Onthophagus osculatii Guérin-Méneville, 1855

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 : 553-558

Onthophagus osculatii Guérin-Méneville, 1855

( Figures 1 (a), 1(b), 1(g), 1(h), 1(k – m), 5(a))

Onthophagus osculatii Guérin-Méneville, 1855: 589

Onthophagus osculatii Guérin-Méneville : Harold 1869: 1034; Harold 1880: 30; Boucomont and Gillet 1927: 207; Boucomont 1932: 305; Blackwelder 1944: 212; Balthasar 1951: 338; Vulcano and Pereira 1967: 564; Martínez 1987: 69 (erroneously cited) 1; Zunino and Halffter 1997: 161; Escobar 2000 b: 209; Vaz-de-Mello 2000: 194; Medina et al. 2001: 140; Pulido-Herrera and Zunino 2007: 112; Hamel-Leigue et al. 2008: 43; Medina and Pulido-Herrera 2009: 61; Ratcliffe et al. 2015; Silva et al. 2017: 4

Diagnosis

Onthophagus osculatii has the clypeus triangular, obtuse to slightly truncate at middle, elytral punctation distinct. Male with cephalic horns not carinate at the base, anteromedial pronotal protuberance narrowly rounded between horns. This species appears to be related to the Onthophagus included in the rubrescens complex but differs in having the elytral surface distinctly and finely microreticulated and slightly opaque.

Redescription

Body length 6 – 8 mm. Colour. Body completely black and weakly shining, pronotum sericeous on the posteromedial region, elytra slightly duller with faint metallic green iridescence, pygidium brownish, ventral side of the body dark-brown, mahogany coloured, femurs slightly lighter, last two abdominal sternites brownish. Mouthparts, antennal articles and tarsi brownish, antennal club brownish and opaque. Head. Clypeus elongated forward, triangular-shaped and obtusely truncated at middle, clypeal margin very weakly reflexed, genal and clypeal margins continuous, barely sinuated at the clypeo-genal junction. Fronto-clypeal region without transversal carina but feebly and transversally swollen, frontal horns straight and parallel, slightly higher than upper side of pronotum, tips of horns slightly curved backward. Surface of head with a dense and simple punctation, clypeus more distinctly punctated. Thorax. Anteromedial pronotal protuberance hump-like and rounded between cephalic horns, superior side of protuberance weakly convex (in lateral view). Pronotum slightly convex, lateral margins evenly curved, anterior angles obtusely acuminated, posterior margin of pronotum with border evanescent near posterior angles, pronotal punctation dense, very close (punctures separated by one diameter or less) and shallow, most of the punctures with a central pit, anterior angles with simple and fine punctures, punctation of the posteromedial region of pronotum obsolete and shallow.

Elytra slightly swollen, striae very shallow, interstriae flat to weakly convex at the base and apex. Elytral surface very finely microreticulated, interstrial punctation shallow and simple, punctures sparse and devoid of conspicuous setae, lateral interstriae (VI – VII) and apex of interstriae V and III with deeper punctures associated to short, straight and brownish setae. Propleuron feebly convex to very slightly concave on the anterior side, ventral side of the anterior angles of pronotum with a weak and rounded tubercle. Propleural carina thin, sides of propleuron with strong punctures associated with long and brownish setae curved upward (setae visible from above). Prosternum obtusely acuminated at middle, surface finely microreticulated and with long setae at middle. Mesosternal surface with coarse and deep punctures, and very short and light setae, central region of mesosternum smooth and slightly swollen, mesosternal margins smooth. Metasternum swollen and hump-like between middle legs, metasternal disc flattened, punctation fine, dense and widespread, punctures slightly deeper near the inferior margin of metasternum. Abdomen. Pygidium weakly convex and completely bordered, shining, with basal region finely microreticulated. Pygidial punctures of different sizes and evenly distributed across the surface, most of the punctures with a central pit. Legs. Foretibiae with four external teeth separated by small denticles, basal and external margin of protibiae with a series of serrate denticles, internoapical margin of protibiae with an obtuse and curved tooth, apical spur weakly curved downward and obtuse at the apex. Middle and hind legs unmodified. Male genitalia. In dorsal view, distal-superior area of the paramerites curved at the apex, external margin straight to very feebly curved. In lateral view, distal-inferior area straight to barely sinuate near the apex, tip of the paramerites slightly curved downward ( Figure 1 (g), 1(h)). Superior margin of the lamella copulatrix widely curved, especially on the right side. Superior left lobe narrow and slightly curved, external margin widely excavated. Basolateral margin of the inferior left lobe with a longitudinal, curved carina ( Figure 1 (k)), inferior right lobe slightly longer than inferior left lobe. Secondary lamella longitudinally elongated and sub-trapezoidal, with a complex keel on the superior margin. Female genitalia. Ventral sclerotization of the vagina distinct, central and inferior margin almost straight, very slightly excavated upward at middle ( Figure 1 (m)).

