962BFFBCB987FC441EAE6668.taxon	discussion	Taxonomic discussion Although the external morphology and the parameres of Phanaeus divisor sp. nov. are similar to P. cambeforti (Figs 5, 8, 13 A, C), differences in the external morphology and endophallites observed between the two taxa gave us the confidence to describe this new species from Serra do Divisor National Park. Externally, they can be easily distinguished by the punctations inside of posteromedial pronotal fossae in P. divisor, absent in P. cambeforti. In P. divisor, the anteromedial prominence of the metaventrite terminates bears a convex tip in lateral view, with distinct carinae. In contrast, P. cambeforti exhibits a flat tip of the anteromedial prominence of the metaventrite, without carinae. In addition, the ME and AE differ in shape and size (Fig. 14 D – E, G – H).	en	Bordin, Bruna Ribeiro, Arias-Buriticá, Jorge, Maldaner, María Eduarda, Vaz-De-Mello, Fernando Z. (2026): Taxonomic revision of the Phanaeus chalcomelas species group and a description of the new Phanaeus achilles species group (Coleoptera: Scarabaeidae: Scarabaeinae: Phanaeini). European Journal of Taxonomy 1080: 1-65, DOI: 10.5852/ejt.2026.1080.3310, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3310/14554
03B6F8059625FFBEB95FFDE0188F61BF.taxon	materials_examined	Figs 1 H, 2 B, 9, 13 E, 14 C, F, I, L, 16 A, 17 A, 18 A, 19 A	en	Bordin, Bruna Ribeiro, Arias-Buriticá, Jorge, Maldaner, María Eduarda, Vaz-De-Mello, Fernando Z. (2026): Taxonomic revision of the Phanaeus chalcomelas species group and a description of the new Phanaeus achilles species group (Coleoptera: Scarabaeidae: Scarabaeinae: Phanaeini). European Journal of Taxonomy 1080: 1-65, DOI: 10.5852/ejt.2026.1080.3310, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3310/14554
03B6F8059625FFBEB95FFDE0188F61BF.taxon	diagnosis	Diagnosis Dorsal and ventral color similar, weakly lustrous, dark blue or green with faded sheen (Fig. 9). Triangular clypeal teeth (Fig. 9). Cephalic horn of major males expanded laterally at base and gradually narrowing towards apex (Fig. 16 A). Pronotum of females with three tubercles arranged in a triangle, with the lateral tubercles longer latitudinally than the middle one, which is rounded (Figs 9 B, D, 19 A). Variation: length: 19 – 24 mm; the coloration varies from purplish blue to emerald green; pronotum and elytra may show different colors, one blue, the other green. Endophallites: ME in dorsal view strongly curved, hookshaped, with a broad basal region and a long, narrow, apically tapering process directed anteriorly; basal and right margin together form a single, strongly convex arch; left margin with a boomerang-shaped fold, occupying the basal half (Fig. 14 F). AE long, with U-shaped base; right margin (dorsal view) straight, with a small tooth-like projection in the middle, more visible in lateral view (Fig. 14 I). SRP circular formed by two sclerotized parts connected by a membrane (Fig. 14 L).	en	Bordin, Bruna Ribeiro, Arias-Buriticá, Jorge, Maldaner, María Eduarda, Vaz-De-Mello, Fernando Z. (2026): Taxonomic revision of the Phanaeus chalcomelas species group and a description of the new Phanaeus achilles species group (Coleoptera: Scarabaeidae: Scarabaeinae: Phanaeini). European Journal of Taxonomy 1080: 1-65, DOI: 10.5852/ejt.2026.1080.3310, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3310/14554
03B6F8059625FFBEB95FFDE0188F61BF.taxon	materials_examined	Type material Holotype of Phanaeus lecourti BOLIVIA • ♂; “ (LA PAZ) N. Yungas, Caranavi-Carasco, 700 - 1200 m; 11.91; G. Lecourt leg. ”; CPFA. Paratypes of Phanaeus lecourti BOLIVIA • 1 ♂, 1 ♀; (examined from photographs provided in Arnaud 2002: 87). Holotype of Phanaeus lecourti peruanus PERU • ♂; “ (HUA), Pucalpa, III. 76; Monguillon leg. ”; CPFA. Paratypes of Phanaeus lecourti peruanus PERU • 1 ♂, 1 ♀; (examined from photographs provided in Arnaud 2002: 87). Non-type material examined (25 ♂♂, 19 ♀♀) BOLIVIA – Cochabamba • 1 ♂, 1 ♀; Huayllas; Dec. 2006; CEMT. – La Paz • 1 ♂; Nor Yungas, rt. Caranavi-Carrasco; P. Arnaud leg.; CEMT • 1 ♂; Nor Yungas, Cotapata; 19 – 30 Sep. 2000; Y. Gareca and P. Blacut leg.; MHNNKM • 1 ♀; Nor Yungas, E. B. T.; 30 May 2001; P. Coro leg.; MHNNKM. PERU – Ayacucho • 2 ♂♂; Moyobamba; 13 ° 05 ′ 14 ″ S, 73 ° 30 ′ 23 ″ W; elev. 1918 m; 28 Feb. – 1 Mar. 2022; M. Alvarado leg.; MUSM. – Cusco • 1 ♂; La Convención, Encharate; 12 ° 32 ′ 40 ″ S, 73 ° 8 ′ 10 ″ W; elev. 1600 m; 21 Aug. 2022; E. Quispe leg.; MUSM • 1 ♂, 3 ♀♀; same data as for preceding; CC. Ozanampiato; 12 ° 39 ′ 38 ″ S, 73 ° 09 ′ 18 ″ W; elev. 1708 m; 22 – 24 Jan. 2010; C. Carranza and C. Rossi leg.; MUSM • 1 ♀; same data as for preceding; 12 ° 39 ′ 32 ″ S, 73 ° 09 ′ 32 ″ W, elev. 1681 m; MUSM • 5 ♂♂, 4 ♀♀; same data as for preceding; CC. Santa Rosa; 12 ° 34 ′ 15 ″ S, 73 ° 05 ′ 10 ″ W; elev. 1693 m; 26 – 29 Jan. 2010; MUSM • 2 ♂♂, 1 ♀; same data as for preceding; elev. 1685 m; MUSM • 1 ♀; same data as for preceding; 12 ° 34 ′ 10 ″ S, 73 ° 05 ′ 15 ″ W; elev. 1693 m; 18 – 21 Sep. 2010; M. Alvarado and J. Peralta leg.; pitfall; MUSM • 1 ♂; same data as for preceding; 12 ° 34 ′ 12 ″ S, 73 ° 05 ′ 13 ″ W; elev. 1709 m; MUSM • 1 ♀; San Pedro; 13 ° 03 ′ 15 ″ S, 71 ° 32 ′ 54 ″ W; elev. 300 m; 19 – 20 Nov. 1999; T. Larsen leg.; pitfall human dung; MUSM • 1 ♂; Union, along road; 13 ° 5 ′ 24 ″ S, 71 ° 34 ′ 53 ″ W; elev. 1900 m; Dec. 1996; N. Quinte leg.; cloud forest; pitfall human dung; CEMT. – Huánuco • 1 ♀; Leoncio Prado Tingo Maria; May 2008; CEMT • 1 ♂, 1 ♀; PN Cordillera Azul; 39 km NE of Tingo Maria; elev. 1700 m; 11 – 14 Jan. 1983; A. Newton and M. Thayer leg.; pitfall human dung; TAMU. – Junín • 2 ♂♂, 1 ♀; Calabaza; elev. 2000 m; 1 – 20 Sep. 2004; Callegari leg.; TAMU • 2 ♂♂, 1 ♀; near Satipo; elev.> 1000 m; Jul. 2004; A. Santibáñez leg.; pitfall human dung; TAMU • 2 ♂♂; Santuario Nacional Pampa Hermosa; 10 ° 59 ′ 51 ″ S, 75 ° 25 ′ 35 ″ W; elev. 1879 m; 30 May 2011; D. Silva leg.; TAMU • 1 ♂; Satipo; May 2005; TAMU • 1 ♂; Satipo; elev. 600 m; 3 – 16 Apr. 2003; A. Santibáñez leg.; TAMU • 1 ♂; Satipo, Huerto Edén; Dec. 2004; CEMT • 1 ♂; same data as for preceding; Apr. 2007; CEMT • 1 ♂; same data as for preceding; elev. 1400 m; Dec. 2007; CEMT • 1 ♀; Satipo, Puerto Ocopa; elev. 450 m; Jul. 2004; CEMT • 1 ♀; Satipo, Santos Atahualpa, Río Tambo; 12 Dec. 2007; R. Poerster leg.; CEMT • 2 ♂♂; Satipo; Nov. 2004; CEMT. Type locality Bolivia, La Paz.	en	Bordin, Bruna Ribeiro, Arias-Buriticá, Jorge, Maldaner, María Eduarda, Vaz-De-Mello, Fernando Z. (2026): Taxonomic revision of the Phanaeus chalcomelas species group and a description of the new Phanaeus achilles species group (Coleoptera: Scarabaeidae: Scarabaeinae: Phanaeini). European Journal of Taxonomy 1080: 1-65, DOI: 10.5852/ejt.2026.1080.3310, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3310/14554
