Influence of carrion smell and rebaiting time on the efficiency of pitfall traps to dung beetle sampling

Dung beetles (Coleoptera: Scarabaeidae: Scarabaeinae) are very useful insects, as they improve the chemo-physical properties of soil, clean pastures from dung pads, and help control symbovine flies associated with bovine cattle. Their importance makes it fundamental to sample and survey them adequately. The objectives of the present study were to determine the influence of decaying insects trapped in pitfalls on the attractiveness of Moura pig Sus scrofa L. (Suidae) and collared peccary Tayassu tajacu (L.) (Tayassuidae) dung used as baits to lure dung beetles, and to establish how long these baits remain attractive to dung beetles when used in these traps. Some dung beetle species seemed to be able to discriminate against foul smell from decaying insects within the first 24 h, hence decreasing trap efficiency. This was more evident in peccary dung-baited traps, which proved to be the least attractive bait. Attractiveness lasted only 24 h for peccary dung, after which it became unattractive, whereas the pig dung bait was highly attractive for 48 h, after which its attractiveness diminished but was not completely lost.     

Co-declining mammals and dung beetles: an impending ecological cascade

Biodiversity loss can precipitate extinction cascades and impair ecological processes. These 'downstream' effects will be exacerbated if functionally important taxa are tightly linked with species threatened by extinction or population decline. We review the current evidence that such a scenario is currently playing out in the linked declines of persistently hunted mammal populations and the dung beetles communities (Coleoptera: Scarabaeidae: Scarabaeinae) that depend on them for adult and larval food resources. Through a close evolutionary association, mammal assemblages have played a fundamental role in structuring extant dung beetle communities. Today many game mammal species' populations are severely depleted by subsistence or commercial hunting, especially in tropical forest systems. Multiple lines of evidence from temperate and tropical systems indicate that the regional-scale decline or extirpation of medium and large bodied mammal faunas can severely disrupt the diversity and abundance of dung beetle communities through alterations in the composition and availability of dung resources. These observed community disassemblies have significant short- and long-term implications for the maintenance of key ecosystem processes including nutrient recycling and secondary seed dispersal. Identifying the species- and community-level traits that buffer or exacerbate these species and functional responses is essential if we are to develop a better understanding of the cascading ecological consequences of hunting in tropical forests.     

Using dung beetles to evaluate the effects of urbanization on Atlantic Forest biodiversity

We used dung beetles to evaluate the impact of urbanization on insect biodiversity in three Atlantic Forest fragments in Londrina, Paraná, Brazil. This study provides the first empirical evidence of the impact of urbanization on richness, abundance, composition and guild structure of dung beetle communities from the Brazilian Atlantic Forest. We evaluated the community aspects (abundance, richness, composition and food guilds) of dung beetles in fragments with different degrees of immersion in the urban matrix using pitfall traps with four alternative baits (rotten meat, rotten fish, pig dung and decaying banana). A total of 1 719 individuals were collected, belonging to 29 species from 11 genera and six Scarabaeinae tribes. The most urban‐immersed fragment showed a higher species dominance and the beetle community captured on dung presented the greatest evenness. The beetle communities were distinct with respect to the fragments and feeding habits. Except for the dung beetle assemblage in the most urbanized forest fragment, all others exhibited contrasting differences in species composition attracted to each bait type. Our results clearly show that the degree of urbanization affects Atlantic Forest dung beetle communities and that the preservation of forest fragments inside the cities, even small ones, can provide refuges for Scarabaeinae.     

