Phanaeus endymion Harold, 1863

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 : 13-18

Phanaeus endymion Harold, 1863

Figs 1D , 2B , 4 , 18E , 19E , 20

Phanaeus endymion Harold, 1863: 163 .

Phanaeus endymion – Gemminger & Harold 1869: 1017. — Bates 1887: 58. — Nevinson 1892: 3, 10; 1892 b: 34. — Olsoufieff 1924: 37, 91, 150, 164, pl. VI, fig. 7. — Islas 1942: 314, 318–319, 336, pl. 1 (in part). — Blackwelder 1944 –1957: 209 (in part). — Halffter & Matthews 1966: 40, 68, 70, 74 (in part). — Edmonds 1972: 830, figs 218–220 (in part); 1979: 99, 102–103, figs 4, 9–10 (in part); 2006: 31–32, 35–36, figs 2, 4, 6–7 (in part). — Morón 1979: 381, 423, 434, 449, fig. 20b; 1985: 63, 88, 91, 97, 107, fig. 20. — Arnaud 1982: 114; 2001: 4 (in part). — Palacios-Ríos et al. 1990: 59. — Estrada & Coates-Estrada 1991: 465. — Halffter et al. 1992: 139, 149, 153–155; 1995: 162, 164–166, 173. — Estrada et al. 1993: 48–49; 1998: 586–587, 589. — Deloya & Morón 1998: 19. — López-Guerrero & Halffter 2000: 241 (in part). — Montes de Oca 2001: 118, 123. — Avendaño-Mendoza et al. 2005: 810, 813, 815, 817. — Price 2005: 197 (in part); 2007: 17, figs 33, 52–54 (in part); 2009: 139, figs 2–4 (in part). — Krajcik 2006: 150 (in part). — Gillett 2008: 5. — Navarrete & Halffter 2008: 2880, 2888, 2893, 2895. — Halffter & Halffter 2009: 6, 8. — Díaz et al. 2010: 6–8. — Inward et al. 2011: 1677, fig. 2, appendix s2. — Edmonds & Zídek 2012: 1, 5–6 (in part). — Barragán et al. 2011: fig. 1. — Delgado et al. 2012b: 209. — Alvarado et al. 2014: tables s1–2; 2019: 175; 2020: 5. — Cancino-López et al. 2014: 137 (in part). — Bourg et al. 2016: 493– 494. — Huerta et al. 2016: 16 (in part). — Sánchez-de-Jesús et al. 2016: table A4. — Halffter 2017: fig. 7 (in part). — Lizardo et al. 2017: 273–275, 292–293, fig. 12 (in part). — Moctezuma & Halffter 2017: 47, 51–52, 55 figs 17–23 (in part). — Moctezuma et al. 2017: 114, 123, 131–132 (in part); 2019: 251–254, fig. 5 (in part). — Kohlmann et al. 2018: 79, 83, 88 (in part). — Sánchez-Hernández et al. 2018: 149–150; 2020: 229 (in part). — Santos-Heredia et al. 2018: appendix 1. — Gillett & Toussaint 2020: 11, fig. 3 (in part). — Salomão et al. 2020: 3–4, 8–9 (in part).

Phanaeus (Notiophanaeus) endymion – Edmonds 1994: 2, 8–9, 12, 19, 36, 39, 41–46, 54, 74, 101, figs 17, 209, 215–216, 221 (in part); 2003: 65 (in part). — Arnaud 2002b: 94–95 (in part). — Delgado et al. 2012a: 327. — Edmonds & Zídek 2012: 3, 13, figs 132–133, 136, 140, 143, 152–155 (in part). — Halffter & Morrone 2017: 21. — Lizardo et al. 2017: 272 (in part). — Capello & Halffter 2019: 113, figs 41a–b.

Phanaeus (Phanaeus) endymion – Edmonds 2003: 63, Fig. 2.20 (in part).

