Dichotomius (Luederwaldtinia) agenor ( Harold, 1869 )

Montoya-Molina, Santiago & Vaz-De, Fernando Z., 2021, Taxonomic review of the Dichotomius (Luederwaldtinia) agenor species group (Coleoptera: Scarabaeidae: Scarabaeinae), European Journal of Taxonomy 734, pp. 1-64 : 8-12

Dichotomius (Luederwaldtinia) agenor ( Harold, 1869)

Fig. 3 , Map 1

Pinotus agenor Harold, 1869: 141 (original description).

Pinotus agenor – Luederwaldt 1929: 99. — Blackwelder 1944: 69.

Dichotomius agenor – Howden & Young 1981: 13. — Medina et al. 2001: 138. — Ratcliffe 2002: 15 — Sarmiento & Amat 2014: 89–90.

Diagnosis

Dichotomius agenor is separated from the other species in the group by the following combination of characters: large male fronto-clypeal carina with lower part wider than high, with median tubercle bifurcate apically, concave between central and lateral tubercles. Pronotum with pronounced, truncate anterior declivity, when viewed laterally. External margin of hypomeron bearing multiple rows of disorganized long setae, all approximately the same length, that can be seen in dorsal view. Posterior edge of metacoxa bearing two rows of disorganized ocellate punctures, separated by half the diameter of a puncture.

Material examined

Lectotype (here designated) COLOMBIA • ♂; “Columbia. P Agenor” [white with red margins, hw] / “ Ex-Musaeo E.Harold ” [white with black margins, p]/ “LECTOTYPE ♂ Pinotus agenor Harold des. F.Z. Vaz-de-Mello, 2014” [red with black margin, phw]; MNHN.

Paralectotypes (1 ♂, 1 ♀)

COLOMBIA • 1 ♂; “ Agenor Colomb.” [white, hw]/ “Ex-Musaeo E. Harold” [white with black margins, p]/ “ ♂ Pinotus agenor Harold des. F.Z. Vaz-de-Mello, 2014” [yellow with black margin, phw]; MNHN • 1 ♀; “ Agenor Colomb.” [white, hw]/ “Ex-Musaeo E. Harold” [white with black margins, p]/ “ ♀ Pinotus agenor Harold des. F.Z. Vaz-de-Mello, 2014” [yellow with black margin, phw]; MNHN.

