Title: scielo.br_j_bn_a_n9K4yTRkN4DYyHYGzRCV4Xy__format=pdf&lang=en
Author: Richard Delahaye (LLE - Staff)

Biota Neotrop., 25(1): e20251770, 2025	

Dung beetles of Serra do Divisor National Park, Brazil

Biota Neotrop.. 25(1): e20251770. 2025

Bach A. et al.

Biota Neotrop.. 25(1): e2025 1770. 2025

Bach A. et al.

Biota Neotropica 25(1): e20251770, 2025
www.scielo.br/bn

biotaneotropica
ISSN 1676-0611 (online edition)
Inventory
Dung beetles (Coleoptera: Scarabaeidae: Scarabaeinae) of Serra do
Divisor National Park, Brazil
Andressa Bach^1·^2*®, Torbjørn Haugaasen®, Carlos A. Peres^34 , Emanuelly F. Lucena^5®,
José Victor A. Ferreira , Joseph E. Hawes^3^ , Bruna R. Bordin^2 , Mario Cupello” ,
Renato A. de Azevedo^9 & Fernando Z. Vaz-de-Mello^2®
‘Norwegian University of Life Sciences, Faculty of Environmental Sciences and Natural Resource Management,
As, Norway.
“Universidade Federal de Mato Grosso, Instituto de Biocièncias, Laboratorio de Scarabaeoidologia,
Cuiaba, MT, Brasil.
‘University of East Anglia, School of Environmental Sciences, Nonvich, NR4 7TJ, United Kingdom.
‘Instituto Juruå, 69083-300 Manaus, AM, Brazil.
'Universidade Federal da Paraiba, Centro de Cièncias Exatas e da Natureza, Departamento de Sistemàtica e
Ecologia, Laboratòrio de Termitologia, Joào Pessoa, PB, Brasil.
^6 Universidade Estadual de Santa Cruz, Programa de Pós- Graduaςào em Ecologia e Conservaςào da
Biodiversidade, Ilhéus, BA. Brasil.
'University of Cumbria, Institute of Science and Environment, LA22 9BB, Ambleside, Cumbria, United Kingdom.
‘Texas A&M University, Department of Entomology, 77843-2475, College Station, Texas, USA.
‘Instituto Nacional de Pesquisas da Amazonia, Manaus, AM, Brasil.
*Corresponding author: andressabach04@gmail.com
BACH, A., HAUGAASEN, T., PERES, C.A., LUCENA, E.F., FERREIRA, J.V.A., HAWES, J.E.,
BORDIN, B.R., CUPELLO, M., AZEVEDO, R.A., VAZ-DE-MELLO, F.Z. Dung beetles (Coleoptera: Scarabaeidae: Scarabaeinae) of Serra do Divisor National Park, Brazil. Biota Neotropica 25(1): e20251770.
https://doi.org/10.1590/1676-0611-BN-2025-1770
Abstract: Serra do Divisor National Park (SDNP) is a protected area located in the western Brazilian Amazon that is threatened by a pending bill proposing to downgrade its status and to build a road that would effectively slice the park in half. The biodiversity of SDNP is poorly understood and no dung beetle inventories have been conducted within the park to date, despite being one of the most studied insect groups in the Neotropics. We sampled dung beetles with flight interception traps (FITs) and pitfall traps baited with human faeces to provide the first comprehensive dung beetle species list for SDNP. In total, we collected 4,909 dung beetle individuals belonging to 83 species. Of these, 18 were new species records for the state of Acre and two for Brazil. We also found species with Andean affinity and five species new to science. This study contributes to our understanding of dung beetle assemblages in the western Brazilian Amazon, showing that SDNP is a unique locality due to the proximity of the Andes, and highlighting that biodiversity in many Amazonian protected areas remains severely under sampled.
Keywords: Amazon; Flight interception traps; Pitfall; Species list.
Os besouros rola-bostas (Coleoptera: Scarabaeidae: Scarabaeinae) do Parque
Nacional da Serra do Divisor, Brasil
Resunio: 0 Parque Nacional da Serra do Divisor (PNSD) é uma unidade de conservano localizada no sudoeste da Amazonia brasileira. 0 parque é extremamente biodiverso, mas està amea^ado por um projeto de lei que propòe diminuir sua categorizaςào para Àrea de Protendo Ambientai (APA) e construir uma rodovia que cortaria ο parque ao meio. Contudo, a biodiversidade do PNSD ainda é pouco estudada. Um exemplo disso éque nenhum inventàrio de besouros rola-bostas havia sido realizado nesta àrea até entào, mesmo tratando- se de um dos grupos de insetos mais estudados na regiào Neotropical. Os besouros foram amostrados usando armadilhas de intercepta’ de voo (Flight Interception Traps - FIT) e armadilhas pitfall iscadas com fezes humanas para conipor a primeira lista de espécies de besouros rola-bostas do PNSD. Nós coletamos um total de 4.909 individuos de 83 espécies, sendo 18 espécies novos registros de ocorréncia para ο estado do Acre e duas para ο Brasil. Tambérn capturamos espécies com afinidade Andina e cinco espécies novas para a ciència. Este traballio amplia ο conhecimento atual das assembleias de rola-bostas no sudoeste amazónico, mostra que PNSD é urna localidade singular devido à sua proximidade coni os Andes e evidencia que muitas areas de proteqào na Amazonia ainda sào pouco estudadas.
Palavras-chave: Amazonia; Armadilha de interceptagào de νοο; Lista de espécies; Pitfall.

