Pinotus belus Harold, 1880: 25 Pinotus belus Luederwaldt 1929: 98 1936: 213 Blackwelder 1944: 80 Dichotomius belus Roze 1955: 44 Medina et al. 2001: 138 Taxonomic review of the Dichotomius (Luederwaldtinia) agenor species group (Coleoptera: Scarabaeidae: Scarabaeinae) Montoya-Molina, Santiago Vaz-De, Fernando Z. European Journal of Taxonomy 2021 2021-02-11 734 1 64 35NB8 (Harold, 1880) Harold 1880 [477,1110,1897,1924] Insecta Scarabaeidae Dichotomius Animalia Coleoptera 14 15 Arthropoda species belus Luederwaldtinia   Fig. 5, Map 3      Pinotus belus Harold, 1880: 25(original description).  Pinotus belus–  Luederwaldt 1929: 98;  1936: 213. —  Blackwelder 1944: 80.    Dichotomius belus–  Roze 1955: 44. —  Medina et al.2001: 138. — Sarmiento & Amat 2014: 93–95.    Diagnosis   Dichotomius belusis separated from other species in the group by the following combination of characters: setae on body surface sparse or absent. Clypeal teeth generally reduced in large males and in most cases completely absent in females. Male cephalic carina conical, antero-posteriorly flattened and curved anteriorly, higher than wide. Ventral clypeal process greatly emarginate.    Material examined    Lectotype(here designated) COLOMBIA• ♂, “ Medellín” [white, hw]/ “  BelusHarold.” [white, hw] /“Ex-Musaeo E. Harold” [white with black margins, p]/ “LECTOTYPE ♂  Pinotus belusHarolddes. F.Z. Vaz-de-Mello, 2014” [red with black margin, phw]; MNHN.   Paralectotypes(3 ♂♂, 3 ♀♀)   COLOMBIA• 1 ♀; “Columbia  Pinotus BelusHarold” [white, hw] / “Ex-Musaeo E. Harold” [white with black margins, p]/ “ PARALECTOTYPE♀  Pinotus belusHarolddes. F.Z. Vaz-de-Mello, 2014” [yellow with black margin, phw]; MNHN•  1 ♂; “ Medellín” [white, hw] / “Ex-Musaeo E. Steinheil” [white with black margins, p]/ “PARALECTOTYPE ♂  Pinotus belusHarolddes. F.Z. Vaz-de-Mello, 2014” [yellow with black margin, phw]; MNHN•  1 ♀; “ Medellín” [white, hw]/ “Ex-Musaeo E. Steinheil” [white with black margins, p]/ “ PARALECTOTYPE♀  Pinotus belusHarolddes. F.Z. Vaz-de-Mello, 2014” [yellow with black margin, phw]; MNHN•  1 ♀; “ Medellín” [white, hw] / “Ex-Musaeo E. Steinheil” [white with black margins, p]/ “ PARALECTOTYPE♀  Pinotus belusHarolddes. F.Z. Vaz-de-Mello, 2014” [yellow with black margin, phw]; MNHN•  1 ♂; “ Ocaña Landoldt” [white, hw]/ “Ex-Musaeo E. Steinheil” [white with black margins, p]/ “ PARALECTOTYPE♂  Pinotus belusHarolddes. F.Z. Vaz-de-Mello, 2014” [yellow with black margin, phw]; MNHN•  1 ♂; “ Ambalema” [white, hw]/ “Ex- Musaeo E. Steinheil” [white with black margins, p]/ “ PARALECTOTYPE♂  Pinotus belusHarolddes. F.Z. Vaz-de-Mello, 2014” [yellow with black margin, phw]; MNHN.  