Abstract

The Pantanal is the largest seasonal freshwater wetland on Earth, characterized by the seasonal flooding and complex mosaic vegetation, which determines its biodiversity. Among this biodiversity, dung beetles (Coleoptera: Scarabaeidae: Scarabaeinae) are a group of insects that perform important ecological functions, such as: nutrient cycling, seed dispersion and parasite control. In order to mitigate the lack of information on dung beetle fauna of the Brazilian Pantanal, we conducted a bibliographic search of virtually all literature published until november-2020 on dung beetles sampled in the Pantanal. In addition, we had accessed to the records the largest collection of the dung beetle species from Brazilian Pantanal. We recorded 68 dung beetle species of 30 genera. The genera Canthon Holffmanseg 1847 is the most diverse with 13 species recorded. Although our knowledge about the dung beetle fauna in this ecosystem is still incipient, our results demonstrated a high richness of dung beetles in the Brazilian Pantanal. In addition, our study provides first list of dung beetle species and an illustrated dichotomy key to identify genera and some species occurring in the Brazilian Pantanal. Thus, the use of this guide for identification of dung beetle species and a list of species can be important tools to help researchers and provide incentive for new inventories on dung beetle fauna in the Brazilian Pantanal.

Keywords
Checklist; insect diversity; humid zones; dichotomous key; wetlands

Resumo

O Pantanal é a maior área úmida sazonal de água doce Neotropical da Terra, caracterizada pelas inundações sazonais e pela complexa vegetação em mosaico, que determina sua biodiversidade. Dentre essa biodiversidade, os besouros rola-bosta (Coleoptera: Scarabaeidae: Scarabaeinae) são um grupo de insetos que desempenham importantes funções ecológicas, tais como: ciclagem de nutrientes, dispersão de sementes e controle de parasitas. A fim de mitigar a falta de informações sobre a fauna de besouros rola-bosta do Pantanal brasileiro, realizamos uma pesquisa bibliográfica de praticamente toda a literatura publicada até novembro de 2020 sobre besouros rola-bosta amostrados no Pantanal. Além disso, acessamos os registros da maior coleção da espécie de rola-bosta do Pantanal brasileiro. Registramos 68 espécies de besouros rola-bosta de 30 gêneros. O gênero Canthon Holffmanseg 1847 é o mais diverso com 13 espécies registradas. Embora nosso conhecimento sobre a fauna de rola-bostas neste ecossistema ainda seja incipiente, nossos resultados demonstraram uma alta riqueza de besouros rola-bosta no Pantanal brasileiro. Além disso, nosso estudo fornece a primeira lista de espécies de besouros rola-bosta e uma chave de dicotomia ilustrada para identificar gêneros e algumas espécies encontrados no Pantanal brasileiro. Assim, a utilização deste guia para identificação das espécies de besouros rola-bostas e uma lista de espécies podem ser ferramentas importantes para auxiliar pesquisadores e incentivar novos inventários sobre a fauna de besouros rola-bostas no Pantanal brasileiro.

Palavras-chave
Checklist; diversidade de insetos; áreas úmidas; chaves dicotomias; áreas alagadas

Introduction

The Pantanal is the largest seasonal freshwater wetland on Earth (ca. 160,000 km²), and also considered a World Heritage Site and Biosphere Reserve (UNESCO, 2000UNESCO. 2000. World heritage. United Nations Educational, Scientific and Cultural Organization, Cairns.). This extensive wetland is located in Brazil (states of Mato Grosso and Mato Grosso do Sul) with approximately 130,000 km², 15,000 km² in Bolivia and 5,000 km² in Paraguay (Cunha et al. 2014CUNHA, C.N., PIEDADE, M.T.F. & JUNK,W.J. 2014. Classificação e delineamento das áreas úmidas brasileiras e de seus microhabitats. EdUFMT, Cuiabá, p.1–156.) and has a vegetation system arranged in a mosaic (i.e., different plant communities scattered side-by-side across space) (Pott et al. 2011POTT, A., OLIVEIRA, A., DAMASCENO-JUNIOR, G. & SILVA, J.S.V. 2011. Plant diversity of the Pantanal wetland. Braz. J. Biol. 71:265–273.). Furthermore, two well-defined ecohydrology cycles can be identified in the Pantanal: dry and rainy. In the dry season, the surface water becomes scarce, being restricted to the perennial rivers (with defined beds) and large ponds (Alho & Sabino 2011ALHO, C.J.R. & SABINO, J. 2011. A conservation agenda for the Pantanal’s biodiversity. Braz. J. Biol. 71:327–335.; Nunes et al. 2014). Thus, the seasonal flooding and complex mosaic vegetation determine a biodiversity unique adapted to Brazilian Pantanal (Junk et al. 2006JUNK, W.J., CUNHA, C.N., WANTZEN, K.M., PETERMANN, P., STRÜSSMANN, C., MARQUES, M.I. et al. 2006. Biodiversity and its conservation in the Pantanal of Mato Grosso, Brazil. Aquat. Sci. 69:278–309.; Tomas et al. 2019TOMAS, W.M., DE OLIVEIRA, R.F., MORATO, R.G., MEDICI, P.E., CHIARAVALLOTI, R., TORTATO, F. et al. 2019. Sustainability agenda for the Pantanal wetland: perspectives on a collaborative interface for science, policy, and decision-making. Trop. Conserv. Sci. 12:1940082919872634.).

Among this biodiversity, insects, such as dung beetles (Coleoptera: Scarabaeidae: Scarabaeinae), are essential for the maintenance of ecosystem functioning in the Brazilian Pantanal (Correa et al. 2020CORREA, C.M.A., AUDINO, L.D., HOLDBROOK, R., BRAGA, R.F., MENÉNDEZ, R. & LOUZADA, J. 2020. Successional trajectory of dung beetle communities in a tropical grassy ecosystem after livestock grazing removal. Biodivers. Conserv. 29:2311–2328.). Dung beetles are broadly distributed and highly diverse taxonomically and functionally (Halffter & Edmonds 1982HALFFTER, G. & EDMONDS, W.D. 1982. Nesting Behavior of Dung Beetles (Scarabaeinae). Man and Biosphere Program – UNESCO, Mexico, p.1–176.; Hanski & Cambefort 1991HANSKI, I. & CAMBEFORT, Y. (1991) Dung beetle ecology. Princeton, NJ: Princeton University Press, p.1–481.). They are represented by more than 6,837 species worldwide which belong to 278 genera (Schoolmeesters 2023SCHOOLMEESTERS, P. 2023. World Scarabaeidae Database. In: Bánki O., Roskov Y., Döring M., Ower G., Vandepitte L., Hobern D., Remsen D., Schalk P., DeWalt R.E., Keping M., Miller J., Orrell T., Aalbu R., Abbott J., Adlard R., Adriaenssens E.M., Aedo C., Aescht E., Akkari N., et al. (eds) Catalogue of Life Checklist (Version 2023–03-06). Available from https://www.catalogueoflife.org/data/dataset/1027 [last accessed 03 Apr 2023].
https://www.catalogueoflife.org/data/dat... ). Their feeding behavior is quite varied, most species being coprophagous, but there are also species that feed on decaying carcasses, fungi and fruits, and can even be predators or generalists (Halffter & Matthews 1966HALFFTER, G. & MARTINEZ, A. 1966. Revision monographica de los Canthonina americanos (Coleoptera, Scarabaeidae) (1a. parte). Rev. Soc. Mex. Hist. Nat. 27:89–177.). They can be classified into three functional groups based on food allocation behavior and/or nesting behavior: (1) roller (telecoprid) species that remove portions of dung, which are rolled away from the food resource and then buried; (2) tunneler (paracoprid) species that construct tunnels below or adjacent to the food resource and transport dung into them; and (3) dweller (endocoprid) species that nest within a dung pad and do not exhibit resource allocation (Halffter & Matthews 1966HALFFTER, G. & MATTHEWS, E.G. 1966. The natural history of dung beetles of the subfamily Scarabaeinae (Coleoptera: Scarabaeidae). Folia Entomol. Mex. 12:1–312.; Hanski & Cambefort 1991HANSKI, I. & CAMBEFORT, Y. (1991) Dung beetle ecology. Princeton, NJ: Princeton University Press, p.1–481.).

When dung beetles bury food resources (e.g. feces, carcass and fruits), they provide important ecological functions and services in natural and anthropogenic ecosystems, such as: increased water infiltration and soil porosity, improved herbaceous plant growth, reduction of livestock gastrointestinal parasite and dung-fly availability (Nichols et al. 2008NICHOLS, E., SPECTOR, S., LOUZADA, J.N.C., LARSEN, T., AMEZQUITA, S. & FAVILA, M.E. 2008. Ecological functions and ecosystem services provided by Scarabaeinae dung beetles. Biol. Conserv. 141:1461–1474.) and greenhouse gas emissions from dung pats (Slade et al. 2016SLADE, E.M., RIUTTA, T., ROSLIN, T. & TUOMISTO, H.L. 2016. The role of dung beetles in reducing greenhouse gas emissions from cattle farming. Sci. Rep. 6:1–8.; Piccini et al. 2017PICCINI, I., ARNIERI, F., CAPRIO, E., NERVO, B., PELISSETTI, S., PALESTRINI, C., ROSLIN, T. & ROLANDO, A. 2017. Greenhouse gas emissions from dung pats vary with dung beetle species and with assemblage composition. PLoS ONE 12:e0178077.). In addition, these beetles have been widely used as both ecological and biodiversity indicators because they are sensitive to natural and anthropogenic changes (Halffter & Favila 1993HALFFTER, G. & FAVILA, M.E. 1993. The Scarabaeinae (Insecta, Coleoptera), an animal group for analyzing, inventorying, and monitoring biodiversity in tropical rain forest and modified landscapes. Biol. Int. 27:15–21.; McGeoch et al. 2002McGEOCH, M.A., van RENSBURG, B.J. & BOTES, A. 2002. The verification and application of bioindicators: a case study of dung beetles in a savanna ecosystem. J. Appl. Ecol. 39:661–672.; Nichols et al. 2007NICHOLS, E., LARSEN, T., SPECTOR, S., DAVIS, A.L., ESCOBAR, F., FAVILA, M. & VULINEC, K. 2007. Global dung beetle response to tropical forest modification and fragmentation: a quantitative literature review and meta-analysis. Biol. Conserv. 137:1–19.), are easily sampled with standardized protocols (Correa et al. 2018CORREA, C.M.A., BRAGA, R.F., PUKER, A., ABOT, A.R. & KORASAKI, V. 2018. Optimising methods for dung beetle (Coleoptera: Scarabaeidae) sampling in Brazilian pastures. Environ. Entomol. 47:48–54.; Gardner et al. 2008GARDNER, T.A., BARLOW, J., ARAUJO, I.S., ÁVILA-PIRES, T.C., BONALDO, A.B., COSTA, J.E. et al. 2008. The cost-effectiveness of biodiversity surveys in tropical forests. Ecol. Lett. 11:139–150.; da Silva & Hernández 2015da SILVA, P.G. & HERNÁNDEZ, M.I.M. 2015. Spatial patterns of movement of dung beetle species in a tropical forest suggest a new trap spacing for dung beetle biodiversity studies. PLoS ONE 10:e0126112.) and are diverse and relatively well characterized taxonomically (Vaz-de-Mello et al. 2011VAZ-DE-MELLO, F.Z., EDMONDS, W.D., OCAMPO, F.C. & SCHOOLMEESTERS, P. 2011. A multilingual key to the genera and subgenera of the subfamily Scarabaeinae of the New World (Coleoptera: Scarabaeidae). Zootaxa 2854:1–73.).

