Taxonomic revision of the Phanaeus endymion species group (Coleoptera: Scarabaeidae), with the descriptions of five new species Author Moctezuma, Victor B672EF02-4113-480E-B297-F0158D11DF4A 0000-0002-4532-0302 Red de Ecoetología, Instituto de Ecología, A. C. Carretera antigua a Coatepec 351, El Haya, Xalapa, Veracruz 91073, Mexico. abadonjvpm@hotmail.com Author Halffter, Gonzalo CF2EF74E-878A-4BC8-82A2-01F669049650 0000-0003-1430-7049 Red de Ecoetología, Instituto de Ecología, A. C. Carretera antigua a Coatepec 351, El Haya, Xalapa, Veracruz 91073, Mexico. gonzalohalffter@inecol.mx text European Journal of Taxonomy 2021 2021-04-28 747 1 71 http://zoobank.org/3f0b6eaf-c616-4865-811a-414a094b590c journal article 10.5852/ejt.2021.747.1333 aca67cc8-96ca-4f24-b8af-2b532ecd355c 2118-9773 4836744 3F0B6EAF-C616-4865-811A-414A094B590C Phanaeus olsoufieffi Balthasar, 1939 stat. rev. Figs 1J , 2F , 8 , 17 , 18K , 19K Phanaeus blanchardi Olsoufieff, 1924: 92 (not P. blanchardi Harold, 1871: 114 ). Phanaeus olsoufieffi Balthasar, 1939: 242 . Phanaeus bothrus Blackwelder, 1944: 209 . Phanaeus ( Phanaeus ) blanchardi Martínez & Pereira 1967: 68 (as synonym of P. funereus ). Phanaeus ( Phanaeus ) olsoufieffi Martínez & Pereira 1967: 68 (as synonym of P. funereus ). Phanaeus bothrus Martínez & Pereira 1967: 68 (as synonym of P. funereus ). — Edmonds 1994: 8 , 45 (as synonym of P. pyrois ). — Arnaud 2002b: 97 (as synonym of P. olsoufieffi ). Krajcik 2006: 152 (as synonym of P. pyrois ). Edmonds & Zídek 2012: 3 , 5 (as synonym of P. pyrois ). Phanaeus blanchardi Vulcano & Pereira 1967: 575 . — Martínez & Pereira 1967: 68 (as synonym of P. funereus ). — Edmonds 1972: 830 , fig. 256; 1994: 3, 8, 45–46 (as synonym of P. pyrois ). — Arnaud 1982: 116 (as synonym of P. funereus ). — Krajcik 2006: 152 (as synonym of P. pyrois ). — Edmonds & Zídek 2012: 5 , 13 (as synonym of P. pyrois ). — Chamorro et al . 2019: 220–221 (as synonym of P. pyrois ). Phanaeus olsoufieffi Edmonds 1994: 8 , 45–46 (as synonym of P. pyrois ). — Arnaud 2002b: 96 . — Krajcik 2006: 152 (as synonym of P. pyrois ). — Edmonds & Zídek 2012: 3 , 5–6 (as synonym of P. pyrois ). Phanaeus ( Notiophanaeus ) pyrois olsoufieffi Arnaud 2002b: 96 . Phanaeus pyrois olsoufieffi Arnaud 2002b: 98 ; 2018: 4 , pl. 1, figs d–e. Non Phanaeus pyrois (error). — Medina et al . 2001: 140 (in part). — Moctezuma & Halffter 2017: 55 (in part). — Moctezuma et al . 2017: 114, 130 (in part). Diagnosis This is the largest species within the P. endymion species group, frequently attaining 21–24 mm in length. Phanaeus olsoufieffi is diagnosed by the black colour with bright metallic red-green sheen; and elytral striae not strongly impressed basally ( Fig. 8 ). This species is easily separated from the closely related P. panamensis sp. nov. by the larger body size and endophallite copulatrix ( Fig. 1 ). Additionally, the major males of P. olsoufieffi are diagnosed by the distinctly developed keel in the middle of anterior pronotal margin; and posterolateral angles rounded, strongly developed, and projected laterally ( Figs 2F , 8A ). Type material (studied from photographs, 2 ♂♂ , 3 ♀♀ ) Lectotype COLOMBIA Valle del Cauca , Arnaud 1982: 116 ( Fig. 8C ); “M. de Mathan 1898 / Ph. blanchardi ♂ Olsuf. det. G. Olsufiew. / Phanaeus apollinaris Muzo 1928 / Muséum Paris 1952 Coll R. Oberthur / P. blanchardi Ols. LECTOTYPE ♂ P. ARNAUD DET 1980/ Lectotype / Phanaeus pyrois Bates, Det. W. D. Edmonds ’83 / MNHN EC10566 ”; MNHN . Paralectotypes COLOMBIA1 ♂ ; “ExMus ӕo VAN LANSBERGE /Museum Paris ex. Coll. R. Oberthur / PARALECTOTYPE/ Phanaeus blanchardi Ols. PARALECTOTYPE ♂ P. ARNAUD DET 1981/ Phanaeus pyrois Bates / Det. W. D. Edmonds ‘83”/ MNHN EC10568 ”; MNHN . – Boyacá 1 ♀ ; “ Muzo / Coll.E.Steinheil /MuseumParisex. Coll.R.Oberthur / PARALECTOTYPE/ PARALECTOTYPE Phanaeus ( Phanaeus ) blanchardi Olsoufieff, 1924 / MNHN EC10570 ”; MNHN EC10570 . – Valle del Cauca 1 ♀ ; “ Santa Rosa entre S. Francisco & Carthago . Eujenio Garzon Aout 1878 ”; MNHN . – Unknown locality 1 ♀ ; “(Illegible data) / Ex-mus ӕo D. Sharp 1890/ Ph. blanchardi ♀ Olsuf. det. G. Olsoufiew. / Museum Paris ex Coll. R. Oberthur / Phanaeus blanchardi ♀’s. PARALECTOTYPE ♀ P. ARNAUD DET 1981/ PARALECTOTYPE / Phanaeus pyrois Bates Det. W. D. Edmonds ’83/ MNHN EC10567 ”; MNHN . Non-type material revised ( 24 ♂♂ , 16 ♀♀ ) COLOMBIA Chocó 3 ♂♂ , 1 ♀ ; “ Q.Taparal B.San Juan . CNF. Nov 3–12–92 . L.C. Pardo leg”; TAMU 1 ♂ , 1 ♀ ; “ Pacurita , 53m . 25-XI-01. 5°41’N , 76°40’W , bosque. Excrem. J.C. Neita col”; IEXA 5 ♂♂ , 5 ♀♀ ; same collection data as for preceding; TAMU 1 ♂ , 1 ♀ ; same collection data as for preceding; UVGC 1 ♂ ; same collection data as for preceding; VMC 1 ♀ ; “ Tutunendó , ( 20 km NE Quibdó ), 60m , 26-XI-01. J.C. Neita col”; TAMU 3 ♀♀ ; “ Unión Panamericana , 115m . 5°32’45N , 76°44’33’’W (No date) J.C. Neita col”; TAMU 1 ♂ , 1 ♀ ; “ Lloró , ( 3k km S Quibdó ) 5°30’N , 76°33.5’W 90 m . J.C. Neita col (no date)”; TAMU 2 ♂♂ ; “ Lloró , 90m , 20-II-03. 5°31’N , 76°33’W . Olaya & Mosquero ; TAMU 1 ♀ ; same collection data as for preceding; VMC . – Tolima 1 ♂ ; “ Honda ”; GHC . – Valle del Cauca 1 ♂ ; “Escalerete CF. Jul 19–21–91 . L.C. Pardo leg”; TAMU 1 ♂ ; same collection data as for preceding; VMC 1 ♂ , 1 ♀ ; “ B/Ventura B.Calima , C.F. 50 msnm. Col. ME Hitchcox ”; TAMU 1 ♂ , 1 ♀ ; “S30- 290. L.C. Pardo Locarno Leg.”; GHC 2 ♂♂ ; same collection data as for preceding; VMC . ECUADOR Esmeraldas 2 ♂♂ ; “ Charco Vicente . Ex. Pitfall heces humanas . 16/05/01. J. Quito F. Añapa ”; TAMU . Type locality Colombia , Valle del Cauca . Redescription Major male HEAD. Clypeus bidentate, black anteriorly, bright metallic red posteriorly, with green sheen; roughened sculpture. Genae bright metallic red, with green sheen; roughened sculpture. Front black. Cephalic horn black, curved posteriorly over pronotum ( Figs 2F , 8A ). PRONOTUM. Carinate, distinctly developed keel in the middle of anterior pronotal margin. Disc triangular, flat, with two distinctly developed tubercles on anterior portion. Triangle completely dull black, with bright metallic red sheen; completely smooth, scabriculous, impunctate. Sides bright metallic red, with green sheen; smooth sculpture, scabriculous, with effaced to almost effaced punctures. Lateral lines of pronotal triangle straight. Posterolateral angles rounded, strongly developed, projected laterally. Lateral fossae distinctly impressed. Basal fossae obtusely oval, deeply impressed to effaced. Posterior margin impunctate, becoming occasionally bright metallic red ( Figs 2F , 8A ). ELYTRA. Striae fine, smooth, dull black, with superficially impressed to effaced punctation, scabriculous, not strongly impressed basally. Interstriae dull black, smooth, scabriculous, impunctate. Sutural margin without apical tooth ( Fig. 8A ). PROTIBIAE. Quadridentate with apical spine. TERGITE VIII. Bright metallic red, with