Introduction
The Neotropical region is estimated to host at least a third of the world’s biodiversity (Raven et al. 2020), with the majority found in the Amazon rainforest (Antonelli et al. 2018). However, this high- biodiversity region is under immense threat from deforestation (Hoang & Kanemoto 2021) largely driven by agribusiness expansion (Pereira et al. 2020, Seymour & Harris 2019). Protected Areas (PAs) have therefore become increasingly important to mitigate deforestation and safeguard biodiversity (Soares-Filho et al. 2010, Barber et al. 2014, Goncalves-Souza et al. 2021). In fact, 47% of Amazonia lies within a PA (IUCN categories I - VI), collectively spanning 198 million hectares (Soares-Filho et al. 2023) and deforestation rates are much lower inside PAs than outside (Qin et al. 2022).
Serra do Divisor National Park (SDNP) is an ~0.84 Mha PA located in the extreme western Brazilian Amazonia in the state of Acre, adjacent to the border with Peru (Brazil 1989, 2000, ISA 2023). SDNP and its contiguous sister-park (the ~ 1.35 Mha Sierra del Divisor National Park) in Peru, form a substantial transnational biological corridor spanning more than two million hectares across the Brazil-Peru border (MMA 2024, SERNANP 2024). SDNP is extremely biodiverse, hosting at least 43 species of large mammals and more than 100 amphibian, 30 reptile, 480 bird, and 55 bat species (UNESCO 2024). Less than 1% of the park’s primary forest has been altered since its creation (Koga et al. 2022) and SDNP is therefore an important refuge for biodiversity including many species of high conservation value (Gonzalez & Acuy 2017, MMA 2024).
Despite its importance, SDNP is under increasing pressure (Ruaro & Laurance 2021, Koga et al. 2022). Tire PA is currently threatened by a pending bill (n. 6024/2019) that proposes to downgrade its status to an Environmental Protection Area (Area de Ρηοίεςάο Ambientai in Portuguese - APA) and extend the BR-364 highway through its territory (Brazil 2019). These proposed actions would allow activities prohibited within a National Park, leading to economic exploitation, accelerated deforestation and habitat degradation with significant environmental and societal impacts (Ruaro & Laurance 2021, Koga et al. 2022). The highway could also intensify illegal logging, promote illegal settlements, mining, and hunting, and benefit drug trafficking and criminal factions (Laurance & Balmford 2013, Faleiro 2023). For a map of the proposed highway through the contiguous PAs in Brazil and Peru, refer to Koga et al. (2022).
Research on the biodiversity of SDNP has increased in recent years, particularly for vertebrates (Calouro 1999, Lopes & Rehg 2003, Whitney et al. 2004, Lemos et al. 2018, Almeida et al. 2022b) and insects including butterflies (Dolibaina et al. 2012, 2015, 2016, Kaminski 2020), leafhoppers (Creào-Duarte et al. 2023), wasps (Azevedo et al. 2002, Morato et al. 2008), and termites (Ferreira et al. 2023). However, invertebrate biodiversity inventories generally remain insufficient, and, to the best of our knowledge, no comprehensive inventory of dung beetles (Coleoptera: Scarabaeidae: Scarabaeinae) has been conducted in the park, even though dung beetles are among the most studied insect groups in the Neotropics, especially South America (Arellano et al. 2023, 2024).
The Neotropical realm harbours the greatest dung beetle diversity and richness in the world (Favila et al. 2023), with Brazil alone hosting at least 787 species (Vaz-de-Mello & Bordin 2024). These beetles portray a wide range of feeding habits which have undoubtedly contributed to their high species richness in this region (Gill 1991). While dung beetles are commonly known for being coprophagous, feeding mainly on mammal faeces (Filgueiras et al. 2009, Tsuji et al. 2021), there are also species that are partially or strictly necrophagous (feeding on carrion) or saprophagous (consuming decaying fruit or fungi; Halffter & Matthews 1966, Halffter & Halffter 2009), and a few may even be predators (Vaz-de-Mello et al. 1998, Oliveira et al. 2022).
Many dung beetles are known for their reproductive and feeding behaviour, which involves rolling some of the food and building nests underground (Cambefort & Hanski 1991). Although this behaviour varies among species (Halffter & Edmonds 1982), dung beetles play a crucial role in nutrient recycling and increasing soil nutrient concentrations (e.g. including phosphorus, ammonia, and nitrogen; Maldonado et al. 2019) by burying decaying organic matter. They also contribute to soil aeration, secondary seed dispersal, and parasite suppression (Louzada 2008, Nichols et al. 2008, Almeida et al. 2022a). Furthermore, dung beetles are a high-performance group in biodiversity assessment and monitoring (Gardner et al. 2008) and can be used to evaluate changes in the habitat integrity (Verdù et al. 2011, Viegas et al. 2014, Carvalho et al. 2020, Noriega et al. 2021).
In this study, we present a checklist of dung beetle species from Serra do Divisor National Park in the state of Acre, Brazil. This is a fundamental first step in monitoring biodiversity and establishing further conservation practices (Jones et al. 2013). By providing information on the occurrence of dung beetle species, we contribute to the broader understanding of biodiversity in this region. We also discuss our findings in relation to the park’s geographic location, sampling methods used, and the proposed actions of bill 6024/2019.
Material and Methods
1.		Study area
Serra do Divisor National Park (SDNP) is situated in the upper Juruà River basin in the state of Acre, westernmost Brazilian Amazonia. Encompassing an area of 837,554 ha, the park spans five municipalities: Cruzeiro do Sul, Måncio Lima, Marechal Thaumaturgo, Porto Walter, and Rodrigues Alves (MMA 2024). The average elevation is 300 m, and
the main primary vegetation is open ombrophilous forest, accounting for 84.99% of the park, followed by dense ombrophilous forest, making up the remaining 15.01% (ISA 2023). Annual precipitation in the park is 2,200 mm, with average temperatures ranging between 23-26°C throughout the dry and rainy seasons (ISA 2023).
2.		Sampling design
Dung beetle sampling was carried out during December 2019 in terra firme forest sites in the northern sector of SDNP as part of our 3rd Amazon Biodiversity & Carbon (ABC) expedition (www. abc-expeditions.com/). The permanent license for zoological material collection was authorized by Ministerio do Meio Ambiente (MMA), Instituto Chico Mendes de Conservando e Biodiversidade (ICMBio), and Sistema de Autorizanào e Informando em Biodiversidade (SISBIO) process η. 72874, on November 4, 2019.
Sampling was conducted along three 1-kilometer transects referred to as plot A (07°24’44”S 73°39’49” W), B (07°29'26"S 73°39’06” W), and C (07°25‘46"S 73°37’39” W; Figure 1). In each plot, four- flight interception traps (FIT) of 2 x 1.5 m were installed 250 m apart and left open for 12 days (the spatial configuration of sampling along eaclr plot is shown in Figure 1). Samples were collected every four- days, resulting in a total of 12 FIT samples per plot. Additionally, for each of the same transects, 10 pitfall traps baited witli human faeces were installed 100 m apart, filled with a mixture of water, salt, and detergent. Pitfall traps were left open for 48 hours and resulted in a total of 10 samples per plot (Larsen & Forsyth 2005, Da Silva and Hernandez 2015; Mora-Aguilar et al. 2023).
3.		Dung beetle identification
Collected specimens were fixed with 96% alcohol in the field and later deposited in Colendo Entomològica de Mato Grosso Eurides Furtado (CEMT), at Universidade Federal de Mato Grosso, Cuiaba, state of Mato Grosso, Brazil. Dung beetle identification was achieved using taxonomic bibliography and the identification keys therein (Génier 1996, Cook 1998, 2000, Edmonds & Zidek 2004, Génier 2009, Edmonds & Zidek 2010, Vaz-de-Mello et al. 2011, Edmonds & Zidek 2012, Génier & Arnaud 2016, Rossini & Vaz-de-Mello 2017, Cupello & Vaz-de-Mello 2018, Rossini et al. 2018, Silva & Valois 2019, Rossini & Vaz-de-Mello 2020), and by comparing with specimens previously identified and deposited at CEMT. When species identification was not possible, specimens were sorted based on external morphology.

Figure 1. Serra do Divisor National Park (SDNP), state of Acre, westernmost Brazilian Amazon, where dung beetles were sampled in plots A, B, and C (inset square) in the northern portion of the park using flight interception traps and pitfall traps baited with human faeces. The spatial configuration of sampling along each plot is also shown.

-/. Data analysis
To assess sampling efficiency, we performed individual-based rarefaction curves of species richness at plot-level (q = 0; Chao et al. 2014) and calculated sample coverage using the ‘iNEXT’ package (Hsieh & Chao 2024) and ‘ggplot2’ (Wickham 2016) in R (R Core Team 2023). To visualize species relative abundance sampled by FITs and pitfall traps we plotted rank abundance curves using the 'BiodiversityR' package (Kindt & Coe 2005).

Results
Tn total, we collected 4,909 dung beetle individuals from 83 species (Table 1): 50 were identified to species level (60.24%) and 33 were not associated to existing species names (39.76%). Among the 20 genera captured, Canthidium was the most species-rich with 12 species and 361 individuals, followed by Ateuchus with 10 species and 139 individuals, and Eurysternus with nine species and 1,171 individuals. The most abundant species was Onthophagus rubrescens

Table 1.List of 83 dung beetle species from Serra do Divisor National Park (SDNP), sampled with flight interception traps (FIT) and pitfall traps baited with human faeces (PT). Literature citations are added when the species have been previously recorded for the state of Acre. * = first species record for Acre; ** = first species record for Brazil; - = no information found; Ν = number of individuals captured in our study.
Species | Capture method | Literature records | Site | Ν
Anomiopus pictus (Harold, 1862) | FIT | * | SDNP | 1
Anomiopus sp. 1 | FIT | - | SDNP | 3
Anomiopus sp. 2 | FIT | - | SDNP | 1
Ateuchus aeneomicans (Harold, 1868) | FIT; PT | * | SDNP | 16
Ateuchus aff. murrayi | FIT; PT | - | SDNP | 100
Ateuchus aff. ovalis | FIT | - | SDNP | 2
Ateuchus aff. pygidialis | PT | - | SDNP | 1
Ateuchus cereus (Harold, 1868) | FIT; PT | * | SDNP | 4
Ateuchus columbianus (Harold, 1868) | FIT; PT | ** | SDNP | 4
Ateuchus connexus (Harold, 1868) | FIT | * | SDNP | 4
Ateuchus globulus (Boucomont, 1928) | FIT | * | SDNP | 3
Ateuchus murrayi (Harold, 1868) | FIT; PT | * | SDNP | 4
Ateuchus substriatus (Harold, 1868) | PT | * | SDNP | 1
Bdelyrus sp. 1 | FIT | - | SDNP | 1
Canthidium aff. dohrni | FIT; PT | - | SDNP | 5
Canthidium aff. fùnebre | PT | - | SDNP | 4
Canthidium aff. lentum | FIT; PT | - | SDNP | 16
Canthidium aff. melanocephalum | FIT; PT | - | SDNP | 5
Canthidium aff. onthophagoides | FIT | - | SDNP | 4
Canthidium aff. orbiculatum | FIT; PT | - | SDNP | 132
Canthidium aff. rufinum | PT | - | SDNP | 2
Canthidium sp. 1 | PT | - | SDNP | 1
Canthidium sp. 2 | PT | - | SDNP | 4
Canthidium sp. 3 | FIT | - | SDNP | 1
Canthidium sp. 4 | FIT; PT | - | SDNP | 77
Canthidium sp. 5 | FIT; PT | - | SDNP | 10
Canthon aff. angustatus | PT | - | SDNP | 78
Canthon brunneus Schmidt, 1922 | PT | * | SDNP | 15
Canthon luteicollis Erichson, 1847 | FIT; PT | Vaz-de-Mello, 1999 | SDNP; Zoobotànico Park - Universidade Federal do Acre | 142
Continue...