Although studies on dung beetle assemblages have been growing recently in the Brazilian Pantanal (Tissiani et al. 2015TISSIANI, A.S.O., SOUSA, W.O., SANTOS, G.G., IDE, S., BATTIROLA, L. & MARQUES, M.I. 2015. Environmental influence on coprophagous Scarabaeidae (Insecta, Coleoptera) communities in the Pantanal of Mato Grosso. Braz. J. Biol. 75:136–142.; Daniel & Vaz-de-Mello 2016DANIEL, G.M. & VAZ-DE-MELLO, F.Z. 2016. Biotic components of dung beetles (Insecta: Coleoptera: Scarabaeidae: Scarabaeinae) from Pantanal – Cerrado Border and its implications for Chaco regionalization. J. Nat. Hist. 50:1159–1173., Pessôa et al. 2017PESSÔA, M.B., IZZO, T.J. & VAZ-DE-MELLO, F.Z. 2017. Assemblage and functional categorization of dung beetles (Coleoptera: Scarabaeinae) from Pantanal. PeerJ 5:e3978.; Correa et al. 2016CORREA, C.M.A., PUKER, A., FERREIRA, K.R., CRISTALDO, C.M., FERREIRA, F.N.F., ABOT, A.R. & KORASAKI, V. 2016. Using dung beetles to evaluate the conversion effects from native to introduced pasture in the Brazilian Pantanal. J. Insect Conserv. 20:447–456., 2019CORREA, C.M.A., BRAGA, R.F., LOUZADA, J. & MENÉNDEZ, R. 2019. Dung beetle diversity and functions suggest no major impacts of cattle grazing in the Brazilian Pantanal wetlands. Ecol. Entomol. 44:524–533., 2020CORREA, C.M.A., AUDINO, L.D., HOLDBROOK, R., BRAGA, R.F., MENÉNDEZ, R. & LOUZADA, J. 2020. Successional trajectory of dung beetle communities in a tropical grassy ecosystem after livestock grazing removal. Biodivers. Conserv. 29:2311–2328., 2021CORREA, C.M.A., da SILVA, P.G., PUKER, A. & ABOT, A. 2021. Spatiotemporal patterns of taxonomic and functional β-diversity of dung beetles in native and introduced pastures in the Brazilian Pantanal. Austral Ecol. 46:98–110., 2022aCORREA, C.M.A., AUDINO, L.D., VAZ-DE-MELLO, F.Z. & LOUZADA, J. 2022a. Diversity and structure of dung beetle (Coleoptera: Scarabaeidae) assemblage in natural grasslands of the Brazilian Pantanal. Int. J. Trop. Insect Sci. 42:3253–3261., bCORREA, C.M.A., FERREIRA, K.R., ABOT, A.R., LOUZADA, J. & VAZ-DE-MELLO, F.Z. 2022b. Ivermectin impacts on dung beetle diversity and their ecological functions in two distinct Brazilian ecosystems. Ecol. Entomol. 47:736–748.; Correa & da Silva 2022CORREA, C.M.A. & da SILVA, P.G. 2022. Environmental drivers of taxonomic and functional diversity of dung beetles across a chronosequence of tropical grasslands with different cattle grazing removal ages. Austral Ecol. 47:928–938., Gonçalves et al. 2022GONÇALVES, T.F., CORREA, C.M.A., AUDINO, L.D., VAZ-DE-MELLO, F.Z., FONTOURA, F.M. & GUEDES, N.M.R. 2022. Quantifying the post-fire recovery of taxonomic and functional diversity of dung beetles in the Brazilian Pantanal. Ecol. Entomol. 47:601–612.), our knowledge on dung beetle fauna in this biome is incipient. Indeed, the Pantanal is one of the least known ecosystems in terms of biodiversity of Brazil (Lewinsohn et al. 2005LEWINSOHN, T.M., FREITAS, A.V.L. & PRADO, P.I. 2005. Conservation of terrestrial invertebrates and their habitats in Brazil. Conserv. Biol. 19:640–645.). Therefore, new inventories of dung beetles in different localities are essential to increase new records of dung beetle species in the Brazilian Pantanal. In this study, we bring a list of dung beetle species of the Brazilian Pantanal and an illustrated identification key.

Material and Methods

We used the Scopus and Web of Science databases to search for literature on the dung beetles in the Brazilian Pantanal, following the PRISMA methodology (Moher et al. 2009MOHER, D., LIBERATI, A., TETZLAFF, J., ALTMAN, D.G., & PRISMA GROUP. 2009. Preferred reporting items for systematic reviews and meta-analyses: the PRISMA statement. Ann. Intern. Med. 151:264–269.), which only considers indexed articles. The following search terms were used: ((“Dung beetle*” OR “Scarabaeinae*” OR “coprophagous beetles*”) AND (“Wetland*” OR “Pantanal*” OR “humid zones”)). We complemented the taxonomic diversity search by including articles published in Portuguese from the authors’ collection. The search window of time covered articles published between January 2007 to November 2020.

Our search returned 40 articles, regarding dung beetle species in wetlands. The relevant articles were selected using the following criteria: i) the study includes species from Scarabaeinae subfamily, ii) the study is based partly or entirely in Brazilian Pantanal (e.g., Mato Grosso and Mato Grosso do Sul states). Under these criteria, 10 articles were retained for data extraction (e.g. Louzada et al. 2007LOUZADA, J.N.C., LOPES, F.S. & VAZ-DE-MELLO, F.Z. 2007. Structure and composition of a dung beetle community (Coleoptera, Scarabaeinae) in a small forest patch from Brazilian Pantanal. Rev. Bras. Zoociencias 9:199–203.; Rodrigues et al. 2010RODRIGUES, S.R., BARROS, A.T.M., PUKER, A. & TAIRA, T.L. 2010. Diversidade de besouros coprófagos (Coleoptera, Scarabaeidae) coletados com armadilha de interceptação de voo no Pantanal Sul-Mato-Grossense, Brasil. Biota Neotrop. 10:123–127.; Tissiani et al. 2015TISSIANI, A.S.O., SOUSA, W.O., SANTOS, G.G., IDE, S., BATTIROLA, L. & MARQUES, M.I. 2015. Environmental influence on coprophagous Scarabaeidae (Insecta, Coleoptera) communities in the Pantanal of Mato Grosso. Braz. J. Biol. 75:136–142.; Daniel & Vaz-de-Mello 2016DANIEL, G.M. & VAZ-DE-MELLO, F.Z. 2016. Biotic components of dung beetles (Insecta: Coleoptera: Scarabaeidae: Scarabaeinae) from Pantanal – Cerrado Border and its implications for Chaco regionalization. J. Nat. Hist. 50:1159–1173.; Pessôa et al. 2017PESSÔA, M.B., IZZO, T.J. & VAZ-DE-MELLO, F.Z. 2017. Assemblage and functional categorization of dung beetles (Coleoptera: Scarabaeinae) from Pantanal. PeerJ 5:e3978.; Vaz-de-Mello et al. 2017; Correa et al. 2016CORREA, C.M.A., PUKER, A., FERREIRA, K.R., CRISTALDO, C.M., FERREIRA, F.N.F., ABOT, A.R. & KORASAKI, V. 2016. Using dung beetles to evaluate the conversion effects from native to introduced pasture in the Brazilian Pantanal. J. Insect Conserv. 20:447–456., 2019CORREA, C.M.A., BRAGA, R.F., LOUZADA, J. & MENÉNDEZ, R. 2019. Dung beetle diversity and functions suggest no major impacts of cattle grazing in the Brazilian Pantanal wetlands. Ecol. Entomol. 44:524–533., 2020CORREA, C.M.A., AUDINO, L.D., HOLDBROOK, R., BRAGA, R.F., MENÉNDEZ, R. & LOUZADA, J. 2020. Successional trajectory of dung beetle communities in a tropical grassy ecosystem after livestock grazing removal. Biodivers. Conserv. 29:2311–2328.), only species identified at a specific level were considered. In addition, 10 genera have been added to the identification key that have a good chance of being collected in the Brazilian Pantanal region (Vaz-de-Mello personal communication). The genera that are under taxonomic revision are not included in the identification keys with their respective species that occur in the Pantanal.