Additional material

COLOMBIA – Antioquia • 1 spec.; Sonsón, San Miguel, Fca. La Rica ; 160–250 m a.s.l.; 21 Aug. 2006; J. Arias leg.; IAvH • 1 ♂; Antioquia; BMNH 5438 . – Atlántico • 1 spec.; Juan de Acosta, tierra arena ; 250 m a.s.l.; 10º16′28″ N, 75º57′1.3″; 21–22 Sep. 2009; J. Rangel and O. Blanco leg.; IAvH • 3 specs; Piojó, Fca. La Cristalina ; 50 m a.s.l.; 10º44′59″ N, 75º9′42″ W; bosque seco tropical; Aug. 2015; A. Lopera and N. Castillo leg.; IAvH • 1 spec.; Piojó , Fca. La Cristalina ; 50 m a.s.l.; 10º44′59″ N, 75º9′42″ W; bosque seco tropical; Aug. 2015; A. Lopera and N. Castillo leg.; ECC. – Bolivar • 1 spec.; Isla de Tierra Bomba ; 100 m a.s.l.; 10º21′36″ N, 75º34′30″ W; Aug. 1996; F. Escobar leg.; ECC • 9 specs.; San Juan de Nepomuceno , Fca. Mis Deseos ; 482 m a.s.l.; 9º55′8.4″ N, 75º9′33.7″ W; bosque seco tropical; Aug. 2015; A. Lopera and N. Castillo leg.; IAvH • 1 spec.; San Juan de Nepomuceno , Fca. Mis Deseos ; 482 m a.s.l.; 9º55′8.4″ N, 75º9′33.7″ W; bosque seco tropical; Aug. 2015; A. Lopera and N. Castillo leg.; ECC • 1 spec.; Zambrano , Had. Moterrey ; 120 m a.s.l.; 9º37′48″ N, 74º54′44″ W; Aug. 1996; F. Escobar leg.; ECC. – Caldas • 1 ♂, 2 ♀♀; Guarinocitos; 200 m a.s.l.; 5º22′00.6″ N, 74º47′34.6″ W; Aug. 2015; S. Montoya-M. leg.; CEMT • 1 ♂; Norcasia , Vda. San Roque-Reserva Rio Natural Manso ; 220 m a.s.l.; 5º39′40″ N, 74º46′98″ W; bosque; 1 Aug. 2004; L. Arango and A. Montes leg.; ECC . – Caquetá • 1 ♂; Yari ; 16 Oct. 1986; CEMT . – Cesar • 110 specs; Codazzi ; 165 m a.s.l.; 9º58′42″ N, 73º56′44″ W; 17 May 2016; C. Giraldo and A. León, J leg.; MUSENUV • 1 spec.; Valledupar ; 160 m a.s.l.; 10º21.747′ N, 73º23.26′ W; Potrero; Aug. 2013; S. Montoya leg.; ECC • 1 spec.; Velledupar, Ecoparque Los Besotes ; 500 m a.s.l.; 10º34′30″ N, 73º18′38″ W; Dec. 2002; F. Forero leg.; IAvH . – Cordoba • 1 spec.; Monteria, Las Palmeras ; 8º30′37.1″ N, 76º6′12.9″ W; bosque; Sep. 2011 spec.; A. Vargas-Pérez leg.; ECC • 1 spec.; Monteria, Vda. Sierra Chiquita ; 125 m a.s.l.; 8º44′22″ N, 75º54′40″ W; bosque seco tropical; Dec. 2011 spec.; R. Leidys Murillo leg.; ECC • 8 specs; Monteria, Vda. Sierra Chiquita ; 125 m a.s.l.; 8º44′22″ N, 75º54′40″ W; bosque seco tropical; Dec. 2011 spec.; R. Leidys Murillo leg.; IAvH • 1 spec.; Tierra Alta, Vereda Tuis-Tuis, Reserva Tuis-Tuis ; 125 m a.s.l.; 8º2′23.7″ N, 76º5′59.6″ W; Apr. 2011 spec.; R. Leidys Murillo leg.; ECC . – Cundinamarca • 2 ♀♀; San Juan de Rio Seco ; bosque seco tropical; 1300 m a.s.l.; 12 Oct. 1997; Pulido-Riveros leg.; ECC • 2 specs; Bogotá; Nevinson leg.; BMNH 1918-14 . – Huila • 7 specs; Desierto de la Tatacoa; 480 m a.s.l.; 3º13′50″ N, 75º9′25″ W; Nov. 2012; D. Martínez leg.; IAvH . – La Guajira • 252 specs; Jagua del Pilar ; 181 m a.s.l.; 10º30.88′ N, 73º5.31′ W; Aug. 2013; S. Montoya leg.; IAvH • 282 specs; Uribia, PNN La Macuira- Corregimiento de Nazareth Kajashiwou ; 70 m a.s.l.; 12º11′37″ N, 71º21′30″ W; bosque seco tropical; 19 May 2014; C. Medina and A. González leg.; IAvH . – Magdalena • 1 spec.; Santa Marta, PNN Sierra Nevada , C. Minca , Cuenca del rio Gaira , Sector Pozo Azúl ; 740 m a.s.l.; 11º8′17″ N, 74º8′29″ W; 1–2 Aug. 2004; H. García and D. Ospino leg.; ECC • 1 spec.; Santa Marta, PNN Sierra Nevada ; C. Gaira leg.; 50 m a.s.l.; 11º10′26″ N, 74º10′37″ W; Jul. 2004; H. García leg.; ECC • 2 ♂♂, 2 ♀♀; PNN Tayrona Neguanje ; 155 m a.s.l.; 11º17′41″ N, 74º06′15″ W; bosque seco excr. humano; Sep. 1996; F. Escobar leg.; CEMT • 2 ♂♂; Stn Martha Colombia ; Nevinson leg.; BMNH 1918-14 . – Meta • 2 ♀♀; Santander, Encino, Sector La Gorda, Finca San Agustín ; Pastizal ; 2000 m a.s.l.; trampa de excremento humano; Dec. 2009; J. Arias-Buriticá leg.; CEMT. – Sucre • 94 specs; San Benito Abad, Vda. Santiago Apostol ; 30 m a.s.l.; 9º2′19″ N, 75º54′25″ W; Palmar leg.; Oct. 2009; L.E. Franco leg.; IAvH • 1 spec.; San Benito Abad, Vda. Santiago Apostol ; 30 m a.s.l.; 9º2′19″ N, 75º54′25″ W; Palmar leg.; Oct. 2009; L.E. Franco; ECC • 3 specs; Colosó, Montes de María, Sector La Cascada , Estación Primatológica ; 28–30 Apr. 2009; L.E. Franco leg.; IAvH . – Tolima • 20 specs; Alvarado, Fca. La Brasilia ; 537 m a.s.l.; 5º11′34″ N, 75º39′33″ W; 7 Mar. 2016; C. Giraldo and A. León J leg.; MUSENUV • 1 ♂; Armero ; bosque; 260 m a.s.l.; Nov. 2000; J. Noriega leg.; CEMT • 1 ♂; Guayabal. Armero. Oct. 2000; J. Noriega leg.; CEMT • 1 ♀; Armero, Cerros de Santo Tomás ; 250 m a.s.l.; Nov. 2000; J. Noriega leg.; CEMT • 1 spec.; Armero, Guayabal, Cerros de Santo Tomás ; 300 m a.s.l.; 5º5′38″ N, 75º46′4″ W; bosque de cerro; Nov. 1995; F. Escobar leg.; ECC • 183 specs; Armero, Guayabal, Cerros de Santo Tomás ; 300 m a.s.l.; 5º5′38″ N, 75º46′4″ W; bosque de cerro; Nov. 1995; F. Escobar leg.; IAvH • 1 ♂; Armero, Guayabal, Mendez Had. Cardonal ; 300 m a.s.l.; 5º5′38″ N, 74º46′39″ W; excr. humano; Borde Bo-Po [border between forest-pasture]; Sep. 1995; F. Escobar leg.; CEMT • 45 specs; Armero, Guayabal, Mendez Had. Cardonal ; 300 m a.s.l.; 5º5′38″ N, 74º46′39″ W; excr. humano; Borde Bo-Po; Sep. 1995; F. Escobar leg.; IAvH • 1 ♀; Armero, Guayabal Mendez Had. Bremen ; 300 m a.s.l.; 5º5′31″ N, 74º45′81″ W; excr. humano; Nov. 1995; F. Escobar leg.; CEMT • 116 specs; Armero, Guayabal Mendez Had. Bremen; 300 m a.s.l.; 5º5′31″ N, 74º45′81″ W; excr. humano; Nov. 1995; F. Escobar leg.; IAvH . – Valle del Cauca • 1 spec.; Buga, Fca. Los Chagualos ; 29 Feb. 2016; C. Giraldo and A. León, J; MUSENUV • 1 ♀; Colombia, Nevinson leg.; BMNH 1918-14 .