Minor males and females differ from major males by the following combination of characters: fronto-clypeal region of minor male with a very feeble carina or simply swollen as in major male, cephalic horns shorter, pronotal and elytral punctures stronger; female with clypeal margin either evenly curved or more distinctly truncated at middle, clypeo-genal junction more distinct, even though always feebly indicated, fronto-clypeal carina low and evenly developed, frons with a very low and sub-trapezoidal carina between eyes, either weakly curved backward or straight at middle (from above), pronotal and elytral punctation stronger and denser.

Distribution and ecology

Onthophagus osculatii encompasses a large geographic area, which comprises Bolivia, Peru, Ecuador, Colombia, Brazil, Venezuela, Guyana, French Guiana and Surinam ( Figure 5 (a)). Within its geographical range, O. osculatii appears to be mostly associated with Amazonian forests. However, this species also penetrates the Bolivian Chaco (Pando, Santa Cruz) and Brazilian cerrado (Mato Grosso), but it appears to maintain strong association with forested areas (e.g. semideciduous forests). According to the collection data reported on the labels of the specimens here examined, O. osculatii is mainly captured with pitfall traps baited with human excrement and carrion.

Morphological variation

Onthophagus osculatii exhibits a considerable and distinct chromatic variation, which appears to match the geographical distribution of different populations. Most of the specimens collected in Colombia, Ecuador, Peru, Brazil (Amazonas, Acre, Rondônia and Mato Grosso) and Bolivia have pygidium distinctly lighter (brownish, alutaceous or yellowish) than individuals collected in Venezuela, north-eastern Brazilian Amazonia (Roraima, Pará and Amapá) and the Guianas, where the pygidium is normally dark brown. The ground colour of the body can be variable too, but apparently does not follow any geographical pattern. Specimens can be either completely black and feebly shining or with pronotum shining and metallic green. Seldom pronotal surface either bronze-coloured or metallic green with cupreous to bronze sheen ( Bolivia: Pando). The elytra are reddish to mahogany-brown, with strong greenish to cupreous iridescence on the interstriae. Some individuals collected in north-eastern Brazilian Amazonia (Pará: Monte Dourado) have body completely brownish, with pronotum distinctly darker.

Remarks

In 1847 – 1848, the Italian naturalist Gaetano Osculati (1808 – 1894) undertook a long voyage in Equatorial America across the Napo and Amazonas rivers, departing from Guayaquil and reaching Belém, Pará. At the Yanayacu river ( Ecuador), right in the middle of the rain forest, Osculati was abandoned by the local guides, who took away most of his possessions. Over the following 14 days, the Italian entomologist carried on his collecting, but his poor health conditions and the bad weather forced Osculati to look for help, losing most of the collected specimens. Finally, he managed to reach Baeza, Archidona and Puerto Napo, where he recovered his health. Osculati stayed for about three months at the house of Dr Villavicencio, an amateur naturalist with whom he collected several birds and insects throughout this period. At the end of October 1847, Osculati left again to complete his trip, reaching the easternmost regions of Brazil on 30 March 1848 ( Papavero 1973).

According to this information, it is very likely that most of the still available specimens collected by Osculatii need to be referred to the second part of his trip, from Rio Misahualli ( Ecuador) to Belém ( Brazil).

Guérin-Méneville (1855) examined and identified the beetles collected by the Italian entomologist, describing several new species. Among them, he provides a very vague description of O. osculatii , whose diagnostic characters might actually lead one to any of the Onthophagus species included in the hircus group. And, more importantly, in no part of his work did Guérin-Méneville (1855) cite the geographical provenance of the examined specimens of O. osculatii .

Throughout our research, we visited several European natural history collections, including those where it was likely to find Osculati and Guérin-Méneville ’ s specimens. To date, information on the entomological collection of G. Osculati are very scarce. Horn et al. (1990) but also Rondani (1850, p. 357 – 358) state that the Diptera collected by Osculati during his South American voyage are housed at the Museo Civico di Storia Naturale of Milan. In addition, according to Bottoni (1929, p. 8, quoting Calidoni 1924), Osculati donated to this institute only the European Coleoptera included in his natural history collection. Our visit to the Coleoptera section of the Museum of Milan did not yield any scarab beetles attributable to the voyage of G. Osculati nor to the description of O. osculatii subsequently provided by Guérin-Méneville (1855). The curators of the Museum of Milan suppose the Osculati collection could have been destroyed and lost in the bombardments of the Second World War, as already reported by Pulido-Herrera and Zunino (2007) discussing the taxonomy of O. compressus Guérin-Méneville. However , the question about the collection of this Italian entomologist is still rather unclear.

As Guérin-Méneville (1855) worked directly on the Neotropical beetles collected by G. Osculati, the syntypes of the newly described species should be deposited in his collection. However, over the years the Guérin-Méneville collection was broken up and distributed to several institutes and probably private collections. Cambefort (2006) states that most of the Coleoptera specimens originally included in the Guérin-Méneville collection ended up in Paris ( MNHN) through different natural history collections (René Oberthür, Félix Monchicourt, Charles Jacob Piochard de La Brûlerie and Maurice Sédillot ) . According to Horn et al. (1990), the Lamellicornia beetles are today held in the Monchicourt collection, whose first choice was sold by Emile Deyrolle to Roger Comte de Bonneuil . Part of the Bonneuil collection was bought by Eugène Le Moult and is currently incorporated in the collection of the Institut Royal des Sciences Naturelles de Belgique, Brussels .