03B6F8059625FFBEB95FFDE0188F61BF.taxon	distribution	Distribution Yungas Province in Bolivia and Peru (Fig. 20) at elevations between 400 and 1100 m. a. s. l.	en	Bordin, Bruna Ribeiro, Arias-Buriticá, Jorge, Maldaner, María Eduarda, Vaz-De-Mello, Fernando Z. (2026): Taxonomic revision of the Phanaeus chalcomelas species group and a description of the new Phanaeus achilles species group (Coleoptera: Scarabaeidae: Scarabaeinae: Phanaeini). European Journal of Taxonomy 1080: 1-65, DOI: 10.5852/ejt.2026.1080.3310, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3310/14554
03B6F8059627FFBBB952FA481F4C6061.taxon	description	urn: lsid: zoobank. org: act: 22700 B 47 - 85 BB- 4 C 1 F- 91 B 5 - 83 D 6 CF 3 DFAA 2 Figs 2 C, 4 C, 10, 13 F, 15 C, F, I, L, 16 F, 17 F, 18 F, I, 19 F	en	Bordin, Bruna Ribeiro, Arias-Buriticá, Jorge, Maldaner, María Eduarda, Vaz-De-Mello, Fernando Z. (2026): Taxonomic revision of the Phanaeus chalcomelas species group and a description of the new Phanaeus achilles species group (Coleoptera: Scarabaeidae: Scarabaeinae: Phanaeini). European Journal of Taxonomy 1080: 1-65, DOI: 10.5852/ejt.2026.1080.3310, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3310/14554
03B6F8059627FFBBB952FA481F4C6061.taxon	diagnosis	Diagnosis This species is diagnosed by the following combination of character states: ventral color iridescent greenish red (Fig. 10 C); dorsum dull, dark olive green to dark olive brown (Fig. 10 A – B, D); posteromedial pronotal fossae always present (Figs 4 C, 10 A – B, D); pair of anterior pronotal tubercles of males small, distinguishable from the black elevations by the bright spot (Figs 10 A, D, 16 F); semicircular clypeal teeth in females (Fig. 10 B).	en	Bordin, Bruna Ribeiro, Arias-Buriticá, Jorge, Maldaner, María Eduarda, Vaz-De-Mello, Fernando Z. (2026): Taxonomic revision of the Phanaeus chalcomelas species group and a description of the new Phanaeus achilles species group (Coleoptera: Scarabaeidae: Scarabaeinae: Phanaeini). European Journal of Taxonomy 1080: 1-65, DOI: 10.5852/ejt.2026.1080.3310, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3310/14554
03B6F8059627FFBBB952FA481F4C6061.taxon	etymology	Etymology This species is dedicated to Mario Jardim Cupello in recognition of his significant contributions to the knowledge of Scarabaeidae.	en	Bordin, Bruna Ribeiro, Arias-Buriticá, Jorge, Maldaner, María Eduarda, Vaz-De-Mello, Fernando Z. (2026): Taxonomic revision of the Phanaeus chalcomelas species group and a description of the new Phanaeus achilles species group (Coleoptera: Scarabaeidae: Scarabaeinae: Phanaeini). European Journal of Taxonomy 1080: 1-65, DOI: 10.5852/ejt.2026.1080.3310, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3310/14554
03B6F8059627FFBBB952FA481F4C6061.taxon	materials_examined	Type material (13 ♂♂, 31 ♀♀) Holotype BRAZIL • ♂; Mato Grosso, Araputanga, Fazenda Araputanga; 15 ° 23 ′ 15 ″ S, 58 ° 24 ′ 43 ″ W; elev. 298 m; 7 – 9 Jan. 2024; F. Z. Vaz-de-Mello, M. E. Maldaner and B. R. Bordin leg.; pitfall human dung; CEMT. Paratypes BRAZIL – Mato Grosso • 6 ♂♂, 13 ♀♀; Araputanga, Fazenda Bandeirantes; 15 ° 23 ′ 26 ″ S, 58 ° 24 ′ 48 ″ W; elev. 296 m; 18 – 20 Jan. 2013; R. J. Silva leg.; pitfall human + pig dung; CEMT • 1 ♂, 2 ♀♀; same data as for preceding; TAMU • 1 ♀; same data as for preceding; 15 ° 23 ′ 06 ″ S, 58 ° 26 ′ 02 ″ W; elev. 308 m; 19 – 21 Jan. 2013; CEMT • 2 ♂♂; same data as for preceding; 15 ° 23 ′ 04 ″ S, 58 ° 26 ′ W; elev. 296 m; 21 – 23 Jan. 2013; CEMT • 1 ♀; same data as for preceding; 15 ° 22 ′ 18 ″ S, 58 ° 26 ′ 23 ″ W; elev. 297 m; CEMT • 2 ♂♂, 8 ♀♀; Araputanga, Fazenda Sudam; 15 ° 15 ′ 22 ″ S, 58 ° 27 ′ 12 ″ W; elev. 352 m; 28 – 30 Jan. 2013; R. J. Silva leg.; pitfall human + pig dung; CEMT • 3 ♀♀; same data as for preceding; 15 ° 15 ′ 45 ″ S, 58 ° 26 ′ 23 ″ W; 29 – 31 Jan. 2013; CEMT • 2 ♂♂, 1 ♀; Indiavaí, Fazenda Alto Jaurú; 15 ° 27 ′ 18 ″ S, 58 ° 33 ′ 12 ″ W; elev. 231 m; 14 – 16 Jan. 2013; R. J. Silva leg.; pitfall human + pig dung; CEMT • 1 ♀; Mirassol d’Oeste, Fazenda Santa Helena; 15 ° 33 ′ 34 ″ S, 57 ° 58 ′ 13 ″ W; elev. 240 m; 25 – 27 Jan. 2013; R. J. Silva leg.; pitfall human + pig dung; CEMT. Type locality Brazil, Mato Grosso state, Araputanga.	en	Bordin, Bruna Ribeiro, Arias-Buriticá, Jorge, Maldaner, María Eduarda, Vaz-De-Mello, Fernando Z. (2026): Taxonomic revision of the Phanaeus chalcomelas species group and a description of the new Phanaeus achilles species group (Coleoptera: Scarabaeidae: Scarabaeinae: Phanaeini). European Journal of Taxonomy 1080: 1-65, DOI: 10.5852/ejt.2026.1080.3310, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3310/14554
03B6F8059627FFBBB952FA481F4C6061.taxon	description	Description Major male (holotype) MEASUREMENTS. Length 16.8 mm. Width 10.7 mm HEAD. Clypeus bidentate with striated sculpture (Fig. 10 A). Anterior margin of clypeus and horn black; olive green on paraocular area and posterior portion of clypeus (Fig. 10 A). Cephalic horn black, curved posteriorly over pronotum. Soft sheen around the eyes (Fig. 10 A). PRONOTUM. Disc flat, triangular in dorsal view; dark olive green, with irregularly black elevations over the whole surface (Fig. 10 A). Pronotal punctures strong, clearly visible over entire pronotal surface with 20 × magnification. Pair of anterior pronotal tubercles small, distinguishable from black rugosities by bright spot. Posteromedial fossae as entire deep circles, aligned with the first elytral striae (Figs 4 C, 10 A). ELYTRA. Dark olive green with scabriculous surface, giving an opaque appearance (Fig. 10 A). Elytral striae punctate, basally impressed as distinct fossae (Fig. 10 A). Anterior margin with bronze-green sheen, lighter than the rest of the surface (Fig. 10 A). METAVENTRITE. Anteromedial prominence long and narrow, slightly expanded at the apex (Fig. 2 C). Surface with large, deep and dense punctures. Central portion black, with a greenish-red coloration near the insertion of the mesocoxae and metacoxae, spreading to the legs (Fig. 10 C). PYGIDIUM. Olive brown, bronze in the middle; scabriculous surface with waves on the sides and smooth in the middle, with strong punctures (Fig. 18 F). ENDOPHALLITES. ME in dorsal view elongate and blade-like, with a broad, rounded basal portion tapering gradually into a narrow, curved apex; left margin with a pronounced fold, occupying the basal half portion; left margin slightly concave, with a sinuosity near apex; right margin almost straight (Fig. 15 F). AE with tapered base; right margin (dorsal view) with a diagonal projection, more visible in lateral view (Fig. 15 I). SRP square, with a diagonal projection (dorsal view) (Fig. 15 L). Variation The dorsal color varies from dark olive green to dark reddish brown. The ventral color varies from redder to greener. Male sizes: length 12.3 – 20 mm; width 8.2 – 12.6 mm. Female sizes: length 15.2 – 19.8 mm; width 8.6 – 11.5 mm. Females: head with transversal carina and semi-circular clypeal teeth (Fig. 10 B); pronotum with three small tubercles arranged in a triangle, middle tubercle posterior to lateral ones (Figs 10 B, 19 F); triangular posteromedial pronotal fossae (Fig. 10 B); pygidium of big females with a black spot on the posterior portion (Fig. 18 I); smaller females with uniformly colored pygidium, as males. Minor males: short horn or without horn; if hornless, only with a transverse carina on the head; triangular pronotal disc, but without lateral expansions and without a pair of anterior pronotal tubercles.	en	Bordin, Bruna Ribeiro, Arias-Buriticá, Jorge, Maldaner, María Eduarda, Vaz-De-Mello, Fernando Z. (2026): Taxonomic revision of the Phanaeus chalcomelas species group and a description of the new Phanaeus achilles species group (Coleoptera: Scarabaeidae: Scarabaeinae: Phanaeini). European Journal of Taxonomy 1080: 1-65, DOI: 10.5852/ejt.2026.1080.3310, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3310/14554
03B6F8059627FFBBB952FA481F4C6061.taxon	distribution	Distribution Brazil, southwestern region of the Mato Grosso state, in remnants of the Amazon rainforest (Fig. 20).	en	Bordin, Bruna Ribeiro, Arias-Buriticá, Jorge, Maldaner, María Eduarda, Vaz-De-Mello, Fernando Z. (2026): Taxonomic revision of the Phanaeus chalcomelas species group and a description of the new Phanaeus achilles species group (Coleoptera: Scarabaeidae: Scarabaeinae: Phanaeini). European Journal of Taxonomy 1080: 1-65, DOI: 10.5852/ejt.2026.1080.3310, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3310/14554
03B6F8059627FFBBB952FA481F4C6061.taxon	discussion	Taxonomic discussion Phanaeus mariocupelloi sp. nov. is similar to P. cambeforti, which has not been recorded in this region. Phanaeus cambeforti has the posteromedial fossae of pronotum shallow or absent, while P. mariocupelloi always has conspicuous posteromedial pronotal fossae. Additionally, P. cambeforti is smaller, with fewer black elevations on the pronotum of males, and with a dark green, almost black ventral color. Phanaeus mariocupelloi is the only species of the P. chalcomelas group with red ventral color that occurs in Brazil and the only species of the group with semicircular clypeal teeth in females. Although the parameres resemble P. cambeforti (Fig. 13 A, F), the endophallites are very different between the two species. The ME is very similar to P. minos (Fig. 15 E – F). Despite having a distinctive dorsal projection, the SRP has a general shape similar to P. meleagris (Fig. 15 J, L).	en	Bordin, Bruna Ribeiro, Arias-Buriticá, Jorge, Maldaner, María Eduarda, Vaz-De-Mello, Fernando Z. (2026): Taxonomic revision of the Phanaeus chalcomelas species group and a description of the new Phanaeus achilles species group (Coleoptera: Scarabaeidae: Scarabaeinae: Phanaeini). European Journal of Taxonomy 1080: 1-65, DOI: 10.5852/ejt.2026.1080.3310, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3310/14554
03B6F8059622FF84B93DFBE61DC563A4.taxon	description	Figs 2 F, 3 B, 11, 13 B, 15 A, D, G, J, 16 D, 17 D, 18 D, 19 D	en	Bordin, Bruna Ribeiro, Arias-Buriticá, Jorge, Maldaner, María Eduarda, Vaz-De-Mello, Fernando Z. (2026): Taxonomic revision of the Phanaeus chalcomelas species group and a description of the new Phanaeus achilles species group (Coleoptera: Scarabaeidae: Scarabaeinae: Phanaeini). European Journal of Taxonomy 1080: 1-65, DOI: 10.5852/ejt.2026.1080.3310, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3310/14554
03B6F8059622FF84B93DFBE61DC563A4.taxon	diagnosis	Diagnosis Elytra completely matte black, with no other color on striae punctation (Fig. 11 A – B, D). Black ventral color, except for bright red femora with green reflections (Fig. 11 C). Pronotum covered with highly anastomosed black elevations (Fig. 11 A – B, D). Pronotum with bright metallic red, yellow, and green coloration between elevations, more visible on the sides (Figs 11 A – B, D, 16 D, 17 D, 19 D). Females with carina adjacent to anteromedial pronotal margin subarcuate in frontal view, almost straight, without sinuosity in the middle (Fig. 3 B). Variation: length 16 – 21 mm. The color between the pronotal elevations can be red, yellow, and green or just red. The black elevations can almost completely cover the pronotum. Endophallites: ME in dorsal view elongate and blade-like, with a broad, rounded basal portion tapering gradually into a narrow, curved apex; left margin concave, right margin arched; left margin with a pronounced fold, occupying the basal half portion (Fig. 15 D). AE with tapered base; right margin (in dorsal view) with a tooth-like projection at the middle, more visible in lateral view; left margin, in ventral view, interrupted by the projection (Fig. 15 G). SRP rectangular with rounded angles (Fig. 15 J).	en	Bordin, Bruna Ribeiro, Arias-Buriticá, Jorge, Maldaner, María Eduarda, Vaz-De-Mello, Fernando Z. (2026): Taxonomic revision of the Phanaeus chalcomelas species group and a description of the new Phanaeus achilles species group (Coleoptera: Scarabaeidae: Scarabaeinae: Phanaeini). European Journal of Taxonomy 1080: 1-65, DOI: 10.5852/ejt.2026.1080.3310, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3310/14554
