Phanaeus (Notiophanaeus) zoque Moctezuma and Halffter, 2017

Moctezuma, V. & Halffter, G., 2017, A New Species ofPhanaeusMacleay (Coleoptera: Scarabaeidae: Scarabaeinae) from Los Chimalapas, Oaxaca, Mexico, The Coleopterists Bulletin 71 (1), pp. 47-56 : 48-55

Phanaeus (Notiophanaeus) zoque Moctezuma and Halffter , new species

( Figs. 1–15 )

Type Material. Holotype male pinned with genitalia in microvial (label data in Spanish): “Chocomanatlán, Santa María Chimalapa, OaxMx, 25/IX/2015 coprotrp t8c31, 16° 40’ 31.3” N, 94° 3’ 57.3” O, Bosque pino, 1219 m, Victor Moctezuma Col.” . Paratypes (108 males, 106 females): 2 males, 1 female, same data as the holotype ; 4 males, 5 females, “ Benito Juárez, San Miguel Chimalapa, Oaxaca , Mex. 17/X/2015 coprtrp t15c20, 16° 45’ 3.6” N, 94° 9’ 12.3” O, Bosque pino, 1091 m, Victor Moctezuma Col.” ; 1 male, 1 female, “ Benito Juárez, San Miguel Chimalapa, Oaxaca, Mex. 24/IX/2015 coprotrap t7c2, 16° 42’ 46.5” N, 94° 6’ 51.3” O, pino-encino, 1263 m, Victor Moctezuma Col.” ; 1 male, 1 female, “ Benito Juárez, San Miguel Chimalapa, Oaxaca, Mex. 24/IX/2015 coprotrap t7c8, 16° 42’ 58.1” N, 94° 6’ 59.5” O, pino-encino, 1211 m, Victor Moctezuma Col.” ; 3 males, 1 female, “ Chocomanatlán, Santa María Chimalapa, OaxMx , 25/IX/2015 coprotrp t8c32, 16° 40’ 32.5” N, 94° 3’ 57” O, Bosque pino, 1219 m, Victor Moctezuma Col.” ; 1 male, “ Chocomanatlán, Santa María Chimalapa, OaxMx , 25/IX/2015 coprotrp t8c14, 16° 40’ 10.2” N, 94° 4’ 0.2” O, Bosque pino, 1273 m, Victor Moctezuma Col.” ; 1 male, “ Benito Juárez, San Miguel Chimalapa, Oaxaca, Mex. 28/IX/2015 coprotrp t9c18, 16° 41’ 41.3” N, 94° 7’ 45.6” O, pino-encino, 1191 m, Victor Moctezuma Col.” ; 2 males, 1 female, “ Chocomanatlán, Santa María Chimalapa, OaxMx , 25/IX/2015 coprotrp t8c30, 16° 40’ 30.1” N, 94° 3’ 56.7” O, Bosque pino, 1218 m, Victor Moctezuma Col.” ; 2 males, 2 females, “ Benito Juárez, San Miguel Chimalapa, Oaxaca, Mex. 24/IX/2015 coprotrp t7c10, 16° 42’ 57.2” N, 94° 6’ 56.7” O, pinoencino, 1224 m, Victor Moctezuma Col.” ; 6 males, 1 female, “ Benito Juárez, San Miguel Chimalapa, Oaxaca, Mex. 1/X/2015, coprotrp t10c5, 16° 44’ 4.8” N, 94° 8’ 15.6” O, pino-encino, 918 m, Victor Moctezuma Col.” ; 3 males, 3 females, “ Benito Juárez, San Miguel Chimalapa, Oaxaca, Mex. 24/ IX/2015 coprotrp t7c12, 16° 42’ 55.4” N, 94° 6’ 54.4” O, pino-encino, 1232 m, Victor Moctezuma Col.” ; 2 males, 1 female, “ Benito Juárez, San Miguel Chimalapa, Oaxaca, Mex. 17/X/2015 coprtrp t15c9, 16° 45’ 3.1” N, 94° 8’ 58.2” O, Bosque pino, 996 m, Victor Moctezuma Col.” ; 1 male, “ Benito Juárez, San Miguel Chimalapa, Oaxaca, Mex. 24/IX/2015 coprotrp t7c19, 16° 42’ 46.5” N, 94° 6’ 51.3” O, pino-encino, 1263 m, Victor Moctezuma Col.” ; 1 female, “ Benito Juárez, San Miguel Chimalapa, Oaxaca, Mex. 17/X/2015 