We visited the Coleoptera section of the MNHN but we did not find any type specimens of O. osculatii . However, in a recent visit to the IRSNB, FZVM rediscovered in the Bonneuil collection some Scarabaeinae specimens belonging to the syntypic series of species described by Guérin-Méneville (1855). However, even among these specimens there were no syntypes of O. osculatii .

Thus, according to the information provided above, specimens belonging to the type series of O. osculatii are considered lost, like most of the insect specimens supposed to be in the Guérin-Méneville collection. Therefore, in order to provide an objective identification of O. osculatii Guérin-Méneville, 1855 , we decided to fix a name-bearing type specimen by designating a neotype (ICZN 1999, Art. 75, Recommendation 75A). This nomenclatural act will allow researchers interested in South American Onthophagus to easily distinguish O. osculatii from sympatric and morphologically very similar species.

The specimen designated as neotype was selected according to the following criteria: (i) it is a well-preserved male specimen, as the high morphological similarity between female specimens belonging to different species could introduce further taxonomic confusion; (ii) it matches as much as possible the description provided by Guérin- Méneville (1855); and (iii) it was collected in a locality as close as possible to the ones reached by G . Osculati in the second part of his South American expedition (see Papavero 1973) . Indeed, as the type locality of O. osculatii was not mentioned by Guérin-Méneville (1855), at the moment it is not possible to identify the exact locality of collection of this species. So, on the basis of the available material, a male specimen collected in Amazonas ( Brazil) was designated as neotype (see below) . The neotype has been kindly donated by the curator of the CEMT (FZVM) and is now deposited at the Institut Royal des Sciences Naturelles de Belgique, Brussels .

Type specimens examined

Neotype here designated (♂ IRSNB): 1: Brasil. Amazonas. BR319. Km350. 05°12 ’ 56.4 “ S, 61°50 ’ 22.6 “ W. 30.vii. 07 -2.viii.2007. H. Gasca leg. (printed on white label with black border). 2: NEOTYPE, Onthophagus osculatii Guérin-Méneville, 1855 , Des. M. Rossini, 2016 (printed on red label with black border).

Additional material examined

COLOMBIA: Amazonas: W Leticia . Tarapacá-Monila-Amena (1 ♂ CEMT) . Leticia . 700 ’. 20- 25-1972 (2 ♂♂, 2 ♀♀ CMNC) . Same locality . Los Alpes . 215 m (13 ♂♂, 2 ♀♀ CMNC) . Guaviare: between San José del Guaviare and El Retorno . Elí farm . 02°21 ’ 46 ” N, 72° 38 ’ 29 ” W. 15-VII-2007 (1 ♂ CEMT) . BOLIVIA: Pando: Santa Rosa in Manuripi National Wildlife Reserve . 12°00 ’ S, 68°52 ’ W. 180- 190 m . 24-X-2004 (2 ♂♂, 3 ♀♀ OUMNH) . Cobija . Reserva San Sebastian Tahuamanu . Distb Amaz frst . 11°24 ’ 27 ” S, 69°01 ’ 04 ” W. 20-XII-2003 (4 ♂♂, 3 ♀♀ OUMNH) . Santa Cruz: Pua Sara . Santa Rosa . XI-1972 (1 ♂ CMNC) . BRAZIL: Acre: Xapuri . Reserva Chico Mendes . Floresta amazônica . 10°18.199 ’ S, 68°41.479 ’ W. 16-X-2008 (1 ♀ CEMT). Same locality . 10°18.337 ’ S, 68°41.564 ’ W. 18-X-2008 (2 ♀♀ CEMT). Same locality . 10°18.199 ’ S, 68°41.479 ’ W. 16-X-2008 (1 ♀ CEMT). Amapá: Serra do Navio . Cava Urucum - Amapari. 00°53 ’ 06 ” S, 51°52 ’ 53 ” W. 05-VII-2000 (1 ♂, 1 ♀ CEMT) . Same locality . X-1957 (1 ♂ CMNC) . Amazonas: Manaus. Reserva Adolpho Ducke . 16-V-2012 (1 ♂, 1 ♀ CEMT) . BR 319

