Coprophanaeus, Olsoufieff, 1924

Edmonds, W. D. & Zidek, J., 2010, A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae), Insecta Mundi 2010 (129), pp. 1-111 : 68-69

Coprophanaeus (C.) pluto ( Harold, 1863)

Fig. 151-153, 161 , 166-167, 169-170 , 172 , 195-199

Phanaeus pluto Harold, 1863: 164

Coprophanaeus pluto (Harold) (recomb. by Edmonds 1972: 843)

Phanaeus morio LeConte, 1863: 36 (syn. by Gillet 1911: 85)

Coprophanaeus pluto nogueirai Arnaud, 2002b: 2 New Synonymy

Type. P. pluto – lectotype male (des. by Arnaud 1982a: 115), Muséum National d’Histoire Naturelle, Paris (examined by photo; see Comments); P. morio – holotype male, Museum of Comparative Zoology, Harvard University, Cambridge (examined by photo). C. nogueirai – holotype male, private collection of Patrick Arnaud, Saintry sur Seine, France.

Diagnosis. General – Black except for green to yellowish-green metallic coloring on pronotum and pygidium. Clypeal teeth normal, length about equal to width at base ( Fig.161 ). Length of frons about equal to that of clypeus. Eyes normal, width of upper portion one-fourth to one-fifth of interocular distance. Frons smooth, at most with faint puncturing behind transverse carina. Paraocular area completely sculptured, sculpturing weakening closer to eye. Posteromedian portion of pronotum distinctly punctured, punctures reaching posterior margin; basal pronotal fossae distinct. Anterior angle of metasternum with rounded cap (seen in profile, Fig. 167 ). Elytral interstriae at most only weakly convex ( Fig. 166 ). Pygidium with wide basal groove. Length 14-25 mm.

Male ( Fig. 151-153 , 169-170 , 197-199 ) – Lateral tubercles of cephalic carina small, acute; middle tubercle variable: a) same size and shape as laterals ( Fig. 151 ), b) elongate, about twice as high as laterals ( Fig. 152 ), or c) lengthened, broadened and truncate or weakly emarginate apically ( Fig. 153 ). Pronotal prominence of large male cleat-shaped ( Fig. 197-198 ), width greater than that of emarginate anteromedian portion of pronotal margin. Parameres lacking subapical tooth ( Fig. 169 , 199 ).

Female – Transverse pronotal carina bidentate medially, followed by transverse depression with summit weakly bitumid.

Specimens examined – 377.

Distribution. All Mesoamerican provinces of Mexico ( Fig. 172 ).

Collection Records. GUATEMALA: Suchitepéquez – Variedades, 300 m. MEXICO: Aguascalientes – Auguascalientes (Sep) . Chiapas – 19 km N Mapastepec , 550 m (Aug) ; Tapachula ; Rosario Izapa (May) . Colima – Manzanillo (Aug) ; Plantanarilla , 1200 m (Jul) ; 1 km S El Limón , 870 m (Jul) . Guerrero – 7 km N Taxco (Jun) ; 45 km NE Ixtapa (Aug) . Hidalgo – Laguna Azteca (Sep) . Jalisco – El Tuito, 600 m (Jul) ; Guadalajara (Jul) ; Estación Biológica Chamela; Sierra de Manantlán , La Calera, 760 m (Oct) ; Sierra de Quila, Tecolotlán , 1580 m (Jul) ; Toanya , 950 m (Jul) ; Sierra de Manantlán , El Tigre, 700 m (Jul) ; Ejutla (la Estancia), 960 m (Oct) ; Autlán , 875 m (Jun) ; Puerto Los Mazos , 1350 m (Jul) ; Mezquitlán , 1000 m (Jul) ; El Parotal, 456 m (Sep); Zenzontla , 1000 m (Jun) ; ~ 5 km WNW Talpa de Allende , 20 o 23.893’N 104 o 51.980’W, 1263 m (Jul) . Mexico – Valle de Bravo (Sep) ; 10 km SW Valle de Bravo (Jun) . Michoacan – road between La Huacana and Ario de Rosales (Jul) . Morelos – Cuernavaca ; Tlaltizapan (Jul) ; Acamilpa (Jul) . Nayarit – Tepic (Jul) ; Mesa de Nayar (Jul) ; Jesus María. Nuevo Leon – El Diente (Oct) . Oaxaca – near Monte Albán , 17 o 03’34.0”N 96 o 45’53.4”W, 1690 m (Jul) . Queretaro – 2 km S Valle Verde , 21.501 oN 99.177 oW, 1300 m (Aug) . Sinaloa – 13 km E Villa Unión , 100 m (Jul) . San Luis Potosí – 6.5 km E Cd.Maiz (Aug) ; 80 km NNW Cd. Valles ( El Salto) (Jul) . Sonora – 29 km NW Yécora , 28 o 23’34.7”N 109 o 05’30.8”W, 1350 m (Jul) ; 32 km NW Yécora , 28 o 24’03.6”N 109 o 06’24.4”W, 1190 m (Jul) ; Alamos , 400 m (Sep) ; 75 km W Yécora , 28 o 29’09”N 109 o 21’32”W, 935 m (Aug) ; 140 km W Yécora , 28 o 36’12”N 109 o 48’06”W, 550 m (Aug) ;. Tamaulipas – 75 km E Cd. Victoria [ Villa de Casas ] (Jul) ; Gómez Farías [Estación Biológica Los Cedros], 350 m (Jul) ; 4.5 km W Gómez Farías [Altas Cimas], 900 m (Jul) ; Villa Aldama (Aug) ; Villa Allende, 90 m (Aug) . Veracruz – Palma Sola (Jun) ; 11 km W Palma Sola , 19 o 46’N 96 o 25’W, 120 m (Aug- Sep) ; Río Playa ; Córdoba (Jul) ; Actopan (Aug) ; Cerro Sonpaso , 690 m (Aug) ; Apazapan , 19 o 19’18”N 96 o 42’39”W, 280 m (Aug) ; 14 km WSW Papantla [ Plan de Hidalgo ] (May) ; Alto Lucero , 690 m (Sep) . UNITED STATES: Arizona – Santa Cruz Co., Nogales (Sep) . Texas – Cameron Co. - Sabal Palm Grove Sanctuary (Oct) ; 12.5 km N junction hwy. 106 and FM 2925 ; Laguna Atascosa National Wildlife Refuge (Nov) ; Harlingen ( Nov ). Hidalgo Co. – Edinburg .