coprtrp t15c26, 16° 45’ 8.1” N, 94° 9’ 18.8” O, Bosque pino, 1126 m, Victor Moctezuma Col.” ; 8 males, 4 females, “ Benito Juárez, San Miguel Chimalapa, Oaxaca, Mex. 17/X/2015 coprtrp t15c15, 16° 45’ 2.3” N, 94° 9’ 6.1” O, Bosque pino, 1050 m, Victor Moctezuma Col.” ; 2 males, 4 females, “ Benito Juárez, San Miguel Chimalapa, Oaxaca, Mex. 17/X/2015 coprtrp t15c11, 16° 45’ 2” N, 94° 9’ 0.7” O, Bosque pino, 1020 m, Victor Moctezuma Col.” ; 5 males, 4 females, “ Benito Juárez, San Miguel Chimalapa, Oaxaca, Mex. 17/X/ 2015 coprtrp t15c34, 16° 45’ 16.4” N, 94° 9’ 23” O, Bosque pino, 1233 m, Victor Moctezuma Col.” ; 2 males, 3 females, “ Benito Juárez, San Miguel Chimalapa, Oaxaca, Mex. 17/X/2015 coprtrp t15c30, 16° 45’ 12.3” N, 94° 9’ 22.1” O, Bosque pino, 1168 m, Victor Moctezuma Col.” ; 4 males, 1 female, “ Benito Juárez, San Miguel Chimalapa, Oaxaca, Mex. 24/IX/2015 coprotrp t7c4, 16° 42’ 59” N, 94° 7’ 5.1” O, pino-encino, 1196 m, Victor Moctezuma Col.” ; 2 males, “ Chocomanatlán, Santa María Chimalapa, OaxMx , 25/IX/2015 coprotrp t8c21, 16° 40’ 19.1” N, 94° 4’ 0.5” O, Bosque pino, 1259 m, Victor Moctezuma Col.” ; 4 females, “ Benito Juárez, San Miguel Chimalapa, Oaxaca, Mex. 17/X/2015 coprtrp t15c21, 16° 45’ 4.7” N, 94° 9’ 13” O, Bosque pino, 1094 m, Victor Moctezuma Col.” ; 5 males, 5 females, “ Benito Juárez, San Miguel Chimalapa, Oaxaca, Mex. 17/X/2015 coprtrp t15c39, 16° 45’ 20.8” N, 94° 9’ 27.8” O, Bosque pino, 1250 m, Victor Moctezuma Col.” ; 5 males, 7 females, “ Benito Juárez, San Miguel Chimalapa, Oaxaca, Mex. 17/X/2015 coprtrp t15c14, 16° 45’ 2.3” N, 94° 9’ 6.1” O, Bosque pino, 1050 m, Victor Moctezuma Col.” : 1 female, “ Benito Juárez, San Miguel Chimalapa, Oaxaca, Mex. 24/IX/2015 coprotrp t7c33, 16° 42’ 34” N, 94° 6’ 35.9” O, pino-encino, 1325 m, Victor Moctezuma Col.” ; 1 male, 2 females, “ Benito Juárez, San Miguel Chimalapa, Oaxaca, Mex. 17/ X/2015 coprtrp t15c40, 16° 45’ 21.7” N, 94° 9’ 28.8” O, Bosque pino, 1261 m, Victor Moctezuma Col.” ; 2 males, 2 females, “ Benito Juárez, San Miguel Chimalapa, Oaxaca, Mex. 17/X/2015 coprtrp t15c3, 16° 45’ 4.7” N, 94° 8’ 50.5” O, Bosque pino, 934 m, Victor Moctezuma Col.” ; 1 female, “ Benito Juárez, San Miguel Chimalapa, Oaxaca, Mex. 17/X/2015 coprtrp t15c37, 16° 45’ 19” N, 94° 9’ 25.8” O, Bosque pino, 1249 m, Victor Moctezuma Col.” ; 1 male, 3 females, “ Benito Juárez, San Miguel Chimalapa, Oaxaca, Mex. 28/IX/2015 coprotrp t9c12, 16° 41’ 45.5” N, 94° 7’ 52.2” O, pinoencino, 1244 m, Victor Moctezuma Col.” ; 6 males, 14 females, “ Benito Juárez, San Miguel Chimalapa, Oaxaca, Mex. 17/X/2015 coprtrp t15c13, 16° 45’ 2” N, 94° 9’ 3.4” O, Bosque pino, 1037 m, Victor Moctezuma Col.” ; 10 males, 3 females, “ Benito Juárez, San Miguel Chimalapa, Oaxaca, Mex. 17/ X/2015 coprtrp t15c4, 16° 45’ 2” N, 94° 9’ 