Additional material   COLOMBIA• 1 ♂;  1–4 Mar. 1972; dung and forest; S. and J. Peckleg.; CEMT•  1 ♀;  22 Apr. 1964; CEMT•  1 ♀;  Sep. 1957; Gallego; CEMT•  1 ♀;  21 Apr. 1964; CEMT. –  Antioquia• 1 ♂; Medellín; BMNH•  7 ♂♂, 5 ♀♀; Medellín; CEMT•  1 ♀; Medellín;  15 Dec. 1962; CEMT•  1 ♀; Medellín; Encorredo;  Apr. 1947; Gallegoleg.; CEMT•  6 ♂♂, 1 ♀; Puerto Berrio;  300 ma.s.l.; 6º29′31.20″ N, 74º34′39.84″ W;  Mar. 2015; S. Montoya-M leg.; CEMT•  3 specs; Puerto Berrio, Vda. La Carlota;  590 ma.s.l.; 6.518° N, -74.582° W;  Oct. 2012; J. Leónleg.; IAvH. –  Boyacá• 1 ♂; Villa de Leyva; Pastizal;  Dec. 1998; J. Noriegaleg.; CEMT•  1 spec.; Villa de Leyva, Rio Cane;  2200 ma.s.l.;  Jul. 1996; F. Escobarleg.; IAvH•  1 spec.; Villa de Leyva;  2200 ma.s.l.;  Sep. 1996; M. Álvaresleg.; IAvH] •  1 spec.; Villa de Leyva, Claustro San Agustín;  2200 ma.s.l.;  Aug. 1998; K. Schutzeleg.; IAvH. –  Cauca• 1 spec.; Bolivar, Buenos Aires, Área de reserva Lerma;  1700 ma.s.l.; 2º0′0″ N, 76º57′ W;  Jan. 2010; W. Guzmánleg.; ECC. –  Cundinamarca• 1 spec.; Beltrán, Insp. Paquiló, Fca. Santana;  650 ma.s.l.; 4.6678° N, 74.7596° W;  Nov. 2014; A. Loperaleg.; ECC•  1 spec.; Chinauta, Fusagasugá;  1900 ma.s.l.;  Jan. 1995; A. Amézquitaand A. Loperaleg.; ECC•  1 ♂; Fusagasugá;  6 Sep. 1962; CEMT•  1 ♀; Cachipay, alrededores colégio Fátima;  1500 ma.s.l.;  14 Apr. 1991; A. Meraand R. Lozanoleg.; CEMT•  1 spec.; Juan de Rio Seco;  1300 ma.s.l.;  Oct. 1997; H. Pulido-Riverosleg.; IAvH•  1 ♂; Bogotá; S9S2; BMNH. –  Huila• 12 specs; Garzón, Vda. El Espinal;  1000 ma.s.l.; 2º17′45″ N, 75º35′37″ W;  Sep. 2002; M. Ospinaleg.; IAvH•  1 ♀; La Guajira, Fonseca; 10º49.488′ N, 72º49.476′ W; S. Montoya-M.leg.; CEMT. –  Meta• 1 ♂; 33 Km E of Villavicencio;  2–4 Mar. 1972; S. and J. Peck; CEMT. –  Nariño• 1 ♀; Aguada; BMNH•  2 ♂♂, 1 ♀; Buesaco, RN El Charmolan, Vda. Hatotongosoy;  2500 ma.s.l.; 1º22′38″ N, 77º13′9″ W; pitfall, Potrero Ganadero;  22–26 Aug. 2008; A.N.L. Hidalgoleg.; CEMT•  1 spec.; R.N. La Planada.  1800 ma.s.l.;  Apr. 1994; F. Escobarleg.; IAvH•  7 specs; Taminango, Vda. Algodonal;  628 ma.s.l.; 1º40′6″ N, 77º17′15″ W;  Oct. 2013; J. Salazarand J. Castilloleg.; IAvH. –  Quindío• 1 spec.; Córdoba, La Tribuna; 4º25′40″ N, 75º40′22″ W;  Jul. 1999; J. Molinaand J. Elejaldeleg.; ECC. –  Risaralda• 1 spec.; La Celia, Vda. La Secreta;  1720 ma.s.l.; 5º1′1.3″ N, 76º1′13.6″ W;  Jul. 2008; CultidACM leg.; IAVH. –  Santander• 2 ♂♂, 2 ♀♀; Encino, Sector La Gorda, Fca. San Agustín;  2000 ma.s.l.; 6º7′12″ N, 73º1′59″ W; Pastizal; trampa de caída con excrement humano;  29 Nov.–1 Dec. 2009; J. Ariasleg.; CEMT•  1 ♀; Santandercito;  May 1960; CEMT•  1 spec.; El Encino, Vda. Las Minas, Fca. Los Nogales;  2280 ma.s.l.; 6º6′28″ N, 73º3′7.6″ W;  Dec. 2014; A. Loperaand R. Sarmiento; ECC•  1 spec.; Virolin, SFF Guanenta Alto rio Fonce;  2450 ma.s.l.;  2 Feb. 1998; J. Martínezleg.; IAvH. –  Tolima• 1 ♂; Ambalema; Nevinsonleg.; BMNH 1918-14•  7 specs; Armero, Guayabal, Loma Santo Tomás;  300 ma.s.l.; 4º55′70″ N, 49º52′ W;  Nov. 1995; F. Escobarleg.; IAvH • 6 specs; Armero, Guayabal Mendez Had. Bremen;  300 ma.s.l.; 5º5′31″ N, 74º45′81″ W; excr. humano;  Nov. 1995; F. Escobarleg.; IAvH•  1 spec.; Mariquita, Bosque Municipal;  650 ma.s.l.;  28 Sep. 2003; V. Fuentesleg.; ECC•  1 spec.; R.N. La Victoria;  700 ma.s.l.; 4º14′45.9″ N, 74º35′37.9″ W;  Apr. 2015; A. Loperaleg.; ECC. –  Valle del Cauca• 1 spec.; Buga, La Magdalena;  1600 ma.s.l.;  Mar. 1991spec.; C. Medinaleg.; ECC•  1 spec.; Cali;  1000 ma.s.l.;  Oct. 1990; Jorge Aldanaleg.; ECC•  1 spec.; Calima;  900 ma.s.l.;  Nov. 1993; R. Bravoleg.; ECC•  1 ♀;  19 mi.S of Cartago;  960 ma.s.l.;  18 Mar. 55; E.I. Schlingerand E.S. Rossleg.; CEMT•  1 spec.; Dagua, Chicoral;  1900 ma.s.l.;  Jun. 1990; C. Medinaand F. Escobarleg.; ECC•  1 ♂; Pance, Farallones; Luz.Mar;  Apr. 1988; L.C. Reyesleg.; CEMT•  2 ♂♂; Pance, Farallones; Luz;  25 Dec.