Identification guide of dung beetle species (Coleoptera: Scarabaeidae: Scarabaeinae) of the Brazilian Pantanal

Guia de identificação de espécies de besouros rola-bostas (Coleoptera: Scarabaeidae: Scarabaeinae) do Pantanal Brasileiro

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, 2000). 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. 2014) and has a vegetation system arranged in a mosaic (i.e., different plant communities scattered side-by-side across space) (Pott et al. 2011). 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 2011; Nunes et al. 2014). Thus, the seasonal flooding and complex mosaic vegetation determine a biodiversity unique adapted to Brazilian Pantanal (Junk et al. 2006; Tomas et al. 2019).

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. 2020). Dung beetles are broadly distributed and highly diverse taxonomically and functionally (Halffter & Edmonds 1982; Hanski & Cambefort 1991). They are represented by more than 6,837 species worldwide which belong to 278 genera (Schoolmeesters 2023). 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 1966). 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 1966; Hanski & Cambefort 1991).

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. 2008) and greenhouse gas emissions from dung pats (Slade et al. 2016; Piccini et al. 2017). 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 1993; McGeoch et al. 2002; Nichols et al. 2007), are easily sampled with standardized protocols (Correa et al. 2018; Gardner et al. 2008; da Silva & Hernández 2015) and are diverse and relatively well characterized taxonomically (Vaz-de-Mello et al. 2011).

Although studies on dung beetle assemblages have been growing recently in the Brazilian Pantanal (Tissiani et al. 2015; Daniel & Vaz-de-Mello 2016, Pessôa et al. 2017; Correa et al. 2016, 2019, 2020, 2021, 2022a, b; Correa & da Silva 2022, Gonçalves et al. 2022), 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. 2005). 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. 2009), 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. 2007; Rodrigues et al. 2010; Tissiani et al. 2015; Daniel & Vaz-de-Mello 2016; Pessôa et al. 2017; Vaz-de-Mello et al. 2017; Correa et al. 2016, 2019, 2020), 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. 2017). The dung beetle species mentioned in the present study are classified according to their nesting behavior, following the classification by Hanski & Cambefort (1991).

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), 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).

Table 1.
List of dung beetle species that occur in the Pantanal with bibliography data separated by their functional group.

