Onthophagus curvicornis Latreille, 1812

Rossini, Michele, 2021, Additional mislabeling in African Onthophagus Latreille, 1802 (Coleoptera: Scarabaeidae: Scarabaeinae): the case of Onthophagus viviensis and Onthophagus laevatus, Zootaxa 5032 (2), pp. 262-274 : 266-268

Onthophagus curvicornis Latreille, 1812

Fig. 2A–H .

Onthophagus curvicornis Latreille, 1812: 220 (also illustrated in plate XXIII, Fig. 2 ) (actual publication date from Bousquet 2016); Harold 1869: 1028; Taschenberg 1870: 184; Harold 1880: 29; Campos 1921: 57; Boucomont & Gillet 1927: 205; Boucomont 1932: 308, 323; Blackwelder 1944: 211; Guêrin 1953: 262; Roze 1955: 45; Howden & Cartwright 1963: 24 (as Onthophagus curvicornis var. incensus ); Vulcano & Pereira 1967: 565; Deloya 1992: 22; Zunino & Halffter 1997: 161; Blanco Casanova 1998: 47; Escobar 2000: 209; Vaz-de-Mello 2000: 194; Kohlmann & Solis 2001: 210 (as Onthophagus curvicornis var. incensus ); Medina et al. 2001: 139; Cancino & Blanco 2002: 120; Medina et al. 2002: 182–185; Zunino 2003: 71; Noriega 2004: 40; Ramírez & Locarno 2004: 61–63; Krajcik 2006: 99; Pulido-Herrera & Zunino 2007: 101, 106 (as Onthophagus curvicornis var. incensus ); Pulido-Herrera et al. 2007: 307; Orozco & Pérez 2008: 38–39; Concha-Lozada et al. 2010: 43–44, 47–49, 52; Carvajal et al. 2011: 322–323; Deloya 2011: 386; Giraldo et al. 2011: 118; Cultid-Medina et al. 2012: 32, 51, 134, 155, 180; Delgado et al. 2012: 331; Noriega et al. 2012: 43, 46–47; Ferrer-Paris et al. 2013: 99–100, 109; Martínez-Quintero et al. 2013: 450; Martínez-Revelo & Lopera-Toro 2014: 65–68; Noguera-Urbano et al. 2014: 18; Pardo-Locarno & Camero 2014: 210, 212-213, 215; Cultid-Medina & Medina 2015: 133, 201 (Appendix 8), 201 (Appendix 11), 203 (Appendix 13); Cultid-Medina et al. 2015: 624; Molina et al. 2016: 128–130; Alburqueque et al. 2017: 211; Rodríguez 2017: 20–22, 24–26; Torres et al. 2017: 48–49, 67, 69; Villada-Bedoya et al. 2017: 197, 200, 203; Chamorro et al. 2018: 97; Giraldo et al. 2018: 104, 111, 132; Girón et al. 2018: 83; Rossini et al. 2018b: 9; Chamorro et al. 2019: 186 ; Escobar et al. 2020 (pages not numbered); Martínez & Morales 2020: 1689; Martínez-Revelo et al. 2020: 6; Nieto et al. 2020: 136; Carrión-Paladines et al. 2021: 9–10, 14, 16–17.

Onthophagus minax Kirsch, 1866: 215 ; Harold 1869: 1028; Boucomont & Gillet 1927: 205; Blackwelder 1944: 211; Pulido- Herrera & Zunino 2007: 101; Chamorro et al. 2019:186

Onthophagus laevatus d’Orbigny, 1902: 69 new synonym; d’Orbigny 1905: 437; Peringuey 1908: 587; d’Orbigny 1913: 174 (identification key); d’Orbigny 1914: 640; Boucomont & Gillet 1927: 177; Ferreira 1972: 666; Daniel & Génier 2019: 28 (mentioned as Onthophagus (s.l.) laevatus )

Remarks. d’Orbigny (1905) considered O. viviensis to be closely related to O. laevatus , which was described by himself three years before from Delagoa, Mozambique: “ Cette espèce est extrêmement voisine du laevatus d’Orbigny (1902 , in Ann. Soc. ent. Fr., p. 69), décrit du Mozambique; elle en diffère seulement par sa coloration en grande partie bronzée ou verdâtre (au lieu d’être entièrement d’un noir d’ébène), la massue des antennes testacée, les stries des élytres pas plus enfoncées à la base, le pygidium à ponctuation moins régulièrement espacée ”. According to d’Orbigny (1902), the type series of O. laevatus is composed by two female specimens that the author received from Otto Staundinger (1830–1900) and Andreas Bang-Haas (1846–1925). The two syntypes are currently housed in the MNHN of Paris and their study revealed that they belong to the well-known South American O. curvicornis Latreille, 1812 . Therefore, O. laevatus d’Orbigny, 1902 (new synonymy) is established as junior subjective synonym of O. curvicornis and thus removed from Afrotropical Onthophagus . One of the two specimens from the type series of O. laevatus is designated as lectotype, while the remaining specimen is labeled as paralectotype (see type material examined below).