...Continuation
Species | Capture method | Literature records | Site | N
Canthon quadrimaculatus Schmidt 1922 | PT | * | SDNP | I
Canthon rufocoeruleus Martinez, 1948 | PT | * | SDNP | 5

Canthon semiopacus Harold, 1868

PT baited witli human faeces or carrion (bovine liver)

Bitencourt et al. 2019a;
Bitencourt et al. 2019b

SDNP; Senador Guiomard, Catuaba Experimental Reserve; Antimary State Forest

3

Canthon xanthopus Blanchard 1846 | FIT; PT baited with human faeces or carrion (chicken heart) | Bitencourt et al. 2019b | SDNP; Floresta Estadual do Antimary | 8
Canthonella sp. 1 | FIT | - | SDNP | 2
Canthonella sp. 2 | PT | - | SDNP | 1
Coprophanaeus sp. 1 | FIT | - | SDNP | 3

Coprophanaeus telamon Erichson 1847

FIT; PT baited with human faeces or carrion (chicken heart or bovine liver)

Bitencourt et al. 2019a;
Bitencourt et al. 2019b;
Cupello & Vaz-de-Mello, 2013; Edmonds & Zidek, 2010; Vaz-de-Mello, 1999

SDNP; Senador Guiomard, Catuaba Experimental Reserve; Antimary State Forest; Xapuri, Chico Mendes Reserve; Zoobotànico
Park - Universidade Federal do Acre

34

Cryptocanthon peckorum Howden 1973

FIT; pan traps; PT baited with human faeces or carcass

Cook, 2002

SDNP; Rio Branco, Catuaba Experimental
Reserve

12

Deltochilum aff. carinatum | FIT; PT | - | SDNP | 4
Deltochilum aff. guyanense | FIT; PT | - | SDNP | 16
Deltochilum gr. aspericolle | FIT; PT | - | SDNP | 37
Deltochilum gr. barbipes | FIT; PT | - | SDNP | 21
Deltochilum gr. femorale | FIT | - | SDNP | 4

Deltochilum orbiculare Lansberge 1874

FIT; PT baited with human faeces or carrion (bovine liver)

Bitencourt et al. 2019a;
Bitencourt et al. 2019b

SDNP; Senador Guiomard, Catuaba Experimental Reserve; Antimary State Forest

15

Deltochilum orbignyi amazonicum Bates, 1887 | FIT; PT baited with human faeces or carrion (chicken heart) | Bitencourt et al. 2019b | SDNP; Antimary State Forest | 37

Deltochilum scheffierorum Silva, Louzada
& Vaz-de-Mello, 2015

FIT

*

SDNP

1

Dendropaemon lydiae Génier & Arnaud 2016 | FIT | Génier & Arnaud, 2016 | SDNP; Zoobotànico Park - Universidade Federal do Acre | 1
Dichotomius aff. lucasi | FIT; PT | - | SDNP | 18
Dichotomius divisor Moura et al. 2025 | FIT; PT | - | SDNP | 70

Dichotomius batesi (Harold, 1869)

FIT; PT

Valois, Vaz-de-Mello &
Silva, 2023

SDNP; Xapuri, Chico Mendes Reserve

141

Dichotomius gandinii Rossini &
Vaz-de-Mello, 2020

PT baited with human faeces or fish carrion

Rossini & Vaz-de-Mello, 2020

SDNP; Xapuri, Chico Mendes Reserve;
Zoobotànico
Park - Universidade Federal do Acre

2

Continue...

...Continuation
Species | Capture method | Literature records | Site | N

Dichotomius mamillatus (Felsche, 1901)

FIT; PT baited with human faeces or carrion (chicken heart or bovine liver)

Bitencourt et al. 2019a;
Bitencourt et al. 2019b;
Vaz-de-Mello, 1999

SDNP; Senador Guiomard, Catuaba
Experimental Reserve; Antimary State Forest; Xapuri,
Chico Mendes Reserve; Zoobotànico Park - Universidade
Federal do Acre

9

Dichotomius ohausi (Luederwaldt, 1923)

FIT; PT baited with human faeces or carrion (chicken heart or bovine liver)

Bitencourt et al. 2019a;
Bitencourt et al. 2019b;
Vaz-de-Mello, 1999

SDNP; Senador Guiomard, Catuaba Experimental Reserve; Antimary State Forest; Zoobotànico Park - Universidade Federal do Acre

2

Dichotomius mhustus (Luederwaldt, 1935)

PT baited with human faeces or carrion (chicken heart)

Bitencourt et al. 2019b;
Vaz-de-Mello. 1999

SDNP; Antimary State Forest; Zoobotànico Park - Universidade Federal do Acre

3

Eurysternus caribaeus (Herbst, 1789)

FIT; PT baited with several mammal faeces (including human); several carcasses; occasionally in light traps

Bitencourt et al. 2019a;
Bitencourt et al. 2019b;
Génier, 2009; Vaz-de-
Mello, 1999

SDNP; Senador
Guiomard, Catuaba Experimental Reserve; Antimary State Forest;
Cruzeiro do Sul;
Zoobotànico
Park - Universidade Federal do Acre

438

Eurysternus cayennensis Castelnau, 1840 | FIT; PT | * | SDNP | 172

Eurysternus foedus Guérin-Méneville, 1844

FIT; PT baited with several mammal faeces (including humans’) or carcasses (as rotten bovine liver); rarely in light traps

Bitencourt et al. 2019a;
Génier, 2009

SDNP; Senador Guiomard, Catuaba Experimental Reserve;
Cruzeiro do Sul

10

Eurysternus hamaticollis Balthasar, 1939 | PT | * | SDNP | 9
Eurysternus hypocrita Balthasar, 1939 | FIT; PT baited witli human faeces or carrion (chicken heart) | Bitencourt et al. 2019b | SDNP; Antimary State Forest | 226
Eurysternus plebejus Harold, 1880 | FIT; PT baited witli human faeces | * | SDNP | 53
Eurysternus strigilatus Génier, 2009 | FIT; PT baited with human faeces or carrion (chicken heart) | Bitencourt et al. 2019b | SDNP; Antimary State Forest | 68
Eurysternus ventricosus Gill 1990 | PT | * | SDNP | 10

Eurysternus wittmerorum Martinez 1988

FIT; PT baited with human faeces, carrion (chicken heart), fish faeces or carcasses (especially snail carcasses)

Bitencourt et al. 2019b, Génier, 2009

SDNP; Antimary State
Forest, Feijó

185

Isocopris rossinii Arias-Buriticå et al. 2023 | FIT; PT baited with human faeces | Arias-Buriticå et al. 2023 | SDNP | 7
Ontherus aff. alexis | PT | - | SDNP | 38
Continue...

...Continuation
Species | Capture method | Literature records | Site | Ν
Ontherus raptor Génier, 1996 | FIT | ** | SDNP | 1
Onthophagus digitifer Boucomont 1932 | PT | * | SDNP | 4
Onthophagus onorei Zunino & Halffter 1997 | PT | * | SDNP | 262

Onthophagus osculata Guérin-Méneville, 1855

FIT; PT baited with human faeces or carcass

Rossini, Vaz-de-Mello &
Zunino, 2018

SDNP; Xapuri, Chico Mendes Reserve

370

Onthophagus rubrescens Blanchard, 1846 | FIT; PT baited with human faeces | * | SDNP | 1457

Onthophagus xanthomerus Bates 1887

FIT; PT baited with human faeces or carrion (bovine liver)

Bitencourt et al. 2019a

SDNP; Senador Guiomard, Catuaba
Experimental
Reserve

17

Oxysternon conspicillatum (Weber, 1801)

FIT; PT baited witli human faeces or carrion (bovine liver)

Bitencourt et al. 2019a;
Vaz-de-Mello, 1999

SDNP; Senador Guiomard, Catuaba Experimental Reserve;
Zoobotànico Park - Universidade Federal do Acre

36

Oxysternon silenus Castelnau, 1840

FIT; PT baited with human faeces or carrion (bovine liver)

Bitencourt et al. 2019a

SDNP; Senador Guiomard, Catuaba
Experimental
Reserve

25

Phanaeus hispinus Bates, 1868 | FIT; PT baited witli human faeces or carrion (bovine liver) | Bitencourt et al. 2019a | SDNP; Senador Guiomard, Catuaba Experimental Reserve | 4
Phanaeus cambeforti Arnaud, 1982 | FIT; PT baited witli human faeces or carrion (chicken heart) | Bitencourt et al. 2019b | SDNP; Antimary State Forest | 23
Phanaeus chalcomelas (Perty, 1830) | FIT; PT baited with human faeces or carrion (bovine liver) | Bitencourt et al. 2019a | SDNP; Senador Guiomard, Catuaba Experimental Reserve | 45
Phanaeus sp. 1 | FIT; PT baited with human faeces | - | SDNP | 20
Scybalocanthon aereus (Schmidt, 1922) | FIT; PT baited with human faeces or carcass | Silva & Valois, 2019 | SDNP; Màncio Lima; Xapuri, Chico Mendes Reserve | 1

Scybalocanthon uniplagiatus
(Schmidt, 1922)

FIT; PT baited with human faeces or fungi

Silva & Valois, 2019

SDNP; Rio
Branco,Catuaba Experimental Reserve

27

Sylvicanthon bridarolhi (Martinez, 1949) | FIT; PT baited with human faeces or carrion (bovine liver) | Bitencourt et al. 2019a | SDNP; Senador Guiomard, Catuaba Experimental Reserve | 12
Sylvicanthon proseni (Martinez 1949) | FIT; PT baited with human faeces, human+pig dung, cow dung, cow spleen, banana, chicken meat or rotten bovine liver; black light traps | Bitencourt et al. 2019a; Cupello & Vaz-de-Mello, 2018 | SDNP; Senador Guiomard, Catuaba Experimental Reserve; Màncio Lima; Manoel Urbano, Chandless State Park Xapuri, Chico Mendes Reserve | 235
Uroxys sp. 1 | FIT; PT | - | SDNP | 62
Uroxys sp. 2 | FIT | - | SDNP | 1

Figure 2. Individual-based rarefaction curves for dung beetle species sampled by both pitfall traps and FITs in plots A. B. and C at Serra do Divisor National Park (SDNP) in the state of Acre, Brazil. Shaded areas represent the 95% confidence intervals of each rarefaction curve.