All previously existing records were reviewed for their identifications (when possible, we used specimen vouchers) for correct species identification. We also list the records of the Entomology Section of the Zoological Collection at the UFMT (CEMT, curator F. Z. Vaz-de-Mello). Currently, it is considered one of the most important collections of dung beetles in the world, with specimens from all regions of Brazil and across the globe, making it possible to assemble a list of species from the different Brazilian biomes (Tissiani et al. 2017TISSIANI, A.S.O., VAZ-DE-MELLO, F.Z. & CAMPELO-JÚNIOR, J.H. 2017. Besouros rola-bostas das pastagens brasileiras e chave para identificação dos gêneros (Coleoptera: Scarabaeidae). Pesq. Agropec. Bras. 52:401–418.). The dung beetle species mentioned in the present study are classified according to their nesting behavior, following the classification by Hanski & Cambefort (1991)HANSKI, I. & CAMBEFORT, Y. (1991) Dung beetle ecology. Princeton, NJ: Princeton University Press, p.1–481..

Finally, an identification key was constructed to identify the genera and species of dung beetles present in Brazilian Pantanal. Its main structure from Vaz-de-Mello et al. (2011)VAZ-DE-MELLO, F.Z., EDMONDS, W.D., OCAMPO, F.C. & SCHOOLMEESTERS, P. 2011. A multilingual key to the genera and subgenera of the subfamily Scarabaeinae of the New World (Coleoptera: Scarabaeidae). Zootaxa 2854:1–73., and this was refined based on the analysis of the external morphology of the species, including – where necessary – the secondary sexual characteristics.

Leica M205A stereomicroscope coupled with a Leica DMC4500 and a Leica Application Suite V4.10.0 Interactive Measurements, Montage was used for the photos.

Results

The dung beetle fauna of the Brazilian pantanal is composed of 68 species in 30 genera. The most diverse genera are: Canthon Hoffmannsegg, 1817 (13 species), Dichotomius Hope, 1838 (nine species) and Ontherus Erichson, 1847 (seven species) (Table 1). Paracoprid beetles correspond to 48.35% of all species recorded, whereas telecoprid and endocoprid accounted for 34.37% and 17.28% of all species recorded, respectively (Table 1).

Key to identification of dung beetle genera

1. Scutellum visible dorsally (Figure 1A)..............................................2

   – Scutellum not visible dorsally (Figure 1B)....................................3

2. (1) Body flattened dorsally, elongated, with parallel sides. Labial palpi with two palpomeres; Mesocoxae parallel to the longitudinal axis of the body, positioned externally in relation to the metaventrite. Pronotum with basal margination (Figure 2A;C).................................................................................................Eurysternus Dalman, 1824

   – Body slightly convex dorsally, without parallel sides, usually oval. Labial palpi with more then two palpomeres; Mesocoxae perpendicular or oblique to the longitudinal axis of the body. Pronotum without basal margination (Figure 2B;D)......................................................................... Malagoniella Martínez, 1961

3. (1) Anterior leg with trochantofemoral fovea (Figure 3A)............4

   – Anterior leg without trochantofemoral fovea (Figure 3B).......................................................................................................11

4. (3) Last abdominal ventrite greatly expanded in the middle, covering the entire disc of the abdomen. The other ventrites visible only on the sides of the abdomen (Figure 4A)...........................................5

   – Last abdominal ventrite not covering the entire disc, other ventrites visible and clearly distinguishable in the middle of the abdomen (Figure 4B)................................................................................9

5. (4) Pseudepipleuron forming two lateral sinuosities, posterior partially (which is at the height of the metacoxa) covers the epipleuron, which is sharply narrowed towards the apex (Figure 5A)........................................................................................................6

   – Pseudepipleuron not forming two lateral sinuosities, epipleura gradually narrowed to apex (Figure 5B).....................................8

6. (5) Straight or slightly curved edge of the clypeus on each side between the clypeal tooth and the clypeogenal suture (Figure 6B)..................................................................TrichillumHarold, 1868HAROLD, E. 1868. Die arten der gattung Choeridium. Coleopterologische Hefte 4, 32–76.

   – Edge of clypeus with strong rounded lobe or angulation between clypeal tooth and clypeogenal suture (Figure 6A).........................7

7. (6) Clypeogenal margin with an incision over the suture, clypeus and gena appear separately rounded (Figure 7A).............................................................................................Eutrichilum Martínez, 1969

   – Clypeogenal margin straight to slightly sinuous (Figure 7B).............................................................. BesourengaVaz-de-Mello, 2008VAZ-DE-MELLO, F.Z. 2008. Synopsis of the new subtribe Scatimina (Coleoptera: Scarabaeidae: Scarabaeinae: Ateuchini), with descriptions of twelve new genera and review of Genieridium, new genus. Zootaxa 1955:1–75.

8. (5) Head with four clypeal teeth, rounded body (Figure 8A)............................................................. TrichillidiumVaz-de-Mello, 2008VAZ-DE-MELLO, F.Z. 2008. Synopsis of the new subtribe Scatimina (Coleoptera: Scarabaeidae: Scarabaeinae: Ateuchini), with descriptions of twelve new genera and review of Genieridium, new genus. Zootaxa 1955:1–75.

   – Head with two clypeal teeth (Figure 8B)................................................................................... GenieridiumVaz-de-Mello, 2008VAZ-DE-MELLO, F.Z. 2008. Synopsis of the new subtribe Scatimina (Coleoptera: Scarabaeidae: Scarabaeinae: Ateuchini), with descriptions of twelve new genera and review of Genieridium, new genus. Zootaxa 1955:1–75.

9. (4) Side of pronotum with deep longitudinal sulcus. Absent lateral pronotal fossae or callus. (Figure 9A)...............................................................................................................Uroxys Westwood, 1842

   – Side of pronotum without deep longitudinal sulcus, with lateral pronotal fossae or callus (Figure 9B).......................................10

10. (9) Pygidium with deep transverse sulcus on disc. Last meso- and metatarsomere without dentiform process above claw insertion (Figure 10A;B)..........................................Agamopus Bates, 1887

    – Pygidium with disc without sulcus. Last meso- and metatarsomere with dentiform process above claw insertion (Figure 10C;D)...............................................................Zonocopris Arrow, 1932

11. (3) Length of the first metatarsomere greater than the combined length of the next three metatarsomeres, metatarsal with five tarsomeres (Figure 11A).............................................................12

    – Length of the first metatarsomere less than the combined length of the next three metatarsomeres together; or metatarsi with less than five tarsomeres (Figure 11B)....................................13

12. (11) Propleuron with oblique carina that reaching lateral margin next to the anterior angle, forming an anterolateral tooth, rounded in males and acute in females; (African species introduced); size7-13mm (Figure 12A)...................Digitonthophagus Balthasar, 1959

    – Propleuron without anterolateral tooth near the anterior angle, insertion of the longitudinal (hypomeral) propleural carina exactly under the anterior angle, or not reaching the pronotal edge; size 4-12mm (Figure 12B)....................................................................................................Onthophagus Latreille, 1807

13. (11) Meso- and metatarsus lacking claws (Figure 13A)...............14

    – Meso- and metatarsus with claws (Figure 13B).......................20

14. (13) Meso- and metatarsus with two to four tarsomeres (Figure 14A)...............................................................Dendropaemon Perty, 1830

    – Meso- and metatarsus with five tarsomeres (Figure 14B)…................................................................................................15

15. (14) Basal antennomere of antennal club not concave apically; apical two antennomeres clearly separated from basal antennomere (Figure 15A). Metepisternum simple, without posterior extension ...................................................................Gromphas Brullé, 1838

    – Basal antennomere of antennal club large, strongly concave apically, receiving in the concavity the two apical lamellae (Figure 15B). Metepisternum with posterior extension covering lateral margin of elytra............................................................16

16. (15) Clypeal margin deeply, acutely emarginate medially, emarginations producing two acute teeth that are separated from adjacent clypeal border by external emargination (Figure 16A)..................................................Coprophanaeus d’Olsoufieff, 1924

    – Clypeal margin without deep and sharp emargination, with at most two conspicuous middle teeth (Figure 16B)..................17

17. (16) Head with transverse frontal carina (sometimes raised as pair of horns in male), in addition to frontoclypeal carina (Figure 16B).....................................................................Diabroctis Gistel, 1857

    – Head with single horn or carina, or bare.................................18

18. (17) Metasternum with long, dorsally curved, acute spiniform process extending between apices of procoxae (Figure 17A)...............................................................Oxysternon Laporte, 1840

    – Metarsternum simply angled anteromedially, without spiniform process (Figure 17B)..............................................................19

19. (18) Anterior portion of circumnotal ridge entire, not interrupted behind each eye (Figure 18A)............................................................................................................Sulcophanaeus Olsoufieff, 1924

    – Anterior portion of circumnotal ridge interrupted behind each eye (Figure 18B)...............................Phanaeus Macleay, 1819

20. (13) Pronotum with two distinct, posteromedian fossae. Body metallic red color and length greater than 15mm (Figure 19A).....................................................................BolbitesHarold, 1868HAROLD, E. 1868. Die arten der gattung Choeridium. Coleopterologische Hefte 4, 32–76.