The taxonomic questions related to O. curvicornis , however, are far from being completely resolved. This species occurs from a wide elevational gradient in the Andes, and the study of a large sample of specimens collected from northern Peru to Venezuela, including also specimens from Trinidad ( Trinidad and Tobago), allowed me to preliminary isolate four different morphological forms. These morphotaxa differ by the color and texture of their dorsal tegument (from black to metallic green or cupreous; from shiny and very little microsculptured to opaque, slightly bright and clearly microsculptured); and shape of the pronotal protuberance in male (from conical to rounded in dorsal view). Very slight differences have been noted in the male genitalia and particularly in the endophallites. The taxonomic separation of these morphological forms will be treated in a separate paper.

According to Horn et al. (1990), many insect specimens of the Pierre A. Latreille (1762–1833) original collection were incorporated in the Pierre F.M.A. Dejean (1780–1845) collection. Subsequently, the latter was split into several lots and sold to museums and private entomologists. A considerable part of Dejean collection is today housed at the MNHN of Paris, while a second and significant lot of insects was acquired by Massimiliano Spinola (1780–1857) and now preserved at the Museo Regionale di Scienze Naturali of Turin (MRSN). However, some Latreille’s type specimens may be also found elsewhere (e.g., The Natural History Museum of London, see Platycoelia flavostriata (Latreille, 1813) in Smith 2003). The extensive examination of historical collections did not allow me to find any potential specimen to be considered the Latreille’s type of O. curvicornis . The type material of this species was collected by Alexander von Humboldt and Aimé Bonpland during their famous voyage across northern South America, and O. curvicornis was apparently collected in Quito, in cattle or horse dung. It is very likely that the type material of O. curvicornis (unknown the exact number of specimens that were studied by Latreille) is lost. Therefore, a neotype of O. curvicornis is here designated. Among the examined material no specimens from Quito, Ecuador, were studied, and the only specimens that satisfied the original description and type locality were collected in San Francisco de Las Pampas and 112 km west of Latacunga (see material examined below). According to Recommendation 76A.1.2 of the code ( International Commission on Zoological Nomenclature 1999), to ascertain and clarify a type locality the author should take into account “ collector’s notes, itineraries, or personal communications ”. Thus, one specimen collected in Latacunga would have been the best choice, as Humboldt and Bompland visited this locality during their travel in Ecuador ( Sandwith 1926; Papavero 1971). However, this location is also part of the distribution range of one of the four morphological forms of O. curvicornis and geographically more distant to Quito than San Francisco de Las Pampas. Thus, a male specimen collected in San Francisco de Las Pampas, whose morphology matches the original description of O. curvicornis is selected as neotype.

Type material examined. Of O. curvicornis : neotype here designated (1 ♂, MZUF): ECUADOR: COTO- PAXI, San Francisco de Las Pampas (1300-1500 m), II.1993, L. Bartolozzi (N. Mag. 1406) / NEOTYPE, Onthophagus curvicornis Latreille, 1812 , Des. M. Rossini, 2021.

Of O. minax : lectotype here designated (1 ♂, SMTD): Bogota, Kirsch / Staatl. Museum für Tierkunde Dres- den / LECTOTYPE, Onthophagus minax Kirsch, 1866 , Des. M. Rossini, 2021. Paralectotypes (2 ♂, 2 ♀ SMTD): same data as lectotype / same data as lectotype / PARALECTOTYPE, Onthophagus minax Kirsch, 1866 , Des. M. Rossini, 2021

Of O. laevatus : lectotype here designated (1 ♀, MNHN): Delagoa / MUSEUM PARIS, Collection H. d’Orbigny 1915 / TYPE / O. laevatus d’Orb. / LECTOTYPE, Onthophagus laevatus d’Orbigny, 1905 , Des. M. Rossini, 2021. Paralectotype (1 ♀, MNHN): Delagoa / green rectangular label / MUSEUM PARIS, Coll. H. d’Orbigny 1915 / PARALECTOTYPE, Onthophagus laevatus d’Orbigny, 1905 , Des. M. Rossini, 2021

