Phanaeus zoque Moctezuma & Halffter, 2017 Phanaeus endymion (Edmonds, 1994)

Moctezuma, Victor & Halffter, Gonzalo, 2021, Taxonomic revision of the Phanaeus endymion species group (Coleoptera: Scarabaeidae), with the descriptions of five new species, European Journal of Taxonomy 747, pp. 1-71 : 50-54

Phanaeus zoque Moctezuma & Halffter, 2017

Figs 1Q , 15 , 18R , 19R

Phanaeus zoque Moctezuma & Halffter, 2017: 47 , 48, 52, 55–56, figs 1–15, 23.

Phanaeus zoque – Moctezuma et al. 2017: 113–114, 126, 128, 131–132, fig. 19; 2019: 252–253, fig. 5. — Kohlmann et al. 2018: 83, 88.

Type material

Holotype

MEXICO • ♂, Moctezuma & Halffter 2017: 48 (revised); Oaxaca, Santa María Chimalapa, Chocomanatlán ; IEXA.

Type locality

Mexico, Oaxaca, Santa María Chimalapa, Chocomanatlán (José López Portillo).

Distribution

Temperate mountains of Los Chimalapas, eastern Oaxaca and western Chiapas ( Fig. 15 ).

Remarks

Mean length 15.8 mm ( 13.2–18.4 mm). To the original description we add the elytral striae impressed basally as distinct fossae; right and left lobes of endophallite copulatrix similar in size; right lobe obtusely triangular in shape, distinctly rounded superiorly; left lobe strongly developed, obtusely lobed; central ridge less developed than central column ( Fig. 1Q ). For the female, the head showing a cephalic trituberculate carina; with conical, similar in size tubercles; middle tubercle slightly more frontally projected than lateral tubercles; frons with almost effaced to effaced punctures; pronotal sculpture completely smooth, with almost effaced to effaced punctures; pronotal process trituberculate, with posterior concavity; with dentiform middle tubercle, strongly more developed and projected frontally than lateral tubercles; carinate or rounded lateral tubercles; posterior pronotal midline almost completely effaced.

After having described P. zoque ( Moctezuma & Halffter 2017) , the following paratypes were deposited by us in additional collections: CEMT: 5 ♂♂, 5 ♀♀; CDC: 5 ♂♂, 5 ♀♀; CMNC: 1 ♂, 1 ♀; CPFA: 6 ♂♂, 6 ♀♀; UVGC: 1 ♂, 1 ♀. These paratypes were previously housed at VMC (see the full list of paratypes in Moctezuma & Halffter 2017).

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.

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.

Figures

Fig. 1. Endophallite copulatrix of the Phanaeus endymion species group (except for P. arletteae Arnaud, 2018). A. Phanaeus bravoensis Moctezuma, Sánchez-Huerta & Halffter, 2017 (IEXA). B. Phanaeus chiapanecus sp. nov. (TAMU). C. Phanaeus edmondsi Moctezuma, Deloya & Halffter, 2019 (IEXA). D. Phanaeus endymion Harold, 1863 (VMC). E. Phanaeus funereus Balthasar, 1939 stat. rev. (VMC). F. Phanaeus halffterorum Edmonds, 1979 (TAMU). G. Phanaeus huichol Moctezuma, Sánchez- Huerta & Halffter, 2017 (IEXA). H. Phanaeus jackenioi sp. nov. (UVGC). I. Phanaeus malyi Arnaud, 2002 (VMC). J. Phanaeus olsoufieffi Balthasar, 1939 stat. rev. (VMC). K. Phanaeus pacificus sp. nov. (UVGC). L. Phanaeus panamensis sp. nov. (TAMU). M. Phanaeus porioni Arnaud, 2001 stat. rev. (VMC). N. Phanaeus pyrois Bates, 1887 (VMC).O. Phanaeus rzedowskii sp. nov. (TAMU).P. Phanaeus zapotecus Edmonds, 2006 (VMC). Q. Phanaeus zoque Moctezuma & Halffter, 2017 (IEXA).

Fig. 15. Distribution of the Phanaeus endymion species group in North and Central America (in part). Phanaeus bravoensis Moctezuma, Sánchez-Huerta & Halffter, 2017;P. edmondsi Moctezuma, Deloya & Halffter, 2019; P. halffterorum Edmonds, 1979; and P. huichol Moctezuma, Sánchez-Huerta & Halffter, 2017 by Jose Luis Sánchez-Huerta.

Fig. 18. Pronotum of females within the Phanaeus endymion species group. A. Phanaeus arletteae Arnaud, 2018 (modified from Arnaud 2018). B. Phanaeus bravoensis Moctezuma, Sánchez-Huerta & Halffter, 2017 (by Jose Luis Sánchez-Huerta).C. Phanaeus chiapanecus sp. nov. D. Phanaeus edmondsi Moctezuma, Deloya & Halffter, 2019 (by Jose Luis Sánchez-Huerta). E. Phanaeus endymion Harold, 1863. F. Phanaeus funereus Balthasar, 1939 stat. rev. G. Phanaeus halffterorum Edmonds, 1979 (by Jose Luis Sánchez-Huerta).H. Phanaeus huichol Moctezuma, Sánchez-Huerta & Halffter, 2017 (by Jose Luis Sánchez-Huerta). I. Phanaeus jackenioi sp. nov. J. Phanaeus malyi Arnaud, 2002. K. Phanaeus olsoufieffi Balthasar, 1939 stat. rev. L. Phanaeus pacificus sp. nov. M. Phanaeus panamensis sp. nov. N. Phanaeus porioni Arnaud, 2001 stat. rev. O. Phanaeus pyrois Bates, 1887. P. Phanaeus rzedowskii sp. nov. Q. Phanaeus zapotecus Edmonds, 2006. R. Phanaeus zoque Moctezuma & Halffter, 2017.

