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First report on dung beetles in intra-Amazonian savannahs in Roraima, Brazil

Primeiro registro de besouros rola-bosta em savanas intra-amazônicas de Roraima, Brasil

Abstracts

This is the first study to address the dung beetle (Coleoptera: Scarabaeidae: Scarabaeinae) diversity in intra-Amazonian savannahs in the state of Roraima, Brazil. Our aim was to survey the dung beetle fauna associated with these savannahs (regionally called 'lavrado'), since little is known about the dung beetles from this environment. We conducted three field samples using pitfall traps baited with human dung in savannah areas near the city of Boa Vista during the rainy seasons of 1996, 1997, and 2008. We collected 383 individuals from ten species, wherein six have no previous record in intra-Amazonian savannahs. The most abundant species were Ontherus appendiculatus (Mannerheim, 1829), Canthidium aff. humerale (Germar, 1813), Dichotomius nisus (Olivier, 1789), and Pseudocanthon aff. xanthurus (Blanchard, 1846). We believe that knowing the dung beetles diversity associated with the intra-Amazonian savannahs is ideal for understanding the occurrence and distribution of these organisms in a highly threatened environment, it thus being the first step towards conservation strategy development.

Tropical environments; new species occurrence; 'lavrado'; Scarabaeinae


Este é o primeiro estudo da diversidade de besouros rola-bosta (Coleoptera: Scarabaeidae: Scarabaeinae) nas savanas intra-amazônicas de Roraima, Brasil. Nosso objetivo foi inventariar a fauna de besouros associada com savanas (conhecidas regionalmente como 'lavrado'), uma vez que pouco se sabe sobre esses besouros nesse ambiente. Nós amostramos em savanas próximas a cidade de Boa Vista durante a estação chuvosa nos anos 1996, 1997 e 2008, utilizando armadilhas pitfall iscadas com fezes humanas. Coletamos 383 indivíduos de dez espécies, das quais seis não haviam ocorrência prévia para savanas intra-amazônicas. As espécies mais abundantes foram Ontherus appendiculatus (Mannerheim, 1829), Canthidium aff. humerale (Germar, 1813), Dichotomius nisus (Olivier, 1789) e Pseudocanthon aff. xanthurus (Blanchard, 1846). Nós acreditamos que conhecendo a diversidade de besouros associada ès savanas intra-Amazônicas se torna ideal para o entendimento da ocorrência e distribuição desses organismos em um ambiente altamente ameaçado, sendo o primeiro passo para a tomada de estratégias de conservação.

Ambientes tropicais; nova ocorrência de espécies; lavrado; Scarabaeinae


Introduction

The Amazon environment comprises a mosaic of ecosystems, including deciduous forests, 'campinaramas' (meadows), semi-deciduous seasonal forests, rain and montane forests, dry land forests, floodplain forests, woodlands and intra-Amazonian savannahs (Junk 1983JUNK, W.J. 1983. As águas da região Amazônica. In: Amazônia: Desenvolvimento, Integração E Ecologia (Salati, E. Junk, W.J. Shubart, H.O.R. & Oliveira, A. eds.). Editora Brasiliense, São Paulo, pp. 45-100., Ribeiro et al. 1999RIBEIRO, J.E.L.S., HOPKINS, M.J.G., VICENTINI, A., SOTHERS, C.A., COSTA, M.A.S., BRITO, J.M., SOUZA, M.A.D., MARTINS, L.H.P., LOHMANN, L.G., ASSUNÇÃO, P.A.C.L., PEREIRA, E.C., SILVA, C.F., MESQUITA, M.R., & PROCOPIO, L.C. 1999. Flora da reserva Ducke: guia de identificação das plantas vasculares de uma floresta de terra-firme na Amazônia Central. DFID - INPA, Manaus.). The Brazilian intra-Amazonian savannah landscape encompasses a predominantly grassy-woody vegetation (e.g. grasses, sedges, and small herbs), with low arboreal-shrub density (Barbosa et al. 2007BARBOSA, R.I., CAMPOS, C., PINTO, F. & FEARNSIDE, P. 2007. The “Lavrados” of Roraima: Biodiversity and conservation of Brazil's Amazonian savannas. Funct. Ecosyst. Communities. 1(1):29-41., Sarmiento 1984SARMIENTO, G. 1984. The Ecology of Neotropical Savannahs. Harvard University Press, Cambridge, UK.) and specific edaphic characteristics (Cavalcante et al. 2014CAVALCANTE, C.O., FLORES, A.S. & BARBOSA, R.I. 2014. Fatores edáficos determinando a ocorrência de leguminosas herbáceas em savanas amazônicas. Acta Amaz. 44(3):379-386, 10.1590/1809-4392201300954.
https://doi.org/10.1590/1809-43922013009...
, Sanaiotti et al. 2002SANAIOTTI, T.M., MARTINELLI, L.A., VICTORIA, R.L., TRUMBORE, S.E. & CAMARGO, P.B. 2002. Past vegetation changes in Amazon savannahs determined using carbon isotopes of soil organic matter. Biotropica 34:2-16, 10.1111/j.1744-7429.2002.tb00237.x.
https://doi.org/10.1111/j.1744-7429.2002...
). These savannahs are located in the extreme north of the Brazilian Amazon forest, mainly in the triple border between Venezuela, Brazil and Guyana (Barbosa et al. 2007, Barbosa & Fearnside 2005BARBOSA, R.I. & FEARNSIDE, P.M. 2005. Fire frequency and area burned in the Roraima savannas of Brazilian Amazonia. For. Ecol. Manage. 204:371-384, 10.1016/j.foreco.2004.09.011.
https://doi.org/10.1016/j.foreco.2004.09...
, Prance 1996PRANCE, G.T. 1996. Islands in Amazonia. Philos. Trans. R. Soc. B Biol. Sci. 351, 823-833, 10.1098/rstb.1996.0077.
https://doi.org/10.1098/rstb.1996.0077...
). This eco-region known as Guyana's shield (Hubber 2006HUBBER, O. 2006. Herbaceous ecosystems on the Guyana Shield, a regional overview. J. Biogeogr. 33:464-475, 10.1111/j.1365-2699.2005.01454.x.
https://doi.org/10.1111/j.1365-2699.2005...
) comprises the largest contiguous area of intra-Amazonian savannah (around 54.000 km2), of which approximately 72% are located in the northeastern region of the Roraima State in Brazil (Barbosa & Fearnside 2005BARBOSA, R.I. & FEARNSIDE, P.M. 2005. Fire frequency and area burned in the Roraima savannas of Brazilian Amazonia. For. Ecol. Manage. 204:371-384, 10.1016/j.foreco.2004.09.011.
https://doi.org/10.1016/j.foreco.2004.09...
, Barbosa & Campos 2011BARBOSA, R.I. & CAMPOS, C. 2011. Detection and geographical distribution of clearing areas in the savannas ('lavrado') of Roraima using Google Earth web tool. J. Geogr. Reg. Plann. 4(3):122-136., Copobianco et al. 2001COPOBIANCO, J.P., VERĺSSIMO, A., MOREIRA, A., SAWER, D., IKEDA, S. & PINTO, L.P. 2001. Biodiversidade na Amazônia Brasileira: Avaliação de Ações Prioritárias para a Conservação, Uso Sustentável e Repartição de Benefícios. Estação Liberdade, São Paulo. COSTA, C.M.Q., BARRETTO, J.W. & DE MOURA, R.D.C. 2014. Changes in the dung beetle community in response to restinga forest degradation. J. Insect Conserv. 18:895-902, 10.1007/s10841-014-9697-6.
https://doi.org/10.1007/s10841-014-9697-...
COSTA, C.M.Q., SILVA, F.A.B., FARIAS, Â.I. & MOURA, R.D.C. 2009. Diversidade de Scarabaeinae (Coleoptera, Scarabaeidae) coletados com armadilha de interceptação de vôo no Refúgio Ecológico Charles Darwin, Igarassu-PE, Brasil. Rev. Bras. Entomol. 53:88-94, 10.1590/S0085-56262009000100021.
https://doi.org/10.1590/S0085-5626200900...
COSTA, F.C., PESSOA, K.K.T., LIBERAL, C.N., FILGUEIRAS, B.K.C., SALOMAO, R.P. & IANNUZZI, L. 2013. What is the importance of open habitat in a predominantly closed forest area to the dung beetle (Coleoptera, Scarabaeinae) assemblage? Rev. Bras. Entomol. 57:329-334, 10.1590/S0085-56262013000300012.
https://doi.org/10.1590/S0085-5626201300...
, Ferreira 2001FERREIRA, L.V. 2001. Identificação de áreas prioritárias para a conservação da biodiversidade por meio da representatividade das unidades de conservação e tipos de vegetação nas ecorregiões da Amazônia brasileira, In: Biodiversidade na Amazônia Brasileira (Veríssimo, A. Moreira, A. Sawer, D. Santos, I. Pinto, L.P. & Copobianco, J.P. eds.). Editora Liberdade, Instituto Socioambiental, São Paulo, pp. 268-286.). Furthermore, those savannahs are regionally known as 'lavrado' (Barbosa et al. 2007, Sanaiotti 1997SANAIOTTI, T.M. 1997. Comparação fitossociológica de quatro savanas de Roraima. In: Homem, Ambiente e Ecologia no Estado de Roraima (Barbosa, R.I Ferreira, E.J.G & Castellón, E. eds.). INPA, Manaus, pp. 481-488.), which means 'place where the trees are absent' (Vanzolini & Carvalho 1991VANZOLINI, P.E. & CARVALHO, C.M. 1991. Two sibling and sympatric species of Gymnophthalmus in Roraima, Brazil (Sauria Teiidae). Pap. Avulsos Zool. 37:173-226.) and reflects its typical non-forest vegetation.

Despite a review about the Phanaeini tribe (Pacheco & Vaz-de-Mello 2015PACHECO, T.L. & VAZ-DE-MELLO, F.Z. 2015. Dung beetles of the tribe Phanaeini (Coleoptera: Scarabaeidade: Scarabaeinae) from Roraima state, Northern Brazil: checklist and key to species. Biota Neotrop. 15(2): 1-9, 10.1590/1676-06032015014514.
https://doi.org/10.1590/1676-06032015014...
), no work to our knowledge has reported the dung beetle fauna from the intra-Amazonian savannahs in Roraima. The dung beetles are a responsive taxonomic group (Bicknell et al. 2014BICKNELL, J.E., PHELPS, S.P., DAVIES, R.G., MANN, D.J., STRUEBIG, M.J. & DAVIES, Z.G. 2014. Dung beetles as indicators for rapid impacts assessments: Evaluating best practice forestry in the neotropics. Ecol. Indic. 43:154-161, 10.1016/j.ecolind.2014.02.030.
https://doi.org/10.1016/j.ecolind.2014.0...
), with more than 600 species recorded in Brazil (Vaz-de-Mello 2000VAZ-DE-MELLO, F.Z. 2000. Estado Atual de conhecimentos dos Scarabaeidae s. str. (Coleoptera: Scarabaeoidea) do Brasil. In: Hacia Un Proyecto CYTED Para El Inventario Y Estimación de La Diversidad Entonológica Em Iberoamérica: PrIBES-2000 (Martín-Piera, F. Morrone, J.J. & Melic, A. eds.). Sociedad Entomologica Aragonesa & CYTED, Zaragoza, pp. 181-195.) and distributed throughout all Brazilian terrestrial ecosystems. Regarding their diversity within Brazilian opened areas, studies have recorded between 13 and 66 species from the Brazilian southeastern savannahs (known as 'cerrado') and pasturelands (Abot et al. 2012ABOT, A.R., PUKER, A., TAIRA, T.L., RODRIGUES, S.R., KORASAKI, V. & OLIVEIRA, H.N. 2012. Abundance and diversity of coprophagous beetles (Coleoptera: Scarabaeidae) caught with a light trap in a pasture area of the Brazilian Cerrado. Stud. Neotrop. Fauna Environ. 47:53-60, 10.1080/01650521.2012.662846.
https://doi.org/10.1080/01650521.2012.66...
, Almeida et al. 2011ALMEIDA, S., LOUZADA, J., SPERBER, C. & BARLOW, J. 2011. Subtle Land-Use Change and Tropical Biodiversity: Dung Beetle Communities in Cerrado Grasslands and Exotic Pastures. Biotropica 43:704-710, 10.1111/j.1744-7429.2011.00751.x.
https://doi.org/10.1111/j.1744-7429.2011...
, Durães et al. 2005DURÃES, R., MARTINS, W.P. & VAZ-DE-MELLOS, F.Z. 2005. Dung beetle (Coleoptera: Scarabaeidae) assemblages across a natural forest-cerrado ecotone in Minas Gerais, Brazil. Neotrop. Entomol. 34:721-731, 10.1590/S1519-566X2005000500003.
https://doi.org/10.1590/S1519-566X200500...
, Gries et al. 2012GRIES, R., LOUZADA, J., ALMEIDA, S., MACEDO, R. & BARLOW, J. 2012. Evaluating the impacts and conservation value of exotic and native tree afforestation in Cerrado grasslands using dung beetles. Insect Conserv. Divers. 5:175-185, 10.1111/j.1752-4598.2011.00145.x.
https://doi.org/10.1111/j.1752-4598.2011...
, Milhomem et al. 2003MILHOMEM, M.S., MELLO, F.Z.V. & DINIZ, I.R. 2003. Técnicas de coleta de besouros copronecrófagos no Cerrado. Pesqui. Agropecuária Bras. 38:1249-1256, 10.1590/S0100-204X2003001100001.
https://doi.org/10.1590/S0100-204X200300...
, Silva & Audino 2011SILVA, P. & AUDINO, L.D. 2011. Escarabeíneos (Coleoptera: Scarabaeidae) atraídos a diferentes iscas em campo nativo de Bagé, Rio Grande do Sul, Brasil. Rev. Bras. Zoociências 13:241-247., Silva et al. 2010SILVA, F.A.B., COSTA, C.M.Q., MOURA, R.C. & FARIAS, A.I. 2010. Study of the dung beetle (Coleoptera: Scarabaeidae) community at two sites: Atlantic forest and clear-cut, Pernambuco, Brazil. Environ. Entomol. 39:359-367, 10.1603/EN09180.
https://doi.org/10.1603/EN09180...
, Vieira & Silva 2012VIEIRA, L. & SILVA, F.A.B. 2012. List Dung beetles (Coleoptera: Scarabaeidae: Scarabaeinae) of the Floresta Nacional Contendas do Sincorá, Bahia, Brazil. Check List 8:733-739.). Furthermore, 15 dung beetle species were reported from the intra-Amazonian savannahs near to Alter do Chão village, state of Pará, Brazil (Louzada et al. 2010LOUZADA, J., GARDNER, T., PERES, C. & BARLOW, J. 2010. A multi-taxa assessment of nestedness patterns across a multiple-use Amazonian forest landscape. Biol. Conserv. 143:1102-1109, 10.1016/j.biocon.2010.02.003.
https://doi.org/10.1016/j.biocon.2010.02...
, Matavelli & Louzada 2008MATAVELLI, R.A. & LOUZADA, J.N.C. 2008. Invasão de áreas de savana intra-amazônicas por Digitonthophagus gazella (Fabricius, 1787) (Insecta: Coleoptera: Scarabaeidae). Acta Amaz. 38:153-158, 10.1590/S0044-59672008000100017.
https://doi.org/10.1590/S0044-5967200800...
). Hence, considering the dung beetles' ecological importance in many ecosystem processes (Nichols et al. 2008NICHOLS, E., SPECTOR, S., LOUZADA, J., LARSEN, T., AMEZQUITA, S. & FAVILA, M.E. 2008. Ecological functions and ecosystem services provided by Scarabaeinae dung beetles. Biol. Conserv. 141:1461-1474, 10.1016/j.biocon.2008.04.011.
https://doi.org/10.1016/j.biocon.2008.04...
) and the poor knowledge about their fauna from the largest intra-Amazonian savannah area in Brazil, here we aimed to present the dung beetle species that were sampled within three lavrado savannahs of Roraima state, Brazil.

