The dung beetles of Venezuela (Coleoptera: Scarabaeidae: Scarabaeinae): catalogue and updated distribution

1 European Journal of Taxonomy 959: 1–272 ISSN 2118-9773 https://doi.org/10.5852/ejt.2024.959.2677 www.europeanjournaloftaxonomy.eu 2024 · Lozano de la Rosa C. et al. This work is licensed under a Creative Commons Attribution License (CC BY 4.0). Monograph urn:lsid:zoobank.org:pub:30872C13-516F-42FA-AFA7-30ADC6BF1BAF The dung beetles of Venezuela (Coleoptera: Scarabaeidae: Scarabaeinae): catalogue and updated distribution Cecilia LOZANO DE LA ROSA 1,*, Mario CUPELLO 2 & Fernando Z. VAZ-DE-MELLO 3 ¹ Programa de Pós-graduação em Ecologia e Conservação da Biodiversidade, Universidade Federal de Mato Grosso, Cuiabá, Mato Grosso, Brazil. 1 Instituto Venezolano de Investigaciones Científicas, Centro de Estudios Botánicos y Agroforestales, Laboratorio de Ecología Espacial, Maracaibo, estado Zulia, Venezuela. 2 Laboratório de Sistemática e Bioecologia de Coleoptera, Departamento de Zoologia, Universidade Federal do Paraná, Curitiba, Paraná, Brazil. Currently at Department of Entomology, Texas A&M University, College Station, TX, 77843, USA. 3 Departamento de Biologia e Zoologia, Instituto de Biociências, Universidade Federal de Mato Grosso, Cuiabá, Mato Grosso, Brazil. * Corresponding author: [email protected] 2 Email: [email protected] 3 Email: [email protected] 1 urn:lsid:zoobank.org:author:36CFEA67-71C5-4F3C-97F8-B57E43C811CC 2 urn:lsid:zoobank.org:author:BDB03C18-7095-4EAA-8BDD-03CB4F79676D 3 urn:lsid:zoobank.org:author:2BC093C2-3E2B-466B-B31C-9F3D9FA844BC Abstract. Venezuela’s diverse land ecosystems are grouped into four major regions (coast-islands, low plains, hills and mountains), ranging from sea level up to 4978 m. The Scarabaeinae (Scarabaeidae, Coleoptera) currently encompass 278 genera and 6837 species worldwide, but are poorly inventoried in Venezuela. We reviewed the literature and the material housed at the entomological collection of the Universidade Federal de Mato Grosso (CEMT), Cuiabá, Brazil, and found 32 genera and 149 species of dung beetles as certainly present in Venezuela. Twenty-four of these species are, as far as current knowledge goes, endemic to the country, while another 34 are restricted to Venezuela and the neighbouring countries of Colombia, Brazil, Trinidad and Tobago, and Guyana. Additionally, 36 species are deemed potential inhabitants of the country, whilst 14 others previously recorded in the literature as part of the Venezuelan fauna are here concluded not to be actually present there. Complete literature is listed for each genus and species, and information on type material, material examined, worldwide distribution, and Venezuelan records is also presented. Keywords. Scarabaeoidea, Neotropics, South America, endemic species, type specimen, type locality. Lozano de la Rosa C., Cupello M. & Vaz-de-Mello F.Z. 2024. The dung beetles of Venezuela (Coleoptera: Scarabaeidae: Scarabaeinae): catalogue and updated distribution. European Journal of Taxonomy 959: 1–272. https://doi.org/10.5852/ejt.2024.959.2677 European Journal of Taxonomy 959: 1–272 (2024) 2 Introduction The Coleoptera Linnaeus, 1758 subfamily Scarabaeinae Latreille, 1802 comprises 278 genera and 6837 species worldwide (Schoolmeesters 2023). Dung beetles, as they are popularly known, are used as a bioindicator group whose highest diversity is found in tropical regions and whose ecosystem functions play a key role in environmental balance (Halffter & Matthews 1966; Hanski & Cambefort 1991; Stokstad 2004; Nichols et al. 2008; Nichols & Gardner 2011; Philips 2011; Ridsdill-Smith & Edwards 2011; Noriega et al. 2021a, 2021b). Catalogues and checklists are invaluable sources of bibliographic information (Papavero 1994) and provide insight into species diversity at both local and global scales. Since 1869, many checklists have been published for scarab beetles at a worldwide level (e.g., Harold 1869d; Gillet 1911b; Gillet & Boucomont 1927; Blackwelder 1944; Krajcik 2012), but in recent decades this type of work has progressively shifted its focus in Central and South America to country-level studies. Among these are the checklists of Vaz-de-Mello (2000, 2022) for Brazil, Escobar (2000) and Medina et al. (2000, 2001) for Colombia, Ratcliffe (2002) for Panama, Morón (2003) for Mexico, Hamel-Leigue et al. (2006) for Bolivia, Carvajal et al. (2011) and Chamorro et al. (2018, 2019) for Ecuador, Solís & Kohlmann (2012) for Costa Rica, Ratcliffe et al. (2015) for Peru, Hielkema & Hielkema (2019) for the Guianas, Maes et al. (2020) for Nicaragua, Pablo-Cea et al. (2023) for El Salvador, and Mondaca (2023) for Chile. For Venezuela, the first reports of dung beetles date back to Harold (1869d), who recorded four genera and five species for the country. Candèze (1891) subsequently presented a checklist from Venezuela entitled “Voyage de M.E. Simon au Venezuela (Décembre 1887–Avril 1888)”, in which he recorded eight genera and fourteen species of dung beetles, among other beetle families. Later, Roze (1955), in a checklist of the Scarabaeinae fauna of the country, reported 15 genera and 72 species based on the holdings of Venezuelan museums. Finally, in a study focused on species monitoring, Ferrer-Paris et al. (2013) recorded 23 genera and 115 species from Venezuela, including 54 morphospecies not identified to the species level. The first and only regional catalogue for the Venezuelan dung beetles was compiled by Blanco (1987, 1988) for the species occurring in the state of Táchira. Available knowledge on Venezuelan dung beetles is patchy and focused on some particular regions of the country, especially the Andes and areas crossed by the Orinoco River. As for the conservation status of the species, six of them, namely Ateuchus ambiguus Martínez & Martínez, 1990, Eurysternus impressicollis Castelnau, 1840, Canthonella gomezi (Halffter & Martínez, 1968), Cryptocanthon nebulinus Howden, 1973, Cryptocanthon punctatus Cook, 2002 and Sulcophanaeus auricollis (Harold, 1880), were included in the latest edition of the Red Book of the Fauna of Venezuela (Rodríguez et al. 2015). However, this number may be underestimated due to the limited knowledge of the fauna. Given the importance of species catalogues for the study of biodiversity and its conservation, here, we present an updated catalogue of the dung beetles (Scarabaeinae) found in Venezuela, including a distribution map for each species inhabiting the country. Material and methods Venezuela is located in northern South America (CIA 2020) and is administratively divided into 23 states, a Capital District (Caracas and metropolitan area) and Federal Dependencies (72 islands) (Fig. 1). Its ecosystem diversity is classified into four major ecoregions ‒ coastal-islands, low plains, hills and mountains (Huber & Oliveira-Miranda 2010) ‒ encompassing a wide variety of landscapes (Fig. 2), with elevation ranging from 0 to 4978 m above sea level (Pérez et al. 2005). Dung beetle records from Venezuela were compiled through an exhaustive review of the literature, covering both taxonomic and ecological studies. These records were then categorized into three groups: those reliable (i.e., the species is confirmed to occur in Venezuela), dubious (the species may occur in LOZANO DE LA ROSA C. et al., Dung beetles (Scarabaeinae) of Venezuela 3 Venezuela, but this has yet to be confirmed), and erroneous (the species is certainly not present in the country and the record was usually based on misidentified material). In the section on each species for which the Venezuelan records were classed as dubious or erroneous, we present the rationale for our decision to categorize them as such. The literature review was supplemented by the examination of material housed in the collection of the institution where the first and third authors reside, the Federal University of Mato Grosso (CEMT), Cuiabá, Brazil. This enabled us to make our own new species Fig. 1. Macrolocation of Venezuela on the South American continent and its states and federal dependencies. European Journal of Taxonomy 959: 1–272 (2024) 4 Fig. 2. Landscape units of Venezuela (from Huber & Oliveira-Miranda 2010). LOZANO DE LA ROSA C. et al., Dung beetles (Scarabaeinae) of Venezuela 5 records for the country as well as to assess records that, based solely on the information provided in the literature, would be dubious. We encourage colleagues to complement our work by examining further collections holding rich Venezuelan material. These include those of the Canadian Museum of Nature, Ottawa, Canada, the Museu de Zoologia, Universidade de São Paulo, São Paulo, Brazil, and other museums in Europe and Venezuela itself. We expect that the list of Venezuelan dung beetles will increase once these collections have been thoroughly reviewed. Given the current taxonomic instability at the tribal, subtribal, and subgeneric levels, our catalogue ignores them and organizes the genera in alphabetical order without consideration of their tribal placement, and the species within each genus are also arranged alphabetically without consideration of the subgenera. Those interested in a classification of the New World fauna including tribes, subtribes, and subgenera should refer to Cupello et al. (2023b) and relevant works published afterwards (Daniel & Davis 2023; Bouchard et al. 2024; Génier & Darling 2024). Global distribution data for each species were retrieved from the literature, particularly from taxonomic revisions, as well as from the examination of the material housed in CEMT. The catalogue presents a synonymic list for each species and specifies the collection where the type specimens are housed, informing whether or not we have examined them. For type material housed in CEMT, label contents are transcribed verbatim, using a forward slash (“/”) to separate information between the lines of each label. Non-type material is cited in the following order: country (given in capital letters), state, number of specimens examined, specific locality, geographic coordinates, elevation, date, collector(s), additional data such as collecting method, and repository abbreviation. Corrections for state, locality, and geographic coordinates are enclosed in square brackets, “[ ]”. The distribution of the species in Venezuela is given according to the subregions of the country as delimited by Huber & Oliveira-Miranda (2010). Distribution maps were made using Quantum GIS (QGis ver. 3.16.8 – Hannover; QGIS Development Team 2020) and the R software for the final figure. Records plotted on the maps are those we found on specimen labels and in the literature and which we deem reliable. For localities cited in both sources without coordinates, these were obtained from Geonames (http://www.geonames.org/). The following collections are mentioned in the catalogue (curator or person responsible, when known, in parentheses): BDGC = Bruce D. Gill personal collection, Woodlawn, Ontario, Canada BMNH = The Natural History Museum, London, UK (Maxwell V.L. Barclay) CEMT = Coleção Entomológica de Mato Grosso Eurides Furtado, Universidade Federal de Mato Grosso, Cuiabá, Brazil (Fernando Z. Vaz-de-Mello) CMNC = Canadian Museum of Nature, Ottawa, Canada (François Génier) CPFA = Patrick and Florent Arnaud personal collection, Saintry-sur-Seine, France CUIC = Cornell University Insect Collection, Ithaca, NY, USA (Jason J. Dombroskie) FGIC = François Génier personal collection, Gatineau, Canada HAHC = Henry & Anne Howden collection, Ottawa, Canada (now housed in the CMNC) MACN = Museo Argentino de Ciencias Naturales “Bernardino Rivadavia”, Buenos Aires, Argentina (Pablo R. Mulieri) MAMU = MacLeay Collections, Chau Chak Wing Museum, University of Sydney, Sydney, Australia MCNZ = Museu de Ciências Naturais da Fundação Zoobotânica do Rio Grande do Sul, Porto Alegre, Brazil (Luciano de Azevedo Moura) MCZC = Museum of Comparative Zoology, Harvard University, Cambridge, MA, USA (Crystal Maier and Brian Farrell) MFNB = Museum für Naturkunde Berlin, Leibniz-Institut für Evolutionsund Biodiversitätsforschung, Berlin, Germany (Joachim Willers, Bernd Jaeger, and Johannes Frisch) European Journal of Taxonomy 959: 1–272 (2024) 6 MIZA = Museo del Instituto de Zoología Agrícola, Universidad Central de Venezuela, Maracay, Venezuela (Luis J. Joly) MNHN = Muséum national d’histoire naturelle, Paris, France (Olivier Montreuil and Antoine Mantilleri) MQCAZ = Museo de Zoología de la Pontificia Universidad Católica, Quito, Ecuador (Álvaro Barragán, Carlos Capio and Fernanda Salazar) MVMA = Museums Victoria, Melbourne, Australia MXAL = Miguel Ángel Morón personal collection, Xalapa, México MZSP = Museu de Zoologia, Universidade de São Paulo, São Paulo, Brazil (Sonia Casari and Carlos Campaner) MZUF = Sezione di Zoologica “La Specola”, Museo di Storia Naturale dell’Università degli Studi di Firenze, Florence, Italy (Luca Bartolozzi) NHRS = Naturhistoriska Riksmuseet, Stockholm, Sweden (Johannes Bergsten) NMPC = National Museum (Natural History), Prague, Czech Republic (Jiří Hájek) OUMNH = Hope Entomological Collections, Oxford University Museum of Natural History, Oxford, UK (Darren Mann) SDEI = Senckenberg Deutsches Entomologisches Institut, Müncheberg, Germany (Vinicius Ferreira) SMTD = Museum für Tierkunde, Senckenberg Naturhistorische Sammlungen Dresden, Dresden, Germany (Olaf Jäger and Klaus-Dieter Klass) USNM = Smithsonian National Museum of Natural History, Washington D.C., USA ZMAS = Zoological Institute, Russian Academy of Sciences, Saint Petersburg, Russia (Andrey V. Frolov) ZMUK = Zoologisches Museum, Universität Kiel, Kiel, Germany (Michael Kuhlmann) ZSM = Zoologische Staatssammlung München, Munich, Germany (Michael Balke and Ditta A. Balke) Results We record 32 genera and 149 species of Scarabaeine dung beetles as certainly present in Venezuela. Twenty-four of the species are endemic to the country and another 34 are restricted to Venezuela and the neighbouring countries of Colombia, Brazil, Trinidad and Tobago, and Guyana. Additionally, 36 species are deemed to be potentially present in Venezuela: all of them have Venezuelan records available in the literature, their presence in the country seems to be at least plausible based on the rest of the known range of the species (e.g., known to occur in neighbouring areas of Colombia, Brazil, or the Guianas), but we ourselves failed to positively confirm the presence in the country (usually because we have not seen any specimens from there). Finally, 14 other species are considered to have been erroneously recorded from the country in the literature, in most cases based on misidentified material. Updated catalogue of the Scarabaeinae (Coleoptera: Scarabaeinae) from Venezuela Class Insecta Linnaeus, 1758 Order Coleoptera Linnaeus, 1758 Family Scarabaeidae Latreille, 1802 Subfamily Scarabaeinae Latreille, 1802 I – Confirmed records Genus Agamopus Bates, 1887 Agamopus Bates, 1887: 42 (original description). Type species: Agamopus lampros Bates, 1887, by original monotypy. LOZANO DE LA ROSA C. et al., Dung beetles (Scarabaeinae) of Venezuela 7 Agamopus – Gillet 1911b: 49 (catalogue, cited for Panama). — Blackwelder 1944: 203 (catalogue). — Pereira 1947a: 1–5 (contribution); 1954a: 56 (key). — Vulcano & Pereira 1967: 577 (key). — Halffter & Martínez 1968: 210 (revision); 1977: 34, 43 (key). — Howden & Young 1981: 40 (diagnosis). — Medina & Lopera-Toro 2000: 301, 306 (key). — Medina et al. 2001: 133 (checklist for Colombia). — Ratcliffe 2002: 32 (checklist for Panama). — Halffter 2003: 24 (redescription). — Vaz-de-Mello et al. 2011a: 4, 8 (key). — Solís & Kohlmann 2012: 2, 4 (checklist for Costa Rica). — Boilly & Vaz-de-Mello 2013: 106 (key for French Guiana). — Cupello & Vaz-de-Mello 2018: 17 (nomenclator). — Hielkema & Hielkema 2019: 58 (catalogue for the Guianas). Agamopus lampros Bates, 1887 Fig. 3A Agamopus lampros Bates, 1887: 42 (original description). Type locality: Panama: Chiriquí: David. Name-bearing type: lectotype (BMNH), designated by Costa-Silva et al. (2022); examined by FZVM. Agamopus lampros – Gillet 1911b: 49 (catalogue, cited for Panama). — Boucomont 1928a: 187 (cited). — Balthasar 1938: 218 (comments). — Blackwelder 1944: 203 (catalogue); 1973: 6 (catalogue). — Pereira 1947a: 4 (key). — Halffter & Martínez 1968: 236 (comments, female description). — Howden & Young 1981: 41 (contributions, distribution). — Morón et al. 1988: 320 (record from Mexico). — Thomas 1993: 395 (record from Mexico). — Schoolmeesters 1996: 20 (record from Nicaragua). — Medina & Lopera-Toro 2000: 310 (cited as “lamprus”, fig. 8e). — Navarrete-Heredia et al. 2001: 52 (list). — Medina et al. 2001: 135 (checklist for Colombia). — Ratcliffe 2002: 13 (checklist for Panama). — Halffter 2003: 24 (redescription). — Kohlmann & Wilkinson 2003: 221–222 (cited). — Kohlmann et al. 2007: 27 (atlas). — Vaz-de-Mello et al. 2011a: 49, figs 1–2 (key). — Solís & Kohlmann 2012: 4 (checklist for Costa Rica). — Arellano et al. 2013: 655 (citation). — Noriega et al. 2013: 470 (checklist from Colombian Caribbean). — Cupello & Vaz-de-Mello 2018: 17 (nomenclator). — Sánchez-Hernández et al. 2020: 226 (list for Mexico). — Costa-Silva et al. 2022: 73–75, 83 (revision, designation of lectotype, key). Material examined VENEZUELA ‒ Guárico • 4 specs; Altagracia de Orituco, NM 08-T-044; 23 Aug. 2006; curso NM2006 leg.; faeces, 09:49, 41h; CEMT. ‒ Táchira • 4 specs; Libertador, San Joaquín de Navay; 7.7741° N, 71.6675° W; 550 m a.s.l.; Aug. 2006; T. Good leg.; CEMT • 2 specs; Libertador, San Joaquín de Navay; 7.6622° N, 71.7104° W; 200 m a.s.l.; Aug. 2006; T. Good leg.; CEMT. Distribution Mexico, Guatemala, El Salvador, Nicaragua, Costa Rica, Panama, Colombia and Venezuela. Subregions of Venezuela Andes mountains and Central Coast Mountain Range. Literature record Costa-Silva et al. 2022: 73–75, 83 (Venezuela: Guárico and Táchira). Genus Anomiopus Westwood, 1842 Anomiopus Westwood, 1842: 59 (original description). Type species: Anomiopus virescens Westwood, 1842, by subsequent designation of Canhedo (2006). European Journal of Taxonomy 959: 1–272 (2024) 8 Fig. 3. Species distribution. A. Agamopus lampros Bates, 1887. B. Anomiopus alexandrei Canhedo, 2006. Green diamond = CEMT collection data; black circle = literature data. LOZANO DE LA ROSA C. et al., Dung beetles (Scarabaeinae) of Venezuela 9 Onthocharis Erichson, 1847b: 189 (original description). Type species: Scatonomus smaragdinus Westwood, 1842, by subsequent designation of Lacordaire (1855). Hypocanthidium Balthasar, 1938: 214 (original description). Type species: Hypocanthidium globulum Balthasar, 1938, by original monotypy. Anomiopus – Erichson 1843: 189 (list); 1847b: 761 (cited as synonym of Onthocharis). — Westwood 1843: 62 (redescription); 1847: 231 (redescription). — Lacordaire 1855: 94 (cited as synonym of Onthocharis, see footnote 2). — Harold 1869d: 1002 (catalogue, synonym of Onthocharis). — Gillet 1911a: 50 (catalogue, synonym of Onthocharis). — Lucas 1920: 100 (catalogue, synonym of Onthocharis). — Blackwelder 1944: 204 (cited as synonym of Onthocharis). — Medina & Lopera, 2000: 306 (key). — Vaz-de-Mello 2000: 190 (checklist for Brazil). — Medina et al. 2001: 137 (list). — Canhedo 2006: 354 (revision). — Hamel-Leigue et al. 2006: 12 (list). — Vaz-de-Mello et al. 2011a: 25 (key). — Carvajal et al. 2011: 124, 318 (description, list). — Krajcik 2012: 28 (list). — Solís & Kohlmann 2012: 2 (checklist for Costa Rica). — Boilly & Vaz-de-Mello 2013: 107 (key). — Chamorro et al. 2018: 75 (checklist for Ecuador); 2019: 9–10 (catalogue). — Hielkema & Hielkema 2019: 58 (catalogue for the Guianas). Hypocanthidium – Pereira 1954a: 56 (key). — Halffter & Mathews 1966: 256 (catalogue, distribution). — Vaz-de-Mello et al. 2011a: 3 (junior synonym of Anomiopus). — Solís & Kohlmann 2012: 2 (synonym of Anomiopus). Anomiopus alexandrei Canhedo, 2006 Fig. 3B Anomiopus alexandrei Canhedo, 2006: 434 (original description). Type locality: Brazil: Roraima: Ilha de Maracá. Name-bearing type: holotype (MCNZ) (Canhedo 2006), not examined. Anomiopus alexandrei – Canhedo 2004a: 189 (key; nomen nudum). — Hielkema & Hielkema 2019: 58 (catalogue for the Guianas). Type material examined Paratype BRAZIL • 1 ♂; “Roraima: Ilha de Maracá-RR/ — .IX.1996 / Ribeiro & / Vaz-de-Mello / leg. / Anomiopus / alexandrei sp. n. / PARÁTIPO ♂ / V.L Canhedo det. 1999 / CEMT / CUIABÁ / 00078205”; CEMT. Distribution Venezuela and Brazil (Canhedo 2006). Subregions of Venezuela Peneplain of the Caura and Paragua rivers, System of hills and low piedmont mountains of the Guiana Shield, and Guiana Shield. Literature records Canhedo 2004a: 189 (Venezuela: Bolívar); 2006: 349, 434 (Venezuela: Bolívar: Río Caura). Anomiopus edmondsi Canhedo, 2006 Fig. 4A Anomiopus edmondsi Canhedo, 2006: 431 (original description). Type locality: Venezuela: Bolívar: Río Caura, Puerto Cabello. Name-bearing type: holotype (BDGC) (Canhedo 2006), not examined. European Journal of Taxonomy 959: 1–272 (2024) 16 Subregions of Venezuela Peneplain of the Caura and Paragua rivers, and Guiana Shield. Literature records Canhedo 2004a: 188 (Venezuela: Bolívar); 2006: 349–400 (Venezuela: Bolívar). Genus Ateuchus Weber, 1801 Ateuchus Weber, 1801: 10 (original description). Type species: Ateuchus histeroides Weber, 1801, by original monotypy. Choeridium Le Peletier de Saint-Fargeau & Audinet-Serville, 1828: 356 [cited under two alternative spellings: Choeridium and Chaeridium, the former the correct original spelling (Cupello et al. 2021a)] (original description). Type species: Choeridium simplex Le Peletier de Saint-Fargeau & Audinet-Serville, 1828, by subsequent designation of Chevrolat (1843). Ateuchus – Fabricius 1801: 54 (redescription). — Latreille 1829: 532 (redescription). — Castelnau 1840: 63 (redescription). — Reiché 1841: 212 (key). — Lacordaire 1855: 66 (redescription). — Lucas 1920: 125 (catalogue, distribution). — Chapin 1946: 79 (synonym of Ateuchus Weber). — Pereira 1954b: 56 (key). — Roze 1955: 43 (checklist for Venezuela). — Martínez 1959: 76 (catalogue for Argentina). — Halffter & Mathews 1966: 256 (catalogue, distribution). — Vulcano & Pereira 1967: 577 (key for the Amazon, cited as Atheuchus Weber, 1801). — Howden & Young 1981: 13, 68 (key, redescription). — Halffter & Edmonds 1982: 137 (catalogue, distribution). — Kohlmann 1984: 25 (redescription); 1997: 178 (redescription). — Medina & Lopera-Toro 2000: 306 (illustrated key). — Vaz-de-Mello 2000: 190 (checklist for Brazil). — Medina et al. 2001: 137 (checklist for Colombia). — Ratcliffe et al. 2002: 49 (key). — Ratcliffe 2002: 13 (checklist for Panama). — Morón 2003: 52 (list). — Hamel-Leigue et al. 2006: 12 (list for Bolivia). — Vaz-de-Mello et al. 2011a: 4, 13, 21, 28, 36, 43–44 (key). — Carvajal et al. 2011: 126, 318 (description, list for Ecuador). — Krajcik 2012: 49 (list). — Solís & Kohlmann 2012: 4 (checklist for Costa Rica). — Boilly & Vazde-Mello 2013: 108 (key). — Chamorro et al. 2018: 77 (list for Ecuador); 2019: 9, 13 (catalogue). — Hielkema & Hielkema 2019: 40 (catalogue for the Guianas). Choeridium – Castelnau 1840: 83 (redescription). — Lacordaire 1855: 93 (redescription, synonym of Ateuchus). — Harold 1867a: 9 (key); 1868b: 32 (redescription); 1868c: 55 (key); 1869d: 1006 (catalogue). — Gillet 1911a: 52 (catalogue). — Lucas 1920: 182 (catalogue, distribution). — Dawson 1922: 61 (key). — Luederwaldt 1929: 11 (key); 1931a: 369 (key). — Paulian 1938: 234 (key). — Balthasar 1939c: 44 (comment). — Pessôa & Lane 1941: 437 (key). — Blackwelder 1944: 204 (list). Ateuchus aeneomicans (Harold, 1868) Fig. 7B Choeridium aeneomicans Harold, 1868d: 82 (original description). Type locality: Brazil: Amazonas: São Paulo de Olivença. Name-bearing type: syntypes (MNHN), examined by MC and FZVM. Choeridium aeneomicans – Harold 1868b: 37, 66 (key, redescription); 1869d: 1006 (catalogue, cited for the Amazon). — Gillet 1911b: 52 (catalogue, cited for Amazonas and Panama). — Balthasar 1939c: 63 (key, cited for Brazil and Panama). — Blackwelder 1944: 204 (list, cited for Panama and Brazil). Choeridium æneo-micans – Bates 1887: 46‒47. Ateuchus aeneomicans – Vulcano & Pereira 1967: 589 (key). — Howden & Young 1981: 68–69 (contributions). — Kohlmann 1997: 178–179 (key, redescription). — Feer 2000: 31 (checklist for French Guiana). — Vaz-de-Mello 2000: 190 (checklist for Brazil). — Medina et al. 2001: 137 (checklist for Colombia). — Ratcliffe 2002: 13 (checklist for Panama). — Kohlmann et al. 2007: 27 (atlas). — Arias-Buriticá et al. 2011: 875 (cited for Chocó, Colombia). — Krajcik 2012: 49 (list). — LOZANO DE LA ROSA C. et al., Dung beetles (Scarabaeinae) of Venezuela 17 Fig. 7. Species distribution. A. Anomiopus smaragdinus (Westwood, 1842). B. Ateuchus aeneomicans (Harold, 1868). Green diamond = CEMT collection data; black circle = literature data. European Journal of Taxonomy 959: 1–272 (2024) 18 Solís & Kohlmann 2012: 4 (checklist for Costa Rica). — Ratcliffe et al. 2015: 196 (checklist for Peru). — Chamorro et al. 2018: 91 (list for Ecuador); 2019: 14 (catalogue). — Hielkema & Hielkema 2019: 44 (catalogue for the Guianas). — Nieto et al. 2020: 136 (report). Material examined VENEZUELA ‒ Apure • 1 ♂; Muñoz, Mantecal; 1 Jun. 1993; Hornburg leg.; CEMT. ‒ Portuguesa • 2 ♀♀; Guanare; Oct. 1985; no collector; CEMT • 1 ♂; Guanare; 15 Oct. 1985; no collector; CEMT. ‒ Táchira • 1 ♀; Libertador, San Joaquín de Navay; 07°47′59″ N, 71°40′24.24″ W; 655 m a.s.l.; 26 Aug. 2006; T. Good et al. leg.; human faeces; CEMT. Distribution Costa Rica, Panama, Colombia, Venezuela (new country record), Brazil, Ecuador, Peru and Bolivia (MC, pers. obs.). Subregions of Venezuela Plains, and Andes mountains. Literature record No previous reports for Venezuela. Ateuchus alvarezi (Martínez & Halffter, 1986) Fig. 8A Canthidium alvarezi Martínez & Halffter, 1986: 21 (original description). Type locality: Brazil: Roraima: Pacaraima: Surumu: Serra do Marari. Name-bearing type: holotype (MACN), confirmed to be there by MC in 2014. Ateuchus alvarezi – Génier & Cupello 2018: 2 (new combination). — Hielkema & Hielkema 2019: 40 (catalogue for the Guianas). Material examined from Venezuela in CEMT Paratype VENEZUELA • 1 ♂; “PARATIPO”; “Canthidium / superbum / sp. nov / A. Martínez-DET. 1982”/; “oct: 974 / VENEZUELA / Bolívar / Canaima / Carcavallo leg. / coll. Martínez / ♂”; CEMT-00078192. Distribution Venezuela, Guyana, and Brazil (Génier & Cupello 2018). Subregion of Venezuela Guiana Shield. Literature records Martínez & Halffter 1986: 21 (Venezuela: Bolívar: Canaima). — Génier & Cupello 2018: 2 (Venezuela). Ateuchus ambiguus Martínez & Martínez, 1990 Fig. 8B Ateuchus ambiguus Martínez & Martínez, 1990: 161 (original description). Type locality: Venezuela: Mérida: San Rafael del Páramo de Mucuchíes, Laguna Grande, 3800 m. Name-bearing type: holotype (MACN), confirmed to be there by MC in 2014. Ateuchus ambiguus – Krajcik 2012: 49 (list). LOZANO DE LA ROSA C. et al., Dung beetles (Scarabaeinae) of Venezuela 19 Fig. 8. Species distribution. A. Ateuchus alvarezi (Martínez & Halffter, 1986). B. Ateuchus ambiguus Martínez & Martínez, 1990. Green diamond = CEMT collection data; black circle = literature data. European Journal of Taxonomy 959: 1–272 (2024) 20 Distribution Venezuela (endemic) (Martínez & Martínez 1990). Subregion of Venezuela Andes mountains. Literature records Martínez & Martínez 1990: 161 (Venezuela: Mérida: Páramo de Mucuchíes, Laguna Grande). — Krajcik 2012: 49 (Venezuela: Mérida). Ateuchus bordoni Martínez & Martínez, 1990 Fig. 9A Ateuchus bordoni Martínez & Martínez, 1990: 159 (original description). Type locality: Venezuela: Mérida: Pico El Águila. Name-bearing type: holotype (MACN), confirmed to be there by MC in 2014. Ateuchus bordoni – Krajcik 2012: 49 (list). Distribution Venezuela (endemic) (Martínez & Martínez 1990). Subregion of Venezuela Andes mountains. Literature records Martínez & Martínez 1990: 159 (Venezuela: Mérida: Pico El Águila). — Krajcik 2012: 49 (Venezuela: Mérida). Ateuchus calcaratus (Harold, 1868) Fig. 9B Choeridium calcaratum Harold, 1868b: 76 (original description). Type locality: originally said to be French Guiana, but this is incorrect (MC, pers. obs.). Name-bearing type: syntypes, ambiguous status (see Cupello 2020 for details). Choeridium calcaratum – Harold 1869d: 1007 (catalogue, cited for Brazil). — Gillet 1911b: 52 (catalogue, cited for Brazil). — Blackwelder 1944: 204 (list, cited for Brazil). Ateuchus calcaratus – Vulcano et al. 1976: 522 (cited for French Guiana and Panama). — Vaz-deMello 2000: 190 (checklist for Brazil). — Ratcliffe 2002: 13 (checklist for Panama). — Hielkema & Hielkema 2019: 44 (catalogue for the Guianas). Material examined VENEZUELA ‒ Bolívar • 1 ♂, 1 ♀; Caroní, Ciudad Guayana; Jan. 1996; Petr Pokorný leg.; CEMT • 2 ♀♀; El Manteco; 2 Aug. 2006; no collector; in dung; CEMT. Distribution Venezuela (new country record); the French Guiana record is incorrect (MC, pers. obs.). LOZANO DE LA ROSA C. et al., Dung beetles (Scarabaeinae) of Venezuela 21 Fig. 9. Species distribution. A. Ateuchus bordoni Martínez & Martínez, 1990. B. Ateuchus calcaratus (Harold, 1868). Green diamond = CEMT collection data; black circle = literature data. European Journal of Taxonomy 959: 1–272 (2024) 22 Subregion of Venezuela System of hills and low piedmont mountains of the Guiana Shield. Literature record No previous reports for Venezuela. Ateuchus columbianus (Harold, 1868) Fig. 10A Choeridium columbianum Harold, 1868b: 58 (original description). Type locality: Colombia. Namebearing type: syntypes lost (MC, pers. obs.). Choeridium columbianus – Harold 1869d: 1007 (catalogue, cited for Colombia). — Gillet 1911b: 52 (catalogue, cited for Colombia). — Balthasar 1939c: 62 (key, cited for Colombia). — Blackwelder 1944: 204 (list, cited for Colombia). Ateuchus columbianus – Vulcano & Pereira 1967: 590 (key). — Medina et al. 2001: 137 (checklist for Colombia). — Ratcliffe et al. 2015: 196 (checklist for Peru). Material examined VENEZUELA ‒ Barinas • 5 ♂♂, 1 ♀; Bolívar, Barinitas (“BARINITAS ENV.”); Jan. 1996; Petr. Pokorny leg.; ex A. Dostal collection; CEMT. ‒ Mérida • 1 ♂, 1 ♀; “Finca Campo Alegre”; Jan. 2001; R. Acconcia leg. CEMT. ‒ Táchira • 1 ♂, 1 ♀; San Cristóbal, Frio River [“Río Frio”]; 1 Aug. 1984; B. Joffre leg.; CEMT. Distribution Colombia, Venezuela (new country record) and Peru. Subregion of Venezuela Andes mountains. Literature record No previous reports for Venezuela. Ateuchus hoplopygus (Harold, 1868) Fig. 10B Choeridium hoplopygum Harold, 1868b: 57 (original description). Type locality: Colombia. Namebearing type: holotype lost (MC, pers. obs.). Choeridium hoplopygum – Harold 1869d: 1007 (catalogue, cited for Colombia). — Gillet 1911b: 53 (catalogue, cited for Colombia). — Balthasar 1939c: 62 (key to species, cited for Colombia). — Blackwelder 1944: 204 (checklist of species, cited for Colombia). Ateuchus hoplopygus – Vulcano & Pereira 1967: 590 (key). — Medina et al. 2001: 137 (checklist for Colombia). Material examined VENEZUELA ‒ Zulia • 1 ♂; Rosário de Perijá; 19 Jul. 2006; no collector; in dung; CEMT. LOZANO DE LA ROSA C. et al., Dung beetles (Scarabaeinae) of Venezuela 23 Fig. 10. Species distribution. A. Ateuchus columbianus (Harold, 1868). B. Ateuchus hoplopygus (Harold, 1868). Green diamond = CEMT collection data. European Journal of Taxonomy 959: 1–272 (2024) 24 Distribution Colombia and Venezuela (new country record). Subregion of Venezuela Maracaibo Depression. Literature record No previous reports for Venezuela. Ateuchus persplendens (Balthasar, 1939) Choeridium persplendens Balthasar, 1939c: 50 (original description). Type locality: Colombia: Valle del Cauca: Dagua River. Name-bearing type: holotype (SMTD), examined by MC and FZVM. Distribution Colombia and Venezuela. Literature record Roze 1955: 43 (Venezuela: Sucre: Carúpano). Remarks Although the CEMT houses no specimens of A. persplendens, MC confirms the presence of the species in Venezuela as reported by Roze (1955) based on his study of material from other collections for his forthcoming revision of Ateuchus. Ateuchus simplex (Le Peletier de Saint-Fargeau & Audinet-Serville, 1828) Fig. 11A Choeridium simplex Le Peletier de Saint-Fargeau & Audinet-Serville, 1828: 356 (original description). Type locality: French Guiana. Name-bearing type: syntypes lost (MC, pers. obs.). Choeridium simplex – Harold 1869d: 1007 (catalogue, cited for Cayenne). — Gillet 1911b: 53 (catalogue, cited for Cayenne). — Balthasar 1939c: 64 (cited for French Guiana, Amazonas, Peru and Ecuador); 1941: 347 (cited for Peru). — Blackwelder 1944: 204 (checklist of species, cited for French Guiana). Ateuchus simplex – Roze 1955: 44 (checklist for Venezuela). — Vulcano & Pereira 1967: 588 (cited for northern South America). — Feer 2000: 31 (list, cited for French Guiana); 2008: 56, 62 (ecology); 2013: 766 (list French Guiana). — Feer & Pincebourde 2005: 29 (ecological study). — Brûlé et al. 2011a: 193 (list, cited for French Guiana). — Larsen 2011: 97 (cited for Suriname); 2013: 95 (cited for Suriname). — Brûlé & Dalens 2012: 37 (list, cited for French Guiana). — Price & Feer 2012: 327 (sampling). — Feer & Boissier 2015: 169 (list). — Ratcliffe et al. 2015: 196 (checklist for Peru). — Hielkema & Hielkema 2019: 44 (catalogue for the Guianas). Material examined VENEZUELA ‒ Bolívar • 2 ♂♂; Sifontes, Cuyuní River, Isla de Anacoco; 6 Aug. 2006; no collector; in dung; CEMT. Distribution Venezuela, Suriname, French Guiana, Brazil, and Peru. LOZANO DE LA ROSA C. et al., Dung beetles (Scarabaeinae) of Venezuela 25 Fig. 11. Species distribution. A. Ateuchus simplex (Le Peletier & Serville, 1828). B. Ateuchus substriatus (Harold, 1868). Green diamond = CEMT collection data; black circle = literature data. European Journal of Taxonomy 959: 1–272 (2024) 32 Coprobius – Brullé 1838: 294 (diagnosis). — Castelnau 1840: 68 (synonym of Canthon). — Reiche 1841: 213 (key, type species designation). — Agassiz 1846: 282 (catalogue). — Harold 1869d: 989 (synonym of Canthon). — Burmeister 1873: 410 (synonym of Canthon). — Gillet 1911b: 27 (synonym of Canthon). — Lucas 1920: 201 (synonym of Canthon). — Blackwelder 1944: 198 (synonym of Canthon). — Pereira & Martínez 1956a: 112 (synonym of Canthon). — Martínez 1959: 27 (synonym of Canthon). — Vulcano & Pereira 1964: 602 (synonym of Canthon). — Solís & Kohlmann 2002: 2 (synonym of Canthon); 2012: 2 (synonym of Canthon). — Ratcliffe 2002: 12 (synonym of Canthon). — Cupello & Vaz-de-Mello 2018: 9-16 (history, nomenclator). Canthon – Castelnau 1840: 68 (redescription). — Agassiz 1846: 184 (catalogue). — Lacordaire 1855: 77 (redescription). — LeConte 1861: 125 (key). — Harold 1868a: 1 (redescription); 1869d: 989 (catalogue). — Blanchard 1885: 163 (redescription). — Blatchley 1910: 912 (key). — Gillet 1911b: 27 (catalogue). — Lucas 1920: 164 (catalogue, distribution). — Dawson 1922: 61 (key). — Paulian 1938: 235 (key); 1939: 22 (redescription, type species designation). — Pessôa & Lane 1941: 414 (diagnosis). — Islas 1942: 303 (redescription). — Blackwelder 1944: 198 (list). — Lane 1947: 110 (comment). — Roze 1955: 41 (checklist for Venezuela). — Pereira & Martínez 1956a: 96 (key). — Martínez 1959: 27 (catalogue for Argentina). — Halffter 1961: 231, 258 (key, redescription). — Vulcano & Pereira 1964: 602 (catalogue); 1967: 549 (key for the Amazon). — Halffter & Matthews 1966: 261 (catalogue, distribution). — Halffter & Martínez 1968: 265 (diagnosis); 1977: 38, 69 (key, redescription). — Howden & Young 1981: 14, 19 (key, redescription). — Halffter & Edmonds 1982: 139 (catalogue, distribution). — Medina & Lopera-Toro 2000: 311 (key). — Vaz-de-Mello 2000: 191 (checklist for Brazil). — Medina et al. 2001: 135 (checklist for Colombia); 2003: 64 (distribution). — Solís & Kohlmann 2002: 2 (redescription); 2012: 2 (checklist for Costa Rica). — Ratcliffe 2002: 13 (checklist for Panama). — Ratcliffe et al. 2002: 49 (key). — Morón 2003: 30 (redescription). — Hamel-Leigue et al. 2006: 13 (list). — Vaz-de-Mello et al. 2011a: 26 (key). — Carvajal et al. 2011: 113, 314 (diagnosis, list). — Krajcik 2012: 63 (list). — Boilly & Vaz-de-Mello 2013: 108 (key). — França et al. 2016: 3 (comments). — Chamorro et al. 2018: 76, 92 (list for Ecuador); 2019: 40 (catalogue). — Cupello & Vaz-de-Mello 2018: 9–16 (history, nomenclator). — Hielkema & Hielkema 2019: 61 (catalogue for the Guianas). Coeloscelis – Agassiz 1846: 268 (catalogue). — Lacordaire 1855: 76 (redescription). — Harold 1869d: 989 (cited as synonym of Canthon). — Gillet 1911b: 27 (synonym of Canthon). — Blackwelder 1944: 198 (synonym of Canthon). — Pereira & Martínez 1956a: 112 (synonym of Canthon). — Martínez 1959: 27 (synonym of Canthon). — Vulcano & Pereira 1964: 602 (synonym of Canthon). — Solís & Kohlmann 2002: 2 (synonym of Canthon); 2012: 2 (synonym of Canthon). — Ratcliffe 2002: 12 (synonym of Canthon, cited as Coeloschelis). — Cupello & Vaz-de-Mello 2018: 9-16 (history, nomenclator). Paedhyboma – Paulian 1938: 235 (key); 1939: 21 (redescription). — Vulcano & Pereira 1964: 636 (catalogue); 1967: 549 (key). — Halffter & Martínez 1977: 38 (synonym of Canthon). — Halffter & Edmonds 1982: 139 (cited as synonym of Canthon). — Solís & Kohlmann 2002: 2 (cited as synonym of Canthon); 2012: 3 (synonym of Canthon, cited as Paedohyboma). — Ratcliffe 2002: 12 (synonym of Canthon, cited as Paedohyboma). — Cupello & Vaz-de-Mello 2018: 1–205 (history, nomenclator). Canthomoechus – Halffter 1961: 231 (key). — Vulcano & Pereira 1964: 590 (catalogue). — Halffter & Matthews 1966: 261 (catalogue, distribution). — Halffter & Martínez 1977: 38 (synonym of Canthon). — Halffter & Edmonds 1982: 139 (cited as synonym of Canthon). — Solís & Kohlmann 2002: 2 (cited as synonym of Canthon); 2012: 3 (cited as synonym of Canthon). — Ratcliffe 2002: 13 (synonym of Canthon). — Cupello & Vaz-de-Mello 2018: 9–16 (history, nomenclator). Canthon aberrans (Harold, 1868) Fig. 14A Deltochilum aberrans Harold, 1868a: 8 (original description). Type locality: Colombia. Name-bearing type: syntype (MNHN), examined by FZVM. LOZANO DE LA ROSA C. et al., Dung beetles (Scarabaeinae) of Venezuela 33 Canthon bifurcatus Robinson, 1948a: 37 (original description). Type locality: Venezuela: Mérida. Namebearing type: holotype (USNM) (originally in Mark Robinson’s personal collection; Robinson 1948a), not examined. Canthon juanae Martínez, 1949b: 176 (original description). Type locality: Bolivia: La Paz: Nor Yungas: Choro River at its confluence with the Coroico River, camp “Dalen”, 700 m a.s.l. Namebearing type: having been originally deposited in Martínez’s personal collection (Martínez 1949b), the holotype would now be expected to be in MACN (see Cupello & Vaz-de-Mello 2018), but MC did not find it there in 2014. MC located in the museum solely six specimens labelled as paratypes. Deltochilum aberrans – Harold 1869d: 995 (catalogue, distribution); 1880: 18 (distribution). — Gillet 1911b: 35 (catalogue). — Campos 1921: 55 (list, cited for Ecuador). — Blackwelder 1944: 202 (checklist). — Martínez 1947b: 113 (notes, correction). — Pereira & d’Andretta 1955: 48 (synonym). — Vulcano & Pereira 1964: 636 (catalogue). — Solís & Kohlmann 2002: 5 (redescription). Paedhyboma aberrans – Kolbe 1893: 191 (transferred to the genus). — Shipp 1897: 195 (cited). — Paulian 1939: 21 (redescription, distribution). — Balthasar 1941: 345 (catalogue, cited for Peru); 1951: 330 (cited for Peru). — Martínez 1947b: 113 (corrections to the work of Blackwelder 1944). — Gacharná 1951: 221 (catalogue for Colombia). — Pereira & d’Andretta 1955: 48 (synonymy). — Vulcano & Pereira 1964: 636 (catalogue of species); 1967: 551 (key). — Halffter & Martínez 1977: 87 (monograph). — Solís & Kohlmann 2002: 5 (synonym). Canthon juanae – Martínez 1951: 23 (cited as synonym of Canthon bifurcatus). — Pereira 1953: 394 (cited as synonym of Canthon plicatipennis). — Pereira & d’Andretta 1955: 48 (cited as synonym of Canthon aberrans). — Vulcano & Pereira 1964: 636 (cited as synonym of Paedhyboma aberrans). — Solís & Kohlmann 2002: 5 (cited as synonym of Canthon aberrans). Canthon aberrans – Pereira & d’Andretta 1955: 48 (transferred to the genus). — Vulcano & Pereira 1964: 636 (cited). — Solís & Kohlmann 2002: 5 (redescription, distribution). — Hamel-Leigue et al. 2006: 13 (cited for Bolivia). — Ratcliffe et al. 2015: 195 (checklist for Peru). Canthon bifurcatus – Pereira & d’Andretta 1955: 48 (cited as synonym of Canthon aberrans). — Vulcano & Pereira 1964: 636 (cited as synonym of Paedhyboma aberrans). — Solís & Kohlmann 2002: 5 (cited as synonym of Canthon aberrans). Canthon (Canthon) aberrans – Halffter & Martínez 1977: 87 (transferred to the subgenus). — Vaz-deMello 2000: 191 (checklist for Brazil). — Medina et al. 2001: 135 (checklist for Colombia); 2003: 64 (Appendix 1). — Ratcliffe 2002: 12 (checklist for Panama). — Carvajal et al. 2011: 314–315 (cited for Ecuador). — Krajcik 2012: 63 (checklist). — Solís & Kohlmann 2002: 5 (redescription, distribution). — Chamorro et al. 2018: 85, 87, 92 (figs 8a, 10h, key for Ecuador). — Chamorro et al. 2019: 43 (catalogue). Deltochilum (Paedhyboma) aberrans –Solís & Kohlmann 2002: 5 (redescription). Distribution Costa Rica, Panama, Colombia, Venezuela, Brazil, Ecuador, Peru, and Bolivia (Solís & Kohlmann 2002). Subregion of Venezuela Andes mountains. Literature records Robinson 1948a: 37 (Venezuela: Mérida). — Martínez 1951: 23 (Venezuela). — Vulcano & Pereira 1964: 636 (Venezuela). — Solís & Kohlmann 2002: 5 (Venezuela). — Chamorro et al. 2019: 42 (Venezuela). European Journal of Taxonomy 959: 1–272 (2024) 34 Fig. 14. Species distribution. A. Canthon aberrans (Harold, 1868). B. Canthon balachowskyi (Martínez & Halffter, 1972). Black circle = literature data. LOZANO DE LA ROSA C. et al., Dung beetles (Scarabaeinae) of Venezuela 35 Canthon balachowskyi (Martínez & Halffter, 1972) Fig. 14B Scybalocanthon balachowskyi Martínez & Halffter, 1972: 46 (original description). Type locality: Venezuela: Aragua: Parque Nacional Henri Pittier, 1200 m a.s.l. Name-bearing type: holotype (MACN), examined by MC. Scybalocanthon balachowskyi – Halffter & Martínez 1977: 68 (list, monograph). Canthon balachowskyi – Silva & Valois 2019: 301–302 (revision, new combination). Distribution Venezuela (endemic) (Martínez & Halffter 1972). Subregion of Venezuela Central Coast Mountain Range. Literature records Martínez & Halffter 1972: 46 (Venezuela: Aragua). Canthon bicolor Castelnau, 1840 Fig. 15A Canthon bicolor Castelnau, 1840: 69 (original description). Type locality: French Guiana. Name-bearing type: lectotype (OUMNH), designated by Nunes et al. (2018), examined by FZVM. Canthon bicolor – Harold 1868a: 14, 71 (key, redescription); 1869d: 990 (catalogue, cited for Cayenne). — Gillet 1911b: 28 (catalogue). — Schmidt 1922: 64, 73 (distribution). — Balthasar 1939a: 189 (key). — Blackwelder 1944: 198 (checklist). — Vulcano & Pereira 1964: 604 (catalogue). — Halffter & Martínez 1977: 39, 70 (key, revision). — Forsyth & Gill 1993: 70 (list for French Guiana). — Feer 2000: 32 (list for French Guiana); 2008: 56, 62 (ecology); 2013: 766 (list for French Guiana). — Vaz-de-Mello 2000: 191 (checklist for Brazil). — Feer & Pincebourde 2005: 30 (list). — Brûle et al. 2011a: 193 (list); 2014: 183 (list). — Larsen 2011: 98 (list for Suriname); 2013: 96 (list for Palumeu River, Suriname). — Price & Feer 2012: 327 (list). — Feer & Boissier 2015: 169 (list). Canthon (Goniocanthon) bicolor – Nunes et al. 2018: 17 (revision). Distribution Venezuela, Guyana, Suriname, French Guiana, and Brazil (Nunes et al. 2018). Subregions of Venezuela System of hills and low piedmont mountains of the Guiana Shield and System of low mountains and hills Imataca-Cuyuní del Escudo Northeast Guiana. Literature records Nunes et al. 2018: 17 (Venezuela: Bolívar). Canthon cyanellus LeConte, 1859 Fig. 15B Canthon cyanellus LeConte, 1859: 11 (original description). Type locality: United States: Texas. Namebearing type: syntype (MCZC), examined by FZVM. European Journal of Taxonomy 959: 1–272 (2024) 36 Canthon sallei Harold, 1863: 174 (original description). Type locality: Nicaragua: Granada: near Granada city. Name-bearing type: syntypes (BMNH ‒ ex Sallé collection), examined by FZVM. Canthon spinosus Harold, 1863: 174 (original description). Type locality: Mexico: Veracruz: Tuxpan. Name-bearing type: a single known syntype (MNHN ‒ ex Harold collection), examined by FZVM. Canthon speciosus Harold, 1868a: 41 (original description). Type locality: Mexico. Name-bearing type: a single known syntype (MNHN ‒ ex Harold collection), examined by FZVM. Canthon sallei gutticollis Schmidt, 1920: 123–124 (original description). Type locality: Colombia: “La Garita”; see Botero & Monné (2012) for details on this ambiguous locality. Name-bearing type: lectotype (NHRS), designated by Vaz-de-Mello & Cupello (2018), examined by FZVM. Canthon sallei triangulatus Schmidt, 1920: 123–124 (original description). Type locality: Colombia: “La Garita”; see comments for the previous name. Name-bearing type: lectotype (NHRS), designated by Vaz-de-Mello & Cupello (2018), examined by FZVM. Canthon cyanellus havranekae Martínez, 1988a: 89 (original description). Type locality: Venezuela: Táchira: Cárdenas: Táriba: Finca La Alameda, “Carretera Vieja Cordero-San Cristóbal”. Namebearing type: the holotype, having been originally deposited in Martínez’s personal collection (Martínez 1988a), would now be expected to be in MACN (see Cupello & Vaz-de-Mello 2018), but it was not found there by MC in 2014. Canthon spinosus – Harold 1868a: 42 (monograph, redescription); 1869d: 990 (cited for Mexico). — Bates 1887: 27 (list, cited for Mexico). — Gillet 1911b: 29 (catalogue, cited for Mexico). Canthon cyanellus – Harold 1868a: 141 (monograph); 1869d: 990 (cited for Kansas). — Horn 1870: 44, 45 (key, cited for Mexico and Texas). — Blanchard 1885: 164, 166 (key, cited for Texas). — Gillet 1911b: 29 (catalogue, cited for Texas). — Leng 1920: 248 (catalogue, cited Mexico and Texas). — Schmidt 1920: 123 (contribution, comments); 1922: 74 (distribution). — Balthasar 1939a: 211 (key, distribution). — Islas 1942: 304–305, 333 (key, redescription, list). — Roze 1955: 41 (checklist for Venezuela). — Halffter 1961: 260 (redescription, distribution). — Vulcano & Pereira 1964: 609 (catalogue). — Martínez 1988a: 89 (list). — Amézquita et al. 1999: 119 (biodiversity). — Escobar 2000: 206 (checklist for Colombia). — Barbero 2001: 2 (list, cited for Nicaragua). — Estrada & Coates-Estrada 2002: 1911 (ecology). — Halffter & Arellano 2002: 147–148 (ecology). — Solís & Kohlmann 2002: 11 (redescription, distribution); 2012: 3 (checklist for Costa Rica). — Hernández et al. 2003: 96 (diversity). — Kohlmann et al. 2007: 28 (atlas). — Noriega et al. 2007: 81 (list). — Pulido-Herrera et al. 2007: 307 (cited for Andean region of Colombia). — Larsen et al. 2008: 1294 (list). — Medina & Pulido-Herrera 2009: 58 (diversity). — Cultid-Medina et al. 2012: 67 (guide). — Ferrer-Paris et al. 2013: 108 (list). Canthon sallei – Harold 1868a: 39 (monograph, redescription); 1869d: 993 (catalogue, cited for Nicaragua). — Kirsch 1873: 340 (comments, cited for Peru). — Bates 1887: 26 (list, distribution). — Gillet 1911b: 33 (catalogue, cited for Central America and Colombia). — Balthasar 1939a: 211 (key, distribution). — Martínez 1947b: 113 (correction: C. cyanellum var. sallei). — Robinson 1948b: 85 (revision, key). — Solís & Kohlmann 2012: 3 (checklist for Costa Rica, cited as synonym of Canthon cyanellus LeConte, 1859). Canthon speciosus – Bates 1887: 27 (list, cited for Mexico and Guatemala). — Gillet 1911b: 29 (catalogue, cited for Mexico and Guatemala). — Solís & Kohlmann 2012: 3 (checklist for Costa Rica, cited as synonym of Canthon cyanellus). Canthon cyanellus sallei – Schmidt 1922: 74 (distribution). — Halffter 1961: 260 (redescription, distribution), 262, 265 (distribution, redescription). — Escobar 1997: 423 (list). — Martínez 1988a: 89 (list). — Barbero 2001: 2 (list, distribution, cited for Nicaragua). — Bustos-Gómez & LoperaToro 2003: 61 (diet). — Arias-Buriticá et al. 2011: 876 (cited). Canthon cyanellus gutticollis – Schmidt 1922: 74 (distribution). — Roze 1955: 41 (checklist for Venezuela). — Halffter 1961: 262 (distribution). Canthon cyanellus triangulatus – Schmidt 1922: 74 (distribution). — Halffter 1961: 262 (distribution). LOZANO DE LA ROSA C. et al., Dung beetles (Scarabaeinae) of Venezuela 37 Fig. 15. Species distribution. A. Canthon bicolor Castelnau, 1840. B. Canthon cyanellus LeConte, 1859. Green diamond = CEMT collection data; black circle = literature data. European Journal of Taxonomy 959: 1–272 (2024) 38 Canthon cyanellus speciosus – Balthasar 1939a: 211 (key, distribution). — Balthasar 1941: 343 (list). — Balthasar 1951: 328 (list). — Halffter 1961: 262 (distribution). Canthon sallei gutticollis – Balthasar 1939a: 212 (key, distribution). — Vaz-de-Mello & Cupello 2018: 51 (designation of lectotype). Canthon sallei triangulatus – Balthasar 1939a: 212 (key, distribution). Canthon cyanellum – Blackwelder 1944: 199 (checklist, cited for Mexico). Canthon cyanellum gutticollis – Blackwelder 1944: 199 (list). Canthon cyanellum sallei – Blackwelder 1944: 199 (list). Canthon cyanellum triangulatus – Blackwelder 1944: 199 (list). Canthon cyanellus havranekae – Blanco 1987: 42 (catalogue). — Havranek 1989: 61 (list). Canthon (Canthon) cyanellus – Padilla-Gil & Halffter 2007: 88 (comments). — Delgado et al. 2012: 327 (list). — Hielkema & Hielkema 2019: 61 (catalogue for the Guianas). Material examined VENEZUELA ‒ Barinas • 1 spec.; Otopún; 28 Aug. 2006; in faeces; CEMT. ‒ Zulia • 1 spec.; Rosario de Perijá; 18 Jul. 2016; in faeces; CEMT. ‒ Cojedes • 4 specs; Hato Piñero, Reserva Natural La Candelaria; 30 Aug.