0.7” O, Bosque pino, 1020 m, Victor Moctezuma Col.” ; 1 male, “ Benito Juárez, San Miguel Chimalapa, Oaxaca, Mex. 17/X/2015 coprtrp t15c32, 16° 45’ 14.1” N, 94° 9’ 23.9” O, Bosque pino, 1202 m, Victor Moctezuma Col.” ; 1 male, 1 female, “ Benito Juárez, San Miguel Chimalapa, Oaxaca, Mex. 24/ IX/2015 coprotrp t7c14, 16° 42’ 52.7” N, 94° 6’ 53.5” O, pino-encino, 1226 m, Victor Moctezuma Col.” ; 2 males, 8 females, “ Benito Juárez, San Miguel Chimalapa, Oaxaca, Mex. 17/X/2015 coprtrp t15c18, 16° 45’ 2.5” N, 94° 9’ 10” O, Bosque pino, 1084 m, Victor Moctezuma Col.” ; 7 males, 1 female, “ Chocomanatlán, Santa María Chimalapa, OaxMx , 25/IX/2015 coprotrp t8c36, 16° 40’ 37.1” N, 94° 3’ 54.6” O, Bosque pino, 1206 m, Victor Moctezuma Col.” ; 3 females, “ Benito Juárez, San Miguel Chimalapa, Oaxaca, Mex. 28/IX/2015 coprotrp t9c30, 16° 41’ 30.7” N, 94° 7’ 34.6” O, pino-encino, 1215 m, Victor Moctezuma Col.” ; 1 male, 2 females, “ Benito Juárez, San Miguel Chimalapa, Oaxaca, Mex. 24/ IX/2015 coprotrp t7c16, 16° 42’ 50.1” N, 94° 6’ 52.7” O, pino-encino, 1233 m, Victor Moctezuma Col.” ; 1 male, “ Benito Juárez, San Miguel Chimalapa, Oaxaca, Mex. 17/X/2015 coprtrp t15c10, 16° 45’ 2.5” N, 94° 8’ 59.4” O, Bosque pino, 1012 m, Victor Moctezuma Col.” ; 1 male, 4 females, “ Benito Juárez, San Miguel Chimalapa, Oaxaca, Mex. 24/ IX/2015 coprotrap t7c9, 16° 42’ 57.5” N, 94° 6’ 58.1” O, pino-encino, 1214 m, Victor Moctezuma Col.” ; 3 males, 1 female, “ Chocomanatlán, Santa María Chimalapa, OaxMx , 25/IX/2015 coprotrp t8c24, 16° 40’ 22.8” N, 94° 3’ 59.3” O, Bosque pino, 1251 m, Victor Moctezuma Col.” ; 2 males, 1 female, “ Chocomanatlán, Santa María Chimalapa, OaxMx , 25/IX/2015 coprotrp t8c1, 16° 39’ 53.3” N, 94° 4’ 3.3” O, Bosque pino, 1325 m, Victor Moctezuma Col.” ; 1 male, 1 female, “ Benito Juárez, San Miguel Chimalapa, Oaxaca, Mex. 24/ IX/2015 coprotrp t7c22, 16° 40’ 20.3” N, 94° 3’ 59.9” O, pino-encino, 1259 m, Victor Moctezuma Col.” ; 2 females, “ Benito Juárez, San Miguel Chimalapa, Oaxaca, Mex. 17/X/2015 coprtrp t15c29, 16° 45’ 11.3” N, 94° 9’ 21.1” O, Bosque pino, 1154 m, Victor Moctezuma Col.” ; 1 female, “ Chocomanatlán, Santa María Chimalapa, OaxMx , 25/IX/2015 coprotrp t8c5, 16° 39’ 57.6” N, 94° 3’ 59.9” O, Bosque pino, 1295 m, Victor Moctezuma Col.” ; 1 male, 1 female, “ Benito Juárez, San Miguel Chimalapa, Oaxaca, Mex. 17/X/2015 coprtrp t15c12, 16° 45’ 1.8” N, 94° 9’ 2.1” O, Bosque pino, 1026 m, Victor Moctezuma Col.” ; 2 males, 2 females, “ Benito Juárez, San Miguel Chimalapa, Oaxaca, Mex. 28/IX/2015 coprotrp t9c24, 16° 41’ 37.4” N, 94° 7’ 38.8” O, pinoencino, 1181 m, Victor Moctezuma Col.” ; 3 males, 2 females, “ Benito Juárez, San Miguel Chimalapa, Oaxaca, Mex. 24/IX/2015 coprotrap t7c2, 16° 42’ 46.5” N, 94° 6’ 51.3” O, pino-encino, 1263 m, Victor Moctezuma Col.” .