Diagnosis

Highly variable species, diagnosed by the following character combination: major male dark blue ( Fig. 4A ), dark metallic blue-green or dark metallic green dorsally ( Fig. 4B ); sometimes with carinate, distinctly developed keel in the middle of anterior pronotal margin ( Fig. 2B ); posterolateral angles of pronotum short, widened, sometimes lightly projected posteriorly ( Figs 4A–B ); elytral striae impressed basally as distinct fossae, with distinctly impressed to effaced punctures ( Fig. 4 ); endophallite copulatrix as in Fig. 1D . Major female with pronotal process concave posteriorly; middle pronotal tubercle more developed, dentiform or rounded, always more projected frontally than lateral tubercles; lateral tubercles obtusely rounded or carinate; posterior pronotal midline superficially impressed to completely effaced ( Fig. 4C ).

Type material

Lectotype (studied from photographs)

MEXICO • ♂, Arnaud 1982: 114 ( Fig. 4D ); Veracruz, Córdoba; “ P. endymion Harold. Ex-Mus ӕo E. Harold. Muséum Paris 1952 Coll. R. Oberthur. MNHN EC10553 ”; MNHN.

Paralectotype (studied from photographs)

MEXICO • 1 ♀; Veracruz; “Ex-Mus ӕo E. Harold. Museum Paris ex Coll. R. Oberthur. MNHN EC10554 ”; MNHN .

Non-type material revised (212 ♂♂, 247 ♀♀)

BELIZE – Cayo • 1 ♂; “ Chiquibul Forest Reserve, Las Cuevas Research Station . 16°44′N, 88°59′W. June 2006. BMNH {E} 2006-141. C. Gillett & J. Kitson ”; TAMU • 1 ♀; “ San Ignacio, Chiquibul Fst Res. Las Cuevas Res. Station. 16°43′N, 88°59′W, 02–17.iv.2004. Deciduous forest , baited pitfall. coll. R. Paterson. Deciduous tropical forest. Baited pitfall trap. Cattle dung. Sub-site: Grano de Oro. Selective logged forest. Trap: 5. Date: 8/4/04 ”; TAMU . – Orange Walk • 1 ♂, 1 ♀; “ 17°50′04″N; 89°03′04″W, Rio Bravo Cons Area , La Milpa , Arqueolog. Site , (#12), 11–17.VII.1996, P. Kovarik ex. Flight intercept trap.”; JLSHC • 1 ♂; “ Rio. Bravo Cons, Area , env. La Milpa ruins , 19–20.VII.1996, P.W. Kovarik, collector. La Milpa transect site #12, flight intercept trap ”; JLSHC . – Stann creek • 1 ♂; “ 16°48′40″N, 88°30′34″W. Cockscomb Basin Wildlife Sanctuary , Outlier Trail nr. Campground. 3.VIII.2006. P. W. Kovarik, coll., FIT”; JLSHC .