Additional material examined. COLOMBIA: Antioquia: Santa Rosa de Osos , 21.III.1937 (2 ♂, 1 ♀ MZc) ; Medellin (3 ♂, 2 ♀ CEMT) ; La Ceja, V.1958 (1 ♂ CMNC) . Bolívar: El MantecO, 2.VIII.2006 (1 ♀ CEMT) . Boyacá: Cusiana Camijoque , 05°26’05’’N, 72°41’30’’W, 2200 m, VI.1997 (1 ♀ CMNC) . Cali: near Pichinde, 5000 feet, 18.VII.1970 (1 ♂, 3 ♀ CMNC) . Caquetá: Parque Nacional Natural Picachos , 02°47’51’’N, 74°51’18’’W, 1250 m, XI–XII.1997 (1 ♂ CMNC) . Cauca: Cauca (1 ♂, 1 ♀ SMTD) , 20 km W Silvia , 6000 feet, 22.II.1970 (10 ♂, 13 ♀ CMNC) ; Parque Nacional Puaracé, 2300 m, 1.V.1972 (1 ♂ CMNC) . Cundinamarca: Pacho (1 ♀ SMTD) ; Laguna Pedro Palo, W Bogotá — La Mesa, 1.X.1994 (2 ♂ CEMT) ; Fusagasugá, 8.V.1946 (1 ♂, 1 ♀ CMNC) ; Tequendama, 8000 feet, 6.VII.1970 (1 ♀ CMNC) . Meta: Villavicencio, IV.1907 (1 ♂ CMNC; 2 ♂, 3 ♀ SMTD) . Nariños: Pasto (2 ♂, 1 ♀ SMTD) . Norte de Santander: 3 km N Chinácota , 1000 feet, 10–12.V.1974 (36 ♂, 35 ♀ CMNC) ; 25 km S Chinácota , 2300 m, 10.V.1974 (1 ♂, 1 ♀ CMNC) ; 30 km S Cucuta , 700 m, 13.V.1974 (7 ♂, 5 ♀ CMNC) . Santander: Fusa , 6°26’21.26”N, 73°04’54.63”W, 17.III.1981 (1 ♂ CEMT) . Tolima: San Antonio (2 ♀ SMTD) ; Tolima (3 ♂, 4 ♀ SMTD) . Valle del Cauca: Restrepo, 6–7.VIII.1983 (1 ♂, 1 ♀ BGc) . Doubtful localities: Aguatal (2 ♂ SMTD) ; Villa Elvira, 6.VII.1908 (1 ♂ SMTD) . No data: (1 ♀ CMNC; 2 ♂, 2 ♀ NHRS) . ECUADOR: Azuay: Pucay, Bucay, 300 m, 2.II.1905 (1 ♀ NHRS) . Cañar: 5 km E Zhud , 3000 m, 2.III.1981 (1 ♂ CMNC) . Chimborazo: 2 km S Puela , 2315 m, 5.V.1998 (3 ♂, 1 ♀ CEMT) . Cotopaxi: 112 km W Latacunga, 14 km W Pilalo , 5100 feet, 21–31.VI.1976 (5 ♂, 5 ♀ CMNC) . El Oro: Huairapongo, 17.X.1905 (1 ♂, 1 ♀ NHRS) . Loja: 10 km N Zambi , 1850 m, 10.V.1998 (1 ♂, 1 ♀ CEMT) ; Gonzanamá , 2000 m, 28.VIII.1997 (1 ♀ CEMT) ; Las Chinchas, 28.XII.1996 (2 ♂, 1 ♀ CEMT) ; Calvario , 3.8500ºS, 79.3833ºW (1 ♀ NHRS) ; Villonaco , 2600 m, 03°59’40’’S, 79°15’37’’W, II.2012 (2 ♂ CEMT) ; Villonaco, 5.IX.1906 (1 ♂ CMNC) ; Loja (1 ♂ NHRS; 1 ♀ CNC) ; Pucará, 3.VIII.1905 (1 ♂ NHRS) ; Catamayo (1 ♂, 1 ♀ NHRS) . Pichincha: Quito-Santo Domingo Tandapi, 1500 m, 8.XI.1983 (1 ♂ MZc) . Tungurahua: Baños de Aguas Santa , 25.VII.1992 (2 ♂ CEMT) . ZamoraChinchipe: LojaZamora, 1800–2200 m, 24–25.VIII.1977 (14 ♂, 16, ♀ CMNC) ; 5 km NE Gonzanama Pass , 2740 m, 3.XI.1987 (1 ♂ CMNC) . PERU: Cajamarca: camino a Cumbemayo, 6–8.VI.2003, 3000 m (4 ♂, 1 ♀ CEMT) . Piura: Ayabaca, Pacaipampa, San Juan, 04°57’16.94’’S, 79°29’19.11’’W, 19.VI.2009 (2 ♂, 3 ♀ CEMT) . TRINIDAD AND TOBAGO: Saint George: 19 km N Arima, Lalaja Trace , 650 m, VI–VII.1993 (3 ♂, 1 ♀ CMNC) ; Tunapuna-Piarco, Maracas Valley , 10°43’43’’N, 61°24’62’’W, 6.VIII.2012 (1 ♂ OUMNH) . VENEZUELA: