Biodiversity Data Journal 11: e96101 OO) doi: 10.3897/BDJ.11.e96101 open access Taxonomy & Inventories Dung beetles from two sustainable-use protected forests in the Brazilian Amazon Edrielly C. CarvalhotS, Maria Eduarda Maldaner', Vinicius Costa-Silva+1, Heivanice Sehn*, Carol Franquinit, Vinicius O. Campos, Vinicius P. Sebat, Lais F. Maia", Fernando Z. Vaz-de-Mellot, Filipe Machado Franga*"“ + Departamento de Biologia e Zoologia, Instituto de Biociéncias, Universidade Federal de Mato Grosso. Laboratorio de Scarabaeoidologia. Instituto de Biociéncias - UFMT, Cuiaba, Brazil § Programa de Pos-Graduagao em Entomologia. Instituto Nacional de Pesquisas da Amazénia — INPA, Manaus, Brazil | Programa de Pos-Graduagao em Ecologia e Conservagao da Biodiversidade (PPGECB), Universidade Federal de Mato Grosso - UFMT, Cuiaba, Brazil 4 Laboratory of Integrative Entomology, Department of Animal Biology, Institute of Biology, University of Campinas, Campinas, Brazil # Programa de Pés-Graduagao em Zoologia. Instituto de Biociéncias, Universidade Federal de Mato Grosso - UFMT, Cuiaba, Brazil = School of Biological Sciences, University of Bristol, Queens Road, BS8 1QU, UK, Bristol, United Kingdom « Programa de Pés-Graduagdo em Ecologia (PPGECO), Universidade Federal do Para, Belém, PA, 66075-110, Brazil, Belém, Brazil Corresponding author: Edrielly C. Carvalho (edrielly.carol@gmail.com) Academic editor: Matthias Seidel Received: 07 Oct 2022 | Accepted: 14 Feb 2023 | Published: 16 Mar 2023 Citation: Carvalho EC, Maldaner ME, Costa-Silva V, Sehn H, Franquini C, Campos VO, Seba VP, Maia LF, Vaz- de-Mello FZ, Franga FM (2023) Dung beetles from two sustainable-use protected forests in the Brazilian Amazon. Biodiversity Data Journal 11: e96101. https://doi.org/10.3897/BDJ.11.e96101 Abstract Background The Amazon Forest is one of the world's most biodiverse ecosystems and yet its protected areas are understudied concerning insects and other invertebrates. These organisms are essential for tropical forests due to their ecological processes, with some species being very sensitive to habitat disturbances. Dung beetles (Coleoptera, Scarabaeidae, Scarabaeinae) have been used as bioindicators for more than 30 years and were surveyed to assess the insect biodiversity of two sustainable-use forest reserves in the Brazilian Amazon. © Carvalho E et al. This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. 2 Carvalho E et al New information We report inventories of dung beetles from two Amazonian forest reserves in Para State, Brazil: the Tapajos National Forest and the Carajas National Forest. Surveys were carried out with baited-pitfall traps installed in 2010, 2016, 2017 and 2019. We collected a total of 3,772 individuals from 19 genera and 96 species. We highlight the importance of Amazonian protected areas as refugia for insect biodiversity, particularly dung beetles, which contribute to many key ecosystem processes. Keywords Amazonia, biodiversity, Coleoptera, dung beetles, Scarabaeinae, sustainable-use forests, tropical ecosystems Introduction The Amazon Forest has global importance for biodiversity, being amongst the world’s most diverse tropical ecosystems (Barlow 2018). The region's warm and humid climate makes Amazonian forests a unique ecosystem with extraordinary biodiversity levels, particularly for insects (Fearnside 2008, Sobral-Souza and Lima-Ribeiro 2017). With the human footprint and climate extremes increasing within tropical regions (Franga et al. 2020a), protected areas are increasingly becoming the final refuges for biodiversity, including many restricted-range and highly threatened species (Rylands and Brandon 2005 , sollmann et al. 2008). Until 2009, around 54% of the remaining Amazon Forest in Brazil was part of a protected area network, ranging from strictly protected areas and indigenous lands to sustainable use forests (Soares-Filho et al. 2010). The importance of these protected areas goes beyond biodiversity conservation by sustaining local livelinoods (Naughton-Treves et al. 2005, Spinola et al. 2020), preventing climate-tipping points (Walker et al. 2009) and supporting the mitigation of climate changes through carbon accumulation and reduction of Greenhouse Gases (GHG) emissions from deforestation (Soares-Filho et al. 2010, Walker et al. 2020). Despite the unparalleled contribution of insects to the