Species	Functional group	References
Canthidium angulicolle Balthasar, 1939	Paracoprid	Vaz-de-Mello et al. 2017
Canthidium barbacenicum Preudhomme de Borre, 1886	Paracoprid	Louzada et al. 2007; Pessôa et al. 2017; Vaz-de-Mello et al. 2017; Correia et al. 2020, 2022b
Canthidium breve (Germar, 1824)	Paracoprid	Louzada et al. 2007; Vaz-de-Mello et al. 2017
Canthidium cuprinum Harold, 1867	Paracoprid	Pessôa et al. 2017
Canthidium kelleri (Martinez, Halfter & Pereira)	Paracoprid	Daniel and Vaz-de-Mello 2015
Canthidium viride (Lucas, 1859)	Paracoprid	Pessôa et al. 2017; Vaz-de-Mello et al. 2017; Correia et al. 2022b
Canthon conformis (Harold, 1868)	Telecoprid	Correia et al. 2022b
Canthon chalybaeus Blanchard 1845	Telecoprid	Daniel and Vaz-de-Mello 2015
Canthon curvodilatatus Schmidt, 1922	Telecoprid	Tissiani et al. 2015; Correa et al. 2016, 2019, 2020; Vaz-de-Mello et al. 2017; Pessôa et al. 2017; Correira et al. 2022b
Canthon daguerrei Martínez, 1951	Telecoprid	Pessôa et al. 2017
Canthon edentulus Harold, 1868	Telecoprid	Daniel and Vaz-de-Mello 2015; Vaz-de-Mello et al. 2017; Correa et al. 2016, 2020
Canthon histrio (Lepeletier de Saint-Fargeau & Audinet-Serville, 1828)	Telecoprid	Louzada et al. 2010; Correa et al. 2016, 2019, 2020, 2022b; Pessôa et al. 2017; Vaz-de-Mello et al. 2017
Canthon mutabilis Lucas, 1859	Telecoprid	Correa et al. 2016, 2020; Vaz-de-Mello et al. 2017
Canthon maldonadoi (Pereira & Martinez, 1960)	Telecoprid	Daniel and Vaz-de-Mello 2015; Pessôa et al. 2017; Vaz-de-Mello et al. 2017
Canthon ornatus ornatus Redtenbacher, 1867	Telecoprid	Tissiani et al. 2015; Correa et al. 2016, 2020; Vaz-de-Mello et al. 2017
Canthon quadratus Blanchard, 1846	Telecoprid	Vaz-de-Mello et al. 2017
Canthon quinquemaculatus Castelnau, 1840	Telecoprid	Louzada et al. 2007; Pessôa et al. 2017; Vaz-de-Mello et al. 2017
Canthon substriatus Harold, 1868	Telecoprid	Correa et al. 2016, 2020; Vaz-de-Mello et al. 2017
Canthon unicolor Blanchard, 1843	Telecoprid	Vaz-de-Mello et al. 2017; Correa et al. 2022b
Canthon virens (Mannerheim, 1828)	Telecoprid	Daniel and Vaz-de-Mello 2015
Coprophanaeus bonariensis (Gory, 1844)	Paracoprid	Correa et al. 2016, 2019, 2020; Pessôa et al. 2017
Coprophanaeus cyanescens (d’Olsoufieff, 1924)	Paracoprid	Vaz-de-Mello et al. 2017; Correa et al. 2019; Correa et al. 2022b
Coprophanaeus ensifer (Germar, 1824)	Paracoprid	Daniel and Vaz-de-Mello 2015; Correa et al. 2016, 2020
Coprophanaeus spitzi (Pêssoa, 1935)	Paracoprid	Correia et al. 2022b
Coprophanaeus milon (Blanchard, 1846)	Paracoprid	Correa et al. 2016, 2020; Pessôa et al. 2017; Vaz-de-Mello et al. 2017
Dendropaemon nitidicollis d’Olsoufieff, 1924	Unknown	Correa et al. 2022a
Deltochillum cupreicolle (Blanchard)	Telecoprid	Daniel and Vaz-de-Mello 2015
Deltochilum silphoides Balthasar, 1939	Telecoprid	Pêssoa et al. 2017; Vaz-de-Mello et al. 2017
Deltochilum icaroides Balthasar, 1939	Telecoprid	Vaz-de-Mello et al. 2017
Deltochilum pseudoicarus Balthasar, 1939	Telecoprid	Daniel and Vaz-de-Mello 2015; Correa et al. 2016, 2019, 2020; Vaz-de-Mello et al. 2017
Dichotomius carbonarius (Mannerheim, 1929)	Paracoprid	Vaz-de-Mello et al. 2017
Dichotomius bos (Blanchard, 1845)	Paracoprid	Louzada et al. 2007; Rodrigues et al. 2010; Correa et al. 2016, 2019, 2020, 2022b; Pessôa et al. 2017; Vaz-de-Mello et al. 2017
Dichotomius cuprinus (Felsche, 1901)	Paracoprid	Daniel and Vaz-de-Mello 2015; Correia et al. 2022b
Dichotomius glaucus (Harold, 1869)	Paracoprid	Correa et al. 2016, 2019, 2020; Vaz-de-Mello et al. 2017