–1 Sep. 1990; G.C. McGavin leg.; frog carcass pitfall trap; CEMT. ‒ Táchira • 2 specs; Rubio, La Tuquerena; 14 Apr. 1982; D. Havranek leg.; CEMT • 2 specs; Palo Gordo, near San Cristobal, La Alameda Farm; 1000 m a.s.l.; Apr. 1987; D. Havranek leg.; CEMT. Distribution From southern Texas (United States) to Brazil (Solís & Kohlmann 2002). Subregions of Venezuela Maracaibo Depression, Plains, System of hills and low sierras Lara-Falcón, System of hills and low piedmont mountains of the Guiana Shield, System of low mountains and hills Imataca-Cuyuní of Northeast Guiana Shield, Andes mountains, and Central Coast Mountain Range. Literature records Schmidt 1922: 74 (Venezuela: Distrito Capital: Caracas). — Blackwelder 1944: 199 (Venezuela). — Roze 1955: 41 (Venezuela). — Vulcano & Pereira 1964: 609 (Venezuela). — Martínez 1988a: 89 (Venezuela: Táchira: Finca “La Alameda”). — Barbero 2001: 2 (Venezuela). — Solís & Kohlmann 2002: 11 (Venezuela: Táchira). — Padilla-Gil & Halffter 2007: 88 (Venezuela: Táchira: Táriba and San Cristóbal). — Larsen et al. 2008: 1294 (Venezuela: Bolívar: Lago Guri). — Arias-Buriticá et al. 2011: 876 (Venezuela). — Ferrer-Paris et al. 2013: 108 (Venezuela: Bolívar: Isla de Anacoco; Miranda: Altos de Pipe; Sucre: Araya; Yaracuy: Hacienda Guáquira; and Zulia: Rosario de Perijá). Canthon juvencus Harold, 1868 Fig. 16A Canthon juvencus Harold, 1868a: 111 (original description). Type locality: Colombia: Bolívar: Cartagena de Indias. Name-bearing type: syntypes (MFNB and MNHN), examined by FZVM. Canthon raripilus Bates 1887: 30 (original description). Type locality: Panama: Panamá: in the districts of Balboa (Pearl Islands: Isla del Rey: San Miguel) and Taboga (Isla Taboga). Name-bearing type: syntypes (BMNH and MNHN), examined by FZVM. Canthon juvencus – Harold, 1869d: 991 (cited for Cartagena); 1880: 17 (cited for Muzo, Colombia). — Candèze 1891: 329 (list). — Schmidt 1922: 76 (list, distribution). — Gillet 1911b: 30 (cited for Colombia). — Balthasar 1939a: 206 (key, cited for Colombia and Mexico). — Howden & Young, 1981: 28 (revision). — Escobar 1997: 423 (list); 2000: 207 (checklist for Colombia). — Amézquita LOZANO DE LA ROSA C. et al., Dung beetles (Scarabaeinae) of Venezuela 39 et al. 1999: 119 (biodiversity). — Solís & Kohlmann 2002: 27 (redescription); 2012: 7 (checklist for Costa Rica). — Bustos-Gómez & Lopera-Toro 2003: 61 (diet). — Noriega 2004: 40 (checklist for Tinigua Park, Colombia). — Hamel-Leigue et al. 2006: 13 (list). — Kohlmann et al. 2007: 29 (atlas). — Pulido-Herrera et al. 2007: 307 (cited for Andean region of Colombia). — Larsen et al. 2008: 1294 (list). — Medina & Pulido-Herrera 2009: 58 (diversity). — Arias-Buriticá et al. 2011: 876 (cited for Brazil, Colombia and Panama). — Cultid-Medina et al. 2012: 67 (guide). — FerrerParis et al. 2013: 108 (list). — Ratcliffe et al. 2015: 195 (checklist for Peru). — Nieto et al. 2020: 136 (report). Canthon raripilus – Gillet 1911b: 32 (cited for Panama). — Schmidt 1920: 125 (synonymized with Canthon juvencus, revision). Canthon juvencum – Blackwelder 1944: 199 (list). — Roze 1955: 41 (checklist for Venezuela, cited as C. juvenicum). Canthon juvencum raripilus – Gacharná 1951: 221 (catalogue). Glaphyrocanthon (Glaphyrocanthon) juvencus – Pereira & Martínez 1956a: 127 (new combination, key). Glaphyrocanthon juvencus – Martínez et al. 1964a: 10, 12 (distribution, key). — Vulcano & Pereira 1964: 662 (catalogue). Canthon (Canthon) juvencus — Halffter & Martínez 1977: 90 (monograph, key). — Hielkema & Hielkema 2019: 61 (catalogue for the Guianas). Material examined VENEZUELA ‒ Guárico • 1 spec.; near Paso Real Guárico; 20 Aug. 1959; C.J. Rosales leg.; CEMT. ‒ Yaracuy • 1 spec.; Bolívar, Aroa; 10°0′0″ N, 68°0′0″ W; 702 m a.s.l.; 19 Jul. 2009; M. Asmüssen, P. Colmenares, and H. Martínez leg.; in human faeces; CEMT • 1 spec.; same data as for the preceding except for elevation; 710 m a.s.l.; CEMT. Distribution Costa Rica, Panama, Colombia, Venezuela, Suriname, Brazil, Peru and Bolivia. Subregions of Venezuela Maracaibo Depression, System of hills and low piedmont mountains of the Guiana Shield, System of low mountains and hills Imataca-Cuyuní of Escudo Northeast Guiana, and Central Coast Mountain Range. Literature records Candèze 1891: 329 (Venezuela: [actually, Carabobo]: San-Esteban). — Schmidt 1922: 76 (Venezuela). — Blackwelder 1944: 199 (Venezuela). — Pereira & Martínez 1956a: 127 (Venezuela). — Martínez et al. 1964a: 10 (Venezuela). — Vulcano & Pereira 1964: 662 (Venezuela). — Howden & Young 1981: 28 (Venezuela). — Solís & Kohlmann 2002: 27 (Venezuela). — Larsen et al. 2008: 1294 (Venezuela: Bolívar state). — Ferrer-Paris et al. 2013: 108 (Venezuela: Bolívar: Isla de Anacoco; Miranda: Altos de Pipe; Sucre: Araya; Yaracuy: Hacienda Guáquira; Zulia: Rosario de Perijá). Canthon rufocoeruleus (Martínez, 1948) Fig. 16B Glaphyrocanthon rufocoeruleus Martínez, 1948a: 47 (original description). Type locality: Venezuela: Distrito Capital: Pico del Naiguatá. Name-bearing type: holotype (MACN), examined by MC. Glaphyrocanthon rufocoeruleus – Martínez 1949a: 282 (key). — Roze 1955: 43 (checklist, cited). European Journal of Taxonomy 959: 1–272 (2024) 40 Fig. 16. Species distribution. A. Canthon juvencus Harold, 1868. B. Canthon rufocoeruleus (Martínez, 1948). Green diamond = CEMT collection data; black circle = literature data. LOZANO DE LA ROSA C. et al., Dung beetles (Scarabaeinae) of Venezuela 41 Glaphyrocanthon (Coprocanthon) rufocoeruleus – Martínez 1950: 160, 171 (new subgenus, catalogue). — Pereira & Martínez 1956a: 127 (list, distribution, identification key). — Martínez et al. 1964a: 2‒3, 6, 10, 12 (comments, identification key). — Vulcano & Pereira 1964: 666 (catalogue); 1967: 560 (key, cited for Brazil). Canthon (Glaphyrocanthon) rufocoeruleus – Halffter & Martínez 1977: 78, 80 (list). Distribution Venezuela and Brazil. Subregions of Venezuela Peneplain of the Casiquiare River–Upper Orinoco, and Central Coast Mountain Range. Literature records Martínez 1948a: 47 (Venezuela: Distrito Capital: Pico Naiguatá and Amazonas: Yavita); 1949a: 282 (Venezuela); 1950: 160 (Venezuela). — Pereira & Martínez 1956a: 127 (Venezuela). — Martínez et al. 1964a: 10 (Venezuela). — Vulcano & Pereira 1964: 666 (Venezuela); 1967: 560 (Venezuela). Remarks Rivera-Cervantes & Halffter (1999) treated Glaphyocanthon rufocoeruleus as a junior synonym of Canthon subhyalinus subhyalinus Harold, 1867. Their argument was limited to the observation that, according to them, no “significant difference” existed between rufocoeruleus, originally described from Venezuela, and “typical” (their word) subhyalinus from Colombia. Indeed, Martínez (1948a) gave no comparison between the two in the original description, indicating that he was at that point most likely unaware of the identity of C. subhyalinus and could have easily described the species again under his new name rufocoeruleus. Later, however, Martínez included both as separate species in identification keys (Pereira & Martínez 1956a; Martínez et al. 1964a), and pointed out the relative length between the mesotarsi and the mesotibiae as a distinguishing character (tarsi longer than the tibiae in rufocoeruleus, as long as or shorter than the tibiae in subhyalinus). Such was the importance that he gave to this character that, initially, Martínez even classified them in different subgenera of Glaphyrocanthon (Pereira & Martínez 1956a), even though he later changed his mind and not only transferred them to the same subgenus, but even entertained the idea that they could be conspecific subspecies under Glaphyrocanthon quadriguttatus (Olivier, 1789) (Martínez et al. 1964b). While the conspecifity with quadriguttatus was ruled out by Rivera-Cervantes & Halffter (1999), the purported difference in the legs was not even mentioned, prompting one to wonder whether the authors may have overlooked it and this compromised their judgement over the synonymy. Yet, no one has so far challenged RiveraCervantes & Halffter’s (1999) decision to invalidate rufocoeruleus. Here, we reinstall Martínez’s original classification, listing rufocoeruleus as valid and treating C. subhyalinus as a separate species, based on the third author’s still unpublished morphological and geographical observations that, according to him, dismiss Rivera-Cervantes & Halffter’s synonymy. Both C. rufocoeruleus and C. subhyalinus subhyalinus occur in Venezuela. Canthon septemmaculatus linearis Schmidt, 1920 Fig. 17A Canthon septemmaculatus linearis Schmidt, 1920: 115 (original description). Type locality: Colombia. Name-bearing type: lectotype (NHRS), designated by Vaz-de-Mello & Cupello (2018). Canthon septemmaculatus maculicollis Schmidt, 1920: 115 (original description). Type locality: Peru. Name-bearing type: lectotype (NHRS), designated by Vaz-de-Mello & Cupello (2018). European Journal of Taxonomy 959: 1–272 (2024) 48 Glaphyrocanthon (Glaphyrocanthon) subhyalinus – Pereira & Martínez 1956a: 132 (new combination, distribution). — Halffter & Martínez 1977: 80 (revision). — Estrada & Coates-Estrada 2002: 1911 (ecology). Canthon (Glaphyrocanthon) quadrimaculatus – Martínez et al. 1964a: 2 (revision). — Halffter & Martínez 1977: 80 (revision). Glaphyrocanthon (Coprocanthon) subhyalinus – Martínez et al. 1964a: 12 (key). — Vulcano & Pereira 1967: 560 (key). Canthon (Glaphyrocanthon) subhyalinus – Halffter & Martínez 1977: 80 (list). — Krajcik 2012: 63 (list, cited for Nova Granada, Bogotá). — Vaz-de-Mello 2000: 191 (checklist for Brazil). — Medina et al. 2003: 64 (cited for Colombia). — Chamorro et al. 2018: 93 (list for Ecuador); 2019: 59 (catalogue). Canthon (Glaphyrocanthon) subhyalinus subhyalinus – Rivera-Cervantes & Halffter 1999: 63 (diagnosis). — Barbero 2001: 3 (catalogue, cited for Nicaragua). — Solís & Kohlmann 2002: 5 (key). — Morón 2003: 33 (cited for Mexico). — Carvajal et al. 2011: 314–315 (cited for Ecuador). — Hielkema & Hielkema 2019: 68 (bibliographic review for Guianas). Canthon subhyalinus subhyalinus – Solís & Kohlmann 2002: 5, 47 (key, redescription); 2012: 3 (checklist for Costa Rica). — Kohlmann et al. 2007: 29 (checklist). Material examined VENEZUELA ‒ Aragua • 1 spec.; Parque Nacional Henri Pittier, Estación Biológica Rancho Grande; 1200 m; 20 Mar. 1966; Bordon leg.; CEMT. Distribution Mexico, Costa Rica, Panama, Colombia, and Venezuela. Subregion of Venezuela Central Coast Mountain Range. Literature records Harold 1869d: 994 (Venezuela). — Roze 1955: 43 (Venezuela: Aragua). — Vulcano & Pereira 1964: 665 (Venezuela). — Rivera-Cervantes & Halffter 1999: 63 (Venezuela: Aragua; La Guaira: Naiguatá). — Chamorro et al. 2019: 59 (Venezuela). Canthon triangularis triangularis (Drury, 1773) Fig. 20A Scarabaeus triangularis Drury, 1773: [“index to the first volume”] (the species-group name was made available by being published accompanied by an indication in the sense of Articles 12.2.1 and 12.2.7 of the Code (ICZN 1999), namely a reference to the description and illustration given in the first volume of Drury’s work [Drury 1770: 82, pl. 36 fig. 7]. See ICZN (1957) for details on the publication of this work, its index, and the availability and date of publication of new names first appeared in it). Type locality: Suriname. Name-bearing type: unknown typification status and whereabouts. Canthon triangularis caliginosus Schmidt, 1920: 122 (description). Type locality: French Guiana. Name-bearing type: lectotype (NHRS), designated by Vaz-de-Mello & Cupello (2018), examined by FZVM. Scarabaeus triangularis – Fabricius 1775: 30 (list, characteristics); 1781: 33 (cited for “America meridionali”); 1787a: 18 (characteristics); 1792: 69 (cited for Suriname) 1801: 63–64 (list). — Herbst 1789: 298 (catalogue, redescription). — Olivier 1789: 166 (characteristics, cited for Cayenne; Suriname). LOZANO DE LA ROSA C. et al., Dung beetles (Scarabaeinae) of Venezuela 49 Fig. 19. Species distribution. A. Canthon subcyaneus Erichson, 1848. B. Canthon subhyalinus subhyalinus Harold, 1867. Green diamond = CEMT collection data; black circle = literature data. European Journal of Taxonomy 959: 1–272 (2024) 50 Canthon triangularis – Castelnau 1840: 69 (list). — Erichson 1848: 563 (list, cited as Ateuchus triangularis, cited for Guyana). — Harold 1868a: 46 (monograph); 1869d: 994 (catalogue, cited for Cayenne). — Heyne & Taschenberg 1908: 61 (list). — Gillet 1911b: 34 (catalogue, cited for Suriname). — Schmidt 1922: 66, 81 (characteristics, distribution). — Boucomont 1928b: 1 (list). — Balthasar 1939a: 197 (key); 1941: 342 (distribution); 1951: 327 (list, distribution). — Pessôa & Lane 1941: 418 (comments). — Blackwelder 1944: 202 (list). — Gacharná 1951: 221 (list, cited for Colombia). — Roze 1955: 43 (checklist, cited as C. triangulare). — Vulcano & Pereira 1964: 632 (catalogue). — Halffter & Martínez 1977: 70 (revision). — Blanco 1987: 42 (catalogue). — Forsyth & Gill 1993: 70 (list). — Escobar 2000: 207 (checklist for Colombia). — Quintero & Roslin 2005: appendix A (ecology). — Feer 2008: 56, 62 (ecology); 2013: 766 (list for French Guiana). — Larsen et al. 2008: 1294 (list). — Medina & Pulido-Herrera 2009: 59 (diversity). — Vaz-de-Mello et al. 2011b: 86 (list). — Brûlé et al. 2011a: 193 (list, cited for French Guiana); 2014: 183 (list for Montagne Pelée). — Ferrer-Paris et al. 2013: 108 (list). — Ratcliffe 2013: 493 (list, cited for Brazil). — Nunes et al. 2014: 408–410 (cited, list). — Feer & Boissier 2015: 169 (list). — Hielkema & Hielkema 2019: 72 (catalogue for the Guianas). — Nieto et al. 2020: 136 (report). — Storck-Tonon et al. 2020: 2426 (diversity). Canthon triangularis caliginosus – Schmidt 1922: 81 (key). — Balthasar 1939a: 197 (key); 1941: 342 (distribution); 1951: 327 (list, distribution). — Pessôa & Lane 1941: 418 (comments). Material examined VENEZUELA ‒ Táchira • 2 specs; Libertador, San Joaquín de Navay; 7°47′59.99″ N, 71°40′24.24″ W; 655 m a.s.l.; 26 Aug. 2006; T. Good et al. leg.; human faeces; CEMT • 2 specs; La Morita del Chururu; 30 Sep. 1985; D. Havranek leg.; pig dung trap; CEMT • 4 specs; Río Frío; Mar 1988; J. Blanco leg.; CEMT. Distribution Colombia, Venezuela, Trinidad and Tobago, Suriname, French Guiana, Brazil, and Bolivia. Subregions of Venezuela Maracaibo Depression, System of hills and low piedmont mountains of the Guiana Shield, Andes mountains, and Guiana Shield. Literature records Pessôa & Lane 1941: 418 (Venezuela). — Blackwelder 1944: 202 (Venezuela). — Roze 1955: 43 (Venezuela). — Vulcano & Pereira 1964: 632 (Venezuela). — Blanco 1987: 42 (Venezuela: Táchira). — Larsen et al. 2008: 1294 (Venezuela: Bolívar: Lago Gurí). — Ferrer-Paris et al. 2013: 108 (Venezuela: Bolívar). Canthon variabilis (Martínez, 1948) Fig. 20B Glaphyrocanthon variabilis Martínez, 1948a: 43 (original description). Type locality: Venezuela: La Guaira: Pico Naiguatá. Name-bearing type: holotype (MACN), examined by MC. Glaphyrocanthon (Glaphyrocanthon) variabilis – Martínez 1949a: 282 (key); 1950: 170 (notes, distribution). — Pereira & Martínez 1956a: 134 (distribution). — Martínez et al. 1964a: 16 (key). — Vulcano & Pereira 1964: 665 (catalogue). — Vaz-de-Mello 2000: 191 (checklist for Brazil). — Hielkema & Hielkema 2019: 68 (catalogue for the Guianas). Glaphyrocanthon variabilis – Roze 1955: 43 (checklist for Venezuela). LOZANO DE LA ROSA C. et al., Dung beetles (Scarabaeinae) of Venezuela 51 Fig. 20. Species distribution. A. Canthon triangularis triangularis (Drury, 1770). B. Canthon variabilis (Martínez, 1948). Green diamond = CEMT collection data; black circle = literature data. European Journal of Taxonomy 959: 1–272 (2024) 52 Canthon (Glaphyrocanthon) variabilis – Halffter & Martínez 1977: 80 (new combination, revision). Canthon variabilis – Forsyth et al. 1998: 369 (list). — Hamel-Leigue et al. 2006: 14 (inventory). — Noriega et al. 2007: 82 (list). — Krajcik 2012: 64 (list). — Ferrer-Paris et al. 2013: 109 (list). Material examined Paratype VENEZUELA ‒ 1 spec; PARÁTIPO/ “Venezuela. [illegible] / [illegible]. 7-VIII-/34/ Lichy leg.”; “Canthon (Glaphyrocan / thon)a. variabilis Mar- / tinez G. Halffter y /A. Martínez det 78”; “ Glaphyrocanthon / variabilis / sp. n./ A. Martínez -DET 1948”; CEMT-00068510. Additional material VENEZUELA ‒ Barinas • 1 spec.; Barinitas; 23–25 Mar. 1967; Bordón leg.; CEMT. Distribution Colombia, Venezuela, Suriname, Brazil, and Bolivia. Subregions of Venezuela Maracaibo Depression, Andes mountains, Central Coast Mountain Range, Oriental Coast Range, and Guiana Shield. Literature records Martínez 1948a: 43 (Venezuela: La Guaira: Pico Naiguatá); 1950: 170 (Venezuela: La Guaira: Pico Naiguatá). — Roze 1955: 43 (Venezuela: Sucre: Puerto de Hierro). — Pereira & Martínez 1956a: 134 (Venezuela). — Vulcano & Pereira 1964: 665 (Venezuela); 1967: 562 (Venezuela). — Ferrer-Paris et al. 2013: 109 (Venezuela: Bolívar: Isla de Anacoco; Miranda: Altos de Pipe; Yaracuy: Guáquira farm; and Zulia: Rosario de Perijá). Genus Canthonella Chapin, 1930 Canthonella Chapin, 1930: 1 (original description). Type species: Canthonella parva Chapin, 1930, by original designation. Ipselissus d’Olsoufieff, 1935: 35 (original description). Type species: Epilissus silphoides Harold, 1867, by original monotypy. Ipsepilissus Paulian, 1939: 29 (original description). Type species: Epilissus silphoides Harold, 1867, by original designation. Canthonella – Blackwelder 1944: 198 (list). — Martínez 1954: 64 (comments). — Pereira & Martínez 1956a: 94, 99 (key). — Halffter 1961: 230 (key). — Vulcano & Pereira 1964: 581 (catalogue). — Matthews 1965: 433, 447 (key, redescription); 1966: 7, 75 (key, redescription). — Zayas & Matthews 1966: 3, 16 (key, redescription). — Halffter & Matthews 1966: 260 (catalogue, distribution). — Halffter & Martínez 1967: 90 (redescription); 1977: 34, 58 (key, comments). — Halffter & Edmonds 1982: 139 (catalogue, distribution). — Ratcliffe & Smith 1999: 2 (comments, list). — Medina & Lopera-Toro 2000: 311 (key). — Vaz-de-Mello 2000: 192 (checklist for Brazil). — Medina et al. 2001: 136 (checklist for Colombia); 2003: 65 (distribution). — Vaz-de-Mello et al. 2011a: 25 (key). — Carvajal et al. 2011: 115, 314–315 (diagnosis, list). — Krajcik 2012: 64 (list). — Boilly & Vaz-de-Mello 2013: 107 (key). — Chamorro et al. 2018: 75, 76, 84, 93 (list for Ecuador); 2019: 9 (catalogue). — Hielkema & Hielkema 2019: 73 (catalogue for the Guianas). Ipsepilissus – Martínez 1947b: 113 (correction). — Matthews 1966: 75 (cited). Ipselissus – Martínez 1954: 59 (redescription). — Pereira & Martínez 1956a: 94, 99 (key, distribution). — Halffter 1961: 230 (key). — Vulcano & Pereira 1964: 581 (catalogue). — Halffter & Matthews LOZANO DE LA ROSA C. et al., Dung beetles (Scarabaeinae) of Venezuela 53 1966: 260 (catalogue, distribution). — Halffter & Martínez 1967: 103 (redescription); 1968: 211 (comments); 1977: 34, 51 (key, synonym of Canthonella). Canthonella gomezi (Halffter & Martínez, 1968) Fig. 21A‒B Ipselissus gomezi Halffter & Martínez, 1968: 221 (original description). Type locality: Venezuela: Miranda: Altos de Pipe. Name-bearing type: holotype (MACN), examined by MC. Canthonella gomezi — Halffter & Martínez 1977: 58 (new synonym, key). — Medina et al., 2001: (list); 2003: 65 (phylogenetic analysis). — Krajcik 2012: 64 (list). — Ferrer-Paris et al. 2013: 109 (list). Material examined Paratype VENEZUELA ‒ Miranda • 1 spec.; “Venezuela / E° Miranda / Altos de Pipe / a la luz / coll. Martínez / Jun. 948”; “Ipselissus gomezi / Halffter y / Martínez”; “HALFFTER collection [red label]”; CEMT00078136. Additional material VENEZUELA ‒ Miranda • 1 spec.; Altos de Pipe, Instituto Venezolano de Investigaciones Científicas (“IVIC”); 12 Sep. 2008; “pasantes BDV2008” leg.; “BV-T-021, cebo:(?)”; CEMT • 1 spec.; same data as for the preceding; “BV-T-035”; CEMT. Distribution Colombia and Venezuela. Possibly in Peru (Ratcliffe et al. 2015). Subregion of Venezuela Guiana Shield. Literature records Halffter & Martínez 1968: 221 (Venezuela: Aragua and Miranda). — Medina et al. 2003: 65 (Venezuela). — Krajcik 2012: 64 (Venezuela: Miranda: Alto de Pipe). — Ferrer-Paris et al. 2013: 109 (Venezuela: Miranda: Altos de Pipe). Genus Coprophanaeus d’Olsoufieff, 1924 Phanaeus (Coprophanaeus) d’Olsoufieff, 1924: 22 (original description). Type species: Scarabeus jasius Olivier, 1789, by original designation. Phanaeus (Coprophanaeus) – Pessôa 1934: 295 (redescription). — Pessôa & Lane 1941: 476 (key). — Martínez 1959: 100 (catalogue for Argentina). — Halffter & Matthews 1966: 258 (checklist). — Vulcano & Pereira 1967: 570 (key). — Krajcik 2012: 204 (list). Coprophanaeus – Blackwelder 1944: 209 (list). — Edmonds 1972: 820, 839 (key, redescription). — Howden & Young 1981: 12 (key). — Halffter & Edmonds 1982: 136 (catalogue, distribution). — Gámez & Mora 2000: 17 (list). — Medina & Lopera-Toro 2000: 303 (key). — Vítolo 2000: 593 (key); 2004: 289 (redescription). — Vaz-de-Mello 2000: 192 (checklist for Brazil). — Medina et al. 2001: 140 (checklist for Colombia). — Arnaud 2002b: 13 (key). — Ratcliffe et al. 2002: 49 (key). — Ratcliffe 2002: 16 (checklist for Panama). — Morón 2003: 59 (redescription). — Edmonds 1994: 17 (key). — Hamel-Leigue et al. 2006: 17 (list); 2009: 56 (distribution). — Edmonds & Zídek 2010: 8 (revision). — Vaz-de-Mello et al. 2011a: 24 (key). — Carvajal et al. 2011: 138, 320 (diagnosis, European Journal of Taxonomy 959: 1–272 (2024) 54 Fig. 21. Species distribution. A. Canthonella gomezi (Halffter & Martínez, 1968). B. Dorsal view of C. gomezi. C. Coprophanaeus abas (MacLeay, 1819). Green diamond = CEMT collection data; black circle = literature data. LOZANO DE LA ROSA C. et al., Dung beetles (Scarabaeinae) of Venezuela 55 list). — Solís & Kohlmann 2012: 7 (checklist for Costa Rica). — Cupello & Vaz-de-Mello 2013b: 359–360 (cited); 2014: 435–436 (cited). — Boilly & Vaz-de-Mello 2013: 107 (key). — França et al. 2016: 5 (comments). — Chamorro et al. 2018: 75, 93 (list for Ecuador); 2019: 69–70, (catalogue). — Hielkema & Hielkema 2019: 94 (catalogue for the Guianas). Coprophanaeus abas (MacLeay, 1819) Fig. 21C Phanaeus abas MacLeay, 1819: 126 (original description). Type locality: Trinidad and Tobago: Trinidad Island. Name-bearing type: neotype (BMNH), designated by Arnaud (2002d), not examined. Phanaeus (Coprophanaeus) rex Balthasar, 1939f: 239 (original description). Type locality: South America. Name-bearing type: holotype (NMPC), not examined. Coprophanaeus (Coprophanaeus) abas – Arnaud 2002b: 28 (key, distribution). — Edmonds & Zídek 2010: 43, 50 (key, comments, revision). — Gámez 2010: 15 (comments). — Pacheco & Vaz-deMello 2015: 2–3, 7 (list, distribution, key). — Gámez & Acconcia 2018: 73 (key). — Hielkema & Hielkema 2019: 94 (catalogue for the Guianas). Coprophanaeus jasius abas – Arnaud 2002c: 3 (designation of neotype). Coprophanaeus rex – Arnaud 2002c: 3 (cited as synonym of Coprophanaeus jasius abas). Coprophanaeus abas – Arnaud 2002b: 28 (new combination, key, distribution). — Edmonds & Zídek 2010: 43, 50 (key, comments, revision). — Gámez 2010: 15 (comments). — Gámez & Acconcia 2018: 73 (key). Material examined VENEZUELA ‒ Amazonas • 2 specs; Caño Piojo; Apr. 2006; D. Garcia leg.; CEMT. ‒ Bolívar • 1 spec.; Isla de Anacoco; 6 Jun. 2006; curso NM2006 leg.; faeces, 23h, 16:20; CEMT. Distribution Colombia, Venezuela, Trinidad and Tobago, and Brazil. Subregions of Venezuela Peneplain of the Casiquiare River–Upper Orinoco, System of hills and low piedmont mountains of the Guiana Shield, System of low mountains and hills Imataca-Cuyuní of Northeast Guyana Shield, Andes mountains, and Central Coast Mountain Range. Literature records Arnaud 2002b: 28 (Venezuela). — Edmonds & Zídek 2010: 43, 50 (Venezuela: Amazonas, Aragua, Barinas and Bolívar). — Gámez 2010: 15, 19 (Venezuela: Llanos del Orinoco). — Gámez & Acconcia 2018: 73 (Venezuela: Cordillera de Mérida). Coprophanaeus corythus (Harold, 1863) Fig. 22A Phanaeus corythus Harold, 1863: 163 (original description). Type locality: Mexico: Veracruz: Córdoba. Name-bearing type: lectotype (MNHN), designated by Arnaud (1982), not examined. Phanaeus perseus Harold, 1880: 27 (original