GUATEMALA – Alta Verapaz • 1 ♂; “ Sanahu. Fca. El Volcán. 14.VII.1988. J.P. Mauger & K. Meyer. 1200 msnm”; UVGC • 1 ♂; “ Senahú , Fca. El Volcán. 850 msnm. 19-VII-88. Col: F. Herrera ”; VMC • 2 ♀♀; “ Colector Carlos García . Heces de humano. Sta. Lucia Lachua. Cobán. 16-09-00 ”; UVGC • 1 ♂, 1 ♀; “ Santa Lucia Lachua. Junio 2001, heces humanas. Col. A. Higueros ”; UVGC • 1 ♀; same collection data as for preceding; VMC . – Huehuetenango • 1 ♀; “ Barillas, aldea Malpais . 1263m, N.1584733, W091.22285. Bosque nuboso. 21 VI 2009. Col. F. Camposeco, UV+M-Arc400w.”; VMC • 1 ♂; “ Octubre 1992. Colector J. Monzón ”; VMC . – Izabal • 1 ♂; “ Livingston. Biotopo Chocón. Machacas. 28–29 V 2002. Heces de vaca. Guamil. Col. E.B. Cano.”; UVGC • 1 ♂; “ Livingston. Biotopo Chocón. Machacas. 17–23 VI 1997. col. C.Avendaño. B. tropical húmedo inundable ”; TAMU • 1 ♀; “ Río Dulce, Fca. Talismán. 9 III 1997. G. Pereira ”; TAMU • 1 ♀; “ Cerro San Gil, Río Carbonera. 1390 msnm. III- 2 IV 2007. Col. R. Granajo ”; UVGC • 1 ♀; “ El Estor , Cuiscoyol , GCN. 27 III 2010. J.Rivers ”; UVGC • 1 ♂, 2 ♀♀; “ Cerro San Gill, Carboneras , estación biol 400 msnm. 08 IX 1999. Bosque tropical. Trampa de pescado. Col. G. Goemans ”; UVGC • 1 ♂; same collection data as for preceding; VMC • 1 ♀; “ El Estor , CGN sector 215 con tala. 24 III 2010. J.Rivers ”; UVGC • 2 ♀♀; same collection data as for preceding; VMC • 1 ♂; “ El Estor , CGN sector pino guiscoyol . 1–3 IX 2009. J.Rivers ”; VMC . – Petén • 1 ♀; “ La Libertad , Bethel. 18 IX 1995. M.Jolon. En trampeo ratones había caído. Oryxomis melesotis ”; UVGC • 1 ♀; “ La Libertad , Bethel. 14 XII 1995. Trampa de roedores. M.Jolon. ”; UVGC • 1 ♂; “ San Miguel la Palotada. 15-V-1999. Col. Mjolon ”; UVGC • 1 ♀; same collection data as for preceding; VMC • 1 ♀; “ San Miguel la Palotada. 16-III-1999. Col. Mjolon ”; UVGC • 1 ♂; “ Dolores. 17-VIII- 2003. Col. D. Ramírez ”; UVGC • 1 ♂; same collection data as for preceding; VMC • 1 ♂; “ Yaxhá. Area nucleo . 2 XII 1996. X. Leiva ”; TAMU • 1 ♂; “ Flores , El Remate. Biot. Cerro Cahuí . 7.VIII.1996. E.Cano ”; UVGC .