Materials and Methods

The city of Boa Vista in the state of Roraima has a humid tropical climate, characterized as a rainy summer, which extends from April to September, and a dry season from October to March. The average annual rainfall and temperature are respectively 1612 ± 400 mm and 27.4 °C (Barbosa & Fearnside 2005BARBOSA, R.I. & FEARNSIDE, P.M. 2005. Fire frequency and area burned in the Roraima savannas of Brazilian Amazonia. For. Ecol. Manage. 204:371-384, 10.1016/j.foreco.2004.09.011.
https://doi.org/10.1016/j.foreco.2004.09...
, Meneses et al. 2007MENESES, M.E.N.S., COSTA, M.L. & COSTA, J.A.V. 2007. Os lagos do lavrado de Boa Vista - Roraima?: fisiografia , físico-química das águas , mineralogia e química dos sedimentos. Rev. Bras. Geociências 37:478-489.). We used dung-baited pitfall traps to sample dung beetles during September 1996, July 1997, and November 2008 in three savannah areas near Boa Vista, Roraima, Brazil.

The first sampling period (1996) was carried out in a small savannah area in the municipality of Cantá (2°46' N 60° 38' W) located on the margin of the 'Rio Branco' river. The region is a large lavrado area continuous with small anthropogenic areas and subjected to floods during the rainy season. We installed five pitfall traps in this area, separated at least 50 m apart from each other, baited with human dung and exposed for 48 h to the dung beetle community. In the second sampling period (1997), a large area of lavrado in the 'Serra da Moça' region (north of Boa Vista, approx. 6°12' N 60°41' W) was sampled. We placed seven pitfall traps, located at least 50 m apart, baited with human dung and collected daily during five days with two exchanges of bait during the sampling period. In the third sampling period (2008), we sampled a large area of lavrado located on the opposite side of the Rio Branco river (02°62' N 60°72' W). We used seven pitfall traps, separated at least 50 m apart from each other, baited with human dung and left in the field for 48 h.

Pitfall traps were plastic containers (14 cm in diameter; 9 cm deep) buried in the ground with the opening at ground level, and with approximately 30 g of bait. We covered each pitfall with a lid to protect it from the rain and filled each one with 250 ml of a saline solution and detergent. After the exposition in the field, we collected, sorted, mounted and identified all dung beetles to the lowest possible taxonomic level. All voucher specimens were deposited in the Reference Collection of Neotropical Scarabaeinae in the Insect Ecology and Conservation Laboratory, Universidade Federal de Lavras and in the Entomology section of the Zoological Collection at Universidade Federal de Mato Grosso, Brazil. To further discussions about the geographical distribution and feeding habits of each genera and species here collected, we reviewed published and unpublished bibliographical references about the species ecology and distribution. Furthermore, we present an illustrated guide from each identified species we have found.

Results and Discussion

We collected 383 dung beetles belonging to ten species and eight genera (Table 1). From the collected species, only Canthidium sp. could not be identified and it is a possible new species. Ontherus appendiculatus, Canthidium aff. humerale, Dichotomius nisus, and Pseudocanthon aff. xanthurus were the most abundant species. As such, we have sampled six species without any previous occurrence from intra-Amazonian savannahs, as only the dung beetle species O. appendiculatus, D. nisus, P. aff. xanthurus and C. mutabilis have been previously recorded in these environments (Génier 1996GÉNIER, F. 1996. A revision of the Neotropical genus Ontherus Erichson (Coleoptera: Scarabaeidae, Scarabaeinae). Mem. Entomol. Soc. Canada 128:1-170, 10.4039/entm128170fv.
https://doi.org/10.4039/entm128170fv....
, Louzada et al. 2010, Matavelli & Louzada 2008MATAVELLI, R.A. & LOUZADA, J.N.C. 2008. Invasão de áreas de savana intra-amazônicas por Digitonthophagus gazella (Fabricius, 1787) (Insecta: Coleoptera: Scarabaeidae). Acta Amaz. 38:153-158, 10.1590/S0044-59672008000100017.
https://doi.org/10.1590/S0044-5967200800...
).

Table 1
Dung beetle species collected in three intra-Amazonian savannahs close to Boa Vista, Roraima, Brazil. In each sampling period, we sampled a different area.

Most species we found belong to species-clusters that have been collected within dry, open and highly disturbed environments on the American continent, some of them with records from Argentina (Damborsky et al. 2015DAMBORSKY, M.P., ALVAREZ BOHLE, M.C., IBARRA POLESEL, M.G., PORCEL, E.A. & FONTANA, J.L. 2015. Spatial and Temporal Variation of Dung Beetle Assemblages in a Fragmented Landscape at Eastern Humid Chaco. Neotrop. Entomol. 44:30-39, 10.1007/s13744-014-0257-2.
https://doi.org/10.1007/s13744-014-0257-...
), Mexico (Novelo et al. 2007NOVELO, E.R., DELFĺN-GONZÁLEZ, H. & MOR=N, M.Á. 2007. Copro-necrophagous beetle (Coleoptera: Scarabaeidae) diversity in an agroecosystem in Yucatan, Mexico. Rev. Biol. Trop. 55:83-99, 10.15517/rbt.v55i1.6059.
https://doi.org/10.15517/rbt.v55i1.6059...
), Ecuador (Carpio et al. 2009CARPIO, C., DONOSO, D.A., RAM=N, G. & DANGLES, O. 2009. Short term response of dung beetle communities to disturbance by road construction in the Ecuadorian Amazon. Ann. la Société Entomol. Fr. 45:455-469, 10.1080/00379271.2009.10697629.
https://doi.org/10.1080/00379271.2009.10...
), Venezuela (Lozano 2010LOZANO, C. 2010. Nuevos registros de escarabajos coprófagos para la Orchila, Sierra de Perijá, Venezuela. Boletín Cent. Investig. Biológicas 44:83-89.), Colombia (Noriega et al. 2007NORIEGA, J.A., SOLIS, C., ESCOBAR, F. & REALPE, E. 2007. Escarabajos coprófagos (Coleoptera: Scarabaeidae) de la provincia de la Sierra Nevada de Santa Marta. Biota Colomb. 8:77-86.) and Costa Rica (Padilla-Gil & Halffter 2007PADILLA-GIL, D.N. & HALFFTER, G. 2007. Biogeography of the areas and Canthonini (Coleoptera: Scarabaeidae) of dry tropical forests in mesoamerica and colombia. Acta Zoológica Mex. 23:73-108.). In Brazil, those have been registered within the semi-arid scrub forest (known as 'caatinga') (Hernandéz 2007HERNANDÉZ, M.I.M. 2007. Besouros Escarabeíneos (Coleoptera: Scarabaeidae) da Caatinga Paraibana, Brasil. Oecologia Bras. 11:356-364, 10.4257/oeco.2007.1103.06.
https://doi.org/10.4257/oeco.2007.1103.0...
, Lopes et al. 2006LOPES, P.P., LOUZADA, J.N.C. & VAZ-DE-MELLO, F.Z. 2006. Organization of dung beetle communities (Coleoptera: Scarabaeidae) in areas of vegetation re-establishment in Feira de Santana, Bahia, Brazil. Sitientibus Série Ciências Biológicas 6:261-266., Silva et al. 2007SILVA, F.A.B., HERNÁNDEZ, M.I.M., IDE, S. & MOURA, R.C. 2007. Comunidade de escarabeíneos (Coleoptera, Scarabaeidae) copronecrófagos da região de Brejo Novo, Caruaru, Pernambuco, Brasil. Rev. Bras. Entomol. 51:228-233, 10.1590/S0085-56262007000200014.
https://doi.org/10.1590/S0085-5626200700...
), coastal sandy vegetation ('restinga') (Costa et al. 2014), coastal low land forests ('Tabuleiro' forests) (Endres et al. 2007) and native and exotic pasturelands (Almeida et al. 2011, Costa et al. 2009, 2013, Silva et al. 2010SILVA, R.J., DINIZ, S. & VAZ-DE-MELLO, F.Z. 2010. Heterogeneidade do habitat, riqueza e estrutura da assembléia de besouros rola-bostas (Scarabaeidae: Scarabaeinae) em áreas de cerrado na Chapada dos Parecis, MT. Neotrop. Entomol. 39:934-940, 10.1590/S1519-566X2010000600014.
https://doi.org/10.1590/S1519-566X201000...
, Silva et al. 2011SILVA, P.G., VAZ-DE-MELLO, F.Z. & MARE, R.A. 2011. Guia de identificação das espécies de Scarabaeinae (Coleoptera: Scarabaeidae) do município de Santa Maria, Rio Grande do Sul, Brasil. Biota Neotrop. 11(4):329-345, 10.1590/S1676-06032011000400027.
https://doi.org/10.1590/S1676-0603201100...
, 2014). In addition, the species reported here were also found within highly impacted Brazilian forests, such as northeastern fragments of Atlantic forest (Salomão & Iannuzzi 2015SALOMÃO, R.P. & IANNUZZI, L. 2015. Dung beetle (Coleoptera, Scarabaeidae) assemblage of a highly fragmented landscape of Atlantic forest: from small to the largest fragments of northeastern Brazilian region. Rev. Bras. Entomol. 59:126-131, 10.1016/j.rbe.2015.03.008.
https://doi.org/10.1016/j.rbe.2015.03.00...
, Viegas et al. 2014VIEGAS, G., STENERT, C., SCHULZ, U.H. & MALTCHIK, L. 2014. Dung beetle communities as biological indicators of riparian forest widths in southern Brazil. Ecol. Indic. 36:703-710, 10.1016/j.ecolind.2013.09.036.
https://doi.org/10.1016/j.ecolind.2013.0...
), planted Eucalyptus sp. forests (Audino et al. 2011AUDINO, L.D., SILVA, P.G., NOGUEIRA, J.M., MORAES, L.P. & VAZ-DE-MELLO, F.Z. 2011. Scarabaeinae (Coleoptera, Scarabaeidae) de um bosque de eucalipto introduzido em uma região originalmente campestre. Iheringia. Série Zool. 101:121-126.) and Amazonian secondary forests, savannahs and agricultural areas (Korasaki et al. 2012KORASAKI, V., VAZ-DE-MELLO, F.Z., BRAGA, R.F., ZANETTI, R. & LOUZADA, J. 2012. Taxocenose de Scarabaeinae (Coleoptera: Scarabaeidae) em Benjamin Constant, AM. Acta Amaz. 42:423-432, 10.1590/S0044-59672012000300015.
https://doi.org/10.1590/S0044-5967201200...
, Louzada et al. 2010, Matavelli & Louzada 2008MATAVELLI, R.A. & LOUZADA, J.N.C. 2008. Invasão de áreas de savana intra-amazônicas por Digitonthophagus gazella (Fabricius, 1787) (Insecta: Coleoptera: Scarabaeidae). Acta Amaz. 38:153-158, 10.1590/S0044-59672008000100017.
https://doi.org/10.1590/S0044-5967200800...
).