Dichotomius luctuosioides (Harold, 1869)	Paracoprid	Vaz-de-Mello et al. 2017
Dichotomius lycas (Felsche, 1901)	Paracoprid	Pessôa et al. 2017
Dichotomius nisus (Olivier, 1789)	Paracoprid	Louzada et al. 2007; Rodrigues et al. 2010; Tissiani et al. 2015; Correa et al. 2016, 2019, 2020, 2022b; Pessôa et al. 2017; Vaz-de-Mello et al. 2017
Dichotomius opacipennis (Luederwaldt, 1931)	Paracoprid	Tissiani et al. 2015; Correa et al. 2016, 2019, 2020, 2022b; Pessôa et al. 2017; Vaz-de-Mello et al. 2017
Dichotomius sexdentatus (Luederwaldt, 1925)	Paracoprid	Daniel and Vaz-de-Mello 2015
Digitonthophagus gazella (Fabricius, 1787)	Paracoprid	Correa et al. 2016, 2019, 2020, 2022b
Eurysternus aeneus Génier, 2009	Endocoprid	Vaz-de-Mello et al. 2017; Correa et al. 2019
Eurysternus caribaeus (Herbst, 1789)	Endocoprid	Correa et al. 2016, 2019, 2020; Pessôa et al. 2017; Vaz-de-Mello et al. 2017
Eurysternus jessopi Martinez, 1988	Endocoprid	Daniel and Vaz-de-Mello 2015
Eurysternus nigrovirens Génier, 2009	Endocoprid	Daniel and Vaz-de-Mello 2015; Correa et al. 2016, 2020, 2022b; Pessôa et al. 2017; Vaz-de-Mello et al. 2017
Eurysternus plebejus Harold, 1880	Endocoprid	Vaz-de-Mello et al. 2017
Genieridium cryptops (Arrow, 1913)	Endocoprid	Daniel and Vaz-de-Mello 2015; Pessôa et al. 2017
Genieridium bidens (Balthasar, 1938)	Endocoprid	Correa et al. 2016, 2020
Gromphas inermis Harold, 1869	Paracoprid	Daniel and Vaz-de-Mello 2015; Correa et al. 2016, 2020; Pessôa et al. 2017; Vaz-de-Mello et al. 2017
Isocopris foveolatus (Luederwaldt, 1931)	Paracoprid	CEMT
Malagoniella astyanax (Oliver, 1789)	Telecoprid	Daniel and Vaz-de-Mello 2015; Correa et al. 2016, 2020; Vaz-de-Mello et al. 2017
Malagoniella punctatostriata (Blanchard, 1846)	Telecoprid	Vaz-de-Mello et al. 2017; Correa et al. 2019; Correa et al. 2022b
Malagoniella puncticollis (Blanchard, 1846)	Telecoprid	Correa et al. 2016, 2019, 2020; Vaz-de-Mello et al. 2017
Ontherus azteca Harold, 1869	Paracoprid	CEMT
Ontherus aphodioides Burmeister, 1874	Paracoprid	CEMT
Ontherus appendiculatus (Mannerheim, 1829)	Paracoprid	Louzada et al. 2007; Rodrigues et al. 2010; Tissiani et al. 2015; Correa et al. 2016, 2019, 2020, 2022b; Pessôa et al. 2017; Vaz-de-Mello et al. 2017
Ontherus dentatus Luederwaldt, 1930	Paracoprid	Vaz-de-Mello et al. 2017
Ontherus digitatus Harold, 1869	Paracoprid	Vaz-de-Mello et al. 2017; Pessôa et al. 2017
Ontherus erosioides Luederwaldt, 1930	Paracoprid	Vaz-de-Mello et al. 2017
Ontherus sulcator (Fabricius, 1775)	Paracoprid	Louzada et al. 2007; Tissiani et al. 2015; Correa et al. 2016, 2020; Pessôa et al. 2017; Vaz-de-Mello et al. 2017
Onthophagus hircus Billberg, 1815	Paracoprid	Correira et al. 2022b
Phanaeus kirbyi Vigors, 1825	Paracoprid	Daniel and Vaz-de-Mello 2015
Phanaeus palaeno Blanchard, 1846	Paracoprid	Daniel and Vaz-de-Mello 2015; Vaz-de-Mello et al. 2017
Pseudocanthon xanthurus (Blanchard, 1845)	Telecoprid	Louzada et al. 2007; Vaz-de-Mello et al. 2017
Sulcophanaeus menelas (Castelnau, 1840)	Paracoprid	Vaz-de-Mello et al. 2017
Trichillidium quadridens Arrow, 1932	Endocoprid	Pessôa et al. 2017; Vaz-de-Mello et al. 2017
Trichillum externepunctatum Preudhomme de Borre, 1886	Endocoprid	Louzada et al. 2007; Tissiani et al. 2015; Correa et al. 2016, 2020, 2022b; Pessôa et al. 2017; Vaz-de-Mello et al. 2017
Zonocopris gibbicollis (Harold, 1868)	Endocoprid	Vaz-de-Mello et al. 2017
Uroxys corporaali Harold, 1868	Endocoprid	CEMT

