Onthophagus confusus Boucomont, 1932

Rossini, Michele, Vaz-de-Mello, Fernando Z. & Zunino, Mario, 2018, A taxonomic revision of the New World Onthophagus Latreille, 1802 (Coleoptera: Scarabaeidae: Scarabaeinae) of the osculatii species-complex, with description of two new species from South America, Journal of Natural History 52 (9 - 10), pp. 541-586 : 573-577

Onthophagus confusus Boucomont, 1932 stat. nov.

( Figures 4 (a – d), 4(j – n), 5(a))

Onthophagus ophion var. confusus Boucomont 1932: 306

Onthophagus nabeleki Balthasar 1939: 43 syn. nov. ( Figure 4 (e), 4(f))

Onthophagus nabeleki Balthasar : Martínez 1947: 112; Zunino & Halffter 1997: 161; Escobar 2000: 209; Medina et al. 2001: 140; Pulido-Herrera et al. 2007: 307 Diagnosis

Onthophagus confusus is undoubtedly related to O. insularis , from which it can be distinguished by the following combination of characters: foretibiae of male with a distinct internoapical tooth, posterolateral margins of pronotum slightly flattened, anteromedial pronotal protuberance conical and slightly extended forward. The male ’ s frontal horns are flattened (the transversal section of the horns is not circular as in many other American Onthophagus ), strongly curved and encircling the pronotal protuberance.

Redescription

Medium-sized species, body length 6 – 8 mm. Colour. Head and pronotum usually metallic green to bronze and shining, sometimes reddish with strong cupreous and metallic-green sheen, elytra opaque, completely black or testaceous, often with paler spots at the base and apex, pygidium metallic green, testaceous or bicoloured (disc dark, sides clearly paler). Ventral side of the body light to dark brown, femurs and metasternal surface, along the inner margins of mesocoxae yellowish, tibiae black to brownish, tarsi, mouthparts, scape and antennal articles I – V brownish to dark brown, antennal club yellow to testaceous. Head. Clypeus of male sub-trapezoidal, with anterior margin truncated to feebly curved and slightly reflexed, genae always clearly expanded beyond clypeus. Fronto-clypeal region without carina, frons with two horns strongly curved and embracing the pronotal protuberance, apexes of horns almost in contact, head surface very finely and evenly punctated. Thorax. Anteromedial pronotal protuberance of male low (close to the pronotal anterior margin) and horizontally protruded between cephalic horns, apex curved to obtusely conical, posterolateral margin of pronotum slightly flattened (in dorsal view) near posterior angles, pronotal punctation simple and evenly distributed across the surface. Elytra opaque, lateral interstriae (VI – VII) shinier, surface very finely microreticulated, striae shallow, interstriae flat and faintly punctated, punctures sparse, interval VII with few short and straight setae associated with the basal punctures, epipleura with setae longer. Abdomen. Pygidium flat to very weakly convex, completely bordered, surface feebly shining to opaque and completely microreticulated, punctures very shallow and evenly distributed. Legs. Foretibiae distinctly elongated and slender, evenly curved inward, with four external teeth separated by very small denticles, internoapical margin of protibiae with an acuminated and distinct tooth, apical spur of foretibiae with apex obtuse and curved downward. Middle and hind legs unmodified. Male genitalia. In dorsal view, distal-superior area of the paramerites obtusely acuminated at the apex, lateral margin straight and abruptly angled on the posterior side. In lateral view, distal-inferior area straight to barely excavated near the apex of the paramerites, tip of the paramerites very feebly curved downward ( Figure 4 (j), 4(k)). Superior margin of the lamella copulatrix excavated at middle but often hardly visible, superior left lobe narrow and rather straight, apical margin curved, external margin deeply excavated. Inferior right lobe slightly longer than inferior left lobe. Secondary lamella square-shaped and wide, margins often indistinct. Female genitalia. Ventral sclerotization of the vagina distinct, central and inferior margin deeply and abruptly excavated ( Figure 4 (n)).

Females differ from males in having the fronto-clypeal region very finely carinate, carina evenly curved forward and interrupted at the clypeo-genal suture, frons with a straight and transverse, ridge-like carina, which appears sub-trapezoidal from above, carina always depressed at middle. Clypeus finely wrinkled, fronto-clypeal region, vertex and genae with simple but stronger punctures than male. Anteromedial pronotal protuberance obsolete, feebly swollen to transversally truncated, foretibiae larger than male and unmodified. Minor males have cephalic horns shorter and straight, anteromedial pronotal protuberance weakly curved and slightly swollen.

Distribution and ecology

Known from the western side of the Ecuadorian Andes and northern Peru (new record) ( Figure 5 (a)), from the Pacific coast to higher elevations (ca. 2000 m). According to the collection data reported in the labels of the specimens here examined, O. confusus is normally collected with pitfall traps baited with human excrement in primary and secondary rain forests.

