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

Mota, Adriana Andrade; Carvalho, Edrielly; Correa, César M. A.; Vaz-de-Mello, Fernando Z.

Adriana Andrade Mota ^* * Corresponding author: adrianaandrademota@gmail.com

specimens identification

photo production

elaboration of the keys and writing of the manuscript

Universidade Rural Federal de Pernambuco, Departamento de Agronomia/Fitossanidade, 52171-900, Recife, PE, Brasil.

Universidade Federal do Paraná, Departamento de Zoologia, Setor de Ciências Biológicas, 81531-980, Curitiba, PR, Brasil.

https://orcid.org/0000-0001-9766-3967

Edrielly Carvalho

photo production and editing

elaboration of the keys and writing of the manuscript

Instituto Nacional de Pesquisas da Amazônia, Programa de Pós-Graduação em Entomologia, 69011-970, Manaus, AM, Brasil.

César M. A. Correa

manuscript writing and review

Universidade Estadual de Mato Grosso do Sul, Laboratório de Bioecologia de Scarabaeoidea, 79200-000, Aquidauana, MS, Brasil.

Fernando Z. Vaz-de-Mello

specimens identification

Universidade Federal de Mato Grosso, Instituto de Biociências, Departamento de Biologia e Zoologia, Av. Fernando Correia da Costa, Coxipó, 78060-900, Cuiabá, MT, Brasil.

* Corresponding author: adrianaandrademota@gmail.com

Associate Editor

José Mermudes

Author Contributions

Adriana Andrade Mota: specimens identification; photo production; elaboration of the keys and writing of the manuscript.

Edrielly Carvalho: photo production and editing; elaboration of the keys and writing of the manuscript.

César Murilo de Albuquerque Correa: manuscript writing and review.

Fernando Vaz-de-Mello: specimens identification.

Conflicts of Interest

All authors have seen and agree with the contents of the manuscript and there is no conflict of interest, including specific financial interest and relationships and affiliations relevant to the subject of manuscript.

Figuras (65)
Tabelas (1)

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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.

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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.

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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.

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

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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.

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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.

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

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

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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.

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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.

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

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

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

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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.

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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.

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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.

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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.

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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.

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

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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.

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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.

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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.

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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.

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

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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.

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

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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.

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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.

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

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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.

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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.

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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.

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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.

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

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

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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.

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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.

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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.

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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.

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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.

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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.

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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.

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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.

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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.

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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.

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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.

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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.

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

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

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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.

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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.

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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.

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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.

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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.

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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.

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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.

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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.

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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.

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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.

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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.

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

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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.

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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.

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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.

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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.