Blanchard, 1846 with 1,457 individuals, followed by Eurysternus caribaeus (Herbst, 1789) with 438 individuals and Onthophagus osculata Guérin-Méneville, 1855 with 370 individuals.
We sampled 1,438 individuals and 51 species in plot A (mean ± standard deviation of individuals per species: 34.65 ± 21.81); 1,516 individuals and 62 species in plot B (36.53 ± 17.11); and 1,955 individuals and 57 species in plot C (47.11 ± 27.89), with a mean sample coverage of 99.27% among plots (Supplementary Material, Table Sl). Rarefaction curves also confirmed that our sampling effort was reasonable, although a few additional species may have been added witli continued sampling (Figure 2). Eighteen species (21.69%) were captured solely by FITs, 20 species (24.1%) were captured only by pitfall traps, and 45 species (54.22%) were captured by both methods. Pitfall traps captured 4,415 individuals (89.94%; mean ± standard deviation of individuals per species: 106.39 ± 6.87) while FITs sampled only 494 individuals (10.06%; 11.9 ± 2.14) (Supplementary Material, Figure Sl).
Our expedition uncovered 18 new dung beetle species records for the state of Acre: Anomiopus pictus (Harold, 1862), Ateuchus aeneomicans (Harold, 1868), Ateuchus cereus (Harold, 1868), Ateuchus connexus (Harold, 1868), Ateuchus globulus (Boucomont, 1928), Ateuchus murrayi (Harold, 1868), Ateuchus substriatus (Harold, 1868), Canthon brunneus Schmidt, 1922, Canthon quadrimaculatus Schmidt, 1922, Canthon rufocoeruleus (Martinez, 1948), Deltochilum schefflerorum Silva, Louzada & Vaz-de-Mello, 2015, Eurysternus cayennensis Castelnau, 1840, Eurysternus hamaticollis Balthasar. 1939, Eurysternus plebejus Harold, 1880, Eurysternus ventricosus Gill 1990, Onthophagus digitifer Boucomont, 1932, Onthophagus onorei Zunino & Halffter 1997, and Onthophagus rubrescens Blanchard, 1845; and two new records for Brazil: Ateuchus columbianus (Harold, 1868) and Ontherus raptor Génier, 1996 (Figures 3-4). Eurysternus hamaticollis was reported for state of Acre by Schoolmeesters (2024) without reporting the source of the data, thus we confirm the occurrence of this species in the state.
In addition, we collected five dung beetle species new to science in SDNP: Isocopris rossinii (already described by Arias-Buriticå et al. 2023), Phanaeus sp. 1 (FZVM & BRB pers, obs.), Ateuchus aff. pygidialis, Ateuchus aff. murrayi, and Ateuchus aff. ovalis (MC pers. obs.).
Discussion
This study presents the first list of dung beetle species from Serra do Divisor National Park. We documented 83 species using flight interception traps and pitfall traps baited with human faeces, of which 18 were unrecorded for the state of Acre, two were new records to Brazil, and five were new to science.
SDNP lies in the transition zone between the Ucayali and Rondonia biogeographical provinces (Μοιτοηβ et al. 2022), on the border witli Peru, and relatively close to the Andean piedmonts within pre-Andean Amazonia (ISA 2023). Indeed, the presence of two dung beetle species that were only previously captured in the Andes (Onthe. aff. alexis and Onthe. raptor) underlines this proximity. We could not unequivocally confirm the identification of the former as Onthe. alexis. However, this species is typically reported in the Andean region at elevations ranging from 1,100 to 2,000 m in Bolivia and over 730 m in Colombia and Peru (Génier 1996, Larsen 2004, Pulido-Herrera et al. 2007, Hamel-Leigue et al. 2008). This study presents the first record of Onthe. raptor in Brazil, previously found at 1,000 m in Bolivia (Génier 1996) and on the eastern slopes of the Peruvian Andes up to 3,200 m (Larsen et al. 2006). All these previous records are at elevations substantially higher than the elevations reached within SDNP (max = 650 m; mean = 300 m; ISA 2023). This supports the notion that SDNP is under some Andean
influence. We also obtained the first record of At. columbianus for Brazil, a species that to date has only been reported from Colombia and Peru (COL 2024).
The geographic location of SDNP may explain the high number of species recorded in the current study, as there is a high likelihood of recording elements from both biogeographical provinces (Morrone 2009). Indeed, a study from the middle section of the Jurua River, (~790 km downstream on a straight line from SDNP), detected 83 species in terra firme forest, of which only 21 species also occurred at our study site despite sampling the same river basin and interfluvial region (i.e. Juruå-Javari; Nichols et al. 2013). Other studies from Acre have registered between 36 to 58 species and have only been conducted in the eastern parts of the state (Vaz-de-Mello 1999, Bitencourt et al. 2019a, b). However, the high species richness may also be due to our use of two sampling techniques. The high proportion of species (45.79%) that we sampled exclusively by either FITs or pitfall traps shows that these two methods are complementary (Andrade et al. 2011, Mora-Aguilar et al. 2023).

Figure 3. New dung beetle species records for the state of Acre (*) and Brazil (**) sampled during our ABC Expedition to Serra do Divisor National Park, Brazil. The bars indicate a 2 mm scale. A) Anomiopus pictus*; B) Ateuchus aeneomiccms*; C) Ateuchus cereus*; D) Ateuchus columbianus**; Ε) Ateuchus connexus*; F) Ateuchus globulus*; G) Ateuchus murrayi*; Η) Ateuchus substriatus*; I) Canthon brunneus*.

In addition to the new species records, we also confirmed at least five dung beetle species new to science in SDNP: 1. rossinii, Phanaeus sp. 1, At. aff. pygidialis, At. aff. murrayi, and At. aff. ovalis. The first species has already been described by Arias-Buriticå et al. (2023), the second is part of the chalcomelas species group currently undergoing revision (FZVM & BRB unpubl. data), and the last three are within a genus that is also under revision (Cupello et al. 2023). These three Ateuchus species were already known from collections elsewhere in the Amazon, but the other two species were sampled for the first time by our ABC Expedition to SDNP.

Figure 4. New dung beetle species records for the state of Acre (*) and Brazil (**) sampled during our ABC Expedition to Serra do Divisor National Park, Brazil. The bars indicate a 2 mm scale. A) Canthon rufocoeruleus*; B) Canthon quadrimaculatus*; C) Deltochilum schefflerorum*; D) Isocopris rossinii (new species sampled in our expedition and described by Arias-Buriticå et al. 2023); E)Eurysternus cayannensis*; F) Eurysternus hamaticollis*'. G) Eurysternus plebejus*·, H) Eurysternus ventricosus*; I) Ontherus raptor**; J) Onthophagus digitifer*; K) Onthophagus onorei*; L) Onthophagus rubrescens*.