Description. Holotype, major male. Length 13.9 mm (including head), width at bases of elytra 8.5 mm ( Fig. 1 ). Head: Clypeus black with metallic green, anterior margin strongly bidentate, clypeal process transverse ( Figs. 2–3 ). Genae metallic green with rugose sculpturing. Frons with black cephalic horn, slender and strongly curved over pronotum ( Fig. 3 ); lateral parts of frons metallic green, with weakly impressed rugose sculpturing and fine, well-spaced punctures. Pronotum: Triangular pronotal disc with lateral fossae, metallic green with black spots anterolaterally ( Figs. 2–3 ); anterior margin projecting upwards, acute posterolateral angles projecting laterally ( Fig. 4 ); disk sculpturing shagreened, becoming weakly impressed anterolaterally; basal fossae oval, deeply impressed ( Fig. 5 ). Elytra: Striae shagreened, metallic green, with punctures ( Fig. 1 ); black interstriae with green reflections and smooth sculpturing. Pygidium: Metallic green, with coarse punctures ( Figs. 6, 7 ); basal margin with almost imperceptible setae and rounded angles. Protibia: Quadridentate with a moreor-less straight apical spur ( Figs. 3, 4 ). Aedeagus: Similar ( Fig. 14 ) to that of Phanaeus endymion Harold ; internal sack with lamella copulatrix in medial position ( Fig. 15 ). Lamella copulatrix not displaying any intraspecific morphological variation.