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, 1868

   – 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, 2008

8. (5) Head with four clypeal teeth, rounded body (Figure 8A)............................................................. TrichillidiumVaz-de-Mello, 2008

   – Head with two clypeal teeth (Figure 8B)................................................................................... GenieridiumVaz-de-Mello, 2008

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, 1868

    – 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.

Figure 1.
(A) Eurysternus sp. scutellum visible dorsally with the elytra closed; (B) Malagoniella sp. scutellum not visible dorsally with closed elytra. Scale: A, 5mm; B, 3mm.

Figure 2.
(A) Eurysternus sp. in dorsal view; (B) Malagoniella sp. in dorsal view; (C) Metasterno de Eurysternus sp; (D) Metasterno de Malagoniella sp. Scale: A, 1mm; B, 5mm; C, 5mm; D, 5mm.

Figure 3.
(A) Trichillum sp. anterior leg with anterior trochantofemoral fóvea; (B) Eurysternus sp. anterior leg without anterior trochantofemoral fovea. Scale: A, 200µm; B, 1,19mm.

Figure 4.
(A) Trichillum sp. view of sternites; (B) Uroxys sp. view of sternites. Scale: A, 500µm; B, 1,19mm.

Figure 5.
(A) Trichillum sp. pseudepipleuron forming two lateral sinuosities, posterior partially (white arrows); (B) Genieridium sp. pseudepipleuron not forming two lateral sinuosities. Scale: A, 500µm; B, 1mm.

Figure 6.
(A) Trichillum sp. the clypeal tooth and the clypeogenal suture; (B) Eutrichilum sp. the clypeal tooth and the clypeogenal suture. Scale: A, 500µm; B, 2mm.

Figure 7.
(A) Eutrichilum sp. clypeogenal margin; (B) Besourenga sp. clypeogenal margin. Scale: A-B, 1mm.

Figure 8.
(A) Trichillidium sp. head with two orfour clypeal teeth; (B) Genieridium sp. head with two orfour clypeal teeth. Scale: A-B, 1mm.

Figure 9.
(A) Uroxys sp. side of pronotum with deep longitudinal sulcus; (B) Agamopus sp. side of pronotum without deep longitudinal sulcus. Scale: A-B, 1mm.

Figure 10.
(A) Agamopus sp. pygidium with deep transverse sulcus on disc; (B) Agamopus sp. Last mesotarsomere and metatarsomere without dentiform process above claw insertion; (C) Zonocopris sp. pygidium with disc without sulcus; (D) Zonocopris sp. last mesotarsomere and metatarsomere with dentiform process above claw insertion. Scale: A, 500µm; B-D 1mm.

Figure 11.
(A) Digitonthophagus sp. the first metatarsomere; (B) Dendropaemon sp. the first metatarsomere. Scale: A, 500µm; B, 1mm.

Figure 12.
(A). Digitonthophagus sp.; (B) Onthophagus sp. Scale: A-B, 1mm.

Figure 13.
(A) Dendropaemon sp. mesotarsus and metatarsos lacking claws; (B) Bolbites sp. mesotarsus and metatarsos with claws. Scale: A-B, 500µm.

Figure 14.
(A) Dendropaemon sp. mesotarsus and metatarsus with two to four tarsomeres; (B) Bolbites sp. mesotarsus and metatarsus with five tarsomeres. Scale: A, 500µm; B, 1mm.

Figure 15.
(A) Gromphas sp. basal antennomere of antennal club not concave apically; (B) Coprophanaeus sp. basal antennal club Lamela large, strongly concave apically. Scale: A-B, 1mm.

Figure 16.
(A) Coprophanaeus sp. clypeal margin with two teeth; (B) Diabroctis sp. clypeal margin with two teeth conspicuous middle teeth. Scale: A-B, 9mm.

Figure 17.
(A) Oxysternon sp. metasternum with long, dorsally curved, acute spiniform process; (B) Phanaeus sp. Metarsternum simply angled anteromedially, without spiniform process. Scale: A-B, 6mm.