    – Posteromedian pronotal fossae absent. Body browdish to black and length smaller than 15mm (Figure 19B).........................21

21. (20) Tarsal claws reduced, straight or only weakly curved (Figure 20A). Hypomeron convex or weakly concave, never strongly excavated. Head without transverse carina...............................................................................Anomiopus Westwood, 1842

    – Tarsal claws large, strongly curved, falciform or angulate (Figure 20B); if only weakly developed, then either hypomeron deeply excavated anteriorly or head with transverse carina or both.......................................................................................22

22. (21) Metatibia not enlarged toward apex, or only weakly and gradually (Figure 21A)................................................................23

    – Metatibia, abruptly widened toward apex (Figure 21B)...........27

23. (22) Elytal interstriae with short carina or tubercles at the apex (Figure 22A).................................Deltochilum Eschscholtz, 1822

    – Elytral interstriae without carina or tubercles at the apex, at most with a lateral carina that may be almost complete (Figure 22B)................................................................................................24

24. (23) Posterior edge of head not margined between eyes (Figure 23A). Mesosternum relatively long, not narrowed medially, completely horizontal............................................Pseudocanthon Bates, 1887

    – Posterior edge of head completely margined between eyes (Figure 23B). Mesosternum shorter in the middle than laterally, or positioned vertically and weakly visible............................25

25. (24) Mentum completely divided longitudinally (Figure 24A)............................................Holocanthon Martínez & Pereira, 1956

    – Mentum not completely divided, at most with deep U- or V-shaped emargination (Figure 24B)......................................26

26. (25) Pronotum with regular sculpturing, regularly convex, at most with posterior mean depression (scutellar impression) (Figure 25A)........................................................Canthon Hoffmannsegg, 1817

    – Pronotum with strong striated microsculpture, flat, and distributed depressions and elevations (Figure 25B)........................................................................Anisocanthon Martínez & Pereira, 1956

27. (22) Hypomeron deeply excavated anteriorly, excavation delimited posteriorly by a vertical area separated from the unexcavated part by a transversal carina (Figure 26A). Apical internal angle of the tibia ~ 90º or acute, anterior edge of continuous apical tooth (without forming an angle) with apical truncation.................................................................................................Ateuchus Weber, 1801

    – Hypomeron weakly excavated anteriorly, excavation not clearly delimited posteriorly; Transverse carina absent (Figure 26B). Apical internal angle of the protibial obliquely truncated (>90º); if ~90° or weakly acute, then the anterior edge of the apical tooth forms an angle with margin of apical truncation....................28

28. (27) Apical internal angle of the protibiae ~90° or acute (Figure 27A). Mesosternum usually very short, vertically positioned; metasternum usually convex (Figure 27B)....................Canthididium Erichson, 1847

    – Apical internal angle of the protibiae clearly obtuse, apical edge in continuation with apical tooth (Figure 27C). Mesosternum very well developed, horizontal; metasternum usually flat (Figure 27D)..............................................................................29

29. (28) Medial ventral carina of protibiae interrupted by punctures (Figure 28A). Body elongated, abdomen very short in relation to the body. Ventral clypeal process absent (Figure 28B)........................................................................................Ontherus Erichson, 1847

    – Medial ventral carina of protibiae complete (Figure 28C). Rounded body, abdomen not shortened. Ventral clypeal process, usually coniform and apically bifurcated, sometimes inserted into a longitudinal carina; rarely otherwise, but never with simple transverse carina (Figure 28D)................................................30

30. (29) Antennae with eight antennomeres (Figure 29A)........................................................................Isocopris Pereira & Martínez, 1960

    – Antennae with nine antennomeres (Figure 29B)...................................................................................Dichotomius Hope, 1838.

Identification key for the species of Eurysternus

1. Metaventrite disc with round fovea in the middle. Pronotum dark with a light spot anteriorly, just behind the head. Size 6-10mm (Figure 30A;B)..............................................................................E. plebejus

   – Metaventrite disk flat, wavy or with longitudinal sulcus, without round fovea in the middle. Pronotum totally dark or mostly light with dark spots. Size 9-20mm (Figure 30C;D)...........................2

2. (1) Light colored posterior femurs, at least in the apical half. Pronotum mostly lighter, with well-defined dark spots (Figure 31A;C).............................................................................................................3

   – Posterior femurs totally dark like the pronotum (Figure 31B;D) ...................................................................................................4

3. (2) Posterior femurs with teeth at the posterior edge, darker at the base. Size 10-20mm (Figure 31A).................................E. caribaeus

   – Posterior femurs without teeth at the anterior edge. Size 9-13mm (Figure 31B).................................................................E. jessopi

4. (2) Middle and posterior femurs claviform. Dorsal body without any metallic luster. Elytra apex bristles straight. Size 9-14mm (Figure 31B).........................E. parallelus (Figure 32A)......................................

   – Middle and posterior femurs not claviform. Body dorsally with metallic sheen at least in the elytral sutural striae and anterior part of the clypeus. Bristles at apex of elytra strongly curved in apical half (Figure 32B)........................................................................5

5. (4) Head surface with most no metallic reflections, except for a narrow metallic green band anteriorly on the clypeus. Lateral edge of male hind tibia curved in half in dorsal view. (Figure 33A;C).....................................................................................E. nigrovirens

   – Head surface mostly glossy and with strong metallic reflections between ocellates. Lateral edge of male hind tibia almost straight or straight in the middle in dorsal view (Figure 33B;D).......................................................................................E. aeneus

Identification key for the species of Canthon

1. Pygidium and propygidium separated by carina (Figure 34A)............2

   – Pygidium and propygidium not separated by carina (Figure 34B).....................................................................................................8

2. (1) Head with quadridented clypeus (Figure 35A).............................3

   – Head with bidented clypeus (Figure 35B)....................................4

3. (2) Pronotum, in dorsal view, with smooth lateral margin and serrated in ventral view. (Figure 36A).......................C. mutabilis Lucas, 1859

   – Pronotum with smooth lateral margin, in dorsal and ventral view. (Figure 36B)................................C. curvodilatatus Schmidt, 1920

4. (2) Gena anteriorly with visible tooth (Figure 37A black arrow).........................................................................................................5

   – Gena anteriorly, prominent, but without tooth.............................7

5. (4) Clypeus with two long teeth bulging, rounded, or subtriangular at apex, not acute (Figure 37A)…................C. substriatusHarold, 1868HAROLD, E. 1868. Die arten der gattung Choeridium. Coleopterologische Hefte 4, 32–76.

   – Clypeus with two long, triangular teeth, acute, not bulging or rounded at the apex (Figure 37B)…............................................6

6. (5) Pronotum completely punctated, being more subtle on the disc than on the sides; shiny elytra with well-marked striations; subtly punctuated interstriae. Metafemora with complete anterior and incomplete posterior margination (Figure 38A;C)….................................................................C. virens (Mannerheim, 1829)

   – Pronotum with almost imperceptible punctation on the disc; opaque elytra, striae not as marked as in C. virens; interstriae with microreticulated punctation; metafemora with complete anterior and posterior margination (Figure 38B;D)…..................................................C. chalybaeus Blanchard, 1845

7. (4) Short clypeal teeth; complete hypomeral carina; pronotum and elytra dark (Figure 39A;B).....................C. daguerrei Martinez, 1951

   – Clypeal teeth not as short as above; incomplete hypomeral carina; light pronotum with black stain and green reflections and dark elytra (Figure 39C;D)….................C. ornatus Redtenbacher, 1858

8. (1) Head with quadridented clypeus (Figure 40A)........................................................................C. quinquemaculatus Castelnau, 1840

   – Head with bidented clypeus, which can be very or poorly defined (Figure 40B)..............................................................................9

9. (8) Central clypeal teeth separated by U-shaped emargination or poorly defined teeth (Figure 40B black arrow)............................10

   – Central clypeal teeth separated by open V-shaped emargination. (Figure 40C)…..........................................................................12

10. (9) Central clypeal teeth separated by well-delimited U-shaped emargination; pronotum and elytra light with dark outline. (Figure 41A)......................................C. maldonadoi Martínez, 1951

    – Clypeal teeth are little delimited, pronotum and elytra dark. (Figure 41B)............................................................................11

11. (10) Clypeal teeth short and rounded; pygidium longer than wide and with thick punctation; size 5‒6,5mm (Figure 42A;B).........................................................................C. edentulousHarold, 1868HAROLD, E. 1868. Die arten der gattung Choeridium. Coleopterologische Hefte 4, 32–76.

    – Clypeal teeth only weakly indicated, not salient, separated by wide emargination; Pygidium wider than long and fine punctation compared to the previous one; size 10‒12mm. (Figure 42C;D)................................................................................. C. quadratus

12. (9) Pronotum and elytra dark; metafemur with anterior and posterior margination. (Figure 43A;C)................C. unicolor Blanchard, 1845

    – Pronotum and elytra may be yellow with black spots; metafemur only with anterior margination. (Figure 43B;D)...............................................C. histrio (Saint-Fargeau & Audinet-Serville, 1828)

Identification key for the species of Coprophanaeus

1. Elytral striae smooth; interstriae with subtle punctation (Figure 44A)...........................................................................................................2

   – Elytral striae sculpted; rugopunctated interstriae (Figure 44B)........3

2. (1) Anterior portion of circumnotal ridge interrupted behind eyes (Figure 45A)............................................C. milon (Blanchard, 1845)

   – Anterior portion of circumnotal ridge entire, not broken behind eyes (Figure 45B).....................C. cyanescens (d’Olsoufieff, 1924)

3. (1) Elytral striae 2‒4 only slightly wavy; width of striations at narrowings at least three-quarters of that at widest points; size 25‒35 mm (Figure 46A)..........................C. bonariensis (Gory, 1844)

   – Elytral striae strongly wavy; width of striations at narrowest points not more than half (often much less than half) that at widest points; size 30‒55mm (Figure 46B)...................................................................................................................C. ensifer (Germar, 1824)

Identification key for the species of Dichotomius

1. Clypeal margin rounded or weakly emarginated; if bidentate, small, non-margined teeth (Figure 47A)...................................................2

   – Clypeus distinctly bidentate, teeth usually margined (Figure 47B)......................................................................................................3

2. (1) Elytra with smooth striations and no punctation. Pronotum with median longitudinal sulcus (Figure 48A;B).......................................................................................................D. bos (Blanchard, 1845)

   – Elytra with punctiform striations. Pronotum without median sulcus (Figure 49A;B)....................D. luctuosioides (Luederwaldt, 1922)

3. (1) Metallic elytra (satin gloss), coloration green, reddish or almost burgundy brown (Figure 50A)........................................................4

   – Pronotum and elytra staining black, opaque or glossy (Figure 50B)......................................................................................................6

4. (3) Pronotum mesoanteriorly not flattened; base of the pronotum without evident punctuation in the central region (Figure 51A)…................................................................................5

   – Pronotum mesoanteriorly flattened; base of pronotum with two rows of ocellar punctuation evident (Figure 51B)….....................................................................D. lycas (Felsche, 1901)

5. (4) Dorsal surface monochromatic, glossy greenish satin; teeth margined, triangular and pointed; pronotum with flat anterior lateral edge; anterior tibia spur, spiniform (Figure 52A)............................................................................................D. glaucus (Harold, 1869)