–2 Jan. 1991; L.C. Pardoleg.; CEMT•  1 ♂, 1 ♀; Buenvª [possibly Buenaventura]; Rosenberg. Fryleg.; BMNH 1905-100•  1 ♂, 2 ♀♀; Cauca Valley; Colombia; BMNH.    Redescription  Male MEASUREMENTS. Length: 17.9 mm. Setae on body surface sparse or absent. HEAD. Anterior third surface rugose, not extending posteriorly to base of cephalic carina. Clypeus with central weak emargination separating two short rounded clypeal teeth ( Fig. 5A, D). Cephalic carina higher than wide, antero-posteriorly flattened, looking like triangular-shaped plate in frontal view. Anterior surface of cephalic carina smooth, with microscopic punctures. Median tubercle twice the high of lateral tubercles. Lateral tubercles acuminate apically ( Fig. 5D, G). Ventral clypeal process greatly emarginate ( Fig. 2C) PRONOTUM. Gibbous, imperceptibly emarginate at antero-medial portion. Wider than long, as wide as elytra; pronotal disc shiny, with fine indistinct punctures. Medial sulcus weakly impressed. Ocellate punctures along posterior and anterior margins ( Fig. 5A, D, G), absent in lateral margins. Hypomeron with broad ocellate setigerous punctures separated by less than half the width of a puncture. Medial disc smooth, without punctures. External margin bearing multiple rows of disorganized long setae when seen in dorsal view (see Fig. 2E). ABDOMEN. Mesosternum covered by large ocellate setigerous punctures, setae as long as in mesosternum. Metepisternum smooth with some weakly marked punctures. Metasternum setigerous punctures restricted mainly to lateral areas. Antero-median portion with some feebly marked punctures next to apical margins of coxa. Lateral surface of metasternum with ocellate punctures separated by two or three times width of a puncture, some punctures with long setae. Metepisternum suture clearly visible. Ocellate punctures along anterior margin of each sternite, denser in lateral areas, without reaching the centre. Pygidium surface covered by fine punctures, basal margin with wide punctures that merge at the centre.   Fig. 5.  Dichotomius belus( Harold, 1880). A–B. Dorsal habitus. A. ♂. B. ♀. C. Lateral view of the aedeagus. D. Anterior view of the shape of the cephalic carina. E. Ventral view of the aedeagus. F–G. Lamella copulatrix. H.Lateral view, showing the pronotum and the cephalic carina flattened antero-posteriorly and curved anteriorly. ELYTRA. Interstriae smooth (see Fig. 2I) and shiny with fine randomly separated punctures. Striae well delimited, punctures separated by once or twice diameter of a puncture on elytral disc. LEGS. Metacoxa glabrous. Fore calcar bifurcate. PARAMERES AND LAMELLA COPULATRIX. As in Fig. 5C, E–F.   