Remarks

Boucomont (1932) proposed confusus as a new variety of O. ophion Erichson , which made the first name available as the subspecific category (nec infrasubspecific; ICZN 1999, Art. 45.6.4). However, on examining the external and genital morphology of the type specimens of O. ophion (ZMHB) and O. ophion confusus , we could confirm they belong to distinct species. The shape of the horns in male adult specimens of confusus are strongly curved and encircling the anteromedial pronotal protuberance, the posterolateral pronotal angles are slightly flattened, while the anterior angles are obtusely acute, with lateral margin straight to slightly curved (often slightly excavated in O. ophion ). Furthermore, strong differences were identified in the male genital lamellae, and especially in the shape of the superior left lobe of the lamella copulatrix. Therefore, maintaining the subspecific name originally proposed by Boucomont, we elevate confusus at species level under the following new status: O. confusus Boucomont, 1932 .

As mentioned by Boucomont (1932, p. 328), within the type series of O. confusus , which is today incorporated in the René Oberthür collection at the MNHN, we found two males labelled ‘ Bolivie, Suapi ’. Both specimens were here identified as O. ophion Erichson (compared with the syntype of this species, which is deposited at the ZMHB) and for this reason excluded from the geographic distribution of O. confusus .

On discussing the relationships of O. transisthmius, Howden and Young (1981) mentioned O. ophion confusus and O. nabeleki as close relatives, alluding to the possible synonymy between these two names. Throughout our research, we examined the type specimens of all the American Onthophagus so far assigned to the hircus group, and among them, O. nabeleki Balthasar, 1939 , described from Ecuador. The detailed analysis of the morphology of its type specimens led us to confirm this species name as junior synonym under O. confusus (ICZN 1999, Art. 23). So, to maintain the nomenclatural stability for O. confusus and O. nabeleki , we designated two well-preserved male specimens as lectotypes of both species names. These name-bearing type specimens are deposited at the MNHN and NMPC, respectively.

Type specimens examined

Of O. confusus : Lectotype here designated (♂ MNHN): 1: TYPE (printed on red label). 2: MUSÉUM PARIS, 1936, COLL. A. BOUCOMONT (printed on light green label with black

border). 3: Huigra, 1000 m, F. Ohs, 10.7.05 (printed in purple on white label) 4: O. ophion Er., V. confusus Bouc. (handwritten in italics on white label). 5: LECTOTYPE, Onthophagus ophion var. confusus Bouc. , ♂, des. F.Z. Vaz-de-Mello, 2014 (printed and handwritten on red label with black border). 6: LECTOTYPE, Onthophagus confusus Bouc., Des. M. Rossini, 2016 (printed on red label with black borded). Paralectotypes (1 ♂, 1 ♀ MNHN): 1: Huigra , 1000 m, F . Ohs, 10.7.05 (printed in purple on white label). 2: MUSÉUM PARIS, 1936, COLL. A. BOUCOMONT (printed on light green label with black border). 3: PARALECTOTYPE, Onthophagus ophion var. confusus Bouc. , des. F.Z. Vaz-de- Mello, 2014 (printed and handwritten on yellow label with black border). 4: PARALECTOTYPE, Onthophagus confusus Bouc., Des. M. Rossini, 2016 (printed on yellow label with black borded). (♀ MNHN): 1: ECUADOR, Loja, E .W. (printed on white label). 2: MUSÉUM PARIS, 1936, COLL. A. BOUCOMONT (printed on light green label with black border). 3: PARALECTOTYPE, Onthophagus ophion var. confusus Bouc. , des. F.Z. Vaz-de- Mello, 2014 (printed and handwritten on yellow label with black border). 4: PARALECTOTYPE, Onthophagus confusus Bouc., Des. M. Rossini, 2016 (printed on yellow label with black borded). (♂ MNHN): 1: Balzapamba ( Ecuad. ), R .Haensch S. (printed in black on cream label with black border). 2: MUSÉUM PARIS, 1936, COLL. A. BOUCOMONT (printed on light green label with black border). 3: PARALECTOTYPE, Onthophagus ophion var. confusus Bouc. , des. F.Z. Vaz-de-Mello, 2014 (printed and handwritten on yellow label with black border). 4: PARALECTOTYPE, Onthophagus confusus Bouc., Des. M. Rossini, 2016 (printed on yellow label with black borded). (1 ♂, 1 ♀ MNHN): 1: BOLIVIE, SUAPI (printed on cream label). 2: EX MUSEO N. VAN DE POLL (printed on cream label with black border). 3: MUSÉUM PARIS, 1936, COLL. A. BOUCOMONT (printed on light green label with black border). 4: PARALECTOTYPE, Onthophagus ophion var. confusus Bouc. , des. F.Z. Vaz-de-Mello, 2014 (printed and handwritten on yellow label with black border). 5: PARALECTOTYPE, Onthophagus confusus Bouc., Des. M. Rossini, 2016 (printed on yellow label with black borded). (♀ MNHN): 1: S. Ecuador, Landangui M . W. (printed on white label). 2: MUSÉUM PARIS, 1936, COLL. A. BOUCOMONT (printed on light green label with black border). 3: PARALECTOTYPE, Onthophagus ophion var. confusus Bouc. , des. F.Z. Vaz-de-Mello, 2014 (printed and handwritten on yellow label with black border). 4: PARALECTOTYPE, Onthophagus confusus Bouc., Des. M. Rossini, 2016 (printed on yellow label with black borded).

