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 : 80-82

Coprophanaeus (C.) dardanus ( MacLeay, 1819)

Fig. 6 , 213-216 , 237 , 239-243

Phanaeus dardanus MacLeay, 1819: 126

Phanaeus bitias Harold, 1863: 163 (syn. by Nevinson 1892: 3)

Phanaeus jasion Felsche, 1901: 153 (syn. by Pessôa 1934: 300)

Phanaeus arrowi Olsoufieff, 1924: 68 (syn. by Arnaud 2002c: 32)

Coprophanaeus dardanus (MacLeay) (recomb. by Edmonds 1972: 843)

Type. P. dardanus – unknown to us (see Comments); P. bitias – unknown to us; P. jasion – holotype male, Staatliches Museum für Tierkunde, Dresden; P. arrowi – lectotype male (des. by Arnaud 1982: 116), Muséum National d’Histoire Naturelle, Paris (examined by photo).

Diagnosis. General – Dorsum dark, sombre, metallic reflections restricted to pronotum and head ( Fig. 239-242 ). Clypeal margin distinctly angulate adjacent to median teeth. Length of clypeus about equal to that of frons. Pronotal disk lacking midlongitudinal sulcus, posterior portion smooth to very weakly punctured. Cap of anterior metasternal angle usually simple, sometimes slightly forked; anterior surface of metasternum smooth above anterior margin of disk. Elytral striae not distinctly carinulate. Length 14- 29 mm.

Male – Head horn ( Fig. 213-215 ) erect, plate-like, gently curved posteriorly; width approaching interocular distance, sides parallel to slightly divergent; apex bidentate, teeth elongate, widely separated by deep, rounded emargination. Pronotal prominence ( Fig. 241-242 ) cleat-like, strongly bilobate, lobes directed anterolaterally and flanked by deep, oval concavities, lateral margin of each concavity with weak tumosity above lateral fossa; width of prominence slightly greater than distance between outer margins of eyes. Apical processes of parameres strong, hook-like.

Female – Cephalic carina ( Fig. 216 ) widely bidentate, apical margin between teeth widely raised, weakly angulate. Transverse crest of pronotum rounded, distinctly bidentate medially, followed by weak depression with feebly bitumose summit.

Specimens examined – 535.

Distribution. Northern Amazonian subregion and coastal Brazil ( Fig. 237 ).

Collection Records. BRAZIL: Amapá – Serra do Navio [Cava Urucum-Amapari], 00 o 53’06”N 51 o 52’53”W (Jan, Sep). Amazonas – 60 km N Manaus [Fazenda Esteio] Bahia – Porto Seguro (Jan) . Ceará – Maranguape, Piraponga , 38 o 41’59"W 03 o 53’26"S, 200 m ; Baturite, Uirapuru , 38 o 54’22"W 04 o 17’27"S, 620 m (Mar) . Goiás – Jaraguá. Pará – Redenção , 7 o 46’S 51 o 58’W (Oct) ; Belêm (Jan-Mar); Alidos (Mar); Monte Dourado , 0 o 42’S 52 o 38’W, 100 m (Mar-Apr) ; 45 km E Canindé (Dec) . Paraíba – João Pessoa [ Mata do Buraquinho ] (Aug) ; Aréia [Mata do Pau Ferro], 6 o 58’S 35 o 42’W (Apr) . Paraná – Londrina (Apr). Pernambuco – Cabo [Reserva Ecologica Gurjaú] (May, Oct). Rio de Janeiro – Itatiaia , 700 m (Oct, Dec) ; Rio de Janeiro [Parque Lage] (Feb); Botafogo (Mar, Nov); Floresta da Tijuca (Dec) ; Petropolis (Sep, Dec). Roraima – Caracaraí (Jul). Santa Catarina – Corupá (Oct-Nov); Joinville (Apr, Jun). CO- LOMBIA: Guaviaré – San José [Finca La Esmeralda], 2 o 33’N 72 o 38’W, 240 m (Oct) . FRENCH GUIANA: 8.4 km SSE Roura , 4 o 40’41”N 52 o 13’25”W, 200 m (May-Jun) ; 7 km N Saül [ Les Eaux Claires ], 3 o 39’46”N 53 o 13’19”W (May-Jun) ; Wanaboo [Marowijne River], 4 o 43’35”N 54 o 26’36”W, 40 m (Jun) ; 9.7 km NW – 4.3 km SE Patawa [ Kaw Mountain ], 4 o 32’40.1”N 52 o 09’08.1”W (Dec) ; Mana , 5 o 39.2’N 53 o 49.9’W (Jul) ; 100 km S Cayenne [ Nouragues Research Station ], 4 o 05’N 52 o 40’W . GUYANA: Mazaruni-Potaro – Takutu Mts., 6 o 15’N 59 o 05’W (Dec). Potaro-Siparuni – Iwokrama Forest Reserve , 4 o 40’19”N 58 o 41’04”W, 100- 200 m (Jun) . SURINAME: Brokopondo – Stoneiland , ~ approx. 4 o 59’N 55 o 09’W, 30 m (Apr-May) ; Rosebel gold mining area, ~ 5 o 07' N ; 55 o 17 W (May). Para – ~ 20 km SSE Joden Savanne, 5 o 16’17”N 54 o 55’15”W, 40 m (Jun-Jul). Paramaribo – Pallisadenweg (Jul). Sipaliwini – Central Suriname Nature Reserve , environs of Voltzberg research station, 4 o 40.90”N 56 o 11.13'’ W, 100 m (Mar) ; Central Suriname Nature Reserve, environs of Lolopasi field station, 4 o 42.91’N 56 o 12.83’W, 80 m (Mar) ; Central Suriname Nature Reserve, environs of Conservation International field station, 50 m (Mar) ; Central Suriname Nature Reserve, Raleighvallen, Fungu Island (= Foengoe Island ), ~ 4 o 33’30"N 56 o 12’21"W (Jul) ; Bakhuis mountains , ~ 4 o 27’13"N 56 o 57’39"W (Nov) ; Palumeu,~3 o 20’46.3'’N 55 o 26’30.8'’W (Jun). Wanica – Lelydorp (Feb); Zanderij, Hannover (Jun). TRINIDAD-TOBAGO: Port-of-Spain [ Capuro Valley ] ; 16 km N Arima , 650 m (Jun) ; 8 km N Arima , 260 m ; Maracas Valley , 600 m ; 11 km SE Arima [ Arena Forest Reserve ], 80 m (Jun) . VENEZUELA: Amazonas – Puerto Ayacucho (May) ; El Infierno (Jun) ; Atabapo (Jun); Camani (Jun-Jul). Aragua – Maracay [Rancho Grande] (Jun). Bolívar – 50 km SE El Manteco [ Río El Supamo ] (Dec) ; Paují , 1100 m ; 10 km E San Francisco Yuruani , 1300 m (Jul) ; 20 km S Luepa , 1500 m (Jul) ; 10 km E Kavanayen , 1500 m (Jun) ; 22 km SE El Dorado (Jul) ; 10 km S El Dorado , 200 m (Jul-Aug) ; 100 km S El Dorado , 350 m (Jul-Aug) ; 6 km S San Isidro (Jun-Jul) ; Parupa , 1500 m (Jun-Jul). Miranda – 33 km N Altagracia de la Montaña [Guatopo National Park, La Crucita], 400 m (Jun). Monagas – 27 km SW Caripe, 300 m (Jun) ; Caripe , 700 m (Jul) .

