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 Key to species of the Phanaeus endymion species group 1. Sutural margin of each elytron upturned to form a sharp ridge, which is progressively more elevated posteriorly and prolonged into a small, sharp tooth at apical angle; elytral margin slightly excised adjacent to this tooth.......................................................................................................................... 2 – Sutural margin of elytra simple (running straight, not upturned)...................................................... 3 2. Male with dentiform keel in the middle of anterior pronotal margin, lateral lines of pronotal triangle straight. Southern Mexico State and Morelos ( Fig. 15 ).................... P. halffterorum Edmonds, 1979 – Male with keel absent in the middle of anterior pronotal margin, lateral lines of pronotal triangle curved. Sierra Madre del Sur, central Guerrero ( Fig. 15 ) ................................................................... ............................................................. P. bravoensis Moctezuma, Sánchez-Huerta & Halffter, 2017 3. Pronotal disc of male evenly and densely but finely granulate, granules in most specimens larger and becoming squamose along lateral margins of disc and extending onto posterolateral angles; sides of pronotum strongly roughened ( Figs 2L , 14A, C ). Female pronotum slightly roughened; with distinctly impressed midline ( Figs 14B , 18Q ). Sierra Madre del Sur and Sierra Norte, central Oaxaca ( Fig. 15 ) ................................................................................................. P. zapotecus Edmonds, 2006 – Pronotal disc of male lacking distinct granulation, or, if granules present, these are minute and restricted along lateral margins of disc; sides of pronotum smooth. Female pronotum smooth; with superficially impressed to completely effaced midline...................................................................... 4 4. Elytral striae deeply punctate; all strial punctures forming a distinct fossa, giving a completely roughened surface to striae ( Figs 1K , 2G , 9 , 19L ). Pacific slope of the Sierra Madre de Chiapas and El Salvador ( Fig. 15 )............................................................................................. P. pacificus sp. nov. – Elytral striae distinctly punctate to impunctate. If strial punctures distinctly impressed, forming a distinct fossa giving a completely roughened surface to first and second striae, or strial punctures forming a distinct fossa giving a partially roughened surface to basal half of striae ....................... 5 5. Pronotum of female with anteromedial concavity bounded anteriorly by three variable on shape tubercles ( Figs 4C , 11B , 13B , 18E, N, P ).......................................................................................... 6 – Pronotum of female evenly convex, lacking anteromedial concavity even in largest specimens, bearing three round, smooth tubercles in transverse line near anterior margin ( Figs 3B , 5B , 6B , 7B , 8B , 10B , 12C , 18C, F, I–K, M, O )................................................................................................... 10 6. Pronotum of female with concavity bounded anteriorly by a raised U- or V-shaped process; middle pronotal tubercle dentiform or rounded, always more developed and frontally projected than lateral tubercles; pronotal concavity interrupted posteriorly by a small rounded bump or strongly developed dentiform tubercle ( Figs 4C , 13B , 18E, P ). Endophallite copulatrix variable................................... 7 – Pronotum of female with concavity bounded anteriorly by three isolated, round, almost aligned tubercles; middle tubercle sometimes dentiform and more strongly developed than lateral tubercles ( Figs 11B , 18N ). Right and left lobes of endophallite copulatrix similar in size; right lobe obtusely triangular in shape, weakly developed superiorly; left lobe obtusely lobed, strongly developed; central ridge distinctly developed ( Fig. 1M ). Caribbean Honduras and Belize ( Fig. 16 ) .......................................................................................... P. porioni Arnaud, 2002 stat. rev. 7. Pronotal disc of male with superficially impressed midline ( Fig. 2K ). Right lobe of endophallite copulatrix more developed than left lobe; right lobe obtusely triangular in shape, sharply acute frontally, rounded superiorly; left lobe strongly developed, obtusely lobed; central ridge strongly developed ( Fig. 1O ). Sierra Madre del Sur, southern Michoacán ( Fig. 15 ) ...... P. rzedowskii sp. nov. – Pronotal disc of male with completely effaced midline ( Fig. 2B ). Central ridge of endophallite copulatrix distinctly developed ( Fig. 1D, G, Q ) ................................................................................ 8 8. Pronotum of male uniformly dark blue, dark metallic blue-green or dark metallic green ( Figs 2B , 4A–B ). Weakly developed keel close to anterior margin of pronotum, or keel completely effaced ( Fig. 2b ). Right lobe of endophallite copulatrix more developed than left lobe; right lobe obtusely triangular in shape; left lobe obtusely lobed inferiorly, obtusely triangular superiorly; central ridge less developed than central column ( Fig. 1D ). Lowlands and midlands of the Gulf of Mexico slope, Yucatán Peninsula, Belize , northern Guatemala ( Fig. 15 )......................... P. endymion Harold, 1863 – Pronotum of male uniformly bright metallic green, bright metallic green-blue, sometimes showing a red or golden sheen. Keel always absent on anterior margin of pronotum. Endophallite copulatrix variable .............................................................................................................................................. 