Variation. Minor male: Similar to major male except cephalic horn smaller or completely reduced to a frontal carina with a small indentation; frons with weakly impressed fine punctures; posterolateral angles of pronotum reduced ( Figs. 8, 9 ). Female: Head black with green reflections in posterior part of frons and genae; cephalic carina trituberculate; pronotum dull black, trituberculate, with metallic green towards border, smooth, with fine, evenly spaced punctures ( Figs. 10, 11, 13 ). Some specimens may be completely black. Chromatic variation is very rare (10 paratypes = 4.7% of type material). Males and females can show an olive green color with golden ( Fig. 12 ) or reddish ( Fig. 13 ) reflections. Size of paratypes: Length 14.1– 10.2 mm, width 8.7– 5.8 mm.

Etymology. We dedicate this new species to the Zoque ethnic group, which inhabits the region of Los Chimalapas, where the type series was collected.

Diagnosis. Phanaeus zoque is easily recognized from other species of the endymion speciesgroup by its geographic distribution ( Fig. 23 ) and typical metallic green color. Phanaeus halffterorum Edmonds is bright green or dark blue with a strongly granulated pronotal sculpturing. Phanaeus zapotecus Edmonds is usually black with posterolateral angles projected backwards. Phanaeus pyrois Bates has a velvety smooth sculpturing, green, red, or blue pronotum, but the elytra are always black. Phanaeus zoque closely resembles the green form of P. endymion , which lacks the golden/reddish reflections ( Figs. 12, 13 ). A constant character that distinguishes both males and females of the two species is that the pygidium of P. endymion has thick setae on the basal margin ( Fig. 22 ), whereas in P. zoque the pygidium bears almost unapparent setae ( Fig. 7 ). The endophallic sclerites of both species also show differences (compare Figs. 15 and 16 ). In addition, P. zoque is smaller (14.1– 10.2 mm; Fig. 1 ) than P. endymion (11–20 mm; Fig. 17 ), and its cephalic horn is more slender (compare Figs. 2 and 18 ). Also, the anterior margin of the pronotum of P. zoque projects upwards, while that of P. endymion projects forwards (compare Figs. 2 and 18 ). The posterolateral angles of pronotum are acute in P. zoque ( Figs. 3, 4 ), but rounded in P. endymion ( Figs. 19, 20 ). And the basal margin of the pygidium has two acute angles in P. endymion , while in P. zoque these are rounded (compare Figs. 6 and 21 ).

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

Distribution and Ecology. This new species is known from two localities, Chocomanatlán (José López Portillo) and Benito Juárez, in the region of Los Chimalapas, Oaxaca ( Fig. 23 ). Phanaeus zoque has been collected only in dung-baited pitfall traps in temperate pine and pine-oak forests between 918 and 1,325 m elevation. According to preliminary data, P. zoque is the dominant species of dung beetle that inhabits the surveyed localities at which it represents 76% of collected dung beetles. Associated scarab taxa are Canthidium Erichson (<1% of abundance), Canthon Hoffmannsegg (4%), Deltochilum scabriusculum Bates (1%), Eurysternus magnus Castelnau (1%), Ontherus mexicanus Harold 1%), Onthophagus Latreille (12%), Phanaeus tridens Laporte (<1%), and Scatimus ovatus Harold (4%).

KEY TO SPECIES OF THE ENDYMION SPECIES- GROUP OF PHANAEUS (modified from Edmonds and Zídek 2012)

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 the apical angle; elytral margin slightly excised adjacent to this tooth. Color bright green or dark blue. South-central Mexico (states of Estado de México and Guerrero) ............. P. halffterorum Edmonds

1′. Sutural margin of elytra simple. Color and distribution variable ................................... 2

2. Triangular pronotal disk of male evenly and densely but finely granulate (10X), granules in most specimens larger and becoming squamose along lateral margins of disk and extending onto posterolateral angles (when distinctly developed); sides of pronotum roughened (10X), lacking distinct punctures except behind lateral fossae. Female pronotum minutely roughened, evenly, distinctly punctate (10X), punctures becoming smaller dorsally but not disappearing altogether; disk impressed medially as a distinct furrow visible to unaided eye, extending forward from posterior margin to near middle of disk. Sierra Madre del Sur of south-central Oaxaca ............... ........................................... P. zapotecus Edmonds