A new species of Isocopris was surprising because this genus was recently reviewed by Rossini & Vaz-de-Mello (2017), making it a focus of intense research across several scientific collections. Isocopris rossinii has also been sampled within the sister-park of SDNP and the Peruvian Amazon (Sierra del Divisor National Park; Arias-Buriticå et al. 2024). Its geographical distribution therefore ranges from SDNP to the slopes of the Andes. A new species of Phanaeus was similarly unexpected since they are large (10-28 mm) and well-represented in scientific collections due to their cephalic horns and iridescent metallic colors (Edmonds 1994, Hamel-Leigue et al. 2009, Edmonds & Zidek 2012).
Our results support the idea that SDNP is a particularly high- biodiversity area within the Amazon, and we are deeply concerned about the consequences of actions proposed by bill 6024/2019. These actions are part of an approach that threatens PAs called ‘protected areas downsizing, downgrading and degazetting’ (PADDD; Bernard et al. 2014, Ruaro & Laurance 2021). Construction of the BR-364 highway would result in ‘downsizing’ by effectively slicing SDNP in half and reducing its official size (Mascia & Paille 2010). The construction of roads in forested areas generally leads to ecosystem impacts like habitat loss and fragmentation, and biodiversity impacts like traffic mortality and stress (Robinson et al. 2010). Regarding dung beetles, Carpio et al. (2009) recorded that rare species vanished close to a new road in the Ecuadorian Amazon, althougli there is a shortage of studies that investigate these effects on dung beetles. Furthermore, the bill proposes to ‘downgrade' the SDNP status of protection from a National Park to an Environmental Protection Area, flexing several legal restrictions on human activities (Brazil 2019). Both actions occur prior to ‘degazetting’ process, which entails the complete removal of protection from an area (Mascia & Pailler 2010). This would clearly have drastic consequences for the biodiversity of this region.
Conclusions
This study contributes to our understanding of dung beetle assemblages in the westernmost Brazilian Amazon, shows that SDNP is under Andean influence due to its proximity to these mountainous regions, and highlights that the insect fauna in many protected areas in the Amazon remains poorly known. This lack of knowledge hampers conservation actions and makes it difficult to understand the impacts of human disturbance and encroachment in remote and relatively pristine regions such as SDNP. We therefore advocate for the continued protection of SDNP and are critical of the proposed road construction and plans to downgrade the park to an Environmental Protection Area (bill 6024/2019).
Supplementary Material
The following online material is available for this article:
Figure Sl - Rank abundance curves of dung beetle species sampled with flight interception traps (FIT) and pitfall traps baited with human feces at Serra do Divisor National Park (SDNP), Acre, Brazil. Y-axis was calculated with species relative abundance on a logarithmic scale (log^).
Table Sl - Sample size, species richness and sample coverage of dung beetles sampled at three transects (plots A, B, and C) with flight interception traps (FITs) and pitfall traps baited with human feces at Serra do Divisor National Park (SDNP), Brazil.
Acknowledgments
We are grateful to the Research Council of Norway (project no. 288086) for financial and logistic support. We thank all members of the Amazon Biodiversity and Carbon (ABC) Expeditions (abc-expeditions. com) for logistic and field support, and the residents and managers of SDNP for their support. We thank Arturo Gonzalez-Alvarado and Jorge A. Arias-Buriticå for confirming the identities of some of the dung beetle species. This study derives from the first author PhD thesis and was financed by Coordenagao de Aperfe^oamento de Pessoal de Nivel Superior - Brazil (CAPES) (n. 88887 686421/2022-00 and n. 88881 934586/2024-01). We are grateful to all reviewers for their insiglitful comments on this paper. This is publication #05 of the ABC Expeditions.
Associate Editor
Carlos Joly
Author Contributions
Andressa Bach: conceptualization; data curation; formal analysis; investigation; methodology; writing - original draft.
Torbjørn Haugaasen: conceptualization; investigation; funding acquisition; project administration; supervision; writing - review & editing.
Carlos A. Peres: conceptualization; funding acquisition; project administration; writing - review & editing.
Emanuelly F. Lucena: investigation; methodology; writing - review & editing.
José Victor A. Ferreira: investigation; methodology; writing - review & editing.
Joseph E. Hawes: conceptualization; funding acquisition; project administration; writing - review & editing.
Bruna R. Bordin: investigation; methodology.
Mario Cupello: investigation; methodology; writing - review & editing.
Renato A. de Azevedo: methodology.
Fernando Z. Vaz-de-Mello: conceptualization; investigation; supervision; resources; writing - review & editing.
Conflicts of Interest
The authors declare that they have no conflict of interest related to the publication of this manuscript.
Ethics
This study did not involve human beings and/or clinical trials that should be approved by one Institutional Committee.
Data Availability
Supporting data are available at <https://doi.org/10.48331/ scielodata.QT4KDV>.
References
ALMEIDA, H.A., ANTONINI, Y., TAVARES JUNIOR, C., BRAGA, R.F., DA SILVA, P.G. & BEIROZ, W. 2022a. Dung beetles can sow: the potential of secondary seed dispersers to assist ecological restoration. Ecol. Entomol. 47:181-191. https://doi.org/10.1111/een.13100.
ALMEIDA, M.R.N., ROCHA, V.J. & BERNARDE, P.S. 2022b. Notas sobre a ocorréncia de pequenos mamiferos pouco conhecidos no Parque Nacional da Serra do Divisor, estado do Acre, Brasil. SAREM 4:1-8. https://doi. org/10.31687/SaremNMS22.9.3.
ANDRADE, R.B., BARLOW, J., LOUZADA, J., VAZ-DE-MELLO. F.Z., SOUZA, M., SILVEIRA, J.M., COCHRANE, Μ.Α. 2011. Quantifying responses of dung beetles to fire disturbance in Tropical Forests: The importance of trapping method and seasonality. Plos One. 6:e26208. https:// doi.org/10.1371/journal.pone.0026208.
ANTONELLI, A., ZIZKA, A., CARVALHO. F.A., SCHARN, R., BACON, C.D., SILVESTRO, D. & CONDAMINE, F.L. 2018. Amazonia is the primary source of Neotropical biodiversity. PNAS 115(23):6034-6039, http://www. pnas.org/cgi/doi/10.1073/pnas.1713819115.
ARELLANO, L., NORIEGA, J.A., ORTEGA-MARTINEZ, I.J., RIVERA, J.D., CORREA. C.M.A., GOMEZ-CIFUENTES, A., RAMIREZ-HERNANDEZ, A. & BARRAGAN, F. 2023. Dung beetles (Coleoptera: Scarabaeidae) in grazing lands of the Neotropics: A review of patterns and research trends of taxonomic and functional diversity, and functions. Front. Ecol. Evol. 11:1084009. https://doi.org/10.3389/fevo.2023.1084009.
ARELLANO, L., ORTEGA-MARTINEZ, I.J., RIVERA, J.D. & ALVARADO, F. 2024. Declining dung beetle (Coleoptera: Scarabaeidae) abundance and Diversity in the Neotropics: Causes and Conservation strategies. In Insect Decline and Conservation in the Neotropics (J.L. León-Cortés & A. Cordoba-Aguilar, eds.). Springer, Cham. p. 75-115. http://dx.doi. org/10.1007/978-3-03l-49255-6_5.
ARIAS-BURITICÅ, J.A., BACH, A., PERES, C.A., HAUGAASEN, T., HAWES, J.E., AZEVEDO, R.A. & VAZ-DE-MELLO, F.Z. 2023. A New Species of Isocopris Pereira and Martinez, 1960 (Coleoptera: Scarabaeidae: Scarabaeinae) from the Southwest Brazilian Amazon. Coleopt. Bull. 77(4):629-635. https://doi.Org/10.1649/0010-065X-77.4.629.