Comments. We assume that the type, or a candidate for the lectotype of this species, will be found in the MacLeay Collection at the University of Sydney. We were unable to secure specimens from Sydney for this study. However, there is little room for doubt about the identity of C. dardanus even from the brief description of MacLeay (1819: 126-127): “P. nigro-viridis, capitis cornubus duobus brevibus basi connatis, thorace antice carinâ bidentatâ instructo … Mas. Thorax fossulâ utrinque sub carinae dentibus unidentatâ …” (Free translation: Black with green, head horn with two teeth broadly connected at the base; thorax with bidentate process … Male: Lateral concavity beneath median process with single tooth …”). His diagnosis highlights the bidentate structure of the male head horn ( Fig. 213-214 ), which is unique to the genus and seen otherwise in phanaeines only in Diabroctis . The length of the teeth varies, but usually amounts to at least the height of the base. In large individuals the upper margin of the base, between the lateral teeth, is slightly raised medially. In very small males ( Fig. 215 ) the horn is reduced to a weakly bidentate swelling reminiscent of the small males of Oxysternon and certain Phanaeus . The cephalic carina of the female ( Fig. 216 ), even in small individuals, is also unique and highly diagnostic. It bears a small denticle at each end, separated by a broad, rounded or slightly angulate elevation.

We regard C. dardanus as a fundamentally Amazonian species, but it has managed to spread southward via the Atlantic coast of Brazil into portions of the Paranaian subregion as far south as Santa Catarina as well as into the central highlands (via gallery forests?). Remarkably, as far as we can see, the vast occupation of its combined range has not resulted in any detectable geographic variation. We have not seen Arnaud’s (2002a) species, C. terrali , which was based on a single male from Mato Grosso and considered a close ally of C. dardanus . The single reported specimen (holotype) is part of a private collection that was not available for study. The shape of the cephalic carina of the holotype is strikingly different from that expected for C. dardanus , and the possibility that the shape is a teratologic form or a distinct morph in otherwise normal populations needs to be ruled out in any assessment of its validity.

Figures

Figure 237. Approximate geographic distribution of Coprophanaeus (s. str.) species in the dardanus species group A.

Figure 5-12. Characters of Coprophanaeus. 5) Ventral view of front left tibia of C. ensifer (arrow marks elongate tuft of setae). 6) Same, C. dardanus. 7) Left anterolateral pronotal margin of C. spitzi (arrow indicates unbroken circumnotal carina. 8) Same, C. magnoi (arrow indicates broken circumnotal carina). 9) Base of left hind wing of C. cerberus. 10) Same, C. horus. 11) Lateral view left occipital lobe of C. ensifer (asterisk marks swelling). 12) Same, C. spitzi (asterisk marks flat area).

Figure 213-227. Characters of the Coprophanaeus (s. str.) dardanus species group. 213-214) C. dardanus, anterior view of cephalic process of large male. 215) Same, small male. 216) Same, female. 217) C. telamon, aedeagus (lateral view above; dorsal view below). 218-219) C. corythus, anterior view of male cephalic process. 220) C. telamon, same. 221-222) C. telamon, lateral view of male cephalic process (arrow in 222 indicates posterior tubercle). 223) C. corythus, lateral view of male cephalic carina. 224) C. degallieri, metasternal shield. 225-226) C. milon, anterior view of male cephalic process. 227) C. magnoi, same.

Figure 239-243. Coprophanaeus dardanus. 239-240) Female habitus. 241-242) Male habitus. 243) Aedeagus (dorsal view on left; lateral view on right).