description). Type locality: Colombia: Antioquia: Medellin. Name-bearing type: lectotype (MNHN), designated by Arnaud (1982), not examined. Coprophanaeus (Coprophanaeus) telamon nevinsoni Arnaud & Gámez, 2002: 10 (original description). Type locality: Colombia: Magdalena: Parque Nacional Tayrona. Name-bearing type: holotype (CPFA), not examined. European Journal of Taxonomy 959: 1–272 (2024) 56 Phanaeus corythus – Harold 1869c: 65 (comments); Harold 1869d: 65 (catalogue, cited for Mexico). — Bates 1887: 55, 482 (distribution, figs 16–17). — Nevinson 1892: 3 (list). — Kolbe 1905: 581 (comments). — Gillet 1911b: 82 (catalogue). — Islas 1942: 314 (comments). Phanaeus (Coprophanaeus) corythus – d’Olsoufieff 1924: 26, 68, 141 (key, distribution). Phanaeus telamon corythus – Pereira & Martínez 1956b: 234 (new combination). — Arnaud 2002b: 35 (cited as synonym of Coprophanaeus telamon). Coprophanaeus telamon corythus – Howden & Young 1981: 141 (redescription). — Deloya 1992: 21 (list). — Estrada & Coates-Estrada 2002: 1911 (ecology). — Hernández et al. 2003: 96 (diversity). — Kohlmann et al. 2007: 29 (checklist). — Arnaud 2002b: 35 (cited as synonym of Coprophanaeus telamon). Coprophanaeus telamon nevinsoni – Arnaud 2002b: 36 (redescription). — Gámez 2004: 48, 58 (ecology). — Gámez & Acconcia 2009: 388, 392–394 (ecology). — Edmonds & Zídek 2010: 93 (revision, synonym of C. telamon). Coprophanaeus (Coprophanaeus) corythus – Edmonds & Zídek 2010: 93 (revision, new status). — Gámez & Acconcia 2016: 57–60 (note). Phanaeus perseus – Edmonds & Zídek 2010: 93 (revision, as synonym of Coprophanaeus corythus). Distribution Mexico, Belize, Guatemala, El Salvador, Honduras, Nicaragua, Costa Rica, Panama, Colombia, and Venezuela. Subregions of Venezuela Maracaibo Depression, System of hills and low sierras Lara-Falcón, Serrania de Perija, Andes mountains and Central Coast Mountain Range. Literature records Howden & Young 1981: 141 (Venezuela). — Arnaud 2002b: 36 (Venezuela: Mérida and Zulia states). — Arnaud & Gámez 2002: 10 (Venezuela: Mérida State). — Gámez 2004: 48, 58 (Venezuela: Mérida and Zulia states). — Gámez et al. 2006: 96, 99–102 (Venezuela: Mérida: Zea). — Gámez & Acconcia 2009: 388, 392–394 (Venezuela: Zulia: Colón: Depresión de Maracaibo). — Edmonds & Zídek 2010: 93 (Venezuela). — Gámez & Acconcia 2016: 57–60 (Carabobo, Falcón and Zulia states). Coprophanaeus dardanus (MacLeay, 1819) Fig. 22B Phanaeus dardanus MacLeay, 1819: 126 (original description). Type locality: Brazil. Name-bearing type: Unknown typification status and whereabouts (Edmonds & Zídek 2010). Phanaeus bitias Harold, 1863: 163 (original description). Type locality: Somewhere in the Amazon or the Atlantic Forest. Originally said to be Mexico: Oaxaca: Tuxtepec: “hot regions” (“Mexique (Tustepec), régions chaudes”), but this is mistaken. The species is exclusively South American (Edmonds & Zídek 2010; Cupello & Vaz-de-Mello 2013a). The type locality is here corrected following Recommendation 76A.2 of the Code. Name-bearing type: a male holotype fixed by monotypy and originally from Chevrolat’s personal collection (Harold 1863), but whose current whereabouts is unknown (Edmonds & Zídek 2010). It was not found by Arnaud (1982) in the MNHN, where most of the Harold collection material has been housed since the 1950s (see Cupello 2020), nor by Olaf Jäger in the collection under his care in SMTD (pers. com. to MC, 3 March 2023), where the Chevrolat Scarabaeinae collection has been preserved since the 1910s via the Carl Felsche collection (Horn et al. 1990a; Cupello 2020). Bates (1887) informed that the holotype (cited as “the type-specimen”) was in Sallé’s collection. If correct, Sallé would have somehow LOZANO DE LA ROSA C. et al., Dung beetles (Scarabaeinae) of Venezuela 57 Fig. 22. Species distribution. A. Coprophanaeus corythus (Harold, 1863). B. Coprophanaeus dardanus (MacLeay, 1819). Black circle = literature data. European Journal of Taxonomy 959: 1–272 (2024) 64 Coprophanaeus lichyi Arnaud, 2002 Fig. 24B Coprophanaeus lichyi Arnaud, 2002a: 3 (original description). Type locality: Venezuela: Zulia: Sierra de Perijá, Rio Yaza. Name-bearing type: holotype (CPFA), not examined. Coprophanaeus lichyi – Arnaud 2002b: 45 (key). — Edmonds & Zídek 2010: 104 (comments). Distribution Venezuela (endemic). Subregion of Venezuela Maracaibo Depression. Literature records Arnaud 2002a: 3 (Venezuela: Zulia: Sierra de Perijá); 2002b: 45 (Venezuela: Zulia). Coprophanaeus parvulus (d’Olsoufieff, 1924) Fig. 25A Phanaeus parvulus d’Olsoufieff, 1924: 67 (original description). Type locality: French Guiana: Passoura. Name-bearing type: lectotype (MHNH), designated by Arnaud (1982), not examined. Coprophanaeus parvulus – Blackwelder 1944: 209 (checklist). — Larsen 2011: 98 (list for Suriname). — Escobar 2000: 207 (checklist for Colombia). — Feer 2000: 32 (list for French Guiana); 2008: 62 (ecology, list); 2013: 766 (list for French Guiana). — Vítolo 2000: 599 (key for Colombia). — Medina et al. 2001: 140 (checklist for Colombia). — Feer & Pincebourde 2005: 30 (list). — Quintero & Roslin 2005: appendix A (ecology). — Gámez & Acconcia 2009: 394 (comments). — Brûle et al. 2011a: 193 (inventory); 2014: 183 (inventory). — Brûle & Dalens 2012: 18 (cited). — Feer & Boissier 2015: 169 (list). Coprophanaeus (Coprophanaeus) parvulus – Martínez & Clavijo 1990: 154 (notes, cited as parrulus). — Escobar 2000: 207 (checklist for Colombia). — Feer 2000: 32 (list for French Guiana); 2008: 62 (ecology, list); 2013: 766 (list for French Guiana). — Vítolo 2000: 599 (key for Colombia). — Medina et al. 2001: 140 (checklist for Colombia). — Arnaud 2002b: 38 (key). — Feer & Pincebourde 2005: 30 (list). — Quintero & Roslin 2005: appendix A (ecology). — Gámez et al. 2006: 101 (comments). — Gámez & Acconcia 2009: 394 (comments). — Edmonds & Zídek 2010: 3–5, 77, 89 (key, revision). — Brûle et al. 2011a: 193 (inventory); 2014: 183 (inventory). — Larsen 2011: 98 (list for Suriname). — Brûle & Dalens 2012: 18 (cited). — Feer & Boissier 2015: 169 (list). — Boilly et al. 2016: 89, 93 (key, catalogue). — Hielkema & Hielkema 2019: 95 (catalogue for the Guianas). Distribution Colombia, Venezuela, Guyana, Suriname, French Guiana, Brazil, and Peru (Edmonds & Zídek 2010; Ratcliffe et al. 2015). Subregions of Venezuela Maracaibo Depression, Peneplain of the Caura and Paragua rivers, Peneplain of the Casiquiare River– Upper Orinoco and System of hills and low piedmont mountains of the Guiana Shield. LOZANO DE LA ROSA C. et al., Dung beetles (Scarabaeinae) of Venezuela 65 Literature records Martínez & Clavijo 1990: 8 (Venezuela: Amazonas: Atures). — Arnaud 2002b: 38 (Venezuela: Bolívar). — Vítolo 2004: 292 (Venezuela: Amazonas: Puerto Ayacucho). — Gámez et al. 2006: 101 (Venezuela: Amazonas). — Gámez & Acconcia 2009: 394 (Venezuela: Zulia: Colón: Depresión de Maracaibo). — Edmonds & Zídek 2010: 77, 89 (Venezuela: Amazonas and Bolívar). Coprophanaeus telamon (Erichson, 1847) Fig. 25B Phanaeus telamon Erichson, 1847a: 106 (original description). Type locality: Peru. Name-bearing type: holotype (MFNB), not examined. Phanaeus telamon – Harold 1869d: 1019 (list, cited for Peru). — Kirsch 1873: 341 (cited for Peru). — Bates 1887: 56 (diagnosis). — Nevinson 1892: 7 (list, revision, cited for Brazil and Peru). — Kolbe 1905: 581 (comments). — Gillet 1911b: 86 (catalogue, cited for Peru and Brazil). — Balthasar 1941: 350 (distribution); 1951: 336 (distribution). — Blackwelder 1944: 210 (list, cited for Brazil and Peru). — Pereira 1953: 391 (notes). Phanaeus (Coprophanaeus) telamon – d’Olsoufieff 1924: 26, 68, 135, 142 (key, distribution). — Pessôa 1934: 296, 299 (key, redescription). — Martínez 1947b: 114 (cited). — Vulcano & Pereira 1967: 572 (key). — Krajcik 2012: 204 (list). — Gámez & Mora 2000: 17 (list). Coprophanaeus (Coprophanaeus) telamon telamon – Pereira & Martínez 1956b: 233 (new combination). — Blanco 1988: 43 (catalogue). — Havranek 1989: 61 (list). — Martínez & Clavijo, 1990: 7 (notes of Phanaeina). — Arnaud 2002b: 35 (monograph, redescription). — Gámez 2004: 59 (distribution). — Hamel-Leigue et al. 2006: 17 (list); 2009: 57 (distribution for Bolivia). — Gámez & Acconcia 2009: 394 (ecological information). — Carvajal et al. 2011: 318, 320 (list). Coprophanaeus (Coprophanaeus) telamon – Forsyth et al. 1998: 370 (conservation, list). — Amézquita et al. 1999: 119 (biodiversity). — Escobar 2000: 207 (checklist for Colombia). — Vaz-de-Mello 2000: 192 (checklist for Brazil). — Vítolo 2000: 599 (key); 2004: 290 (diagnosis, distribution, cited for Colombia). — Barbero 2001: 6 (note for Nicaragua). — Medina et al. 2001: 140 (checklist for Colombia). — Halffter & Arellano 2002: 148 (ecology). — Pulido-Herrera et al. 2003: 54 (list for Caqueta, Colombia); 2007: 307 (cited for Andean region of Colombia). — Carpio et al. 2009: 469 (ecology). — Medina & Pulido-Herrera 2009: 61 (diversity). — Gámez 2010: 16, 19 (comments). — Edmonds & Zídek 2010: 77, 91 (key, diagnosis). — Carvajal et al. 2011: 318, 320 (list). — Vaz-de-Mello et al. 2011b: 86 (list). — Cultid-Medina et al. 2012: 57 (guide). — Cupello & Vaz-de-Mello 2013a: 358, 361–364, 367 (comments, distribution). — Ferrer-Paris et al. 2013: 109 (list). — Figueroa et al. 2014: 127 (list, distribution). — Pardo-Locarno & Camero 2014: 211 (list for Chocó, Colombia). — Silva et al. 2014: 348 (diversity); 2016: 7 (ecology, biogeography). — Ratcliffe et al. 2015: 197 (checklist for Peru). — Chamorro et al. 2018: 93 (list for Ecuador); 2019: 77 (catalogue). — Gámez & Acconcia 2018: 65, 69, 73 (comments, cited for Peru). — Giraldo et al. 2018: 70 (guide). — Santos et al. 2018: 46 (list for Acre state, Brazil). — Hielkema & Hielkema 2019: 95 (catalogue for the Guianas). — Nieto et al. 2020: 136 (report). Material examined VENEZUELA ‒ Aragua • 1 spec.; Cuyagua; 50 m a.s.l.; 6 Jul. 2004; D. García leg.; CEMT. ‒ Táchira • 2 specs; San Cristóbal, Parque Nacional Natural Paramillo; Jul. 1993; J. Blanco leg.; CEMT. ‒ Yaracuy • 1 spec.; Bolívar, Aroa; 10.3867° N, 68.6471° W; 1380 m a.s.l.; Jul. 2009; Asmüssen, Colmenares and Martínez leg.; CEMT. Distribution Colombia, Venezuela, Brazil, Ecuador, Peru, and Bolivia. European Journal of Taxonomy 959: 1–272 (2024) 66 Fig. 25. Species distribution. A. Coprophanaeus parvulus (d’Olsoufieff, 1924). B. Coprophanaeus telamon (Erichson, 1847). Green diamond = CEMT collection data; black circle = literature data. LOZANO DE LA ROSA C. et al., Dung beetles (Scarabaeinae) of Venezuela 67 Subregions of Venezuela Maracaibo Depression, Plains, System of hills and low sierras Lara-Falcón, System of low mountains and hills Imataca-Cuyuní of Northeast Guyana Shield, Andes mountains, Central Coast Mountain Range, and Guiana Shield. Literature records Blanco 1988: 43 (Venezuela: Táchira: Carretera San Cristóbal–Chorro del Indio, Río Negro, Paramillo, Palo Gordo and Morretón). — Havranek 1989: 61 (Venezuela: Táchira). — Martínez & Clavijo 1990: 7 (Venezuela: Aragua: Parque Nacional Henri Pittier, Estación Biológica Rancho Grande). — Gámez & Mora 2000: 17 (Venezuela). — Arnaud 2002b: 35 (Venezuela: Táchira). — Barbero 2001: 6 (Venezuela). — Gámez 2004: 59 (Venezuela: Mérida: Guaraque: Mesa de Quintero; and Táchira: San Cristóbal: Hacienda Pánaga); 2010: 16, 19 (Venezuela: Andean plains flank). — Gámez & Acconcia 2009: 394 (Venezuela: Zulia: Colón: Depresión de Maracaibo); 2018: 65, 69, 73 (Venezuela: Barinas: Bolívar and Andrés Eloy Blanco). — Hamel-Leigue et al. 2009: 57 (Venezuela). — Edmonds & Zídek 2010: 77, 91 (Venezuela: Amazonas [Táchira?]: La Tuquerena Rubio; Aragua; Bolívar; Falcón; Mérida; and Táchira). — Cupello & Vaz-de-Mello 2013a: 367 (Venezuela: Aragua; Táchira: San Cristóbal; La Guaira; and Yaracuy: Bolívar). — Ferrer-Paris et al. 2013: 109 (Venezuela: Yaracuy: Hacienda Guáquira). — Chamorro et al. 2019: 77 (Venezuela). Genus Cryptocanthon Balthasar, 1942 Cryptocanthon Balthasar, 1942: 36 (original description). Type species: Cryptocanthon paradoxus Balthasar, 1942, by original designation. Cryptocanthon — Pereira & Martínez 1956a: 96, 181 (key, distribution). — Halffter 1961: 231 (key); 2003: 24 (redescription). — Vulcano & Pereira 1964: 673 (catalogue); 1967: 548 (key). — Halffter & Matthews 1966: 261 (catalogue, distribution). — Howden 1973: 39 (redescription). — Halffter & Martínez 1977: 35, 60 (key, list). — Howden & Young 1981: 13, 39 (key, redescription). — Halffter & Edmonds 1982: 139 (catalogue, distribution). — Medina & Lopera-Toro 2000: 306 (key). — Vazde-Mello 2000: 192 (checklist for Brazil). — Medina et al. 2001: 136 (checklist for Colombia). — Cook 2002: 4 (revision). — Ratcliffe 2002: 13 (checklist for Panama). — Medina et al. 2003: 65 (distribution). — Hamel-Leigue et al. 2006: 14 (list). — Vaz-de-Mello et al. 2011a: 23 (key). — Carvajal et al. 2011: 118, 316 (diagnosis, list). — Krajcik 2012: 82 (list). — Solís & Kohlmann 2012: 3 (checklist for Costa Rica). — Boilly & Vaz-de-Mello 2013: 106 (key). — Chamorro et al. 2018: 74, 93 (list for Ecuador); 2019: 79 (catalogue). Cryptocanthon andersoni Cook, 2002 Fig. 26A Cryptocanthon andersoni Cook, 2002: 11 (original description). Type locality: Venezuela: Merida: Santos Marquina: Tabay, “Mucay, Send. Lago Suero” (?), 2700 m a.s.l. Name-bearing type: holotype (CMNC), not examined. Cryptocanthon andersoni — Krajcik 2012: 82 (list). Distribution Venezuela (endemic). Subregion of Venezuela Cordillera de los Andes. European Journal of Taxonomy 959: 1–272 (2024) 68 Literature records Cook 2002: 11 (Venezuela: Mérida, Lara and Trujillo). — Krajcik 2012: 82 (Venezuela). Cryptocanthon gilli Cook, 2002 Fig. 26B Cryptocanthon gilli Cook, 2002: 12 (original description). Type locality: Venezuela: Bolívar: 120 km S of El Dorado. Name-bearing type: holotype (CMNC), not examined. Cryptocanthon gilli – Krajcik 2012: 82 (list). — Hielkema & Hielkema 2019: 74 (catalogue for the Guianas). Distribution Venezuela (endemic). Subregion of Venezuela System of low mountains and hills Imataca-Cuyuní Northeast Guyana Shield, and Guiana Shield. Literature records Cook 2002: 10, 12 (Venezuela: Bolívar state). — Krajcik 2012: 82 (Venezuela). Cryptocanthon nebulinus Howden, 1973 Fig. 27A–B Cryptocanthon nebulinus Howden, 1973: 40, 45 (original description). Type locality: Venezuela: Aragua: Parque Nacional Henri Pittier, Estación Biológica Rancho Grande. Name-bearing type: holotype (CMNC), not examined. Cryptocanthon nebulinus — Cook 2002: 10, 14 (revision). — Krajcik 2012: 82 (list). Material examined VENEZUELA ‒ Aragua • 1 spec.; Parque Nacional Henri Pittier, Estación Biológica Rancho Grande; 1500 m a.s.l.; 21‒25 Feb. 1971; S. Peck leg.; human dung trap; forest; CEMT • 1 spec.; Parque Nacional Henri Pittier, Estación Biológica Rancho Grande, Portachuelo; 1100 m a.s.l.; 21‒24 Feb. 1971; S. Peck leg.; human dung trap; CEMT. Distribution Venezuela (endemic). Subregion of Venezuela Central Coast Mountain Range. Literature records Cook 2002: 10, 14 (Venezuela: Aragua). — Krajcik 2012: 82 (Venezuela: Aragua). Cryptocanthon punctatus Cook, 2002 Fig. 27C LOZANO DE LA ROSA C. et al., Dung beetles (Scarabaeinae) of Venezuela 69 Fig. 26. Species distribution. A. Cryptocanthon andersoni Cook, 2002. B. Cryptocanthon gilli Cook, 2002. Black circle = literature data. European Journal of Taxonomy 959: 1–272 (2024) 70 Cryptocanthon punctatus Cook, 2002: 83 (original description). Type locality: Venezuela: Trujillo: old road to Trujillo city. Name-bearing type: holotype (CMNC), not examined. Cryptocanthon punctatus — Krajcik 2012: 82 (list). Distribution Venezuela (endemic). Subregion of Venezuela Andes mountains. Literature records Cook 2002: 11, 83 (Venezuela: Trujillo). — Krajcik 2012: 82 (Venezuela). Genus Deltochilum Eschscholtz, 1822 Deltochilum Eschscholtz, 1822: 37 (original description). Type species: Deltochilum dentipes Eschscholtz, 1822, by original monotypy. Note: Bouchard et al. (2024) recently stated that D. dentipes was fixed as the type species of Deltochilum by the subsequent designation of Thon (1832), but this is incorrect. Deltochilum dentipes was the sole nominal species that Eschscholtz originally included in the genus without hesitation. It is true that he also said that Ateuchus gibbosus (Fabricius, 1775) (currently, Deltochilum gibbosum) was “closely related” to D. dentipes and “seems to also belong to the genus” (“[...] welcher ebenfalls zu dieser Gattung zu gehören scheint”). However, because this was at most a doubtful assignment of the species to Deltochilum, gibbosus cannot be considered a second originally included nominal species per Article 67.2.5 of the Code (ICZN 1999) and has, therefore, never been eligible for type fixation. Deltochilum dentipes, having been the only originally included nominal species, was automatically fixed as the type species by original monotypy (Art. 68.3). After we brought this to his attention, Patrice Bouchard (pers. com. to MC, 12th June 2024) now agrees with us. Anamnesis Vigors, 1826: 510 (original description). Type species: Anamnesis macleayii Vigors, 1826, by original monotypy. Meghyboma Kolbe, 1893: 192 (original description). Type species: Deltochilum dentipes Eschscholtz, 1822, by subsequent designation of Shipp (1897). Telhyboma Kolbe, 1893: 192 (original description). Type species: Deltochilum orbiculare Van Lansberge, 1874, by original monotypy. Deltochilum – Agassiz 1846: 341 (catalogue). — Burmeister 1848: 134 (redescription); 1873: 408 (redescription). — Lacordaire 1855: 79 (redescription). — LeConte 1861: 125 (key). — Harold 1869d: 995 (catalogue, distribution). — Van Lansberge 1874: 188 (characteristics). — Kolbe 1893: 191 (redescription). — Shipp 1897: 194 (comments). — Gillet 1911b: 35 (catalogue). — Lucas 1920: 228 (catalogue, distribution). — Paulian 1938: 235, 237 (key, redescription). — Pessôa & Lane 1941: 411, 426 (key, redescription). — Blackwelder 1944: 202 (list). — Lane 1946: 171 (comments). — Roze 1955: 43 (checklist for Venezuela). — Pereira & Martínez 1956a: 96 (key). — Martínez 1959: 50 (catalogue for Argentina). — Halffter 1961: 231 (key); 2003: 26 (redescription). — Vulcano & Pereira 1964: 639 (catalogue). — Halffter & Matthews 1966: 261 (catalogue, distribution). — Vulcano & Pereira 1967: 549 (key). — Halffter & Martínez 1977: 36 (key). — Howden & Young 1981: 14, 36 (key, redescription). — Halffter & Edmonds 1982: 139 (catalogue, distribution). — Medina & Lopera-Toro 2000: 311 (key). — Vaz-de-Mello 2000: 192 (checklist for Brazil). — Medina et al. 2001: 136 (checklist for Colombia). — Ratcliffe et al. 2002: 49 (key); 2015: 195 (checklist for Peru). — Ratcliffe 2002: 13 (checklist for Panama). — Medina et al. 2003: 64 (distribution). — LOZANO DE LA ROSA C. et al., Dung beetles (Scarabaeinae) of Venezuela 71 Fig. 27. Species distribution. A. Cryptocanthon nebulinus Howden, 1973. B. Dorsal view of C. nebulinus. C. Cryptocanthon punctatus Cook, 2002. Green diamond = CEMT collection data; black circle = literature data. European Journal of Taxonomy 959: 1–272 (2024) 72 Hamel-Leigue et al. 2006: 14 (list). — Vaz-de-Mello et al. 2011a: 25 (key). — Carvajal et al. 2011: 123, 316 (diagnosis, list). — Krajcik 2012: 88 (list). — Solís & Kohlmann 2012: 4 (checklist for Costa Rica). — Boilly & Vaz-de-Mello 2013: 107 (key, characteristics). — Chamorro et al. 2018: 76, 93–94 (list for Ecuador); 2019: 9, 82 (catalogue). — Hielkema & Hielkema 2019: 74 (catalogue for the Guianas). Anamnesi – Agassiz 1846: 58 (catalogue). — Harold 1869d: 995 (cited as synonym, cited as Annamesis). — Shipp 1897: 194 (comments, cited as synonym of Deltochilum). — Gillet 1911b: 35 (cited as synonym of Deltochilum, cited as Annamesis). — Paulian 1938: 237 (cited as synonym of Deltochilum, cited as Annamesis). — Blackwelder 1944: 202 (cited as synonym of Deltochilum, cited as Annamesis). — Lane 1946: 171 (comments). — Pereira & Martínez 1956a: 120 (cited as synonym of Deltochilum). — Martínez 1959: 50 (cited as synonym of Deltochilum). — Vulcano & Pereira 1964: 639 (cited as synonym of Deltochilum). — Ratcliffe 2002: 13 (cited as synonym of Deltochilum). — Solís & Kohlmann 2012: 4 (cited as synonym of Deltochilum, cited as Annamesis). Meghyboma – Shipp 1897: 194 (comments, cited as synonym of Deltochilum). — Paulian 1938: 243, 246 (key, review). — Blackwelder 1944: 202 (cited synonym of Deltochilum). — Lane 1946: 174 (cited synonym of Deltochilum). — Halffter & Edmonds 1982: 139 (cited as subgenus of Deltochilum). — Solís & Kohlmann 2012: 4 (cited as synonym of Deltochilum). Telhyboma – Paulian 1938: 219–220, 224 (cited, comments). — Génier 2012: 25–26, 30–31 (revision, new synonymy). — Gonzalez-Alvarado & Vaz-de-Mello 2021a: 2‒3 (revision). Deltochilum abdominale Martínez, 1947 Fig. 28A Deltochilum abdominalis [sic] Martínez, 1947a: 269 (original description). Type locality: Venezuela: Distrito Capital: Pico Naiguatá. Name-bearing type: holotype (MACN), examined by MC. Note: as explained by Harold (1869d), the genus-group name Deltochilum is formed from the combination of the Ancient Greek nouns δέλτᾰ (‘délta’), the name for the fourth letter in the Ancient Greek alphabet (Δ) and, by extension, used to refer to anything shaped like that letter, and χεῖλος (‘kheîlos’), meaning ‘lip’, ‘mouth’, in reference to the labrum of the single species originally assigned to the genus, a structure that Eschscholtz (1822) described as being “cordiform” (i.e., heart-shaped, or an inverted letter delta). Since, in his Latinisation of the new name, Eschscholtz changed the ending of ‘kheîlos’ to ‘-um’, a neuter suffix in Latin, to render ‘-chilum’, the noun Deltochilum, too, is to be regarded as neuter in gender in accordance with Article 30.1.3 of the Code (ICZN 1999). Latin or Latinised adjectival specific and subspecific names combined with it must, therefore, be declined accordingly (Article 31.2). Martínez (1947a), however, formed the binomen originally as Deltochilum abdominalis, with the two-termination New Latin adjective used as the specific name ‒ meaning ‘abdominal’ ‒ incorrectly declined in the masculine/feminine. We fix this in the heading above by declining the name in the neuter; i.e., by replacing the desinence ‘-is’ with ‘-e’ as grammatically required. Deltochilum abdominale – Roze 1955: 43 (checklist, cited for Venezuela). — Havranek 1989: 61 (list). — Medina et al. 2001: 136 (checklist for Colombia). — Krajcik 2012: 88 (list). Deltochilum (Deltohyboma) abdominale – Vulcano & Pereira 1964: 652 (catalogue). — Blanco 1987: 43 (catalogue). — González-Alvarado & Vaz-de-Mello 2021a: 62, 74 (revision, comments). — Martínez 1954: 47 (comments, cited for Colombia). Material examined Paratype VENEZUELA ‒ 1 ♂; “PARATIPO”; “VENEZUELA / D.F. 720 mts. / C° NAIGUATÁ / R. LICHYLEG. / Coll-Martínez. / Jul 943”; “Deltochilum (D.) / abdominalis ♂ / sp.n. / A. Martínez-DET.19 46.”; CEMT00068431. LOZANO DE LA ROSA C. et al., Dung beetles (Scarabaeinae) of Venezuela 73 Fig. 28. Species distribution. A. Deltochilum abdominale Martínez, 1947. B. Deltochilum bolivariense González-Alvarado & Vaz-de-Mello, 2021. Green diamond = CEMT collection data; black circle = literature data. European Journal of Taxonomy 959: 1–272 (2024) 80 Fig. 31. Species distribution. A. Deltochilum lindemannae Balthasar, 1967 B. Deltochilum nonstriatum González-Alvarado & Vaz-de-Mello, 2021. Black circle = literature data. LOZANO DE LA ROSA C. et al., Dung beetles (Scarabaeinae) of Venezuela 81 193 (inventory); 2014: 183 (list). — Price & Feer 2012: 327 (list for French Guiana). — FerrerParis et al. 2013: 109 (list). — Ratcliffe 2013: 493 (list, ecology). — Boilly 2015: 86–89 (key, fig. 4, comments). — Feer & Boissier 2015: 169 (list). — Hielkema & Hielkema 2019: 76 (catalogue for the Guianas). — González-Alvarado & Vaz-de-Mello 2021a: 58 (taxonomic revision). Material examined VENEZUELA ‒ Bolivar • 1 spec.; Anacoco; 6 Aug. 2006 15: 35; col. curso NM2006; faeces; CEMT. Distribution Venezuela, Guyana, Suriname, French Guiana, and Brazil. Subregions of Venezuela System of low mountains and hills Imataca-Cuyuní of Northeast Guyana Shield, and Guiana Shield. Literature records Ferrer-Paris et al. 2013: 109 (Venezuela: Bolívar: Isla de Anacoco). — González-Alvarado & Vaz-deMello 2021a: 58 (Venezuela: Bolívar). Deltochilum tenuistriatum González-Alvarado & Vaz-de-Mello, 2021 Fig. 32B Deltochilum tenuistriatum González-Alvarado & Vaz-de-Mello, 2021b: 102 (original description). Type locality: Venezuela: Bolívar: 10 km E of San Francisco Yuruani. Name-bearing type: holotype (CMNC), not examined. Distribution Venezuela (endemic). Subregion of Venezuela Guiana Shield. Literature record González-Alvarado & Vaz-de-Mello 2021b: 102, 104 (Venezuela: Bolívar). Genus Dendropaemon Perty, 1830 Dendropaemon Perty, 1830: 38 (original description). Type species: Eurysternus piceus Perty, 1830, by subsequent designation of Blut (1939). Dendropemon Agassiz, 1846: 119 (unjustified emendation of Dendropaemon Perty). Type species: Eurysternus piceus Perty, 1830, in accordance with Articles 33.2.3 and 67.8 of the Code (ICZN 1999) (see Cupello & Génier 2017). Dendropaemon – Lacordaire 1855: 102 (redescription). — d’Olsoufieff 1924: 19, 121 (key, redescription). — Blut 1939: 267 (redescription). — Pessôa & Lane 1941: 470 (key). — Martínez 1959: 106 (catalogue for Argentina). — Halffter & Matthews 1966: 258 (catalogue, distribution). — Vulcano & Pereira 1967: 566 (key). — Edmonds 1972: 843 (description); 1994: 17 (key). — Halffter & Edmonds 1982: 136 (catalogue, distribution). — Gámez & Mora 2000: 17 (list). — Medina & LoperaToro 2000: 301 (key). — Vítolo 2000: 593 (key); 2004: 292 (redescription). — Vaz-de-Mello 2000: 192 (checklist for Brazil). — Medina et al. 2001: 140 (checklist for Colombia). — Arnaud 2002b: European Journal of Taxonomy 959: 1–272 (2024) 82 Fig. 32. Species distribution. A. Deltochilum septemstriatum Paulian, 1938. B. Deltochilum tenuistriatum González-Alvarado & Vaz-de-Mello, 2021. Green diamond = CEMT collection data; black circle = literature data. LOZANO DE LA ROSA C. et al., Dung beetles (Scarabaeinae) of Venezuela 83 14 (key). — Hamel-Leigue et al. 2006: 17 (list); 2009: 59 (distribution for Bolivia). — Vaz-deMello et al. 2011a: 24 (key). — Carvajal et al. 2011: 142 (diagnosis). — Krajcik 2012: 89 (list). — Boilly & Vaz-de-Mello 2013: 107 (key). — Figueroa et al. 2014: 136 (distribution for Peru). — Boilly et al. 2016: 93 (list, comments). — Génier & Arnaud 2016: 6 (monograph). — Cupello & Génier 2017: 821–824 (revision). — Chamorro et al. 2018: 75, 94 (list for Ecuador); 2019: 10, 106 (catalogue). — Hielkema & Hielkema 2019: 98 (catalogue for the Guianas). Dendropemon – Harold 1869d: 1020 (catalogue, distribution). — Lucas 1920: 230 (nomenclator). — Gillet 1911b: 88 (catalogue). — Blackwelder 1944: 210 (list). Dendropaemon angustulus Génier & Arnaud, 2016 Fig. 33A Dendropaemon angustulus Génier & Arnaud, 2016: 26 (original description). Type locality: Venezuela: Bolívar: 26 km N Guasipati. Name-bearing type: holotype (CMNC), not examined. Distribution Venezuela, Guyana, French Guiana, and Brazil. Subregion of Venezuela System of hills and low piedmont mountains of the Guiana Shield. Literature record Génier & Arnaud 2016: 26 (Venezuela: Bolívar). Dendropaemon fredericki (Klages, 1906) Fig. 33B Eurypodea fredericki Klages 1906: 45 (original description). Type locality: Venezuela: Bolívar: Suapure, Caura River. Name-bearing type: holotype (CUIC) (Génier & Arnaud 2016), not examined. Tetramereia frederickii – Klages 1907: 141 (redescription). — Blut 1939: 298 (cited). — Edmonds 1972: 851 (nomenclatural comments). Eurypodea fredericki – Felsche 1908 (synonym). — d’Olsoufieff 1924: 120 (cited as synonym of Dendropaemon convexus). — Janssens 1940: 7 (contribution, cited as synonym). — Blackwelder 1944: 211 (checklist, cited as synonym of Eurypodea convexus). Dendropaemon fredericki – Gillet 1911b: 88 (catalogue, cited as synonym of Dendropaemon convexus). Dendropaemon (Eurypodea) fredericki – Génier & Arnaud 2016: 45, 86 (revision, key). — Hielkema & Hielkema 2019: 100 (catalogue for the Guianas). Distribution Colombia, Venezuela, Suriname, French Guiana, and Brazil. Subregions of Venezuela System of hills and low piedmont mountains of the Guiana Shield. Literature records Klages 1906: 45 (Venezuela: [Bolívar]: Suapure); 1907: 141(Venezuela: Bolívar: Suapure). — Gillet 1911b: 88 (Venezuela). — d’Olsoufieff 1924: 120 (Venezuela?). — Blackwelder 1944: 211 European Journal of Taxonomy 959: 1–272 (2024) 84 Fig. 33. Species distribution. A. Dendropaemon angustulus Génier & Arnaud, 2016. B. Dendropaemon fredericki (Klages, 1906). Black circle = literature data. LOZANO DE LA ROSA C. et al., Dung beetles (Scarabaeinae) of Venezuela 85 (Venezuela). — Arnaud 2002b: 16 (Venezuela: Bolívar: Suapure). — Noriega et al. 2008a: 133 (Venezuela). — Génier & Arnaud 2016: 45, 85 (Venezuela: Bolívar). Dendropaemon inemarginatus Génier & Arnaud, 2016 Fig. 34A Dendropaemon (Glaphyropaemon) inemarginatus Génier & Arnaud, 2016: 52 (original description). Type locality: Venezuela: Amazonas: Atures: Coromoto. Name-bearing type: holotype (CMNC), not examined. Dendropaemon (Glaphyropaemon) inemarginatus – Hielkema & Hielkema 2019: 100 (comments). non Dendropaemon (D.) refulgens (misidentification) – Martínez & Clavijo 1990: 155 (biology). Distribution Venezuela (endemic). Subregion of Venezuela System of hills and low piedmont mountains of the Guiana Shield. Literature records Martínez & Clavijo 1990: 155 (Venezuela: Amazonas: Atunes). — Génier & Arnaud 2016: 52 (Venezuela: Amazonas: Atures: Coromoto). Dendropaemon nigritulus Génier & Arnaud, 2016 Fig. 34B Dendropaemon (Nigropaemon) nigritulus Génier & Arnaud, 2016: 54 (original description). Type locality: Brazil: Roraima: Pacaraima, Surumu, Serra do Marari. Name-bearing type: holotype (CMNC), not examined. Dendropaemon (Nigropaemon) nigritulus – Boilly et al. 2016: 94 (list, comments). — Hielkema & Hielkema 2019: 100 (catalogue for the Guianas). Material examined VENEZUELA — Amazonas • 1 spec.; Atabapo, Cano Piojo; 130 m; Apr 2006; D.G. Fagre leg.; CEMT. Distribution Colombia, Venezuela, Guyana, Suriname, French Guiana, and Brazil. Subregions of Venezuela Peneplain of the Casiquiare River–Upper Orinoco, and System of hills and low piedmont mountains of the Guiana Shield. Literature record Génier & Arnaud 2016: 54, 86 (Venezuela: Bolívar). European Journal of Taxonomy 959: 1–272 (2024) 86 Fig. 34. Species distribution. A. Dendropaemon inemarginatus Génier & Arnaud, 2016. B. Dendropaemon nigritulus Génier & Arnaud, 2016. Black circle = literature data. LOZANO DE LA ROSA C. et al., Dung beetles (Scarabaeinae) of Venezuela 87 Genus Diabroctis Gistel, 1857 Diabroctis Gistel, 1857: 92 (original description). Type species: Scarabaeus mimas Linnaeus, 1758, by original designation. Taurocopris d’Olsoufieff, 1924: 18, 61 (original description). Type species: Scarabaeus mimas Linnaeus, 1758, by subsequent designation of Edmonds (1972). Taurocopris – Pessôa 1934: 282 (cited). — Janssens 1940: 2 (cited). — Pessôa & Lane 1941: 473 (key). — Blackwelder 1944: 208 (list). — Martínez 1959: 96 (catalogue for Argentina). — Martínez & Pereira 1967: 58 (nomenclatural comments). — Vulcano & Pereira 1967: 567 (key). Diabroctis – Martínez & Pereira 1967: 58 (nomenclatural comments). — Edmonds 1972: 820 (key, redescription). — Arnaud 1982: 115 (list of types). — Gámez & Mora 2000: 17 (list). — Medina & Lopera-Toro 2000: 300, 303 (key). — Vaz-de-Mello 2000: 186 (checklist for Brazil). — Vítolo 2000: 595 (key); 2004: 279 (list). — Medina et al. 2001: 134, 140 (list). — Arnaud 2002b: 14, 18 (key). — Gámez 2004: 51, 52 (distribution for Venezuela, key). — Hamel-Leigue et al. 2006: 3 (list); 2009: 60 (cited). — Noriega et al. 2007: 80 (list). — Vaz-de-Mello 2011a: 5, 10, 17, 24, 32, 39, 44 (list, key). — Ferrer-Paris et al. 2013: 96 (list). — Boilly et al. 2016: 94 (list, comments). — Valois et al. 2018: 351 (diagnosis). — Hielkema & Hielkema 2019:100 (catalogue for the Guianas). Diabroctis cadmus (Harold, 1868) Fig. 35A Phanaeus cadmus Harold, 1868e: 82 (original description). Type locality: French Guiana. Name-bearing type: lectotype (MNHN), designated by Arnaud (1982), not examined. Phanaeus mimaeformis Ancey, 1880: 205 (original description). Type locality: unknown; incorrectly stated as Costa Rica by Ancey (1880) based on mislabelled material, but the species does not occur there (Valois et al. 2018). Name-bearing type: lectotype (RBINS), designated by Valois et al. (2018), not examined. Phanaeus cadmus – Harold 1869d: 1017 (catalogue). — Nevinson 1892: 2 (list). — Heyne & Taschenberg 1908: 65 (list). — d’Olsoufieff 1924: 15 (comments). — Gillet 1911b: 81 (catalogue, cited for Colombia, Guianas, Costa Rica). Phanaeus mimaeformis – Bates 1887: 387 (cited for Costa Rica). — Nevinson 1892: 2 (as junior synonym of D. cadmus). — d’Olsoufieff 1924: 63 (cited as junior synonym of D. cadmus). Taurocopris cadmus – d’Olsoufieff 1924: 22, 62, 140 (key, comments, distribution). — Janssens 1940: 2, 4 (cited, key). — Blackwelder 1944: 208 (list, cited for Costa Rica). — Vulcano & Pereira 1967: 567 (key). Diabroctis cadmus – Edmonds 1972: 821 (cited). — Arnaud 1982: 115 (list of types, MNHN). — Martínez & Clavijo 1990: 151 (key, comments). — Escobar 2000: 208 (checklist for Colombia). — Gámez & Mora 2000: 17 (list). — Vítolo 2000: 595 (key); 2004: 279 (diagnosis, distribution). — Arnaud 2002b: 18 (key, distribution). — Gámez 2004: 48, 51, 52, 58 (list, distribution, key). — Noriega et al. 2007: 83 (list). — Giraldo et al. 2018: 22 (guide). — Valois et al. 2018: 351, 365 (key, diagnosis). — Hielkema & Hielkema 2019:100, 292 (catalogue for the Guianas). — Nieto et al. 2020: 136 (report). Material examined VENEZUELA ‒ Zulia • 1 spec.; Rosario de Perijá; 20 Jul. 2006; curso NM2006 leg.; faeces, 13h – 00:00; CEMT. European Journal of Taxonomy 959: 1–272 (2024) 88 Fig. 35. Species distribution. A. Diabroctis cadmus (Harold, 1868). B. Diabroctis venezuelensis Martínez & Clavijo, 1990. Green diamond = CEMT collection data; black circle = literature data. LOZANO DE LA ROSA C. et al., Dung beetles (Scarabaeinae) of Venezuela 89 Distribution Colombia and Venezuela. Subregions of Venezuela Maracaibo Depression, and System of hills and low sierras Lara-Falcón. Literature records d’Olsoufieff 1924: 62, 140 (Venezuela). — Roze 1955: 45 (Venezuela: Falcón: Buena Vista). — Martínez & Clavijo 1990: 5 (Venezuela: Zulia: Maracaibo, El Guaco). — Gámez & Mora 2000: 17 (Venezuela). — Arnaud 2002b: 18 (Venezuela: Zulia). — Gámez 2004: 58 (Venezuela: Zulia: Mara and El Guaco). — Vítolo 2004: 279 (Venezuela: Zulia). — Ferrer-Paris et al. 2013: 109 (Venezuela: Zulia: Rosario de Perijá). — Valois et al. 2018: 350, 365 (Venezuela: Zulia). Diabroctis venezuelensis Martínez & Clavijo, 1990 Fig. 35B Diabroctis mimas venezuelensis Martínez & Clavijo, 1990: 147 (original description). Type locality: Venezuela: Portuguesa: Araure. Name-bearing type: holotype (MIZA) (Valois et al. 2018), examined from photos by FZVM. Diabroctis mimas venezuelensis –Arnaud 2002b: 18 (cited as Diabroctis mimas venezuelina Martínez & Clavijo, 1990, cited as synonym of Diabroctis mimas). — Gámez 2004: 48, 51–52, 58 (key, list). — Hamel-Leigue et al. 2009: 60 (cited as synonym of Diabroctis mimas). Diabroctis venezuelensis – Valois et al. 2018: 357 (revision). — Hielkema & Hielkema 2019:100, 292 (catalogue for the Guianas). Material examined VENEZUELA ‒ Bolívar • 1 ♂, 2 ♀♀; near Río Paraguaz, 100 m a.s.l., 15 May 2004; CEMT. ‒ Portuguesa • 1 ♂; Araure; 1978; CEMT. ‒ Cojedes • 1 ♀; Tinaquillo; 15 May 1978; CEMT. Distribution Venezuela (endemic). Subregions of Venezuela Plains, System of hills and low piedmont mountains of the Guiana Shield, Andes mountains, and Central Coast Mountain Range. Literature records Martínez & Clavijo 1990: 147 (Venezuela: Cojedes; Portuguesa: Araure; Lara [actually, Yaracuy]: Sabana de Parra). — Arnaud 2002b: 18 (Venezuela). — Gámez 2004: 48 (subregion of Llanos). — Valois et al. 2018: 357–358 (Venezuela: Cojedes, Bolívar and Portuguesa). Genus Dichotomius Hope, 1838 Dichotomius Hope, 1838: 321 (original description). Type species: Scarabaeus boreus Olivier, 1789, by original designation; see Martínez (1951). Pinotus Erichson, 1847a: 108 (original description). Type species: uncertain. Usually claimed to be Scarabaeus carolinus Linnaeus, 1767 (e.g., Lacordaire 1855; Luederwaldt 1929; Nunes & Vazde-Mello 2019), but, as argued by Martínez (1951), this nominal species was not among the two European Journal of Taxonomy 959: 1–272 (2024) 96 Fig. 38. Species distribution. A. Dichotomius costaricensis (Luederwaldt. 1935). B. Dichotomius deyrollei (Harold, 1869). Green diamond = CEMT collection data; black circle = literature data. LOZANO DE LA ROSA C. et al., Dung beetles (Scarabaeinae) of Venezuela 97 Literature records Kohlmann & Solís 1997: 360 (Venezuela: Falcón). — Solís & Kohlmann 2012: 12 (Venezuela). Dichotomius deyrollei (Harold, 1869) Fig. 38B Pinotus deyrollei Harold, 1869c: 139 (original description). Type locality: Brazil. Name-bearing type: lectotype (MNHN), designated by Montoya-Molina & Vaz-de-Mello (2021), examined by FZVM. Pinotus deyrollei – Luederwaldt 1929: 99 (key, cited for Trinidad Island). — Blackwelder 1944: 206 (list, cited for Brazil). Dichotomius (Luederwaldtinia) deyrollei – Roze 1955: 44 (checklist for Venezuela). — Vulcano & Pereira 1967: 586 (key). — Blanco 1988: 40–41 (catalogue, comments). — Escobar 2000: 208 (checklist for Colombia). — Medina et al. 2001: 138 (checklist for Colombia). — Noriega 2004: 39 (checklist for Tinigua Park, Colombia). — Medina & Pulido-Herrera 2009: 60 (diversity). — Sarmiento-Garcés & Amat-García 2014: 89, 91 (key, diagnosis, distribution for Colombia). — Pardo-Díaz et al. 2019: 6, 13 (comments). — Montoya-Molina & Vaz-de-Mello 2021: 7, 22 (key, diagnosis, distribution, redescription). Material examined VENEZUELA ‒ Mérida • 2 specs; Arzobispo Chacón, Mucutuy environs; 08°14′18″ N, 71°17′12″ W; 1380 m a.s.l.; 13 Mar. 2015; R. Acconcia leg.; light; CEMT. ‒ Táchira • 2 specs; Río Negro; 25 Mar. 1984; Joffre B. leg.; CEMT • 1 spec.; 42 km SE of San Cristóbal; 700 m a.s.l.; 19 May 1974; S. Peck leg.; dung trap 30; CEMT • 4 specs; San Cristóbal, Parque Nacional Natural Paramillo; Jul. 1993; J. Blanco leg.; CEMT. ‒ Yaracuy • 1 spec.; Bolívar, Aroa; 10°0′0″ N, 68°0′0″ W; 710 m a.s.l.; 20 Jul. 2009; M. Asmüssen, P. Colmenares and H. Martínez leg.; human faeces; CEMT • 1 spec.; Bolívar, Aroa; 10°22′8.21″ N, 68°50′18.52″ W; 1626 m a.s.l.; 21 Jul. 2009; M. Asmüssen, P. Colmenares and H. Martínez leg.; human faeces; CEMT • 1 spec.; Bolívar, Aroa; 10°20′21.98″ N, 68°50′6.03″ W; 1362 m a.s.l.; 19 Jul. 2009; M. Asmüssen, P. Colmenares and H. Martínez leg.; human faeces; CEMT. Distribution Colombia, Venezuela, and Trinidad and Tobago. Subregions of Venezuela Andes mountains and Central Coast Mountain Range. Literature records Roze 1955: 44 (Venezuela: Distrito Capital: Caracas). — Vulcano & Pereira 1967: 586 (Venezuela); Blanco 1988: 40–41 (Venezuela: Táchira: Río Negro: Parque Nacional El Tamá); Montoya-Molina & Vaz-de-Mello 2021: 7, 22 (Venezuela: Mérida, Miranda, Táchira and Yaracuy). Dichotomius fallax (Harold, 1880) Fig. 39A Pinotus fallax Harold, 1880: 26 (original description). Type locality: Colombia: Cundinamarca: Tequendama: La Mesa; and Venezuela: Distrito Capital: Caracas. Name-bearing type: syntypes (MNHN), examined by FZVM. Pinotus (Selenocopris) fallax – Luederwaldt 1929: 64 (key). European Journal of Taxonomy 959: 1–272 (2024) 98 Pinotus fallax – Gillet 1911b: 60 (catalogue). — Gacharná 1951: 222 (list for Colombia). — Balthasar 1941: 349 (list, characteristics); 1951: 334 (list). — Blackwelder 1944: 207 (checklist). Dichotomius fallax – Roze 1955: 44 (cited, checklist for Venezuela). — Vulcano & Pereira 1967: 587 (key). — Medina et al. 2001: 138 (checklist for Colombia). — Larsen et al. 2008: 1294 (list). Material examined VENEZUELA ‒ Bolívar • 1 spec.; El Manteco; 5 Aug. 2006; curso NM2006 leg.; heces, 24h, 08:00; CEMT • 2 specs; Campo Minero, Río Yuruani, El Dorado; 190 m; 23‒30 May 1987; CEMT • 1 spec.; Las Piñas, 30 km E of Paragua; 12 Apr. 1983; C. Bordon leg.; CEMT. ‒ Yaracuy • 1 spec.; Bolívar, Aroa; 10°0′0″ N, 68°0′0″ W; 483 m; 19 Jul. 2009; M. Assmüssen, P Colmenares and H. Martínez leg.; human faeces; CEMT. ‒ Zulia • 1 spec.; Rosario de Perijá; 18 Jul. 2006; curso NM2006 leg.; faeces, 23h, 13:09; CEMT. Distribution Colombia, Venezuela, Trinidad and Tobago (new country record, CEMT), Brazil (new country record, CEMT) and Peru. Subregions of Venezuela Maracaibo Depression, System of hills and low piedmont mountains of the Guiana Shield, System of low mountains and hills Imataca-Cuyuní del Escudo Northeast Guiana, and Central Coast Mountain Range. Literature records Harold 1880: 26 (Venezuela: Caracas). — Gillet 1911b: 60 (Venezuela). — Luederwaldt 1929: 64 (Venezuela). — Balthasar 1941: 349 (Venezuela); 1951: 334 (Venezuela).— Blackwelder 1944: 207 (Venezuela). — Vulcano & Pereira 1967: 587 (Venezuela). — Larsen et al. 2008: 1294 (Venezuela: Bolívar: Lago Guri). Dichotomius henripittieri Montoya-Molina & Vaz-de-Mello, 2021 Fig. 39B‒D Dichotomius (Luederwaldtinia) henripittieri Montoya-Molina & Vaz-de-Mello, 2021: 31 (original description). Type locality: Venezuela: Aragua: Mario Briceño Iragorry: Parque Nacional Henri Pittier: Estación Biológica Periquito. Name-bearing type: holotype (CEMT), examined by CL and FZVM. Material examined Holotype VENEZUELA ‒ Aragua • ♂; “Mario / Briceño Iragorry, PN. Henri / Pittier, Est. Bio. Rancho Grande, / way to Pico Periquito, 10°20’ / 51”N, 67°41’17”W, 1145m, / human feces; 18–20, viii.2010, / R. Acconcia & J. Gámez”; CEMT. Paratypes VENEZUELA • 3 ♂♂, 3 ♀♀; same data as for holotype; CEMT. Distribution Venezuela (endemic). LOZANO DE LA ROSA C. et al., Dung beetles (Scarabaeinae) of Venezuela 99 Fig. 39. Species distribution. A. Dichotomius fallax (Harold, 1880). B. Dichotomius henripittieri Montoya-Molina & Vaz-de-Mello, 2021. C. Male of D. henripittieri. D. Female of D. henripittieri. Green diamond = CEMT collection data; black circle = literature data. European Journal of Taxonomy 959: 1–272 (2024) 100 Subregion of Venezuela Central Coast Mountain Range. Literature record Montoya-Molina & Vaz-de-Mello 2021: 31 (Venezuela: Aragua). Dichotomius inachoides (Felsche, 1901) Fig. 40A Pinotus inachoides Felsche, 1901: 144 (original description). Type locality: Venezuela: Mérida. Namebearing type: holotype (SMTD), examined by FZVM. Pinotus inachoides – Gillet 1911b: 61 (catalogue). — Blackwelder 1944: 207 (list). Pinotus (Cephagonus) inachoides – Luederwaldt 1929: 109 (key, characteristics). Dichotomius (Luederwaldtinia) inachoides – Roze 1955: 44 (cited, checklist for Venezuela). — Vulcano & Pereira 1967: 585 (key for the Amazon). — Krajcik 2012: 91 (list). — Montoya-Molina & Vaz-de-Mello 2021: 7, 33 (key, diagnosis, distribution). Material examined VENEZUELA ‒ Aragua • 1 spec.; Cuyagua; 50 m a.s.l.; 6 Jul. 2004; D. García leg.; CEMT. ‒ Barinas • 7 specs; Bolívar, Barinitas, Paguey, Sector El Cacao, Andean foothills; 1080 m a.s.l.; 3‒4 Sep. 2015; R. Acconcia and J. Gámez leg.; CEMT. ‒ Merida • 1 ♂; Monte Zerpa, La Hechicera; Nov. 1987; Joffre B. leg.; CEMT • 1 ♀; Santa Rosa, La Hechicera; Jun. 1986; Joffre B. leg.; CEMT • 2 specs; Sucre, Jají; 8°40′20.49″ N, 71°24′38.59″ W; 1999 m a.s.l.; 8 Jul. 2009; D. Mora, P. Colmenares, M.Córdova and M. Nuñez leg.; human faeces; CEMT. ‒ Táchira • 2 ♀♀; “La Flautera Palmira” (?); 15 Jul. 1985; Joffre B. leg.; CEMT. Distribution Colombia and Venezuela. Subregions of Venezuela Coastal mainland, Plains, Andes mountains and Oriental Coast Range. Literature records Gillet 1911b: 61 (Venezuela). — Luederwaldt 1929: 109 (Venezuela). — Blackwelder 1944: 207 (Venezuela). — Vulcano & Pereira 1967: 585 (Venezuela). — Krajcik 2012: 91 (Venezuela). — MontoyaMolina & Vaz-de-Mello 2021: 7, 33 (Venezuela: Aragua; Barinas; Mérida; Sucre and Táchira). Dichotomius latilobatus Boilly & Vaz-de-Mello, 2021 Fig. 40B Dichotomius latilobatus Boilly & Vaz-de-Mello, 2021: 22 (original description). Type locality: French Guiana: Saint-Laurent-du-Maroni: Enéné Patatpe, Mitaraka Massif. Name-bearing type: holotype (MNHN), examined by FZVM. Material examined VENEZUELA ‒ Bolívar • 1 spec.; Isla de Anacoco; 6 Aug. 2006; curso NM2006 leg.; faeces, 23h; CEMT. LOZANO DE LA ROSA C. et al., Dung beetles (Scarabaeinae) of Venezuela 101 Fig. 40. Species distribution. A. Dichotomius inachoides (Felsche, 1901). B. Dichotomius latilobatus Boilly & Vaz-de-Mello, 2021. Green diamond = CEMT collection data; black circle = literature data. European Journal of Taxonomy 959: 1–272 (2024) 102 Distribution Venezuela, Suriname, French Guiana, and Brazil. Subregions of Venezuela System of hills and low piedmont mountains of the Guiana Shield, System of low mountains and hills Imataca-Cuyuní of Northeast Guyana Shield, and Guiana Shield. Literature record Boilly & Vaz-de-Mello 2021: 22 (Venezuela: Bolívar). Dichotomius mamillatus (Felsche, 1901) Fig. 41A Pinotus mamillatus Felsche, 1901: 143 (original description). Type locality: Ecuador. Name-bearing type: lectotype (SMTD), designated by Rossini & Vaz-de-Mello (2020), examined by FZVM. Pinotus calcaratus Arrow, 1913: 457 (original description). Type locality: Brazil: Rondônia: Madeira River, “Madeira-Mamoré Railway Company Camp 41” (“Matto Grosso”, “Madeira-Mamore / R.R.Co.Camp 41”). Name-bearing type: holotype (BMNH) (Rossini & Vaz-de-Mello 2020), examined by FZVM. Pinotus mamillatus – Gillet 1911b: 61 (catalogue). — Balthasar 1941: 349 (checklist); 1951: 334 (checklist). — Blackwelder 1944: 207 (checklist). — Sarmiento-Garcés & Amat-García 2009: 291 (cited). — Rossini & Vaz-de-Mello 2020: 60 (cited). Pinotus calcaratus – Luederwaldt 1929: 97, 102–103 (key, diagnosis); 1936: 213 (cited). — Rossini & Vaz-de-Mello 2020: 60 (cited as synonym of Dichotomius mamillatus). Dichotomius (Dichotomius) mamillatus – Vulcano & Pereira 1967: 585 (key, synopsis). — Luederwaldt 1929: 53 (key). — Amézquita et al. 1999: 119–120, 123 (list for Orinoquía, Colombia). — Escobar 2000: 208 (checklist for Colombia). — Medina & Lopera-Toro 2000: 309 (key for the Amazon). — Vaz-de-Mello 2000: 193 (checklist for Colombia). — Medina et al. 2001: 138 (checklist for Colombia). — Pulido-Herrera et al. 2003: 54 (list for Caqueta, Colombia); 2007: 307 (list for Andean Region Colombia). — Celi et al. 2004: 44 (list for Morona Santiago, Ecuador). — Noriega-Alvarado 2004: 39 (list for La Macarena, Colombia). — Noriega & Botero-Trujillo 2008: 451 (comments). — Noriega et al. 2008b: 79 (ecology). — Carpio et al. 2009: 469 (ecology). — Medina & PulidoHerrera 2009: 60 (list for Orinoquia, Colombia). — Sarmiento-Garcés & Amat-García 2009: 287, 291, 295 (diagnosis, key). — Larsen 2011: 98, 102 (list for Suriname); 2013: 98 (list for Suriname). — Vaz-de-Mello et al. 2011b: 87 (list). — Krajcik 2012: 91 (checklist). — Noriega & Navarrete-Heredia 2013: 191(comments). — Ratcliffe 2013: 493, 504, 510–511 (fig. 48, cited as Dichotomius nr. mamillatus). — Martínez et al. 2014: 17–18 (comments). — Silva et al. 2014: 349 (ecology); 2015: 611 (list for Mato Grosso state, Brazil); 2016: 554 (ecology, biogeography). — Ratcliffe et al. 2015: 196 (checklist for Peru). — Chamorro et al. 2018: 89 (key for Ecuador); 2019: 114–115 (catalogue). — Giraldo et al. 2018: 61 (guide). — Santos et al. 2018: 46 (list for Acre, Brazil). — Rossini & Vaz-de-Mello 2020: 59–61 (key, distribution, diagnosis, redescription). Dichotomius (Dichotomius) calcaratus – Vulcano et al. 1976: 517 (new combination). — Vaz-de-Mello 2000: 193 (checklist for Brazil). — Rossini & Vaz-de-Mello 2020: 60 (as synonym of Dichotomius mamillatus). Dichotomius (Dichotomius) mamillatus – Vaz-de-Mello 2000: 193 (checklist for Colombia). — Vazde-Mello et al. 2011b: 87 (list). — Chamorro et al. 2018: 89 (key for Ecuador); 2019: 114–115 (catalogue). — Rossini & Vaz-de-Mello 2020: 59–61 (key, distribution, diagnosis, redescription). LOZANO DE LA ROSA C. et al., Dung beetles (Scarabaeinae) of Venezuela 103 Fig. 41. Species distribution. A. Dichotomius mamillatus (Felsche, 1901). B. Dichotomius nisus (Olivier, 1789). Green diamond = CEMT collection data; black circle = literature data. European Journal of Taxonomy 959: 1–272 (2024) 104 Distribution Colombia, Venezuela, Guyana, Suriname, French Guiana, Brazil, Ecuador, Peru, and Bolivia. Subregions of Venezuela Peneplain of the Casiquiare River–Upper Orinoco, System of hills and low piedmont mountains of the Guiana Shield, and Guiana Shield. Literature record Rossini & Vaz-de-Mello 2020: 60 (Venezuela: Amazonas and Bolívar). Dichotomius nisus (Olivier, 1789) Fig. 41B Scarabaeus nisus Olivier, 1789: 139 (original description). Type locality: French Guiana. Name-bearing type: lectotype (MNHN), designated by Cassenote et al. (2020), examined by FZVM. Pinotus garbei Luederwaldt, 1922: 11 (original description). Type locality: French Guiana: Cayenne. Name-bearing type: lectotype (MZSP), designated by Cassenote et al. (2020), examined by FZVM. Pinotus taunay var. pilosus Luederwaldt, 1931b: 303 (original description). Type locality: Brazil: São Paulo: “Alto da Serra”. Name-bearing type: holotype by monotypy (MZSP), not examined. Copris nisus – Olivier 1790: 166 (list, characteristics). — Fabricius 1801: 44 (list, comments). — Nunes & Vaz-de-Mello 2019: 2239 (comments). Copris (Selenocopris) nisus – Burmeister 1846: unpaged [p. 93 of 220] (key). Pinotus nisus – Harold 1869c: 141 (list, revision). — Candèze 1891: 330 (list). — Gillet 1911b: 61 (catalogue, cited for Guyana, Brazil, Argentina). — Luederwaldt 1914: 369 (comments); 1929: 84 (key, comments, cited for Brazil). — Blackwelder 1944: 207 (list, distribution). Pinotus garbei – Luederwaldt 1929: 84 (synonym of D. nisus). Dichotomius (Luederwaldtinia) nisus – Martínez 1951: 140 (comments). — Roze 1955: 44 (checklist for Venezuela). — Halffter & Matthews 1966: 98 (comments). — Blanco 1988: 40–41 (catalogue, comments). — Escobar 2000: 208 (checklist for Colombia). — Vaz-de-Mello 2000: 193 (checklist for Brazil). — Krajcik 2006: 54 (list); 2012: 91 (list, cited for “Cayenne”). — Pulido-Herrera et al. 2007: 307 (cited for Andean region of Colombia). — Feer 2008: 56, 62 (ecology). — Larsen et al. 2008: 1294 (list). — Medina & Pulido-Herrera 2009: 60 (diversity). — Lozano 2010: 86 (list). — Ferrer-Paris et al. 2013: 109 (list). — Nunes & Vaz-de-Mello 2013: 411, 413, 416–418 (comments, fig. 2, key); 2019: 2238, 2240 (comments). — França et al. 2016: 2, 4 (list, comments). — Silva et al. 2016: 7 (ecology, biogeography). — Giraldo et al. 2018: 68 (guide). — Hielkema & Hielkema 2019: 55 (catalogue for the Guianas). — Pardo-Díaz et al. 2019: 1, 5, 7–8, 12–14 (comments). — Nieto et al. 2020: 136 (report). Scarabaeus nisus – Nunes & Vaz-de-Mello 2019: 2237, 2239 (comments). Dichotomius (Selenocopris) nisus – Cassenote et al. 2020: 2–6 (distribution, diagnosis). Material examined VENEZUELA ‒ Aragua • 4 specs; Cuyagua; 50 m a.s.l.; 6 Jul. 2004; D. García leg.; CEMT. ‒ Bolívar • 1 ♀; Rio Pargaza; 15 May 2004; D. Garcia leg.; CEMT. ‒ Táchira • 1 ♂, 1 ♀; La Fría, Hacienda La Libertad; 21 Jul. 1984; Joffre B. leg.; CEMT. Distribution Colombia, Venezuela, French Guiana, Brazil, Bolivia, Paraguay, Argentina, and Uruguay. LOZANO DE LA ROSA C. et al., Dung beetles (Scarabaeinae) of Venezuela 105 Subregions of Venezuela Coastal island, coastal mainland, Maracaibo Depression, System of hills and low sierras Lara-Falcón, System of hills and low piedmont mountains of the Guiana Shield, Andes mountains and Central Coast Mountain Range. Literature records Candèze 1891: 330 (Venezuela: Falcón: Corosal; and Carabobo: San-Esteban). — Roze 1955: 44 (Venezuela: Distrito Capital: Caracas; Aragua; and Nueva Esparta: Isla Margarita). — Blanco 1988: 40–41 (Venezuela: Táchira). — Larsen et al. 2008: 1294 (Venezuela: Bolívar: Lago Guri). — Lozano 2010: 86 (Venezuela: Zulia). — Ferrer-Paris et al. 2013: 109 (Venezuela: Aragua [Guárico]: Altagracia de Orituco). — França et al. 2016: 4 (Venezuelan dry forests and pastures). — Cassenote et al. 2020: 2 (Venezuela: Bolívar). Dichotomius ocellatopunctatus (Felsche, 1901) Pinotus ocellatopunctatus Felsche, 1901: 141 (original description). Type locality: Venezuela: Mérida. Name-bearing type: lectotype (SMTD), designated by Vaz-de-Mello & Nunes (2016), examined by FZVM. Pinotus ocellatopunctatus – Gillet 1911b: 61 (catalogue). — Luederwaldt 1929: 74 (key). — Blackwelder 1944: 207 (list). — Pereira 1947b: 318, 319 (key, comments). Dichotomius ocellatopunctatus – Roze 1955: 44 (cited, checklist for Venezuela). — Vulcano & Pereira 1967: 586 (key). — Krajcik 2012: 91 (list). — Vaz-de-Mello & Nunes 2016: 297 (redescription, new combination). — Valois et al. 2022: 119‒120, 123 (revision, key). Material examined Paralectotype VENEZUELA • 1 ♀; “PARALECTOTYPE / Pinotus ♀/ ocellatopunctatus / Felsche / des. F.Z. Vaz-deMello, 2014 [yellow label]”; “staatl. Museum für / Tierkunde Dresden”; “Coll. C. Felsche / Kauf 20, 1918 [green label]”; “Merida [white label]”; CEMT-00078242. Distribution Venezuela (endemic). Literature records Gillet 1911b: 61 (Venezuela). — Luederwaldt 1929: 74 (Venezuela). — Blackwelder 1944: 207 (Venezuela). — Vulcano & Pereira 1967: 586 (Venezuela). — Krajcik 2012: 91 (Venezuela). — Vaz-deMello & Nunes 2016: 297 (Venezuela: Mérida). — Valois et al. 2022: 123 (Venezuela). Dichotomius protectus (Harold, 1867) Fig. 42A Pinotus protectus Harold, 1867d: 98 (original description). Type locality: Colombia. Name-bearing type: syntypes (MNHN), examined by FZVM. Pinotus protectus tridentatus Luederwaldt, 1923: 10 (original description). Type locality: Ecuador and Venezuela. Name-bearing type: syntypes (MZSP), not examined. Pinotus protectus – Harold 1869b: 130 (redescription, cited for Colombia); 1869d: 1010 (catalogue, cited for Colombia). — Candèze 1891: 330 (list). — Heyne & Taschenberg 1908: 62 (list, cited for European Journal of Taxonomy 959: 1–272 (2024) 112 Fig. 44. Species distribution. A. Dichotomius validipilosus (Luederwaldt, 1931). B. Dichotomius worontzowi (Pereira, 1942). Green diamond = CEMT collection data; black circle = literature data. LOZANO DE LA ROSA C. et al., Dung beetles (Scarabaeinae) of Venezuela 113 2001: 138 (checklist for Colombia). — Pulido-Herrera et al. 2003: 54 (list for Caqueta, Colombia); 2007: 307 (list for Andean region, Colombia). — Celi et al. 2004: 44 (diversity). — NoriegaAlvarado 2004: 39 (list for La Macarena, Colombia). — Hamel-Leigue et al. 2006: 16 (catalogue for Bolivia). — Medina & Pulido-Herrera 2009: 60 (list for Orinoquia, Colombia). — SarmientoGarcés & Amat-García 2009: 287, 291, 294 (diagnosis, key). — Quintero & Halffter 2009: 649 (ecology, list). — Larsen 2013: 96, 100 (list for Suriname). — Nunes et al. 2014: 408–411 (cited, list). — Silva et al. 2014: 349 (ecology); 2016: 555 (ecology, biogeography, list); 2017: 490 (list). — Feer & Boissier 2015: 169 (list). — Ratcliffe et al. 2015: 196 (checklist for Peru). — Silva et al. 2015: 611 (list for Mato Grosso, Brazil). Distribution Colombia, Venezuela, Guyana, Suriname, French Guiana, Brazil, Peru, and Bolivia. Subregions of Venezuela System of hills and low piedmont mountains of the Guiana Shield, and Guiana Shield. Literature record Rossini & Vaz-de-Mello 2020: 65 (Venezuela: Bolívar). Genus Digitonthophagus Balthasar, 1959 Onthophagus (Digitonthophagus) Balthasar, 1959: 464 (original description). Type species: Scarabaeus bonasus Fabricius, 1775, by original designation. Digitonthophagus — Zunino 1981: 413 (new status). — Huchet 1992: 300 (key). — Kohlmann 1994: 35‒41 (distribution for Mexico). — Ripa et al. 1995: 429 (list). — Lobo & Montes de Oca 1997: 17‒32 (comments). — Montes de Oca & Halffter 1998: 37‒45 (comments, distribution). — Koller et al. 1999: 403‒404 (comments); 2007: 84 (comments). — Aidar et al. 2000: 817 (comments). — Vaz-de-Mello 2000: 193 (checklist for Brazil). — Marchiori et al. 2003: 174–175 (comments, list). — Noriega et al. 2006: 379‒381 (continental record for Colombia); 2007: 80 (list for Colombia); 2010: 451‒456 (cited for Peru); 2011: 35‒40 (revision); 2017: 371 (distribution); 2020: 3 (distribution). — Matavelli & Louzada 2008: 153‒158 (intra-Amazonian distribution). — Álvarez et al. 2009: 373‒376 (cited for Argentina). — Lozano 2010: 86 (list). — Arias-Buriticá et al. 2011: 875‒878 (cited). — Vaz-de-Mello et al. 2011a: 5, 9, 16, 23, 31, 38, 58 (key). — Ferrer-Paris 2014: 41‒46 (distribution for Venezuela, comments). — Génier & Krell 2017: 79 (nomenclature). — Hielkema & Hielkema 2019: 89 (catalogue for the Guianas). — Gámez et al. 2021: 46–50 (short note). Digitonthophagus gazella (Fabricius, 1787) Fig. 45A Scarabaeus gazella Fabricius, 1787b: 377 (original description). Type locality: Zimbabwe: Manicaland: Mutare: Mutare (formerly, Umtali). Name-bearing type: neotype (ZMUK), designated by ICZN (2020), not examined. Scarabaeus dorcas Olivier, 1789: 121 (original description). Type locality: Madagascar: Diana: Antsiranana (formerly, Diego-Suárez). Name-bearing type: neotype (MNHN), designated by Génier & Moretto (2017), not examined. Scarabaeus gazella – Fabricius 1792: 56 (list, characteristics). Copris gazella – Fabricius 1801: 47 (list). European Journal of Taxonomy 959: 1–272 (2024) 114 Onthophagus gazella – Dejean 1821: 53 (catalogue). — Blume & Aga 1978: 190‒191 (comments). — Fincher et al. 1983: 159‒162 (distribution). — Nascimento et al. 1990: 1‒5 (cited). — Ripa et al. 1995: 429 (list). — Maes 1998: 654 (list for Nicaragua). — Miranda et al. 1998: 681–685 (contribution). — Koller et al. 1999: 404, 405 (comments). — Larsen et al. 2008: 1294 (list). — Génier & Krell 2017: 78 (nomenclature). Onthophagus (Digitonthophagus) gazella – Zunino 1979: 292 (cited for Saudi Arabia). — Maes et al. 1997: 44 (list for Nicaragua). — Bianchin et al. 1998: 276‒278 (comments). Digitonthophagus gazella – Zunino 1981: 413‒414 (comments). — Barbero & López-Guerrero 1992: 115‒120 (contribution). — Deloya 1992: 2 (list). — Huchet 1992: 300‒302 (cited for Antillas). — Kohlmann 1994: 35‒41 (distribution for Mexico). — Montes de Oca & Halffter 1995: 167 (ecology); 1998: 37‒45 (comments, distribution). — Lobo & Montes de Oca 1997: 17‒32 (comments). — Koller et al. 1999: 405‒407 (list); 2007: 81‒83, 85‒86 (comments, list). — Aidar et al. 2000: 818‒819 (comments). — Vaz-de-Mello 2000: 193 (checklist for Brazil). — NavarreteHeredia et al. 2001: 56 (list). — Estrada & Coates-Estrada 2002: 1911 (ecology). — Noriega 2002b: 213‒215 (cited for Colombia). — Marchiori et al. 2003: 174‒175 (comments, list). — Noriega et al. 2006: 379‒381 (cited for Colombia); 2007: 82 (list for Colombia); 2010: 451‒456 (cited for Peru); 2011: 35‒40 (revision); 2017: 371 (distribution); 2020: 3 (distribution). — Matavelli & Louzada 2008: 153‒158 (distribution intra-Amazonian). — Vidaurre et al. 2008: 217‒220 (cited for Bolivia). — Álvarez et al. 2009: 373‒376 (cited for Argentina). — Medina & Pulido-Herrera 2009: 61 (diversity). — Lozano 2010: 86 (list). — Arias-Buriticá et al. 2011: 875‒878 (cited). — Vaz-de-Mello et al. 2011b: 88 (list). — Delgado et al. 2012: 327 (list). — Ferrer-Paris 2014: 42, 45 (distribution for Venezuela). — Peck et al. 2014: 43 (diversity). — Silva et al. 2014: 349 (diversity); 2017: 490 (list). — Génier & Moretto 2017: 39 (taxonomy, systematics, phylogeny). — Génier & Davis 2017: 497‒500 (revision). — Génier & Krell 2017: 78‒82 (revision, diagnosis). — Giraldo et al. 2018: 36 (guide). — Hielkema & Hielkema 2019: 89 (catalogue for the Guianas). — Gámez et al. 2021: 46‒50 (distribution). — Nieto et al. 2020: 136 (report). Material examined VENEZUELA ‒ Yaracuy • 1 spec.; Bolívar, Aroa; 10°0′0″ N, 68°0′0″ W; 459 m a.s.l.; 19 Aug. 2009; M. Asmüssen, P. Colmenares and H. Martínez leg.; human faeces; CEMT. ‒ Zulia • 1 spec.; Rosario de Perijá; 20 Jul. 2006; curso NM2006 leg.; faeces, 00:00, 14h; CEMT. Distribution Native to continental Africa. Introduced and naturalized all over the the Americas, in Australia, and on several islands worldwide (Génier & Moretto 2017). Subregions of Venezuela Maracaibo Depression, Plains, System of low mountains and hills Imataca-Cuyuní of Northeast Guiana Shield, Andes mountains, and Central Coast Mountain Range. Literature records Larsen et al. 2008 (Venezuela: Bolívar: Lago Guri). — Lozano 2010: 86 (Venezuela: Zulia). — Noriega et al. 2010: 452 (Venezuela); 2011: 34–36 (Venezuela). — Ferrer-Paris et al. 2013: 109 (Venezuela: Aragua [Guárico]: Altagracia de Orituco and Zulia: Rosario de Perijá). — Ferrer-Paris 2014: 41–46 (Venezuela: Aragua and Zulia). — Génier & Davis 2017b: 497 (Venezuela). — Génier & Krell 2017: 79 (Venezuela: Ferrer-Paris 2014). — Noriega et al. 2017: 371 (Venezuela: Apure; Bolívar; Cojedes; Guárico; and Táchira); 2020: 3 (Venezuela). — Gámez et al. 2021: 46–50 (Venezuela: Apure; Barinas; Mérida; and Zulia). LOZANO DE LA ROSA C. et al., Dung beetles (Scarabaeinae) of Venezuela 115 Fig. 45. Species distribution. A. Digitonthophagus gazella (Fabricius, 1787). B. Eurysternus atrosericus Génier, 2009. Green diamond = CEMT collection data; black circle = literature data. European Journal of Taxonomy 959: 1–272 (2024) 116 Genus Eurysternus Dalman, 1824 Eurysternus Dalman, 1824: 8 (original description). Type species: Eurysternus planus Dalman, 1854, by subsequent designation of Jessop (1985). Aeschrotes Le Peletier de Saint-Fargeau & Audinet-Serville, 1828: 357 (original description). Type species: Aeschrotes planus Dalman, 1824, by subsequent designation of Jessop (1985). Eurysternodes Martínez, 1988b: 281 (original description), junior homonym of Eurysternodes Schuster & Summers, 1978 (Acari: Diarthrophallidae). Type species: Eurysternus velutinus Bates, 1887, by original designation. Pareurysternus Martínez, 1988b: 282 (original description). Type species: Eurysternus navajasi Martínez, 1988, by original designation. Tropicalia Koçak & Kemal, 2008: 7 (new replacement name for Eurysternodes Martínez). Type species: Eurysternus velutinus Bates, 1887, in accordance with Article 72.7 of the Code (ICZN 1999). Amartinezus Ozdikmen, 2009: 143 (new replacement name for Eurysternodes Martínez). Type species: Eurysternus velutinus Bates, 1887, in accordance with Article 72.7 of the Code (ICZN 1999). Eurysternus – Latreille 1829: 535 (redescription). — Castelnau 1840: 92 (redescription). — Agassiz 1846: 436 (catalogue). — Lacordaire 1855: 106 (redescription). — Harold 1869d: 1023 (catalogue). — Bruch 1911: 182 (list). — Gillet 1911b: 25 (catalogue). — Lucas 1920: 289 (catalogue, distribution). — Paulian 1938: 232 (key). — Pessôa & Lane 1941: 406 (redescription). — Blackwelder 1944: 197 (list). — Martínez 1959: 19 (catalogue for Argentina); 1988a: 281 (key). — Halffter & Matthews 1966: 259 (catalogue, distribution). — Halffter & Halffter 1976: 47 (redescription). — Howden & Young 1981: 11, 14 (key, redescription). — Halffter & Edmonds 1982: 137 (catalogue, distribution). — Jessop 1985: 1089 (redescription). — López-Guerrero & Halffter 2000: 244 (monograph). — Medina & Lopera-Toro 2000: 301 (key). — Vaz-de-Mello 2000: 193 (checklist for Brazil). — Medina et al. 2001: 135 (checklist for Colombia). — Ratcliffe 2002: 11 (checklist for Panama). — Morón 2003: 44 (list). — Huerta et al. 2003: 6 (biology). — Hamel-Leigue et al. 2006: 16 (list). — Génier 2009: 22 (revision). — Camero 2010: 149 (distribution for Colombia). — Vazde-Mello et al. 2011a: 21 (key). — Carvajal et al. 2011: 111, 314 (diagnosis, list). — Krajcik 2012: 107 (list). — Solís & Kohlmann 2012: 7 (checklist for Costa Rica). — Boilly & Vaz-de-Mello 2013: 106 (key). — Chamorro et al. 2018: 73, 95 (list for Ecuador); 2019: 133 (catalogue). — Hielkema & Hielkema 2019: 85 (catalogue for the Guianas). Aeschrotes – Castelnau 1840: 92 (cited as synonym). — Agassiz 1846: 26 (catalogue). — Lacordaire 1855: 106 (synonym of Eurysternus, see footnote). — Harold 1869d: 1023 (synonym of Eurysternus). — Gillet 1911b: 25 (synonym of Eurysternus). — Pessôa & Lane 1941: 406 (cited as synonym of Eurysternus). — Blackwelder 1944: 197 (cited as synonym of Eurysternus). — Martínez 1959: 19 (cited as synonym of Eurysternus). — Howden & Young 1981: 14 (synonym of Eurysternus). — Jessop 1985: 1089 (synonym of Eurysternus). — Ratcliffe 2002: 11 (synonym of Eurysternus). — Génier 2009: 23 (synonym of Eurysternus). — Solís & Kohlmann 2012: 7 (synonym of Eurysternus). Eurysternodes – Vaz-de-Mello 2000: 193 (checklist for Brazil, cited as subgenus of Eurysternus). — Génier 2009: 23 (synonym of Eurysternus). — Solís & Kohlmann 2012: 7 (cited as synonym of Eurysternus). Pareurysternus – Vaz-de-Mello 2000: 193 (checklist for Brazil; cited as subgenus of Eurysternus). — Génier 2009: 23 (synonym of Eurysternus). — Solís & Kohlmann 2012: 7 (cited as synonym of Eurysternus). — Krajcik 2012: 107 (cited as synonym of Eurysternus). AMartínezus – Génier 2009: 23 (synonym of Eurysternus). — Carvajal et al. 2011: 110 (description). — Solís & Kohlmann 2012: 7 (cited as synonym of Eurysternus). Eurysternus atrosericus Génier, 2009 Fig. 45B Eurysternus atrosericus Génier, 2009: 88 (original description). Type locality: Brazil: Pará: Óbidos. Name-bearing type: holotype (MZSP), not examined. LOZANO DE LA ROSA C. et al., Dung beetles (Scarabaeinae) of Venezuela 117 Eurysternus atrosericus – Camero 2010: 149, 153 (key, diagnosis). — Larsen 2011: 98 (list for Suriname); 2013: 97 (list). — Vaz-de-Mello et al. 2011b: 88 (list). — Krajcik 2012: 107 (checklist of species). — Ratcliffe 2013: 493 (pitfall trap study). — Bicknell et al. 2014: supp. file. — Silva et al. 2014: 348 (diversity); 2015: 611 (list for Mato Grosso state, Brazil); 2016: 7 (ecology, biogeography). — Nunes et al. 2014: 408–410 (list). — Chamorro et al. 2018: 95 (cited for Ecuador); 2019: 135 (catalogue). — Hielkema & Hielkema 2019: 85 (catalogue for the Guianas). Distribution Venezuela, Guyana, Suriname, Brazil, and Ecuador. Subregions of Venezuela Peneplain of the Casiquiare River–Upper Orinoco, and System of hills and low piedmont mountains of the Guiana Shield. Literature records Génier 2009: 86 (Venezuela: Amazonas: La Esmeralda, Puerto Ayacucho, Atunes, Río Negro and Río Baria). — Chamorro et al. 2019: 135 (Venezuela). Eurysternus balachowskyi Halffter & Halffter, 1976 Fig. 46A Eurysternus balachowskyi Halffter & Halffter 1976: 65 (original description). Type locality: French Guiana: Cayenne: Camopi, Oyapock River, Îlet Massikiri. Name-bearing type: holotype (MNHN) (Génier 2009), not examined. Eurysternus (Eurysternus) balthasari Martínez, 1988b: 294 (original description). Type locality: Brazil: Amapá: Serra do Navío. Name-bearing type: holotype (MACN), examined by MC. Eurysternus balachowskyi – Halffter et al. 1980: 613 (biology, cited for “Guiana”). — Halffter & Edmonds 1982: 46 (biology). — Huerta et al. 2003: 15 (biology). — Quintero & Roslin 2005: appendix A (ecology). — Feer 2008: 62 (ecology, list); 2013: 767 (list for French Guiana). — Génier 2009: 103 (revision). — Krajcik 2012: 63 (list). — Ferrer-Paris et al. 2013: 109 (list). — Ratcliffe 2013: 493 (list, cited for Brazil). — Feer & Boissier 2015: 169 (list). — Boilly 2018: 36, 37 (key, distribution). — Hielkema & Hielkema 2019: 85 (catalogue for the Guianas). — Storck-Tonon et al. 2020: 2426 (diversity). Eurysternus (Eurysternus) balthasari – Génier 2009: 103 (cited as synonym). Material examined VENEZUELA ‒ Bolívar • 2 specs; Isla de Anacoco; 9 Aug. 2006; curso. NM2006 leg.; faeces, 52h, 13:02; CEMT. Subregions of Venezuela Peneplain of the Caura and Paragua rivers, System of low mountains and hills Imataca-Cuyuní of Northeast Guyana Shield, and Guiana Shield. Distribution Venezuela, Guyana, Suriname, French Guiana, and Brazil (Génier 2009). European Journal of Taxonomy 959: 1–272 (2024) 118 Literature records Génier 2009: 103 (Venezuela: Bolívar). — Ferrer-Paris et al. 2013: 109 (Venezuela: Bolívar: Isla de Anacoco). — Boilly 2018: 37 (Venezuela). Eurysternus caribaeus (Herbst, 1789) Fig. 46B Scarabaeus caribaeus Herbst, 1789: 300 (original description). Type locality: French Guiana. Namebearing type: neotype (MFNB), designated by Jessop (1985), not examined. Note: see Iannuzzi et al. (2023) for a discussion on the authorship of the name. Eurysternus planus Dalman, 1824: 10 (original description). Type locality: French Guiana. Namebearing type: holotype (NHRS) (Génier 2009), not examined. Eurysternus nebulosus Kirsch, 1871: 361 (original description). Type locality: Colombia: Cundinamarca: Bogotá. Name-bearing type: lectotype (SMTD), designated by Jessop (1985), not examined. Eurysternus peruanus Harold, 1875a: 137 (original description). Type locality: Peru. Name-bearing type: lectotype (MNHN), designated by Jessop (1985), not examined. Eurysternus planus – Dejean 1836: 160 (catalogue, cited as “Eurysternus planus Dej”). — Castelnau 1840: 92 (redescription). — Harold 1869d: 1023 (synonym of Eurysternus caribaeus, catalogue). — Bruch 1911:182 (cited as synonym of E. caribaeus). — Gillet 1911b: 25 (cited as synonym of E. caribaeus). — Blackwelder 1944: 197 (cited as synonym of E. caribaeus). — Martínez 1959: 20 (cited as synonym of E. caribaeus). — Jessop 1985: 1102 (cited as synonym of E. caribaeus). — Génier 2009: 214 (cited as synonym of E. caribaeus). — Camero 2010: 164 (cited as synonym of E. caribaeus). — Solís & Kohlmann 2012: 7 (cited as synonym of E. caribaeus). Eurysternus caribaeus – Harold 1869d: 1023 (catalogue, distribution); 1880: 13 (distribution). — Bruch 1911: 182 (cited for Argentina). — Gillet 1911b: 25 (catalogue). — Boucomont 1928b: 1 (list). — Blackwelder 1944: 197 (list). — Guérin 1953: 256 (diagnosis). — Roze 1955: 41 (checklist for Venezuela). — Martínez 1959: 20 (catalogue for Argentina). — Vulcano & Pereira 1967: 547 (key for the Amazon). — Halffter & Halffter 1976: 57 (redescription). — Halffter et al. 1980: 600 (biology). — Howden & Young 1981: 14, 17 (key, characteristics, redescription). — Jessop 1985: 1093, 1102 (key, distribution). — Deloya 1992: 2 (list). — Amézquita et al. 1999: 119 (biodiversity). — Forsyth & Gill 1993: 70 (list). — Forsyth et al. 1998: 370 (conservation, list). — Escobar 2000: 208 (checklist for Colombia). — Vaz-de-Mello 2000: 193 (checklist for Brazil). — Medina et al. 2001: 135 (cited for Colombia). — Estrada & Coates-Estrada 2002: 1911 (ecology). — Ratcliffe 2002: 11 (checklist for Panama). — Kohlmann 2003: 45 (cited for Mexico). — Huerta et al. 2003: 16 (biology). — Pulido-Herrera et al. 2003: 54 (list for Caquetá, Colombia); 2007: 307 (cited for Andean region of Colombia). — Noriega 2004: 40 (checklist for Tinigua Park, Colombia). — Hamel-Leigue et al. 2006: 16 (cited for Bolivia); 43 (diversity). — Kohlmann et al. 2007: 29 (checklist). — Noriega et al. 2007: 82 (list for Colombia). — Feer 2008: 56, 62 (ecology); 2013: 767 (list for French Guiana). — Orozco & Pérez 2008: 38–39 (list for Colombia). — Carpio et al. 2009: 469 (ecology). — Génier 2009: 215, 219 (redescription, key). — Medina & Pulido-Herrera 2009: 60 (diversity). — Camero 2010: 150, 164 (key, diagnosis). — Brûlé et al. 2011a: 193 (list, cited for French Guiana). — Carvajal et al. 2011: 314 (cited for Ecuador). — Vaz-de-Mello et al. 2011b: 88 (list). — Krajcik 2012: 107 (list). — Solís & Kohlmann 2012: 7 (checklist for Costa Rica). — Ratcliffe 2013: 493 (list, cited for Brazil). — Nunes et al. 2014: 408–410 (list). — Silva et al. 2014: 348 (diversity); 2016: 554 (ecology, biogeography); 2017: 490 (list). — Feer & Boissier 2015: 169 (list). — Ratcliffe et al. 2015: 195 (checklist for Peru). — Boilly 2018: 36, 39 (key). — Chamorro et al. 2018: 95 (list for Ecuador); 2019: 135 (catalogue). — Giraldo et al. 2018: 72 (guide). — Santos et al. 2018: 46 (list for Acre, Brazil). — Hielkema & Hielkema 2019: 86 (catalogue for the Guianas). — Nieto et al. 2020: 136 (report). — Storck-Tonon et al. 2020: 2426 (diversity). LOZANO DE LA ROSA C. et al., Dung beetles (Scarabaeinae) of Venezuela 119 Fig. 46. Species distribution. A. Eurysternus balachowskyi Halffter & Halffter, 1976. B. Eurysternus caribaeus (Herbst, 1789). Green diamond = CEMT collection data; black circle = literature data. European Journal of Taxonomy 959: 1–272 (2024) 120 Eurysternus nebulosus – Harold 1880: 13 (synonym of E. caribaeus). — Bates 1887: 40 (comments, distribution). — Gillet 1911b: 25 (cited as synonym of E. caribaeus). — Blackwelder 1944: 197 (cited as synonym of E. caribaeus). — Martínez 1959: 20 (cited as synonym of E. caribaeus). — Jessop 1985: 1102 (cited as synonym of E. caribaeus). — Génier 2009: 214 (cited as synonym of E. caribaeus). — Camero 2010: 164 (cited as synonym of E. caribaeus). — Solís & Kohlmann 2012: 7 (cited as synonym of E. caribaeus). Eurysternus peruanus – Balthasar 1941: 340 (cited for Peru); 1951: 325 (cited for Peru). — Blackwelder 1944: 197 (list). — Jessop 1985: 1102 (synonym of E. caribaeus). — Génier 2009: 213 (cited as synonym of E. caribaeus). — Camero 2010: 164 (cited as synonym of E. caribaeus). — Solís & Kohlmann 2012: 7 (cited as synonym of E. caribaeus). Material examined VENEZUELA ‒ 2 specs; Caracas, Cerro Verde; 1100 m a.s.l.; 15 Apr. 2004; D. García leg.; CEMT. ‒ Táchira • 1 spec.; near Pregonero, Presa La Honda (Trampita); 1100 m a.s.l.; 21 Dec. 1988; D. Havranek leg.; mini-cup trap; CEMT. Distribution Mexico, Belize, Guatemala, Honduras, Nicaragua, Costa Rica, Panama, Colombia, Venezuela, Trinidad and Tobago, Guyana, Suriname, French Guiana, Brazil, Ecuador, Peru, Bolivia, Paraguay, and Argentina (Génier 2009). Subregions of Venezuela Plains, Delta plain of the Orinoco River and coastal swamp of the San Juan River, Peneplain of the Casiquiare River–Upper Orinoco, system of hills and low piedmont mountains of the Guiana Shield, System of low mountains