03B6F8059622FF84B93DFBE61DC563A4.taxon	materials_examined	Type material Lectotype (studied from photographs) BOLIVIA • ♂; “ long. 18.5 mm / P. meleagris, Blanch, Yungas, M. d’Orbigny / 6275 / 1276 / M. P. C. G. /. ”; MNHN. Designated by Arnaud (1982 a: 114). Paralectotypes (not studied) BOLIVIA • 3 ♂♂, 3 ♀♀; MNHN. Non-type material examined (310 ♂♂, 492 ♀♀) BOLIVIA – Chuquisaca • 2 ♂♂, 22 ♀♀; Hernando Siles, Monteagudo; 19 ° 43 ′ 35 ″ S, 63 ° 55 ′ 03 ″ W, elev. 1300 m; 18 – 21 Apr. 2004; M. Aliaga leg.; MHNNKM • 4 ♂♂, 49 ♀♀; same data as for preceding; 19 ° 46 ′ 15 ″ S, 63 ° 53 ′ 11 ″ W; 9 – 12 Apr. 2004; MHNNKM • 1 ♀; Monteagudo, Ticucha; 19 ° 96 ′ 15 ″ S, 63 ° 53 ′ 11 ″ W; elev. 1300 m; M. Aliaga leg.; pitfall human dung; TAMU. – Cochabamba • 34 ♂♂, 35 ♀♀; Ayopaya, Altamachi; 16 ° 02 ′ S, 66 ° 40 ′ W; elev. 1150 m; 25 – 27 Sep. 2004; C. Hamel leg.; MHNNKM • 1 ♂, 1 ♀; Chapare, Cristal Mayu; 17 ° 00 ′ 26 ″ S, 65 ° 38 ′ 32 ″ W; Nov. 1949; TAMU • 1 ♂; Chapare, Locotal; 1100 m; Nov. 1954; Martínez leg.; ex-Gonzalo Halffter Salas; IEXA • 1 ♀; Mosetenes, Isiboro-Securé N. P.; 16 ° 13 ′ 51 ″ S, 66 ° 25 ′ 06 ″ W; elev. 1250 m; 1 – 4 Sep. 2003; A. C. Hamel leg.; pitfall human dung; TAMU • 1 ♂; same data as for preceding; 16 ° 13 ′ 58 ″ S, 66 ° 24 ′ 54 ″ W; TAMU • 1 ♂; same data as for preceding; 14 – 17 Sep. 2003; 16 ° 14 ′ 04 ″ S, 66 ° 25 ′ 06 ″ W; TAMU • 1 ♀; Yungas del Palmar, km 135; elev. 1600 m; Feb. 1952; Martínez leg.; TAMU. – La Paz • 5 ♀♀; Caranavi; Nov. 2005; P. Wagner leg.; CEMT • 2 ♀♀; same data as for preceding; Nov. 2009; CEMT • 1 ♂; Coroico; Feb. 1984; P. Arnaval leg.; CEMT • 1 ♂; Nor Yungas, Río Coroico; elev. 1200 m; Jan. 1949; Martínez leg.; ex-Gonzalo Halffter Salas; IEXA • 1 ♂; same data as for preceding; Dec. 1948; elev. 1700 m; IEXA • 1 ♀; Nor Yungas, Tunquini Biological Station; 30 May 2001; P. Coro leg.; MHNNKM. – Santa Cruz • 9 ♂♂, 14 ♀♀; Andrés Ibáñez, Espejillos; 27 – 29 Apr. 2007; M. Amaya leg.; MHNNKM • 16 ♂♂, 24 ♀♀; Andrés Ibáñez, Laguna Volcanes; Feb. 2010; T. Vidaurre leg.; MHNNKM • 4 ♀♀; Andrés Ibáñez, Potrerillos del Güendá; 17 ° 40 ′ 21 ″ S; 63 ° 27 ′ 1 ″ W; elev. 370 m; 15 Oct. 2010; T. Vidaurre leg.; MHNNKM • 2 ♂♂, 2 ♀♀; Bermejo, Refugio Los Volcanes; 18 ° 06 ′ S, 63 ° 36 ′ W; elev. 1060 m; 10 Dec. 2003; semi-deciduous forest; D. J. Mann and A. C. Hamel leg.; pitfall human dung; CEMT • 1 ♂, 1 ♀; same data as for preceding; pitfall chicken carrion; CEMT • 2 ♂♂, 2 ♀♀; same data as for preceding; 18 ° 06 ′ S, 63 ° 35 ′ W; elev. 1045 m; 23 Feb. 2010; T. Vidaurre leg.; pitfall human dung; CEMT • 8 ♂♂, 15 ♀♀; Cordillera, Incahuasi; 13 – 16 Mar. 2008; T. Vidaurre leg.; MHNNKM • 1 ♂, 1 ♀; same data as for preceding; TAMU • 9 ♂♂, 23 ♀♀; Cordillera, Parabano; 24 – 28 Mar. 2003; J. Ledezma et al. leg.; MHNNKM • 9 ♂♂, 13 ♀♀; same data as for preceding; 23 – 27 Mar. 2004; J. L. Aramayo, et al. leg.; MHNNKM • 44 ♂♂, 33 ♀♀; same data as for preceding; 28 Sep. – 2 Oct. 2004; MHNNKM • 16 ♂♂, 25 ♀♀; Cordillera, R. M. Parabano; 19 – 30 May 2004; T. Vidaurre leg.; MHNNKM • 2 ♀♀; Cordillera, Serranía Parabado I; 13 Apr. 2002; L. Céspedes leg.; MHNNKM • 1 ♂, 1 ♀; Cordillera, Serranía Parabado II; 15 – 17 Apr. 2002; L. Céspedes leg.; MHNNKM • 36 ♂♂, 58 ♀♀; Cordillera, Yaguapoa; 19 ° 53 ′ 40.7 ″ S, 63 ° 40 ′ 25.3 ″ W; 20 – 21 Nov. 2007; T. Vidaurre leg.; MHNNKM • 17 ♂♂, 54 ♀♀; Cordillera, Yaguapoa; 19 ° 51 ′ 36 ″ S, 63 ° 40 ′ 35 ″ W; 26 Nov. 2007; T. Vidaurre leg.; MHNNKM • 2 ♂♂, 7 ♀♀; Cordillera, Yupa; 19 ° 47 ′ 47.1 ″ S, 63 ° 41 ′ 59.7 ″ W; 29 Nov. 2007; T. Vidaurre leg.; MHNNKM • 1 ♂; Florida, Alta Florida; elev. 1697 m; 16 Dec. 2011; A. Alcoba et al. leg.; MHNNKM • 13 ♂♂, 16 ♀♀; Florida, Bella Vista; 14 – 15 Nov. 2007; J. Ledezma et al. leg.; MHNNKM • 1 ♂, 1 ♀; Florida, Bella Vista; 25 Sep. 2011; A. Alcoba et al. leg.; MHNNKM • 1 ♂, 8 ♀♀; Florida, Buena Vista; 8 Mar. 2005; T. Vidaurre leg.; MHNNKM • 3 ♂♂, 1 ♀; Florida, Cuevas; 18 Nov. 2011; A. Alcoba et al. leg.; MHNNKM • 1 ♀; Florida, La Negra; 18 Sep. 2011; A. Alcoba et al. leg.; MHNNKM • 1 ♂, 1 ♀; Florida, Lagunillas; 28 Dec. 2011; A. Alcoba et al. leg.; MHNNKM • 3 ♂♂, 4 ♀♀; Florida, near Samaipata; 18 ° 8.4 ′ S, 63 ° 48.1 ′ W; elev. 1730 m; 19 Nov. 2006; W. D. Edmonds and P. Reyes leg.; pitfall human dung; TAMU • 2 ♂♂, 6 ♀♀; same data as for preceding; 18 ° 8.9 ′ S, 63 ° 48.8 ′ W; elev. 1380 m; TAMU • 2 ♂♂, 2 ♀♀; Florida, Pacay; elev. 1614 m; 18 Dec. 2011; A. Alcoba et al. leg.; MHNNKM • 1 ♂; Florida, Palermo; elev. 1467 m; 18 Dec. 2011; A. Alcoba et al. leg.; MHNNKM • 3 ♂♂, 2 ♀♀; Florida, Samaipata; 18 ° 08.4 ′ S; 63 ° 48.8 ′ W; elev. 1380 m; 19 Nov. 2006; W. D. Edmonds leg.; MHNNKM • 1 ♀; Florida, Samaipata; 26 Nov. 2011; A. Alcoba et al. leg.; MHNNKM • 1 ♂; Ichilo, Bella Vista; 18 ° 17 ′ 04 ″ S, 63 ° 40 ′ 38 ″ W; 31 Dec. 2004; T. Vidaurre leg.; MHNNKM • 4 ♂♂, 5 ♀♀; same data as for preceding; 2 – 5 Jan. 2005; T. Vidaurre leg.; MHNNKM • 10 ♂♂, 4 ♀♀; Ichilo, Elias Meneses; 9 Nov. 2003; L. Acosta leg.; MHNNKM • 5 ♂♂, 6 ♀♀; Refugio Los Volcanes; 18 ° 6.3 ′ S, 63 ° 35.9 ′ W; elev. 1050 m; 21 – 25 Feb. 2010; W. D. Edmonds and T. Vidaurre; pitfall human dung; TAMU • 2 ♂♂, same data as for preceding; 18 ° 06 ′ 19 ″ S, 63 ° 35 ′ 49 ″ W; elev. 1060 m; 6 Feb. 2003; A. C. Hamel L. and S. K. Herzog leg.; TAMU • 1 ♂; same data as for preceding; 18 ° 06 ′ 50 ″ S, 63 ° 36 ′ 15 ″ W; elev. 1200 m; 22 Apr. 2003; TAMU • 9 ♂♂, 3 ♀♀; Vallegrande, Algodonales; 18 ° 47 ′ 11 ″ S, 63 ° 50 ′ 49 ″ W; 13 Apr. 2016; M. Amaya et al. leg.; MHNNKM • 9 ♂♂, 10 ♀♀; Vallegrande, Algodonales; 18 ° 47 ′ 18 ″ S, 63 ° 50 ′ 49 ″ W; 22 Oct. 2011; A. Alcoba et al. leg.; MHNNKM • 2 ♀♀; Vallegrande, Churcos; 5 Dec. 2011; A. Alcoba et al. leg.; MHNNKM • 4 ♂♂, 2 ♀♀; Vallegrande, Churo Citano; 18 ° 51 ′ 30 ″ S, 64 ° 49 ′ 53 ″ W; elev. 1761 m; 16 Apr. 2011; A. Vidaurre leg.; MHNNKM • 6 ♀♀; Vallegrande, Islas; 18 ° 42 ′ 03 ″ S, 63 ° 58 ′ 40 ″ W; 19 Apr. 2011; A. Alcoba leg.; MHNNKM • 2 ♂♂, 1 ♀; Vallegrande, Llorenti; 5 Oct. 2011; A. Alcoba and M. Pérez leg.; MHNNKM • 1 ♂, 1 ♀; Vallegrande, Mosquera; 18 ° 42 ′ 43 ″ S, 63 ° 42 ′ 43 ″ W; 5 Oct. 2011; A. Alcoba et al. leg.; MHNNKM • 5 ♂♂, 6 ♀♀; Vallegrande, Plan Citano; 16 May 2011; T. Vidaurre leg.; MHNNKM • 1 ♀; Vallegrande, Plan Citano; 30 Aug. 2011; A. Alcoba et al. leg.; MHNNKM • 1 ♀; Vallegrande, Postrervalle, Pampas; 18 ° 27 ′ 12 ″ S, 63 ° 42 ′ 36 ″ W; elev. 1234 m; 6 Dec. 2011; A. Alcoba et al. leg.; MHNNKM • 2 ♂♂, 2 ♀♀; Vallegrande, Postrervalle, Quebrada el Palo; 1 Dec. 2011; A. Alcoba et al. leg.; MHNNKM • 1 ♂; Vallegrande, Rancho Novillero; 9 Nov. 2011; T. Vidaurre leg.; MHNNKM • 1 ♂, 1 ♀; Vallegrande, Rio Villca; 18 ° 38 ′ 39 ″ S, 63 ° 47 ′ 11 ″ W; 6 Dec. 2011; A. Alcoba et al. leg.; MHNNKM • 1 ♀; same data as for preceding; 18 Sep. 2011; MHNNKM. PERU – Cusco • 1 ♂; Quispicanchi Marcapata; TAMU. – Junín • 1 ♂; Satipo Calabaza; elev. 2000 m; 1 – 20 Sep. 2004; Callegari leg.; TAMU. Type locality Bolivia, somewhere visited by Alcide d’Orbigny in the Yungas region.	en	Bordin, Bruna Ribeiro, Arias-Buriticá, Jorge, Maldaner, María Eduarda, Vaz-De-Mello, Fernando Z. (2026): Taxonomic revision of the Phanaeus chalcomelas species group and a description of the new Phanaeus achilles species group (Coleoptera: Scarabaeidae: Scarabaeinae: Phanaeini). European Journal of Taxonomy 1080: 1-65, DOI: 10.5852/ejt.2026.1080.3310, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3310/14554