PANAMA • 3 ♂♂, 2 ♀♀; Panamá City, Cerro Ancon ; 8º57′ N, 79º33′ W; light; Apr. 2014; B. Becker leg.; CEMT • 1 ♂; La Chorrera , Panamá; BMNH 98-146 .

VENEZUELA – Mérida • 4 ♂♂, 4 ♀♀; Libertador, Meseta de Zumba, Had. Los Arcos ; 1181 m a.s.l.; 8º33′14″ N, 71º13′20″ W; pitfall human faeces; 6 May 2016; R. Acconcia and J. Gámez leg.; CEMT • 2 ♂♂; Merida ( Seilbahn ); 220 m a.s.l.; 11 Apr. 1995; Hornburg and Krause leg.; CEMT . – Táchira • 3 ♂♂, 3 ♀♀; Libertador, San Joaquín de Navay ; 200 m a.s.l.; 7.6622º N, 71.7104º W; Aug. 2006; T. Good leg.; CEMT . – Yaracuy • 1 ♀; Bolivar Aroa ; 468 m a.s.l.; 10º0′0″ N, 68º0′0″ W; heces humanas; 21 Jul. 2009; M. Asmüssen, P. Colmenares and H. Martínez leg.; CEMT • 2 ♂♂, 4 ♀♀; Venez [Venezuela]; Nevinson leg. BMNH 1918-14 • 2 ♂♂; Veneza. [Venezuela]; Fry leg.; BMNH 1905-100 • 1 ♂; Nevinson leg.; BMNH 1918-14 • 1 ♂; BMNH 3521 .

Redescription

Male

MEASUREMENTS. Length: 20.5 mm.

HEAD. Anterior surface rugose, not extending posteriorly to base of cephalic carina. Cephalic carina wider than high, anterior surface smooth, dorsally concave between central and lateral tubercles ( Fig. 3C );

median tubercle twice as high as lateral tubercles, conical in shape and apically emarginate. Interocular space shiny, smooth. Ventral clypeal process cordate (see Fig. 2A ).