Of O . nabeleki: Lectotype here designated (♀ SMTD): 1: Ecuador (printed on green label) . 2: TYPUS (printed on red label with black border). 3: Onthophagus nabeleki , ♂, n. sp., Typ , Dr . V. Balthasar det. (printed and handwritten on white label). 4: Muesum für Tierkunde, Dresden ( SMTD) (printed on white label) . Paralectotypes (♂ NMPC): 1: W Ecuador . Guayaquil, F. Ohaus S. (printed on white label). 2: TYPUS (printed on red label with black border). 3: Onthophagus nabeleki n.sp., Dr . V. Balthasar det. (printed and handwritten on white label). 4: Mus. Nat . Pragae , 26225, Inv. (printed and handwritten on orange label) . 5: ex. coll. V . Balthasar , National Museum Prague, Czech Republic (printed on white label) . 6: PARALECTOTYPE, Onthophagus nabeleki Balth. , des. F.Z. Vaz-de-Mello , 2013 (printed and handwritten on yellow label with black border). (♂ NMPC) : 1: S. Ecuador, Ciano M.W. (printed on white label). 2: TYPUS (printed on red label with black border). 3: Mus. Nat . Pragae , 26226, Inv. (printed and handwritten on orange label) . 4: nabeleki m. (handwritten on green label with black border). 5: ex. coll. V . Balthasar , National Museum Prague, Czech Republic (printed on white label) . 6: PARALECTOTYPE, Onthophagus nabeleki Balth. , des. F.Z. Vaz-de-Mello , 2013 (printed and handwritten on yellow label with black border). (♀ NMPC) : S Ecuador, Huairapongo, F. Ohs ., 17.10.05 (printed in purple on white label). 2: Typus (handwritten on red label) . 3: Onthophagus nabeleki n.sp., Dr . V. Balthasar det. (printed and handwritten on white label). 4: Mus. Nat . Pragae , 26227, Inv. (printed and handwritten on orange label) . 5: ex. coll. V . Balthasar , National Museum Prague, Czech Republic (printed on white label) . 6: PARALECTOTYPE, Onthophagus nabeleki Balth. , des. F.Z. Vaz-de-Mello , 2013 (printed and handwritten on yellow label with black border). (♀ NMPC) : 1: Ecuador (printed on green label). 2: TYPUS (printed on red label with balck border). 3: Onthophagus nabeleki n.sp., Dr . V. Balthasar det. (printed and handwritten on white label). 4: Mus. Nat . Pragae , 26228, Inv. (printed and handwritten on orange label) . 5: ex. coll. V . Balthasar , National Museum Prague, Czech Republic (printed on white label) . 6: PARALECTOTYPE, Onthophagus nabeleki Balth. , des. F.Z. Vaz-de-Mello , 2013 (printed and handwritten on yellow label with black border). (♂ NMPC) : 1: W Ecuador, Huigra, F . Ohaus S. / 10.7.05 (printed and handwritten on white label). 2: Huigra , 1000 m ., F. Ohs . 16.7.05 (printed in purple on white label). 3: Typus (handwritten on red label) . 4: Onthophagus nabeleki n.sp., Dr . V. Balthasar det. (printed and handwritten on white label). 5: PARALECTOTYPE, Onthophagus nabeleki Balth. , des. F.Z. Vaz-de-Mello , 2013 (printed and handwritten on yellow label with black border) .