   – Dorsal surface bicolor, black pronotum and glossy satin burgundy elytra; rounded and flat teeth; pronotum with anterior lateral edge forming an almost triangular tip; protibial spur quadrangular (Figure 52B).......................................D. cuprinus (Felsche, 1901)

6. (3) Sides of metaventrite and hypomeron with dense and long setae, red to yellowish-brown (Figure 53A).............................................7

   – Sides of metaventrite and hypomeron without setae as described above (Figure 53B)...................................................................................................................D. opacipennis (Luederwaldt, 1931)

7. (6) Clypeus with six teeth; pronotum with median anterior slope with ocellated punctuations (Figure 54A)..............................................................................................D. sexdentatus (Luederwaldt, 1925)

   – Clypeus with two teeth; posterior margin of the pronotum with ocellated punctuations (Figure 54B)........................................................................................................D. nisus (Olivier, 1789)

Identification key for the species of Ontherus

1. Frontoclypeal suture without carina and/or tubercles (Figure 55A).....................................................................O. digitatus Harold, 1869

   – Frontoclypeal suture with carina and/or tubercles (Figure 55B).......2

2. (1) Abdominal ventrites medially set, bristles similar to those on the lateral aspect of the metasternum (Figure 56A)..............................3

   – Abdominal ventrites glabrous medially, with at most one row of short bristles along the posterior border of the metacoxa (Figure 56B)..................................................................................4

3. (2) Posterior border of metacoxa smooth. Metaventrite with most of the punctation of the oval lateral edges, with little defined edges (Figure 57A;B).....................O. appendiculatus (Mannerheim, 1829)

   – Posterior edge of metacoxa slightly serrated. Metaventrite with most of the punctation of the lateral edges rounded, clearly delimited (Figure 57C;D).............................................................................................................................O. azteca Harold, 1869

4. (2) Median metaventral lobe length less than twice its width (Figure 58A)......................................................................................5

   – Median metaventral lobe length twice its maximum width (Figure 58B).................................................................................6

5. (4) Pygidium wider than long, punctures of moderate size and deeply impressed; space between glossy scores; smaller than 12mm (Figure 59A)…...................................O. aphodioides Burmeister, 1874

   – Pygidium as wide as long, punctures small and weakly impressed; greater than 12mm (Figure 59B).......................................................................................................O. sulcator (Fabricius, 1775)

6. (4) Posterior edge of the metatrochanter complete with sulcus; pygidium with subtle punctation (Figure 60A;B)....................................................................................O. dentatus Luederwaldt, 1930

   – Posterior edge of metatrochanter partially with sulcus; pygidium with well-marked punctation (Figure 60C;D)................................................................................O. erosioides Luederwaldt, 1930

Identification key for the species of Deltochilum

1. Bidentate clypeus (Figure 61A)......................................................2

   – Clypeus quadridentate, medial teeth stronger and longer than others (Figure 61B).....................D. pseudoicarusBalthasar, 1939BALTHASAR, V. 1939. Megathoposoma n.Gen. und neue Arten der Gattung Deltochilum Eschz. 59. Beitrag zur Kenntnis der Scarabaeiden (Col.). Cas. Ces. Spol. Entomol 36:5–19.

2. (1) Head longer than wide (Figure 62A)........................................3

   – Head much wider than long (Figure 62B)...........................................................................................................D. silphoides (Balthasar, 1939BALTHASAR, V. 1939. Megathoposoma n.Gen. und neue Arten der Gattung Deltochilum Eschz. 59. Beitrag zur Kenntnis der Scarabaeiden (Col.). Cas. Ces. Spol. Entomol 36:5–19.) (and possibly other species of the irroratum group)

3. (2) Pronotum and elytra of the same color, dark with greenish reflections (Figure 63A).........................D. icaroidesBalthasar, 1939BALTHASAR, V. 1939. Megathoposoma n.Gen. und neue Arten der Gattung Deltochilum Eschz. 59. Beitrag zur Kenntnis der Scarabaeiden (Col.). Cas. Ces. Spol. Entomol 36:5–19.

   – Pronotum yellow with dark spot in the center, elytra of uniform dark coloration (Figure 63B)..........................................................................................................D. cupreicoile (Castelnau, 1840)

Discussion

In this study, we provide an updated list of species of dung beetles of the Brazilian Pantanal. No such list had hitherto been published. The closer parallel had been Vaz-de-Mello et al.’s (2017) list of the species of Mato Grosso do Sul state, which indicated those which were present in its Pantanal area. Our study mitigates the lack of information on dung beetles species of the Brazilian Pantanal as a whole, increasing the number of species recorded from 43 (see Vaz-de-Mello et al. 2017) to 68 species (Figure 64A-P and Figure 65A-J). Below is an up-to-date commentary on the taxonomic status of the genera that occur in the Pantanal.

Ateuchus Weber, 1801

Genus with paracoprid representatives and 101 species currently recognized as valid distributed in the Americas (Schoolmeesters 2023SCHOOLMEESTERS, P. 2023. World Scarabaeidae Database. In: Bánki O., Roskov Y., Döring M., Ower G., Vandepitte L., Hobern D., Remsen D., Schalk P., DeWalt R.E., Keping M., Miller J., Orrell T., Aalbu R., Abbott J., Adlard R., Adriaenssens E.M., Aedo C., Aescht E., Akkari N., et al. (eds) Catalogue of Life Checklist (Version 2023–03-06). Available from https://www.catalogueoflife.org/data/dataset/1027 [last accessed 03 Apr 2023].
https://www.catalogueoflife.org/data/dat... ). Except for some regional works, the genus was last revised by Harold (1868)HAROLD, E. 1868. Die arten der gattung Choeridium. Coleopterologische Hefte 4, 32–76. and identifying its South American species is currently unfeasible relying on published information. During the data survey for this work, these species were reported: Ateuchus carbonarius (Harold, 1868HAROLD, E. 1868. Die arten der gattung Choeridium. Coleopterologische Hefte 4, 32–76.) (Pessôa et al. 2017PESSÔA, M.B., IZZO, T.J. & VAZ-DE-MELLO, F.Z. 2017. Assemblage and functional categorization of dung beetles (Coleoptera: Scarabaeinae) from Pantanal. PeerJ 5:e3978.), Ateuchus latus (Boucomont, 1928) (Vaz-de-Mello et al. 2007VAZ-DE-MELLO, F.Z. 2007. Revision and phylogeny of the dung beetle genus Zonocopris Arrow 1932 (Coleoptera: Scarabaeidae: Scarabaeinae), a phoretic of land snails. Ann. Soc. Entomol. Fr. 43:231–239.), and Ateuchus pruneus (Boucomont, 1928) (Correira et al. 2022b). However, this genus is under revision and only one species could be confirmed as present in the Brazilian Pantanal: Ateuchus aff. viduus (Blanchard, 1846), a species with wide distribution in the Pantanal and southern Cerrado regions (Mario Cupello, personal communication).

Canthidium Erichson, 1847

Genus with paracoprids representatives and reported exclusively for the Neotropical region. It is one of the most speciose scarabaeine genera in the continent, with about 178 described species (Schoolmeesters 2023SCHOOLMEESTERS, P. 2023. World Scarabaeidae Database. In: Bánki O., Roskov Y., Döring M., Ower G., Vandepitte L., Hobern D., Remsen D., Schalk P., DeWalt R.E., Keping M., Miller J., Orrell T., Aalbu R., Abbott J., Adlard R., Adriaenssens E.M., Aedo C., Aescht E., Akkari N., et al. (eds) Catalogue of Life Checklist (Version 2023–03-06). Available from https://www.catalogueoflife.org/data/dataset/1027 [last accessed 03 Apr 2023].
https://www.catalogueoflife.org/data/dat... ), and at least another 150 awaiting to be described (Vaz-de-Mello personal observation). This genus is divided into two subgenera: Canthidium s. str. and Neocanthidium Martínez et al., 1964, equally diverse (80 and 70 species, respectively), with 31 species deemed incertae sedis (not assigned to any subgenus). All of these taxa need urgent taxonomic revision. In this study, we recorded six species: Canthidium angulicoleBalthasar, 1939BALTHASAR, V. 1939. Megathoposoma n.Gen. und neue Arten der Gattung Deltochilum Eschz. 59. Beitrag zur Kenntnis der Scarabaeiden (Col.). Cas. Ces. Spol. Entomol 36:5–19., C. barbacenicum Preudhomme de Borre, 1886, C. breve (Germar, 1823), C. cuprinum Harold, 1867, C. kelleri (Martinez et al., 1964), C. viride (Lucas, 1859). Currently, the genus is under review, with new species being described singly or being grouped into species groups (Cupello 2018CUPELLO, M. 2018. On the types species of the New World dung beetle genus Canthidium Erichson, 1847 (Coleoptera: Scarabaeidae: Scarabaeinae), with an annotated checklist of species. Zootaxa 4388:451–486.; Kohlmann et al. 2018KOHLMANN, B., ARRIAGA-JIMÉNEZ, A. & RÖS, M. 2018. An unusual new species of Canthidium (Coleoptera: Scarabaeidae: Scarabaeinae) from Oaxaca, Mexico. Zootaxa 4378:273–278.; Moctezuma et al. 2019MOCTEZUMA, V., SÁNCHEZ-HUERTA, J.L. & HALFFTER, G. 2019. New species of Canthidium (Coleoptera: Scarabaeidae: Scarabaeinae) from Mexico. Can. Entomol. 151:432–441.; Carvalho de Santana et al. 2019CARVALHO DE SANTANA, E.C., PACHECO, T.L. & VAZ-DE-MELLO, F.Z. 2019. Taxonomic revision of the Canthidium Erichson, 1847 species of the gigas group (Coleoptera, Scarabaeidae, Scarabaeinae). Eur. J. Taxon. 530:1–24.).