Figure 18.
(A) Sulcophanaeus Anterior portion of circumnotal carina entire, not interrupted behind each eye; (B) Phanaeus Anterior portion of circumnotal ridge interrupted behind each eye. Scale: A-B, 1mm.

Figure 19.
(A) Bolbites sp. pronotum with two distinct, posteromedian fossae white arrows; (B) Anomiopus sp. posteromedian pronotal fossae absent. Scale: A-B, 1mm.

Figure 20.
(A) Anomiopus sp. tarsal claws reduced, straight or only weakly curved; (B) Deltochilum sp. tarsal claws large, strongly curved, falciform or angulate. Scale: A, 500µm; B, 1mm.

Figure 21.
(A) Deltochilum sp. mesotibia and metatibia not enlarged to apex; (B) Ateuchus sp. mesotibia, and usually also metatibia, abruptly widened to apex. Scale: A-B, 1mm.

Figure 22.
(A) Deltochilum sp. apex of some discal elytral interstriae with short carina or tubercles; (B) elytral interstriae without carina or tubercles. Scale: A-B, 1mm.

Figure 23.
(A) Pseudocanthon sp. posterior margin of head not margined between eyes; (B) Canthon sp. posterior border of head completely margined between eyes. Scale: A, 500µm; B, 1mm.

Figure 24.
(A) Holocanthon sp mentum completely divided longitudinally; (B) Canthon sp. mentum not completely divided. (white arrows). Scale: A-B, 1mm.

Figure 25.
(A) Canthon sp. pronotum with regular sculpturing, regularly convex, at most with posterior mean depression; (B) Anisocanthon sp. flat pronotum, with distributed depressions and elevations, and strong striated microsculpture. Scale: A-B, 1mm.

Figure 26.
(A) Ateuchus sp. hypomere deeply excavated anteriorly (B) Canthididium sp. hypomere weakly excavated anteriorly. Scale: A-B, 1mm.

Figure 27.
(A-B) Canthididium sp. apical internal angle of the protibial and mesosternum; (C-D) Ontherus sp apical internal angle of the protibial and mesosternum apical internal angle of the protibial and mesosternum. Scale: A-D, 1mm.

Figure 28.
(A) Ontherus sp. carina of the protibia and transverse ventral clypeal process; (B) Dichotomius sp. carina of the protibia and transverse ventral clypeal process. Scale: A-B, 1mm.

Figure 29.
(A) Isocopris sp. antennas with eight antennomeres; (B) Dichotomius sp. antennas with nine antennomeres. Scale: A-B, 1mm.

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

Figure 30.
(A) Eurysternus plebejus. Pronotum dark with a light spot anteriorly, just behind the head. Size 6-10mm; (B) Metaventrite disc with round fovea in the middle; (C) E. aeneus. Pronotum totally dark or mostly light with dark spots. variable size; (D) E. jessopi. Pronotum totally dark or mostly light with dark spots variable size. Scale: A-C, 1mm.

Figure 31.
(A-C) E. caribaeus. Light colored posterior femurs, at least in the apical Half, E. jessopi Pronotum mostly lighter, with well-defined dark spots; (B-D) E. parallelus. Posterior femurs totally dark like the pronotum. Scale: A, 3mm; B-D, 1mm.

Figure 32.
(A) E. parallelus. Dorsal body without any metallic luster. Elytra apex bristles straight. Size 9-14mm; (B) E. aeneus. 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. Scale: A-B, 1mm.

Figure 33.
(A) E. nigrovirens. Head surface with most no metallic reflections, except for a narrow metallic green band anteriorly on the clypeus; (B) E. aeneus. Head surface mostly glossy and with strong metallic reflections between ocellates; (C) E. nigrovirens. Lateral edge of male hind tibia curved in half in dorsal view; (D) E. aeneus. Lateral edgeof male hind tibia almost straight or straight in the middle in dorsal view. Scale: A-B, 1mm.

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, 1868

   – 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, 1868

    – 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)

Figure 34.
(A) Canthon. Pygidium and propygidium separated by carina; (B) Canthon. Pygidium and propygidium not separated by carina. Scale: A-B, 1mm.

Figure 35.
(A) Canthon mutabilis. Head with quadridented clypeus; (B) Canthon (ornatos) ornatos. Head with bidented clypeus. Scale: A, 1mm; B, 2mm.