The number of dung beetle species and individuals reported here was lower than in other intra-Amazonian savannahs (Louzada et al. 2010, Matavelli & Louzada 2008MATAVELLI, R.A. & LOUZADA, J.N.C. 2008. Invasão de áreas de savana intra-amazônicas por Digitonthophagus gazella (Fabricius, 1787) (Insecta: Coleoptera: Scarabaeidae). Acta Amaz. 38:153-158, 10.1590/S0044-59672008000100017.
https://doi.org/10.1590/S0044-5967200800...
), southern cerrado (Abot et al. 2012, Almeida et al. 2011, Durães et al. 2005, Gries et al. 2012, Silva et al. 2010, Vieira & Silva 2012VIEIRA, L. & SILVA, F.A.B. 2012. List Dung beetles (Coleoptera: Scarabaeidae: Scarabaeinae) of the Floresta Nacional Contendas do Sincorá, Bahia, Brazil. Check List 8:733-739.) and native grasslands of the 'Pampa' region (Silva et al. 2011). We believe this lower species-rich sample can be due the historical isolation of the intra-Amazonian from the cerrado savannahs in central Brazil during the Pleistocene period (Sanaiotti et al. 2002), besides some natural specific habitat characteristics, such as edaphic factors, vegetation structure and fire occurrence (Barbosa & Fearnside 2005BARBOSA, R.I. & FEARNSIDE, P.M. 2005. Fire frequency and area burned in the Roraima savannas of Brazilian Amazonia. For. Ecol. Manage. 204:371-384, 10.1016/j.foreco.2004.09.011.
https://doi.org/10.1016/j.foreco.2004.09...
, Louzada et al. 2010). Henceforward, we will discuss the knowledge concerning the distribution and ecology of each identified species.

Canthidium Erichson (1847):

Considered as one of the most diverse dung beetle groups (Halffter & Martínez 1966HALFFTER, G. & MARTĺNEZ, A. 1966. Revisión monográfica de los Canthonina americanos (Coleoptera, Scarabaeidae). Rev. Soc. Mex. Hist. Nat. 27:89-117.), this genus needs urgent revision, which may result in the creation of new genera. It currently includes about 170 described species, mainly recorded with dung-baited pitfalls in Neotropical forests and savannahs (Gill 1991GILL, B.D. 1991. Dung beetles in tropical American forests. In: Dung Beetle Ecology (Cambefort, I & Hanski, Y. eds.), Princeton University Press, Princeton, pp. 211-229., Silva et al. 2014SILVA, R.J., COLETTI, F., COSTA, D. & VAZ-DE-MELLO, F.Z. 2014. Rola-bostas (Coleoptera: Scarabaeidae: Scarabaeinae) de florestas e pastagens no sudoeste da Amazônia brasileira: Levantamento de espécies e guildas alimentares. Acta Amaz. 44:345-352, 10.1590/1809-4392201304472.
https://doi.org/10.1590/1809-43922013044...
, Vaz-de-Mello & Louzada 1997VAZ-DE-MELLO, F.Z. & LOUZADA, J.N.C. 1997. Considerações sobre o forrageio arbóreo por Scarabaeidae (Coleoptera, Scarabaeoidea) e dados sobre sua ocorrência em floresta tropical do Brasil. Acta Zool. Mex. Nueva Ser. 72:55-61. VAZ-DE-MELLO, F.Z., LOUZADA, J.N.C. & SCHOEREDER, J.H. 1998. New data and comments on Scarabaeidae (Coleoptera: Scarabaeoidea) associated with Atini (Hymenoptera: Formicidae). Coleopt. Bull. 52:209-216. VAZ-DE-MELLO, F.Z. 1999a. Scarabaeidae do Estado de Roraima. Memórias da IV Reunião Latino-Americana de Scarabaeoidologia, Viçosa, MG, p. 78-79., Vaz-de-Mello et al. 1998). There are also some records of species sampled with carrion-baited pitfalls or consuming rooting fruit, fungi and dead insects (Medri & Lopes 2001MEDRI, ĺ.M. & LOPES, J. 2001. Scarabaeidae (Coleoptera) do Parque Estadual Mata dos Godoy e de área de pastagem, no norte do Paraná, Brasil. Rev. Bras. Zool. 18:135-141, 10.1590/S0101-81752001000500011.
https://doi.org/10.1590/S0101-8175200100...
, Silva & Audino 2011SILVA, P. & AUDINO, L.D. 2011. Escarabeíneos (Coleoptera: Scarabaeidae) atraídos a diferentes iscas em campo nativo de Bagé, Rio Grande do Sul, Brasil. Rev. Bras. Zoociências 13:241-247., Silva et al. 2014). Hence, species have been suggested as myrmecophilous associated with nests of Atta ants Fabricius, 1804FABRICIUS, J.C. 1804. Systema Piezatorum secundum ordines, genera, species, adjectis, synonymis, locis, observationnibus, descriptionibus. (Reichard, C). Brunswick XIV, pp. 15-439., probably exploiting rotting fungi (Falqueto et al. 2005FALQUETO, S.A., VAZ-DE-MELLO, F.Z. & SCHOEREDER, J.H. 2005. Are fungivorous Scarabaeidae less specialist? Ecol. Austral 15:17-22, S1667-782X2005000100003.). Canthidium aff. humerale (Figure 1A) belongs to a species-cluster that contains species considered as coprophagous (Silva et al. 2007) and distributed within many opened environments. Species in this cluster have been recorded in northeastern Brazilian grassland areas (Almeida et al. 2011, Costa et al. 2009, Silva et al. 2007), Tabuleiro forests (Endres et al. 2007ENDRES, A.A., CREÃO-DUARTE, A.J. & HERNÁNDEZ, M.I.M. 2007. Diversidade de Scarabaeidae s. str. (Coleoptera) da Reserva Biológica Guaribas, Mamanguape, Paraíba, Brasil: uma comparação entre Mata Atlântica e Tabuleiro Nordestino. Rev. Bras. Entomol. 51:67-71, 10.1590/S0085-56262007000100012.
https://doi.org/10.1590/S0085-5626200700...
), secondary regeneration zones between caatinga and cerrado (Lopes et al. 2006) and transitional regions between the Amazon forest and cerrado (Andrade et al. 2011ANDRADE, R.B., BARLOW, J., LOUZADA, J., VAZ-DE-MELLO, F.Z., SOUZA, M., SILVEIRA, J.M. & COCHRANE, M.A. 2011. Quantifying Responses of Dung Beetles to Fire Disturbance in Tropical Forests: The Importance of Trapping Method and Seasonality. PLoS One 6:e26208, 10.1371/journal.pone.0026208.
https://doi.org/10.1371/journal.pone.002...
). The other Canthidium species we found could not be identified accurately, thus no further comments are possible about its ecology and distribution.

Figure 1
Figure 1. Dung beetle species sampled from intra-Amazonian savannahs in the state of Roraima: (A) Canthidium aff. humerale; (B) Canthon aff. scrutator; (C) Canthon aff. mutabilis; (D) Malagoniella astyanax (extracted from Vieira & Silva, 2012VIEIRA, L. & SILVA, F.A.B. 2012. List Dung beetles (Coleoptera: Scarabaeidae: Scarabaeinae) of the Floresta Nacional Contendas do Sincorá, Bahia, Brazil. Check List 8:733-739.); (E) Pseudocanthon aff. xanthurus; (F) Dichotomius nisus; (G) Ontherus appendiculatus (extracted from Vidaurre et al., 2008VIDAURRE, T., GONZALES, L. & LEDEZA, J.M. 2008. Escarabajos coprófagos (Scarabaeidae: Scarabaeinae) del Palmar de las Islas, Santa Cruz, Bolívia. Kempffiana 4:3-20.); (H) Onthophagus aff. hirculus; (I) Coprophanaeus gamezi (extracted from Pacheco & Vaz-de-Mello, 2015PACHECO, T.L. & VAZ-DE-MELLO, F.Z. 2015. Dung beetles of the tribe Phanaeini (Coleoptera: Scarabaeidade: Scarabaeinae) from Roraima state, Northern Brazil: checklist and key to species. Biota Neotrop. 15(2): 1-9, 10.1590/1676-06032015014514.
https://doi.org/10.1590/1676-06032015014...
).

Canthon Hoffmannsegg (1817):

Featuring approximately 200 species, this genus has been studied at a supraspecific level, except for some species that are not yet grouped into any subgenera. It is an American group, with species mainly found from Argentina to Canada (Vaz-de-Mello 2000VAZ-DE-MELLO, F.Z. 2000. Estado Atual de conhecimentos dos Scarabaeidae s. str. (Coleoptera: Scarabaeoidea) do Brasil. In: Hacia Un Proyecto CYTED Para El Inventario Y Estimación de La Diversidad Entonológica Em Iberoamérica: PrIBES-2000 (Martín-Piera, F. Morrone, J.J. & Melic, A. eds.). Sociedad Entomologica Aragonesa & CYTED, Zaragoza, pp. 181-195.). Although most species are considered as copro-necrophagous, other feeding habits, including fungi and debris feeders (Vaz-de-Mello 1999bVAZ-DE-MELLO, F.Z. 1999b. Scarabaeidae s. str. (Coleoptera: Scarabaeoidea) de um fragmento de Floresta Amazônica no estado do Acre, Brasil. 1. Taxocenose. An. da Soc. Entomológica do Bras. 28:439-446, 10.1590/S0301-80591999000300009.
https://doi.org/10.1590/S0301-8059199900...
) and ant predation (Hertel & Colli 1998HERTEL, F. & COLLI, G.R. 1998. The use of leaf-cutter Ants, Atta laevigata (Smith) (Hymenoptera: Formicidae), as a substrate for oviposition by the dung beetle Canthon virens Mannerheim (Coleoptera: Scarabaeidae) in central Brazil. Coleopt. Bull. 52:105-108., Vaz-de-Mello et al. 1998, Villalobos et al. 1998VILLALOBOS, F.J., DIAZ, A. & FAVILA, M.E. 1998. Two species of Canthon feed on dead and live invertebrates. Coleopt. Bull. 52:101-104.) were also recorded. The subgenus Canthon (Canthon) Hoffmannsegg, 1817 includes mainly coprophagous or necrophagous dung beetles, despite species preying on ants (Cantil et al. 2014CANTIL, L.F., SÁNCHEZ, M.V. & GENISE, J.F. 2014. The Nest and Brood Ball of Canthon (Canthon) virens aff. paraguayanus Balthasar (Coleoptera: Scarabaeidae: Scarabaeinae). Coleopt. Bull. 68:384-386, 10.1649/072.068.0306.
https://doi.org/10.1649/072.068.0306...
, Forti et al. 2012FORTI, L.C., RINALDI, I.M.P., CAMARGO, R., DA S. & FUJIHARA, R.T. 2012. Predatory Behavior of Canthon virens (Coleoptera: Scarabaeidae): A Predator of Leafcutter Ants. Psyche. 2012:1-5, 10.1155/2012/921465.
https://doi.org/10.1155/2012/921465...
, Villalobos et al. 1998) and feeding on dead insects and millipedes (Villalobos et al. 1998). Canthon (Canthon) aff. scrutator Balthasar, 1939 (Figure 1B) belongs to the Canthon virens species-group (Halffter & Martínez 1977HALFFTER, G. & MARTĺNEZ, A. 1977: Revision monografi ca de los Canthonina Americanos, IV parte. Clava parageneros y subgeneros. Folia Entomológica Mex. 38:29-107), which is distributed throughout the northern Amazon Basin, Guyana and Atlantic Forest region. This species-group comprises copro-necrophagous dung beetles, also recorded consuming insect carcasses (Forti et al. 2012) or sampled with carrion-baited pitfall traps (Salomão & Iannuzzi 2015SALOMÃO, R.P. & IANNUZZI, L. 2015. Dung beetle (Coleoptera, Scarabaeidae) assemblage of a highly fragmented landscape of Atlantic forest: from small to the largest fragments of northeastern Brazilian region. Rev. Bras. Entomol. 59:126-131, 10.1016/j.rbe.2015.03.008.
https://doi.org/10.1016/j.rbe.2015.03.00...
). We believe the species Canthon (Canthon) aff. mutabilis Lucas, 1859 (Figure 1C) needs urgent revision, given that individuals identified as C. mutabilis may comprise a complex group of different species. This species-group have been recorded within open environments in America, such as Colombia (Noriega et al. 2007), Costa Rica (Padilla-Gil & Halffter 2007PADILLA-GIL, D.N. & HALFFTER, G. 2007. Biogeography of the areas and Canthonini (Coleoptera: Scarabaeidae) of dry tropical forests in mesoamerica and colombia. Acta Zoológica Mex. 23:73-108.), Argentina (Boito et al, 2009BOITO, G.T., GIUGGIA, J.A., ORNAGHI, J.A., GERARDO, U.A. & GIOVANINI, D. 2009. Uso de trampas “Barber” para determinar la diversidad de coleópteros epígeos asociados al cultivo de maní (Arachis hypogaea L.). FCA UNCuyo XLI N°1:23-31.) and primary forests from Suriname (Larsen 2011LARSEN, T.H. 2011. Dung beetles of the Kwamalasamutu region, Suriname (Coleoptera: Scarabaeidae: Scarabaeinae). In: . eds.). A Rapid Biological Assessment of the Kwamalasamutu Region, Southwestern Suriname. RAP Bulletin of Biological Assessment 63. Conservation International, Arlington, pp. 91-103.). In Brazil, there are samples from flight intercept traps (Audino et al. 2011, Costa et al. 2009) and pitfalls baited with dung and carrion (Costa et al. 2013, 2014, Silva et al. 2007, 2014, Silva & Audino 2011SILVA, P. & AUDINO, L.D. 2011. Escarabeíneos (Coleoptera: Scarabaeidae) atraídos a diferentes iscas em campo nativo de Bagé, Rio Grande do Sul, Brasil. Rev. Bras. Zoociências 13:241-247.).