Importance of dung incorporation activity by three species of coprophagous beetle (Coleoptera: Scarabaeidae: Scarabaeinae) macrofauna in pastureland on "La Michilía" biosphere reserve in Durango, Mexico

The dung incorporation activity of three species of coprophagous beetle macrofauna (Coleoptera: Scarabaeidae: Scarabaeinae), Dichotomius colonicus (Say), Phanaeus quadridens (Say), and Copris klugi sierrensis Matthews, was studied on a hectare of pastureland on "La Michilía" Biosphere Reserve. In total, 223 specimens of the three species were recorded. On the basis of abundance, the dominant species was D. colonicus. A substantial percentage (31% in September and 51% in October) of dung pats remained on the surface, untouched by beetles. Most of the specimens observed (92%) were found amassing dung in galleries. The dung accumulation capacity of D. colonicus varied from 5 to 135 g of dung (dry weight)/pair, whereas C. klugi sierrensis amassed an average of 24.3 +/- 8.5 g/pair and P. quadridens amassed an average of 12 +/- 2.5 g/pair. Based on dung burying efficiency, D. colonicus was considered the most beneficial of the three species in the study area. It dug deeper, wider galleries and incorporated the largest amount of dung into the soil.     

Optimising Bait for Pitfall Trapping of Amazonian Dung Beetles (Coleoptera: Scarabaeinae)

The accurate sampling of communities is vital to any investigation of ecological processes and biodiversity. Dung beetles have emerged as a widely used focal taxon in environmental studies and can be sampled quickly and inexpensively using baited pitfalls. Although there is now a wealth of available data on dung beetle communities from around the world, there is a lack of standardisation between sampling protocols for accurately sampling dung beetle communities. In particular, bait choice is often led by the idiosyncrasies of the researcher, logistic problems and the dung sources available, which leads to difficulties for inter-study comparisons. In general, human dung is the preferred choice, however, it is often in short supply, which can severely limit sampling effort. By contrast, pigs may produce up to 20 times the volume. We tested the ability of human and pig dung to attract a primary forest dung beetle assemblage, as well as three mixes of the two baits in different proportions. Analyses focussed on the comparability of sampling with pig or human-pig dung mixes with studies that have sampled using human dung. There were no significant differences between richness and abundance sampled by each bait. The assemblages sampled were remarkably consistent across baits, and ordination analyses showed that the assemblages sampled by mixed dung baits were not significantly different from that captured by pure human dung, with the assemblages sampled by 10% and 90% pig mixes structurally most similar to assemblages sampled by human dung. We suggest that a 10:90 human:pig ratio, or similar, is an ideal compromise between sampling efficiency, inter-study comparability and the availability of large quantities of bait for sampling Amazonian dung beetles. Assessing the comparability of assemblage samples collected using different baits represents an important step to facilitating large-scale meta-analyses of dung beetle assemblages collected using non-standard methodology.     

A comparison of dung beetle (Coleoptera: Scarabaeidae) attraction to native and exotic mammal dung

Although the preference of dung beetles (Coleoptera: Scarabaeidae) for specific types and conditions of dung has been given substantial attention, little has been done to investigate the potential effects of exotic mammal introduction for game farms or rewilding projects. We used pitfall traps baited with various native and exotic herbivore, carnivore, and omnivore dung to evaluate dung beetle preference in the Great Plains of North America. Additionally, we analyzed of the nutrient quality of each dung type. In total, 9,089 dung beetles from 15 species were captured in 2 yr of sampling. We found significant differences (P < 0.05) in mean dung beetle capture among omnivore, herbivore, and carnivore dung, as well as differences in individual species preference for dung type. Omnivore dung was the most attractive with chimpanzee and human dung having the highest mean capture (291.1 ± 27.6 and 287.5 ± 28.5 respectively). Carrion also was highly attractive with a mean of 231.9 ± 20.6 beetles per trap (N = 8). Our results suggest definitive local preference of carrion in Phanaeus vindex Macleay and Onthophagus hecate (Panzer), while the congener, O. pennsylvanicus (Harold), was rarely captured in carrion and highly preferred omnivore dung. Preference for a specific bait type does not appear to be correlated with dung quality, mammalian diet, or origin of mammal. Results suggest niche segregation by dung type among dung beetle species.     