Figure 36.
(A) C. mutabilis. Pronotum in dorsal view, with straight lateral margin and serrated in ventral view; (B) C. curvodilatatus. Pronotum with straight lateral margin, in dorsal and ventral view. Scale: A-B, 1mm.

Figure 37.
(A) C. substriatus. Gena anteriorly with visible tooth and clypeus with two long teeth bulging, rounded, or subtriangular at apex, not acute; (B) C. virens. Clypeus with two long, triangular teeth, acute, not bulging or rounded at the apex. Scale: A, 1mm; B, 2mm.

Figure 38.
(A) C. virens. Pronotum is completely punctuated, being more subtle on the disc than on the sides; shiny elytra with well-marked striations; subtly punctuated interstriae; (B) C. chalybaeus. Pronotum with almost imperceptible punctuation on the disc; opaque elytra, striae not as marked as before; (C) C. virens. Metafemur with complete anterior and incomplete posterior margination; (D) C. chalybaeus. interstria with micro reticulated punctuation; metafemur with complete anterior and posterior margination. Scale: A-D, 1mm.

Figure 39.
(A-B) C. daguerrei. Short clypeal teeth; complete hypomeral carina; pronotum and elytra dark; (C-D) C. ornatos Clypeal teeth not as short as before; incomplete hypomeral carina; light pronotum with black stain and green reflections and dark elytra. Scale: A-D, 1mm.

Figure 40.
(A) C. quinquemaculatus. Head with quadridented clypeus; (B) C. maldonadoi. Head with bidented clypeus, can be very or poorly defined; (C) C. edentulus. Clypeal teeth are short and rounded; “V” shape. Scale: A-B, 1mm.

Figure 41.
(A) C. maldonadoi. Central clypeal teeth separated by well-delimited U-shaped emargination; pronotum and elytra light with dark outline; (B) C. edentulus Clypeal teeth are little delimited, pronotum and elytra dark. Scale: A-B, 1mm.

Figure 42.
(A-B) C. edentulus. Clypeal teeth are short and rounded; pygidium longer than wide and with thick punctuation; size 5-6,5mm; (C-D) C. quadratus. Clypeal teeth only weakly indicated, not salient, separated by wide emargination; Pygidium wider than long and fine punctuation compared to the previous one; size 10-12mm. Scale: A-D, 1mm.

Figure 43.
(A-C) C. unicolor. Pronotum and elytra dark; metafemur with anterior and posterior margination; (C-D) C. histrio. Pronotum and elytra may be yellow with black spots; metafemur only with anterior margination. Scale: A-D, 1mm.

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)

Figure 44.
(A) C. milon. Elytral striae smooth; interstriae with subtle punctuation; (B) C. cyanescens. Elytral striae sculpted; rugopuncted interstriae.Scale: A, 7mm; B, 10mm.

Figure 45.
(A) C. milon. Margin behind eye absent; head with horn trident laminated; (B) C. cyanescens. Margin behind the eye present; head with cylindrical horn. Scale: A, 1mm; B, 5mm.

Figure 46.
(A) C. bonariensis. Elytral striae 2-4 only slightly wavy; width of striations at narrowings at least three-quarters of that at widest points; size 25-35mm; (B) C. ensifer. 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. Scale: A-B, 10mm.

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)

Figure 47.
(A) D. bos. Clypeal margin rounded or weakly emarginated; if bidentate, small, non-margined teeth; (B) D. luctuosioides. Clypeus distinctly bidentate, teeth usually margined. Scale: A-B, 1mm.

Figure 48.
(A-B) D. bos. Elytra with smooth striations and no punctuation; pronotum with median longitudinal sulcus. Scale: A-B, 1mm.

Figure 49.
(A-B) D. luctuosioides. Elytra with punctiform striations; pronotum without median sulcus. Scale: A-B, 1mm.

Figure 50.
(A) D. lycas. Metallic elytra (satin gloss) coloration in green, reddish or almost burgundy brown; (B) D. opacipennis. Pronotum and elytra staining black, opaque or glossy. Scale: A-B, 1mm.