Malagoniella Martínez, 1961:

Halffter & Martínez (1966)HALFFTER, G. & MARTĺNEZ, A. 1966. Revisión monográfica de los Canthonina americanos (Coleoptera, Scarabaeidae). Rev. Soc. Mex. Hist. Nat. 27:89-117. reviewed this genus, which includes nine colorful metallic dung beetle species distributed across the Neotropical region, from USA (Texas) to Argentina (Padilla-Gil & Halffter 2007PADILLA-GIL, D.N. & HALFFTER, G. 2007. Biogeography of the areas and Canthonini (Coleoptera: Scarabaeidae) of dry tropical forests in mesoamerica and colombia. Acta Zoológica Mex. 23:73-108.). Malagoniella astyanax (Oliver 1789) (Figure 1D) belongs to a species complex that is currently being revised. It has been found in the Brazilian caatinga (Hernandéz, 2007HERNANDÉZ, M.I.M. 2007. Besouros Escarabeíneos (Coleoptera: Scarabaeidae) da Caatinga Paraibana, Brasil. Oecologia Bras. 11:356-364, 10.4257/oeco.2007.1103.06.
https://doi.org/10.4257/oeco.2007.1103.0...
, Silva et al. 2007, Vieira & Silva 2012VIEIRA, L. & SILVA, F.A.B. 2012. List Dung beetles (Coleoptera: Scarabaeidae: Scarabaeinae) of the Floresta Nacional Contendas do Sincorá, Bahia, Brazil. Check List 8:733-739.), Ecuadorian Amazon (Carpio et al. 2009), Colombia (Escobar 1997ESCOBAR, F. 1997. Estudio de la comunidad de coleópteros coprófagos (Scarabeidae) en un remanente de Bosque Seco al Norte del Tolima, Colombia. Caldasia 19:419-430.), Costa Rica (Padilla-Gil & Halffter 2007PADILLA-GIL, D.N. & HALFFTER, G. 2007. Biogeography of the areas and Canthonini (Coleoptera: Scarabaeidae) of dry tropical forests in mesoamerica and colombia. Acta Zoológica Mex. 23:73-108.), Mexico (Novelo et al. 2007), Venezuela (Ferrer-Paris et al. 2013FERRER-PARIS, J.R., SÁNCHEZ-MERCADO, A. & RODRĺGUEZ, J.P. 2013. Optimización del muestreo de invertebrados tropicales: Un ejemplo con escarabajos coprófagos (Coleoptera: Scarabaeinae) en Venezuela. Rev. Biol. Trop. 61:89-110, 10.15517/rbt.v61i1.10941.
https://doi.org/10.15517/rbt.v61i1.10941...
), Bolivia (Vidaurre et al. 2008) and Guyana (F.Z. Vaz-de-Mello, unpublished data). Species in this group have been considered as nocturnal (Hernandéz 2007HERNANDÉZ, M.I.M. 2007. Besouros Escarabeíneos (Coleoptera: Scarabaeidae) da Caatinga Paraibana, Brasil. Oecologia Bras. 11:356-364, 10.4257/oeco.2007.1103.06.
https://doi.org/10.4257/oeco.2007.1103.0...
), sampled by pitfall traps baited with dung and carrion (Carpio et al. 2009, Hernandéz 2007HERNANDÉZ, M.I.M. 2007. Besouros Escarabeíneos (Coleoptera: Scarabaeidae) da Caatinga Paraibana, Brasil. Oecologia Bras. 11:356-364, 10.4257/oeco.2007.1103.06.
https://doi.org/10.4257/oeco.2007.1103.0...
, Novelo et al. 2007, Silva et al. 2007).

Pseudocanthon Bates, 1887:

This genus is small, including nine species distributed from the USA to Argentina and Antilles (Halffter & Matthews 1966HALFFTER, G. & MATTHEWS, E.G. 1966. The natural history of dung beetles of the subfamily Scarabaeinae (Coleoptera: Scarabaeidae). Folia Entomológica Mex. 12:1-312., Padilla-Gil & Halffter 2007PADILLA-GIL, D.N. & HALFFTER, G. 2007. Biogeography of the areas and Canthonini (Coleoptera: Scarabaeidae) of dry tropical forests in mesoamerica and colombia. Acta Zoológica Mex. 23:73-108.). Although small, it urgently needs taxonomic revision, at least for the continental species. The species Pseudocanthon aff. xanthurus (Figure 1E) is a complex of species, which is difficult to identify (Korasaki et al. 2012) and, as the genus, it needs taxonomic revision. Although extensively sampled with dung baited pitfalls, the species have been considered as generalist (Andrade et al. 2011), with records from open habitats in Colombia and Venezuela (Ferrer-Paris et al. 2013). In Brazil they have been reported within pastures and grasslands (Costa et al. 2009, Schiffler et al. 2015, Silva et al. 2014), caatinga (Lopes et al. 2006), degraded restinga (Costa et al. 2014), northeastern Atlantic forest (Costa et al. 2013, Salomão & Iannuzzi 2015SALOMÃO, R.P. & IANNUZZI, L. 2015. Dung beetle (Coleoptera, Scarabaeidae) assemblage of a highly fragmented landscape of Atlantic forest: from small to the largest fragments of northeastern Brazilian region. Rev. Bras. Entomol. 59:126-131, 10.1016/j.rbe.2015.03.008.
https://doi.org/10.1016/j.rbe.2015.03.00...
), intra-Amazonian savannahs (Matavelli & Louzada 2008MATAVELLI, R.A. & LOUZADA, J.N.C. 2008. Invasão de áreas de savana intra-amazônicas por Digitonthophagus gazella (Fabricius, 1787) (Insecta: Coleoptera: Scarabaeidae). Acta Amaz. 38:153-158, 10.1590/S0044-59672008000100017.
https://doi.org/10.1590/S0044-5967200800...
) and Amazonian agricultural areas (Korasaki et al. 2012).

Dichotomius Hope (1838):

According to the last taxonomic revision, this genus features approximately 165 valid species widely distributed from the USA to Argentina (Luederwaldt 1929LUEDERWALDT, H. 1929. As espécies brasileiras do gênero Pinotus. (Coleoptera - Lamellicornidae - Coprini), com algumas considerações também sobre outras espécies. Rev. do Mus. Paul. 16, 603-775.), especially in Neotropical forest and savannah areas (Vaz-de-Mello 1999bVAZ-DE-MELLO, F.Z. 1999b. Scarabaeidae s. str. (Coleoptera: Scarabaeoidea) de um fragmento de Floresta Amazônica no estado do Acre, Brasil. 1. Taxocenose. An. da Soc. Entomológica do Bras. 28:439-446, 10.1590/S0301-80591999000300009.
https://doi.org/10.1590/S0301-8059199900...
, 2000VAZ-DE-MELLO, F.Z. 2000. Estado Atual de conhecimentos dos Scarabaeidae s. str. (Coleoptera: Scarabaeoidea) do Brasil. In: Hacia Un Proyecto CYTED Para El Inventario Y Estimación de La Diversidad Entonológica Em Iberoamérica: PrIBES-2000 (Martín-Piera, F. Morrone, J.J. & Melic, A. eds.). Sociedad Entomologica Aragonesa & CYTED, Zaragoza, pp. 181-195.). Although most species are coprophagous, we believe there are some exceptions. Dichotomius nisus (Olivier 1789) (Figure 1F) is considered a nocturnal coprophagous-generalist species (Hernandéz 2007HERNANDÉZ, M.I.M. 2007. Besouros Escarabeíneos (Coleoptera: Scarabaeidae) da Caatinga Paraibana, Brasil. Oecologia Bras. 11:356-364, 10.4257/oeco.2007.1103.06.
https://doi.org/10.4257/oeco.2007.1103.0...
, Silva et al. 2007, Vieira et al. 2008VIEIRA, L., LOUZADA, J.N.C. & SPECTOR, S. 2008. Effects of degradation and replacement of Southern Brazilian Coastal Sandy Vegetation on the dung beetles (Coleoptera: Scarabaeidae). Biotropica 40:719-727, 10.1111/j.1744-7429.2008.00432.x.
https://doi.org/10.1111/j.1744-7429.2008...
), commonly occurring within opened and/or disturbed environments (Filgueiras et al. 2015FILGUEIRAS, B.K.C., TABARELLI, M., LEAL, I.R., VAZ-DE-MELLO, F.Z. & IANNUZZI, L. 2015. Dung beetle persistence in human-modified landscapes: Combining indicator species with anthropogenic land use and fragmentation-related effects. Ecol. Indic. 55:65-73, 10.1016/j.ecolind.2015.02.032.
https://doi.org/10.1016/j.ecolind.2015.0...
, Nunes et al. 2012NUNES, R.V., FRIZZAS, M.R. & VAZ-DE-MELLO, F.Z. 2012. Scarabaeinae (Coleoptera: Scarabaeidae) of a rupestrian field at Caatinga, Distrito Federal, Brazil: commented list of species. Biota Neotrop. 12(4):125-129, 10.1590/S1676-06032012000400013.
https://doi.org/10.1590/S1676-0603201200...
). It is widely distributed, with records from Guyana (Boilly & Vaz-de-Mello 2014BOILLY, O. & VAZ-DE-MELLO, F.Z. 2014. Les Scarabaeinae de Guyane: clé illustrée des genres. ACOREP-France Coléoptàres Guyane. Tome VII 103-112.), Argentinian Chaco (Damborsky et al. 2015), transitional forests in Bolivia (Vidaurre et al. 2008), Colombia Andes (Putumayo region) (Escobar et al. 2005ESCOBAR, F., LOBO, J.M. & HALFFTER, G. 2005. Altitudinal variation of dung beetle (Scarabaeidae: Scarabaeinae) assemblages in the Colombian Andes. Glob. Ecol. Biogeogr. 14:327-337, 10.1111/j.1466-822X.2005.00161.x.
https://doi.org/10.1111/j.1466-822X.2005...
) and Venezuelan dry forests and pastures (Ferrer-Paris et al. 2013, Lozano 2010LOZANO, C. 2010. Nuevos registros de escarabajos coprófagos para la Orchila, Sierra de Perijá, Venezuela. Boletín Cent. Investig. Biológicas 44:83-89.). In Brazil, it has been sampled with both dung and carrion-baited pitfall traps within more than 17 states (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. Zoociências 9:199-203.). Accordingly, it has been extensively collected from the northern intra-Amazonian savannahs (Koller et al. 1999, Matavelli & Louzada 2008MATAVELLI, R.A. & LOUZADA, J.N.C. 2008. Invasão de áreas de savana intra-amazônicas por Digitonthophagus gazella (Fabricius, 1787) (Insecta: Coleoptera: Scarabaeidae). Acta Amaz. 38:153-158, 10.1590/S0044-59672008000100017.
https://doi.org/10.1590/S0044-5967200800...
) to southern Campanha region (Silva et al. 2008SILVA, P.G., ROSA-GARCIA, A.M. & VIDAL, M.B. 2008. Espécies de besouros copro-necrófagos (Coleoptera: Scarabaeidae) da Região da Campanha do Rio Grande do Sul. Rev. Ciências Agroveterinárias 7:143-149.), within non-native forests of Eucalyptus sp. (Audino et al. 2011). Furthermore, there are records from restinga (Costa et al. 2014, Schiffler et al. 2015, Vieira et al. 2008), Tabuleiro forest (Endres et al. 2007), caatinga (Hernandéz 2007HERNANDÉZ, M.I.M. 2007. Besouros Escarabeíneos (Coleoptera: Scarabaeidae) da Caatinga Paraibana, Brasil. Oecologia Bras. 11:356-364, 10.4257/oeco.2007.1103.06.
https://doi.org/10.4257/oeco.2007.1103.0...
, Medina & Lopes 2014MEDINA, A.M. & LOPES, P.P. 2014. Resource utilization and temporal segregation of Scarabaeinae (Coleoptera: Scarabaeidae) community in a caatinga fragment. Neotrop. Entomol. 43:127-133, 10.1007/s13744-014-0198-9.
https://doi.org/10.1007/s13744-014-0198-...
, Santos et al. 2014SANTOS, W., ALVES, A. & CREÃO-DUARTE, A. 2014. Beetles (Insecta, Coleoptera) associated with pig carcasses exposed in a Caatinga area, Northeastern Brazil. Brazilian J. Biol. 74:649-655, 10.1590/bjb.2014.0072.
https://doi.org/10.1590/bjb.2014.0072...
, Vieira & Silva 2012VIEIRA, L. & SILVA, F.A.B. 2012. List Dung beetles (Coleoptera: Scarabaeidae: Scarabaeinae) of the Floresta Nacional Contendas do Sincorá, Bahia, Brazil. Check List 8:733-739.), Atlantic forest fragments (Costa et al. 2013, Flechtmann et al. 2009FLECHTMANN, C.A.H., TABET, V.G. & QUINTERO, I. 2009. Influence of carrion smell and rebating time on the efficiency of pitfall traps to dung beetle sampling. Entomol. Exp. Appl. 132:211-217, 10.1111/j.1570-7458.2009.00885.x.
https://doi.org/10.1111/j.1570-7458.2009...
, Salomão & Iannuzzi 2015SALOMÃO, R.P. & IANNUZZI, L. 2015. Dung beetle (Coleoptera, Scarabaeidae) assemblage of a highly fragmented landscape of Atlantic forest: from small to the largest fragments of northeastern Brazilian region. Rev. Bras. Entomol. 59:126-131, 10.1016/j.rbe.2015.03.008.
https://doi.org/10.1016/j.rbe.2015.03.00...
, Silva et al. 2015SILVA, R.J., RIBEIRO, H.V., SOUZA, M.F. & VAZ-DE-MELLO, F.Z. 2015. Influence of soil granulometry on the structure of functional guilds of dung beetles (Coleoptera: Scarabaeidae: Scarabaeinae) at semideciduos forests in the State of Mato Grosso, Brazil. Biosci. J. 31:601-612, 10.14393/BJ-v31n1a2015-23525.
https://doi.org/10.14393/BJ-v31n1a2015-2...
, Viegas et al. 2014), cerrado (Almeida et al. 2011, 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(2):123-127, 10.1590/S1676-06032010000200015.
https://doi.org/10.1590/S1676-0603201000...
) and many pasturelands and altered grasslands (Costa et al. 2009, Filgueiras et al. 2015, Louzada & Silva 2009LOUZADA, J.N.C. & SILVA, P.R.C.E. 2009. Utilisation of introduced Brazilian pastures ecosystems by native dung beetles: Diversity patterns and resource use. Insect Conserv. Diver. 2:45-52, 10.1111/j.1752-4598.2008.00038.x.
https://doi.org/10.1111/j.1752-4598.2008...
, Louzada et al. 2007, Puker et al. 2013PUKER, A., CORREA, C.M.A., KORASAKI, V., FERREIRA, K.R. & OLIVEIRA, N.G. 2013. Dung Beetles (Coleoptera: Scarabaeidae) Attracted to Dung of the Largest Herbivorous Rodent on Earth: A Comparison with Human Feces. Environ. Entomol. 42:1218-1225, 10.1603/EN13100.
https://doi.org/10.1603/EN13100...
, Scheffler 2005SCHEFFLER, P.Y. 2005. Dung beetle (Coleoptera: Scarabaeidae) diversity and community structure across three disturbance regimes in eastern Amazonia. J. Trop. Ecol. 21:9-19, 10.1017/S0266467404001683.
https://doi.org/10.1017/S026646740400168...
SCHIFFLER, G., VAZ-DE-MELLO, F.Z. & AZEVEDO, C.O. 2003. Scarabaeidae s. str. (Coleoptera) from the Doce river and Suruaca valleu in Linhares municipality, Espírito Santo State, Brazil. Rev. Bras. Zoociências 5:205-211., Silva et al. 2011).

