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.