and hills Imataca-Cuyuní del Escudo Northeast Guiana, Serrania de Perija, Andes mountains, Sierra de San Luis and Cerro Santa Ana, Central Coast Mountain Range, Oriental Coast Range, and Guiana Shield. Literature records Roze 1955: 41 (Venezuela: Aragua: Parque Nacional Henri Pittier, Estación Biológica Rancho Grande). — Jessop 1985: 1103 (Venezuela). — Blanco 1987: 41 (Venezuela: Táchira). — Havranek 1989: 61 (Venezuela: Táchira). — Barbero 2001: 5 (Venezuela). — Larsen et al. 2008: 1294 (Venezuela: Bolívar). — Génier 2009: 214 (Venezuela: Amazonas, Anzoátegui, Aragua, Bolívar, Delta Amacuro, Distrito Capital, Falcón, Lara, Miranda, Monagas, Táchira and Zulia). — Ferrer-Paris et al. 2013: 109 (Venezuela: Bolívar: Isla de Anacoco, Miranda: Altos de Pipe and Yaracuy: Hacienda Guáquira). — Boilly 2018: 39 (Venezuela). — Chamorro et al. 2019: 135 (Venezuela). Eurysternus cayennensis Castelnau, 1840 Fig. 47A Eurysternus cayennensis Castelnau 1840: 93 (original description). Type locality: French Guiana: Cayenne: Mont Paramana. Name-bearing type: neotype (MNHN), designated by Jessop (1985), not examined. Eurysternus confusus Jessop, 1985: 1106 (original description). Type locality: Ecuador: Sucumbíos: Dureno, Aguarico River. Name-bearing type: holotype (CMNC), not examined. Eurysternus cayennensis – Harold 1869d: 1023 (catalogue); 1880: 15 (comment). — Gillet 1911b: 25 (catalogue). — Blackwelder 1944: 197 (checklist). — Halffter & Halffter 1976: 73 (redescription). — Jessop 1985: 1095, 1104 (identification guide, designation of lectotype). — Feer 2000: 32 (list for LOZANO DE LA ROSA C. et al., Dung beetles (Scarabaeinae) of Venezuela 121 French Guiana). — Vaz-de-Mello 2000: 193 (checklist for Brazil). — Medina et al. 2001: 135 (checklist for Colombia). — Celi et al. 2004: 39 (diversity). — Feer & Pincebourde 2005: 30 (list). — Génier 2009: 25 (revision). — Brûlé & Dalens 2012: 37 (inventory). — Price & Feer 2012: 327 (list for French Guiana). — Bicknell et al. 2014: supp. 1 (ecology). — Boilly 2018: 34, 37 (key for French Guiana, comment). — Hielkema & Hielkema 2019: 86 (catalogue for the Guianas). Eurysternus (Eurysternus) cayennensis – Martínez 1988b: 283 (comment). Eurysternus (Eurysternus) confusus – Martínez 1988b: 283 (comments). Eurysternus confusus – Medina et al. 2001: 135 (checklist for Colombia). — Feer & Pincebourde 2005: 30 (list). — Génier 2009: 25 (cited as synonym of E. cayennensis). — Hielkema & Hielkema 2019: 86 (cited as synonym of E. cayennensis Castelnau, 1840). Distribution Colombia, Venezuela, Guyana, Suriname, French Guiana, Brazil, Ecuador, Peru, and Bolivia. Subregions of Venezuela Peneplain of the Casiquiare River–Upper Orinoco, System of hills and low piedmont mountains of the Guiana Shield, and Guiana Shield. Literature records Halffter & Halffter 1976: 73 (Venezuela). — Martínez 1988b: 283 (Venezuela). — Génier 2009: 25 (Venezuela: Amazonas and Bolivar). Eurysternus foedus Guérin-Méneville, 1830 Fig. 47B Eurysternus foedus Guérin-Méneville, 1830: pl. 21 fig. 5, 5.a (species-group name made available by being accompanied by an illustration of a member of the taxon, an indication in the sense of Article 12.2.7 of the Code [ICZN 1999]). Type locality: Brazil: Mato Grosso: 264 km N of Xavantina, Serra do Roncador. Name-bearing type: neotype (BMNH), designated by Jessop (1985), not examined. Eurysternus claudicans Kirsch, 1871: 360 (original description). Type locality: Colombia: Cundinamarca: Bogotá. Name-bearing type: holotype (SMTD), not examined. Eurysternus foedus – Castelnau 1840: 93 (comments). — Guérin-Méneville 1844: 76 (redescription). — Harold 1869d: 1024 (catalogue). — Ohaus 1909: 94 (cited for Ecuador). — Gillet 1911b: 25 (catalogue). — Pessôa & Lane 1941: 409 (redescription). — Blackwelder 1944: 197 (list). — Vulcano & Pereira 1967: 547 (key). — Halffter & Halffter 1976: 78 (redescription). — Jessop 1985: 1093, 1102 (key, distribution, neotype). — Forsyth et al. 1998: 370 (conservation, list). — Maes 1998: 654 (list for Nicaragua). — Escobar 2000: 208 (checklist for Colombia). — Barbero 2001: 4 (cited for Nicaragua). — Medina et al. 2001: 135 (cited for Colombia). — Ratcliffe 2002: 11 (checklist for Panama). — Kohlmann 2003: 45 (cited for Mexico). — Huerta et al. 2003: 24 (biology). — Noriega 2004: 40 (checklist for Tinigua Park, Colombia). — Hamel-Leigue et al. 2006: 16 (cited for Bolivia). — Kohlmann et al. 2007: 29 (checklist). — Pulido-Herrera et al. 2007: 307 (cited for Andean region of Colombia). — Feer 2008: 62 (ecology, list); 2013: 767 (list for French Guiana). — Génier 2009: 146, 288 (diagnosis, key). — Medina & Pulido-Herrera 2009: 60 (diversity). — Camero 2010: 149, 153 (key, diagnosis). — Brûlé et al. 2011a: 193 (list, cited for French Guiana). — Carvajal et al. 2011: 314–315 (cited for Ecuador). — Larsen 2011: 99 (cited for Suriname). — Cultid-Medina et al. 2012: 63 (guide). — Vaz-de-Mello et al. 2011b: 88 (list). — Krajcik 2012: 107 (checklist). — Noriega et al. 2012: 47 (list for Antioquia, Colombia). — Solís & Kohlmann 2012: 7 (checklist for Costa Rica). — Pardo-Locarno & Camero 2014: 210, 211–212 (list for Chocó, Colombia). — Cultid-Medina & Medina 2015: 119–133, 201 (Appendix 11 European Journal of Taxonomy 959: 1–272 (2024) 128 Torres et al. 2017: 48–49, 67 (cited for Boyaca, Colombia). — Villada-Bedoya et al. 2017: 197 (ecology). — Chamorro et al. 2018: 78, 95 (fig. 1A, 1E, list for Ecuador); 2019: 148 (catalogue). — Nieto et al. 2020: 136 (report). Eurysternus pectoralis – Harold 1869d: 1024 (catalogue). — Gillet 1911b: 25 (catalogue, cited for Colombia). — Blackwelder 1944: 197 (catalogue, cited for Brazil). — Jessop 1985: 1106 (comments). — Génier 2009: 194 (synonym of E. marmoreus). — Camero 2010: 171 (cited as synonym of E. marmoreus). Eurysternus (Eurysternus) pectoralis – Vaz-de-Mello 2000: 193 (checklist for Brazil). Material examined VENEZUELA ‒ Mérida • 1 spec.; Sucre, Jaji; 8°39′36.75″ N, 71°23′52.69″ W; 2235 m a.s.l.; 8 Jul. 2009; D. Mora, P. Colmenares, M. Córdova and M. Nuñez leg.; CEMT. ‒ Yaracuy • 2 specs; Bolívar, Aroa; 10°22′10.56″ N, 68°50′16.94″ W; 1639 m a.s.l.; 21 Jul. 2009; M. Asmüssen, P. Colmenares and H. Martínez leg.; CEMT • 1 spec.; same data as for the preceding except for the coordinates and elevation; 10°22′8.21″ N, 68°50′18.52″W; 1626 m a.s.l.; CEMT. Distribution Colombia, Venezuela, Ecuador, Peru, and Bolivia (Génier 2009). Subregions of Venezuela Andes mountains and Central Coast Mountain Range Literature records Harold 1869d: 1024 (Venezuela). — Heyne & Taschenberg 1908: 68 (Venezuela). — Huerta et al. 2003: 5 (Venezuela: Mérida). — Génier 2009: 194 (Venezuela: Aragua, Mérida and Táchira). — Ferrer-Paris et al. 2013: 109 (Venezuela: Mérida: La Azulita-Jají). — Chamorro et al. 2019: 148 (Venezuela). Eurysternus mexicanus Harold, 1869 Fig. 50A Eurysternus mexicanus Harold 1869a: 505 (original description). Type locality: Mexico. Name-bearing type: lectotype (MNHN), designated by Jessop (1985), not examined. Eurysternus mexicanus – Harold 1880: 14 (comment). — Bates 1887: 41 (distribution). — Candèze 1891: 330 (list). — Gillet 1911b: 26 (catalogue). — Blackwelder 1944: 197 (list). — Roze 1955: 41 (checklist for Venezuela). — Halffter & Matthews 1966: 73 (biology). — Vulcano & Pereira 1967: 547 (key). — Howden & Young 1981: 17 (diagnosis). — Halffter & Edmonds 1982: 46 (biology). — Jessop 1985: 1100 (designation of lectotype). — Blanco 1987: 41 (catalogue). — Deloya 1992: 2 (list). — López-Guerrero & Morón 1994: 347 (biology). — Montes de Oca & Halffter 1995: 167 (ecology). — Maes 1998: 654 (list for Nicaragua). — Amézquita et al. 1999: 119 (biodiversity). — Escobar 2000: 208 (checklist for Colombia). — López-Guerrero & Halffter 2000: 245 (morphology). — Estrada & Coates-Estrada 2002: 1911 (ecology). — Huerta et al. 2003: 19 (biology). — Morón 2003: 44 (diagnosis). — Kohlmann et al. 2007: 29 (checklist). — Noriega et al. 2007: 82 (list for Colombia). — Pulido-Herrera et al. 2007: 307 (cited for Andean region of Colombia). — Huerta & Martínez 2008: 123–132 (biology). — Larsen et al. 2008: 1294 (list). — Génier 2009: 198, 261, 326, 427 (diagnosis, distribution, key). — Medina & Pulido-Herrera 2009: 60 (diversity). — Camero 2010: 169 (cited of Colombia). — Lozano 2010: 86 (list). — CultidMedina et al. 2012: 64 (guide). — Delgado et al. 2012: 329 (list). — Solís & Kohlmann 2012: 7 (checklist for Costa Rica). — Ferrer-Paris et al. 2013: 109 (list). — Pardo-Locarno & Camero 2014: LOZANO DE LA ROSA C. et al., Dung beetles (Scarabaeinae) of Venezuela 129 210, 212, 215 (list for Chocó, Colombia). — Molina et al. 2016: 128–130 (cited for Valle del Cauca, Colombia). — Giraldo et al. 2018: 50 (guide). — Nieto et al. 2020: 136 (report). Material examined VENEZUELA ‒ Bolívar • 1 spec.; Isla de Anacoco; 6 Aug. 2006; Curso NM2006 leg.; faeces, 23h -16:22, NM; CEMT. ‒ Táchira • 2 specs; Jun. 1989; J. Blanco leg.; CEMT • 1 spec.; San Cristóbal, Coloncito; 9 Apr. 1995; O. Hillert leg.; CEMT. ‒ Yaracuy • 1 spec.; Bolívar, Aroa; 10°0′0″ N, 68°0′0″ W; 468 m a.s.l.; 21 Aug. 2009; M. Assmüssen, P. Colmenares and H. Martínez leg.; human faeces; CEMT. Distribution Mexico, Belize, Guatemala, Honduras, Nicaragua, Costa Rica, Panama, Colombia, Venezuela, Trinidad and Tobago, and Guyana (Génier 2009). Subregions of Venezuela Maracaibo Depression, Plains, Delta plain of the Orinoco River and coastal swamp of the San Juan River, Peneplain of the Caura and Paragua rivers, System of hills and low sierras Lara-Falcón, System of hills and low piedmont mountains of the Guiana Shield, System of low mountains and hills ImatacaCuyuní of Northeast Guyana Shield, Serrania de Perija, Andes mountains, Central Coast Mountain Range, Oriental Coast Range, and Guiana Shield. Literature records Harold 1880: 14 (Venezuela). — Candèze 1891: 330 (Venezuela: Carabobo: San Esteban). — Gillet 1911b: 26 (Venezuela). — Blackwelder 1944: 197 (Venezuela). — Roze 1955: 41 (Venezuela). — Vulcano & Pereira 1967: 547 (Venezuela). — Jessop 1985: 1101 (Venezuela). — Blanco 1987: 41 (Venezuela: Táchira). — Larsen et al. 2008: 1294 (Venezuela: Bolívar). — Génier 2009: 199 (Venezuela: Aragua; Bolívar; Lara; Mérida; Miranda; Monagas; Sucre; Táchira; Zulia and Distrito Capital). — Lozano 2010: 86 (Venezuela: Zulia). — Ferrer-Paris et al. 2013: 109 (Aragua [Guárico]: Altagracia de Orituco, Bolívar: Isla de Anacoco, Mérida: La Azulita-Jají, Yaracuy: Hacienda Guáquira and Zulia: Rosario de Perijá). Eurysternus plebejus Harold, 1880 Fig. 50B Eurysternus plebejus Harold, 1880: 14 (original description). Type locality: Colombia: Boyacá: Muzo. Name-bearing type: lectotype (MNHN), designated by Jessop (1985), not examined. Eurysternus (Eurysternus) joffrei Martínez, 1988b: 290 (original description). Type locality: Peru: Huánuco: Leoncio Prado: Tingo María: Universidad Nacional Agraria de la Selva. Name-bearing type: holotype (MACN), examined by MC. Eurysternus plebejus – Ohaus 1909: 94 (cited for Ecuador). — Gillet 1911b: 26 (catalogue). — Blackwelder 1944: 197 (list). — Roze 1955: 41 (checklist for Venezuela). — Halffter & Matthews 1966: 146 (cited for Ecuador). — Vulcano & Pereira 1967: 547 (key). — Howden & Young 1981: 14, 15 (key, redescription). — Jessop 1985: 1093, 1100 (key, distribution). — Escobar 1997: 422, 423 (diversity); 2000: 208 (checklist for Colombia). — Maes 1998: 654 (list for Nicaragua). — Barbero 2001: 5 (cited for Nicaragua). — Medina et al. 2001: 135 (cited for Colombia). — Ratcliffe 2002: 11 (checklist for Panama). — Bustos-Gómez & Lopera-Toro 2003: 61 (diet). — Morón 2003: 44 (cited for Mexico). — Noriega 2004: 40 (checklist for Tinigua Park, Colombia). — HamelLeigue et al. 2006: 16 (cited for Bolivia). — Kohlmann et al. 2007: 29 (checklist). — Noriega et al. 2007: 82 (list for Colombia). — Pulido-Herrera et al. 2007: 307 (cited for Andean region of Colombia). — Génier 2009: 172, 289 (diagnosis, key). — Medina & Pulido-Herrera 2009: 61 European Journal of Taxonomy 959: 1–272 (2024) 130 Fig. 50. Species distribution. A. Eurysternus mexicanus Harold, 1869. B. Eurysternus plebejus Harold, 1880. Green diamond = CEMT collection data; black circle = literature data. LOZANO DE LA ROSA C. et al., Dung beetles (Scarabaeinae) of Venezuela 131 (diversity). — Camero 2010: 150, 168 (key, diagnosis). — Carvajal et al. 2011: 314–315 (cited for Ecuador). — Cultid-Medina et al. 2012: 65 (guide). — Krajcik 2012: 107 (list). — Silva et al. 2016: 7 (ecology, biogeography). — Solís & Kohlmann 2012: 7 (checklist for Costa Rica). — Ferrer-Paris et al. 2013: 109 (list). — Ratcliffe et al. 2015: 195 (checklist for Peru). — Boilly 2018: 36–37 (key, distribution). — Chamorro et al. 2018: 95 (list for Ecuador); 2019: 149 (catalogue). — Hielkema & Hielkema 2019: 88 (catalogue for the Guianas). — Nieto et al. 2020: 136 (report). Eurysternus (Eurysternus) plebejus – Vaz-de-Mello 2000: 193 (checklist for Brazil). Eurysternus (Eurysternus) joffrei – Génier 2009: 172 (synonym of E. plebejus). — Camero 2010: 168 (cited as synonym of E. plebejus). — Solís & Kohlmann 2012: 7 (cited as synonym of E. plebejus). Material examined VENEZUELA ‒ Táchira • 1 spec.; Libertador, San Joaquín de Navay; 200 m a.s.l.; Aug. 2006; T. Good leg.; CEMT. ‒ Yaracuy • 1 spec.; Bolívar, Aroa; 10°23′11.9″ N, 68°50′49.41″ W; 1380 m a.s.l.; 19 Jul. 2009; M. Assmüssen, P. Colmenares and H. Martínez leg.; human faeces; CEMT. Distribution Nicaragua, Costa Rica, Panama, Colombia, Venezuela, Guyana, Suriname, French Guiana, Brazil, Ecuador, Peru, and Bolivia. Subregions of Venezuela Plains, System of hills and low sierras Lara-Falcón, System of low mountains and hills Imataca-Cuyuní of Northeast Guyana Shield, Serrania de Perija, Central Coast Mountain Range, and Guiana Shield. Literature records Roze 1955: 41 (Venezuela: Aragua and Zulia). — Howden & Young, 1981: 15 (Venezuela). — Jessop 1985: 1101 (Venezuela). — Martínez 1988b: 290 (Venezuela: Aragua). — Barbero 2001: 5 (Venezuela). — Génier 2009: 172 (Venezuela: Aragua; Bolívar; Carabobo; Distrito Capital and Miranda). — FerrerParis et al. 2013: 109 (Miranda: Altos de Pipe and Yaracuy: Hacienda Guáquira). — Boilly 2018: 36, 37 (Venezuela). — Chamorro et al. 2019: 149 (Venezuela). Eurysternus sanbornei Gill, 1990 Fig. 51A Eurysternus sanbornei Gill, 1990: 355 (original description). Type locality: Venezuela: Bolivar: 15 km E of Kavanayen. Name-bearing type: holotype (MIZA), not examined. Eurysternus sanbornei – Génier 2009: 144 (revision). — Hielkema & Hielkema 2019: 89 (catalogue for the Guianas). Distribution Venezuela and Guyana (Génier 2009). Subregions of Venezuela System of low mountains and hills Imataca-Cuyuní of Northeast Guyana Shield, and Guiana Shield. Literature records Gill 1990: 355 (Venezuela: Bolívar). — Génier 2009: 144 (Venezuela: Bolívar). European Journal of Taxonomy 959: 1–272 (2024) 132 Eurysternus superbus Génier, 2009 Fig. 51B–C Eurysternus superbus Génier, 2009: 113 (original description). Type locality: Venezuela: Aragua: Parque Nacional Henri Pittier, Estación Biológica Rancho Grande, 1100 m a.s.l. Name-bearing type: holotype (CMNC), not examined. Eurysternus superbus — Ferrer-Paris et al. 2013: 109 (list). Material examined VENEZUELA ‒ Aragua • 3 specs; Maracay, “La cumbre”; 1500 m; 14 Jun. 1993; O. Hillert leg.; CEMT • 1 spec.; same data as for preceding except for the date; 27 Jul. 1993; CEMT. ‒ Miranda • 3 specs; Altos de Pipe, Instituto Venezolano de Investigaciones Científicas (“Campus IVIC”); 10°23′51″ N, 66°58′15″ W; 1500 m a.s.l.; Jul. 2009; F.Z. Vaz-de-Mello leg.; CEMT. ‒ Yaracuy • 2 specs; Bolívar, Aroa; 10°22′10.56″ N, 68°50′16.94″ W; 1639 m a.s.l.; 21 Jul. 2009; M.Asmüssen, P. Colmenares and H. Martínez leg.; CEMT. Distribution Venezuela (endemic, Génier 2009). Subregions of Venezuela Andes mountains and Central Coast Mountain Range. Literature records Génier 2009: 113 (Venezuela: Aragua, Carabobo and Lara). — Ferrer-Paris et al. 2013: 109 (Venezuela: Miranda: Altos de Pipe). Eurysternus truncus Génier, 2009 Fig. 52A Eurysternus truncus Génier, 2009: 166 (original description). Type locality: Venezuela: Bolívar: 120 km S of El Dorado. Name-bearing type: holotype (CMNC), not examined. Eurysternus truncus – Hielkema & Hielkema 2019: 89 (catalogue for the Guianas). Distribution Venezuela and Guyana (Génier 2009). Subregions of Venezuela System of hills and low piedmont mountains of the Guiana Shield, System of low mountains and hills Imataca-Cuyuní of Northeast Guyana Shield, and Guiana Shield. Literature records Génier 2009: 167 (Venezuela: Bolívar). Eurysternus ventricosus Gill, 1990 Fig. 52B Eurysternus ventricosus Gill, 1990: 358 (original description). Type locality: Venezuela: Bolívar: 120 km S of El Dorado. Name-bearing type: holotype (MIZA), not examined. LOZANO DE LA ROSA C. et al., Dung beetles (Scarabaeinae) of Venezuela 133 Fig. 51. Species distribution. A. Eurysternus sanbornei Gill, 1990. B. Eurysternus superbus Génier, 2009. C. Dorsal view of E. superbus. Green diamond = CEMT collection data; black circle = literature data. European Journal of Taxonomy 959: 1–272 (2024) 134 Eurysternus ventricosus – Génier 2009: 44 (revision). — Boilly 2018: 34, 37 (key, distribution). — Larsen 2011: 99 (list, cited for Suriname); 2013: 97 (list). — Ferrer-Paris et al. 2013: 109 (list). — Ratcliffe 2013: 493 (list, ecology). — Hielkema & Hielkema 2019: 89, 290 (catalogue for the Guianas). — Storck-Tonon et al. 2020: 2426 (diversity). Eurysternus (Eurysternus) ventricosus – Vaz-de-Mello 2000: 193 (list, cited for Brazil). Distribution Venezuela, Guyana, Suriname, French Guiana, Brazil, and Bolivia. Subregions of Venezuela Peneplain of the Casiquiare River–Upper Orinoco, System of low mountains and hills Imataca-Cuyuní of Northeast Guyana Shield, and Guiana Shield. Literature records Gill 1990: 355, 358 (Venezuela: Bolívar). — Génier 2009: 44 (Venezuela: Amazonas and Bolívar). — Ferrer-Paris et al. 2013: 109 (Venezuela: Bolívar: Isla de Anacoco). — Boilly 2018: 37 (Venezuela). Genus Genieridium Vaz-de-Mello, 2008 Genieridium Vaz-de-Mello, 2008: 26 (original description). Type species: Pedaridium bidens Balthasar, 1942, by original designation. Genieridium – Krajcik 2012: 111 (list). — Hielkema & Hielkema 2019: 45 (catalogue for the Guianas). Genieridium bordoni (Martínez, 1992) Fig. 53A–B Pedaridium bordoni Martínez 1992: 22 (original description). Type locality: Venezuela: Barinas: Santa Bárbara. Name-bearing type: holotype (MACN), examined by FZVM. Pedaridium bordoni – Vaz-de-Mello & Canhedo 1998: 100 (comments, key). Genieridium bordoni – Vaz-de-Mello 2008: 27, 29 (key, distribution, new combination). — Krajcik 2012: 111 (list). — Hielkema & Hielkema 2019: 45 ( catalogue for the Guianas). Material examined VENEZUELA ‒ Anzoategui • 1 spec.; Pariaguan; 13 Aug. 1967; J. and B. Bechyne leg.; CEMT. ‒ Barinas • 3 specs; Otopún; 28 Aug 2006; curso NM2006 leg.; faeces, 15:43, 24h; CEMT. ‒ Bolívar • 2 specs; 15 km E of Caicara; 12 Jun 1996; H. and A. Howden leg.; CEMT. ‒ Monagas • 4 specs; Maturin, 42 km SE of Maturin; 5 Jul. 1958; Arnold Menke leg.; CEMT • 2 specs; Uverito; 23 Jun. 1978; C.J. Rosales and J.A. Gonzales leg.; neon light; CEMT. Distribution Venezuela (endemic). Subregions of Venezuela Plains, Delta plain of the Orinoco River and coastal swamp of the San Juan River, Peneplain of the Caura and Paragua rivers, System of hills and low piedmont mountains of the Guiana Shield, and Central Coast Mountain Range. LOZANO DE LA ROSA C. et al., Dung beetles (Scarabaeinae) of Venezuela 135 Fig. 52. Species distribution. A. Eurysternus truncus Génier, 2009. B. Eurysternus ventricosus Gill, 1990. Black circle = literature data. European Journal of Taxonomy 959: 1–272 (2024) 136 Literature records Vaz-de-Mello & Canhedo 1998: 100 (Venezuela: Guárico, Barinas, Monagas, Anzoátegui and Aragua). — Vaz-de-Mello 2008: 29 (Anzoátegui, Barinas; Bolívar; Delta Amacuro; Guárico and Monagas). — Krajcik 2012: 111 (Venezuela: Guárico). Genus Gromphas Dejean, 1836 Gromphas Dejean, 1836: 159 (genus-group name made available by being accompanied by an available species-group name, an indication as defined in Art. 12.2.5 of the Code [ICZN 1999]; see Cupello [2024] for details). Type species: Coprobius lacordairii Oken, 1834, by original monotypy; see Cupello (2024). Copris (Gromphas) – Brullé 1838: 283, 298, 304 (first description of the genus). Gromphas – Sturm 1843: 108 (catalogue). — Agassiz 1846: 481 (catalogue). — Blanchard 1846: 181 (description of new species). — Erichson 1847b: 760 (comment). — Lacordaire 1855: 100 (redescription). — Harold 1868e: 81 (characteristics); Harold 1869d: 1016 (catalogue). — Bates 1870: 175 (comments). — Chapuis 1876: 276 (catalogue). — Karsch 1887: 1 (cited). — Kolbe 1905: 550 (comments). — Gillet 1911b: 80 (catalogue). — Fabre 1919: 244, 247, 256 (comments). — Lucas 1920: 309 (catalogue, distribution). — d’Olsoufieff 1924: 17 (key, redescription). — Luederwaldt 1929: 614 [12] (characteristics, diagnosis). — Pessoa 1934: 282 (comments). — Pessôa & Lane 1941: 470 (key). — Blackwelder 1944: 208 (list). — Roze 1955: 45 (checklist for Venezuela). — Martínez 1959: 95 (catalogue for Argentina). — Barattini & Sáenz 1961: 21 (contribution); 1964: 173 (comments). — Halffter & Matthews 1966: 257 (catalogue, distribution). — Vulcano & Pereira 1967: 565 (key). — Edmonds 1972: 816 (comment). — Halffter & Edmonds 1982: 137 (catalogue, distribution). — Zunino 1983: 22 (comments); 1985: 104 (comment). — Martínez 1989: 50, 67 (list, distribution). — Hanski & Cambefort 1991: 468 (ecology). — Walsh & Gandolfo 1996: 587 (biology). — Medina & Lopera-Toro 2000: 305 (key). — Vaz-de-Mello 2000: 193 (checklist for Brazil). — Medina et al. 2001: 138 (checklist for Colombia). — Davis et al. 2002: 1240 (biogeography, list). — Philips et al. 2004a: 55–56 (phylogeny); 2004b: 43, 51–54, 58 (phylogeny). — HamelLeigue et al. 2006: 17 (list). — Ocampo & Hawks 2006: 558 (phylogeny). — Noriega et al. 2008a: 131 (cited); 2010: 454 (comments). — Scholtz et al. 2009: 246 (history). — Gillet et al. 2010: 4 (distribution). — Bouchard et al. 2011: 245 (nomenclator). — Vaz-de-Mello et al. 2011a: 24 (key). — Figueroa et al. 2012: 2 (redescription); 2014: 137 (distribution for Peru). — Gatty et al. 2012: 1 (comments). — Krajcik 2012: 117 (list). — Korasaki et al. 2012: 427 (distribution). — Cupello 2013: 15–16. — Cupello & Vaz-de-Mello 2013b: 443 (revision); 2014: 399 (comments); 2015: 3, 11 (key, distribution, phylogeny). — Chamorro et al. 2018: 75 95 (list for Ecuador); 2019: 156 (catalogue). — Hielkema & Hielkema 2019: 93 (catalogue for the Guianas). — Cupello 2024: 23–51. Gromphas lemoinei Waterhouse, 1891 Fig. 53C Gromphas lemoinei Waterhouse, 1891b: 60 (original description). Type locality: Venezuela: La Guaira: La Guaira. Name-bearing type: holotype (BMNH), examined by MC. Gromphas aeruginosa var. lemoinei – Gillet 1911b: 80 (catalogue). — d’Olsoufieff 1924: 138 (catalogue). — Blackwelder 1944: 208 (list). — Roze 1955: 45 (cited, checklist for Venezuela). — Barattini & Sáenz 1961: 23 (contribution); 1964: 177 (description). Gromphas lemoinei – Cupello & Vaz-de-Mello 2013b: 447, 454 (key, revision, redescription); 2015: 12 (phylogeny). — Gámez & Acconcia 2018: 71, 73 (list). — Giraldo et al. 2018: 78 (guide). LOZANO DE LA ROSA C. et al., Dung beetles (Scarabaeinae) of Venezuela 137 Fig. 53. Species distribution. A. Genieridium bordoni (Martínez, 1992). B. Dorsal view of G. bordoni. C. Gromphas lemoinei Waterhouse, 1891. Green diamond = CEMT collection data; black circle = literature data. European Journal of Taxonomy 959: 1–272 (2024) 144 138 (checklist for Colombia). — Pulido-Herrera et al. 2007: 307 (cited for Andean region of Colombia). — Medina & Pulido-Herrera 2009: 60 (diversity). — Ferrer-Paris et al. 2013: 109 (list). — França et al. 2016: 3 (report). — Giraldo et al. 2018: 62 (guide). — Nieto et al. 2020: 136 (report). Ontherus polynice – Harold 1869c: 59 (comments); 1869d: 1008 (catalogue, cited for Bolivia). — Gillet 1911a: 57 (catalogue, cited as synonym of O. appendiculatus). — Luederwaldt 1931b: 389 (revision, cited as synonym of O. appendiculatus). — Pessôa & Lane 1941: 458 (monograph, cited as synonym of O. appendiculatus). — Blackwelder 1944: 206 (list, cited as synonym of O. appendiculatus). — Génier 1996: 78 (revision, cited as synonym of O. appendiculatus). Ontherus (Copris) polynice – Harold 1869c: 59 (comments). Ontherus quadratus – Harold 1869c: 59 (comments); 1869d: 1008 (catalogue, cited for Brazil); 1875b: 181 (comments). — Bruch 1911: 187 (catalogue, cited as synonym of Ontherus rotundatus). — Gillet 1911a: 57 (catalogue, cited