03B6F8059622FF84B93DFBE61DC563A4.taxon	distribution	Distribution Distributed along the eastern slopes of the Andes in southern Peru and Bolivia from 370 to 1700 m a. s. l. Yungas Province (Fig. 20).	en	Bordin, Bruna Ribeiro, Arias-Buriticá, Jorge, Maldaner, María Eduarda, Vaz-De-Mello, Fernando Z. (2026): Taxonomic revision of the Phanaeus chalcomelas species group and a description of the new Phanaeus achilles species group (Coleoptera: Scarabaeidae: Scarabaeinae: Phanaeini). European Journal of Taxonomy 1080: 1-65, DOI: 10.5852/ejt.2026.1080.3310, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3310/14554
03B6F8059622FF84B93DFBE61DC563A4.taxon	discussion	Remarks Differences in external morphology between Phanaeus meleagris and Phanaeus minos Erichson, 1847 stat. rev., such as the color pattern (Figs 11 D, 12 D) and the shape of the carina adjacent to the anteromedial pronotal margin in females (Fig. 3 A – B), as well as differences in male genitalia, including narrower parameres in P. meleagris compared to P. minos (Fig. 13 B, D), and distinct shapes of the endophallites (Fig. 15 D – E, G – H, J – K), gave us the confidence to return P. minos to full species status. The geographical distribution of P. meleagris has also been redefined here: it occurs on the eastern slopes of the Andes in Bolivia and southern Peru, whereas P. minos is found in Peru, Ecuador, Colombia, and Venezuela (Fig. 20).	en	Bordin, Bruna Ribeiro, Arias-Buriticá, Jorge, Maldaner, María Eduarda, Vaz-De-Mello, Fernando Z. (2026): Taxonomic revision of the Phanaeus chalcomelas species group and a description of the new Phanaeus achilles species group (Coleoptera: Scarabaeidae: Scarabaeinae: Phanaeini). European Journal of Taxonomy 1080: 1-65, DOI: 10.5852/ejt.2026.1080.3310, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3310/14554
03B6F805961EFF8FB923FE86181865F3.taxon	description	Figs 2 I, 3 A, 12, 13 D, 15 B, E, H, K, 16 E, 17 E, 18 E, 19 E	en	Bordin, Bruna Ribeiro, Arias-Buriticá, Jorge, Maldaner, María Eduarda, Vaz-De-Mello, Fernando Z. (2026): Taxonomic revision of the Phanaeus chalcomelas species group and a description of the new Phanaeus achilles species group (Coleoptera: Scarabaeidae: Scarabaeinae: Phanaeini). European Journal of Taxonomy 1080: 1-65, DOI: 10.5852/ejt.2026.1080.3310, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3310/14554
03B6F805961EFF8FB923FE86181865F3.taxon	diagnosis	Diagnosis Elytra brown, reddish brown (Fig 12 B), or olive green in the interstriae (Fig 12 A, D). Each strial puncture within a black macula (Fig 12 A – B, D). Pronotum covered with highly anastomosed black elevations (Fig 12 A – B, D). Elevations covering the tones of coppery brown or coppery red, which are more visible on the pronotum sides (Figs 12 A – B, D, 16 E, 17 E, 19 E). Dark coppery brown or black ventral color, except for the femora which are coppery red with green reflections (Fig 12 C). Female carina adjacent to anteromedial pronotal margin with sinuosity in the middle in frontal view (Fig 3 A). Variation: length 16 – 22.4 mm. In specimens from Peru, the metaventrite is black in the central portion and red near the insertion of the mesocoxae and metacoxae, spreading to the femora (Fig. 12 C). The metaventrite is completely black in specimens from Ecuador, Colombia and Venezuela, with red coloration just on the femora. Pygidium completely coppery red in most specimens (Fig. 18 E), some specimens with a black spot on the posterior portion. Endophallites: ME in dorsal view elongate and blade-like, with a broad, rounded basal portion tapering gradually into a narrow, curved apex; left margin with a pronounced fold, occupying the basal half portion; left margin slightly concave, with a sinuosity near apex; right margin almost straight (Fig. 15 E). AE with wide base; in dorsal view, left margin and basal margin form a 45 ° angle; right margin (dorsal view) almost straight, with a sinuous projection, more visible in lateral view (Fig. 15 H). SRP triangular (Fig. 15 K).	en	Bordin, Bruna Ribeiro, Arias-Buriticá, Jorge, Maldaner, María Eduarda, Vaz-De-Mello, Fernando Z. (2026): Taxonomic revision of the Phanaeus chalcomelas species group and a description of the new Phanaeus achilles species group (Coleoptera: Scarabaeidae: Scarabaeinae: Phanaeini). European Journal of Taxonomy 1080: 1-65, DOI: 10.5852/ejt.2026.1080.3310, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3310/14554
03B6F805961EFF8FB923FE86181865F3.taxon	materials_examined	Type material Lectotype (examined from photographs) PERU • ♂; “ Mountains of Peru ”; MFNB. Designated by Edmonds (1994: 31). Paralectotypes (not studied) PERU • 2 ♂♂, 1 ♀; “ Mountains of Peru ”; MFNB. Non-type material examined (39 ♂♂, 40 ♀♀) COLOMBIA – Antioquia • 1 ♀; Amalfi, Cañon del río Porce; 6 ° 45 ′ 42.43 ″ N, 75 ° 5 ′ 28.31 ″ W; elev. 1221 m; Sep. 2006; A. Bedoya leg.; IAvH-E. – Arauca • 2 ♂♂, 2 ♀♀; Tame, PNN El Cocuy; 6 ° 30 ′ 48 ″ N, 71 ° 50 ′ 15 ″ W; elev. 1000 m; 1 Oct. 2007; F. Alvarado leg.; human dung; IAvH-E. – Boyacá • 2 ♂♂; Cusiana, cerca de Pajarito, Finca Guayabetal - El paisano; 5 ° 17 ′ N, 72 ° 40 ′ W; elev. 1300 m; May 1997; F. Escobar leg.; pitfall trap; UPTC-CE • 1 ♀; same data as for preceding; 5 ° 17 ′ N, 72 ° 41 ′ W; elev. 1200 m; UPTC-CE • 1 ♂; Cusiana, cerca de Pajarito; 5 ° 23 ′ 39 ″ N, 72 ° 41 ′ 17 ″ W; elev. 1250 m; F. Escobar leg.; pitfall trap; CEMT • 1 ♂; same data as for preceding; 5 ° 17 ′ N, 72 ° 40 ′ W; elev. 1300 m; 1 Jun. 1997; IAvH-E. – Caquetá • 2 ♂♂, 1 ♀; San Vicente del Caguán, PNN Los Picachos - Alto del Río Pato; 2 ° 47 ′ 51 ″ N, 74 ° 51 ′ 18 ″ W; elev. 1500 m; Nov. – Dec. 1997; pitfall trap; F. Escobar leg.; UPTC-CE • 5 ♂♂, 5 ♀♀; same data as for preceding; 2 ° 44 ′ N, 74 º 53 ′ W; elev. 1300 – 1560 m; UPTC-CE • 1 ♂, 1 ♀; same data as for preceding; elev. 1560 m; flight interception trap; UPTC-CE • 1 ♀; same data as for preceding; 2 º 47 ′ 53 ″ N, 74 º 51 ′ 28 ″ W; elev. 1250 m; pitfall trap; UPTC-CE • 1 ♂, 1 ♀; same data as for preceding; 2 º 47 ′ N, 74 º 51 ′ W; elev. 1280 m; flight interception trap; UPTC-CE • 1 ♀; San Vicente del Caguán, camino a PNN Los Picachos; F. Escobar leg.; cow dung; UPTC-CE. – Casanare • 1 ♂; Salto de Candelas; elev. 1600 m; May 1997; F. Escobar leg.; trap; UPTC-CE. – Cauca • 1 ♀; Sn. Fortunato Bonis; TAMU. – Cundinamarca • 1 ♂, 1 ♀; Medina, Miralindo; 4 º 35 ′ 33 ″ N, 73 º 23 ′ 17 ″ W; elev. 1250 m; Feb. – Mar. 1997; F. Escobar leg.; pitfall human dung; CEMT • 1 ♀; same data as for preceding; elev. 1000 m; IAvH-E • 1 ♀; Medina, Vereda Periquito, finca Miramar, Campamento 2; 4 ° 30 ′ 55 ″ N, 73 ° 26 ′ 28 ″ W; elev. 1408 m; 14 – 16 Mar. 2018; pitfall human dung; M. I. Castro, J. C. Neita and E. Torres leg.; IAvH-E • 2 ♀♀; same data as for preceding; 4 ° 30 ′ 57.7 ″ N, 73 ° 26 ′ 3.1 ″ W; elev. 1194 m; 15 – 16 Mar. 2018; IAvH-E • 1 ♂; same data as for preceding; 4 ° 30 ′ 58.4 ″ N, 73 ° 26 ′ 5.9 ″ W; elev. 1229 m; IAvH-E • 1 ♂; same data as for preceding; 4 ° 30 ′ 59.1 ″ N, 73 ° 26 ′ 7.8 ″ W; elev. 1264 m; IAvH-E • 1 ♂; same data as for preceding; 4 ° 30 ′ 56.8 ″ N, 73 ° 26 ′ 11 ″ W; elev. 1289 m; IAvH-E • 1 ♀; same data as for preceding; 4 ° 30 ′ 58.7 ″ N, 73 ° 26 ′ 10.9 ″ W; elev. 1304 m; IAvH-E • 1 ♂, 1 ♀; same data as for preceding; 4 ° 30 ′ 59.3 ″ N, 73 ° 26 ′ 12.6 ″ W; elev. 1320 m; IAvH-E • 1 ♂, 1 ♀; same data as for preceding; 4 ° 30 ′ 57.8 ″ N, 73 ° 26 ′ 13.4 ″ W; elev. 1341 m; IAvH-E • 1 ♂; same data as for preceding; 4 ° 30 ′ 58 ″ N, 73 ° 26 ′ 14 ″ W; elev. 1346 m; IAvH-E • 1 ♀; same data as for preceding; 4 ° 30 ′ 55 ″ N, 73 ° 26 ′ 28 ″ W; elev. 1408 m; IAvH-E. – Nariño • 1 ♂; Ipiales, cuenca alta de los ríos Rumiyaco-Ranchería; elev. 1250 m; Sep. 1997; F. Escobar leg.; pitfall trap; UPTC-CE • 2 ♀♀; same data as for preceding; 0 ° 03 ′ N, 77 ° 13 ′ W; Oct. 1998; IAvH-E • 1 ♂; same data as for preceding; elev. 1430 m; 24. Sep. 1998; hand collecting at night; UPTC-CE. ECUADOR – Tungurahua • 2 ♂♂; Oriente, Abitagua; elev. 1200 m; 3 Jun. 1937; W. C. MacIntyre leg.; TAMU. – Zamora-Chinchipe • 1 ♂, 1 ♀; Chito, R. San Francisco; elev. 1540 m; 17 Feb. 2008; W. Chamorro and M. Amari leg.; pitfall trap; CEMT • 1 ♀; Libertad; 4 ° 31 ′ 39 ″ S, 79 ° 05 ′ 39 ″ W; elev. 1844 m; 28 – 30 Apr. 2023; M. Tapia leg.; pitfall human dung; LOUNAZ. PERU – Junín • 1 ♂; Calabaza; 11 º 30 ′ 12 ″ S, 74 º 48 ′ 57 ″ W; elev. 2085 m; Nov. 2004; CEMT • 1 ♂; Calabaza; elev. 2000 m; 1 – 20 Sep. 2004; Callegari leg.; TAMU • 1 ♂, 2 ♀♀; ~ 17 km N of San Ramón; 10 ° 58.8 ′ S, 75 ° 26.3 ′ W; elev. 1326 m; Mar. 2010; W. D. Edmonds and L. Figueroa leg.; human dung; TAMU • 2 ♀♀; same data as for preceding; carrion; TAMU • 1 ♂, 1 ♀; ~ 33 km S of San Ramón; 11 ° 24.5 ′ S, 75 ° 15.2 ′ W; elev. 1540 m; 6 Mar. 2010; W. D. Edmonds and L. Figueroa leg.; human dung; CEMT • 1 ♂; same data as for preceding; TAMU • 2 ♀♀; same data as for preceding; carrion; TAMU • 1 ♂, 1 ♀; Satipo; elev. 600 m; 3 – 16 Apr. 2003; A. Santibáñez leg.; TAMU • 1 ♂; Santuario Nacional Pampa Hermosa; 10 ° 59 ′ 51 ″ S, 75 ° 25 ′ 35 ″ W; elev. 1879 m; 30 May 2011; D. Silva leg.; TAMU • 1 ♂; Siete Casas, Santuario Nacional Pampa Hermosa; elev. 2220 m; 24 Oct. 2011; A. Almeyda leg.; CEMT. VENEZUELA – Táchira • 1 ♂, 1 ♀; Vía Chorro del Indio; elev. 1345 m; Aug. 1990; J. Blanco leg.; CEMT • 1 ♂; same data as for preceding; elev. 1100 m; Jun. 1982; C. F. Romero leg.; TAMU • 2 ♀♀; km 2.5 of San Cristóbal to Chorro del Indio; elev. 1100 m; 31 Oct. 1987; D. Havranek leg.; TAMU • 1 ♂; San Cristóbal; 10 – 18 Aug. 1983; B. Gill leg.; TAMU • 1 ♀; San Gil, San Cristóbal; 20 May 1983; G. Borges leg.; TAMU.	en	Bordin, Bruna Ribeiro, Arias-Buriticá, Jorge, Maldaner, María Eduarda, Vaz-De-Mello, Fernando Z. (2026): Taxonomic revision of the Phanaeus chalcomelas species group and a description of the new Phanaeus achilles species group (Coleoptera: Scarabaeidae: Scarabaeinae: Phanaeini). European Journal of Taxonomy 1080: 1-65, DOI: 10.5852/ejt.2026.1080.3310, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3310/14554
03B6F805961EFF8FB923FE86181865F3.taxon	distribution	Distribution Along the eastern slopes of the Andes in Peru, Ecuador, Colombia and Venezuela from 600 to 2100 m a. s. l. (Fig. 20).	en	Bordin, Bruna Ribeiro, Arias-Buriticá, Jorge, Maldaner, María Eduarda, Vaz-De-Mello, Fernando Z. (2026): Taxonomic revision of the Phanaeus chalcomelas species group and a description of the new Phanaeus achilles species group (Coleoptera: Scarabaeidae: Scarabaeinae: Phanaeini). European Journal of Taxonomy 1080: 1-65, DOI: 10.5852/ejt.2026.1080.3310, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3310/14554
03B6F805961EFF8FB923FE86181865F3.taxon	discussion	Remarks Phanaeus minos was considered a synonym of P. meleagris by Harold (1870), Edmonds (1994) and Edmonds & Zídek (2012). However, differences in external and genital morphology were found by us. Regarding external morphology, P. meleagris exhibits entirely black elytra, while in P. minos, only the striae are black (Figs 11 – 12). In females, the carina adjacent to the anteromedial pronotal margin is sinuous medially in P. minos (frontal view), whereas it is nearly straight and lacks medial sinuosity in P. meleagris (Fig. 3 A – B). The cephalic horn of major males is wider and more curved in P. minos than in P. meleagris (Figs 16 D – E, 17 D – E). With respect to genital morphology, the parameres are narrower in P. meleagris than in P. minos in lateral view (Fig. 13 B, D), and the medial, superior right peripheral, and additional endophallites exhibit distinct shapes — differences in the additional endophallite were also previously noted by Manjarres-H. & Molano-R. (2015) (Fig. 15 D – E, G – H, J – K).	en	Bordin, Bruna Ribeiro, Arias-Buriticá, Jorge, Maldaner, María Eduarda, Vaz-De-Mello, Fernando Z. (2026): Taxonomic revision of the Phanaeus chalcomelas species group and a description of the new Phanaeus achilles species group (Coleoptera: Scarabaeidae: Scarabaeinae: Phanaeini). European Journal of Taxonomy 1080: 1-65, DOI: 10.5852/ejt.2026.1080.3310, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3310/14554
03B6F8059617FF88B950FE8619196668.taxon	diagnosis	Diagnosis of the Phanaeus achilles species group Anteromedial margin of clypeus weakly bidentate (Fig. 1 B). Head of females with transverse carina with two elevations in the middle, giving a bituberculate appearance in frontal view (Fig. 1 E). Pronotum bearing contrasted black elevations, variously and irregularly shaped (Fig. 1 G). Pronotum of females with more elevations and those more anastomosed than in males (Fig. 22). Pronotum of females with anterior W-shaped carina (Fig. 1 E). Anteromedial pronotal portion of males with two longitudinal curved carinae, each forming a large posterior tubercle (Figs 23 A – B, 25). Pronotal disc of major males with posterior angles expanded upwards and backwards, the apices of which are curved towards center of pronotum (Figs 23 A – B, 25 A – B). Posteromedial pronotal fossae present (Fig. 22). Elytral striae punctate, basally impressed as distinct fossae (Fig. 1 C). Front tibiae tridentate, with faint, almost imperceptible fourth tooth. Dorsum completely shiny metallic (except for black elevations), red, red with greenish iridescence, green, or bronze green. Occurrence in dry forests and shrublands in southwestern Ecuador and northwestern Peru (Fig. 27).	en	Bordin, Bruna Ribeiro, Arias-Buriticá, Jorge, Maldaner, María Eduarda, Vaz-De-Mello, Fernando Z. (2026): Taxonomic revision of the Phanaeus chalcomelas species group and a description of the new Phanaeus achilles species group (Coleoptera: Scarabaeidae: Scarabaeinae: Phanaeini). European Journal of Taxonomy 1080: 1-65, DOI: 10.5852/ejt.2026.1080.3310, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3310/14554