totality of biodiversity (Zhang 2011) and their important role in many critical ecological functions (Nichols et al. 2008, Campbell et al. 2012, Dangles and Casas 2019), insects are understudied when compared to vertebrates (Dornelas and Daskalova 2020). For example, only 1.67% of the known invertebrate species have been assessed by the IUCN Red List of threatened species compared with 68.9% of all vertebrates (Kitching et al. 2020). The lack of studies with insects and invertebrates is also evidenced within Amazonian protected areas. Dung beetles (Coleoptera, Scarabaeinae) are a key group of detritivore insects frequently used in ecological research linking biodiversity and ecosystem functioning (Franga et al. 2018, Griffiths et al. 2015). Through feeding and nesting in mammal dung, carrion or rotten Dung beetles from two sustainable-use protected forests in the Brazilian ... 3 fruits, dung beetles play important roles in nutrient cycling and other ecosystem functions (Halffter and Matthews 1966, Nichols et al. 2008). Given their quick responses to environmental degradation by anthropogenic and climatic disturbances (e.g. Franga et al. 2020a, Franga et al. 2020b), since the 1990s, dung beetles have been used as an efficient indicator of environmental quality in tropical forests (Halffter and Favila 1993, Davis et al. 2004, Larsen and Forsyth 2005, Spector 2006, Nichols et al. 2007, Gardner et al. 2008, Nichols et al. 2008, Culot et al. 2013). In general, anthropogenic activities lead to changes in dung beetle ‘fitness' (through physiological stress: for example, Franga et al. (2016), Salomao et al. (2018)), species richness and abundance (Klein and Bert 1989, Halffter and Arellano 2002, Escobar et al. 2007). Here, we: (1) present a list of dung beetle species surveyed at two sustainable-use protected forests in the Brazilian Amazon — the Tapajos National Forest and the Carajas National Forest (FLONAS); and (2) discuss insights associated with the species distribution and previous recordings in literature. Materials and methods Study region The Tapajos National Forest and Carajas National Forest (hereafter ‘'Tapajos' and 'Carajas', respectively) cover 527,319 ha and 411,948 ha of Amazonian forests, respectively, spread across multiple municipalities in the State of Para, Brazil (Fig. 1). These FLONAS are located in two Amazonian biogeographical regions with distinct socio-environmental contexts. Specifically, the Tapajos region has a more recent history of agriculture expansion and lower deforestation rates than the Carajas region (Braz 2016), with FLONA Carajas located within a mosaic of federal and state forest reserves, national parks and indigenous protected lands (Pilo et al. 2015). The climate is characterised as hot-humid (K6ppen's classification) and the annual average temperature is 25-26°C in both regions, which have short dry seasons in August-November (average precipitation [mm]: Tapajos = 1405.8) and July-September (Xingu = 84.8). Surveyed sites are within the ‘terra-firm’ forests, with vegetation varying depending on soil and relief. Sampling design Dung beetles were sampled within a total of 13 forest sites (Carajas = 3 and Tapajos = 10). We surveyed Carajas in February—March 2019, while Tapajos forests were surveyed in June-July 2010, June—July 2016, March-April 2017 and Feb—March 2019. These field sites are part of the Long-Term Ecological Research Program of the Sustainable Amazon Network (PELD-RAS). At each of our forest sites, dung beetles were sampled at three sampling points (0, 150 and 300 m) along a 300-m transect. As in Franga et al. (2020b), we used three dung-baited pitfall traps arranged at the ends of a 2-m equilateral triangle at each sampling point, resulting in a total of 117 traps (21 and 96 pitfalls in Carajas and Tapajos, respectively). Pitfall traps were 1-litre plastic containers (14 cm in diameter; 9 cm deep) buried in the ground with the opening at ground level and protected from rain with a 4 Carvalho E et al plastic lid suspended 15 cm above the surface. Each trap was part-filled with a saline killing solution, had a bait container with 35 g of dung (4:1 pig to human ratio, following Marsh et al. (2013)) supported by a wire above the trap and was left in the field for 48 hours. FLONA Carajas be —~ —__ 4 Ee é Qo 40 eure 60 «240 320 T Figure 1. EES Map showing the