Morphological variation  Females( Fig. 5B) As males except for the following characters: HEAD. Rounded, clypeal margin not emarginate or faintly emarginate. Surface almost completely rugose, denser on anterior portion. Fronto-clypeal carina quadridentate. Interocular space shiny and smooth. PRONOTUM. Smooth, shiny. Hypomeron setae longer and denser than in males.    Remarks   Dichotomius belusis endemic to Colombiaand is distributed throughout the dry zones of the Andes mountains and lowlands associated with the interior of the Tropical Dry Forests. Based on the original description, specimens used to describe the species were collected in Medellín, Ambalema and Ocaña. 3037418511 MNHN F. Z. Vaz-de-Mello Colombia Medellin Medellin 15 16 1 1 lectotype 3037418378 MNHN F. Z. Vaz-de-Mello Colombia COLOMBIA 15 16 1 1 paralectotype 3037418345 MNHN F. Z. Vaz-de-Mello Medellin Medellin 15 16 1 1 3037418591 MNHN F. Z. Vaz-de-Mello Medellin Medellin 15 16 1 1 paralectotype 3037418497 MNHN F. Z. Vaz-de-Mello Medellin Medellin 15 16 1 1 paralectotype 3037418602 MNHN F. Z. Vaz-de-Mello Ocana Landoldt Ocana Landoldt 15 16 1 1 paralectotype 3037418563 MNHN F. Z. Vaz-de-Mello Ambalema Ambalema 15 16 1 1 paralectotype 3037418538 [189,1145,1360,1389] dung and forest 1972-03-01 1972-03-04 1972-03-01 CEMT S. and J. Peck Colombia COLOMBIA 15 16 1 1 3037418567 1964-04-22 CEMT Colombia 15 16 1 1 3037418321 [298,687,1398,1425] 1957-09 CEMT Gallego Colombia 15 16 1 1 3037418380 [715,1037,1398,1424] 1964-04-21 CEMT Colombia Antioquia 15 16 1 1 Antioquia 3037418342 1964-04-21 CEMT Medellin Colombia Antioquia 15 16 1 1 1 Antioquia 3037418617 [310,694,1431,1460] CEMT Medellin Colombia 15 16 12 5 7 3037418467 [726,1184,1433,1460] 1962-12-15 CEMT Colombia Medellin 15 16 1 1 3037418351 1947-04 CEMT Gallego Colombia Encorredo Medellin 15 16 1 1 3037418441 2015-03 CEMT Colombia 300 6.4919996 Puerto Berrio 1 -74.577736 15 16 7 1 6 3037418426 2012-10 IAvH J. Leon Colombia 590 6.518 Vda. La Carlota 78 -74.582 Puerto Berrio 15 16 3 Boyaca 3037418566 2012-10 IAvH J. Leon Colombia 590 6.518 Vda. La Carlota 78 -74.582 Puerto Berrio 15 16 3 1 Boyaca 3037418377 1996-07 IAvH F. Escobar Colombia 2200 Rio Cane Villa de Leyva 15 16 1 3037418394 [482,1371,1609,1638] 1996-09 IAvH M. Alvares Colombia 2200 Villa de Leyva 15 16 1 3037418492 [189,1373,1645,1672] 1998-08 IAvH K. Schutze Colombia 2200 Claustro San Agustin Villa de Leyva 15 16 1 Cauca 3037418526 1998-08 IAvH K. Schutze Colombia Bolivar 2200 2.0 Claustro San Agustin 924 -76.95 Villa de Leyva 15 16 1 Cauca 3037418520 2010-01 ECC W. Guzman Colombia Bolivar 1700 2.0 Area de reserva Lerma 924 -76.95 Buenos Aires 15 16 1 Cundinamarca 3037418423 1995-01 ECC A. Amezquita & A. Lopera Colombia 1900 Fusagasuga Chinauta 15 16 1 3037418415 1962-09-06 CEMT Colombia Fusagasuga 15 16 1 1 3037418468 1991-04-14 CEMT A. Mera & R. Lozano Colombia 1500 alrededores colegio Fatima Cachipay 15 16 1 1 3037418516 [444,1382,1857,1884] 1997-10 IAvH H. Pulido-Riveros Colombia 1300 Juan de Rio Seco 15 16 1 