MEXICO – Chiapas • 1 ♀; “ Templo de las Inscripciones , Palenque , 22–VI–1993, Borde-copro-24 hrs. G. Halffter Col. ”; VMC • 3 ♂♂, 4 ♀♀; “ Palenque, Templo Inscripciones , 19/VII/1993, trap fruta #40, 17°29’0.9’’N, 94°2’49.07’’O, selva, 170 m, nocturna, G. y V. Halffter Cols. ”; GHC • 1 ♂; same collection data as for preceding; VMC • 1 ♀; “ Casa arqueólogos , Palenque , 22–VI–1993, Selva B-copro-24 hrs. G. Halffter Col. ”; GHC • 1 ♂; “ Casa Arqueólogos, Palenque , 20–VI–1993, Selva-copro A-24 hrs. G. Halffter Col. ”; GHC • 1 ♂; “ Palenque , 22–V–65, excrem. Humano.”; TAMU • 1 ♂; “ Palenque , 22– V–65, col. Halffter-Reyes. ”; TAMU • 4 ♂♂, 5 ♀♀; “ Bonampak, Archeological Site , 16°44’N, 91°05’W, 300 m, 24–26–IX–77, forest, human feces, W. D. Edmonds, col.”; TAMU • 1 ♂, 1 ♀; same collection data as for preceding; VMC • 1 ♀; “ Bonampak, Archeological Site , 16°44′N, 91°05′W, 300 m, I– IX–77, B. Kohlmann, col.”; TAMU • 1 ♂; “ Bonampak, Archeological Site , 16°44′N, 91°05′W, 300 m, 24–IX– 77, forest, carrion, B-1, W. D. Edmonds, col.”; TAMU • 1 ♂; “ Bonampak, Archeological Site , 16°44′N, 91°05′W, 300 m, 25–IX–77, forest, human feces, W. D. Edmonds, col.”; TAMU • 3 ♂♂, 2 ♀♀; “ Bonampak, Archeological Site , 16°44′N, 91°05′W, 300 m, 8–IX–77, forest, human feces, W. D. Edmonds, col.”; TAMU • 1 ♂; “ Bonampak Archeological Site , 16°44′N, 91°05′W, 8–IX–77, forest, human feces, W. D. Edmonds, col.”; TAMU • 3 ♂♂, 1 ♀; “ Bonampak , Agosto-1977, B. Kohlmann Col. ”; VMC • 1 ♀; “ Laguna Miramar , 14–15–X–1976, G. y V. Halffter, col.”; GHC • 1 ♀; “ Chansayab, Lacanja , 24–I–77, P. Reyes C., Col.”; GHC • 1 ♀; “ Lacanja, Chansayab , 26.IV.77, G. Aguirre, E. Frey, col.”; VMC • 1 ♂; “ Lacanja, Chansayab , 6–II–77, S. Gallina, Col.”; VMC • 1 ♀; “ Lacanjá-Chansayab , 300 m, 6–VIII–1977, camino a La Selva, M. Lamotte & B. Kohlmann col.”; TAMU • 1 ♂; same collection data as for preceding; VMC • 1 ♂; “ Lacanja , VIII-1977, #11, Selva alta, Bert Kohlmann, Col.”; GHC • 1 ♂, 3 ♀♀; “ Boca de Chajul , alt 110 m, 3-II-1984, M.A. Morón, C. Fragoso, F.J. Villalobos, cols.”; IEXA • 1 ♂; “4800’, 6.6 mi. W. El Bosque, VIII.25 –29.1973, Cl. For. A. Newton 542”; TAMU – Oaxaca • 1 ♀; “ Km Valle Nal. 550m 17/ 18-V-94, C. Deloya, R. Arce, coprotrampa ”; IEXA • 1 ♂; “2000’, 6 mi S Valle Nac. VII.71, A. Newton 299”; TAMU • 1 ♂, 1 ♀; “ Concepción Pápalo , 1800 m, 13-VII-97, hongos, G. Nogueira ”; IEXA • 1 ♂; “ La Chinantla , Sierra Norte . Selva mediana. 29-IIX- 2010. 