COMMUNITY AND ECOSYSTEM ECOLOGY Dung Beetles (Coleoptera: Scarabaeidae) Attracted to Dung of the Largest Herbivorous Rodent on Earth: a Comparison With Human Feces ANDERSON PUKER,1,2 CE´ SAR M. A. CORREA,3 VANESCA KORASAKI,4 KLEYTON R. FERREIRA,3 5 AND NAIARA G. OLIVEIRA Environ. Entomol. 42(6): 1218Ð1225 (2013); DOI: http://dx.doi.org/10.1603/EN13100 ABSTRACT The capybara, Hydrochoerus hydrochaeris (L.) (Rodentia: Caviidae), is the largest herbivorous rodent on Earth and abundant in the Neotropical region, which can provide a stable food source of dung for dung beetle communities (Coleoptera: Scarabaeidae: Scarabaeinae). However, the use of capybara dung by dung beetles is poorly known. Here, we present data on the structure of the dung beetle community attracted to capybara dung and compare with the community attracted to human feces. Dung beetles were captured with pitfall traps baited with fresh capybara dung and human feces in pastures with exotic grass (Brachiaria spp.), patches of Brazilian savanna (Cerrado), and points of degraded riparian vegetation along the Aquidauana river in Anasta´cio and Aquidauana, Mato Grosso do Sul, Brazil. In traps baited with human feces, 13,809 individuals of 31 species were captured, and in those baited with capybara dung 1,027 individuals belonging to 26 species were captured. The average number of individuals and species captured by the traps baited with human feces was greater than for capybara dung in all habitats studied. Composition of the communities attracted to human feces and capybara dung formed distinct groups in all habitats. Despite the smaller number of species and individuals captured in capybara dung when compared with human feces, capybara dung was attractive to dung beetles. In Brazil, the legalization of hunting these rodents has been debated, which would potentially affect the community and consequently the ecological functions performed by dung beetles that use the feces of these animals as a resource. In addition, the knowledge of the communities associated with capybaras may be important in predicting the con- sequences of future management of their populations. KEY WORDS biodiversity conservation, capybara dung, coprophagous beetle, feeding preference, Scarabaeoidea Dung beetles (Coleoptera: Scarabaeidae: Scarabae- which are rolled various distances and then buried; inae) comprise Ϸ6,000 species found worldwide, with and tunneler species that construct tunnels below or 1,250 species described for the Neotropical region adjacent to the food resource and transport dung into (Hanski and Cambefort 1991), of which 600 have been the bottom (Halffter and Matthews 1966). This diver- reported in Brazil (Vaz-de-Mello 2000). The feeding sity of feeding habits affects characteristics of behav- behavior of dung beetles is quite varied, including ior, morphology, and development (Halffter and Mat- carcasses, fungi, fruits, and decaying plant material, thews 1966, Hanski and Cambefort 1991), and this may but coprophagy is most frequent (e.g., Halffter and be a determining factor in species richness in the Matthews 1966). Dung beetles also have different re- Neotropical region (Halffter and Favila 1993). source allocation strategies, including dweller species The availability of mammal dung to the dung beetles that live within a dung pat and do not exhibit resource varies in time and space, and is important for the allocation; roller species that remove portions of dung, stabilization of communities (Herna´ndez and Vaz-de- Mello 2009). The attractiveness of this resource may 1 Programa de Po´s-Graduac¸a˜o em Entomologia, Departamento de also vary depending on the feeding habits of these Entomologia, Universidade Federal de Vic¸osa, 36570-000 Vic¸osa-MG, animals (carnivore, herbivore, and omnivore), but Brazil. some studies show that dung beetles are more at- 2 Corresponding author, e-mail: [email protected]. 3 Programa de Po´s-Graduac¸a˜o em Agronomia, Laborato´rio de En- tracted to feces of omnivorous mammals (e.g., Fincher tomologia, Universidade Estadual de Mato Grosso do Sul, Rodovia et al. 1970, Martõ´nÐPiera and Lobo 1996, Filgueiras et Aquidauana/CERA km 12, 79200-000 Aquidauana-MS, Brazil. al. 2009, Whipple and Hoback 2012). In Brazil, dung 4 Departamento de Biologia, Setor de Ecologia, Universidade Fed- beetles have been sampled using traps baited with eral de Lavras, 37200-000 Lavras-MG, Brazil. 