Km 350 . 05°12 ’ 56.4 ” S, 61°50 ’ 22.6 ” W. VII-VIII-2007 (1 ♂, 1 ♀ CEMT) . Mato Grosso: Alta Floresta . Ceplac . 24-V-2009 (12 ♂♂, 3 ♀♀ CEMT) . Same locality . Frag . FAH. 09°52 ’ 52 ” S, 56° 06 ’ 12 ” W. III-2008 (6 ♂♂, 1 ♀ CEMT) . Same locality . Frag . 7. 09°56 ’ 48 ” S, 56°06 ’ 33 ” W. III-2008 (8 ♂♂, 9 ♀♀ CEMT) . Same locality . Frag . 16. 09°41 ’ 24 ” S, 55°56 ’ 28 ” W. VI-2008 (15 ♂♂, 13 ♀♀ CEMT) . Same locality . Frag . 26. 09°53 ’ 40 ” S, 56°16 ’ 34 ” W. VI-2008 (3 ♂♂, 5 ♀♀ CEMT) . Same locality . Frag . 27. 09°49 ’ 44 ” S, 55°19 ’ 51 ” W. V-2008 (8 ♂♂, 9 ♀♀ CEMT) . Same locality . Frag . 29. 09°56 ’ 52 ” S, 56°03 ’ 02 ” W. V-2008 (1 ♂ CEMT) . Same locality . Frag . 42. 09°46 ’ 55 ” S, 56°02 ’ 13 ” W. IV-2008 (7 ♂♂, 1 ♀ CEMT) . Same locality . Frag . 48. 09°58 ’ 46 ” S, 56°05 ’ 31 ” W. V- 2008 (3 ♂♂, 3 ♀♀ CEMT) . Same locality . Frag . 61. V-2008 (13 ♂♂, 11 ♀♀ CEMT) . Same locality . Frag . 62. 09°53 ’ 15 ” S, 55°59 ’ 43 ” W. V-2008 (33 ♂♂, 36 ♀♀ CEMT) . Same locality . Frag . 67. 09°46 ’ 22 ” S, 56°11 ’ 48 ” W. VI-2008 (30 ♂♂, 36 ♀♀ CEMT) . Same locality . Frag . 94. 09° 51 ’ 59 ” S, 55°54 ’ 02 ” W. IV-2008 (1 ♀ CEMT). Same locality . Frag . 114. 09°45 ’ 19 ” S, 55°58 ’ 19 ” W. VI-2008 (1 ♂, 3 ♀♀ CEMT) . Same locality . Frag . 143. 09°54 ’ 27 ” S, 56°03 ’ 18 ” W. III-2008 (2 ♀♀ CEMT). Same locality . Frag . 150. 09°35 ’ 54 ” S, 55°56 ’ 09 ” W. IV-2008 (68 ♂♂, 51 ♀♀ CEMT) . Same locality . Frag . 151. 09°44 ’ 55 ” S, 56°01 ’ 41 ” W. IV-2008 (12 ♂♂, 21 ♀♀ CEMT) . Same locality . Frag . 152. 09°56 ’ 33 ” S, 55°55 ’ 33 ” W. V-2008 (4 ♂♂, 3 ♀♀ CEMT) . Same locality . Frag . 153. 09°50 ’ 22 ” S, 56°00 ’ 21 ” W. IV-2008 (22 ♂♂, 21 ♀♀ CEMT) . Same locality . Frag . 154. 09° 49 ’ 27 ” S, 55°53 ’ 34 ” W. IV-2008 (3 ♂♂, 2 ♀♀ CEMT) . Same locality . Frag . 155. 10°01 ’ 13 ” S, 56° 26 ’ 11 ” W. VI-2008 (4 ♂♂, 4 ♀♀ CEMT) . Same locality . Frag . 156. 09°54 ’ 24 ” S, 56°02 ’ 39 ” W. VI- 2008 (1 ♂ CEMT) . ContriguaÇu . Faz . São Nicolau. Prainha . 09°51 ’ 36 ” S, 58°12 ’ 53 ” W. X-2009 (4 ♂♂, 4 ♀♀ CEMT) . Same locality . 09°49 ’ 27.15 ” S, 58°16 ’ 15.99 ” W. 10-XI-2010 (2 ♂♂, 2 ♀♀ CEMT) . Same locality . 09°51 ’ 18 ” S, 58°13 ’ 20 ” W. X-XII-2009 (3 ♀♀ CEMT). Same locality . 09° 50 ’ 24 ” S, 58°15 ’ 10 ” W. X-2009 (1 ♂, 1 ♀ CEMT) . Same locality . Castanheira . 09°49 ’ 18 ” S, 58° 17 ’ 18 ” W. XII-2009 (10 ♂♂, 11 ♀♀ CEMT) . Same locality . 09°52 ’ 05 ” S, 58°13 ’ 35 ” W. XII-2009 (1 ♂ CEMT) . Same locality . 09°49 ’ 08 ” S, 58°15 ’ 40 ” W. X-2009 (1 ♀ CEMT). Same locality . Matinha . Borda da mata. 09°50 ’ 19 ” S, 58°15 ’ 03 ” W. 13-XII-2009 (8 ♂♂, 4 ♀♀ CEMT) . Same locality . Ilha do Rio Juruena . 