Figure 51.
(A) D. glaucus. Pronotum mesoanteriorly not flattened; base of the pronotum without evident punctuation in the central region; (B) D. lycas. Pronotum mesoanteriorly flattened; base of pronotum with two rows of ocellar punctuation evident. Scale: A-B, 1mm.

Figure 52.
(A) D. glaucus. Dorsal surface monochromatic, glossy greenish satin; teeth margined, triangular and pointed; pronotum with flat anterior lateral edge; (B) D. cuprinus. Dorsal surface bicolor, black pronotum and glossy satin burgundy elytra; rounded and flat teeth; pronotum with anterior lateral edge forming an almost triangular tip. Scale: A-B, 1mm.

Figure 53.
(A) D. sexdentatus. Sides of metasternum and hypomere with dense and long setae, red to yellowish-brown; (B) D. opacipennis. Sides of metasternum and hypomerus without setae as described above. Scale: A-B, 1mm.

Figure 54.
(A) D. sexdentatus. Clypeus with six teeth; pronotum with median anterior slope with ocellated punctuations; (B) D. nisus. Clypeus with two teeth; posterior margin of the pronotum with ocellated punctuations. Scale: A-B, 1mm.

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

Figure 55.
(A) O. digitatus. Fronto clypeal suture without carina and/or tubercles; (B) O. appendiculatus. Posterior border of metacoxa smooth. Metasternum with most of the punctuation of the oval lateral edges, with little defined edges. Scale: A-B, 1mm.

Figure 56.
(A) O. appendiculatus. Abdominal sternites medially set, bristles similar to those on the lateral aspect of the metasternum; (B) O. aphodioides. Abdominal sternites glabrous medially, with at most one row of short bristles along the posterior border of the metacoxa. Scale: A-B, 1mm.

Figure 57.
(A-B) O. appendiculatus. Posterior border of metacoxa smooth. Metasternum with most of the punctuation of the oval lateral edges, with little defined edges; (C-D) O. azteca. Posterior edge of metacoxa slightly serrated. Metasternum with most of the punctuation of the lateral edges rounded, clearly delimited. Scale: A-D, 1mm.

Figure 58.
(A) O. aphodioides. Median metasternum lobe length less than twice its width; (B) O. dentatus. Median metasternum lobe length twice its maximum width. Scale: A-B, 1mm.

Figure 59.
(A-B) O. aphodioides. Pygidium wider than long, pits of moderate size and deeply imprinted; space between glossy scores; (C-D) O. sulcator. Pygidium as wide as long, punctuations small and weakly printed. Scale: A-B, 1mm.

Figure 60.
(A) O. dentatus. Posterior edge of the metatrochanter complete with sulcus; pygidium with final score; (B) O. erosioides. Posterior edge of metatrochanter partially with sulcus; pygidium with thick score. Scale: A-B, 1mm.

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, 1939

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

   – Head much wider than long (Figure 62B)...........................................................................................................D. silphoides (Balthasar, 1939) (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, 1939

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

Figure 61.
(A) D. silphoides. Bidentate clypeus; (B) D. pseudoicarus. Clypeus quadridentate, medial teeth stronger and longer than others. Scale: A-B, 1mm.

Figure 62.
(A) D. icaroides. Head much wider than long; (B) silphoides. Pronotum and elytra of the same color, dark with greenish reflections. Scale: A-B, 1mm.

Figure 63.
(A) D. icaroides, Pronotum and elytra of the same color, dark with greenish reflections; (B) D. cupreicoile, Pronotum yellow with dark spot in the center, elytra of uniform dark coloration. Scale. A-B, 1mm.

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.

Figure 64.
Habitus. (A) Canthon curvodilatatus; (B) C. mutabilis; (C) C. quadratus; (D) C. quinquemaculatus; (E) C. substriatus; (F) Coprophanaeus bonariensis; (G) C. cyanescens; (H) C. ensifer; (I) C. milon; (J) Deitchler pseudoicarus; (K) D. silphoides; (L) Dichotomius bos; (M) D. luctuosioides; (N) D. lycas; (O) D. nisus; (P) D. opacipennis. Scale bar: A-E; M-P 1mm; G, I-L 7mm; H 10mm.