Mitogenome of Coprophanaeus ensifer and phylogenetic analysis of the Scarabaeidae family (Coleoptera)

Several studies about the phylogenetic relationships of the Scarabaeinae subfamily (Coleoptera: Scarabaeidae) have been performed, but some phylogenetic uncertainties persist including the relationship and monophyly of different tribes and some genera. The aim of this study was to characterize the mitogenome of Coprophanaeus ensifer in order to establish its position within the Scarabaeidae family and to contribute to the resolution of some phylogenetic uncertainties. The mitogenome was sequenced on the Illumina HiSeq 4000, assembled using the Mitobim software and annotated in MITOS WebServer. The phylogenetic trees were reconstructed by Bayesian inference. The C. ensifer mitogenome is a molecule of 14,964 bp that contains the number and organization of the genes similar to those of most Coleoptera species. Phylogenetic reconstruction suggests monophyly of the tribe Phanaeini and supports the hypothesis that Coprini is a sister group of Phanaeini. The results also revealed the position of the tribe Oniticellini which is grouped with Onthophagini and Onitini. The geographic distribution of these species that form the most ancestral clade suggests with Scarabaeinae originated in Africa. Keywords: Dung beetle, mitochondrial genome, phylogenomics     

Generalist dung attraction response in a New Zealand dung beetle that evolved with an absence of mammalian herbivores

1. Dung beetles (Scarabaeidae: Scarabaeinae) are integral parts of many ecosystems because of their role in decomposition of dung; particularly mammal dung, which forms the diet of both larvae and adults. 2. New Zealand dung beetles are unusual as they are flightless and evolved on islands with a highly depauperate mammal fauna and thus without the usual dung resource used by dung beetles elsewhere. The diet of New Zealand dung beetles is unknown. 3. We hypothesised (1) that the endemic dung beetle Saphobius edwardsi would be attracted to a broad range of food types, and (2) that S. edwardsi would be able to survive and reproduce on a range of dung types and puriri (Vitex lucens) humus. 4. Laboratory choice tests identified that S. edwardsi was attracted to a range of mammal, bird, invertebrate, and reptile dung types, but not to non‐dung food sources. Five‐month no‐choice tests found that beetle survival rates were lower for beetles fed with humus compared with those fed on mammal, bird, or invertebrate dung. None of the beetles reproduced. 5. This study suggests S. edwardsi have a strong preference for dung, and are likely to be broad dung generalists in their feeding behaviour.     

Relative abundance and species composition of some dung beetles (Coleoptera: Scarabaeinae) in Bangladesh

1. The occurrence and community structure of dung beetle species (Coleoptera: Scarabaeinae) in the faecal deposits of man and some domestic vertebrates varied according to dung type, seasonal conditions and soil type. 2. Of the twenty-four species belonging to six genera, Onthophagus was the prevalent genus and O. ramosellus the most abundant species. 3. Cattle dung was the dominant food type.     

Does an aridity and trophic resource gradient drive patterns of dung beetle food selection across the Botswana Kalahari?

1. Aridity gradients are paralleled by both reductions in resources and decreased species richness of animals. Across the aridity gradient of the Botswana Kalahari, a reduction in mammal species richness leads to reduced density and diversity of dung types, accompanied by reduced dung beetle species richness. We investigated whether this gradient also drives changes in dung beetle food type association and specialisation owing to a loss of some dung types to the arid southwest. 2. Dung beetles were sampled from three study sites in each of the six study areas using 2 × 10 grids of pitfall traps baited with dung (pig, elephant, cattle, and sheep) or carrion (chicken livers). 3. Canonical correspondence analysis (CCA) showed that distributions of dung beetle species between bait types deviated significantly from random associations. 4. Central Kalahari assemblages were more specialist than those at the mesic and arid extremes of the gradient. 5. Patterns of selection and specialisation to bait types differed between mesic northeast and arid southwest study areas. There were specialist faunas on carrion and more generalist faunas on ruminant herbivore dung (cattle and sheep) in each region. However, specialist species associated with elephant dung in the northeast were replaced by a more generalist fauna in the southwest with an opposite trend on pig dung. 6. Reduced species richness and high species turnover from the mesic northeast to the arid southwest is paralleled by a shift in patterns of food association that may reflect changes in the diversity of food types, particularly the absence of elephant dung from the southwest.     