Aragua: Carretera Marcay-Choroni, 980 m, VIII.1940 (1 ♂ CMNC) ; Rancho Grande, VI.1963 (1 ♂, 2 ♀ CMNC) ; Rancho Grande, 1100 m (1 ♂ OUMNH) ; Rancho Grande, Biological Station Pico Periquitos, 10°21’N, 67°41’W, 1100 m, II–III.1995 (1 ♂, 1 ♀ CMNC) ; Tiara, 50 km SW Caracas , 1000–2000 m, 22–25.II.1971 (16 ♂, 20 ♀ CMNC) . Falcón: Cueva Coy Coy de Uria (1 ♂ CMNC) ; Curimagua, 5.VIII.1956 (1 ♂ MSNG) . Mérida: La Muchuy, 2200 m, 2.VIII.1995 (5 ♂, 5 ♀ MSNG) ; La Muchuy, Tabay, 1900–2300 m, VI–VIII (3 ♂, 5 ♀ CMNC; 7 ♂, 1 ♀ MZc) ; La Muchuy, National Park Sierra Nevada, 2400 m, 13.IV.1995 (1 ♀ CEMT) ; El Joque, Jají, 2000 m, 1–15.III.1978 (1 ♀ MZc) ; Santo Domingo, 2200 m, 5.I.1965 (1 ♂, 1 ♀ MZc) . Cacute, 2100 m, 17–24.IV.1975 (2 ♂ MZc) ; Mariño, 1700 m, 1.XII.1984 (2 ♂ MZc) ; Carbonera, 17.IX.1985 (1 ♂ MZc) ; La Trampa, 1700 m, 10.III.1978 (2 ♂, 1 ♀ CMNC) . Carbonera, Lagunilla, 10.III.1978 (1 ♂ MZc) . Mérida, 1700 m, VI.1987 (1 ♂ CMNC) ; Sucre Jají , 08°39’39.05’’N, 71°23’55.85’’W, 2235 m, 8.VII.2009 (1 ♂, 1f CEMT) ; Rio Chama , Tabay, VIII.1942, 1800 m (1 ♀ CMNC) ; La Mucuy (2 ♂ CMNC) ; Timotes, 2200 m, 25.I.1968 (2 ♂, 1 ♀ CMNC; 2 ♂ MZc) ; La Cutata, 4000 m, V.1996 (1 ♀ SMTD) . No locality (3 ♂, 6 ♀ SMTD) . Miranda: Guatopo National Park , 50 km SE Caracas, 400 m, 5–6.III.1971 (1 ♀ OUMNH) . Monagas: Teresen, 800 m, 12.IV.1963 (3 ♂ CMNC) ; Caripe, Cueva Guacharo, 700 m, 20–30.VII.1987 (1 ♀ CMNC) . Yaracuy: Bolivar Aroa , 10°23’06.1’’N, 68°58’45.45’’W, 1415 m, 19.VII.2009 (1 ♂ CEMT) . Táchira: Pregonero, La Honda, 1100 m, 13.II.1989 (1 ♀ CEMT) ; 30 km NE San Cristóbal , 2000 m, 20–22.V.1974 (6 ♂, 12 ♀ CMNC; 1 ♂ OUMNH) ; San Cristobal , 1200 m, 10–18.VIII.1983 (1 ♂ BGc) . Sucre: Elvecia near Mt. Turumquire , 14.I.1930 (1 ♂ CMNC) . Trujillo: Trujillo (1 ♀ CEMT) ; Boconó, 2000–2300 m, V–VI.1981 (5 ♂♂, 2 ♀♀ MSNG) . Vargas: Petaquire, El Junquito, 7.III.1965 (1 ♂ MZc) . Zulia: Miranda, 28-30.XI.2005 (1 ♂ CEMT) . Miranda, 28 km N Guatopo Nacio- nal Park, El Lucero, 700 m, 14.VI–5.VIII.1987 (2 ♂, 3 ♀ CMNC) ; Altagracia, Guatopo National Park , 50 km SE Caracas, 5–6.III.1971 (4 ♂, 1 ♀ CMNC) ; Guatopo National Park , 6–8.III.1988 (1 ♂, 4 ♀ CMNC) . No data: (1 ♂ MCZ; 1 ♂ NHRS) ; (2 ♂, 3 ♀ MCZ: “Mexico” erroneous locality) .

Figures

FIGURE 2. Onthophagus curvicornis. A, Dorsal habitus of the male; B, dorsal habitus of female; C, lateral habitus of the male; D, dorsal habitus of the lectotype of O. laevatus; E, original labels of the lectotype of O. laevatus; F, lateral view of the parameres; G, frontalview of the parameres; H, endophallites. LC: lamella copulatrix; A: axial; SA: subaxial; SRP: superior right peripheral; FLP: fronto-lateral peripheral.