09°53 ’ 42.1 ” S, 58°13 ’ 28.64 ” W. X-2010 (3 ♂♂, 2 ♀♀ CEMT) . Diamantino. Vale da Solidão . 14°32 ’ 13 ” S, 56°07 ’ 12 ” W (5 ♂♂, 2 ♀♀ CEMT) . Same locality . Alto Rio Arinos . I-2001 (6 ♂♂, 5 ♀♀ CEMT) . Same locality . Fazenda São João . 14°14 ’ 10 ” S, 56°08 ’ 11 ” W. 400 m . 11-I-2001 (9 ♂♂, 7 ♀♀ CMNC). Porto Estrela. ESEC Serra das Araras. Trilha da Boca do José . 15°39 ’ 00 ” S, 57°12 ’ 31 ” W. (22 ♂♂, 30 ♀♀ CEMT) . Same locality . Mata ciliar . 15-X-2011 (7 ♂♂, 6 ♀♀ CEMT) . Same locality . Olho d ’ Água . 14-X-2011 (5 ♀♀ CEMT). Same locality . 15°21 ’ 49 ” S, 56°57 ’ 32 ” W. 12-X-2011 (1 ♀ CEMT). Same locality . 15°38 ” 25 ” S, 57°12 ’ 17 “ W (2 ♂♂ CEMT) . Same locality . Vale do Saloba. (5 ♂♂, 2 ♀♀ CEMT) . Same locality . 15°38 ’ 50 ” S, 57°12 ’ 27 ” W (11 ♂♂ CEMT) . Same locality . 15°39.10 ’ S, 57°12.80 ’ W (3 ♂♂, 1 ♀ CEMT) . Same locality . 15°38 ’ 56 ” S, 57°12 ’ 34 ” W (1 ♂ CEMT) . Same locality . 15°38 ’ 59 “ S, 57° 12 ’ 38 ” W (8 ♂♂, 5 ♀♀ CEMT) . Same locality . 15°38 ’ 53 ” S, 57°12 ’ 30 ” W (3 ♂♂, 2 ♀♀ CEMT) . Novo Mundo. Parque Estadual do Cristalino . 09°27 ’ 60 ” S, 55°50 ’ 02 ” W. V-2013 (1 ♂ CEMT) . Same locality . 09°27 ’ 38 ” S, 55°48 ’ 26 ” W. V-2013 (1 ♂ CEMT) . Same locality . 09°27 ’ 31 ” S, 55° 47 ’ 54 ” W. V-2013 (1 ♂ CEMT) . Same locality . 09°28 ’ 16 ” S, 55°48 ’ 47 ” W. V-2013 (1 ♂ CEMT) . Same locality . 09°28 ’ 01 ” S, 55°47 ’ 43 ” W. V-2013 (1 ♂ CEMT) . Same locality . 09°28 ’ 16 ” S, 55° 48 ’ 47 ” W. V-2013 (2 ♂♂ CEMT) . Nova Mutum. Trivelato . XII-1995 (7 ♂♂, 10 ♀♀ CEMT) . Querência . Faz . São Luiz. 12°41.692 ’ S, 52°30.354 ’ W. II-2009 (1 ♂♂ CEMT) . Same locality . 12° 40.49 ’ S, 52°21.94 ’ W (BF 1-500 m). 09-VII-2009 (5 ♂♂, 5 ♀♀ CEMT) . Same locality . 12°41.09 ’ S, 52°30.28 ’ W (UF 4-500 m). 09-VII-2009 (2 ♂♂, 1 ♀ CEMT) . Same locality . 12°39.72 ’ S, 52° 22.74 ’ W (UF 2-200 m). 17-VII-2009 (8 ♂♂, 2 ♀♀ CEMT) . Nova Mutum. 13°48 ’ 07 ” S, 56° 05 ’ 22 ” W. 19-I-2011 (2 ♂♂ CEMT) . Tangará da Serra . Sitio Boa Vista. 14°36 ’ 42 ” S, 57°50 ’ 12 ” W. 328 m . 09-11-II-2012 (1 ♂ CEMT). Same locality . Faz . Sudamata . 14°37 ’ 34 ” S, 57°58 ’ 24 ” W. 10- 17-III-2012 (7 ♂♂, 1 ♀ CEMT) . Same locality . 14°38 ’ 13 ” S, 57°56 ’ 08 ” W. 20-22-II-2012 (8 ♂♂, 5 ♀♀ CEMT) . Same locality . 14°37 ’ 18 ” S, 57°58 ’ 01 ” W (22 ♂♂, 21 ♀♀ CEMT) . Same locality . 14° 37 ’ 19 ” S, 57°58 ’ 04 ” W (3 ♂♂, 1 ♀ CEMT) . Same locality . 14°37 ’ 32 ” S, 57°58 ’ 12 ” W (15 ♂♂, 11 ♀ CEMT) . Same locality . Fazenda Paraiso. 14°41 ’ 46 ” S, 57°24 ’ 40 ” W. 13-15-I-2011 (1 ♂ CEMT) . Same locality . Fazenda Fontosa. 14°35 ’ 52 ” S, 57°50 ’ 13 ” W (2 ♂♂, 1 ♀ CEMT) . Same locality . Fazenda Rosa Branca . 