400 m. Pit-fall exc. hum. T-2. Alvarado, F. Col. ”; FESC • 1 ♂; “ Disto. De Yautepec, Juquila Mixes , VIII.1973, W. Miller ”; VMC • 1 ♀; “ La Nueva Esperanza , I– IX–1980, C. Huerta-G. Quintero-E. Rivera Cols.”; GHC • 102 ♂♂, 123 ♀♀; “ Santa María y San Miguel Chimalapa , Benito Juárez/Santa María Chimalapa/San Antonio/San Francisco La Paz . Victor Moctezuma col.”; VMC . – Quintana Roo • 1 ♂, 1 ♀; “X-Can, VI-77, E. Welling, col.”; TAMU • 1 ♀; “ Puerto Morelos , 3–IX– 1984, G. y V. Halffter, cols. Selva baja-media, cebo excremento humano, noche/día”; GHC • 1 ♂; “ Carrillo Puerto , 25–26–XII–82, G. Halffter y V. Halffter, Col.”; VMC • 1 ♀; “ Carrillo Puerto , Septiembre 1984, A. Martínez, col.”; GHC . – Veracruz • 1 ♂; “ Córdoba , 13–VII–36, E. W. Torrens niger d’Ols.”; GHC • 2 ♂♂, 6 ♀♀; “ Córdoba, Guadalupe del Barreal , alt. 940 m, E. Santos, Col. NTP-1-C. 2-VIII-92. Julio”; IEXA • 4 ♂♂, 5 ♀♀; “ Córdoba, Guadalupe del Barreal , alt. 940 m, E. Santos, Col. NTP-1-C. 2-VII-92. Junio”; IEXA • 1 ♂, 1 ♀; “ Córdoba, Gpe. Barreal , alt. 940 m, E. Santos, Col. NTP- 3. X-91 ”; IEXA • 1 ♂; “ Córdoba, Gpe. Barreal , alt. 940 m, Cafetal, E. Santos, Col. NTP- 3. VIII-91 ”; IEXA • 1 ♂; “ Córdoba, Guadalupe del Barreal , alt. 940 m, E. Santos, Col. N-C. V-92 ”; IEXA • 2 ♂♂, 7 ♀♀; “ Córdoba, Guadalupe del Barreal , VIII-91, cafetal, NTP-80, E. Santos, Col.”; IEXA • 1 ♂, 1 ♀; “ Córdoba, Guadalupe del Barreal , VII-91, cafetal NTP-80, E. Santos, Col.”; IEXA • 1 ♀; “ Córdoba, Guadalupe del Barreal , X-91, cafetal NTP-80, E. Santos, Col.”; IEXA • 3 ♂♂, 3 ♀♀; “ Córdoba, Guadalupe del Barreal , IX-91, cafetal NTP-80, E. Santos, Col.”; IEXA • 1 ♂; “ Cordoba , 18–20/July/62, Terry W. Taylor. ”; TAMU • 1 ♂; “ Cordoba , 15–VII–1965, coll. C. L. Hogue. ”; VMC • 1 ♀; “ Córdoba , cafetal, 900 m. 2/03/91. E. Santos, Col.”; IEXA • 6 ♂♂, 3 ♀♀; “ Fortín de las Flores , September 1984, T. W. Taylor, col.”; TAMU • 2 ♂♂, 2 ♀♀; same collection data as for preceding; VMC • 1 ♂; “ Fortín de las Flores , 14–VII–1973, R. R. Shelling and T. W. Taylor, colls.”; TAMU • 1 ♂; “ Presidio , VII–48, A. Ramírez ”; VMC • 1 ♂; “ Presidio , VII–52, G. H”; GHC • 1 ♀; “ Presidio, V–47, A. Ramírez ”; VMC • 1 ♂; “ Presidio , I. 43 ”; TAMU • 1 ♂, 3 ♀♀; “ 22 km W. Palma Sola (19°46’N, 96°25’W) on road to Plan de las Hayas (± 800 m), 30–31/VIII/1976, W. D. Edmonds and Bert Kohlmann cols. Code: K22-HF-F. Collected in baited pitfall trap: HF=human feces; SF =swine feces; CC= carrion. Habitat: F =remains of forest; P=pasture (clearing).”; TAMU • 3 ♀♀; “ 1 km al E de Jalcomulco, NTP-80, alt. 500 m. 6-XI-91. Luis Quiroz, col.”; IEXA • 1 ♂, 1 ♀; “ Jalcomulco, Cerro Brujo , 06-VIII- 15. 400 m. CT3 Bosque, selva baja, Álvaro Hernández Rivera Col.”; FESC • 1 ♂; “ Xalapa, 30/Julio/80, Col. E. Aranda D.”; IEXA • 1 ♂, 1 ♀; “ Xalapa, Chiltoyac , 20/VI/15, 1000 m. NT2 Selva baja, Álvaro Hernández Rivera Col.”; FESC • 1 ♀; “ Xalapa, Chiltoyac , 1000 m. NT1 Bosque, Álvaro Hernández Rivera Col.”; FESC • 2 ♀♀; “ Xalapa, Chiltoyac , 20/VI/15, 1000 m. CT2 Potrero, Álvaro Hernández Rivera Col.”; FESC • 1 ♂, 1 ♀; “ Xalapa, Chiltoyac , 1000 m. CT3 Bosque, Álvaro Hernández Rivera Col.”; FESC • 2 ♂♂; “ Xalapa, Tiro de Hayas , 1200 m. NT2 Bosque, Álvaro Hernández Rivera Col.”; FESC • 1 ♀; “ Sumidero, Tiro de Hayas. 