Canthon Hoffmannsegg, 1817

It is one of the most diverse genera among the Scarabaeinae, exclusive to the Americas, with currently 163 recognized species (Schoolmeesters 2023SCHOOLMEESTERS, P. 2023. World Scarabaeidae Database. In: Bánki O., Roskov Y., Döring M., Ower G., Vandepitte L., Hobern D., Remsen D., Schalk P., DeWalt R.E., Keping M., Miller J., Orrell T., Aalbu R., Abbott J., Adlard R., Adriaenssens E.M., Aedo C., Aescht E., Akkari N., et al. (eds) Catalogue of Life Checklist (Version 2023–03-06). Available from https://www.catalogueoflife.org/data/dataset/1027 [last accessed 03 Apr 2023].
https://www.catalogueoflife.org/data/dat... ). Its distribution ranges from the United States to Uruguay and central-west Argentina, with representatives in all biomes along this range. This genus is divided into nine subgenera, some of which are under revision (Cupello & Vaz-de-Mello 2018CUPELLO, M. & VAZ-DE-MELLO, F.Z. 2018. A monographic revision of the Neotropical dung beetle genus Sylvicanthon Halffter & Martínez, 1977 (Coleoptera: Scarabaeidae: Scarabaeinae: Deltochilini), with a reappraisal of the taxonomic history of “Canthon sensu lato”. European Journal of Taxonomy 467: 1‒205. and references cited therein). Overall, their representatives are telecoprids, and may be generalists and predators. In this study, we recorded 13 species: C. chalybaeus Blanchard, 1846, C. curvodilatatus Schmidt, 1920, C. daguerrei Martínez, 1951, C. edentulusHarold, 1868HAROLD, E. 1868. Die arten der gattung Choeridium. Coleopterologische Hefte 4, 32–76., C. histrio (LePeletier de Saint-Fargeau & Audinet-Serville, 1828), C. maldonadoi Martínez, 1951, C. mutabilis Lucas, 1859, C. ornatus Redtenbacher, 1868, C. quadratus Blanchard, 1846, C. quinquemaculatus Castelnau, 1840, C. substriatusHarold, 1868HAROLD, E. 1868. Die arten der gattung Choeridium. Coleopterologische Hefte 4, 32–76., C. unicolor Blanchard, 1846 and C. virens (Mannerheim, 1829). Nunes & Vaz-de-Mello (2022), in the revision of the genus Tetraechma Blanchard, 1842, two species of Canthon that were reported for the Brazilian Pantanal were transferred to the genus Tetraechma. They are: Canthon apicalis Lucas, 1859 and Canthon lituratus solutus Schmidt, 1920.

Coprophanaeus d’Olsoufieff 1924

Genus exclusive to the Americas, with 50 described species (Schoolmeesters 2023SCHOOLMEESTERS, P. 2023. World Scarabaeidae Database. In: Bánki O., Roskov Y., Döring M., Ower G., Vandepitte L., Hobern D., Remsen D., Schalk P., DeWalt R.E., Keping M., Miller J., Orrell T., Aalbu R., Abbott J., Adlard R., Adriaenssens E.M., Aedo C., Aescht E., Akkari N., et al. (eds) Catalogue of Life Checklist (Version 2023–03-06). Available from https://www.catalogueoflife.org/data/dataset/1027 [last accessed 03 Apr 2023].
https://www.catalogueoflife.org/data/dat... ). This genus is divided into three subgenera: Coprophanaeus s. str. (38 species), Metallophanaeus d’Olsoufieff, 1924 (eight species) and Megaphanaeus d’Olsoufieff, 1924 (four species). The species are easily identifiable with the recent review by Edmonds & Zídek (2010)EDMONDS, W.D. & ZÍDEK, J. 2010. A taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 129:1–111., and further descriptions and revalidations (Kohlmann & Solís 2012KOHLMANN, B. & SOLÍS, Á. 2012. New species and revalidations of scarab beetles (Coleoptera: Geotrupidae: Athyreini and Coleoptera: Scarabaeidae: Scarabaeinae) from Costa Rica and Panama. Zootaxa 3193:28–52., Cupello & Vaz-de-Mello 2013CUPELLO, M. & VAZ-DE-MELLO, F.Z. 2013a. New evidence for the validity of Coprophanaeus (C.) terrali Arnaud, 2002 (Coleoptera: Scarabaeidae: Scarabaeinae: Phanaeini), a dung beetle from Brazil. Zootaxa 3717:359–368., 2014CUPELLO, M. & VAZ-DE-MELLO, F.Z. 2014. Revalidation of the Brazilian Atlantic Forest dung beetle species Coprophanaeus (Metallophanaeus) machadoi (Pereira & d-Andretta, 1955) (Coleoptera: Scarabaeidae: Scarabaeinae: Phanaeini) based on morphological and distributional evidence. Zootaxa 3869:435–451.). Here, we recorded four species: Coprophanaeus (Megaphanaeus) ensifer (Germar, 1823), C. (Megaphanaeus) bonarienis (Gory, 1844), C. (C.) cyanescens (d’Olsoufieff, 1924), C. (C.) milon (Blanchard, 1845). Usually, the representatives are paracoprids and in dissonance of their name, they are usually necrophagous.

Deltochilum Eschscholtz, 1822

Genus endemic to the Americas, with approximately 114 described species (Schoolmeesters 2023SCHOOLMEESTERS, P. 2023. World Scarabaeidae Database. In: Bánki O., Roskov Y., Döring M., Ower G., Vandepitte L., Hobern D., Remsen D., Schalk P., DeWalt R.E., Keping M., Miller J., Orrell T., Aalbu R., Abbott J., Adlard R., Adriaenssens E.M., Aedo C., Aescht E., Akkari N., et al. (eds) Catalogue of Life Checklist (Version 2023–03-06). Available from https://www.catalogueoflife.org/data/dataset/1027 [last accessed 03 Apr 2023].
https://www.catalogueoflife.org/data/dat... ), divided into eight subgenera, some of which are under revision (namely, Deltohyboma Paulian, 1938, with 42 species, and Calhyboma Kolbe, 1893, with 13), some with recent reviews completed (Génier 2012GÉNIER, F. 2012. A new species and notes on the subgenus Deltochilum (Deltochilum) Eschscholtz, 1822 (Coleoptera: Scarabaeidae: Scarabaeinae: Deltochilini). Zootaxa 3357:25–36.; González-Alvarado & Vaz-de-Mello 2014GONZÁLEZ-ALVARADO, A. & VAZ-DE-MELLO. 2021. Towards a comprehensive taxonomic revision of the Neotropical dung beetle subgenus Deltochilum (Deltohyboma) Lane, 1946 (Coleoptera: Scarabaeidae: Scarabaeinae): Division into species-groups. PLoS ONE 16:e0244657.; Silva et al. 2015SILVA, F.A.B., LOUZADA, J. & VAZ-DE-MELLO, F. 2015. A revision of the Deltochilum subgenus Aganhyboma Kolbe, 1893 (Coleoptera: Scarabaeidae: Scarabaeinae). Zootaxa 3925: 451–504.; Nazaré-Silva & Silva 2021NAZARÉ-SILVA, E.E. & SILVA, F.A.B. 2012. A revision of the subgenera Euhyboma Kolbe, 1893, Parahyboma Paulian, 1938, and Rubrohyboma Paulian, 1939 of Deltochilum Eschscholtz, 1822 (Coleoptera:Scarabaeidae: Scarabaeinae). Revista Brasileira de Entomologia 65(3):1‒19.). The species are telecoprid. Deltochilum is here represented by four species belonging to three different subgenera: Deltochilum (Deltochilum) silphoidesBalthasar, 1939BALTHASAR, V. 1939. Megathoposoma n.Gen. und neue Arten der Gattung Deltochilum Eschz. 59. Beitrag zur Kenntnis der Scarabaeiden (Col.). Cas. Ces. Spol. Entomol 36:5–19., D. (Hybomidium) pseudoicarusBalthasar, 1939BALTHASAR, V. 1939. Megathoposoma n.Gen. und neue Arten der Gattung Deltochilum Eschz. 59. Beitrag zur Kenntnis der Scarabaeiden (Col.). Cas. Ces. Spol. Entomol 36:5–19., Deltochilum (Aganhyboma) cupreicolle (Blanchard, 1846) and Deltochilum (A.) icarioidesBalthasar, 1939BALTHASAR, V. 1939. Megathoposoma n.Gen. und neue Arten der Gattung Deltochilum Eschz. 59. Beitrag zur Kenntnis der Scarabaeiden (Col.). Cas. Ces. Spol. Entomol 36:5–19..

Dichotomius Hope, 1838

Genus with exclusive distribution in the Americas, with 200 described species, divided into four subgenera; Dichotomius s. str. (74 species), Selenocopris Burmeister, 1846 (85 spp.), Cephagonus Luederwaldt, 1929 (40 spp.) and Homocanthonides Luederwaldt, 1929 (1 sp.) (Schoolmeesters 2023SCHOOLMEESTERS, P. 2023. World Scarabaeidae Database. In: Bánki O., Roskov Y., Döring M., Ower G., Vandepitte L., Hobern D., Remsen D., Schalk P., DeWalt R.E., Keping M., Miller J., Orrell T., Aalbu R., Abbott J., Adlard R., Adriaenssens E.M., Aedo C., Aescht E., Akkari N., et al. (eds) Catalogue of Life Checklist (Version 2023–03-06). Available from https://www.catalogueoflife.org/data/dataset/1027 [last accessed 03 Apr 2023].
https://www.catalogueoflife.org/data/dat... ), all under revision or recently revised ( Arias-Buriticá et al. 2013, Maldaner et al. 2015, Nunes et al. 2016a-b, Vaz-de-Mello et al. 2016, Arias-Buriticá et al. 2019, Montoya-Molina et al. 2019, Cassenote et al. 2020, Boilly et al. 2021, Montoya-Molina et al. 2021, Valois et al. 2022, Maldaner et al. 2022, Valois et al. 2023, Arias-Buriticá et al. 2023). We recorded eight species from two subgenera: Dichotomius (Dichotomius) bos (Blanchard, 1846), D. (D.) luctuosioides (Harold, 1869), Dichotomius (Selenocopris) cuprinus (Felsche, 1901), D. (S.) glaucus (Harold, 1869), D. (S.) lycas (Felsche, 1901), D. (S.) nisus (Olivier, 1789), D. (S.) opacipennis (Luederwaldt, 1931) and D. (S.) sexdentatus (Luederwaldt, 1925).