14°33 ’ 50 ” S, 57°52 ’ 16 ” W. 25-27-II-2011 (1 ♂ CEMT) . Same locality . 14° 34 ’ 00 ” S, 57°52 ’ 24 ” W. 18-20-II-2011 (4 ♂♂, 2 ♀♀ CEMT) . Same locality . Fazenda Netolândia. 14°39 ’ 54 ” S, 57°55 ’ 08 ” W. 12-14-III-2012 (7 ♂♂, 5 ♀♀ CEMT) . Same locality . 14°39 ’ 48 ” S, 57° 54 ’ 13 ” W. 19-21-III-2012 (6 ♂♂, 9 ♀♀ CEMT) . Same locality . 14°41 ’ 05 ” S, 57°54 ’ 08 ” W. 25-27- III-2012 (2 ♂♂, 2 ♀♀ CEMT) . Same locality . 14°39 ’ 56 ” S, 57°54 ’ 08 ” W. 20-22-III-2012 (2 ♂♂, 6 ♀♀ CEMT) . Araputanga . Fazenda Bandeirantes. 15°22 ’ 14 ” S, 58°26 ’ 02 ” W. 20-22-I-2013 (6 ♂♂, 4 ♀♀ CEMT) . Indiavai . Fazenda Alto Jaurú . 15°27 ’ 18 ” S, 58°33 ’ 12 ” W. 14-16-I-2013 (1 ♂, 1 ♀ CEMT) . Mirassol d ’ Oeste . Faz . Sta . Helena . 15°36 ’ 34 ” S, 57°58 ’ 12 ” W. 25-27-I-2013 (1 ♀ CEMT). Cláudia . Fazenda Continental. 11°36 ’ 29 ” S, 55°15 ’ 01 ” W. 20-II-2010 (4 ♂♂, 3 ♀♀ CEMT) . Nova Bandeirante. Margem do rio Juruena . 09°52 ’ 47 ” S, 58°14 ’ 07 ” W. X-2010 (2 ♂♂, 1 ♀ CEMT) . Sinop . X-1976 (1 ♂ CNCI) . Pará: Belém . IPEAN (Instituto de Pesquisa e ExperimentaÇão Agropecuárias do Norte ) . X-1984 (2 ♀♀ CEMT). Novo Progresso. Faz . Florentino . 06°51 ’ 58.61 ” S, 55°29 ’ 23.28 ” W. 12-I-2011 (2 ♂♂, 4 ♀♀ CEMT) . Monte Dourado. Jari Cellulose Area 95 . Eucalyptus plantation. 00°41 ’ S, 52°48 ’ W. III-IV (2 ♂♂, 2 ♀♀ CEMT) . Same locality . 00°36 ’ S, 52°39 ’ W (2 ♀♀ CEMT). Same locality . 00°42 ’ S, 52°46 ’ W (2 ♀♀ CEMT). Same locality . 00°35 ’ S, 52°39 ’ W (1 ♂, 2 ♀♀ CEMT) . Same locality . 00°80 ’ S, 52°66 ’ W (1 ♀ CEMT). Same locality . 01°01 ’ S, 52°44 ’ W (1 ♀ CEMT). Same locality . Reserva do Pacanari . 00° 38 ’ S, 52°34 ’ W (2 ♂♂, 4 ♀♀ CEMT) . Bituba . 01°11 ’ S, 52°38 ’ W. 180 m . VII-2004 (1 ♂, 2 ♀♀ CEMT). Same locality . 00°50 ’ S, 53°02 ’ W (1 ♂, 1 ♀ CEMT) . Km 27 Altamira-Marabá . II-III-2001 (1 ♀ CEMT). RedenÇão . Pinkaití-Aik . 07°46 ’ S, 51°58 ’ W (2 ♀♀ CEMT). São Felix do Xingú . Pinkaití Reserve . 07°45 ’ S, 51°57 ’ W. 11-XI-1998 (1 ♀ CEMT). Pau d ’ Arco . Faz . Marajoara . 07° 50 ’ S, 50°16 ’ W. 11-X-1998 (1 ♂ CEMT) . Jacareacanga . Rio Teles Pires . Sete Quedas. 09° 18 ’ 40 ” S, 56°46 ’ 10 ” W. 08-VI-2009 (1 ♂, 3 ♀♀ CEMT) . Paranaíta . Rio Teles Pires . Faz. Argão. 05°13 ’ 37 ” S, 56°59 ’ 56 ” W. 09-VI-2009 (2 ♂♂ CEMT) . Santarém . Reserva Tapajós. 02°46.075 ’ S, 55°40.438 ’ W. 25-XII-2008 (2 ♂♂, 1 ♀ CEMT) . Serra Norte. Carajás . X-XI-1984 (3 ♂♂, 3 ♀♀ CEMT) . Tucurui . 09-17-XII-1985 (1 ♂ CMNC) . Rondônia: Guajará Mirim . 10°22 ’ 16.05 ” S, 64° 44 ’ 47.11 ” W. 27-I-2010 (2 ♂♂, 2 ♀♀ CEMT) . Same locality . Fazenda Agropecuária. 10° 37 ’ 47.59 ” S, 64°59 ’ 52.58 ” W. 15-I-2010 (2 ♂♂ CEMT) . Porto Velho. ESEC-Cuniã. 