green sheen; scabriculous; with rough, superficially impressed punctures. Basal margin with setae variable in size. GENITALIA. Right lobe of endophallite copulatrix slightly more developed than left lobe. Right lobe obtusely triangular in shape; weakly developed, projected frontally. Left lobe concave superiorly, lobed inferiorly. Central ridge less developed than central column ( Fig. 1J ). Minor male Like the major male, except for the reduction of secondary sexual characters (i.e., cephalic horn, pronotal triangle and tubercles, and pronotal posterolateral angles). Female Similar to the male, except for the head showing a cephalic trituberculate carina; with weakly developed tubercles; carinate middle tubercle, slightly more frontally projected and more developed than lateral tubercles; frons with superficially impressed punctures; pronotal sculpture smooth, with almost effaced to effaced punctures; pronotum almost completely black, becoming posteriorly and laterally bright metallic red with green sheen; pronotal process trituberculate, lacking concavities; pronotal tubercles weakly developed, well-spaced, with middle tubercle more developed and posteriorly projected than lateral tubercles; posterior pronotal midline completely effaced ( Fig. 8B ). Variation Mean length 20.1 mm ( 15.9–23.9 mm ). Minor males occasionally show the pronotal disc almost completely bright metallic red, with green sheen. Fig. 8. Phanaeus olsoufieffi Balthasar, 1939 stat. rev. A . ♂ (TAMU). B . ♀ (TAMU). C . Lectotype ♂ (by Christophe Rivier, Muséum national d’histoire naturelle, Paris, France) (MNHN). Distribution Pacific Slope of the Andes, north-central Colombia and northern Ecuador ( Fig. 17 ). Previous authors reported P. olsoufieffi from Panama ( Arnaud 2002b ; Kohlmann et al . 2018 ). Nevertheless, these authors confused P. olsoufieffi with P. panamensis sp. nov. The specimens revised herein are the first accurate records of P. olsoufieffi from Ecuador . The distributions of P. olsoufieffi and P. funereus show a significant area of sympatry. Nevertheless, we did not find any putative hybrid specimens. Remarks Phanaeus olsoufieffi was considered as a synonym of P. pyrois by previous authors ( Edmonds 1994 ; Edmonds & Zídek 2012 ). Nonetheless, a diagnosis and an updated key to separate P. olsoufieffi and closely related species are provided by us. As a consequence, P. olsoufieffi is confidently resurrected herein from previous synonymy and full species status is assigned to it. The lectotype of P. olsoufieffi is a minor male ( Fig. 8C ). Therefore, the redescription mainly relies on the type series and some specimens (n =8) collected from the type locality ( Valle del Cauca , Colombia ). When revising the type material deposited at MNHM, we were not able to find out the locality data for a Colombian female paralectotype of P. olsoufieffi (MNHN EC10567). Its locality data is probably indicated in an illegible label. Additionally, we found out that a paralectotype of P. olsoufieffi pertained to P. malyi ( Fig. 7C , MNHN EC10569). Consequently, we conclude that the type series of P. olsoufieffi lumped together two distinct species. Previous authors ( Arnaud 1982 ; 2002a , 2002b ; Edmonds 1994 ; Edmonds & Zídek 2012 ) did not realize this fact that solves the controversy considering P. malyi as a junior subjective synonym of P. olsoufieffi or not ( Edmonds & Zídek 2012 ; Solís & Kohlmann 2012 ; Kohlmann et al . 2018 ).