localities of Tapajés National Forest (FLONA Tapajos) and Carajas National Forest (FLONA Carajas; both in dark grey) in Para State (medium grey), in the north of Brazil (light grey). Data resources All trapped dung beetles were collected and taken to the laboratory, where they were sorted, mounted and identified to species (using identification key or descriptions) or morphospecies. Voucher specimens were deposited at the Entomological Section of the Zoological Collection (CEMT) at the Federal University of Mato Grosso, Brazil (UFMT). Specimens were photographed using the Leica M250C Photomontage Equipment (UFMT/ Finep) and an Olympus SZX16 stereomicroscope with expandable stream motion imaging software v. 2.5 (UoB/Liv Sidse Hansen Foundation). The morphospecies identification numbers are not indicating the amount of species collected at the sites and are purely reference numbers for species across multiple projects. The abbreviations aff., cf. and gp. are qualifiers used in taxonomy to indicate different degrees of uncertainty of identification. The use of aff. and cf. follows Lucas (2012) and gp. indicates species group affinity. The map showing the localities of Tapajos National Forest and Carajas National Forest was prepared using ArcGIS 10.8 software. Dung beetle data can be found at hittp:// www.gbif.org/tools/data-validator/f1e2a538-5fea-4258-9b0e-27805b684404 (GBIF 2022). Dung beetles from two sustainable-use protected forests in the Brazilian ... 5 List and abundance of species present in FLONA Tapajos and Carajas Subfamily Scarabaeinae Latreille, 1802 Notes: We collected 3,772 dung beetles from 96 species and 19 genera. Only 14 of the 96 identified species were found in both FLONAS (Table 1). Canthidium deyrollei was the most abundant species, with exclusive records from FLONA Tapajos. The three most diverse genera comprise distinct functional strategies in dung beetles: Eurysternus (endocoprids — i.e. residents in the dung resource); Dichotomius (paracoprids — i.e. tunnellers, where all species mostly dig tunnels close to or immediately below the resource) and Canthon (telecoprids — i.e. rollers), which could be an indicator of whole exploitation of dung resources in Amazonian forests. However, if abundance is considered, small paracoprids (especially Onthophagus species) were the most abundant functional group, which is expected for the Amazon Region (FVM, pers. obs). We discuss below the current knowledge about the distribution and ecology of each identified genera. Table 1. List and abundance of species present in FLONA Tapajos and Carajas. Species Abundance FLONA Tapajos FLONA Carajas Anomiopus aff. pereirai 1 - Anomiopus sp. 2 1 - Anomiopus sp. 3 1 - Anomiopus sp. 4 1 - Anomiopus sp. 5 - 1 Ateuchus globulus (Balthasar, 1938) 1 - Ateuchus sp. 2 14 - Ateuchus sp. 3 10 - Ateuchus gp. pygidialis - 1 Ateuchus cf. murrayi 113 - Ateuchus sp. 4 64 - Ateuchus semicupreus (Harold, 1868) - 18 Ateuchus sp. 1 - 39 Ateuchus substriatus (Harold, 1868) 12 - Canthidium deyrollei Harold, 1867 283 - Canthidium melanocephalum (Olivier, 1789) 31 - Canthidium sp. 1 - 13 Species Canthidium sp. Canthidium sp. Canthidium sp. Canthidium sp. Canthidium sp. Canthidium sp. Canthidium sp. Canthidium sp. Canthidium sp. Canthidium sp. Canthidium sp. Canthidium sp. Canthidium sp. Canthidium sp. Canthidium sp. Canthidium sp. Canthidium sp. Canthidium sp. Canthidium sp. Canthidium sp. Canthidium sp. Canthon aff. histrio Canthon aff. sericatus Canthon aff. xanthopus Canthon conformis Harold, 1868 Canthon fulgidus Redtenbacher, 1868 Canthon histrio (Lepeletier de Saint-Fargeau & Audinet-Serville, 1828) N Oo ao fF W DN Carvalho E et al Canthon subhyalinus (Rivera-Cervants & Halffter, 1999) Canthon semiopacus Harold, 1868 Canthon triangularis (Drury, 1770) Coprophanaeus degallieri Arnaud, 1997 Coprophanaeus jasius (Olivier, 1789) Coprophanaeus l/ancifer (Linnaeus, 1767) Cryptocanthon campbellorum Howden, 1973 Deltochilum enceladus Kolbe, 1893 Abundance FLONA Tapajos 4 84 128 FLONA Carajas 3 29 29 35 Dung beetles from two sustainable-use protected forests in the Brazilian ... Species Deltochilum gp. aspericolle Deltochilum gp. guyanense Deltochilum gp. sextuberculatum Deltochilum orbiculare Van Lansberge, 1874 Deltochilum orbignyi amazonicum Bates, 1887 Deltochilum sp. 1 Dichotomius aff. batesi Dichotomius aff. lucasi 1 Dichotomius