ARIAS-BURITICÅ, J.A., FIGUEROA, L. & VAZ-DE-MELLO, F.Z. First record and range extension of Isocopris rossinii Arias-Buriticå et al.. 2023 (Coleoptera: Scarabaeidae: Scarabaeinae) for the Peruvian Amazon. Coleopt. Bull. 78:567-569. https://doi.Org/10.1649/0010-065X-78.4.567.
AZEVEDO, C.O., HELMER, J.L. & MORATO, E. 2002. Diversidade de Bethylidae (Hymenoptera) do Parque Nacional da Serra do Divisor, Acre Brasil e seu uso no plano de manejo e conservarlo da àrea. Acta Amazon. 32(1):71-82. https://doi.org/10.1590/1809-43922002321082.
BARBER, C.P., COCHRANE, Μ.Α.. SOUZA JR, C.M. & LAURENCE. W.F. 2014. Roads, deforestation, and the mitigating effect of protected areas in the Amazon. Biol. Conserv. 177:203—209. https://doi.0rg/10.1016/j, biocon.2014.07.004.
BERNARD. E., PENNA. L.A.O. & ARAÙJO, E. 2014. Downgrading, Downsizing, Degazettement, and Reclassification of Protected Areas in Brazil. Conserv. Biol. 28(4):939-950. http://dx.doi.org/10.1111/cobi.12298.
BITENCOURT, B.S., DA SILVA, P.G., MORATO, E.F. & DE LIMA. Y.G. 2019a. Dung beetle responses to successional stages in the Amazon rainforest. Biodivers. Conserv. 28:2745-2761. https://doi.org/10.1007/ sl0531-019-01791-y.
BITENCOURT. B.S., DIMAS, Τ.Μ., DA SILVA, P.G. & MORATO, E.F. 2019b. Forest complexity drives dung beetle assemblages along an edge-interior gradient in the southwest Amazon rainforest. Ecol. Entomol. 45(2):259-268. https://d0i.0rg/l0.1111/een.12795.
BRAZIL. 1989. Decreto Federal n° 97.839 16 de Junho de 1989. http://www. planalto.gov.br/ccivil_03/dccrcto/antigos/d97839.htm (last access in 02/ June/2024).
BRAZIL. 2000. Lei Federal n° 9.985 de 18 de Julho de 2000. http://www.planalto. gov.br/ccivil_03/leis/19985.htm (last access in 02/June/2024).
BRAZIL. 2019, Camara dos Deputados - Projeto de Lei 6024/2019. https://www. camara.leg.br/proposicoesWeb fichadetramitacao?idProposicao=2229754 &fichaAmigavel=nao (last access in 04/July/2024).
CALOURO, Α.Μ. 1999. Riqueza de mamiferos de grande e mèdio porte do Parque Nacional da Serra do Divisor (Acre, Brasil). Rev. Bras. Zool. 16(2):195-213. https://doi.org/10.1590/S0101-81751999000600020.
CAMBEFORT, Y. & HANSKI, I., 1991. Dung Beetle Population Biology, In Dung Beetle Ecology (I. Hanski & Y. Cambefort, eds). Princeton University Press, United Kingdom, p. 211-229.
CARPIO, C., DONOSO, D.A., RAMÓN, G. & DANGLES, 0.2013. Short term response of dung beetle communities to disturbance by road construction in the Ecuadorian Amazon. Ann. Soc. Entomol. Fr. 45(4):455—469. http:// dx.doi.org/10.l080/00379271.2009.10697629.
CARVALHO, R.L., ANDERSEN, Α.Ν., ANJOS, D.V., PACHECO, R., CHAGAS, L. & VASCONCELOS, H.L. 2020. Understanding what bioindicators are actually indicating: Linking disturbance responses to ecological traits of dung beetles and ants. Ecol. Indic. 108:105767. https:// doi.org/10.1016/j.ecolind.2019.105764.
CHAO, A., GOTELLI, N.J., HSIEH, T.C., SANDER, E.L., MA, Κ.Η., COLWELL, R.K. & ELLISON, A.M. 2014. Rarefaction and extrapolation with Hill numbers: a framework for sampling and estimation in species diversity studies. Ecol. Monogr. 84(l):45-67. https://doi.org/10.1890/13- 0133.1.
COL - CATALOGUE OF LIFE. 2024. https://www.catalogueoflife.org/ (last access in 17/September/2024).
COOK, J. 1998. A revision of the Neotropical genus Bdelyrus Harold (Coleoptera: Scarabaeidae). Can. Entomol. 130:631-689. https://doi. org/10.4039/Entl30631-5.
COOK, J. 2002. Revision of the Neotropical Genus Cryptocanthon Balthasar (Coleoptera: Scarabaeidae: Scarabaeinae). Coleopterists Society Monographs 56:1-96. http://dx.doi.org/10.1649/00l0-065X(2002) 56[3:AROTNG]2.0.CO;2.
CREÀO-DUARTE,A..I., LOURENqO, A., ROTHÉA, R.R.A.D. & PEREIRA- COLAVITE, A. 2023. Treehoppers (Hemiptera, Auchenorryncha: Aetalionidae and Membracidae) from western Acre, Brazil, with emphasis on the fauna of Serra do Divisor National Park. Biota Neotropica 23(3):e20231488. https://doi.org/10.1590/1676-0611-BN-2023-1488 (last access on 24/September/2024).
CUPELLO, M. & VAZ-DE-MELLO, F.Z. 2013. New evidence for the validity of Coprophanaeus (C.) terrali Arnaud, 2002 (Coleoptera: Scarabaeidae: Scarabaeinae: Phanaeini), a dung beetle from Brazil. Zootaxa 3717:359-368.
CUPELLO, M. & VAZ-DE-MELLO, F.Z. 2018. A monographic revision of the Neotropical dung beetle genus Sylvicanthon Halffter & Martinez, 1977 (Coleoptera: Scarabaeidae: Scarabaeinae: Deltochilini), including a reappraisal of the taxonomic history of ‘Canthon sensu lato’. Eur. J. Taxon. 467:1-205. https://doi.org/l0.5852/ejt.2018.467.
CUPELLO. M.. SILVA, F.A.B. & VAZ-DE-MELLO, F.Z. 2023. The Taxonomic Revolution of New World dung beetles (Coleoptera: Scarabaeidae: Scarabaeinae). Front. Ecol. Evol. 1168754:1-42. https://doi.org/10.3389/ fevo.2023.1168754.
DA SILVA. P.G. & HERNANDEZ. Μ.Ι.Μ. 2015. Spatial Patterns of Movement of Dung Beetle Species in a Tropical Forest Suggest a New Trap Spacing for Dung Beetle Biodiversity Studies. PLoS ONE 10(5):c0126112. https:// doi.org/10. I 37I/journal.pone.0126112.
DOLIBAINA, D.R., DIAS, F.M.S., MIELKE, Ο.Η.Η. & CASAGRANDE, M.M. 2015. Argyrogrammana Strand (Lepidoptera: Riodinidae) from Parque Nacional da Serra do Divisor. Acre. Brazil, with description of four new species. Zootaxa 4028(2):227-245. http://dx.doi.org/10.11646/ zootaxa.4028.2.4.
DOLIBAINA, D.R., DIAS, F.M.S., SIEWERT, R.R., MIELKE, Ο.Η.Η. & CASAGRANDE, M.M. 2016. Mesene Doubleday (Lepidoptera: Riodinidae) from Parque Nacional da Serra do Divisor, Acre, Brazil, witli taxonomic notes and the description of a new species. Zootaxa 4175(5): 463-472. http://doi.Org/10.11646/zootaxa.4175.5.4.
DOLIBAINA, D.R., LEITE, L.A.R., DIAS, F.M.S., MIELKE, Ο.Η.Η. & CASAGRANDE, M.M. 2012. An annotated list of Symmachia Hubner, [1819] (Lepidoptera: Riodinidae: Symmachiini) from Parque Nacional da Serra do Divisor, Acre. Brazil, witli the description of a new species. Insecta Mundi 0249:1-11.
EDMONDS, W.D. & ZIDEK, J. 2004. Revision of the Neotropical Dung Beetle Genus Oxysternon (Scarabaeidae: Scarabaeinae: Phanaeini). Folia Heyrovskyana 11:1-58.
EDMONDS, W.D. & ZIDEK, J. 2010. Taxonomic review of the neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 0129:1-111.
EDMONDS, W.D. & ZIDEK, J. 2012. Taxonomy of Phanaens revisited: Revised keys to and comments on species of the New World dung beetle genus Phanaeus Macleay, 1819 (Coleoptera: Scarabaeidae: Scarabaeinae, Phanaeini). Insecta Mundi 0274:1-108.
EDMONDS, W.D. 1994. Contributions in Science: Revision of Phanaeus Macleay. a New World Genus of Scarabaeine Dung Beetles (Coleoptera: Scarabaidae, Scarabaeinae). 1 ed. Natural History Museum, Los Angeles.
FALE1RO, A. 2023. Relatório da comissào da Amazònia e dos povos originarios e tradicionais: Projeto de Lei n° 6.024, de 2019. https://www.camara.leg. br/proposicoesWeb/fichadetramitacao?idProposicao=2229754 (last access in 02/September/2024).
FAVILA, M.E., NORIEGA, J.A., DA SILVA, P.G. & KORASAKI, V. 2023. Editorial: Neotropical dung beetle diversity: ecological, historical, and anthropogenic perspectives. Front. Ecol. Evol. 11:1294471. http://dx.doi. org/10.3389/fevo.2023.1294471.
FERREIRA, R.R., LUCENA, E.F., KOROIVA, R., AZEVEDO, R.A., HAUGAASEN, T., PERES, C.A., HAWES, J.E. & VASCONCELLOS, A. 2023. Amazonian forest termites: a species checklist from the State of Acre, Brazil. Biota Neotropica 23(4):e20231551.http://dx.doi.org/10.1590/1676- 0611-bn-2023-l551 (last access in 02/September/2024).