Digitonthophagus Balthasar, 1959

An Afro-Asian genus, with 16 species (Genier & Moretto 2017GÉNIER, F.A. & MORETTO, P. 2017. Digitonthophagus Balthasar, 1959: taxonomy, systematics, and morphological phylogeny of the genus revealing an African species complex (Coleoptera: Scarabaeidae: Scarabaeinae). Zootaxa 4248:1–110.). A species of African origin, Digitonthophagus gazella (Fabricius, 1787), was introduced in Brazil in the 1980s and is currently present in almost the entire national territory (Tissiani et al. 2017TISSIANI, A.S.O., VAZ-DE-MELLO, F.Z. & CAMPELO-JÚNIOR, J.H. 2017. Besouros rola-bostas das pastagens brasileiras e chave para identificação dos gêneros (Coleoptera: Scarabaeidae). Pesq. Agropec. Bras. 52:401–418.; Génier & Moretto 2017GÉNIER, F.A. & MORETTO, P. 2017. Digitonthophagus Balthasar, 1959: taxonomy, systematics, and morphological phylogeny of the genus revealing an African species complex (Coleoptera: Scarabaeidae: Scarabaeinae). Zootaxa 4248:1–110.). This species is mainly associated with exotic pastures (e.g., African grasses; Urochloa spp.). It benefits from the presence of cattle ranching (Correa et al. 2020CORREA, C.M.A., AUDINO, L.D., HOLDBROOK, R., BRAGA, R.F., MENÉNDEZ, R. & LOUZADA, J. 2020. Successional trajectory of dung beetle communities in a tropical grassy ecosystem after livestock grazing removal. Biodivers. Conserv. 29:2311–2328.), and its occurrence is rare in areas of native vegetation. This species has a high rate of dispersal and fertility and is rapidly established in environments with high solar incidence, including savanna environments and pastures in the Amazon region (Matavelli & Louzada 2008MATAVELLI, R.A. & LOUZADA, J.N.C. 2008. Invasão de áreas de savana intra-amazônicas por Digitonthophagus gazella (Fabricius, 1787) (Insecta: Coleoptera: Scarabaeidae). Acta Amaz. 38:153–158.).

Eurysternus Dalman, 1824

Genus with a Neotropical distribution, with 53 described species (Schoolmeesters 2023SCHOOLMEESTERS, P. 2023. World Scarabaeidae Database. In: Bánki O., Roskov Y., Döring M., Ower G., Vandepitte L., Hobern D., Remsen D., Schalk P., DeWalt R.E., Keping M., Miller J., Orrell T., Aalbu R., Abbott J., Adlard R., Adriaenssens E.M., Aedo C., Aescht E., Akkari N., et al. (eds) Catalogue of Life Checklist (Version 2023–03-06). Available from https://www.catalogueoflife.org/data/dataset/1027 [last accessed 03 Apr 2023].
https://www.catalogueoflife.org/data/dat... ), recently revised by Génier (2009)GÉNIER, F. 2009. Le genre Eurysternus Dalman, 1824 (Scarabaeidae: Scarabaeinae: Oniticellini), révision taxonomique et clés de détermination illustrées. Sofia: Pensoft, (Pensoft series faunistica, 85), p.1–430.. Its representatives have endocoprid behavior. We recorded five species: Eurysternus caribaeus (Herbst, 1789) with distribution from Mexico to and almost all of South America, E. aeneusGénier, 2009GÉNIER, F. 2009. Le genre Eurysternus Dalman, 1824 (Scarabaeidae: Scarabaeinae: Oniticellini), révision taxonomique et clés de détermination illustrées. Sofia: Pensoft, (Pensoft series faunistica, 85), p.1–430., E. jessopi Martínez, 1988, E. nigrovirensGénier, 2009GÉNIER, F. 2009. Le genre Eurysternus Dalman, 1824 (Scarabaeidae: Scarabaeinae: Oniticellini), révision taxonomique et clés de détermination illustrées. Sofia: Pensoft, (Pensoft series faunistica, 85), p.1–430., and E. plebejus Harold, 1880.

Genieridium Vaz-de-Mello, 2008VAZ-DE-MELLO, F.Z. 2008. Synopsis of the new subtribe Scatimina (Coleoptera: Scarabaeidae: Scarabaeinae: Ateuchini), with descriptions of twelve new genera and review of Genieridium, new genus. Zootaxa 1955:1–75.

This genus has seven described species, frequently found in pastures in the Cerrado and Chaco areas. Other species are also associated with natural grasslands and non-forest phytophysiognomies (Vaz-de-Mello 2008VAZ-DE-MELLO, F.Z. 2008. Synopsis of the new subtribe Scatimina (Coleoptera: Scarabaeidae: Scarabaeinae: Ateuchini), with descriptions of twelve new genera and review of Genieridium, new genus. Zootaxa 1955:1–75.). Here represented by two species: Genieridium bidens (Balthasar, 1938) and G. cryptops (Arrow, 1913).

Gromphas Brullé, 1838

This genus has six species and the last revision of this genus was made by Cupello & Vaz-de-Mello (2013bCUPELLO, M. & VAZ-DE-MELLO, F.Z. 2013b. Taxonomic revision of the South American dung beetle genus Gromphas Brullé, 1837 (Coleoptera: Scarabaeidae: Scarabaeinae: Phanaeini: Gromphadina). Zootaxa 3722:439–482., 2015CUPELLO, M. & VAZ-DE-MELLO, F.Z. 2015. A new species and the phylogeny of the South American genus Gromphas Brullé, 1837 (Coleoptera: Scarabaeidae: Scarabaeinae: Phanaeini). Journal of Natural History 50(15-16): 943-969.). Here, it is represented by only one species, Gromphas inermis Harold, 1869. This species occurs in Brazil, Bolivia, Paraguay, Argentina and Uruguay.

Isocopris Pereira & Martínez, 1960

This genus has seven species and the last revision of this genus was made by Rossini & Vaz-de-Mello (2017)ROSSINI, M. & VAZ-DE-MELLO, F.Z. 2017. A taxonomic review of the genus Isocopris Pereira and Martínez, 1960 (Coleoptera: Scarabaeidae: Scarabaeinae), with description of a new Brazilian species. J. Nat. Hist. 51:1091–1117.. In this study, we recorded only Isocopris foveolatus (Luederwaldt, 1931).

Malagoniella Martínez 1961

This genus needs taxonomic revision, since the last was performed by Hallfter & Martínez (1966)HALFFTER, G. & MATTHEWS, E.G. 1966. The natural history of dung beetles of the subfamily Scarabaeinae (Coleoptera: Scarabaeidae). Folia Entomol. Mex. 12:1–312.. It is currently divided into two subgenera Malagoniella s. str. and Megatophomima Martínez, 1961, with 16 described species (Schoolmaster 2023).

Ontherus Erichson, 1847

Genus with Neotropical distribution and 60 species, divided into three subgenera, Ontherus s. str. (34 species), CaelontherusGénier, 1996GÉNIER, F. 1996. A revision of the Neotropical genus Ontherus Erichson (Coleoptera, Scarabaeidae, Scarabaeinae). Mem. Ent. Soc. Can. 128:3–170. (24 species) and PlanontherusGénier, 1996GÉNIER, F. 1996. A revision of the Neotropical genus Ontherus Erichson (Coleoptera, Scarabaeidae, Scarabaeinae). Mem. Ent. Soc. Can. 128:3–170. (two species) (Schoolmeesters 2023SCHOOLMEESTERS, P. 2023. World Scarabaeidae Database. In: Bánki O., Roskov Y., Döring M., Ower G., Vandepitte L., Hobern D., Remsen D., Schalk P., DeWalt R.E., Keping M., Miller J., Orrell T., Aalbu R., Abbott J., Adlard R., Adriaenssens E.M., Aedo C., Aescht E., Akkari N., et al. (eds) Catalogue of Life Checklist (Version 2023–03-06). Available from https://www.catalogueoflife.org/data/dataset/1027 [last accessed 03 Apr 2023].
https://www.catalogueoflife.org/data/dat... ), revised by Génier (1996GÉNIER, F. 1996. A revision of the Neotropical genus Ontherus Erichson (Coleoptera, Scarabaeidae, Scarabaeinae). Mem. Ent. Soc. Can. 128:3–170., 1998GÉNIER, F. 1998. A revision of the Neotropical genus Ontherus Erichson (Coleoptera: Scarabaeidae: Scarabaeinae), supplement 1. The Coleopterists Bulletin 52(3):270–274.; see also González & Medina 2015GONZÁLEZAZ-ALVARADO, F.A. & MEDINA, C.A. 2015. The genus Ontherus Erichson 1847 (Coleoptera: Scarabaeidae: Scarabaeinae): description of a new species, and notes on the genus in Colombia. Zootaxa 3949(1):82–90.). Its representatives present paracoprid habits. We recorded six species: Ontherus appendiculatus (Mannerheim, 1829), O. aphodioides Burmeister, 1874, O. dentatus Luederwaldt, 1930, O. digitatusHarold, 1868HAROLD, E. 1868. Die arten der gattung Choeridium. Coleopterologische Hefte 4, 32–76., O. erosioides Luederwaldt, 1930, and O. sulcator (Fabricius, 1775).

Phanaeus MacLeay, 1819

A genus with 82 species (Edmonds & Zídek 2012EDMONDS, W.D. & ZÍDEK, J. 2012. Taxonomy of Phanaeus revisited: revised keys to and comments on species of the New World dung beetle genus Phanaeus MacLeay, 1819 (Coleoptera: Scarabaeidae: Scarabaeinae: Phanaeini). Insecta Mundi, v.274, p.1–108.; Schoolmeesters 2023SCHOOLMEESTERS, P. 2023. World Scarabaeidae Database. In: Bánki O., Roskov Y., Döring M., Ower G., Vandepitte L., Hobern D., Remsen D., Schalk P., DeWalt R.E., Keping M., Miller J., Orrell T., Aalbu R., Abbott J., Adlard R., Adriaenssens E.M., Aedo C., Aescht E., Akkari N., et al. (eds) Catalogue of Life Checklist (Version 2023–03-06). Available from https://www.catalogueoflife.org/data/dataset/1027 [last accessed 03 Apr 2023].
https://www.catalogueoflife.org/data/dat... ), two of which are inhabitants of grassland and savanna formations in Brazil and are occasionally collected in cultivated pastures (Edmonds 1994EDMONDS, W.D. 1994. Revision of Phanaeus Macleay, a new world genus of Scarabaeinae dung beetles (Coleoptera: Scarabaeidae, Scarabaeinae). Contrib. Sci. 443:1–105.). Here represented by two species: Phanaeus kirbyi Vigors, 1825 and Phanaeus palaeno Blanchard, 1846.