Type specimens examined

ECUADOR: Cotopaxi: Otonga . 00°25 ’ S, 79°00 ’ W. 1800 m (1 ♂, 3 ♀♀ CEMT) . Las Pampas. 1800 m (1 ♂ MECN; 13 ♂♂, 12 ♀♀ MZc) . Esmeraldas: Esmeraldas. I-1956 (1 ♂, 1 ♀ CMNC) . San Mateo (9 ♂♂, 5 ♀♀ TAMU) . El Oro: Piñas. 1200 m (12 ♂♂, 18 ♀♀ CEMT; 3 ♂♂, 1 ♀ MZc) . 10 Km S Portovelo. 750 m . 10-V-1998 (1 ♂ CEMT). Arenillas. 03°34.24 ’ S, 80°08.86 ’ W. 13 m . 15- VI-2012 (1 ♂, 2 ♀♀ CEMT). Manabí: Chone. V-1976 (5 ♀♀ CMNC). 20 Km N Chone. 300 m . 06- 09-VI-1976 (12 ♂♂, 17 ♀♀ CMNC). Guayas: W Ecuador . Rio Pucay . Bucay. 300 m . 21-VI-1905 (1 ♂ NHRS). Palestina. 25 Km N Daule. 30 m . 22-27-VII-1976 (18 ♂♂, 3 ♀♀ CMNC). 30 Km NNE Playas. Tinalandia. 680 m . 24-25-VII-1975 (3 ♂♂, 2 ♀♀ CMNC). B.P. Cerro Blanco. 350 m . 06- IX-1997 (12 ♂♂, 13 ♀♀ CEMT). Loja: Jimbura. 2000-2100 m (3 ♂♂, CEMT). Gonzamaná. 2000 m (3 ♂♂ CEMT) . 5 Km N Zambi. 1300 m. 10-V-1998 (1 ♀ CEMT). Landangui (1 ♂ NHRS) . Macará. Catacocha. 650 m . 14-VIII-1977 (4 ♂♂, 2 ♀♀ CMNC). Pichincha: San José de Minas . “ Quito ”. 00°10 ’ 17 ” S, 78°24 ’ 42 ” W. 2400 m. 21-V-2014 (5 ♂♂, 4 ♀♀ CEMT) . 5 Km SE Nanegalito. 24-IV-1998 (1 ♂, 1 ♀ CEMT) . Nanegalito. 00°03 ’ S, 78°42 ’ W. 1500 m. 10-VII-1998 (1 ♀ CEMT). Mindo. 1200 m. 08-VII-1995 (3 ♀♀ CEMT). Santo Domingo de Los Colorados . IX-1981 (7 ♂♂, 11 ♀♀ MZc) . 34 Km Santo Domingo de los Colorados. 2000 m. 13-16-X-1986 (4 ♂♂, 4 ♀♀ MZc) . 47 Km S Santo Domingo. Rio Palenque Station . 230- 250 m . 1975-1976 (49 ♂♂, 73 ♀♀ CMNC) . 4 Km SE Santo Domingo. 500 ’. 08-22-VI-1976 (1 ♂ CMNC) . Tinalandia. 780 m . 13-16- X-1986 (1 ♀ MZc). Los Rios: Quevado. Pichilingue. I-1977 (6 ♂♂, 4 ♀♀ CMNC) . Mindo. Santiago Espinosa. 1500 m. 08-VII-1995 (1 ♂ CEMT) . Santo Domingo de los Tsáchilas: Puerto Limón. 00°17 ’ 14 ” S, 79°16 ’ 28 ” W. 397 m (1 ♂, 1 ♀ CEMT) . Santa Elena: 27 Km S Puerto López . 76 Km N Santa Elena . 500 ’. 25-27-VII-1976 (13 ♂♂, 8 ♀♀ CMNC) . Manglar Alto. 08-IX- 1981 (1 ♂ CMNC) . Olón. 01°47 ’ 46 ” S, 80°45 ’ 25 ” W. 20-XII-2012 (136 ♂♂, 163 ♀♀ CEMT) . Same locality. 01°46 ’ 54 ” S, 17°82 ’ 05 ” W. 20-XII-2012 (incorrect coordinates; 6 ♂♂, 21 ♀♀ CEMT). PERU: Piura: Morropón . Cacerio de Caracucho . 560 m . 12-XII-2007 (4 ♂♂, 6 ♀♀ CEMT).

Figures

Figure 4. Onthophagus confusus stat. nov. (a, b) Dorsal habitus of male and female. (c, d) Dorsal habitus of the lectotype and original labels (scale bar: 1 mm). (j, k) Lateral and dorsal view of the parameres (scale bar: 0.05 mm). (l, m) Lamella copulatrix and accessory lamellae. (n) Female genitalia (scale bar: 0.02 mm). Onthophagus nabeleki. (e, f) Dorsal habitus of the lectotype and original labels (scale bar: 1 mm). Onthophagus insularis. (g–i) Dorsal habitus of the lectotype and paralectotype, original labels of the name-bearing type (scale bar: 1 mm). (o, p) Lateral and dorsal view of the parameres (scale bar: 0.05 mm). (q, r) Lamella copulatrix and accessory lamellae (scale bar: 0.02 mm).

Figure 5. Geographic distribution of the species of the osculatii complex.