08° 04 ’ 11.82 ” S, 63°28 ’ 34.64 ” W. 14-16-VIII-2012 (13 ♂♂, 7 ♀♀ CEMT) . Roraima: Apiaú . VII- 1997 (3 ♂♂, 2 ♀♀, CEMT). Amajari. Ilha de Maracá . 03°30.762 ’ S, 61°35.261 ’ W. 09-V-2009 (2 ♂♂ CEMT) . Same locality . 03°30.211 ’ S, 61°35.000 ’ W. 09-V-2009 (4 ♂♂, 2 ♀♀ CEMT) . Same locality . 03°31.930 ’ S, 61°36.302 ’ W. 10-V-2009 (2 ♂♂, 1 ♀ CEMT) . Same locality . 03° 21.951 ’ S, 61°27.601 ’ W. 16-V-2009 (1 ♂, 4 ♀♀ CEMT) . Same locality . 03°31.722 ’ S, 61° 36.486 ’ W. 10-V-2009 (2 ♂♂, 4 ♀♀ CEMT) . Vila Pacaraima. 04°27 ’ N, 61°07 ’ W. 820 m . IX- 1996 (12 ♂♂, 13 ♀♀ CEMT). Cantá . Serra Negra. IX-1996 (1 ♂, 1 ♀ CEMT) . Caracaraí . VII- 1997 (1 ♂, 2 ♀♀ CEMT) . ECUADOR: Pastaza: Villano . 01°29 ’ 48 ” S, 77°28 ’ 53 ” W. 03-VII-1996 (2 ♂♂, 1 ♀ CEMT) . Napo: Yasuní . 250 m. 03-IV-1995 (1 ♀ CEMT). 20 Km S Tena . 09-11-VII- 1976 (2 ♀♀ CMNC). Estación Cientifica Yasuní . 215 m . 05-10-IX-1999 (6 ♂♂, 5 ♀♀ TAMU). Jatun Sacha Biological Station . 01°04 ’ S, 77°36 ’ W. 450 m. 10-12-II-1995 (1 ♀ BGc). Same locality (21 Km E Puerto Napo) . 400 m . 08-VII-1994 (2 ♂♂, 3 ♀♀ CMNC). Orellana: Tiputini Biodiversity Station . VII-VIII-2008 (1 ♀ CMNC). FRENCH GUIANA: Cayenne: Cayenne . Paracou Field Station . 05°02 ’ N, 53°00 ’ W (18 ♂♂ CEMT) . Same locality . La Chaumière. IV- 1978 (5 ♂♂, 6 ♀♀ CMNC) . Mont Grand Matoury (1 ♂ CEMT; 1 ♂ BGc). 20 Km SW Cayenne . 04°48 ’ 18 ” N, 52°28 ’ 41 ” W (2 ♂♂ CMNC) . Roura (8.4 Km SSE) . 04°40 ’ 41 ” N, 52°13 ’ 25 ” W. 200 m (4 ♂♂, 2 ♀♀ CMNC) . Same locality (7 Km N) . Same locality (39.4 Km SSE) . 04°32 ’ 43 ” N, 52° 08 ’ 26 ” W. 270 m (5 ♂♂, 4 ♀♀ CMNC) . Kaw road . PK-38. 19-20-VIII-1995 (1 ♀ CMNC; 1 ♂ FSCA) . Crique Plomb. 24-25-IX-1992 (1 ♂ BGc). Saint-Laurent-du-Maroni: Saint Laurent du Maroni. IV-1976 (2 ♂♂, 2 ♀♀ BGc). Saül . 03°37 ’ 55 ” N, 53°11 ’ 52 ” W. VII-1977 (2 ♂♂ CEMT) . Same locality (2 ♂♂ MSNM) . 1 Km NW Les Eaux Claires . 03°39 ’ 46 ” N, 53°13 ’ 19 ” W. 220-280 m (7 ♀♀ CMNC). Wanaboo (near Nason) . Marowijne river . 04°43 ’ 35 ” N, 54°26 ’ 36 ” W. 05-VI- 1999 (5 ♀♀ CMNC). D-5 4k SE Tngmd Jct. 25-27-VIII-1995 (3 ♂♂, 3 ♀♀ CMNC) . No locality: VII-1973 (1 ♂ CEMT) . GUYANA: Potaro-Siparuni: Iwokrama Forest Reserve . 04°40 ’ 19 ” N, 58°41 ’ 04 ” W. 100- 200 m . V-VI-2001 (10 ♂♂, 1 ♀ CMNC) . Sandstone . 4°23 ’ 13.2 “ N, - 58° 55 ’ 12 ” W (37 ♂♂, 13 ♀♀ MRc) Upper Takutu-Upper Essequibo: Mazaruni . Potaro District . Takutu Mountains . 06°15 ’ N, 58°55 ’ W. 08-10-XII-1983 (2 ♂♂, 1 ♀ CMNC) . Kanuku Mountains . 05-II-1993 (1 ♂ BGc). PERU: Loreto: 1.5 Km N Teniente López . 02°35 ’ 39.6 ” S, 76° 06 ’ 55 ” W. 230- 305 m . 18-26-VII-1993 (2 ♂♂, 2 ♀♀ CMNC) . Compamento San Jacinto . 02° 18 ’ 44.85 ” S, 75°51 ’ 46 ” W. 175- 215 m . 