3037418483 [188,529,1890,1919] BMNH Colombia Bogota 15 16 1 1 Bogota 3037418392 2002-09 BMNH M. Ospina Colombia 1000 2.2958333 Bogota 21 -75.59361 15 16 1 1 Bogota 3037418524 CEMT S. Montoya-M. Colombia 10.8248 La Guajira 1 -72.8246 15 16 1 1 La Guajira 3037418438 1972-03-02 1972-03-04 1972-03-02 CEMT S. Montoya-M. Colombia 10.8248 La Guajira 1 -72.8246 15 16 1 1 1 La Guajira 3037418408 [282,665,1998,2025] 1972-03-02 1972-03-04 1972-03-02 CEMT S. Montoya-M. Colombia 10.8248 La Guajira 1 -72.8246 15 16 1 1 1 La Guajira 3037418445 2008-08-22 2008-08-26 2008-08-22 pitfall CEMT A. N. L. Hidalgo Colombia Buesaco 2500 16 17 1.3772222 Potrero Ganadero 21 -77.21917 Vda. Hatotongosoy 15 16 3 1 2 3037418571 [293,1138,301,328] 1994-04 IAvH F. Escobar Colombia 1800 La Planada. 16 17 1 3037418532 2013-10 IAvH J. Salazar & J. Castillo Colombia 628 1.6683334 Vda. Algodonal 21 -77.2875 Taminango 16 17 7 Quindio 3037418431 2013-10 IAvH J. Salazar & J. Castillo Colombia 628 1.6683334 Vda. Algodonal 21 -77.2875 Taminango 16 17 7 Quindio 3037418379 2013-10 IAvH J. Salazar & J. Castillo Colombia 628 1.6683334 Vda. Algodonal 21 -77.2875 Taminango 16 17 7 Quindio 3037418443 2008-07 2009-12-01 2009-11-29 trampa de caida con excrement humano IAVH Cultid Colombia Encino 1720 5.017028 La Celia 1 -76.02045 Sector La Gorda 16 17 1 2 2 Santander 3037418301 [810,1256,520,546] 1960-05 CEMT Colombia Santandercito 16 17 1 1 3037418547 2014-12 ECC A. Lopera & R. Sarmiento Colombia El Encino 2280 6.1077776 Fca. Los Nogales 15 -73.05212 Vda. Las Minas 16 17 1 3037418365 1998-02-02 IAvH J. Martinez Colombia Virolin 2450 SFF Guanenta Alto rio Fonce Guanenta Alto 16 17 1 Tolima 3037418372 [445,1169,627,655] 1998-02-02 IAvH J. Martinez Colombia Virolin 2450 SFF Guanenta Alto rio Fonce Guanenta Alto 16 17 BMNH 1918-14 1 1 Tolima 3037418607 1995-11 excr. humano IAvH F. Escobar Colombia Armero 300 5.0919447 Loma Santo Tomas 921 -49.866665 Guayabal 16 17 13 3037418487 1995-11 excr. humano IAvH F. Escobar Colombia Armero 300 5.0919447 Loma Santo Tomas 921 -49.866665 Guayabal 16 17 13 3037418496 2003-09-28 ECC V. Fuentes Colombia 650 Bosque Municipal Mariquita 16 17 1 3037418508 2015-04 ECC A. Lopera Colombia 700 4.2460833 Valle del Cauca 1 -74.593864 16 17 1 Valle del Cauca 3037418505 2015-04 ECC A. Lopera Colombia 700 4.2460833 Valle del Cauca 1 -74.593864 Buga 16 17 1 Valle del Cauca 3037418463 [431,1179,847,874] 1990-10 ECC Jorge Aldana Colombia 1000 Cali 16 17 1 3037418303 1993-11 ECC R. Bravo Colombia 900 Calima 16 17 1 3037418551 1955-03-18 CEMT E. I. Schlinger & E. S. Ross Colombia 960 19 mi. S of Cartago 16 17 1 1 3037418555 1990-06 ECC C. Medina & F. Escobar Colombia 1900 Chicoral Dagua 16 17 1 3037418314 [504,1302,954,983] 1988-04 CEMT L. C. Reyes Colombia Pance Luz. Mar Farallones 16 17 1 1 3037418546 1991-01-02 1991-12-25 1991-01-02 CEMT L. C. Pardo Colombia Pance Luz Farallones 16 17 2 2 3037418548 BMNH Rosenberg. Fry Colombia Buenaventura 16 17 BMNH 1905-100 2 1 1 3037418464 [856,1390,1027,1056] BMNH Colombia Colombia Cauca Valley 16 17 3 2 1 Cauca