Ontherus Erichson, 1847:

This genus is distributed from Mexico to Argentina and was recently reviewed by Génier (1996)GÉNIER, F. 1996. A revision of the Neotropical genus Ontherus Erichson (Coleoptera: Scarabaeidae, Scarabaeinae). Mem. Entomol. Soc. Canada 128:1-170, 10.4039/entm128170fv.
https://doi.org/10.4039/entm128170fv....
, which resulted in 56 species divided into three subgenera. We believe that most of the species are coprophagous; however, there are some species associated with sediment deposits within ant colonies of AttaFabricius, 1804FABRICIUS, J.C. 1804. Systema Piezatorum secundum ordines, genera, species, adjectis, synonymis, locis, observationnibus, descriptionibus. (Reichard, C). Brunswick XIV, pp. 15-439. and Acromyrmex Mayr, 1865 (Halffter & Halffter 2009HALFFTER, G. & HALFFTER, V. 2009. Why and where coprophagous beetles (Coleoptera: Scarabaeinae) eat sedes, fruits or vegetable detritus. Bol. Soc. Entomol. Aragonesa. 45:1-22.). Ontherus appendiculatus (Mannerhein, 1829) (Figure 1G), although considered as typically coprophagous, has been collected also with carrion (Flechtmann et al. 2009, Rosa et al. 2011ROSA, T.A., BABATA, M.L.Y., SOUZA, C.M., SOUSA, D., MELLO-PATIU, C.A., VAZ-DE-MELLO, F.Z. & MENDES, J. 2011. Arthropods associated with pig carrion in two vegetation profiles of Cerrado in the State of Minas Gerais, Brazil. Rev. Bras. Entomol. 55:424-434, 10.1590/S0085-56262011005000045
https://doi.org/10.1590/S0085-5626201100...
, Silva et al. 2007). Widely distributed in South America (Génier 1996GÉNIER, F. 1996. A revision of the Neotropical genus Ontherus Erichson (Coleoptera: Scarabaeidae, Scarabaeinae). Mem. Entomol. Soc. Canada 128:1-170, 10.4039/entm128170fv.
https://doi.org/10.4039/entm128170fv....
), O. appendiculatus has been reported in Argentina, Bolivia, Brazil, Colombia, Ecuador, French Guyana, Guyana, Paraguay, Uruguay and Venezuela (Génier 1996GÉNIER, F. 1996. A revision of the Neotropical genus Ontherus Erichson (Coleoptera: Scarabaeidae, Scarabaeinae). Mem. Entomol. Soc. Canada 128:1-170, 10.4039/entm128170fv.
https://doi.org/10.4039/entm128170fv....
, Louzada et al. 2007, Vidaurre et al. 2008, Ferrer-Paris et al. 2013). In Brazil, it has a broad distribution, with records from intra-Amazonian savannahs (Matavelli & Louzada 2008MATAVELLI, R.A. & LOUZADA, J.N.C. 2008. Invasão de áreas de savana intra-amazônicas por Digitonthophagus gazella (Fabricius, 1787) (Insecta: Coleoptera: Scarabaeidae). Acta Amaz. 38:153-158, 10.1590/S0044-59672008000100017.
https://doi.org/10.1590/S0044-5967200800...
), Tabuleiro forest (Lima et al. 2013LIMA, M.G.A., SILVA, R.P., DE, A., SOUSA, M.D.F. & COSTA, E.M. 2013. Diversidade de Scarabaeinae (Coleoptera: Scarabaeidae) no Parque Botânico do Ceará, Caucaia - CE, Brasil. Rev. Ago@mbiente On-line 7:89-94, 10.18227/1982-8470ragro.v7i1.970.
https://doi.org/10.18227/1982-8470ragro....
) and Atlantic forest fragments (Silva et al. 2015). Furthermore, it has been recorded in transitional forests between cerrado and Amazon forest (Andrade et al. 2011), and cerrado within Pantanal regions (Louzada et al. 2007, Puker et al. 2013, Rodrigues et al. 2010).

Onthophagus Latreille, 1802:

It is a very diverse and cosmopolitan genus with more than 2000 species (Tarasov & Kabakov 2010TARASOV, S.I. & KABAKOV, O.N. 2010. Two new species of Onthophagus (Coleoptera: Scarabaeidae) from Indochina, with a discussion of some problems with the classification of Serrophorus and similar subgenera. Zootaxa 2344:17-28.). Although considered as coprophagous, there are species attracted by the defense secretions from millipedes (Brühl & Krell 2003BRÜHL, C. & KRELL, F.T. 2003. Finding a rare resource: Bornean Scarabaeoidea (Coleoptera) attracted by defensive secretions of diplopoda. Coleopt. Bull. 57:51-55, 10.1649/0010-065X(2003)057[0051:FARRBS]2.0.CO;2.
https://doi.org/10.1649/0010-065X(2003)0...
, Schmitt et al. 2004SCHMITT, T., KRELL, F.T. & LINSENMAIR, K.E. 2004. Quinone mixture as attractant for necrophagous dung beetles specialized on dead millipedes. J. Chem. Ecol. 30:731-740, 10.1023/B:JOEC.0000028428.53797.cb.
https://doi.org/10.1023/B:JOEC.000002842...
) or associated with caves (Slay et al. 2012SLAY, M.E., SKELLEY, P.E. & TAYLOR, S.J. 2012. Howden and Cartwright (Coleoptera: Scarabaeidae) from Ozark Caves, with a Review of Scarabaeoids Reported from North American Caves. Coleopt. Bull. 66:187-199, 10.1649/072.066.0302.
https://doi.org/10.1649/072.066.0302...
), bird nests (Krištofík et al. 2003KRIŠTOFĺK, J., MAŠÁN, P., ŠUSTEK, Z. & KLOUBEC, B. 2003. Arthropods (Pseudoscorpionida, Acari, Coleoptera, Siphonaptera) in nests of the tengmalm's owl, Aegolius funereus. Biol. Bratislava. 58:231-240.) and rodents (Sánchez-Huerta et al. 2015SÁNCHEZ-HUERTA, J.L., TONELLI, M., ZUNINO, M. & HALFFTER, G. 2015. Redescription of Onthophagus halffteri Zunino (Coleoptera: Scarabaeidae: Scarabaeinae), with Ecological and Distributional Notes. Coleopt. Bull. 69:225-230, 10.1649/0010-065X-69.2.225.
https://doi.org/10.1649/0010-065X-69.2.2...
). Onthophagus aff. hirculus Mannerheim, 1829 (Figure 1H) belongs to a complex of species that is taxonomically close to O. hirculus and needs urgent taxonomic revision. Those species have been considered as coprophagous and generalist (Hernandéz 2007HERNANDÉZ, M.I.M. 2007. Besouros Escarabeíneos (Coleoptera: Scarabaeidae) da Caatinga Paraibana, Brasil. Oecologia Bras. 11:356-364, 10.4257/oeco.2007.1103.06.
https://doi.org/10.4257/oeco.2007.1103.0...
, Silva et al. 2011), collected by pitfall traps baited with different types of dung, rotting fruits and carcass (Audino et al. 2011, Flechtmann et al. 2009, Puker et al. 2013, Rosa et al. 2011, Silva & Audino 2011SILVA, P. & AUDINO, L.D. 2011. Escarabeíneos (Coleoptera: Scarabaeidae) atraídos a diferentes iscas em campo nativo de Bagé, Rio Grande do Sul, Brasil. Rev. Bras. Zoociências 13:241-247., Silva et al. 2008). This species-complex have been recorded in South American open areas at altitudes <1000 m, such as Argentinian Chaco (Damborsky et al. 2015), Colombian secondary dry forests (Noriega et al. 2007) and Bolivian transitional forests (Vidaurre et al. 2008). In Brazil, they have been collected within pasturelands (Puker et al. 2013, Silva et al. 2010, Silva et al. 2014), caatinga (Hernandéz 2007HERNANDÉZ, M.I.M. 2007. Besouros Escarabeíneos (Coleoptera: Scarabaeidae) da Caatinga Paraibana, Brasil. Oecologia Bras. 11:356-364, 10.4257/oeco.2007.1103.06.
https://doi.org/10.4257/oeco.2007.1103.0...
, Santos et al. 2014, Silva et al. 2007, Vieira & Silva 2012VIEIRA, L. & SILVA, F.A.B. 2012. List Dung beetles (Coleoptera: Scarabaeidae: Scarabaeinae) of the Floresta Nacional Contendas do Sincorá, Bahia, Brazil. Check List 8:733-739.) and restinga areas (Costa et al. 2014, Vieira et al. 2008). In addition, they have been recorded in Atlantic forest fragments (Silva et al. 2015) and transitional areas between the Amazon forest and cerrado (Andrade et al. 2011).