03B6F8059611FF94B94CFDE018FA669E.taxon	description	Figs 1 B, E, G, 22 A – B, 23 A, C, E, 24 A, C, 25 A, 26 A, C, E, G, 29 A	en	Bordin, Bruna Ribeiro, Arias-Buriticá, Jorge, Maldaner, María Eduarda, Vaz-De-Mello, Fernando Z. (2026): Taxonomic revision of the Phanaeus chalcomelas species group and a description of the new Phanaeus achilles species group (Coleoptera: Scarabaeidae: Scarabaeinae: Phanaeini). European Journal of Taxonomy 1080: 1-65, DOI: 10.5852/ejt.2026.1080.3310, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3310/14554
03B6F8059611FF94B94CFDE018FA669E.taxon	diagnosis	Diagnosis Dorsum completely shiny metallic red or red with greenish iridescence (Figs 22 A – B, 23 A, C, E). Venter dark green except for black central portion of metaventrite. Pronotum bearing contrasted black elevations, variously and irregularly shaped (Figs 1 G, 22 A – B, 23 A, E, 24 A). Anteromedial pronotal region of males with two longitudinal curved carinae, each forming a large posterior sub-quadrate tubercle (Figs 23 A, 24 A, C). Pronotum of females with anterior W-shaped carina (Figs 1 E, 23 E). Elytral striae punctate, basally impressed as distinct fossae (Fig. 22 A – B). Length 15 – 22.5 mm. Endophallites: ME with a large tab, given triangular appearance in dorsal and ventral view; margins of tab sinuous with a lateral rounded C-shaped angle in ventral view (Fig. 26 C). AE in dorsal view, triangular, with a concave left margin and convex right margin; left margin with lateral projection near the apex (Fig. 26 E). SRP almost horizontal guttiform, with rounded left margin and sharp right margin (dorsal view) (Fig. 26 G).	en	Bordin, Bruna Ribeiro, Arias-Buriticá, Jorge, Maldaner, María Eduarda, Vaz-De-Mello, Fernando Z. (2026): Taxonomic revision of the Phanaeus chalcomelas species group and a description of the new Phanaeus achilles species group (Coleoptera: Scarabaeidae: Scarabaeinae: Phanaeini). European Journal of Taxonomy 1080: 1-65, DOI: 10.5852/ejt.2026.1080.3310, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3310/14554
03B6F8059611FF94B94CFDE018FA669E.taxon	materials_examined	Type material Lectotype of Phanaeus achilles (here designated) ECUADOR • ♂; “ Puna / Kinb. / Riksmuseum Stockholm / 6056 E 92 + / 50 84 ”; NRMS. Paralectotype of Phanaeus achilles ECUADOR • 1 ♂; “ Puna / Kinb. / Phanaeus Mac’Leay / achilles Boh. / Riksmuseum Stockholm / 51 84 ”; NRMS. Holotype of Phanaeus achilles lydiae ECUADOR • ♂; “ Guayas, Los Ceibos Guayaquil; 28 Nov. 1987 ”; CPFA. Paratypes of Phanaeus achilles lydiae (25 ♂♂, 32 ♀♀) ECUADOR – Guayaquil • 1 ♀; same data as for holotype; CPFA • 3 ♂♂, 3 ♀♀; Gua. env. Cerecita. Rte Salinas; Feb. 1990; CPFA • 20 ♂♂, 26 ♀♀ same data as for preceding, 5 – 10 Mar. 1988; CPFA • 2 ♂♂, 2 ♀♀; Guayaquil, Los Ceibos; 29 Feb. 1988; L. Arnaud and P. Arnaud leg.; CEMT. Holotype of Phanaeus foveolatus (studied from photographs) ECUADOR • ♂; “ Guayaquil: Reist. / 38678 / 1405 / foveolatus Harold. ”; GMFNB. Non-type material examined (4 ♂♂, 2 ♀♀) ECUADOR • 1 ♀; L. W. Saylor leg.; ex-Gonzalo Halffter Salas; IEXA. – Guayas • 1 ♂; 40 km SW of Guayaquil; elev. 50 m; 21 – 22 Feb. 1981; B. D. Gill leg.; TAMU. – Manabí • 1 ♂, El Aromo, La Fabril; 16 May 2010; W. Chamorro leg; feces; CEMT • 1 ♂; Machalilla; 1 ° 28 ′ 18 ″ S, 80 ° 45 ′ 48 ″ W; elev. 30 m; 17 Mar. 1999; C. Rivadeneira leg.; CEMT • 1 ♂, 1 ♀; vicinity of Montecristi; 01 ° 02 ′ 22 ″ S, 80 ° 40 ′ 33 ″ W; 8 – 9 Feb. 2003; B. D. Streit leg.; pitfall human dung; TAMU. Type locality Ecuador, Guayas, Isla Puná.	en	Bordin, Bruna Ribeiro, Arias-Buriticá, Jorge, Maldaner, María Eduarda, Vaz-De-Mello, Fernando Z. (2026): Taxonomic revision of the Phanaeus chalcomelas species group and a description of the new Phanaeus achilles species group (Coleoptera: Scarabaeidae: Scarabaeinae: Phanaeini). European Journal of Taxonomy 1080: 1-65, DOI: 10.5852/ejt.2026.1080.3310, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3310/14554
03B6F8059611FF94B94CFDE018FA669E.taxon	distribution	Distribution Dry forests and shrublands of southwestern Ecuador, from 0 to 50 m a. s. l. (Fig. 27).	en	Bordin, Bruna Ribeiro, Arias-Buriticá, Jorge, Maldaner, María Eduarda, Vaz-De-Mello, Fernando Z. (2026): Taxonomic revision of the Phanaeus chalcomelas species group and a description of the new Phanaeus achilles species group (Coleoptera: Scarabaeidae: Scarabaeinae: Phanaeini). European Journal of Taxonomy 1080: 1-65, DOI: 10.5852/ejt.2026.1080.3310, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3310/14554
03B6F805960EFF90B95BFE861F8A6277.taxon	description	urn: lsid: zoobank. org: act: FEF 361 C 8 - 1 CE 1 - 419 D-B 62 B-FB 8 E 76 F 26 E 11 Figs 1 C, 22 C – D, 23 B, D, F, 24 B, D, 25 B, 26 B, D, F, H	en	Bordin, Bruna Ribeiro, Arias-Buriticá, Jorge, Maldaner, María Eduarda, Vaz-De-Mello, Fernando Z. (2026): Taxonomic revision of the Phanaeus chalcomelas species group and a description of the new Phanaeus achilles species group (Coleoptera: Scarabaeidae: Scarabaeinae: Phanaeini). European Journal of Taxonomy 1080: 1-65, DOI: 10.5852/ejt.2026.1080.3310, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3310/14554
03B6F805960EFF90B95BFE861F8A6277.taxon	diagnosis	Diagnosis This species is diagnosed by the following combination of characters: Anteromedial margin of clypeus weakly bidentate (Fig. 22 C – D); major males with long cephalic horn curved over the pronotum (Figs 22 C, 23 B, 24 B); pronotum bearing contrasted black elevations, variously and irregularly shaped (Figs 22 C – D, 23 B, F, 24 B); anteromedial pronotal region of major males with two longitudinal curved carinae, each forming a pronotal tubercle, with the apices pointing upwards (Figs 23 B, 24 B, D); pronotum of females with anterior W-shaped carina (Fig. 23 F); elytral striae punctate, basally impressed as distinct fossae (Fig. 22 C – D); dorsum completely shiny metallic, green or bronze green (Figs 22 C – D, 23 B, D, F); black ventral color, or black with green reflections on the ventral portion of femora.	en	Bordin, Bruna Ribeiro, Arias-Buriticá, Jorge, Maldaner, María Eduarda, Vaz-De-Mello, Fernando Z. (2026): Taxonomic revision of the Phanaeus chalcomelas species group and a description of the new Phanaeus achilles species group (Coleoptera: Scarabaeidae: Scarabaeinae: Phanaeini). European Journal of Taxonomy 1080: 1-65, DOI: 10.5852/ejt.2026.1080.3310, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3310/14554