03-12-VII-1993 (2 ♂♂, 3 ♀♀ CMNC) . Madre de Dios: Rio Palma Real Grande . Limón camp. 12°32 ’ 20 ” S, 68°51 ’ 41 ” W. 400 m 05-06-IV-1999 (1 ♂ CMNC) . Parque Manu. Pakitza . 11°53 ’ S, 70°58 ’ W. 400 m . 16-XI-1990 (1 ♂ CEMT). Pantiacolla Lodge. 5.5 Km NW El Mirador trail . Alto Madre de Dios River. 12°39 ’ 10 ” S, 71°15 ’ 28 ” W. 500 m. 23-26-X-2000 (1 ♀ CMNC). Tambopata . 200 m . X-1983 (6 ♂♂, 8 ♀♀ MZc). Rio Tambopata . Ccolpa de Guacamayos . 13°8,5 ’ S, 69°36,4 ’ W. 300 m X-1995 (1 ♂ BGc). SURINAM: Para: Saramacca W . Suriname Road. 108 Km WSW Zanderij Airport . 05° 13 ’ 37 ” N, 55°52 ’ 54 ” W. 10-14-VI-1999 (5 ♀♀ CMNC). 11 Km SE Zanderij Airport . 18-20-VI- 1999 (1 ♂ CMNC) . Commewijne: Akintosoela . CELOS. Road to Redi Doli . 05°16 ’ 17 ” N, 54° 55 ’ 15 ” W. 40-50 m (4 ♂♂, 2 ♀♀ CMNC) . VENEZUELA: Bolívar: 20 Km E El Palmar . 18-VI- 1996 (1 ♀ CMNC). Anacoco . 09-VIII-2005 (1 ♂ CEMT) . Gransabana. San Francisco de Yuruani . 19-I-1988 (1 ♂, 1 ♀ CMNC) . Santa Elena. Icabaru Road. 04-06-VIII-1986. 1000 m (2 ♀♀ CMNC; 1 ♂, 1 ♀ MZc) . 105 Km S El Dorado . 17-VII, 07-VIII-1986 (1 ♂, 1 ♀ BGc). 120 Km S El Dorado . 20-27-XII-1987 (1 ♂ BGc). 10 Km S El Dorado . 17-VII, 07-VIII-1986 (1 ♂ BGc). 20 Km S El Dorado . 20-23-VII-1986 (3 ♂♂, 1 ♀ CMNC; 1 ♀ BGc) . 22 Km SE El Dorado . VI-VII- 1987 (4 ♀♀ CMNC). Guri. (2 ♂♂, 1 ♀ CMNC; 1 ♂, 2 ♀♀ MZc) . Rio Chicanán . 40 Km SW El Dorado . 22-23-VII-1986 (1 ♂ CMNC) . Nuria . 550 m . 30-V-1975 (2 ♂♂, 1 ♀ MZc). 10 Km E Icabaru . 700 m. 06-VII-1987 (1 ♀ CMNC). 10 Km E San Francisco de Yuruaní . Gransabana. 1300 m. 08-10-VII-1987 (1 ♂, 7 ♀♀ CMNC) . Caicara . Rio Suapure . 150 Km S Caicara . 03-10- IV-1977 (1 ♂, 1 ♀ CMNC) . Yaracuy: Aroa . 600 m . 21-V-1985 (1 ♂ MZc).

Figures

Figure 1. Onthophagus osculatii. (a, b) Dorsal habitus of male (neotype) and female (scale bar: 1 mm). (g, h) Lateral and dorsal view of the parameres (scale bar: 0.05 mm). (k, l) Lamella copulatrix and accessory lamellae; arrow indicating the basolateral carina of the inferior left lobe of O. osculatii. (m) Female genitalia (scale bar: 0.02 mm); arrow indicating the slight medial excavation of the ventral sclerotization of O. osculatii. Onthophagus chacoensis sp. nov. (c, d) Dorsal habitus of male (holotype) and female. (e) Metasternal punctuation (scale bar: 1 mm). (f) Detail of the pronotal punctuation. (i, j) Lateral and dorsal view of the parameres (scale bar: 0.05 mm). (n, o) Lamella copulatrix and accessory lamellae; arrow indicating the deep excavation of the external margin of the left lobe of O. chacoensis. (p) Female genitalia (scale bar: 0.02 mm).

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