aff. lucasi 2 Dichotomius cuprinus (Felshe, 1901) Dichotomius mamillatus (Felshe, 1901) Dichotomius meizeri (Luederwaldt, 1922) Dichotomius nisus (Olivier, 1789) Dichotomius pelamon (Harold, 1869) Dichotomius worontzowi (Pereira, 1942) Eurysternus arnaudi Génier, 2009 Eurysternus atrosericus Génier, 2009 Eurysternus balachowskyi Halffter & Halffter, 1977 Eurysternus caribaeus (Herbst, 1789) Eurysternus cavatus Génier, 2009 Eurysternus cayennensis Castelnau, 1840 Eurysternus cyclops Génier, 2009 Eurysternus fallaciosus Génier, 2009 Eurysternus foedus Guérin-Méneville, 1844 Eurysternus hamaticollis Balthasar, 1939 Eurysternus hypocrita Balthasar, 1939 Eurysternus plebejus Harold, 1880 Eurysternus wittmerorum Martinez, 1988 Eutrichillum sp. 1 Hansreia oxygona (Perty, 1830) lsocopris imitator (Felsche, 1901) lsocopris nitidus (Luederwaldt 1922) Ontherus carinifrons Luederwaldt, 1930 Onthophagus digitifer Boucomont, 1932 Onthophagus gp. rubrescens Abundance FLONA Tapajos 156 13 6 13 91 FLONA Carajas 45 27 36 50 44 17 147 8 Carvalho E et al Species Abundance FLONA Tapajos FLONA Carajas Onthophagus onthochromus Arrow, 1913 - 1 Onthophagus osculatii Guérin-Méneville, 1855 70 10 Oxysternon macleayi Nevinson, 1892 26 11 Oxysternon silenus Castelnau, 1840 1 3 Scybalocanthon sp. 1 - 2 Sulcophanaeus faunus (Fabricius, 1775) 1 Sylvicanthon candezei (Harold, 1869) 1 Sylvicanthon proseni (Martinez, 1949) 116 Uroxys cf. minutus 29 8 Genus Anomiopus Westwood, 1842 Notes: Anomiopus is a Neotropical genus with most species occurring in South America. The latest revision has 48 described species (Canhedo 2004, Canhedo 2006). Most Anomiopus species are collected with flight interception traps (FIT), Malaise traps, pitfalls baited with human dung (Canhedo 2006), light traps and bird faeces (Martinez 1959). Some species were observed in the Colombian Amazon landing on leaves during the day (Canhedo 2006). In our study, we found five species: Anomiopus aff. pereirari, Anomipus sp. 2, Anomiopus sp. 3, Anomiopus sp. 4 and Anomiopus sp. 5. Genus Ateuchus Weber, 1801 Notes: With around 100 species described, this genus needs urgent revision. The last revision of Brazilian Ateuchus species was done by Harold (1868), while Balthasar (1939) represents the last identification key for the genus. Most species occur in North America, Costa Rica and Mexico (Kohlmann 1984, Kohlmann 1997, Genier 2000, Kohlmann and Vaz-de-Mello 2018) and appear to be copro-necrophagous, including species from open areas and species that live associated with ant nests (Vaz-de-Mello et al. 1998). Nine species were identified in our study: Ateuchus globulus (Balthasar, 1938), A. cf. pygidialis, A. cf. murrayi, A. semicupreus (Harold, 1868), A. substriatus (Harold, 1868) and Ateuchus sp. 1, Ateuchus sp. 2, Ateuchus sp. 3 and Ateuchus sp. 4 (Fig. 2A-G). Genus Canthidium Erichson, 1847 Notes: This is one of the most diverse dung beetle genera, comprising around 180 described species (Genier and Cupello 2018, Schoolmeesters 2022). Numerous species were described in small revisions, synopses and regional studies (e.g. Boucomont (1928), Balthasar (1939), Martinez et al. (1964), Howden and Young Dung beetles from two sustainable-use protected forests in the Brazilian ... FS) (1981), Solis and Kohlmann (2004), Kohlmann and Solis (2006)), while new species are expected to be described (Cupello 2018, Kohlmann et al. 2018, Santana et al. 2019 ). Canthidium species have been recorded within Neotropical forests and _ intra- Amazonian savannahs (e.g. Franga et al. (2016)). Although little is known about most species’ habits, specimens are easily collected in traps baited with faeces, rotten fruit and/or light traps (e.g. Medri and Lopes (2001), Silva and Audino (2011), Silva et al. (2014)), while some species were observed feeding on fungus (Falqueto et al. 2005). The specimens were identified from comparison with the original types and descriptions, which were analysed by one of the authors. Two species were identified to the species level in our survey in FLONA Tapajos: Canthidium deyrollei Harold, 1867 and C. melanocephalum (Olivier, 1789) (Fig. 2H-l). Other 22 species are present, but could not be identified. A yy} | Figure 2. EE Dorsal habitus of dung beetle species collected in the Tapajés National Forest and/or Carajas National Forest A Ateuchus globulus (Balthasar, 