Coprophanaeus d'Olsoufieff, 1924:

Reviewed by Edmonds & Zidek (2010)EDMONDS, W.D. & ZIDEK, J. 2010. Taxonomic review of the Neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta mundi Paper 649, 49-63., this genus includes 38 species distributed among three subgenera and eight species recorded from Texas to Argentina. Coprophanaeus gamezi Arnaud, 2002 (Figure 1I) has been found in tropical dry forests in Colombia (Solís et al. 2011SOLĺS, C., NORIEGA, J.A. & HERRERA, G. 2011. Escarabajos coprófagos (Coleoptera: Scarabaeinae) en tres bosques secos del departamento del Atlántico-Colombia. Boletín del Mus. Entomol. la Univ. del Val. 12:33-41.), the coast and provinces in Venezuela (Ferrer-Paris et al. 2013, Lozano 2010LOZANO, C. 2010. Nuevos registros de escarabajos coprófagos para la Orchila, Sierra de Perijá, Venezuela. Boletín Cent. Investig. Biológicas 44:83-89.) and adjacent Amazon regions of Brazil, Colombia and Guyana (Edmonds & Zidek 2010EDMONDS, W.D. & ZIDEK, J. 2010. Taxonomic review of the Neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta mundi Paper 649, 49-63.). Accordingly, it has been suggested as preferring open habitats and readily invading pasturelands (Edmonds & Zidek 2010EDMONDS, W.D. & ZIDEK, J. 2010. Taxonomic review of the Neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta mundi Paper 649, 49-63.), and has already been reported in introduced pastures and banana plantations (Gámez & Acconcia 2009GÁMEZ, J. & ACCONCIA, R. 2009. Ecological informations on Coprophanaeus (Coprophanaeus) gamezi Arnaud (Coleoptera: Scarabaeidae: Phanaeini) in a rural agricultural area in the depression of Maracaibo. Zulia State. Venezuela. Acta Zool. Mex. Nueva Ser. 25:387-396., Gámez 2010GÁMEZ, J. 2010. Escarabajos necrófagos (Coleoptera: Scarabaidae: Scarabainae: Phanaeini y Coleoptera: Silphidae: Nicrophorinae y Silphinae) de la Cordillera de los Andes, depresión de Maracaibo y Llanos de Venezuela. Kempffiana 6:15-19.).

Here we reinforce the unique botanical and faunal characteristics that Amazonian savannahs exhibit, which are considered an important area of endemism within South America (Barbosa et al. 2007). Hence, the studies concerning the fauna of this region should aim to establish conservation strategies associated with this important environment. Our study was the first to document the dung beetle fauna from Roraima savannahs and we believe that knowing the biodiversity is the first step in developing conservation strategies. Thus, since human disturbances threaten these environments (Barbosa et al. 2007, 2011), what leads to biodiversity loss, we reinforce that efforts should be undertaken towards the conservation of intra-Amazonian savannahs. Furthermore, we suggest further studies aiming to understand the effects of human disturbances on the dung beetle diversity in the Brazilian intra-Amazonian savannahs.

Acknowledgments

We thank Lívia Audino, Ross Thomas, the editor and anonymous reviewers for constructive comments on earlier versions of the manuscript. We thank to Fausto França and Edna Macedo for logistical and financial support, also to Daniel Macedo for his invaluable field assistance. We also thank T. Pacheco, L. Vieira and T. Vidaurre for allowing us to use dung beetle photographs from their previous publications; and Amanda Arcanjo and Laís Maia for taking the dung beetle photographs we included in this work. Insect Ecology and Conservation Lab (UFLA) was funded by and CNPq-PELD site 23, and by FAPEMIG, CNPq and CAPES grants. F.F. was awarded a studentship by CAPES (BEX 5528/13-5). F.V.M. was granted by CNPq (405697/2013-9, 302997/2013-0, 484035/2013-4, 202327/2013-2) and thanks very especially to the Assis and Estevam families (especially Joana and Pedro Raimundo - in memoriam - and Pedro Heber), and Adrian Forsyth for logistical support.