9 9. Anterior metasternal angle obtuse in lateral view. Lateral metasternal angles well defined and slightly curved. Left lobe of endophallite copulatrix almost completely lobed ( Fig. 1Q ). Temperate mountains of Los Chimalapas, eastern Oaxaca and western Chiapas ( Fig. 15 ).................................. ................................................................................................. P. zoque Moctezuma & Halffter, 2017 – Anterior metasternal angle almost right angled but with rounded apex in lateral view. Lateral metasternal angles evanescent. Left lobe of endophallite copulatrix straight posteroinferiorly ( Fig. 1G ). Jalisco , northern Michoacán , Nayarit , southern Sinaloa ( Fig. 15 ) ..................................... .................................................................. P. huichol Moctezuma, Sánchez-Huerta & Halffter, 2017 10. Elytral interstriae evenly convex and glossy midlongitudinally; striae impressed basally as distinct fossae ( Figs 3 , 6 , 7 , 18C, I–J ) ...........................................................................................................11 – Elytral interstriae distinctly flattened and uniformly dull; striae not strongly impressed basally ( Figs 5 , 8 , 10 , 12 , 18F, K, M, O )...................................................................................................... 14 11. Almost completely bright black dorsally ( Figs 2E , 7 ). Pacific slope of the Cordillera de Talamanca, Costa Rica and Panama ( Fig. 16 ) ..................................................................... P. malyi Arnaud, 2002 – Pronotum typically bright metallic green, blue-green or dark metallic blue ( Figs 2A, D , 3 , 6 )...... 12 12. Distinctly developed keel on central anterior margin of pronotum of male; right lobe of endophallite copulatrix tapering superiorly and straight apically ( Fig. 1C ). Sierra Madre del Sur, southern Oaxaca ( Fig. 15 ) ............................................................... P. edmondsi Moctezuma, Deloya & Halffter, 2019 – Always with keel absent on central anterior margin of pronotum of male ( Fig. 2A, D ). Endophallite copulatrix variable ........................................................................................................................... 13 13. Larger species, frequently attaining 19–20 mm in length. Posterolateral angles of male widened, slightly projected posteriorly ( Figs 2A , 3 ). Central highlands of Chiapas ( Fig. 15 ) ........................... ........................................................................................................................ P. chiapanecus sp. nov. – Smaller species, never attaining 19 mm in length. Posterolateral angles of male sharply acute, strongly projected posteriorly ( Figs 2D , 6 ). Inner slope of the Sierra Madre de Chiapas, south-central Guatemala and southeastern Chiapas ( Fig. 15 ) ................................................... P. jackenioi sp. nov. 14. Pronotal disc of male completely bright metallic ( Figs 2J , 12 )....................................................... 15 – Pronotal disc of male distinctly black, without metallic reflection ( Figs 2E, J, L , 5A, C , 8A , 10A ) .. ........................................................................................................................................................ 16 15. Pronotal disc typically bright metallic green, rarely dark metallic blue. Pronotal process of female with central tubercle more developed and anteriorly projected than lateral tubercles. Ecuador ( Figs 17 , 18A ).............................................................................................. P. arletteae Arnaud, 2002 – Pronotal disc typically bright metallic red, green, or dark metallic blue ( Figs 2J , 12 ). Pronotal process of female with tubercles almost aligned, central tubercle not projecting anteriorly ( Figs 12C , 18O ). Nicaragua and north Caribbean Costa Rica ( Fig. 16 )......................................... P. pyrois Bates, 1887 16 Pronotum distinctly bright metallic red frontolaterally. Head bright metallic red with green-yellow metallic sheen ( Figs 2F, H , 8 , 10 ).................................................................................................... 17 – Pronotum distinctly dull metallic red frontolaterally. Head dull metallic red without green-yellow metallic sheen ( Figs 2C , 5 ). Pacific slope of the Andes, north-central Ecuador and Colombia ( Fig. 17 ) ..................................................................................... P. funereus Balthasar, 1939 stat. rev. 17. Larger species, frequently attaining from 22 to 24 mm in length. Carinate, distinctly developed keel on central anterior margin of pronotum of male ( Fig. 2F ). Pronotal posterolateral angles of male strongly developed, projected laterally ( Fig. 8A ). Pacific slope of the Andes, south-central Colombia and northern Ecuador ( Fig. 17 )................................................ P. olsoufieffi Balthasar, 1939 stat. rev. – Smaller species, rarely attaining 20 mm in length. Always with keel absent on central anterior margin of pronotum of male ( Fig. 2H ). Pronotal posterolateral angles of male weakly developed, widened, and slightly projected posteriorly ( Fig. 10A ). Caribbean Costa Rica and Panama ( Fig. 16 ) ........................................................................................................... P. panamensis sp. nov.