Figure 65.
Habitus. (A) D. sexdentatus; (B) Eurysternus aeneus; (C) E. caribaeus; (D) Ontherus aphodioides; (E) O. appendiculatus; (F) O. azteca; (G) O. dentatus; (H) O. digitatus; (I) O. erosioides; (J) O. sulcator. Scale bar: B, D-J, 1mm; C, 5mm; A, 7mm.

Ateuchus Weber, 1801

Genus with paracoprid representatives and 101 species currently recognized as valid distributed in the Americas (Schoolmeesters 2023). Except for some regional works, the genus was last revised by Harold (1868) 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, 1868) (Pessôa et al. 2017), Ateuchus latus (Boucomont, 1928) (Vaz-de-Mello et al. 2007), 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 2023), 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, 1939, 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 2018; Kohlmann et al. 2018; Moctezuma et al. 2019; Carvalho de Santana et al. 2019).

Canthon Hoffmannsegg, 1817

It is one of the most diverse genera among the Scarabaeinae, exclusive to the Americas, with currently 163 recognized species (Schoolmeesters 2023). 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 2018 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, 1868, 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, 1868, 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 2023). 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), and further descriptions and revalidations (Kohlmann & Solís 2012, Cupello & Vaz-de-Mello 2013, 2014). 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 2023), 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 2012; González-Alvarado & Vaz-de-Mello 2014; Silva et al. 2015; Nazaré-Silva & Silva 2021). The species are telecoprid. Deltochilum is here represented by four species belonging to three different subgenera: Deltochilum (Deltochilum) silphoidesBalthasar, 1939, D. (Hybomidium) pseudoicarusBalthasar, 1939, Deltochilum (Aganhyboma) cupreicolle (Blanchard, 1846) and Deltochilum (A.) icarioidesBalthasar, 1939.

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 2023), 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 2017). 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. 2017; Génier & Moretto 2017). 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. 2020), 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 2008).

Eurysternus Dalman, 1824

Genus with a Neotropical distribution, with 53 described species (Schoolmeesters 2023), recently revised by Génier (2009). 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, 2009, E. jessopi Martínez, 1988, E. nigrovirensGénier, 2009, and E. plebejus Harold, 1880.

Genieridium Vaz-de-Mello, 2008

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 2008). 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 (2013b, 2015). 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). 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). 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, 1996 (24 species) and PlanontherusGénier, 1996 (two species) (Schoolmeesters 2023), revised by Génier (1996, 1998; see also González & Medina 2015). Its representatives present paracoprid habits. We recorded six species: Ontherus appendiculatus (Mannerheim, 1829), O. aphodioides Burmeister, 1874, O. dentatus Luederwaldt, 1930, O. digitatusHarold, 1868, O. erosioides Luederwaldt, 1930, and O. sulcator (Fabricius, 1775).

Phanaeus MacLeay, 1819

A genus with 82 species (Edmonds & Zídek 2012; Schoolmeesters 2023), two of which are inhabitants of grassland and savanna formations in Brazil and are occasionally collected in cultivated pastures (Edmonds 1994). 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 2023), its representatives are paracoprids. The last revision is by Edmonds (2000), and it was represented here by only one species, Sulcophanaeus menelas (Castelnau, 1840).

Trichillidium Vaz-de-Mello, 2008

Genus with four species, usually found in the Amazon, Chaco and Central America and with coprophagous diet (Vaz-de-Mello 2008). Here represented by Trichillidium quadridens (Arrow, 1932).

Trichillum Harold, 1868

Genus with Neotropical distribution, with 11 described species (Schoolmeesters 2023). 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 2007). In the Brazilian Pantanal, it is represented by Zonocopris gibbicollis (Harold, 1868).

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, 2008, 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. 2022b) and Dendropaemon were recently reported from the Pantanal region in Aquidauana and Miranda – Mato Grosso do Sul, Brazil (see Correa et al. 2022a; Gonçalves et al. 2022).

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. 2022a). 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. 2022b). 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.>.

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Edited by

Associate Editor

José Mermudes

Publication Dates

• Publication in this collection
  30 June 2023
• Date of issue
  2023

History

• Received
  18 Jan 2023
• Accepted
  16 May 2023