03B6F805960EFF90B95BFE861F8A6277.taxon	etymology	Etymology The specific epithet is dedicated to Alain (in memoriam), Patrick, Lydie (in memoriam), Florent and Audrey Arnaud and their descendants, in honor of their contributions to Scarabaeinae collections and taxonomy, and strong friendship to one of us for so many years.	en	Bordin, Bruna Ribeiro, Arias-Buriticá, Jorge, Maldaner, María Eduarda, Vaz-De-Mello, Fernando Z. (2026): Taxonomic revision of the Phanaeus chalcomelas species group and a description of the new Phanaeus achilles species group (Coleoptera: Scarabaeidae: Scarabaeinae: Phanaeini). European Journal of Taxonomy 1080: 1-65, DOI: 10.5852/ejt.2026.1080.3310, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3310/14554
03B6F805960EFF90B95BFE861F8A6277.taxon	materials_examined	Type material (59 ♂♂, 39 ♀♀) Holotype ECUADOR • ♂; El Oro, Arenillas; 3 ° 34 ′ 24 ″ S, 80 ° 08 ′ 36 ″ W; elev. 13 m; 15 Jun. 2012; D. Marín leg.; pitfall dung; CEMT. Paratypes ECUADOR – El Oro • 29 ♂♂, 15 ♀♀; same data as for holotype; CEMT • 2 ♂♂, 1 ♀; same data as for holotype; MGCB • 5 ♂♂, 2 ♀♀; same data as for holotype; CPFA • 1 ♂, 1 ♀; 3 km S of Arenillas; elev. 50 m; 26 May 2002; Curoe leg.; CEMT. – Loja • 1 ♂; Catamayo; 16 Oct. 2005; F. Ohs leg.; TAMU • 1 ♂; Cazaderos; TAMU • 1 ♀; Cantón Zapotillo, Ceiba Chica; 17 Mar. 2018; T. Atocha leg.; LOUNAZ • 1 ♀; Guachanamá, Sabanilla; 4 ° 06 ′ 58 ″ S, 79 ° 48 ′ 59.4 ″ W; elev. 800 m; 8 Mar. 2021; W. Chamorro leg.; pitfall hum dung; LOUNAZ • 11 ♂♂, 15 ♀♀; Loja, ex Abbé Gaujon; MNHN • 1 ♂; Loja; ac: 22623; L. W. Saylor leg.; ex-Gonzalo Halffter Salas; IEXA. PERU – Pasco • 3 ♂♂, 2 ♀♀; Oxapampa; elev. 1800 m; ex Le Moult; CPFA. – Piura • 1 ♀; Ayabaca, C. C. Samanga, Sector Espíndola; 4 ° 42 ′ 16 ″ S, 79 ° 27 ′ 30 ″ W; elev. 2847 m; 16 Feb. 2009; D. Saavedra leg.; CEMT. – Tumbes • 3 ♂♂; Canovitas; PN Cerros de Amotape; Feb. 1995; CPFA • 1 ♂; Tumbes; 16 May 2007; D. Ramírez leg.; CEMT. Type locality Ecuador, El Oro, Arenillas.	en	Bordin, Bruna Ribeiro, Arias-Buriticá, Jorge, Maldaner, María Eduarda, Vaz-De-Mello, Fernando Z. (2026): Taxonomic revision of the Phanaeus chalcomelas species group and a description of the new Phanaeus achilles species group (Coleoptera: Scarabaeidae: Scarabaeinae: Phanaeini). European Journal of Taxonomy 1080: 1-65, DOI: 10.5852/ejt.2026.1080.3310, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3310/14554
03B6F805960EFF90B95BFE861F8A6277.taxon	description	Description Major male (holotype) MEASUREMENTS. Length 20 mm. Width 12.4 mm. HEAD. Anteromedial margin of clypeus weakly bidentate with striated sculpture (Fig. 22 C). Cephalic horn black, long, curved posteriorly over the pronotum, occupying central third of total width of head (Figs 22 C, 23 B, 24 B). Bright green on paraocular area and posterior portion of clypeus (Fig. 22 C). Paraocular ridge distinct, extending along the entire lateral margin of eyes (Fig. 22 C). PRONOTUM. Bright green, with irregularly black elevations over the whole disc (Fig. 22 C). Anteromedial region with two longitudinal curved carinae, each forming a posterior tubercle (Figs 23 B, 24 B, D). Posterior angles expanded upwards and backwards, the apices of which are curved towards center of pronotum (Fig. 23 B). Posteromedial fossae rounded (Fig. 22 C). Pronotal punctures strong, more visible on anterior and lateral portions (10 × magnification). ELYTRA. Bright green, striae punctate, basally impressed as distinct fossae (Fig. 22 C). Interstriae with punctures (10 × magnification) evenly spaced over the entire surface (Fig. 1 C). METAVENTRITE. Completely black with strong punctures (10 × magnification). Tip of anteromedial prominence triangular. PYGIDIUM. Bright green with strong punctures over the whole surface (Fig. 23 D). ENDOPHALLITES. ME with a large tab, given triangular appearance in dorsal and ventral view; margins of tab almost straight with a lateral acute angle (Fig. 26 D). AE, in dorsal view, triangular, with concave lateral margins; left margin with lateral projection near the apex (Fig. 26 F). SRP with rounded left margin and tapered left portion (dorsal view) (Fig. 26 H). Variation Male sizes: length 14 – 22 mm; width 9 – 12 mm. Female sizes: length 17 – 22 mm; width 9.8 – 12 mm. Females: head with a transversal carina. Middle of the cephalic carina with two elevations, giving a bituberculate appearance in frontal view (Fig. 23 F); anterior carina of pronotum W-shaped in frontal view (Fig. 23 F); oval posteromedial pronotal fossae (Fig. 22 D).	en	Bordin, Bruna Ribeiro, Arias-Buriticá, Jorge, Maldaner, María Eduarda, Vaz-De-Mello, Fernando Z. (2026): Taxonomic revision of the Phanaeus chalcomelas species group and a description of the new Phanaeus achilles species group (Coleoptera: Scarabaeidae: Scarabaeinae: Phanaeini). European Journal of Taxonomy 1080: 1-65, DOI: 10.5852/ejt.2026.1080.3310, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3310/14554
03B6F805960EFF90B95BFE861F8A6277.taxon	distribution	Distribution Dry forests and mountains in extreme northwest of Peru and the southwest of Ecuador, from 13 to 1210 m a. s. l. (Fig. 27).	en	Bordin, Bruna Ribeiro, Arias-Buriticá, Jorge, Maldaner, María Eduarda, Vaz-De-Mello, Fernando Z. (2026): Taxonomic revision of the Phanaeus chalcomelas species group and a description of the new Phanaeus achilles species group (Coleoptera: Scarabaeidae: Scarabaeinae: Phanaeini). European Journal of Taxonomy 1080: 1-65, DOI: 10.5852/ejt.2026.1080.3310, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3310/14554
03B6F805960EFF90B95BFE861F8A6277.taxon	discussion	Taxonomic discussion The external morphology and the parameres of P. achilles and P. arnaudorum sp. nov. are very similar (Figs 22, 25). However, in addition to the difference in coloration — P. achilles being currant red with green sheen, and P. arnaudorum being green or bronze green — these species also differ in the shape of the medial and additional endophallites (Fig. 26 C – F). Furthermore, the anterolateral pronotal fossae are larger in major males of P. arnaudorum compared to P. achilles and the pronotal tubercles of major males are smaller in P. arnaudorum than in P. achilles (Fig. 24 C – D).	en	Bordin, Bruna Ribeiro, Arias-Buriticá, Jorge, Maldaner, María Eduarda, Vaz-De-Mello, Fernando Z. (2026): Taxonomic revision of the Phanaeus chalcomelas species group and a description of the new Phanaeus achilles species group (Coleoptera: Scarabaeidae: Scarabaeinae: Phanaeini). European Journal of Taxonomy 1080: 1-65, DOI: 10.5852/ejt.2026.1080.3310, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3310/14554