16 sep - 5 de oct 1989. L. Arellano col. Bosque mesófilo perturbado. 1366 m. NTP 80”; GHC • 1 ♂; “N.C 600”; GHC • 2 ♀♀; “ Veracruz , 1962–63, leg. Epping. ”; GHC • 1 ♂; “ Escoloa , 15 july 1972, Terry W. Taylor, coll.”; TAMU • 3 ♀♀; “ Ixtaxoquitlán, Cerro Chicahuaztla. 1540 m. VI.87. Col. L. Delgado ”; IEXA • 4 ♂♂, 4 ♀♀; “ El Trianón, Cerro La Campana y R. San Gabriel. V-VI-2004. Mesófilo. Col. exc. hum. Escobar y Pineda Cols.”; FESC • 3 ♂♂, 1 ♀; same collection data as for preceding; IEXA • 1 ♀; “ Tatahuicapan de Juárez, López Arias, Magallanes. 21-VI-2002. F. Escobar S. Col. Fragmentos de selva alta. Trampa #8. Cebo-exc. humano ”; FESC • 1 ♀; “ Tatahuicapan de Juárez , El Mirador , Magallanes . 21-VI-2002. F. Escobar S. Col. Fragmentos de selva alta. Trampa #2. Cebo-exc. humano ”; FESC • 1 ♂, 4 ♀♀; “ Catemaco, Pipiapan, Parque de la Flora y Fauna Silvestre Tropical , 600 m., VIII-1991, Selva Med. Per. F. Capistran Col. NTP”; IEXA • 1 ♀; “ Catemaco, Est. Biol. Trop. Los Tuxtlas , Bosq. Trop. Perennifolio , 17 VIII 90, M. F. Favila y A. Díaz. Cols. ”; VMC • 1 ♂; “ Los Tuxtlas. Estación Biológica UNAM. Selva alta. 01-VII- 2010. 200 m. Pit-fall exc. hum. T-12. Alvarado, F. Col. ”; FESC • 3 ♀♀; “ Los Tuxtlas , 21/VIII/87, G. Halffter y V. Halffter, Cols.”; GHC • 1 ♂; “ Guadalupe Victoria. PIA. Tm10.”; FESC • 1 ♀; “T 6m.”; FESC • 1 ♂, 1 ♀; “ Km 7.5, Carr. El Castillo, 15-oct-1994, L. Arellano y R. Sánchez cols. Alt. 1100 m, Bosque de pino/café, trampa necropermanente.”; TAMU • 1 ♀; “ Apazapan , 19°19′18″N, 96°42′39″W, VIII- 2000, 280 m, E. Montes de Oca, col.”; TAMU • 2 ♂♂; “ km 50 Las Choapas-Río Playa, 30-XII-1977, carroña, selva media, Col. B. Kohlmann ”; TAMU • 1 ♀; “ Dos Amates (cerca de Catemaco), V-62, G. Halffter leg.”; TAMU • 2 ♂♂; “ Dos Amates ( Catemaco ), IX–1967, G. Halffter. Leg. ”; GHC • 1 ♂; same collection data as for preceding; VMC • 1 ♀; “ Dos Amates , Alt. 300 m. 29-XI-67. Col. P. Reyes C. Selva tropical perturbada. Cebo excremento humano. Noche-Día.”; VMC • 1 ♀; “ Estación de Biología Tropical Los Tuxtlas , VIII–1984, A. Martínez, Col.”; GHC • 1 ♀; “ 20 km al E de Sontecomapan, 11-IV-69, H. Cabrera, col.”; GHC . – Yucatán • 1 ♂, 1 ♀; “ Municipio de Tizimín , 18-VII- 13, 200 m. Bosque (Zapotal). Colecta trampa copro pit-fall #6. Alvarado, F. Col. ”; FESC • 3 ♂♂, 2 ♀♀; “ Municipio de Tizimín , - VII- 13, 200 m. Bosque (Espita). Colecta trampa copro pit-fall #3. Alvarado, F. Col. ”; FESC • 2 ♂♂, 4 ♀♀; “ Municipio de Tizimín , 28-VII- 13, 200 m. Bosque (Santa Marta). Colecta trampa copro pit-fall #25. Alvarado F. Col. ”; FESC .