Abrupt species loss of the Amazonian dung beetle in pastures adjacent to species-rich forests

Anthropogenic disturbances, such as conversion of forests to pastures, are considered one of the major causes of Amazonian biodiversity loss. Pastures are hostile environments for forest species that are not adapted to the adverse conditions of open areas. The aim of this study was to analyze the effects of forest replacement by cattle pastures in the north-eastern part of the Amazonian rainforest using dung beetles (Scarabaeidae: Scarabaeinae) as indicators. Therefore, we determined edge effects and differences in dung beetle species richness, composition, and abundance between forest and neighbouring cattle pastures. A total of 6723 individuals of 63 species of dung beetle were collected. In comparison to forests, adjacent pastures were sharply less rich and abundant in dung beetles. The forests had 6604 individuals and 59 species, while the pastures had 119 individuals and 12 species. The replacement of forests by pastures results in an increase of dung beetle species turnover. Only 12% of dung beetle species were shared by both forest and pasture environments. We highlight that most Amazonian dung beetles are forest species sensitive to abrupt human-driven modification of habitats and are only moderately affected by edge effects.     

Dung Beetle Assemblages and Seasonality in Primary Forest and Forest Fragments on Agricultural Landscapes in Budongo, Uganda

Very little is known about the diversity of arthropods in the fast-disappearing fragments of natural forests in sub-Saharan Africa. This study investigated: (1) the influence of forest fragment characteristics on dung beetle species richness, composition, abundance, and diversity; and (2) the relationship between dung beetle assemblages and rainfall pattern. Beetles were sampled through 12 mo using dung baited pitfall traps. A total of 18,073 dung beetles belonging to three subfamilies and 45 species were captured. The subfamily Scarabaeinae was the most abundant (99%) and species rich (89%). Fast-burying tunnellers (paracoprids) were the most dominant functional group. Catharsius sesostris, Copris nepos , and Heliocopris punctiventris were the three most abundant species, and had the highest contributions to dissimilarities between forests. With few exceptions, dung beetle abundance, species richness, and diversity were generally higher in larger forest fragments (100–150 ha) than in smaller ones (10–50 ha) and the nature reserve (1042 ha). Forest fragment size had a highly significant positive relationship with beetle abundance, but only when the nature reserve is excluded in the analysis. Dung beetle abundance and species richness showed direct weak relationships with litter depth (positive) and groundcover (negative) but not tree density, tree species richness, and fragment isolation distance. Dung beetle abundance and species richness were strongly correlated with monthly changes in rainfall. Results of this study indicate that forest fragments on agricultural lands in the Budongo landscape, especially medium-sized (100–150 ha) ones, represent important conservation areas for dung beetles.     

Attractiveness of dung beetles (Coleoptera: Scarabaeinae) to faeces from native mammals in different trophic guilds

Dung beetles are predominantly coprophagous and use mammalian faeces as their main food resource and for offspring rearing. However, these resources are generally ephemeral and scarce. The objective of this study was to verify the attractiveness to dung beetles of the faeces of nine native mammal species of the Cerrado biome in different trophic guilds. This study was conducted during the rainy season (October 2016 to March 2017) in two sensu stricto areas of the Cerrado in the Brasília National Park (PNB), Federal District, Brazil. Pitfalls containing faeces of nine mammals native to the region were used as bait (from three herbivores, three carnivores and three omnivores) for the collection of dung beetles. We collected 203 individuals and 19 species, and the most abundant species were Uroxys aff. thoracalis, Oxysternon palemo, Coprophanaeus spitzi and Diabroctis mirabilis. The most attractive faecal bait was from the omnivorous Chrysocyon brachyurus (32.67% of the individuals and 13 species). This study reveals variation in the attraction of dung beetles to native mammal dung of the Cerrado, with more abundance in the faeces of omnivores and carnivores and less in the faeces of herbivores. The species composition in the carnivore baits did not differ from that of the omnivore baits. These data indicate that changes in Scarabaeinae assemblages result from changes in mammal species such as those that have occurred in the Cerrado biome with the conversion to cattle production, which in recent years has caused intense fragmentation of habitats, expansion of agricultural and livestock activities and the introduction of exotic species. The mammal C. brachyurus, vulnerable to extinction in Brazil, should be a focus for future research, since its faeces support a great diversity of dung beetles in the Cerrado.     