5 Universidade Estadual de Mato Grosso do Sul, Rodovia different types of baits, including the feces of domestic Aquidauana/CERA km 12, 79200-000 Aquidauana-MS, Brazil. (e.g., Flechtmann et al. 2009, Louzada and Carvalho e 0046-225X/13/1218Ð1225$04.00/0 ᭧ 2013 Entomological Society of America December 2013 PUKER ET AL.: FEEDING PREFERENCE OF DUNG BEETLES 1219 Silva 2009, Correa et al. 2013) and wild animals (e.g., ness of dung beetles (e.g., Almeida et al. 2011, Abot et Filgueiras et al. 2009, Flechtmann et al. 2009). al. 2012). In each area, we installed two parallel linear The capybara, Hydrochoerus hydrochaeris (L.) (Ro- transects spaced by 20 m from each other. One tran- dentia: Caviidae) is the largest herbivorous rodent on sect received a set of Þve pitfall traps baited with fresh Earth, measuring from 1.0 to 1.3 m in length and the capybara dung (Ϸ 40 g) and the other transect re- weight of adults ranges from 35 to 65 kg (Woods and ceived Þve traps baited with human feces (Ϸ 40 g), Kilpatrick 2005). It is a semiaquatic rodent that in- totaling a sampling effort of 300 traps. The distance habits the margins of water bodies in groups that can between traps within each transect was 100 m to en- reach Ͼ20 individuals (Woods and Kilpatrick 2005). sure independence of the samples (Larsen and For- Capybaras feed exclusively on plants and are distrib- syth 2005). uted from Panama to northern Argentina (Woods and Dung beetles were sampled during 48 h using pitfall Kilpatrick 2005). Several dung beetle species have traps (20 cm in diameter and 15 cm in height) installed been found in the nests of rodents (e.g., Howden et al. at ground level and covered with a plastic lid to reduce 1956, Howden and Cartwright 1963, Halffter and drying of the bait and prevent damage from rainfall. Mathews 1966, Anduaga and Halffter 1991, Anduaga Inside each trap, 250 ml of a saline ϩ neutral detergent 2007), but no studies have been conducted to assess the solution (1.5%) was added, and the baits were placed in community structure of dung beetles attracted to the plastic containers (50 ml) at the center of each trap using feces of the largest herbivorous rodent on the planet. a wire. Fresh capybara dung was collected every morn- Here, we used a wide sampling effort to capture ing along the banks of the Joa˜o Dias creek in the city of dung beetles in pastures with exotic grass (Brachiaria Aquidauana where these animals are commonly found. spp.), patches of the Brazilian savanna (Cerrado), The beetles captured were sent to the Federal and points of degraded riparian forest along the University of Mato Grosso (UFMT; Cuiaba´, Mato Aquidauana river (in Anasta´cio and Aquidauana, Mato Grosso, Brazil) where they were identiÞed by Dr. Grosso do Sul, Brazil) using pitfall traps baited with Fernando Z. Vaz-de-Mello. Vouchers are deposited fresh capybara dung (native herbivores) and human in the State University of Mato Grosso do Sul feces (exotic omnivores) to test the following hypoth- (UEMS; Aquidauana, Mato Grosso do Sul, Brazil) eses: 1) capybara dung attracts lower abundance and and also in the Entomology Section of the Zoolog- richness than human feces in the three systems; and 2) ical Collection of the UFMT. there are differences in community structure and spe- Data Analysis. We performed analyses for each of cies composition attracted to the two feces types in the the three systems separately. We performed a series of three systems. Our null hypothesis is that the number Generalized Linear Models (GLMs) to determine of species, abundance, and species composition at- whether the baits differed with respect to the average tracted to the two feces types is similar in the three number of individuals and species of dung beetles systems. Because humans are omnivores and dung sampled per transect in the three systems. We used the beetles often prefer omnivore dung, we believe that baits as the explanatory variables and richness and the capture will be greater in human feces than in the abundance as response variables. All GLMs were sub- capybara dung. In addition, human feces is among the mitted to residual analysis, so as to evaluate adequacy most attractive types of dung to most of the dung of error distribution (Crawley 2002). All analyses were beetle species (Howden and Nealis 1975), and is avail- performed using the free software R (R Development able at any study site in the world (Larsen and Forsyth Core Team 2013). 2005) being used in most articles with dung beetles Comparisons between the number of accumulated (e.g., Larsen 2012, Korasaki et al. 2013). species per baits in each system were performed vi- sually with the curve of the 95% CI. In an additional attempt to evaluate the efÞciency of our sampling Materials and Methods program, we assessed the completeness of each system Study Sites and Sampling Procedures. The study by calculating the number of observed species (Sobs) was developed in a transitional region of the Cerrado as a percentage of the total species richness, which was and Pantanal ecosystems, in Anasta´cio (20Њ 45Ј45Љ S, estimated based on the average of three abundance- 55Њ 43Ј49Љ W) and Aquidauana (20Њ 27Ј28Љ S, 55Њ 50Ј16Љ based nonparametric estimators: ACE, Chao 1, and W), Mato Grosso do Sul, Brazil, with annual rainfall Jackknife 1, using the formula: Sampling EfÞciency ϭ ranging from 1,200Ð1,300 mm and mean annual tem- [Sobs ϫ 100/((ACEϩChao1ϩJack1)/3)].