FILGUEIRAS, Β.Κ.. LIBERAL, C.N., AGUIAR, C.D.M., HERNANDEZ. M.I.M. & lANNUZZI, L. 2009. Attractivity of omnivore, carnivore and herbivore mammalian dung to Scarabaeinae (Coleoptera, Scarabaeidae) in a tropical Atlantic rainforest remnant. Rev. Bras. Entomol. 53(3):422-427. https://doi.org/10.1590/S0085-56262009000300017.
GARDNER, Τ.Α., BARLOW, J., ARAUJO, I.S., AVILA-PIRES, T.C., BONALDO, Α.Β., COSTA, J.E. et al. 2008. The cost-effectiveness of biodiversity surveys in tropical forests. Ecol. Lett. 11(2):139—150. https:// doi.0rg/lO.llll/j.1461-0248.2007.01133.x.
GÉNIER, F. & ARNAUD, P. 2016. Dendropaemon Perty, 1830: taxonomy, systematics and phylogeny of the morphologically most derived phanaeine genus (Coleoptera: Scarabaeidae, Scarabaeinae, Phanaeini). Zootaxa 4099:1-125. https://doi.0rg/lO.l1646/zootaxa.4099.I.I.
GENIER, F. 1996. A revision of the Neotropical genus Ontherus Erichson (Coleoptera: Scarabaeidae, Scarabaeinae). Men. Entomol. Soc. Can. 170:1—169.
GÉNIER, F. 2009. Le genre Eurysternus Dalman, 1824 (Scarabaeidae: Scarabaeinae: Oniticellini), révision taxonomique et clés de determination illustrées. 1 ed. Sofia, Pensoft. http://dx.doi.Org/10.13140/RG.2.l.1627.9129.
GILL, B.D. 1991. Dung Beetles In Tropical American Forests. In Dung Beetle Ecology (I. HANSKI & Y. CAMBEFORT, eds.). Princeton University Press, United Kingdom, p. 211-229.
GONQALVES-SOUZA, D., VILELA, B., PHALAN, B. & DOBROVOLSKI, R. 2021. The role of protected areas in maintaining the natural vegetation in Brazil. Sci. Adv. 7(38):1-9. https://doi.org/10.1126/sciadv.abh2932.
GONZALEZ, O. & ACUY, M. 2017. A new location in Peru for the poorly- known Acre Antshrike Thamnophilus divisorius endemie of Serra do Divisor. Rev. Bras. Ornitol. 25(2):137-139. http://dx.doi.org/10.1007/BF03544390.
HALFFTER, G. & EDMONDS, W.D. 1982. The nesting behavior of dung beetles (Scarabaeinae): An ecological and evolutive approach. 1 ed. Instituto de Ecologia, México.
HALFFTER, G. & HALFFTER, V. 2009. Why and where coprophagous beetles (Coleoptera: Scabaeinae) eat seeds, fruits or vegetable detritus. Boletin de la S.E.A. 45:1-22.
HALFFTER, G. & MATTHEWS, E.G., 1966. The Natural History of Dung Beetles of the Subfamily Scarabaeinae (Coleoptera: Scarabaeidae), 1 ed. Folia Entomologica Mexicana, México.
HAMEL-LEIGUE, A.C., HERZOG, S.K. & MANN, D.J. 2008. Composición y riqueza de una comunidad de escarabajos peloteros (Coleoptera: Scarabaeinae) en los Youngas bajos de la Cordillera Mosetenes, Bolivia. Revista Boliviana de Ecologia y Conservación Ambientai 23:39—19.
HAMEL-LEIGUE. A.C., HERZOG. S.K., MANN, D.J., LARSEN, Τ.Η., GILL, B.D., EDMONDS, W.D. & SPECTOR, S. 2009. Distribución e Historia Natural de Escarabajos Coprófagos de la tribu Phanaeini (Coleoptera: Scarabaeidae: Scarabaeinae) en Bolivia. Kempffiana 5:43-95.
HOANG, Ν.Τ. & KANEMOTO, Κ. 2021. Mapping the deforestation footprint of nations reveals growing threat to tropical forests. Nat. Ecol. Evol. 5:845-853. https://doi.org/10.1038/s41559-021-01417-z.
HSIEH, T C. & CHAO, A. iNEXT: interpolation and EXTtrapolation for species diversity. R package version 3.0.1. http://chao stat nthu edu tw/wordpress/ software-download/.
ISA. 2023. Instituto Socioambiental - Parque Nacional da Serra do Divisor. https://uc.socioambiental.org/pt-br/arp/598 (last access in 26/June/2023).
JONES, J.P.G., ASNER, G.P., BUTCHART, S.H.M. & KARANTH, K.U. 2013. The ‘why’, ’what’ and ‘how’ of monitoring for conservation. In Key topics in conservation biology 2 (D. MACDONALD & K.J. WILLIS, eds.). Wiley- Blackwell, p. 329-343.
KAMINSKI, L.A., CARNEIRO, E., DOLIBAINA. D.R., CASAGRANDE, M.M. & MIELKE, Ο.Η.Η. 2020. Oviposition of Minstrellus grandis (Lepidoptera: Riodinidae) in a harmful ant-plant symbiosis. Acta Amazon. 50:256-259. http://dx.doi.org/10.1590/1809-4392202001801.
KINDT, R. & COE, R. 2005. Tree diversity analysis. A manual and software for common statistical methods for ecological and biodiversity studies. ICRAF. http://www.worldagroforestry.org/output/tree-diversity-analysis.
KOGA, D.M., BROWN, I.F., FEARNSIDE, Ρ.Μ., SALISBURY, D.S. & DA SILVA, S. 2022, Serra do Divisor National Park: a protected area under threat in the South-western Brazilian Amazon. Environ. Conserv. 49:74-82. https://doi.org/I0.I017/S037689292200009I.
LARSEN, Τ.Η. & FORSYTH, A. 2005. Trap spacing and transect design for dung beetle biodiversity studies. Biotropica 37(2):322-325. http://dx.doi. org/10.1111/j.1744-7429.2005.00042.Χ.
LARSEN, Τ.Η. 2004. Dung beetles (Coleoptera: Scarabaeidae: Scarabaeinae). In Peru: Megantoni. Rapid Biological Inventories Resport 15 (C. VRIESENDORP, L. RIVERA CHAVEZ, D. MOSKOVITS & J. SHOPLAND, eds.). The Field Museum, Chicago, p. 185-192.
LARSEN, Τ.Η., LOPERA, A. & FORSYTH, A. 2006. Extreme Trophic and Habitat Specialization by Peruvian Dung Beetles (Coleoptera: Scarabaeidae: Scarabaeinae). Coleopt. Bull. 60:315-324. http://dx.doi.org/10.1649/0010- 065X(2006)60[315:ETAHSB]2.0.CO:2.
LAURANCE, W.F. & BALMFORD, A. 2013. A global map for road building. Nat. 495:308-309. https://doi.org/10.1038/495308a.
LEMOS, L.P.. BIZRI, H.R.E., DO AMARAL, J.V., SANTOS, A.S., KOGA, D.M. & SILVA, F.E. 2018. Caca de vertebrados no Parque Nacional da Serra do Divisor, Acre. BioBrasil 8(1):69-88. https://doi.org/10.37002/ biodiversidadebrasileira.v8il.795.
LOPES, M.A.O.A. & REHG, J.A. 2003. Observations of Callimico goeldii with Saguinus imperator in the Serra do Divisor National Park, Acre, Brasil. Neotrop. Primates 11(3):181-183. https://doi.org/10.62015/np.2003. vll.557.
LOUZADA, J. 2008. Scarabacinac (Coleoptera: Scarabaeidae) Detritivoros cm Ecossistemas Tropicais: Diversidade e servitos ambientais. In Biodiversidade do Solo em ecossistemas brasileiros (F.M.A. MOREIRA, J.O. SIQUEIRA & L. BRUSSAARD, eds.). Editora UFLA, Lavras, p. 309-332.
MALDONADO, Μ.Β., ARANIBAR, J.N., SERRANO, Α.Μ., CHACOFF, Ν.Ρ. & VÀZQUEZ, D.P. 2019. Dung beetles and nutrient cycling in a dryland environment. Catena 179:66—73. https://doi.0rg/lO.lOl6/j. catena.2019.03.035.
MASCIA, M.B. & PAILLER, S. 2010. Protected area downgrading, downsizing, and degazettement (PADDD) and its conservation implications. Policy Perspective 4(l):9-20. https://doi.Org/10.llll/j.1755-263X.2010.00147.x.
MMA. 2024. Ministério do Meio Ambiente. Relatório do Cadastro Nacional de Unidades de Conservato Parque Nacional da Serra do Divisor, https:// cnuc.mma.gov.br/pesquisar (last access in 02/July/2024).
MORA-AGUILAR, E.F., ARRIAGA-JIMENEZ, A., CORREA, C.M.A., DA SILVA, P.G., KORASAK1, V. LOPEZ-BEDOYA, Ρ.Α., HERNANDEZ, M.I.M., et al. 2023. Toward standardized methodology for sampling dung beetles (Coleoptera: Scarabaeinae) in the Neotropics: A critical review. Front. Ecol. Evol. 11:109608. https://doi.org/10.3389/fevo.2023.1096208.
MORATO, E.F., AMARANTE, S.T. & SILVEIRA. Ο.Τ. 2008. Avaliagao ecologica ràpida da fauna de vespas (Hymenoptera: Aculeata) do Parque Nacional da Serra do Divisor, Acre. Brasil. Acta Amazon. 38(4):789-798. https://doi.org/10.1590/S0044-59672008000400025.
MORRONE, J. 2009. Evolutionary biogeography: an integrative approach with case studies. 1 ed. Columbia University Press.
MORRONE, J.J., ESCALANTE, T., RODRIGUEZ-TAPIA, G., CARMONA, A., ARANA, M. & MERCADO-GÓMEZ, J. D. 2022. Biogeographic regionalization of the Neotropical region: New map and shapefile. An. Acad. Bras. Ciènc. 94:e20211167. https://doi.org/10.1590/0001- 3765202220211167.
