taxonID	type	description	language	source
281287B461138564FF4AFD6906E151AB.taxon	description	(Figs. 1 – 15)	en	Moctezuma, V., Halffter, G. (2017): A New Species ofPhanaeusMacleay (Coleoptera: Scarabaeidae: Scarabaeinae) from Los Chimalapas, Oaxaca, Mexico. The Coleopterists Bulletin 71 (1): 47-56, DOI: 10.1649/0010-065X-71.1.47, URL: http://dx.doi.org/10.1649/0010-065x-71.1.47
281287B461138564FF4AFD6906E151AB.taxon	materials_examined	Type Material. Holotype male pinned with genitalia in microvial (label data in Spanish): “ Chocomanatlán, Santa María Chimalapa, OaxMx, 25 / IX / 2015 coprotrp t 8 c 31, 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 t 15 c 20, 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 t 7 c 2, 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 t 7 c 8, 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 t 8 c 32, 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 t 8 c 14, 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 t 9 c 18, 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 t 8 c 30, 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 t 7 c 10, 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 t 10 c 5, 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 t 7 c 12, 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 t 15 c 9, 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 t 7 c 19, 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 t 15 c 26, 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 t 15 c 15, 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 t 15 c 11, 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 t 15 c 34, 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 t 15 c 30, 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 t 7 c 4, 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 t 8 c 21, 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 t 15 c 21, 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 t 15 c 39, 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 t 15 c 14, 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 t 7 c 33, 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 t 15 c 40, 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 t 15 c 3, 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 t 15 c 37, 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 t 9 c 12, 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 t 15 c 13, 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 t 15 c 4, 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 t 15 c 32, 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 t 7 c 14, 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 t 15 c 18, 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 t 8 c 36, 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 t 9 c 30, 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 t 7 c 16, 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 t 15 c 10, 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 t 7 c 9, 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 t 8 c 24, 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 t 8 c 1, 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 t 7 c 22, 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 t 15 c 29, 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 t 8 c 5, 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 t 15 c 12, 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 t 9 c 24, 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 t 7 c 2, 16 ° 42 ’ 46.5 ” N, 94 ° 6 ’ 51.3 ” O, pino-encino, 1263 m, Victor Moctezuma Col. ”.	en	Moctezuma, V., Halffter, G. (2017): A New Species ofPhanaeusMacleay (Coleoptera: Scarabaeidae: Scarabaeinae) from Los Chimalapas, Oaxaca, Mexico. The Coleopterists Bulletin 71 (1): 47-56, DOI: 10.1649/0010-065X-71.1.47, URL: http://dx.doi.org/10.1649/0010-065x-71.1.47
281287B461138564FF4AFD6906E151AB.taxon	description	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.	en	Moctezuma, V., Halffter, G. (2017): A New Species ofPhanaeusMacleay (Coleoptera: Scarabaeidae: Scarabaeinae) from Los Chimalapas, Oaxaca, Mexico. The Coleopterists Bulletin 71 (1): 47-56, DOI: 10.1649/0010-065X-71.1.47, URL: http://dx.doi.org/10.1649/0010-065x-71.1.47
281287B461138564FF4AFD6906E151AB.taxon	etymology	Etymology. We dedicate this new species to the Zoque ethnic group, which inhabits the region of Los Chimalapas, where the type series was collected.	en	Moctezuma, V., Halffter, G. (2017): A New Species ofPhanaeusMacleay (Coleoptera: Scarabaeidae: Scarabaeinae) from Los Chimalapas, Oaxaca, Mexico. The Coleopterists Bulletin 71 (1): 47-56, DOI: 10.1649/0010-065X-71.1.47, URL: http://dx.doi.org/10.1649/0010-065x-71.1.47
281287B461138564FF4AFD6906E151AB.taxon	diagnosis	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).	en	Moctezuma, V., Halffter, G. (2017): A New Species ofPhanaeusMacleay (Coleoptera: Scarabaeidae: Scarabaeinae) from Los Chimalapas, Oaxaca, Mexico. The Coleopterists Bulletin 71 (1): 47-56, DOI: 10.1649/0010-065X-71.1.47, URL: http://dx.doi.org/10.1649/0010-065x-71.1.47
281287B461138564FF4AFD6906E151AB.taxon	materials_examined	Type Locality. Chocomanatlán (José López Portillo), Santa María Chimalapa, Oaxaca, Mexico.	en	Moctezuma, V., Halffter, G. (2017): A New Species ofPhanaeusMacleay (Coleoptera: Scarabaeidae: Scarabaeinae) from Los Chimalapas, Oaxaca, Mexico. The Coleopterists Bulletin 71 (1): 47-56, DOI: 10.1649/0010-065X-71.1.47, URL: http://dx.doi.org/10.1649/0010-065x-71.1.47
281287B461138564FF4AFD6906E151AB.taxon	distribution	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 %).	en	Moctezuma, V., Halffter, G. (2017): A New Species ofPhanaeusMacleay (Coleoptera: Scarabaeidae: Scarabaeinae) from Los Chimalapas, Oaxaca, Mexico. The Coleopterists Bulletin 71 (1): 47-56, DOI: 10.1649/0010-065X-71.1.47, URL: http://dx.doi.org/10.1649/0010-065x-71.1.47