2′. Pronotal disk of male either lacking distinct granulation, or, if granules present, these are minute and restricted along lateral margins of disk; sides of pronotum smooth, minutely punctate. Female pronotum (fig. 140 in Edmonds and Zídek 2012) smooth, punctures (50X) fine and usually restricted to sides; median furrow lacking or at most indicated by fine, scarcely visible line ...................................... 3

3. Elytral interstriae distinctly flattened and uniformly dull (more convex and shiny in some Central American populations); striae not strongly impressed basally, anterior ends in most specimens bearing deep punctures rather than large fossae. Male: Pronotal disk dull, velvety smooth medially, finely asperate, brighter laterally. Female: Pronotum evenly convex, lacking anteromedial concavity even in largest specimens, bearing 3 round, smooth tubercles in transverse line near anterior margin. Head and pronotum largely highly shiny metallic red to nearly completely dull black with metallic red restricted to ridges and isolated areas on anterior part of pronotum; elytra dull to weakly shiny black; pygidium usually metallic red medially, green peripherally, in some specimens completely red or green. Southern Nicaragua through Central America into western Colombia and Ecuador................................................. P. pyrois Bates

3′. Elytral interstriae evenly convex and glossy mid-longitudinally; striae impressed basally as distinct fossae. Male: Pronotal disk velvety smooth medially, finely aspirate laterally and sometimes also medially. Female: Pronotum with anteromedial concavity bounded anteriorly by a raised U- or V-shaped ridge ...................................... 4

4. Dorsum shiny green or dark blue; in a few specimens, shiny green with strong yellow reflections. Anterior margin of pronotum projecting forwards. Relatively rounded posterolateral angles of pronotum. Basal margin of pygidium with slender setae. Internal sack of aedeagus and lamella copulatrix as in Fig. 16 . Southwestern Mexico and southeastern Mexico to Honduras........................ ..................................... P. endymion Harold

4′. Dorsum metallic green; in a few specimens, olive green with golden/reddish reflections. Anterior margin of pronotum projecting upwards. Posterolateral angles of pronotum acute. Basal margin of pygidium with almost imperceptible setae. Internal sack of aedeagus and lamella copulatrix as in Fig. 15 . Eastern Oaxaca ................. P. zoque Moctezuma and Halffter , new species

Figures

Fig. 1. Phanaeus zoque, holotype, major male.

Figs. 2–7. Phanaeus zoque, holotype. 2) Head and prothorax, lateral view; 3) Head and pronotum, dorsal view; 4) Pronotal posterolateral projection and protibia; 5) Pronotal basal fossae; 6) Pygidium; 7) Basal margin of pygidium with almost imperceptible setae.

Figs. 8–13. Phanaeus zoque, paratypes. 8) Minor male, dorsal view; 9) Minor male head and prothorax, lateral view; 10) Female, dorsal view; 11) Female head and pronotum, dorsal view; 12) Male head and pronotum, dorsal view, with golden reflections; 13) Female head and pronotum, dorsal view, with reddish reflections.

Figs. 14–16. Phanaeus species. 14) P. zoque, aedeagus; 15) P. zoque, internal sac and lamella copulatrix; 16) P. endymion, internal sac and lamella copulatrix.

Fig. 23. Known distribution of the Mexican species of the endymion species-group (modified from Edmonds 2006). Red dots = P. zoque; yellow dots = P. endymion; blue dots = P. zapotecus; white dots = P. halffterorum; solid line = national borders.

Figs. 17–22. Phanaeus endymion. 17) Major male, dorsal view; 18) Head and prothorax, lateral view; 19) Head and pronotum, dorsal view; 20) Pronotal posterolateral projection and front tibia; 21) Pygidium; 22) Basal margin of pygidium with slender setae.