1938); B Ateuchus semicupreus (Harold, 1868); C Ateuchus substriatus (Harold, 1868); D Canthidium deyrollei Harold, 1867; E Canthidium melanocephalum (Olivier, 1789); F Canthon conformis Harold, 1868; G Canthon fulgidus Redtenbacher, 1868; M Canthon histrio (Lepeletier de Saint Fargeau & Audinet- Serville, 1828); | Canthon subhyalinus (Rivera-Cervants & Halffter, 1999); J Canthon semiopacus Harold, 1868; L Canthon triangularis (Drury, 1770). Scale bar: 1 mm. 10 Carvalho E et al Genus Canthon Hoffmannsegg, 1817 Notes: This is also a very diverse genus, comprising more than 170 described species (Halffter and Martinez 1977). Most species are considered copro-necrophagous, although some exhibit predatory behaviour — for example, hunting ants (Halffter and Matthews 1966) — or use dead insects and millipedes (Villalobos et al. 1988, Silva et al. 2014), rotten fruits and fungus as food resources (Vaz-de-Mello 1999). This genus is endemic to the Americas and its distribution ranges from the USA to Uruguay and northern Argentina. Recent revisions have been made for some Canthon subgenera (Nunes et al. 2018, Nunes et al. 2020). Typically, these species are abundant in lowland forest environments, with individuals found perching on leaves exposed to light (Nunes et al. 2018). Another important point to be discussed is the population of Canthon fulgidus Redtenbacher, 1868 with green colour living in eastern Amazonia. According to Nunes et al. (2018), the population with green colour, named Canthon fulgidus martinezi Nunes et al., 2018, is restricted to the western Amazon, while the populations from Carajas and Tapapos regions were expected to have a red metallic colour (named by the authors as Canthon fulgidus pereirai Nunes et al., 2018). This new finding (both green and red populations collected in the same region) suggests that Nunes et al. (2018) may have overlooked the green specimens from eastern Amazonia, as previously mentioned by Cupello et al. (2021), who discuss the colour variation and geographical distribution of distinct Scarabaeinae beetles. For the identification of species, the following works were mainly used: Nunes et al. (2018), Nunes et al. (2020). Nine species were identified: Canthon aff. histrio, C. aff. sericatus, C. aff. xanthopus, C. conformis Harold, 1868, C. fulgidus Redtenbacher, 1868, C. histrio (Lepeletier de Saint-Fargeau & Audinet-Serville, 1828), C. subhyalinus (Rivera- Cervants & Halffter, 1999), C. semiopacus Harold, 1868 and C. triangularis (Drury, 1770) (Fig. 2J-P). Genus Coprophanaeus d’ Olsoufieff, 1924 Notes: A Neotropical genus with approximately 51 known species (Schoolmeesters 2022), which are easily identified using the taxonomic keys published by Edmonds and Zidek (2010). This genus is known to be attracted to carcasses and be captured in flight intercept traps (Vaz-de-Mello 1999). Usually found in fresh carrion at dusk periods (Halffter and Matthews 1966). For the identification of species, the following works were mainly used: Edmonds and Zidek (2010). Three species were identified in our study: Coprophanaeus degallieri Arnaud, 1997, C. jasius (Olivier, 1789) and C. lancifer (Linnaeus, 1767) (Fig. 3A-C). Genus Cryptocanthon Balthasar, 1942 Notes: This genus comprises around 43 species occurring from Brazil to Mexico (Arias and Medina 2014, Martinez-Revelo et al. 2020, Giraldo-Mendonza 2022). The only available information about their habitat describes specimens inhabiting the leaf litter of humid and tropical forests, both in mountains and low altitudes (Cook 2002). For the Dung beetles from two sustainable-use protected forests in the Brazilian ... 11 identification of the species, the following works were mainly used: Cook (2002). Cryptocanthon campbellorum Howden, 1973 (Fig. 3D) was the only species, which is usually collected in leaf litter, with flight interception and pitfall traps baited with human faeces (Cook 2002). | ! ' E F G H 1 @. 