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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, 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.	Paracoprid	Vaz-de-Mello et al. 2017
Canthidium barbacenicum Preudhomme de Borre, 1886	Paracoprid	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.; 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; Correia 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., 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.
Canthidium breve (Germar, 1824)	Paracoprid	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.; Vaz-de-Mello et al. 2017
Canthidium cuprinum Harold, 1867	Paracoprid	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.
Canthidium kelleri (Martinez, Halfter & Pereira)	Paracoprid	Daniel and Vaz-de-Mello 2015
Canthidium viride (Lucas, 1859)	Paracoprid	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; Correia 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.
Canthon conformis (Harold, 1868HAROLD, E. 1868. Die arten der gattung Choeridium. Coleopterologische Hefte 4, 32–76.)	Telecoprid	Correia 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.
Canthon chalybaeus Blanchard 1845	Telecoprid	Daniel and Vaz-de-Mello 2015
Canthon curvodilatatus Schmidt, 1922	Telecoprid	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.; 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.; Vaz-de-Mello et al. 2017; 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.; Correira et al. 2022b
Canthon daguerrei Martínez, 1951	Telecoprid	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.
Canthon edentulus Harold, 1868HAROLD, E. 1868. Die arten der gattung Choeridium. Coleopterologische Hefte 4, 32–76.	Telecoprid	Daniel and Vaz-de-Mello 2015; 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., 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.
Canthon histrio (Lepeletier de Saint-Fargeau & Audinet-Serville, 1828)	Telecoprid	Louzada et al. 2010; 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., 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.; 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
Canthon mutabilis Lucas, 1859	Telecoprid	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., 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.; Vaz-de-Mello et al. 2017
Canthon maldonadoi (Pereira & Martinez, 1960)	Telecoprid	Daniel and Vaz-de-Mello 2015; 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
Canthon ornatus ornatus Redtenbacher, 1867	Telecoprid	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.; 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., 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.; 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. 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.; 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
Canthon substriatus Harold, 1868HAROLD, E. 1868. Die arten der gattung Choeridium. Coleopterologische Hefte 4, 32–76.	Telecoprid	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., 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.; Vaz-de-Mello et al. 2017
Canthon unicolor Blanchard, 1843	Telecoprid	Vaz-de-Mello et al. 2017; 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.
Canthon virens (Mannerheim, 1828)	Telecoprid	Daniel and Vaz-de-Mello 2015
Coprophanaeus bonariensis (Gory, 1844)	Paracoprid	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.; 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.
Coprophanaeus cyanescens (d’Olsoufieff, 1924)	Paracoprid	Vaz-de-Mello et al. 2017; Correa et al. 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.; 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.
Coprophanaeus ensifer (Germar, 1824)	Paracoprid	Daniel and Vaz-de-Mello 2015; 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., 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.
Coprophanaeus spitzi (Pêssoa, 1935)	Paracoprid	Correia 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.
Coprophanaeus milon (Blanchard, 1846)	Paracoprid	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., 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.; 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
Dendropaemon nitidicollis d’Olsoufieff, 1924	Unknown	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.
Deltochillum cupreicolle (Blanchard)	Telecoprid	Daniel and Vaz-de-Mello 2015
Deltochilum 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.	Telecoprid	Pêssoa et al. 2017; Vaz-de-Mello et al. 2017
Deltochilum icaroides 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.	Telecoprid	Vaz-de-Mello et al. 2017
Deltochilum pseudoicarus 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.	Telecoprid	Daniel and Vaz-de-Mello 2015; 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.; 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. 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.; 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., 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.; 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
Dichotomius cuprinus (Felsche, 1901)	Paracoprid	Daniel and Vaz-de-Mello 2015; Correia 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.
Dichotomius glaucus (Harold, 1869)	Paracoprid	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.; 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. 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.
Dichotomius nisus (Olivier, 1789)	Paracoprid	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.; 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., 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.; 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
Dichotomius opacipennis (Luederwaldt, 1931)	Paracoprid	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.; 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., 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.; 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
Dichotomius sexdentatus (Luederwaldt, 1925)	Paracoprid	Daniel and Vaz-de-Mello 2015
Digitonthophagus gazella (Fabricius, 1787)	Paracoprid	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., 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.
Eurysternus aeneus Génier, 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.	Endocoprid	Vaz-de-Mello et al. 2017; Correa et al. 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.
Eurysternus caribaeus (Herbst, 1789)	Endocoprid	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.; 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
Eurysternus jessopi Martinez, 1988	Endocoprid	Daniel and Vaz-de-Mello 2015
Eurysternus nigrovirens Génier, 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.	Endocoprid	Daniel and Vaz-de-Mello 2015; 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., 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., 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.; 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
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. 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.
Genieridium bidens (Balthasar, 1938)	Endocoprid	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., 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.
Gromphas inermis Harold, 1869	Paracoprid	Daniel and Vaz-de-Mello 2015; 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., 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.; 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
Isocopris foveolatus (Luederwaldt, 1931)	Paracoprid	CEMT
Malagoniella astyanax (Oliver, 1789)	Telecoprid	Daniel and Vaz-de-Mello 2015; 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., 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.; Vaz-de-Mello et al. 2017
Malagoniella punctatostriata (Blanchard, 1846)	Telecoprid	Vaz-de-Mello et al. 2017; Correa et al. 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.; 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.
Malagoniella puncticollis (Blanchard, 1846)	Telecoprid	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.; 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. 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.; 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., 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.; 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
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. 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.
Ontherus erosioides Luederwaldt, 1930	Paracoprid	Vaz-de-Mello et al. 2017
Ontherus sulcator (Fabricius, 1775)	Paracoprid	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.; 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.; 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., 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.; 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
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. 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.; 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. 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
Trichillum externepunctatum Preudhomme de Borre, 1886	Endocoprid	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.; 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.; 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., 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., 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.; 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
Zonocopris gibbicollis (Harold, 1868HAROLD, E. 1868. Die arten der gattung Choeridium. Coleopterologische Hefte 4, 32–76.)	Endocoprid	Vaz-de-Mello et al. 2017
Uroxys corporaali Harold, 1868HAROLD, E. 1868. Die arten der gattung Choeridium. Coleopterologische Hefte 4, 32–76.	Endocoprid	CEMT

[1] * Corresponding author: adrianaandrademota@gmail.com

Brasil

Brasil

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

Resumo