References

  • ABOT, A.R., PUKER, A., TAIRA, T.L., RODRIGUES, S.R., KORASAKI, V. & OLIVEIRA, H.N. 2012. Abundance and diversity of coprophagous beetles (Coleoptera: Scarabaeidae) caught with a light trap in a pasture area of the Brazilian Cerrado Stud. Neotrop. Fauna Environ. 47:53-60, 10.1080/01650521.2012.662846.
    » https://doi.org/10.1080/01650521.2012.662846
  • ALMEIDA, S., LOUZADA, J., SPERBER, C. & BARLOW, J. 2011. Subtle Land-Use Change and Tropical Biodiversity: Dung Beetle Communities in Cerrado Grasslands and Exotic Pastures. Biotropica 43:704-710, 10.1111/j.1744-7429.2011.00751.x.
    » https://doi.org/10.1111/j.1744-7429.2011.00751.x
  • ANDRADE, R.B., BARLOW, J., LOUZADA, J., VAZ-DE-MELLO, F.Z., SOUZA, M., SILVEIRA, J.M. & COCHRANE, M.A. 2011. Quantifying Responses of Dung Beetles to Fire Disturbance in Tropical Forests: The Importance of Trapping Method and Seasonality. PLoS One 6:e26208, 10.1371/journal.pone.0026208.
    » https://doi.org/10.1371/journal.pone.0026208
  • AUDINO, L.D., SILVA, P.G., NOGUEIRA, J.M., MORAES, L.P. & VAZ-DE-MELLO, F.Z. 2011. Scarabaeinae (Coleoptera, Scarabaeidae) de um bosque de eucalipto introduzido em uma região originalmente campestre. Iheringia. Série Zool. 101:121-126.
  • BARBOSA, R.I. & CAMPOS, C. 2011. Detection and geographical distribution of clearing areas in the savannas ('lavrado') of Roraima using Google Earth web tool. J. Geogr. Reg. Plann. 4(3):122-136.
  • BARBOSA, R.I., CAMPOS, C., PINTO, F. & FEARNSIDE, P. 2007. The “Lavrados” of Roraima: Biodiversity and conservation of Brazil's Amazonian savannas. Funct. Ecosyst. Communities. 1(1):29-41.
  • BARBOSA, R.I. & FEARNSIDE, P.M. 2005. Fire frequency and area burned in the Roraima savannas of Brazilian Amazonia. For. Ecol. Manage. 204:371-384, 10.1016/j.foreco.2004.09.011.
    » https://doi.org/10.1016/j.foreco.2004.09.011
  • BICKNELL, J.E., PHELPS, S.P., DAVIES, R.G., MANN, D.J., STRUEBIG, M.J. & DAVIES, Z.G. 2014. Dung beetles as indicators for rapid impacts assessments: Evaluating best practice forestry in the neotropics. Ecol. Indic. 43:154-161, 10.1016/j.ecolind.2014.02.030.
    » https://doi.org/10.1016/j.ecolind.2014.02.030
  • BOILLY, O. & VAZ-DE-MELLO, F.Z. 2014. Les Scarabaeinae de Guyane: clé illustrée des genres. ACOREP-France Coléoptàres Guyane. Tome VII 103-112.
  • BOITO, G.T., GIUGGIA, J.A., ORNAGHI, J.A., GERARDO, U.A. & GIOVANINI, D. 2009. Uso de trampas “Barber” para determinar la diversidad de coleópteros epígeos asociados al cultivo de maní (Arachis hypogaea L.). FCA UNCuyo XLI N°1:23-31.
  • BRÜHL, C. & KRELL, F.T. 2003. Finding a rare resource: Bornean Scarabaeoidea (Coleoptera) attracted by defensive secretions of diplopoda. Coleopt. Bull. 57:51-55, 10.1649/0010-065X(2003)057[0051:FARRBS]2.0.CO;2.
    » https://doi.org/10.1649/0010-065X(2003)057[0051:FARRBS]2.0.CO;2
  • CANTIL, L.F., SÁNCHEZ, M.V. & GENISE, J.F. 2014. The Nest and Brood Ball of Canthon (Canthon) virens aff. paraguayanus Balthasar (Coleoptera: Scarabaeidae: Scarabaeinae). Coleopt. Bull. 68:384-386, 10.1649/072.068.0306.
    » https://doi.org/10.1649/072.068.0306
  • CARPIO, C., DONOSO, D.A., RAM=N, G. & DANGLES, O. 2009. Short term response of dung beetle communities to disturbance by road construction in the Ecuadorian Amazon. Ann. la Société Entomol. Fr. 45:455-469, 10.1080/00379271.2009.10697629.
    » https://doi.org/10.1080/00379271.2009.10697629
  • CAVALCANTE, C.O., FLORES, A.S. & BARBOSA, R.I. 2014. Fatores edáficos determinando a ocorrência de leguminosas herbáceas em savanas amazônicas. Acta Amaz. 44(3):379-386, 10.1590/1809-4392201300954.
    » https://doi.org/10.1590/1809-4392201300954
  • COPOBIANCO, J.P., VERĺSSIMO, A., MOREIRA, A., SAWER, D., IKEDA, S. & PINTO, L.P. 2001. Biodiversidade na Amazônia Brasileira: Avaliação de Ações Prioritárias para a Conservação, Uso Sustentável e Repartição de Benefícios. Estação Liberdade, São Paulo.
  • COSTA, C.M.Q., BARRETTO, J.W. & DE MOURA, R.D.C. 2014. Changes in the dung beetle community in response to restinga forest degradation. J. Insect Conserv. 18:895-902, 10.1007/s10841-014-9697-6.
    » https://doi.org/10.1007/s10841-014-9697-6
  • COSTA, C.M.Q., SILVA, F.A.B., FARIAS, Â.I. & MOURA, R.D.C. 2009. Diversidade de Scarabaeinae (Coleoptera, Scarabaeidae) coletados com armadilha de interceptação de vôo no Refúgio Ecológico Charles Darwin, Igarassu-PE, Brasil. Rev. Bras. Entomol. 53:88-94, 10.1590/S0085-56262009000100021.
    » https://doi.org/10.1590/S0085-56262009000100021
  • COSTA, F.C., PESSOA, K.K.T., LIBERAL, C.N., FILGUEIRAS, B.K.C., SALOMAO, R.P. & IANNUZZI, L. 2013. What is the importance of open habitat in a predominantly closed forest area to the dung beetle (Coleoptera, Scarabaeinae) assemblage? Rev. Bras. Entomol. 57:329-334, 10.1590/S0085-56262013000300012.
    » https://doi.org/10.1590/S0085-56262013000300012
  • DAMBORSKY, M.P., ALVAREZ BOHLE, M.C., IBARRA POLESEL, M.G., PORCEL, E.A. & FONTANA, J.L. 2015. Spatial and Temporal Variation of Dung Beetle Assemblages in a Fragmented Landscape at Eastern Humid Chaco Neotrop. Entomol. 44:30-39, 10.1007/s13744-014-0257-2.
    » https://doi.org/10.1007/s13744-014-0257-2
  • DURÃES, R., MARTINS, W.P. & VAZ-DE-MELLOS, F.Z. 2005. Dung beetle (Coleoptera: Scarabaeidae) assemblages across a natural forest-cerrado ecotone in Minas Gerais, Brazil. Neotrop. Entomol. 34:721-731, 10.1590/S1519-566X2005000500003.
    » https://doi.org/10.1590/S1519-566X2005000500003
  • EDMONDS, W.D. & ZIDEK, J. 2010. Taxonomic review of the Neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta mundi Paper 649, 49-63.
  • ENDRES, A.A., CREÃO-DUARTE, A.J. & HERNÁNDEZ, M.I.M. 2007. Diversidade de Scarabaeidae s. str. (Coleoptera) da Reserva Biológica Guaribas, Mamanguape, Paraíba, Brasil: uma comparação entre Mata Atlântica e Tabuleiro Nordestino. Rev. Bras. Entomol. 51:67-71, 10.1590/S0085-56262007000100012.
    » https://doi.org/10.1590/S0085-56262007000100012
  • ESCOBAR, F. 1997. Estudio de la comunidad de coleópteros coprófagos (Scarabeidae) en un remanente de Bosque Seco al Norte del Tolima, Colombia. Caldasia 19:419-430.
  • ESCOBAR, F., LOBO, J.M. & HALFFTER, G. 2005. Altitudinal variation of dung beetle (Scarabaeidae: Scarabaeinae) assemblages in the Colombian Andes. Glob. Ecol. Biogeogr. 14:327-337, 10.1111/j.1466-822X.2005.00161.x.
    » https://doi.org/10.1111/j.1466-822X.2005.00161.x
  • FABRICIUS, J.C. 1804. Systema Piezatorum secundum ordines, genera, species, adjectis, synonymis, locis, observationnibus, descriptionibus. (Reichard, C). Brunswick XIV, pp. 15-439.
  • FALQUETO, S.A., VAZ-DE-MELLO, F.Z. & SCHOEREDER, J.H. 2005. Are fungivorous Scarabaeidae less specialist? Ecol. Austral 15:17-22, S1667-782X2005000100003.
  • FERREIRA, L.V. 2001. Identificação de áreas prioritárias para a conservação da biodiversidade por meio da representatividade das unidades de conservação e tipos de vegetação nas ecorregiões da Amazônia brasileira, In: Biodiversidade na Amazônia Brasileira (Veríssimo, A. Moreira, A. Sawer, D. Santos, I. Pinto, L.P. & Copobianco, J.P. eds.). Editora Liberdade, Instituto Socioambiental, São Paulo, pp. 268-286.
  • FERRER-PARIS, J.R., SÁNCHEZ-MERCADO, A. & RODRĺGUEZ, J.P. 2013. Optimización del muestreo de invertebrados tropicales: Un ejemplo con escarabajos coprófagos (Coleoptera: Scarabaeinae) en Venezuela. Rev. Biol. Trop. 61:89-110, 10.15517/rbt.v61i1.10941.
    » https://doi.org/10.15517/rbt.v61i1.10941
  • FILGUEIRAS, B.K.C., TABARELLI, M., LEAL, I.R., VAZ-DE-MELLO, F.Z. & IANNUZZI, L. 2015. Dung beetle persistence in human-modified landscapes: Combining indicator species with anthropogenic land use and fragmentation-related effects. Ecol. Indic. 55:65-73, 10.1016/j.ecolind.2015.02.032.
    » https://doi.org/10.1016/j.ecolind.2015.02.032
  • FLECHTMANN, C.A.H., TABET, V.G. & QUINTERO, I. 2009. Influence of carrion smell and rebating time on the efficiency of pitfall traps to dung beetle sampling. Entomol. Exp. Appl. 132:211-217, 10.1111/j.1570-7458.2009.00885.x.
    » https://doi.org/10.1111/j.1570-7458.2009.00885.x
  • FORTI, L.C., RINALDI, I.M.P., CAMARGO, R., DA S. & FUJIHARA, R.T. 2012. Predatory Behavior of Canthon virens (Coleoptera: Scarabaeidae): A Predator of Leafcutter Ants. Psyche. 2012:1-5, 10.1155/2012/921465.
    » https://doi.org/10.1155/2012/921465
  • GÁMEZ, J. 2010. Escarabajos necrófagos (Coleoptera: Scarabaidae: Scarabainae: Phanaeini y Coleoptera: Silphidae: Nicrophorinae y Silphinae) de la Cordillera de los Andes, depresión de Maracaibo y Llanos de Venezuela. Kempffiana 6:15-19.
  • GÁMEZ, J. & ACCONCIA, R. 2009. Ecological informations on Coprophanaeus (Coprophanaeus) gamezi Arnaud (Coleoptera: Scarabaeidae: Phanaeini) in a rural agricultural area in the depression of Maracaibo. Zulia State. Venezuela. Acta Zool. Mex. Nueva Ser. 25:387-396.
  • GÉNIER, F. 1996. A revision of the Neotropical genus Ontherus Erichson (Coleoptera: Scarabaeidae, Scarabaeinae). Mem. Entomol. Soc. Canada 128:1-170, 10.4039/entm128170fv.
    » https://doi.org/10.4039/entm128170fv.
  • GILL, B.D. 1991. Dung beetles in tropical American forests. In: Dung Beetle Ecology (Cambefort, I & Hanski, Y. eds.), Princeton University Press, Princeton, pp. 211-229.
  • GRIES, R., LOUZADA, J., ALMEIDA, S., MACEDO, R. & BARLOW, J. 2012. Evaluating the impacts and conservation value of exotic and native tree afforestation in Cerrado grasslands using dung beetles. Insect Conserv. Divers. 5:175-185, 10.1111/j.1752-4598.2011.00145.x.
    » https://doi.org/10.1111/j.1752-4598.2011.00145.x
  • HALFFTER, G. & HALFFTER, V. 2009. Why and where coprophagous beetles (Coleoptera: Scarabaeinae) eat sedes, fruits or vegetable detritus. Bol. Soc. Entomol. Aragonesa. 45:1-22.
  • HALFFTER, G. & MARTĺNEZ, A. 1966. Revisión monográfica de los Canthonina americanos (Coleoptera, Scarabaeidae). Rev. Soc. Mex. Hist. Nat. 27:89-117.
  • HALFFTER, G. & MARTĺNEZ, A. 1977: Revision monografi ca de los Canthonina Americanos, IV parte. Clava parageneros y subgeneros. Folia Entomológica Mex. 38:29-107
  • HALFFTER, G. & MATTHEWS, E.G. 1966. The natural history of dung beetles of the subfamily Scarabaeinae (Coleoptera: Scarabaeidae). Folia Entomológica Mex. 12:1-312.
  • HERNANDÉZ, M.I.M. 2007. Besouros Escarabeíneos (Coleoptera: Scarabaeidae) da Caatinga Paraibana, Brasil. Oecologia Bras. 11:356-364, 10.4257/oeco.2007.1103.06.
    » https://doi.org/10.4257/oeco.2007.1103.06
  • HERTEL, F. & COLLI, G.R. 1998. The use of leaf-cutter Ants, Atta laevigata (Smith) (Hymenoptera: Formicidae), as a substrate for oviposition by the dung beetle Canthon virens Mannerheim (Coleoptera: Scarabaeidae) in central Brazil. Coleopt. Bull. 52:105-108.
  • HUBBER, O. 2006. Herbaceous ecosystems on the Guyana Shield, a regional overview. J. Biogeogr. 33:464-475, 10.1111/j.1365-2699.2005.01454.x.
    » https://doi.org/10.1111/j.1365-2699.2005.01454.x
  • JUNK, W.J. 1983. As águas da região Amazônica. In: Amazônia: Desenvolvimento, Integração E Ecologia (Salati, E. Junk, W.J. Shubart, H.O.R. & Oliveira, A. eds.). Editora Brasiliense, São Paulo, pp. 45-100.
  • KORASAKI, V., VAZ-DE-MELLO, F.Z., BRAGA, R.F., ZANETTI, R. & LOUZADA, J. 2012. Taxocenose de Scarabaeinae (Coleoptera: Scarabaeidae) em Benjamin Constant, AM. Acta Amaz. 42:423-432, 10.1590/S0044-59672012000300015.
    » https://doi.org/10.1590/S0044-59672012000300015
  • KRIŠTOFĺK, J., MAŠÁN, P., ŠUSTEK, Z. & KLOUBEC, B. 2003. Arthropods (Pseudoscorpionida, Acari, Coleoptera, Siphonaptera) in nests of the tengmalm's owl, Aegolius funereus Biol. Bratislava. 58:231-240.
  • LARSEN, T.H. 2011. Dung beetles of the Kwamalasamutu region, Suriname (Coleoptera: Scarabaeidae: Scarabaeinae). In: . eds.). A Rapid Biological Assessment of the Kwamalasamutu Region, Southwestern Suriname. RAP Bulletin of Biological Assessment 63. Conservation International, Arlington, pp. 91-103.
  • LIMA, M.G.A., SILVA, R.P., DE, A., SOUSA, M.D.F. & COSTA, E.M. 2013. Diversidade de Scarabaeinae (Coleoptera: Scarabaeidae) no Parque Botânico do Ceará, Caucaia - CE, Brasil. Rev. Ago@mbiente On-line 7:89-94, 10.18227/1982-8470ragro.v7i1.970.
    » https://doi.org/10.18227/1982-8470ragro.v7i1.970
  • LOPES, P.P., LOUZADA, J.N.C. & VAZ-DE-MELLO, F.Z. 2006. Organization of dung beetle communities (Coleoptera: Scarabaeidae) in areas of vegetation re-establishment in Feira de Santana, Bahia, Brazil. Sitientibus Série Ciências Biológicas 6:261-266.
  • LOUZADA, J., GARDNER, T., PERES, C. & BARLOW, J. 2010. A multi-taxa assessment of nestedness patterns across a multiple-use Amazonian forest landscape. Biol. Conserv. 143:1102-1109, 10.1016/j.biocon.2010.02.003.
    » https://doi.org/10.1016/j.biocon.2010.02.003
  • LOUZADA, 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. Zoociências 9:199-203.