1 | / I J L M \ | | 6 6 & vi | é Figure 3. EE Dorsal habitus of dung beetle species collected in the Tapajés National Forest and/or Carajas National Forest. A Coprophanaeus degallieri Arnaud, 1997; B Coprophanaeus jasius (Olivier, 1789); C Coprophanaeus lancifer (Linnaeus, 1767); D Cryptocanthon campbellorum Howden, 1973; E Deltochilum anceladus Kolbe, 1893; F Deltochilum orbiculare Lansberge, 1874; G Deltochilum orbignyi amazonicum Bates, 1887; H Dichotomius cuprinus (Felsche, 1901); | Dichotomius mamillatus (Felsche, 1901); J Dichotomius melzeri (Luederwaldt, 1922); L Dichotomius nisus (Olivier, 1789); M Dichotomius pelamon (Harold, 1869); N Dichotomius worontzowi (Pereira, 1942); O Eurysternus arnaudi Génier, 2009; P Eurysternus atrosericus Génier, 2009; Q Eurysternus balachowski Halffter & Halffter, 1977. Scale bar: 1 mm. Genus Deltochilum Eschscholtz, 1822 Notes: This is a very diverse genus of the Americas, with approximately 115 described species (Gonzalez-Alvarado and Vaz-de-Mello 2021). Deltochilum individuals are mostly nocturnal and often found in temperate, tropical and subtropical forests (Halffter and Matthews 1966). De/tohyboma is currently the most challenging subgenus, which 12 Carvalho E et al has been recently revised with several new species (Gonzalez-Alvarado and Vaz-de- Mello 2021). For the identification of the species, the following works were mainly used: Gonzalez-Alvarado and Vaz-de-Mello (2021). Seven species were identified through our study: Deltochilum enceladus Kolbe, 1893, D. gp. aspericolle, D. gp. guyanense, D. gp. sextuberculatum, D. orbiculare van Lansberge, 1874, D. orbignyi amazonicum Bates, 1887 and Deltochilum sp. 1 (Fig. 3E-G). Genus Dichotomius Hope, 1838 Notes: According to the most recent taxonomic revision from one of the subgenera of Dichotomius, this Neotropical genus comprises around 190 species widely distributed from the USA to Argentina (Nunes and Vaz-de-Mello 2019). The four subgenera are either being revised or were recently revised (Nunes et al. 2016, Valois et al. 2017, Maldaner et al. 2018, Nunes and Vaz-de-Mello 2019). Dichotomius species occur in all Brazilian biomes and can be collected in pastures, savannahs or forests. The genus as a whole is considered paracoprid — i.e. tunnellers (Nunes and Vaz-de-Mello 2019). For the identification of the species, the following works were mainly used: Nunes et al. (2016), Valois et al. (2017), Maldaner et al. (2018), Nunes and Vaz-de-Mello (2019). We collected nine species: Dichotomius aff. batesi, D. aff. lucasi 1, D. aff. lucasi 2, D. cuprinus (Felshe, 1901), D. mamillatus (Felshe, 1901), D. melzeri (Luederwaldt, 1922), D. nisus (Olivier, 1789), D. pelamon (Harold, 1869) and D. worontzowi (Pereira, 1942) (Fig. 3H-N). Genus Eurysternus Dalman, 1824 Notes: A Neotropical genus with 53 described species (Genier 2009) that are mostly endocoprids (Halffter and Matthews 1966, Cupello and Vaz-de-Mello 2018). Eurysternus species are easily collected in pitfalls baited with faeces, occurring in forests and frequently abundant in flooding-prone areas (Genier 2009). Genier (2009) was used for species identification. Thirteen species were recorded in our surveys: Eurysternus arnaudi Genier, 2009, E. atrosericus Genier, 2009, E. balachowskyi Halffter & Halffter, 1977, E. caribaeus (Herbst, 1789), E. cavatus Genier 2009, E. cayannensis Castelnau, 1840, E. cyclops Génier, 2009, E. fallaciosus Génier, 2009, E. foedus Guerin-Méneville, 1844, E. hamaticollis Balthasar, 1939, E. hypocrita Balthasar, 1939, E. plebejus Harold, 1880 and E. wittmemorum Martinez, 1988 (Fig. 30-Q, Fig. 4 A-J). Genus Eutrichilum Martinez, 1969 Notes: This genus presents a group of species that inhabit South American lowlands, east of the Andes as far south as Buenos Aires in Argentina; one species in Costa Rica (Vaz-De-Mello 2008). Species of this genus are frequently necrophagous and are often attracted to light (Vaz-De-Mello 2008). Eutrichillum sp. 1 was the only species recorded within the Tapajos region. Dung beetles from two sustainable-use protected forests in the Brazilian ... 13 Genus Hansreia Halffter & Martinez, 1977 Notes: This is an Amazonian genus with six species distributed across Brazil, French Guiana and Venezuela (Halffter and Martinez 1977); recently revised by Valois et al. (2015). Valois et al. (2015) was used for species identification. There is not much ecological information about Hansreia dung beetles (Hadara et al. 2020). Only the species Hansreia oxygona (Perty, 1830) was recorded within the Carajas region btEs e386 1260 686 Dorsal habitus of dung beetle species collected in the Tapajés National Forest and/or Carajas National Forest. A Eurysternus caribaeus (Herbst, 1789); B Eurysternus cavatus Génier, 2009; C Eurysternus cayennensis Castelnau, 