Pseudocanthon Bates, 1887

Genus with ten described species present on continental America and the Antilles. Among these species, five occurring in South America, P. perplexus (LeConte, 1847), P. xanthurus (Blanchard, 1847), P. vazdemelloi Nazaré-Silva & Silva, 2021, P. pantanensis Nazaré-Silva & Silva, 2021 and P. chaquensis Nazaré-Silva & Silva, 2021.

Sulcophanaeus d’Olsoufieff, 1924

Genus exclusive to the Americas, with 15 valid species (Schoolmeesters 2023SCHOOLMEESTERS, P. 2023. World Scarabaeidae Database. In: Bánki O., Roskov Y., Döring M., Ower G., Vandepitte L., Hobern D., Remsen D., Schalk P., DeWalt R.E., Keping M., Miller J., Orrell T., Aalbu R., Abbott J., Adlard R., Adriaenssens E.M., Aedo C., Aescht E., Akkari N., et al. (eds) Catalogue of Life Checklist (Version 2023–03-06). Available from https://www.catalogueoflife.org/data/dataset/1027 [last accessed 03 Apr 2023].
https://www.catalogueoflife.org/data/dat... ), its representatives are paracoprids. The last revision is by Edmonds (2000)EDMONDS, W.D. 2000. Revision of the Neotropical dung beetle genus Sulcophanaeus (Coleoptera: Scarabaeidae: Scarabaeinae). Folia Heyrovskyana 6:1–60., and it was represented here by only one species, Sulcophanaeus menelas (Castelnau, 1840).

Trichillidium Vaz-de-Mello, 2008VAZ-DE-MELLO, F.Z. 2008. Synopsis of the new subtribe Scatimina (Coleoptera: Scarabaeidae: Scarabaeinae: Ateuchini), with descriptions of twelve new genera and review of Genieridium, new genus. Zootaxa 1955:1–75.

Genus with four species, usually found in the Amazon, Chaco and Central America and with coprophagous diet (Vaz-de-Mello 2008VAZ-DE-MELLO, F.Z. 2008. Synopsis of the new subtribe Scatimina (Coleoptera: Scarabaeidae: Scarabaeinae: Ateuchini), with descriptions of twelve new genera and review of Genieridium, new genus. Zootaxa 1955:1–75.). Here represented by Trichillidium quadridens (Arrow, 1932).

Trichillum Harold, 1868HAROLD, E. 1868. Die arten der gattung Choeridium. Coleopterologische Hefte 4, 32–76.

Genus with Neotropical distribution, with 11 described species (Schoolmeesters 2023SCHOOLMEESTERS, P. 2023. World Scarabaeidae Database. In: Bánki O., Roskov Y., Döring M., Ower G., Vandepitte L., Hobern D., Remsen D., Schalk P., DeWalt R.E., Keping M., Miller J., Orrell T., Aalbu R., Abbott J., Adlard R., Adriaenssens E.M., Aedo C., Aescht E., Akkari N., et al. (eds) Catalogue of Life Checklist (Version 2023–03-06). Available from https://www.catalogueoflife.org/data/dataset/1027 [last accessed 03 Apr 2023].
https://www.catalogueoflife.org/data/dat... ). Its representatives are probably endocoprids. Here represented by only one species, Trichillum externepunctatum Preudhomme de Borre, 1886.

Zonocopris Arrow, 1932

Genus with two described species associated with large gastropods of the genera Strophocheilus (Müller, 1774) and Megalobulimus (Müller, 1774). This genus is found in Brazil, northern and northeastern Argentina, southern Bolivia and Paraguay (Vaz-de-Mello 2007VAZ-DE-MELLO, F.Z. 2008. Synopsis of the new subtribe Scatimina (Coleoptera: Scarabaeidae: Scarabaeinae: Ateuchini), with descriptions of twelve new genera and review of Genieridium, new genus. Zootaxa 1955:1–75.). In the Brazilian Pantanal, it is represented by Zonocopris gibbicollis (Harold, 1868HAROLD, E. 1868. Die arten der gattung Choeridium. Coleopterologische Hefte 4, 32–76.).

Uroxys Westwood, 1842

Genus with 59 recognized species (Schoolmasters 2023), which are present in forested areas. The last revision was carried out by Arrow (1933). It has size and ecological functions similar to those of the genus Trichillum. Here represented by one species: Uroxys corporaali Balthasar, 1940.

Although not recorded in our study, species of the genera Agamopus Bates, 1887, Anomiopus Westwood, 1846, BesourengaVaz-de-Mello, 2008VAZ-DE-MELLO, F.Z. 2008. Synopsis of the new subtribe Scatimina (Coleoptera: Scarabaeidae: Scarabaeinae: Ateuchini), with descriptions of twelve new genera and review of Genieridium, new genus. Zootaxa 1955:1–75., Dendropaemon Perty, 1830, Diabroctis Gistel, 1857 and Eutrichillum Martínez, 1969 are also expected to occur in the Brazilian Pantanal, due to the record of species of these genera in regions close to the Pantanal (Vaz-de-Mello, personal observation). Indeed, species of the genus Anomiopus (Correa et al. 2022bCORREA, C.M.A., FERREIRA, K.R., ABOT, A.R., LOUZADA, J. & VAZ-DE-MELLO, F.Z. 2022b. Ivermectin impacts on dung beetle diversity and their ecological functions in two distinct Brazilian ecosystems. Ecol. Entomol. 47:736–748.) and Dendropaemon were recently reported from the Pantanal region in Aquidauana and Miranda – Mato Grosso do Sul, Brazil (see Correa et al. 2022aCORREA, C.M.A., AUDINO, L.D., VAZ-DE-MELLO, F.Z. & LOUZADA, J. 2022a. Diversity and structure of dung beetle (Coleoptera: Scarabaeidae) assemblage in natural grasslands of the Brazilian Pantanal. Int. J. Trop. Insect Sci. 42:3253–3261.; Gonçalves et al. 2022GONÇALVES, T.F., CORREA, C.M.A., AUDINO, L.D., VAZ-DE-MELLO, F.Z., FONTOURA, F.M. & GUEDES, N.M.R. 2022. Quantifying the post-fire recovery of taxonomic and functional diversity of dung beetles in the Brazilian Pantanal. Ecol. Entomol. 47:601–612.).

Conclusions

This study provides an updated list and identification key for the identification of species of dung beetles occurring in the Brazilian Pantanal, except for species belonging to genera that are under taxonomic revision. We recorded high richness of dung beetles in the Brazilian Pantanal, with 68 dung beetle species of 30 genera. This underscores the importance of studies on this ecosystem in which the dung beetle fauna is still poorly known (Correa et al. 2022aCORREA, C.M.A., AUDINO, L.D., VAZ-DE-MELLO, F.Z. & LOUZADA, J. 2022a. Diversity and structure of dung beetle (Coleoptera: Scarabaeidae) assemblage in natural grasslands of the Brazilian Pantanal. Int. J. Trop. Insect Sci. 42:3253–3261.). The bibliographical survey and the construction of the identification keys were carried out in 2020, and some species were updated (Table 1) according to new updates: Coprophanaeus spitzi (Pêssoa, 1935), Dendropaemon nitidicollis d’Olsoufieff, 1924, e Onthophagus hircus Billberg, 1815 (Correa et al. 2022bCORREA, C.M.A., FERREIRA, K.R., ABOT, A.R., LOUZADA, J. & VAZ-DE-MELLO, F.Z. 2022b. Ivermectin impacts on dung beetle diversity and their ecological functions in two distinct Brazilian ecosystems. Ecol. Entomol. 47:736–748.). With new inventories carried out in the Pantanal region, this species list will probably increase. Finally, the use of this guide for identification of dung beetle species can be an important tool to help researchers and provide incentive for new studies on the dung beetle fauna in the Brazilian Pantanal.

Acknowledgments

AAM thanks Universidade Federal de Mato Grosso for financing this project project (Process number: 148/2017). AAM thanks A Fundação de Amparo à Ciência e Tecnologia do Estado de Pernambuco (FACEPE) for financial support (Process number: IBPG-0545-2.04/21) and CNPq (Conselho Nacional de Desenvolvimento Científico e Tecnológico) (Process 140194/2023-1).

EC thanks CAPES (Coordenação de Aperfeiçoamento de Pessoal de Nível Superior, Brazil) (Process 88887.654514/2021-00) and the Fundação de Amparo à Pesquisa do Estado do Amazonas (FAPEAM) – POSGRAD/scholarship/ for financing this study. CMAC thanks Coordenação de Aperfeiçoamento de Pessoal de Nível Superior ‘Bolsista CAPES/Brasil’ for postdoctoral grant (Process 88887.603414/2021-00). We also thank the Invertebrate Collection of INPA for allowing the use of photographic equipment. We also thank Vinicius Costa-Silva, Emanuel Ribas Gama and Arturo Gonzalez for providing some of the photos. We thanks to Mário Cupello for english revision.

FZVM, a CNPq PQ1A fellow, thanks Instituto Nacional de Ciencia e Tecnologia em Áreas úmidas (INAU), Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPQ) Process number: 431760/2018-7, and Fundação de Amparo a Pesquisa do Estado de Mato Grosso (FAPEMAT): Process number: 0147956/2017; PRONEM: 568005/2014. The authors thank Istituto Nacional de Ciência e Tecnologia em Áreas Úmidas (INAU) for the financial support.

Data Availability

Supporting data are available at <https://doi.org/10.48331/scielodata.HJNGV1.>.

References

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