as synonym of Ontherus appendiculatus). — Luederwaldt 1931a: 389 (revision, cited as synonym of O. appendiculatus). — Pessôa & Lane 1941: 458 (monograph, cited as synonym of O. appendiculatus). — Blackwelder 1944: 206 (list, cited as synonym of O. appendiculatus). — Génier 1996: 78 (revision, cited as synonym of O. appendiculatus). Ontherus rotundatus – Harold 1869c: 59 (comments); 1869d: 1008 (catalogue, cited for Corrientes); 1875b: 181 (comments). — Bruch 1911: 187 (catalogue, cited for Corrientes). — Gillet 1911a: 57 (catalogue, cited as synonym of Ontherus appendiculatus). — Luederwaldt 1931a: 389 (revision, cited as synonym of O. appendiculatus). — Pessôa & Lane 1941: 458 (monograph, cited as synonym of O. appendiculatus). — Blackwelder 1944: 206 (list, cited as synonym of O. appendiculatus). — Génier 1996: 78 (revision, cited as synonym of O. appendiculatus). Copris rotundatus – Burmeister 1874: 127 (comments, cited as Copris rotundata). — Bruch 1911: 187 (catalogue, cited as synonym of Ontherus rotundatus). — Génier 1996: 78 (revision, cited as synonym of O. appendiculatus). Copris polynice – Bruch 1911: 187 (catalogue, cited as synonym of Ontherus rotundatus). — Génier 1996: 78 (revision, cited as synonym of O. appendiculatus). Ontherus contractus – Bruch 1911: 187 (catalogue, cited for Tucumán). — Luederwaldt 1931a: 389 (revision, cited as synonym of O. appendiculatus). — Blackwelder 1944: 206 (list, cited for Argentina). — Génier 1996: 78 (revision, cited as synonym of O. appendiculatus). Copris appendiculatus – Génier 1996: 78 (revision). Ontherus (Ontherus) appendiculatus – Génier 1996: 78 (revision). — Vaz-de-Mello 2000: 194 (checklist for Brazil). — Vaz-de-Mello et al. 2011b: 88 (list). — Hielkema & Hielkema 2019: 56 (catalogue for the Guianas). Material examined VENEZUELA ‒ Táchira • 1 spec.; Libertador, San Joaquín de Navay; 7.7741° N, 71.6675° W; 550 m a.s.l.; Aug 2006; T. Good leg.; CEMT. Distribution Colombia, Venezuela, Guyana, Suriname, French Guiana, Brazil, Ecuador, Bolivia, Paraguay, Argentina, and Uruguay. Subregions of Venezuela Plains, System of hills and low sierras Lara-Falcón, System of hills and low piedmont mountains of the Guiana Shield, Andes mountains, and Central Coast Mountain Range. LOZANO DE LA ROSA C. et al., Dung beetles (Scarabaeinae) of Venezuela 145 Literature records Blanco 1987: 44 (Venezuela: Táchira). — Génier 1996: 78 (Venezuela: Anzoategui, Apure, Aragua, Bolívar, Carabobo, Distrito Capital, Guárico, Miranda, Monagas, Portuguesa, Tachira and Yaracuy). — França et al. 2016: 4–5 (Venezuela). — Ferrer-Paris et al. 2013: 109 (Venezuela: Aragua [Guárico]: Altagracia de Orituco). Ontherus brevicollis Kirsch, 1871 Fig. 56A Ontherus brevicollis Kirsch, 1871: 356 [340] (original description). Type locality: Colombia: Cundinamarca: Bogotá. Name-bearing type: holotype (SMTD), not examined. Ontherus brevicollis – Harold 1880: 23 (list, comments, cited for Nueva Granada [Colombia]). — Gillet 1911b: 57 (catalog, cited for Colombia). — Luederwaldt 1931a: 393 (key). — Blackwelder 1944: 206 (list, cited for Colombia). — Gacharná 1951: 222 (list for Colombia). — Vulcano & Pereira 1967: 583 (key). — Escobar 2000: 209 (checklist for Colombia). — Medina et al. 2001: 138 (checklist for Colombia); 2002: 182–185 (ecology). — Pulido-Herrera et al. 2007: 307 (cited for Andean region of Colombia). — Medina & Pulido-Herrera 2009: 60 (diversity). — Cultid-Medina et al. 2012: 44 (guide). — Krajcik 2012: 174 (list). — Martínez-Revelo & Lopera-Toro 2014: 65–68 (cited for Nariño department, Colombia). — Cultid-Medina & Medina 2015: 119–133, 203 (Appendix 13). — Giraldo et al. 2018: 116 (guide). Ontherus (Caelontherus) brevicollis – Génier 1996: 31 (revision, redescription). — Vaz-de-Mello 2000: 194 (checklist for Brazil). — Carvajal et al. 2011: 318–319 (list for Ecuador). — Chamorro et al. 2018: 96 (list for Ecuador); 2019: 167 (catalogue). Material examined VENEZUELA ‒ Mérida • 1 spec.; no further data; CEMT • 3 specs; La Mukuy; 2500 m a.s.l.; 14 Apr. 1995; O. Hillert leg.; CEMT • 4 specs; Monte Zerpa; 2400 m a.s.l.; Jul. 1995; A. de Ascençao leg.; dung; CMET • 1 spec.; Mérida; 2200 m a.s.l.; 11 Apr. 1995; Hornburg and, Krause leg.; CEMT. Distribution Colombia, Venezuela, Brazil, Ecuador, and Bolivia. Subregion of Venezuela Andes mountains. Literature records Vulcano & Pereira 1967: 583 (Venezuela). — Génier 1996: 31 (Venezuela: Mérida and Táchira). — Chamorro et al. 2019: 167 (Venezuela). Ontherus gilli Génier, 1996 Fig. 56B–C Ontherus (Caelontherus) gilli Génier, 1996: 28 (original description). Type locality: Venezuela: Táchira: San Cristóbal. Name-bearing type: holotype (CMNC), not examined. Ontherus gilli – Krajcik 2012: 174 (list). — Ferrer-Paris et al. 2013: 109 (list). — Gonzáles & Medina 2015: 80‒90 (new record for Colombia). European Journal of Taxonomy 959: 1–272 (2024) 146 Fig. 56. Species distribution. A. Ontherus brevicollis Kirsch, 1871. B. Ontherus gilli Génier, 1996. C. Dorsal view of O. gilli. Green diamond = CEMT collection data; black circle = literature data. LOZANO DE LA ROSA C. et al., Dung beetles (Scarabaeinae) of Venezuela 147 Material examined VENEZUELA • 1 spec.; no further data; CEMT. ‒ Mérida • 1 spec.; Santo Domingo, NM27-T078; 13 Jul. 2006; curso NM2006 leg.; faeces, 22h-07:41; CEMT • 2 specs; Sucre, Jají; 8°40′27.98″ N, 71°25′3.53″ W; 1849 m a.s.l.; 7 Jul. 2009; D. Mora, P. Colmenares, M. Córdova and M. Nuñez leg.; minced meat; CEMT • 4 specs; same data as for preceding except for the elevation and date; 1864 m a.s.l.; 8 Jul. 2009; CEMT. ‒ Yaracuy • 1 spec.; Bolívar, Aroa; 10°22′9.37″ N, 68°50′19.35″ W; 1624 m a.s.l.; 19 Jul. 2009; M. Asmüssen, P. Colmenares and H. Martínez leg.; human faeces; CEMT. Distribution Colombia and Venezuela. Subregions of Venezuela Andes mountains and Central Coast Mountain Range. Literature records Génier 1996: 28 (Venezuela: Mérida and Táchira). — Krajcik 2012: 174 (Venezuela). — Ferrer-Paris et al. 2013: 109 (Venezuela: Mérida: La Azulita-Jají). Ontherus kirschii Harold, 1867 Fig. 57A Ontherus kirschii Harold, 1867d: 96 (original description). Type locality: Colombia: Cundinamarca: Bogotá. Name-bearing type: lectotype (MNHN), designated by Génier (1996), not examined. Ontherus elongatus Waterhouse, 1891a: 357 (original description). Type locality: Venezuela. Namebearing type: holotype (BMNH), not examined. Ontherus kirschii – Harold 1869a: 503 (comments); 1869c: 59 (comments); 1869d: 1008 (catalogues, cited for “Bogota”); 1880: 21 (distribution, comments). — Kirsch 1871: 357 (comments). — Gillet 1911b: 58 (catalog, cited for Colombia). — Luederwaldt 1931a: 396, 399 (key, cited Colombia). — Blackwelder 1944: 206 (list, cited for Colombia). — Escobar 2000: 209 (checklist for Colombia). — Pulido-Herrera et al. 2007: 307 (cited for Andean region of Colombia). — Orozco & Pérez 2008: 38–39 (list for Colombia). — Medina & Pulido-Herrera 2009: 60 (diversity). Ontherus elongatus – Luederwaldt 1931a: 364, 396, 399 (comments, key). — Blackwelder 1944: 206 (list). — Roze 1955: 44 (cited, checklist for Venezuela). — Havranek 1989: 61 (list). — Génier 1996: 35 (revision, redescription, cited as synonym of O. kirschii). Ontherus (Caelontherus) kirschii – Génier 1996: 35 (revision, redescription). Material examined VENEZUELA ‒ Aragua • 1 spec.; Choroni bei Maracay; 1500 m a.s.l.; 26 Mar. 1995; O. Hillert leg.; CEMT. ‒ Táchira • 2 specs; May 1999; CEMT. ‒ Yaracuy • 2 specs; Bolívar, Aroa; 10°22′9.37″ N, 68°50′19.35″ W; 1624 m a.s.l.; 19 Jul. 2009; M. Asmüssen, P. Colmenares and H. Martínez leg.; human faeces; CEMT. Distribution Colombia and Venezuela. Subregions of Venezuela Coastal mainland, Plains, Andes mountains, and Central Coast Mountain Range. European Journal of Taxonomy 959: 1–272 (2024) 148 Fig. 57. Species distribution. A. Ontherus kirschii Harold, 1867. B. Ontherus lichyi Martínez, 1947. Green diamond = CEMT collection data; black circle = literature data. LOZANO DE LA ROSA C. et al., Dung beetles (Scarabaeinae) of Venezuela 149 Literature records Gillet 1911b: 58 (Venezuela). — Luederwaldt 1931a: 396, 399, 414, 416 (Venezuela). — Blackwelder 1944: 206 (Venezuela). — Havranek 1989: 61 (Venezuela: Táchira). — Génier 1996: 35 (Venezuela: Aragua, Barinas, Distrito Capital, Táchira and Trujillo). Ontherus lichyi Martínez, 1947 Fig. 57B Ontherus lichyi Martínez, 1947c: 45 (original description). Type locality: Venezuela: Carabobo: Borburata River valley, 675 m a.s.l. Name-bearing type: holotype (MACN), examined by MC. Ontherus mirandai Pereira, 1954b: 461 (original description). Type locality: Brazil: Amazonas: Rio Negro. Name-bearing type: holotype (MZSP), not examined. Ontherus lichyi – Roze 1955: 44 (cited, checklist for Venezuela). — Pereira & Martínez 1960: 49 (distribution). — Vulcano & Pereira 1967: 582 (key). — Blanco 1987: 44 (catalogue). — Escobar 2000: 209 (checklist for Colombia). — Medina et al. 2001: 139 (checklist for Colombia). — Noriega et al. 2007: 81 (list for Colombia). — Pulido-Herrera et al. 2007: 307 (cited for Andean region of Colombia). — Medina & Pulido-Herrera 2009: 60 (diversity). — Krajcik 2012: 174 (list). Ontherus mirandai – Pereira & Martínez 1960: 49 (cited as synonym of Ontherus lichyi). Ontherus (Ontherus) lichyi – Génier 1996: 100 (revision, redescription). — Vaz-de-Mello 2000: 194 (checklist for Brazil). — Hielkema & Hielkema 2019: 57 (catalogue for the Guianas). Material examined Paratype VENEZUELA ‒ 1 ♂; “VENEZUELA/ E° CARABOBO/ Valle Rio Borhu_/ rata, 670 m. alt./ R. Lichylegit:/ Jul-941”; “Ontherus/ lichyi/ sp. n./ A. Martínez-DET.1946/ ♂”; “PARATIPO”; CEMT/ CUIABÁ/ 00078190. Additional material VENEZUELA ‒ Miranda • 1 spec.; Altos de Pipe, Instituto Venezolano de Investigaciones Científicas (“Campus IVIC”); 10°23′51″ N, 66°58′15″ W; 1500 m a.s.l.; Jul. 2009; F.Z. Vaz-de-Mello leg.; CEMT. Distribution Colombia, Venezuela, Suriname, French Guiana, and Brazil. Subregion of Venezuela Central Coast Mountain Range. Literature records Vulcano & Pereira 1967: 582 (Venezuela). — Blanco 1987: 44 (Venezuela: Táchira). — Génier 1996: 100 (Venezuela: Carabobo and Distrito Capital). — Krajcik 2012: 174 (Venezuela). Ontherus pubens Génier, 1996 Fig. 58A Ontherus (Ontherus) pubens Génier, 1996: 71 (original description). Type locality: Ecuador: Napo: Puerto Misahuallí: Reserva Biológica Jatun Sacha, 21 km E of Puerto Napo, 400 m. Name-bearing type: holotype (CMNC), not examined. European Journal of Taxonomy 959: 1–272 (2024) 150 Ontherus pubens – Amézquita et al. 1999: 119 (diversity). — Escobar 2000: 209 (list). — Medina et al. 2001: 139 (checklist for Colombia). — Hamel-Leigue et al. 2006: 16 (inventory, cited for Bolivia). — Pulido-Herrera et al. 2007: 307 (list for Andean region, Colombia). — Donoso et al. 2009: Appendix II, 17 (catalogue of types specs at the MQCAZ). — Krajcik 2012: 174 (checklist, cited for Ecuador). — Bezděk & Hájek 2012: 313 (catalogue of type specs). — Ratcliffe et al. 2015: 197 (checklist, cited for Peru). Ontherus (Ontherus) pubens – Vaz-de-Mello 2000: 194 (checklist for Brazil). — Morón 2006: 121 (catalogue of MXAL types). — Carvajal et al. 2011: 318–319 (cited for Ecuador). — Chamorro et al. 2018: 90, 96, fig. 13e (key, cited for Ecuador); 2019: 177 (catalogue). — Hielkema & Hielkema 2019: 57 (catalogue for the Guianas). Distribution Colombia, Venezuela, Brazil, Ecuador, Peru, Bolivia, and Argentina. Subregions of Venezuela Plains and Andes mountains. Literature records Génier 1996: 71 (Venezuela: Apure and Táchira). — Bezděk & Hájek 2012: 313 (Venezuela: Apure). Ontherus sulcator (Fabricius, 1775) Fig. 58B Scarabaeus sulcator Fabricius, 1775: 27 (original description). Type locality: French Guiana. Namebearing type: according to Zimsen (1964), three syntypes in ZMUK, but deemed lost without further explanation by Génier (1996). We ourselves have made no attempt to locate them, and so they may well be in ZMUK as Zimsen said. Copris nisus Castelnau, 1840: 79 (original description). Type locality: French Guiana. Name-bearing type: lectotype (MVMA), designated by Génier (1996), not examined. Copris ovalipennis Blanchard, 1846: 180 (original description). Type locality: Argentina: Corrientes. Name-bearing type: lectotype (MNHN), designated by Génier (1996), not examined. Copris senegalensis Gillet, 1910: 15 (original description). Type locality: Unknown. Originally said to be Senegal (“Sénégal”), but this is incorrect (Gillet 1911a). Name-bearing type: syntypes (RBINS) (Gillet 1910), not examined. Scarabaeus sulcator – Olivier 1789: 142 (redescription). — Fabricius 1792: 61 (redescription). Copris sulcator – Fabricius 1801: 53 (redescription). — Olivier, 1790: 168 (redescription). — Martínez 1959: 71 (cited). Copris nisus – Blanchard, 1846: 180 (comments). — Burmeister 1874: 126 (list for Argentina). — Martínez 1959: 71 (cited as synonym of Ontherus sulcator). Ontherus sulcator – Erichson 1847b: 761 (comments). — Harold 1869c: 59 (comments); 1869d: 1008 (catalogue). — Burmeister 1874: 126 (list for Argentina). — Gillet 1911a: 319 (list); 1911b: 58 (catalogue). — Bruch 1915: 541 (list). — Lucas 1920: 459 (catalogue). — Luederwaldt 1930: 106 (comments); 1931a: 371, 385, 389, 392, 408, 411, 416 (key, synonymy). — Pessôa & Lane 1941: 456, 458 (key). — Blackwelder 1944: 206 (checklist). — Martínez 1959: 71 (catalogue). — Vulcano & Pereira, 1967: 583 (key). — Forsyth & Gill 1993: 70 (as O. suclator [sic], cited for Guyana). — Escobar 2000: 209 (list). — Vaz-de-Mello 2000: 194 (checklist for Brazil). — Medina et al. 2001: 139 (checklist for Colombia). — Hamel-Leigue et al. 2006: 16 (inventory, cited for Bolivia). — Feer 2008: 62 (ecology, list). — Larsen et al. 2008: 1294 (list). — Krajcik 2012: 174 LOZANO DE LA ROSA C. et al., Dung beetles (Scarabaeinae) of Venezuela 151 Fig. 58. Species distribution. A. Ontherus pubens Génier, 1996. B. Ontherus sulcator (Fabricius, 1775). Black circle = literature data. European Journal of Taxonomy 959: 1–272 (2024) 152 (checklist, cited for Cayenne). — Boilly & Vaz-de-Mello 2013: 112 (Fig. 43, key). — Ferrer-Paris et al. 2013: 109 (list). — Ratcliffe et al. 2015: 197 (checklist, cited for Peru). Copris ovalipennis – Harold 1869a: 503 (comments); 1869c: 59 (comments). — Burmeister 1874: 126 (list for Argentina). — Luederwaldt 1931a: 386 (cited as synonym of Ontherus nisus). — Martínez 1959: 71 (cited as synonym of Ontherus sulcator). Ontherus nisus – Harold 1869d: 1008 (catalogue, cited as synonym of O. sulcator). — Gillet 1911b: 58 (catalog, cited as synonym of O. sulcator). — Luederwaldt 1931a: 387, 392, 399, 400, 409, 411, 417 (redescription, distribution, key). — Pessôa & Lane 1941: 457, 458 (key, redescription). — Blackwelder 1944: 206 (cited as synonym of O. sulcator). — Lange 1947: 310 (list). — Martínez 1959: 71 (cited as synonym of O. sulcator). Pinotus ovalipennis – Harold 1869d: 1008 (catalogue, cited for Argentina: Corrientes). Ontherus ovalipennis – Gillet 1911b: 58 (catalog, cited as synonym of O. sulcator). — Luederwaldt 1931a: 387 (cited as synonym of O. nisus). — Pessôa & Lane 1941: 458 (cited as synonym of O. nisus). — Blackwelder 1944: 206 (cited as synonym of O. sulcator). — Martínez 1959: 71 (cited as synonym of O. sulcator). Copris senegalensis – Gillet 1911a: 319 (cited as synonym of O. sulcator); 1911b: 58 (catalog, cited as synonym of O. sulcator). — Bruch 1915: 541 (list, cited as synonym of O. sulcator). — Martínez 1959: 72 (cited as synonym of O. sulcator). Ontherus sulcator – Luederwaldt 1931a: 364 (type species). Ontherus senegalensis – Luederwaldt 1931a: 386, 387 (cited as synonym of Ontherus nisus). — Pessôa & Lane 1941: 458 (cited as synonym of O. nisus). — Blackwelder 1944: 206 (cited as synonym of O. sulcator). Ontherus (Ontherus) sulcator – Génier 1996: 74‒76 (revision). — Hielkema & Hielkema 2019: 57 (catalogue for the Guianas). Distribution Colombia, Venezuela, Trinidad and Tobago, Guyana, Suriname, French Guiana, Brazil, Ecuador, Peru, Bolivia, Paraguay, Argentina, and Uruguay (Génier 1996). Subregions of Venezuela System of hills and low piedmont mountains of the Guiana Shield, System of low mountains and hills Imataca-Cuyuní of Northeast Guyana Shield, and Guiana Shield. Literature records Luederwaldt 1931a: 387, 409, 417 (Venezuela). — Pessôa & Lane 1941: 457–458 (Venezuela). — Vulcano & Pereira 1967: 583 (Venezuela). — Génier 1996: 74 (Venezuela: Bolívar). — Larsen et al. 2008: 1294 (Venezuela: Bolívar). — Ferrer-Paris et al. 2013: 109 (Venezuela: Bolívar: Isla de Anacoco). Genus Onthophagus Latreille, 1802 Onthophagus Latreille, 1802: 141 (original description). Type species: Scarabaeus taurus Schreber, 1759, by original monotypy. Chalcoderus Erichson, 1847b: 763 (original description). Type species: no fixation known to us. Monapus Erichson, 1847b: 763 (original description). Type species: no fixation known to us; no nominal species seem to have been originally included. Chamorro et al. (2019) stated the type species was “probably” Onthophagus mniszechi Harold, 1869, but did not explain why they thought so. Psilax Erichson, 1847b: 764 (original description). Type species: Onthophagus pronus Erichson, 1842, by original monotypy. LOZANO DE LA ROSA C. et al., Dung beetles (Scarabaeinae) of Venezuela 153 Macropocopris Arrow, 1920: 435 (original description). Type species: Macropocopris prehensilis Arrow, 1920, by subsequent designation of Matthews (1972). Onthophagus – Le Peletier de Saint-Fargeau & Audinet-Serville 1828: 353 (redescription). — Agassiz 1846: 749 (catalogue). — Latreille 1829: 536 (redescription). — Brullé 1838: 300 (redescription). — Castelnau 1840: 83 (redescription). — Lacordaire 1855: 107 (redescription). — Harold 1869d: 1024 (catalogue). — Gillet 1911b: 118 (catalogue). — Gillet & Boucomont 1927: 205 (catalogue). — Dawson 1922: 61 (key). — Paulian 1938: 232 (key). — Blackwelder 1944: 211 (list). — Roze 1955: 45 (checklist for Venezuela). — Martínez 1959: 108 (catalogue for Argentina). — Howden & Cartwright 1963: 6 (redescription). — Halffter & Matthews 1966: 254 (catalog, distribution). — Vulcano & Pereira 1967: 562 (key). — Howden & Young 1981: 11, 93 (key, redescription). — Halffter & Edmonds 1982: 135 (catalog, distribution). — Huchet 1992: 300 (key). — Medina & Lopera-Toro 2000: 301 (key). — Vaz-de-Mello 2000: 194 (checklist for Brazil). — Medina et al. 2001: 139 (list for Colombia). — Kohlmann & Solís 2001: 160 (redescription). — Ratcliffe 2002: 17 (checklist for Panama). — Ratcliffe et al. 2002: 49 (key). — Hamel-Leigue et al. 2006: 16 (list for Bolivia). — Pulido-Herrera & Zunino 2007: 94 (catalogue). — Vaz-de-Mello et al. 2011a: 23 (key). — Carvajal et al. 2011: 145, 322 (diagnosis, list). — Krajcik 2012: 174 (list). — Solís & Kohlmann 2012: 8 (checklist for Costa Rica). — Chamorro et al. 2018: 74, 96–97 (list for Ecuador); 2019: 179 (catalogue). — Hielkema & Hielkema 2019: 90 (catalogue for the Guianas). Chalcoderus – Lacordaire 1855: 109 (comments). — Harold 1869d: 1024 (cited as synonym of Onthophagus). — Gillet 1911b: 118 (cited as synonym of Onthophagus). — Blackwelder 1944: 211 (cited as synonym of Onthophagus). — Solís & Kohlmann 2012: 8 (cited as synonym of Onthophagus). Monapus – Lacordaire 1855: 109 (comments). — Harold 1869d: 1024 (cited as synonym of Onthophagus). — Gillet 1911b: 118 (cited as synonym of Onthophagus). — Blackwelder 1944: 211 (cited as synonym of Onthophagus). — Solís & Kohlmann 2012: 8 (cited as synonym of Onthophagus). Psilax – Lacordaire 1855: 109 (comments). — Harold 1869d: 1024 (cited as synonym of Onthophagus). — Gillet 1911b: 118 (cited as synonym of Onthophagus). — Blackwelder 1944: 211 (cited as synonym of Onthophagus). — Solís & Kohlmann 2012: 8 (cited as synonym of Onthophagus). Macropocopris – Halffter & Matthews 1966: 254 (cited as genus). — Halffter & Edmonds 1982: 135 (cited as synonym of Onthophagus). — Solís & Kohlmann 2012: 8 (cited as synonym of Onthophagus). Onthophagus bidentatus Drapiez, 1819 Fig. 59A Onthophagus bidentatus Drapiez, 1819: 134 (original description). Type locality: French Guiana. Namebearing type: lectotype (MNHN), designated by Rossini et al. (2016), examined by FZVM. Onthophagus bicornis Castelnau, 1840: 87 (original description). Type locality: French Guiana. Namebearing type: lectotype (MNHN), designated by Rossini et al. (2016), examined by FZVM. Onthophagus femoralis Kirsch, 1871: 362 (original description). Type locality: Colombia: Cundinamarca: Bogotá. Name-bearing type: unknown to us. Onthophagus semichalcites d’Orbigny, 1902: 149 (original description). Type locality: unknown; the lectotype is mislabelled as though coming from Benin (Rossini et al. 2016). Name-bearing type: lectotype (MNHN), designated by Rossini et al. (2016), examined by FZVM. Onthophagus bidentatus – Harold 1869d: 1026 (catalogue); 1880: 33 (redescription, distribution). — Gillet 1911b: 204 (catalogue). — Bruch 1911: 190 (cited for Argentina). — Gillet & Boucomont 1927: 204 (catalogue of species). — Boucomont 1932: 304, 321 (key, distribution). — Balthasar European Journal of Taxonomy 959: 1–272 (2024) 256 ingenia et mores describuntur et iconibus supra quingentas illustrantur: 245–259. Franciscum Hackium, Leiden [Lugdun] & Elzevirium, Amsterdam [Amstelodami]. https://doi.org/10.5962/bhl.title.48544 Marchiori C.H., Caldas E.R. & Almeida K.G.S. 2003. Succession of Scarabaeidae on bovine dung in Itumbiara, Goiás, Brazil. Neotropical Entomology 32: 173‒176. https://doi.org/10.1590/S1519-566X2003000100029 Martínez A. 1947a. Insectos nuevos o poco conocidos V. El género Pseudocanthon Bates, y algunas especias nuevos o poco conocidos de Scarabaeidae. Revista de la Sociedad Entomológica Argentina 13 (1‒5): 263‒280. Martínez A. 1947b. Addenda y corrigenda al trabajo de Blackwelder “Checklist of the coleopterous insects of Mexico, Central America, the West Indies, and South America” (Scarabaeidae, Coprinae). Arthropoda 1 (1): 109–114. Martínez A. 1947c. Lamellicornia neotropica I. Especies nuevas o poco conocidas de pinotides (Col. Scarab.). Arthropoda 1 (1): 41–53. Martínez A. 1948a. Notas coleopterológicas I. Anales de la Sociedad Científica Argentina 146 (1): 41– 50. Martínez A. 1948b. Insectos nuevos o poco conocidos — VI (Col. Scarabaeidae) dos nuevos generos de Lamellicornia (Coleoptera). Revista de la Sociedad Entomológica Argentina 14 (12): 3–11. Martínez A. 1949a. Notas coleopterológicas III. Anales de la Sociedad Científica Argentina 148 (5): 281–291. Martínez A. 1949b. Insectos nuevos o poco conocidos VII. Revista de la Sociedad Entomólogica Argentina 14 (4): 175–193. Martínez A. 1950. Notas coleopterologicas IV. Nuevo subgénero y especies de Glaphyrocanthon Martínez, 1948 (Col. Scarabaeineae [sic]). Anales de la Sociedad Científica Argentina 150: 159–172. Martínez A. 1951. Notas coleopterológicas V. La invalidez del nombre genérico Pinotus Erichson y dos nuevas sinonimias. Anales de la Sociedad Científica Argentina 152: 138–142. Martínez A. 1954. Fauna de los Parques Nacionales. Natura, Buenos Aires 1 (1): 59–74. Martínez A. 1959. Catálogo de los Scarabaeidae Argentinos. Revista del Museo Argentino de Ciencias Naturales “Bernardino Rivadavia” 5: 1–126. Martínez A. 1961. Scarabaeoidea Neotropica VIII. Notas sobre el complejo Megathopa Eschscholtz, 1822. Neotropica 7 (24): 81–86. Martínez A. 1987. La entomofauna de Scarabaeinae de la provincia de Salta (Col. Scarabaeoidea). Anales de la Sociedad Científica Argentina 216: 45–69. Martínez A. 1988a. Algunos Scarabaeinae neotropicales nuevos o poco conocidos. Revista Científica Unet 2 (1): 85–90. Martínez A. 1988b. Notas sobre Eurysternus Dalman. Entomologica Basiliensia 12: 279–304. Martínez A. 1989. 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Manuscript received: 13 August 2022 Manuscript accepted: 7 November 2023 Published on: 11 October 2024 Topic editor: Tony Robillard Section editor: Max Barclay Desk editor: Pepe Fernández Printed versions of all papers are deposited in the libraries of four of the institutes that are members of the EJT consortium: Muséum national dʼHistoire naturelle, Paris, France; Meise Botanic Garden, Belgium; Royal Museum for Central Africa, Tervuren, Belgium; Royal Belgian Institute of Natural Sciences, Brussels, Belgium. The other members of the consortium are: Natural History Museum of Denmark, Copenhagen, Denmark; Naturalis Biodiversity Center, Leiden, the Netherlands; Museo Nacional de Ciencias Naturales-CSIC, Madrid, Spain; Leibniz Institute for the Analysis of Biodiversity Change, Bonn – Hamburg, Germany; National Museum of the Czech Republic, Prague, Czech Republic; The Steinhardt Museum of Natural History, Tel Aviv, Israël.