Fig. 19. Elytral integument of species within the Phanaeus endymion species group. A. Phanaeus arletteae Arnaud, 2018 (modified from Arnaud 2018). B. Phanaeus bravoensis Moctezuma, Sánchez- Huerta & Halffter, 2017 (by Jose Luis Sánchez-Huerta). C. Phanaeus chiapanecus sp. nov. D. Phanaeus edmondsi Moctezuma, Deloya & Halffter, 2019 (by Jose Luis Sánchez-Huerta). E. Phanaeus endymion Harold, 1863. F. Phanaeus funereus Balthasar, 1939 stat. rev. G. Phanaeus halffterorum Edmonds, 1979 (by Jose Luis Sánchez-Huerta). H. Phanaeus huichol Moctezuma, Sánchez-Huerta & Halffter, 2017 (by Jose Luis Sánchez-Huerta). I. Phanaeus jackenioi sp. nov. J. Phanaeus malyi Arnaud, 2002.K. Phanaeus olsoufieffi Balthasar, 1939 stat. rev. L. Phanaeus pacificus sp. nov. M. Phanaeus panamensis sp. nov. N. Phanaeus porioni Arnaud, 2001 stat. rev. O. Phanaeus pyrois Bates, 1887. P. Phanaeus rzedowskii sp. nov. Q. Phanaeus zapotecus Edmonds, 2006. R. Phanaeus zoque Moctezuma & Halffter, 2017.

Fig. 2. Lateral view of males within the Phanaeus endymion species group. A. Phanaeus chiapanecus sp. nov. (TAMU). B. Phanaeus endymion Harold, 1863 (VMC). C. Phanaeus funereus Balthasar, 1939 stat. rev. (VMC). D. Phanaeus jackenioi sp. nov. (UVGC). E. Phanaeus malyi Arnaud, 2002 (VMC). F. Phanaeus olsoufieffi Balthasar, 1939 stat. rev. (VMC). G. Phanaeus pacificus sp. nov. (UVGC). H. Phanaeus panamensis sp. nov. (TAMU). I. Phanaeus porioni Arnaud, 2001 stat. rev. (VMC). J. Phanaeus pyrois Bates, 1887, green morph (VMC). K. Phanaeus rzedowskii sp. nov. (TAMU). L. Phanaeus zapotecus Edmonds, 2006 (VMC).

Fig. 14. A–D. Phanaeus zapotecus Edmonds, 2006. A. ♂ (TAMU). B. ♀ (TAMU). C. Minor ♂ (VMC). D. Tergite VIII (TAMU). E. Phanaeus dionysius Kohlmann, Arriaga-Jiménez & Rös, 2018 syn. nov., tergite VIII (IEXA).

Fig. 9. Phanaeus pacificus sp. nov. A. Holotype, ♂ (UVGC). B. ♀ (IEXA).

Fig. 4. Phanaeus endymion Harold, 1863. A. ♂, blue morph (typical; VMC). B. ♂, green morph (VMC). C. ♀, blue morph (typical; VMC). D. Lectotype, ♂ (by Christophe Rivier, Muséum national d’histoire naturelle, Paris, France) (MNHN).

Fig. 11. Phanaeus porioni Arnaud, 2001 stat. rev. A. ♂ (TAMU). B. ♀ (TAMU). C. Holotype, ♂ (by Patrick Arnaud) (CPFA).

Fig. 13. Phanaeus rzedowskii sp. nov. A. Holotype, ♂ (TAMU). B. ♀ (VMC).

Fig. 3. Phanaeus chiapanecus sp. nov. A. Holotype, ♂, blue morph (typical; TAMU). B. ♀, green morph (TAMU).

Fig. 5. Phanaeus funereus Balthasar, 1939 stat. rev. A. ♂ (TAMU). B. ♀ (TAMU). C. Holotype, ♂ (by Jiří Hájek, Národní Muzeum, Prague, Czech Republic) (NMPC).

Fig. 6. Phanaeus jackenioi sp. nov. A. Holotype, ♂ (UVGC). B. ♀ (VMC).

Fig. 7. Phanaeus malyi Arnaud, 2002.A. ♂ (TAMU). B. ♀ (TAMU). C. Colombian specimen previously labelled as a paralectotype of P. blanchardi Olsoufieff, 1924 (by Christophe Rivier, Muséum national d’histoire naturelle, Paris, France) (MNHN).

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).

Fig. 10. Phanaeus panamensis sp. nov. A. Holotype, ♂ (TAMU). B. ♀ (TAMU).

Fig. 12. Phanaeus pyrois Bates, 1887. A. ♂, red morph (VCM). B. ♂, blue morph (VMC). C. ♀, green morph (TAMU). D. Lectotype, ♂ (by Mario Cupello, Universidade Federal do Paraná, Brazil) (BMNH).

Fig. 16. Distribution of the Phanaeus endymion species group in Central America (in part).

Fig. 17. Distribution of the Phanaeus endymion species group in South America. Phanaeus malyi Arnaud, 2001 was reported from Colombia without accurate locality. Phanaeus arletteae Arnaud, 2018 was modified from Arnaud (2018).