Type locality

Mexico, Veracruz, Córdoba.

Redescription

Major male

HEAD. Clypeus bidentate, black on anterior margin, dark metallic blue-green or green on posterior portion, rough sculpture. Genae dark metallic blue-green or green with rough sculpture. Front black, with dark metallic blue-green or green on portions adjacent to cephalic horn. Cephalic horn black, curved posteriorly over pronotum ( Figs 2B , 4A–B ).

PRONOTUM. Uniformly dark blue, dark metallic blue-green or dark metallic green, becoming black beneath posterolateral angles. Carinate, distinctly developed keel in the middle of anterior pronotal margin, or keel completely effaced. Disc triangular, flat, with two distinctly developed tubercles on anterior portion. Triangle with lightly granulate or nearly smooth sculpture, scabriculous, impunctate. Sides with almost smooth sculpture, scabriculous, with superficially impressed punctures. Lateral lines of triangle straight. Posterolateral angles short, widened, sometimes lightly projected posteriorly. Lateral fossae distinctly impressed. Basal fossae obtusely oval, superficially to deeply impressed. Posterior margin with superficially to deeply impressed punctures ( Figs 2B , 4A–B ).

ELYTRA. Striae fine, dark metallic blue-green, or dark metallic green, scabriculous, impressed basally as distinct fossae, with distinctly impressed to effaced punctures. If punctures distinctly impressed, each forming a distinct fossa, giving a completely roughened surface to first and second striae. If punctures effaced, interstriae completely smooth. Interstriae black, with smooth surface, scabriculous, superficially impressed to effaced punctation. Sutural margin without apical tooth ( Figs 4A–B ).

PROTIBIAE. Quadridentate with apical spine.

TERGITE VIII. Dark metallic blue-green, or dark metallic green, scabriculous; with rough, distinctly impressed punctures. Basal margin with setae variable in size.

GENITALIA. 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 ).

Minor male

Like the major male, except for the reduction of secondary sexual characters (i.e., cephalic horn, pronotal triangle and tubercles, and pronotal posterolateral angles).

Female

Similar to the male, except for the head showing a cephalic trituberculate carina with similar in size, nearly aligned, conical tubercles; frons with distinctly to superficially impressed punctures; pronotal sculpture completely smooth, with superficially impressed punctures; shining black, variable in size area on central portion of pronotal disc; pronotal process trituberculate, with posterior concavity; middle pronotal tubercle more developed, dentiform or rounded, always more frontally projected than lateral tubercles; lateral tubercles obtusely rounded or carinate; posterior pronotal midline superficially impressed to completely effaced ( Fig. 4C ).

Variation

Mean length 17.9 mm (13.9–20 mm). The typical colour morph is dark metallic blue-green ( Figs 4A, C ), but a less frequent dark metallic green morph ( Fig. 4B ) is also found, while scarce specimens are completely bright metallic green. The interstriae are completely dark metallic blue-green or dark metallic green in some specimens. The rarest specimens from Chiapas and Veracruz show a bright metallic yellow or red sheen on head, pronotum and elytra. Variation in the interstrial punctation is observed in the whole distribution area of P. endymion . The size of the right and left lobes of the endophallite copulatrix is not homogeneous.

Distribution From Veracruz and the Yucatan Peninsula to Belize and northern Guatemala ( Fig. 15 ).

Remarks

The lectotype (a minor male), a female paralectotype and several specimens (n= 42) from the type locality cited by Harold (1863) were used for the redescription of P. endymion , that was complemented with specimens from its whole distribution area. As a consequence, the high intraspecific variation of P. endymion is adequately represented herein. Although P. endymion is a polymorphic species, closely related species are confidently separated by differences in the pronotal morphology of males and females, elytral interstriae, and genital morphology.

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. 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. 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. 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. 20. Lateral view of the aedeagus of the Phanaeus endymion species group (P. endymion Harold, 1863).

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.