Comments. The male cephalic carina of C. pluto is trimorphic. In the “major” state ( Fig. 153 ) the middle tubercle is greatly thickened, much higher than the laterals, and apically truncate or weakly excised. This form is characteristic of very large males along the Gulf coast of Veracruz, Tamaulipas and into southern Texas, where it occurs along with the “minor” form. In the “minor” state ( Fig. 151 ) the three tubercles are equal-sized. The minor state (described as the subspecies “ nogueirai ” by Arnaud 2002b; see below) is found in Gulf Coast populations and is the only known condition in populations along the Pacific coast. In the third state ( Fig. 152 ), occurring infrequently in isolated populations in southern Mexico, the middle tubercle is acute and about twice the length of the laterals.

This is the only species of the genus known to reach the United States, where it has been collected in extreme southern Texas and Arizona. Riley and Wolfe (2003) reported C. pluto from Cameron, Starr and Willacy counties, which, along with Hidalgo County, comprise the southern tip of Texas. Certain individuals from southern Texas exhibit the dark blue metallic color seen also in C. gilli , which they can resemble at first glance. Robinson (1948) reported C. pluto from Arizona.

We regard the lectotype in the Paris museum as the primary type of this species on the assumption that Harold’s many phanaeine types remained there in the nineteenth century in the wake of the dispute between the Paris and London museums. However, there is a specimen in the London museum labeled as holotype; in this case none of the labels is attributable to Harold himself. Both specimens are, to our eyes, assignable to C. pluto .

Arnaud (2002b) based C. pluto nogueirai (original name “ noguerai ” emended to “ nogueirai ” by Zidek 2005) on the structure of the “minor” male cephalic carina. For much of this study we were inclined to recognize Arnaud’s taxon as valid. As far as we know, large males of “ nogueirai ” from southern Mexico and the Pacific coast always have an acute middle tubercle; that is, they are always “minor.” We have found, however, that Gulf coast populations often also include, along with “major” males, individuals identical to C. nogueirai . Our conclusion is that the only defining character of C. p. nogueirai – a male cephalic carina with three acute, usually equal-sized tubercles ( Fig. 151 ) – is an expression of the normal intrapopulational variation of the male of C. pluto that has become fixed in southern and western populations of the species. The case of “ nogueirai ” poses interesting questions about male variation and the genetic status of species “ en statu nascendi ” that would be well worth an intensive population genetics study.

Figures

Figure 172. Approximate geographic distribution of Coprophanaeus (s. str.) species in the pluto species group.

Figure 151-161. Characters of the Coprophanaeus (s. str.) pluto species group. 151-153) Anterior views of male cephalic process of C. pluto. 154-157) Dorsal views of male pronotal prominence of C. chiriquensis. 158-159) Same, C. gilli. 160) Clypeal margin of C. boucardi. 161) Same, C. pluto.

Figure 162-170. Characters of the Coprophanaeus (s. str.) pluto species group. 162-163) Posterior view of pygidium of C. morenoi. 164) Same, C. chiriquensis. 165) Dorsal view left of elytron of C. solisi. 166) Same, C. pluto. 167) Metasternal prominence of C. pluto. 168) Same, C. pecki. 169) Dorsal view of parameres of C. pluto. 170) Lateral view, same.

Figure 195-199. Coprophanaeus pluto. 195-196) Female habitus. 197-198) Male habitus. 199) Aedeagus (lateral view on left; dorsal view on right).