1840; D Eurysternus cyclops Génier, 2009; E Eurysternus fallaciosus Génier, 2009; F Eurysternus foedus Guérin-Méneville, 1844; G Eurysternus hamaticollis Balthasar, 1939; HH Eurysternus hypocrita Balthasar, 1939; | Eurysternus plebejus Harold, 1880; J Eurysternus wittmerorum Martinez, 1988; L Hansreia oxygona (Perty, 1830); M /socopris imitator (Felsche, 1901); N /socopris nitidus (Luederwaldt, 1922): O Ontherus carinifrons Luederwaldt, 1930; P Onthophagus digitifer Boucomont, 1932; Q Onthophagus onthochromus Arrow, 1913. Scale bar: 1 mm. 14 Carvalho E et al Genus Isocopris Pereira e Martinez, 1960 Notes: This Neotropical genus, frequently misidentified as Dichotomius, comprises seven known species recently revised by Rossini and Vaz-de-Mello (2017), the same work being used to identify the species. No biological information for the genus was found. Two species were identified in our study: /socopris imitator (Felsche, 1901) and |. nitidus (Luederwaldt, 1922) (Fig. 4M-N). Genus Ontherus Erichson, 1847 Notes: Occurring from Argentina to Mexico, this genus has approximately 60 species (Genier 1996, Genier 1998). Although most species are considered coprophagous or saprophagous, some complex associations with ants have been previously recorded (Génier 1996). For the identification of the species, the following works were mainly used: Génier (1996). One species was found in the Tapajos region: Ontherus carinifrons Luederwaldt, 1930 (Fig. 40). This species belongs to a group called appendiculatus, which is widely distributed in South America (Genier 1996). Species from this group are usually collected in human or cattle dung, also using flight or light traps in sandy habitats (Genier 1996). Genus Onthophagus Latreille, 1802 Notes: Considered a megadiverse and cosmopolitan genus with approximately 2,000 described species (Tarasov and Kabakov 2010). Some species have been recently revised (Rossini et al. 2018a, Rossini et al. 2018b), while others are under current revision. The species mentioned here have been recorded mainly in primary and secondary forests, through the use of both flight interception traps and pitfalls baited with dung or carrion (Korasaki et al. 2012). For the identification of the species, the following works were mainly used: Rossini et al. (2018a), Rossini et al. (2018b). Four species were identified in our study: Onthophagus digitifer Boucomont, 1932, O. gp. rubrescens, O. onthochromus Arrow, 1913 and O. osculatii Guérin-Meéneville, 1855 (Fig. 5A). Genus Oxysternon Castelnau, 1840 Notes: This Neotropical genus comprises 11 species according to the last taxonomic revision by Edmonds and Zidek (2004) which was used for species identification. Oxysternon beetles are usually found in primary and secondary forests (Gigliotti et al. 2011). The literature on the biology of the genus is scarce, but most species are considered as coprophagous and inhabit moist forests (Edmonds and Zidek 2004). Two species were recorded in this study: Oxysternon macleayi Nevinson, 1892 and O. silenus Castelnau, 1840 (Fig. 5B-C). Dung beetles from two sustainable-use protected forests in the Brazilian ... 15 Genus Scybalocanthon Martinez, 1948 Notes: Scybalocanthon is a widespread genus occurring in South and Central America (Pereira and Martinez 1956, Silva and Valois 2019). The genus comprises 24 valid species, most of which are diurnal and inhabit either moist or dry forests in the Amazon Region, Atlantic Rainforest and the Yungas (Silva and Valois 2019, Silva and Genier 2019). Only the species Scybalocanthon sp. 1 was reported to the Carajas region. Genus Sulcophanaeus d'Olsoufieff, 1924 Notes: This Neotropical genus has approximately 15 described species (Edmonds 2000). Morelli et al. (1996)Noriega (2001) bring information about the life cycle of some Sulcophanaeus species. Edmonds (2000) was used for species identification. Sulcophanaeus faunus (Fabricius, 1775) was the only species recorded in this study (Fig. 5D). Figure 5. EE Dorsal habitus of dung beetle species collected in the Tapajés National Forest and/or Carajas National Forest. A Onthophagus osculatii Guérin-Méneville, 1855; B Oxysternon macleayi Nevison, 1892; C Oxysternon silenus Castelnau, 1840; D Sulcophanaeus faunus (Fabricius, 1775); E Silvycanthon candezei (Harold, 1869); F Sylvicanthon proseni (Martinez, 1949). Scale bar: 1 mm. Genus