PRONOTUM. Wider than long and wider than elytra. Pronotal disc shiny and smooth, lacking deep punctures, with at most fine punctures. Anterior declivity pronounced and truncate when viewed laterally ( Fig. 3D ). Ocellate punctures present along anterior and posterior margins, few scattered ocellate punctures in lateral margins ( Fig. 3A ). Hypomeron medial disc punctures separated by three times the width of a puncture. External margin bearing multiple rows of disorganized long setae (see Fig. 2E ).

ABDOMEN. Mesosternum and metepisternum fully covered by ocellate setigerous punctures, pilosity dense, setae all of same length. Metasternum setigerous punctures restricted mainly to lateral and anterior areas. Anterior lobe pilosity never connecting with lateral pilosity through margin of mesocoxae (see Fig. 2H ). Lateral surface with ocellate setigerous punctures separated by twice the width of a puncture, variation in puncture diameter almost imperceptible. Ocellate punctation along anterolateral margins of sternites. Pygidium surface covered by small, fine punctures. Basal margin bearing few ocellate punctures.

ELYTRA. Interstriae shagreened, with fine, randomly separated punctures with interior shagreened microsculpture. Striae hardly marked, with ocellate punctures separated by twice the diameter of a puncture on elytral disc (see Fig. 2J ).

LEGS. Metacoxa posterior edge bearing two contiguous rows of organized ocellate punctures, separated by half the diameter of a puncture ( Fig. 3E ).

PARAMERES AND LAMELLA COPULATRIX OF AEDEAGUS. As in Fig. 3 F–H.

Morphological variation

Females ( Fig. 3B )

As males except for the following characters:

HEAD. Completely transversely rugose, denser on anterior portion. Cephalic carina bituberculate (see Fig. 1E ).

PRONOTUM. Weak medial knob, only detectable in lateral view. Hypomeron setae longer and denser than in males.

Remarks

Luederwaldt (1929) placed this species in the assifer section which included species from the assifer and agenor groups. Harold’s type material was from Colombia, but the label does not specify the exact locality. This species is known to occur in Panama, Colombia and Venezuela from 100 to 1500 m a.s.l., in the Caribbean subregion as proposed by Morrone (2001). It is found in the Magdalena and Maracaibo Provinces, associated with dry and humid forests ( Morrone 2001). Individuals of this species are easily confused with D. enioi sp. nov., a Central American species.

Figures

Fig. 3. Dichotomius agenor (Harold, 1869). A–B. Dorsal habitus. A. ♂. B. ♀. C. Posterior view of the cephalic carina, concavity between the central and the lateral tubercles, apically emarginate. D. Lateral view of the pronotum, anterior declivity truncate. E. Ventral view of the metacoxa, displaying two rows of punctures on the posterior margin. F. Lateral view of the aedeagus. G. Ventral view of the aedeagus. H–I. Lamella copulatrix.

Fig. 2. General characters that help differentiate the species of the Dichotomius (Luederwaldtinia) agenor species group. A–D. Ventral clypeal process. A. Cordate as in D. agenor (Harold, 1869). B. Truncate as in D. sagittarius (Harold, 1869) stat. rev. C. Bifurcate as in D. belus (Harold, 1880). D. Cylindrical as in D. henripittieri sp. nov. E–F. Hypomeral characters. E. Medial disc smooth, lateral margin with multiple rows of setae as in D. amplicollis (Harold, 1869). F. Medial disc punctate, lateral margin with single row of setae as in D. tristis (Luederwaldt, 1923). G–H. Metasternum characters. G. Setigerous punctures from the medial and lateral edges joining throughout the margin of the mesocaxa as in D. enioi sp. nov. H. Setigerous punctures from the medial and lateral edges not joining throughout the margin of the mesocaxa as in D. centralis (Harold, 1869). I–J. Elytra interstriae. I. Smooth as in D. amplicollis. J. Shagreened as in D. agenor.

Fig. 1. Diagnostic characters of the Dichotomius (Luederwaldtinia) agenor species group. A–D. Male diverse trituberculate fronto-clypeal carina shapes. A. D. tristis (Luederwaldt, 1923). B. D. amplicollis (Harold, 1869). C. D. belus (Harold, 1880). D. D. validipilosus (Luederwaldt, 1931). E–F. Female fronto-clypeal carina. E. Bituberculate carina typical of D. tristis. F. Quadrituberculate carina typical of D. amplicollis.