NICHOLS, E., SPECTOR. S.. LOUZADA, J., LARSEN, T., AMEZQUITA. S.. FAVILA, Μ.Ε. & THE SCARABAEINAE RESEARCH NETWORK. 2008. Ecological functions and ecosystem services provided by Scarabaeinae dung beetles. Biol. Conserv. 141(6):1461-1474. https://doi.0rg/lO.lOl6/j. biocon.2008.04.011.
NICHOLS, E., URIART, M., PERES. C.A.. LOUZADA, J., BRAGA, R.F., SCHIFFLER, G., ENDO, W. & SPECTOR, S.H. 2013. Human-Induced Trophic Cascades along the Fecal Detritus Pathway. PLoS ONE 8(10):e75819. https://doi.org/10.1371/journal.pone.0075819.
NORIEGA, J.A., MARCH-SALAS, M., CASTILLO, S., GARCÌA-Q, H., HORTAL, J. & SANTOS, A.M.C. 2021. Human perturbations reduce dung Beetle diversity and dung removal ecosystem function. Biotropica. 53:735-766. https://doi.org/10.llll/btp.12953.
OLIVEIRA, G.C.S., HENRIQUES, N.R., LOUZADA, J. & VIEIRA, L. 2022. New behavior report by Canthon virens (Mennerheim, 1 829) (Coleoptera: Scarabaeidae: Scarabaeinae) in Southeastern Brazil. Entomol. Commun. 4:ec04023. https://doi.org/10.37486/2675-1305.ec04023.
PEREIRA, E.J.A.L., RIBEIRO, L.C.S., FREITAS, L.F.S. & PEREIRA, H.B.B. 2020. Brazilian policy and agribusiness damage the Amazon rainforest. Land Use Policy 92:104491. https://doi.Org/10.1016/j.landuscpol.2020.104491.
PULIDO-HERRERA, L.A.. MEDINA, C.A. & RIVEROS, R.A. 2007. Nuevos registros de escarabajos coprófagos (Scarabaeidae: Scarabaeinae) para la region andina de Colombia. Parte I. Rev. Acad. Colomb. Cienc. Exactas, Fis. Nat. 31:305-310.
QIN, Υ., XIAO, Χ., LIU, F., SILVA, F.S.. SHIMABUKURO. Y.. ARAI, E. & FEARNSIDE, P.M. 2022. Forest conservation in Indigenous territories and protected areas in the Brazilian Amazon. Nat. Sustain. 6:295-305. https:// doi.org/10.1038/s41893-022-01018-z.
R CORE TEAM. 2023. _R: A Language and Environment for Statistical. Computing_. R Foundation for Statistical Computing, Vienna, Austria. https://www R-project org/.
RAVEN, Ρ.Η., GEREAU, R.E., PHILLIPSON, Ρ.Β., CHATELAIN, C., JENKINS. C.N. & ULLOA, A.U. The distribution of biodiversity richness in the tropics. Sci. Adv. 6(37): l-5, 2020. https://doi.0rg/lO.l l26/sciadv.abc6228.
ROBINSON, C., DUINKER, Ρ.Ν. & BEAZLEY, K.F. 2010. A conceptual framework for understanding, assessing, and mitigating ecological effects of forest roads. Environ. Rev. 18:61-86. http://dx.doi.org/10.1139/A10-002.
ROSSINI, M. & VAZ-DE-MELLO, F.Z. 2017. A taxonomic review of the genus Isocopris Pereira and Martinez, 1960 (Coleoptera: Scarabaeidae, Scarabaeinae), witli description of a new Brazilian species. J. Nat. Hist. 51:1091-1117. http://dx.doi.org/10.1080/00222933.2017.1319517.
ROSSINI, M. & VAZ-DE-MELLO, F.Z. 2020. Taxonomic review of the Dichotomius mamillatus group (Coleoptera: Scarabaeidae), with a description of a new species, Dichotomius (Dichotomius) gandinii sp. nov., from western Amazonia. Aust. J. Entomol. 59:52—73. https://doi. org/lO.llll/aen.12443.
ROSSINI, M., VAZ-DE-MELLO, F.Z. & ZUNINO, M. 2018. A taxonomic revision of the New World Outhophugus Latreille, 1802 (Coleoptera: Scarabaeidae: Scarabaeinae) of the osculatii species-complex, with description of two new species from South America. J. Nat. Hist. 52(9-10): 541-586. https://doi.org/10.1080/00222933.2018.1437230.
RUARO, R. & LAURENCE, W.F. 2021. Pending bill could devastate Brazil’s Serra do Divisor National Park. Nat. Ecol. Evol. 6:120—121. https://doi. org/10. I038/s41559-021-01632-8.
SCHOOLMEESTERS, P. 2024. World Scarabaeidae Database (version 2024-08-05). In Catalogue of Life (Version 2024-08-29) (O. BÀNKI, Y. ROSKOV, M. DÒRING, G. OWER, G., D.R. HERNANDEZ ROBLES, C.A. PLATA CORREDOR, et al., org.). Amsterdam, Netherlands, https:// doi.org/10.48580/dgdwl-38g.
SERNANP. 2024. Servicio Nacional de Areas Naturales Protegidas por el Estado. Parque Nacional Sierra del Divisor, https://www.gob.pe/institucion/sernanp/ informes-publicaciones/1949413-parque-nacional-sierra-del-divisor (last access in 02/July/2024).
SEYMOUR, F. & HARRIS, N.L. 2019. Reducing tropical deforestation. Science 365(6455):756-757. https://doi.org/10.1126/science.aax8546.
SILVA, F.A.B. & VALOIS, M. 2019. A taxonomic revision of the genus Scybalocanthon Martinez, 1948 (Coleoptera: Scarabaeidae: Scarabaeinae: Deltochilini). Zootaxa, 4629(3):301-341. https://doi.org/10.11646/ zootaxa.4629.3.1.
SOARES-FILHO, B., MOUTINHO, P., NEPSTAD. D., ANDERSON. A., RODRIGUES, H., GARCIA, R., DIETZSCH, L., MERRY. F., BOWMAN, M., HISSA, L., SILVESTRINI, R. & MARETTI, C. 2010. Role of Brazilian Amazon protected areas in climate change mitigation. PNAS 107(24):10821-10826. http://www.pnas.org/cgi/doi/10.1073/ pnas.0913048107.
SOARES-FILHO, B.S., OLIVEIRA. U., FERREIRA, Μ.Ν., MARQUES, F.F.C., DE OLIVEIRA, A.R., SILVA, F.R. & BORNER, J. 2023. Contribution of the Amazon protected areas program to forest conservation. Biol. Conserv. 279:109928. https://doi.Org/10.r016/j.biocon.2023.l09928.
TSUJI, Y., SHIRAISHI, K.S.T., MATSUBARA, M. & KOSUGI, J. 2021. Differential Attraction of Large and Small Tunneling Dung Beetles (Coleoptera: Geotrupidae and Scarabaeidae) to Native Mammal Dung in Satoyama Forest in Central Japan. Coleopt. Bull. 75(2):376-381. https:// doi.org/10.1649/0010-065Χ-75.2.376.
UNESCO. 2024. Serra do Divisor National Park, https://whc.unesco.org/en/ tentativelists/l12Ι/ (last access in 02/June/2024).
VALOIS, M.C., VAZ-DE-MELLO, F.Z. & SILVA, F.A.B. 2023. Taxonomic revision of the batesi species group of Dichotomius Hope, 1838, with the description of three new Amazonian species (Coleoptera, Scarabaeidae, Scarabaeinae). Insect Syst. Evol. 54(5):459-490. https://doi. org/10.I163/1876312X-bjal0046.
VAZ-DE-MELLO, F.Z. & BORDIN, B.R. 2024. Scarabaeinae in Catàlogo Taxonòmico da Fauna do Brasil, http://fauna.jbrj.gov.br/fauna/ faunadobrasil/127498 (last access in 01/July/2024).
VAZ-DE-MELLO, F.Z. 1999. Scarabaeidae s. str. (Coleoptera: Scarabaeoidea) de um Fragmento de Floresta Amazònica no Estado do Acre, Brasil. 1. Taxocenose. An. Soc. Entomol. Bras. 28(3):447—453, https://doi. org/10.1590/S0301-80591999000300009.
VAZ-DE-MELLO, F.Z., EDMONDS, W.D., OCAMPO, F.C. & SCHOOLMEESTERS, P. 2011. A multilingual key to the genera and subgenera of the subfamily Scarabaeinae of the New World (Coleoptera: Scarabaeidae). Zootaxa, 2854:1-73. http://dx.doi.org/10.11646/ zootaxa.2854.1.1.
VAZ-DE-MELO, F.Z., LOUZADA, J.N.C. & SCHOEREDER, J.H. 1998. New data and comments on Scarabaeidae (Coleoptera: Scarabaeoidea) associated with Attini (Hymenoptera: Formicidae). Coleopt. Bull. 52(3):209-216. https://www.jstor.org/stable/4009354.
VERDU, J.R., NUMA C. & HERNANDEZ-CUBA, O. 2011. The influence of landscape structure on ants and dung beetles diversity in a Mediterranean savanna-Forest ecosystem. Ecol. Indic. 11(3):831—839. https://doi. org/10.10I6/j.ecolind.2010.10.011.
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. https://doi.0rg/lO.lOl6/j. ecolind.2013.09.036.
WHITNEY, B.M., OREN, D.C. & BRUMFIELD, R.T. 2004. A new species of Thamnophilus antshrike (Aves: Thamnophilidae) from Serra do Divisor, Acre, Brazil. Auk 121(4):1031-1039. https://doi.org/10.1093/ auk/121.4.1031.
WICKHAM, H. 2016. ggplot2: Elegant Graphics for Data Analysis. SpringerVerlag, New York.
Received: 14/02/2025
Accepted: 17/02/2025
Published online: 21/03/2025

https://doi.org/10.1590/1676-0611-BN-2025-1770	http://www.scielo.br/bn

http://www.scielo.br/bn	https://doi.org/10.1590/1676-0611-BN-2025-1770

https://doi.org/10.1590/1676-061 l-BN-2025-1770	http://www.scielo.br/bn