  • LOUZADA, J.N.C. & SILVA, P.R.C.E. 2009. Utilisation of introduced Brazilian pastures ecosystems by native dung beetles: Diversity patterns and resource use. Insect Conserv. Diver. 2:45-52, 10.1111/j.1752-4598.2008.00038.x.
    » https://doi.org/10.1111/j.1752-4598.2008.00038.x
  • LOZANO, C. 2010. Nuevos registros de escarabajos coprófagos para la Orchila, Sierra de Perijá, Venezuela. Boletín Cent. Investig. Biológicas 44:83-89.
  • LUEDERWALDT, H. 1929. As espécies brasileiras do gênero Pinotus. (Coleoptera - Lamellicornidae - Coprini), com algumas considerações também sobre outras espécies. Rev. do Mus. Paul. 16, 603-775.
  • MATAVELLI, R.A. & LOUZADA, J.N.C. 2008. Invasão de áreas de savana intra-amazônicas por Digitonthophagus gazella (Fabricius, 1787) (Insecta: Coleoptera: Scarabaeidae). Acta Amaz. 38:153-158, 10.1590/S0044-59672008000100017.
    » https://doi.org/10.1590/S0044-59672008000100017
  • MEDINA, A.M. & LOPES, P.P. 2014. Resource utilization and temporal segregation of Scarabaeinae (Coleoptera: Scarabaeidae) community in a caatinga fragment. Neotrop. Entomol. 43:127-133, 10.1007/s13744-014-0198-9.
    » https://doi.org/10.1007/s13744-014-0198-9
  • MEDRI, ĺ.M. & LOPES, J. 2001. Scarabaeidae (Coleoptera) do Parque Estadual Mata dos Godoy e de área de pastagem, no norte do Paraná, Brasil. Rev. Bras. Zool. 18:135-141, 10.1590/S0101-81752001000500011.
    » https://doi.org/10.1590/S0101-81752001000500011
  • MENESES, M.E.N.S., COSTA, M.L. & COSTA, J.A.V. 2007. Os lagos do lavrado de Boa Vista - Roraima?: fisiografia , físico-química das águas , mineralogia e química dos sedimentos. Rev. Bras. Geociências 37:478-489.
  • MILHOMEM, M.S., MELLO, F.Z.V. & DINIZ, I.R. 2003. Técnicas de coleta de besouros copronecrófagos no Cerrado. Pesqui. Agropecuária Bras. 38:1249-1256, 10.1590/S0100-204X2003001100001.
    » https://doi.org/10.1590/S0100-204X2003001100001
  • NICHOLS, E., SPECTOR, S., LOUZADA, J., LARSEN, T., AMEZQUITA, S. & FAVILA, M.E. 2008. Ecological functions and ecosystem services provided by Scarabaeinae dung beetles. Biol. Conserv. 141:1461-1474, 10.1016/j.biocon.2008.04.011.
    » https://doi.org/10.1016/j.biocon.2008.04.011
  • NORIEGA, J.A., SOLIS, C., ESCOBAR, F. & REALPE, E. 2007. Escarabajos coprófagos (Coleoptera: Scarabaeidae) de la provincia de la Sierra Nevada de Santa Marta. Biota Colomb. 8:77-86.
  • NOVELO, E.R., DELFĺN-GONZÁLEZ, H. & MOR=N, M.Á. 2007. Copro-necrophagous beetle (Coleoptera: Scarabaeidae) diversity in an agroecosystem in Yucatan, Mexico. Rev. Biol. Trop. 55:83-99, 10.15517/rbt.v55i1.6059.
    » https://doi.org/10.15517/rbt.v55i1.6059
  • NUNES, R.V., FRIZZAS, M.R. & VAZ-DE-MELLO, F.Z. 2012. Scarabaeinae (Coleoptera: Scarabaeidae) of a rupestrian field at Caatinga, Distrito Federal, Brazil: commented list of species. Biota Neotrop. 12(4):125-129, 10.1590/S1676-06032012000400013.
    » https://doi.org/10.1590/S1676-06032012000400013
  • PACHECO, T.L. & VAZ-DE-MELLO, F.Z. 2015. Dung beetles of the tribe Phanaeini (Coleoptera: Scarabaeidade: Scarabaeinae) from Roraima state, Northern Brazil: checklist and key to species. Biota Neotrop. 15(2): 1-9, 10.1590/1676-06032015014514.
    » https://doi.org/10.1590/1676-06032015014514
  • PADILLA-GIL, D.N. & HALFFTER, G. 2007. Biogeography of the areas and Canthonini (Coleoptera: Scarabaeidae) of dry tropical forests in mesoamerica and colombia. Acta Zoológica Mex. 23:73-108.
  • PRANCE, G.T. 1996. Islands in Amazonia. Philos. Trans. R. Soc. B Biol. Sci. 351, 823-833, 10.1098/rstb.1996.0077.
    » https://doi.org/10.1098/rstb.1996.0077
  • PUKER, A., CORREA, C.M.A., KORASAKI, V., FERREIRA, K.R. & OLIVEIRA, N.G. 2013. Dung Beetles (Coleoptera: Scarabaeidae) Attracted to Dung of the Largest Herbivorous Rodent on Earth: A Comparison with Human Feces. Environ. Entomol. 42:1218-1225, 10.1603/EN13100.
    » https://doi.org/10.1603/EN13100
  • RIBEIRO, J.E.L.S., HOPKINS, M.J.G., VICENTINI, A., SOTHERS, C.A., COSTA, M.A.S., BRITO, J.M., SOUZA, M.A.D., MARTINS, L.H.P., LOHMANN, L.G., ASSUNÇÃO, P.A.C.L., PEREIRA, E.C., SILVA, C.F., MESQUITA, M.R., & PROCOPIO, L.C. 1999. Flora da reserva Ducke: guia de identificação das plantas vasculares de uma floresta de terra-firme na Amazônia Central. DFID - INPA, Manaus.
  • RODRIGUES, 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(2):123-127, 10.1590/S1676-06032010000200015.
    » https://doi.org/10.1590/S1676-06032010000200015
  • ROSA, T.A., BABATA, M.L.Y., SOUZA, C.M., SOUSA, D., MELLO-PATIU, C.A., VAZ-DE-MELLO, F.Z. & MENDES, J. 2011. Arthropods associated with pig carrion in two vegetation profiles of Cerrado in the State of Minas Gerais, Brazil. Rev. Bras. Entomol. 55:424-434, 10.1590/S0085-56262011005000045
    » https://doi.org/10.1590/S0085-56262011005000045
  • SALOMÃO, R.P. & IANNUZZI, L. 2015. Dung beetle (Coleoptera, Scarabaeidae) assemblage of a highly fragmented landscape of Atlantic forest: from small to the largest fragments of northeastern Brazilian region. Rev. Bras. Entomol. 59:126-131, 10.1016/j.rbe.2015.03.008.
    » https://doi.org/10.1016/j.rbe.2015.03.008
  • SANAIOTTI, T.M. 1997. Comparação fitossociológica de quatro savanas de Roraima. In: Homem, Ambiente e Ecologia no Estado de Roraima (Barbosa, R.I Ferreira, E.J.G & Castellón, E. eds.). INPA, Manaus, pp. 481-488.
  • SANAIOTTI, T.M., MARTINELLI, L.A., VICTORIA, R.L., TRUMBORE, S.E. & CAMARGO, P.B. 2002. Past vegetation changes in Amazon savannahs determined using carbon isotopes of soil organic matter. Biotropica 34:2-16, 10.1111/j.1744-7429.2002.tb00237.x.
    » https://doi.org/10.1111/j.1744-7429.2002.tb00237.x
  • SÁNCHEZ-HUERTA, J.L., TONELLI, M., ZUNINO, M. & HALFFTER, G. 2015. Redescription of Onthophagus halffteri Zunino (Coleoptera: Scarabaeidae: Scarabaeinae), with Ecological and Distributional Notes. Coleopt. Bull. 69:225-230, 10.1649/0010-065X-69.2.225.
    » https://doi.org/10.1649/0010-065X-69.2.225
  • SANTOS, W., ALVES, A. & CREÃO-DUARTE, A. 2014. Beetles (Insecta, Coleoptera) associated with pig carcasses exposed in a Caatinga area, Northeastern Brazil. Brazilian J. Biol. 74:649-655, 10.1590/bjb.2014.0072.
    » https://doi.org/10.1590/bjb.2014.0072
  • SARMIENTO, G. 1984. The Ecology of Neotropical Savannahs. Harvard University Press, Cambridge, UK.
  • SCHEFFLER, P.Y. 2005. Dung beetle (Coleoptera: Scarabaeidae) diversity and community structure across three disturbance regimes in eastern Amazonia. J. Trop. Ecol. 21:9-19, 10.1017/S0266467404001683.
    » https://doi.org/10.1017/S0266467404001683.
  • SCHIFFLER, G., VAZ-DE-MELLO, F.Z. & AZEVEDO, C.O. 2003. Scarabaeidae s. str. (Coleoptera) from the Doce river and Suruaca valleu in Linhares municipality, Espírito Santo State, Brazil. Rev. Bras. Zoociências 5:205-211.
  • SCHMITT, T., KRELL, F.T. & LINSENMAIR, K.E. 2004. Quinone mixture as attractant for necrophagous dung beetles specialized on dead millipedes. J. Chem. Ecol. 30:731-740, 10.1023/B:JOEC.0000028428.53797.cb.
    » https://doi.org/10.1023/B:JOEC.0000028428.53797.cb
  • SILVA, P.G., VAZ-DE-MELLO, F.Z. & MARE, R.A. 2011. Guia de identificação das espécies de Scarabaeinae (Coleoptera: Scarabaeidae) do município de Santa Maria, Rio Grande do Sul, Brasil. Biota Neotrop. 11(4):329-345, 10.1590/S1676-06032011000400027.
    » https://doi.org/10.1590/S1676-06032011000400027
  • SILVA, F.A.B., COSTA, C.M.Q., MOURA, R.C. & FARIAS, A.I. 2010. Study of the dung beetle (Coleoptera: Scarabaeidae) community at two sites: Atlantic forest and clear-cut, Pernambuco, Brazil. Environ. Entomol. 39:359-367, 10.1603/EN09180.
    » https://doi.org/10.1603/EN09180
  • SILVA, F.A.B., HERNÁNDEZ, M.I.M., IDE, S. & MOURA, R.C. 2007. Comunidade de escarabeíneos (Coleoptera, Scarabaeidae) copronecrófagos da região de Brejo Novo, Caruaru, Pernambuco, Brasil. Rev. Bras. Entomol. 51:228-233, 10.1590/S0085-56262007000200014.
    » https://doi.org/10.1590/S0085-56262007000200014
  • SILVA, P. & AUDINO, L.D. 2011. Escarabeíneos (Coleoptera: Scarabaeidae) atraídos a diferentes iscas em campo nativo de Bagé, Rio Grande do Sul, Brasil. Rev. Bras. Zoociências 13:241-247.
  • SILVA, P.G., ROSA-GARCIA, A.M. & VIDAL, M.B. 2008. Espécies de besouros copro-necrófagos (Coleoptera: Scarabaeidae) da Região da Campanha do Rio Grande do Sul. Rev. Ciências Agroveterinárias 7:143-149.
  • SILVA, R.J., COLETTI, F., COSTA, D. & VAZ-DE-MELLO, F.Z. 2014. Rola-bostas (Coleoptera: Scarabaeidae: Scarabaeinae) de florestas e pastagens no sudoeste da Amazônia brasileira: Levantamento de espécies e guildas alimentares. Acta Amaz. 44:345-352, 10.1590/1809-4392201304472.
    » https://doi.org/10.1590/1809-4392201304472
  • SILVA, R.J., DINIZ, S. & VAZ-DE-MELLO, F.Z. 2010. Heterogeneidade do habitat, riqueza e estrutura da assembléia de besouros rola-bostas (Scarabaeidae: Scarabaeinae) em áreas de cerrado na Chapada dos Parecis, MT. Neotrop. Entomol. 39:934-940, 10.1590/S1519-566X2010000600014.
    » https://doi.org/10.1590/S1519-566X2010000600014
  • SILVA, R.J., RIBEIRO, H.V., SOUZA, M.F. & VAZ-DE-MELLO, F.Z. 2015. Influence of soil granulometry on the structure of functional guilds of dung beetles (Coleoptera: Scarabaeidae: Scarabaeinae) at semideciduos forests in the State of Mato Grosso, Brazil. Biosci. J. 31:601-612, 10.14393/BJ-v31n1a2015-23525.
    » https://doi.org/10.14393/BJ-v31n1a2015-23525.
  • SLAY, M.E., SKELLEY, P.E. & TAYLOR, S.J. 2012. Howden and Cartwright (Coleoptera: Scarabaeidae) from Ozark Caves, with a Review of Scarabaeoids Reported from North American Caves. Coleopt. Bull. 66:187-199, 10.1649/072.066.0302.
    » https://doi.org/10.1649/072.066.0302
  • SOLĺS, C., NORIEGA, J.A. & HERRERA, G. 2011. Escarabajos coprófagos (Coleoptera: Scarabaeinae) en tres bosques secos del departamento del Atlántico-Colombia. Boletín del Mus. Entomol. la Univ. del Val. 12:33-41.
  • TARASOV, S.I. & KABAKOV, O.N. 2010. Two new species of Onthophagus (Coleoptera: Scarabaeidae) from Indochina, with a discussion of some problems with the classification of Serrophorus and similar subgenera. Zootaxa 2344:17-28.
  • VANZOLINI, P.E. & CARVALHO, C.M. 1991. Two sibling and sympatric species of Gymnophthalmus in Roraima, Brazil (Sauria Teiidae). Pap. Avulsos Zool. 37:173-226.
  • VAZ-DE-MELLO, F.Z. & LOUZADA, J.N.C. 1997. Considerações sobre o forrageio arbóreo por Scarabaeidae (Coleoptera, Scarabaeoidea) e dados sobre sua ocorrência em floresta tropical do Brasil. Acta Zool. Mex. Nueva Ser. 72:55-61.
  • VAZ-DE-MELLO, F.Z., LOUZADA, J.N.C. & SCHOEREDER, J.H. 1998. New data and comments on Scarabaeidae (Coleoptera: Scarabaeoidea) associated with Atini (Hymenoptera: Formicidae). Coleopt. Bull. 52:209-216.
  • VAZ-DE-MELLO, F.Z. 1999a. Scarabaeidae do Estado de Roraima. Memórias da IV Reunião Latino-Americana de Scarabaeoidologia, Viçosa, MG, p. 78-79.
  • VAZ-DE-MELLO, F.Z. 1999b. Scarabaeidae s. str. (Coleoptera: Scarabaeoidea) de um fragmento de Floresta Amazônica no estado do Acre, Brasil. 1. Taxocenose. An. da Soc. Entomológica do Bras. 28:439-446, 10.1590/S0301-80591999000300009.
    » https://doi.org/10.1590/S0301-80591999000300009
  • VAZ-DE-MELLO, F.Z. 2000. Estado Atual de conhecimentos dos Scarabaeidae s. str. (Coleoptera: Scarabaeoidea) do Brasil. In: Hacia Un Proyecto CYTED Para El Inventario Y Estimación de La Diversidad Entonológica Em Iberoamérica: PrIBES-2000 (Martín-Piera, F. Morrone, J.J. & Melic, A. eds.). Sociedad Entomologica Aragonesa & CYTED, Zaragoza, pp. 181-195.
  • VIDAURRE, T., GONZALES, L. & LEDEZA, J.M. 2008. Escarabajos coprófagos (Scarabaeidae: Scarabaeinae) del Palmar de las Islas, Santa Cruz, Bolívia. Kempffiana 4:3-20.
  • VIEGAS, G., STENERT, C., SCHULZ, U.H. & MALTCHIK, L. 2014. Dung beetle communities as biological indicators of riparian forest widths in southern Brazil. Ecol. Indic. 36:703-710, 10.1016/j.ecolind.2013.09.036.
    » https://doi.org/10.1016/j.ecolind.2013.09.036
  • VIEIRA, L., LOUZADA, J.N.C. & SPECTOR, S. 2008. Effects of degradation and replacement of Southern Brazilian Coastal Sandy Vegetation on the dung beetles (Coleoptera: Scarabaeidae). Biotropica 40:719-727, 10.1111/j.1744-7429.2008.00432.x.
    » https://doi.org/10.1111/j.1744-7429.2008.00432.x
  • VIEIRA, L. & SILVA, F.A.B. 2012. List Dung beetles (Coleoptera: Scarabaeidae: Scarabaeinae) of the Floresta Nacional Contendas do Sincorá, Bahia, Brazil. Check List 8:733-739.
  • VILLALOBOS, F.J., DIAZ, A. & FAVILA, M.E. 1998. Two species of Canthon feed on dead and live invertebrates. Coleopt. Bull. 52:101-104.

Publication Dates

  • Publication in this collection
    Mar 2016

History

  • Received
    30 Mar 2015
  • Reviewed
    29 Dec 2015
  • Accepted
    20 Jan 2016
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