Sylvicanthon Halffter & Martinez, 1977 Notes: This genus has 15 species, broadly occurring in the Neotropics and was recently revised by Cupello and Vaz-de-Mello (2018). Some species are widespread in the Amazon Basin (Cupello and Vaz-de-Mello 2018). All known species are nocturnal, with most of them considered coprophagous feeding on primate, pig and cattle dung ( 16 Carvalho E et al Cupello and Vaz-de-Mello 2018). For the identification of the species, the following works were mainly used: Cupello and Vaz-de-Mello (2018). Two species were identified in our study: Sylvicanthon candezei (Harold, 1869) and S. proseni (Martinez, 1949) (Fig. 5E-F). Genus Uroxys Westwood, 1842 Notes: Uroxys is an exclusively Neotropical genus with more than 50 described species (Vaz-De-Mello 2008, Korasaki et al. 2012). This genus has species that can be found in grasslands and within primary and secondary forests; it also includes species specialised in sloth (Bradypodidae) dung (Korasaki et al. 2012). Uroxys cf. minutus was the only species reported in this study. Discussion The knowledge of biodiversity is key to providing information for conservation and management strategies, particularly within the hyperdiverse tropics (Barlow 2018). Our dung beetle surveys within FLONA Tapajos and Carajas highlight the importance of Amazonian Sustainable-Use protected forests for conserving insect biodiversity in the tropics. Protected areas have a key role as a thermal buffer against climate changes (Xu et al. 2022) and for the long-term maintenance of Amazonian biodiversity (Laurance 2005). However, Brazilian protected forests are largely underfunded, particularly in Amazonia (Silva et al. 2021) — which hinders their efficacy in protecting biodiversity and raises the urgency for new policies and funding mechanisms to enhance their efficacy. Acknowledgements We are grateful to all the farmers for collaborating to the Long-Term Ecological Research Project of the Sustainable Amazon Network (PELD-RAS); to Victor Hugo de Oliveira, Rodrigo Braga, Julio Louzada, Gilson de Oliveira, Marcos de Oliveira, Elivan Santos, Jaiane da Silva, Janitoni Lima, Renilson de Freitas and Josivan Oliveira for invaluable help during fieldwork and dung beetle identification; and to the Large Scale Biosphere- Atmosphere Program (INPA-LBA) and to Instituto Chico Mendes de Conservagao da Biodiversidade (ICMBio CR-03-Santareém and Carajas) for all provided support. FF acknowledges funding provided by [1] CNPq (National Council for Scientific and Technological Development) through the National Institute of Science and Technology (INCT) “Syntheses of Amazonian Biodiversity (CNPq/INCT 406767/2022-0 [SinBiAm)), PELD-RAS (CNPq-CAPES 441659/2016-0 and 441573/2020-7), SEM-FLAMA (CNPo- PrevFogo-IBAMA 441949/2018-5), and RESFLORA (MCIC-CNPq 420254/2018-8); and [2] NERC (UK Natural Environment Research Council (NERC, NE/P004512/1 [AFIRE]). EC thanks Coordenagao de Aperfeigcoamento de Pessoal de Nivel Superior - Brasil (CAPES) - Finance Code 001 and Fundacgao de Amparo a Pesquisa do Estado do Amazonas (FAPEAM) - POSGRAD/scholarship/ financial support. VCS thanks UNICAMP for providing the licence to use the software ArcGIS (process n° 20-P-21727/2021/Contract n° Dung beetles from two sustainable-use protected forests in the Brazilian ... 17 161/2021). We would like to thank all members of the Rede Amazénia Sustentavel (RAS Network) for their support, in particular Joice Ferreira, Jos Barlow, Erika Berenguer and Fernando Elias Silva for their commitment to establish and maintain the long-term research plots in Santarem and Carajas. Author contributions Fieldwork was conducted by FMF (2016, 2017 and 2019) and LFM (2016 and 2017). Dung beetles were sorted and mounted by CFGC, VOC, VPS, HS with key support from EC and MEM (2019). Beetle identification was conducted by LFM (2016, 2017) and MEM and EC (2019). FZVM revised and validated all dung beetle identifications (2010, 2016, 2017 and 2019). FF contributed for funding acquisition and supervision. All dung beetle photos were made by VCS. The manuscript was written by EC, MEM, FF and VCS. All authors gave final approval for publication. References ° Arias J, Medina C (2014) Tres nuevas especies de Cryptocanthon Balthasar, 1942 (Coleoptera: Scarabaeidae: Scarabaeinae) para Colombia. 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