Dung preference and trophic association of dung beetles (Coleoptera: Scarabaeidae) in the moist forests of the South-western Ghats of the Indian subcontinent

First quantitative dung beetle-feeding trophic network analysis for the Oriental region is carried out by investigating trophic network interaction between dung beetles and mammal dung types in the moist forests of the Western Ghats a global biodiversity hot spot in south-western India. Dung-beetle assemblage associated with the dung of the prominent mammals, such as the macaque, boar, gaur, elephant and deer, showed differences in richness, abundance and composition among different dung types. Most dung beetles were generalists with low resource specificity and community-wide generalist feeding on herbivore and omnivore dung types. Dung beetles in the region displayed high species richness and abundance in boar dung. The high attraction and specificity of dung beetles towards the odoriferous boar dung indicate that the omnivore mammal Sus scrofa has a major role in maintaining the dung beetle community in the forests of the Western Ghats. Network interaction analysis shows that the vast majority in the assemblage are generalist species, and the few specialist species were all with low abundance. Low overall specialisation and low resource partitioning with high species richness is recorded in the assemblage. The assemblage's trophic level preference is reflected in the high dung specificity recorded in the omnivore and herbivore dung types.     

Copro-necrophagous beetle (Coleoptera: Scarabaeidae) diversity in an agroecosystem in Yucatan, Mexico

Scarabaeinae are sensitive to structural habitat changes caused by disturbance. We compared copronecrophagous beetle (Scarabaeinae) community structure in three differently managed zones within an agroeco-system of the northern Yucatan Peninsula, Mexico. We placed dung and carrion traps once a month from June 2004 through May 2005. The beetle community included 17 species from the genera Canthon, Canthidium, Deltochilum, Pseudocanthon, Malagoniella, Onthophagus, Phanaeus, Copris, Uroxys, Sisyphus and Ateuchus. The secondary vegetation had a higher beetle diversity than the other two zones. Species richness was highest in the Brosimum alicastrum plantation. The pasture had the lowest species diversity and richness, but exhibited the highest abundance of Scarabaeinae in the dry season. The two zones with extensive tree cover were the most diverse. Roller beetles were dominant over burrower species and small-sized species outnumbered large species. Our data show two important issues: beetle species in the pasture extended their activity to the beginning of the dry season, while abundances dropped in the other, unirrigated zones; and the possibility that the Scarabaeinae living in neotropical forests are opportunistic saprophages and have specialized habits for resources other than dung. The B. alicastrum plantation is beneficial to the entire ranch production system because it functions as a dispersion and development area for stenotopic species limited to tree cover.     

The volatile organic compounds of introduced and native dung and carrion and their role in dung beetle foraging behaviour

1. The decomposition of biological material produces a plethora of volatile organic compounds (VOCs), which are implicated in the foraging behaviour of coprophagous and necrophagous insects. Dung beetles (Coleoptera: Scarabaeidae: Scarabaeinae) have an acute olfactory system used to locate food resources. Accordingly, identification of food resource VOCs potentially used in food location is integral to understanding dung beetle foraging ecology. 2. In this study, volatile emissions from dung and carrion of native and introduced animals in New Zealand were analysed using solid‐phase microextraction (SPME) and gas chromatography‐mass spectrometry (GC‐MS). Volatile profiles were compared via principal component analyses (PCAs) and cluster solutions based on attractiveness using canonical discriminant analysis (CDA). 3. A total of 115 compounds were detected from 21 food types. Statistical analyses showed that dung and carrion volatile profiles clustered according to attractiveness to the dung beetle Saphobius edwardsi, and that different dung types formed distinct clusters and grouped separately from carrion. 4. This study suggests that volatile profiles emitted by food resources used by dung beetles are complex, producing distinct odours, which potentially mediate foraging decisions.     

Response of a dung beetle assemblage along a reforestation gradient in Restinga forest

Scarabaeinae dung beetles are indicator insects used in the evaluation of the ecological effects caused by changes in habitat structure and ecosystem integrity resulting from environmental degradation. We compared dung beetle diversity in conserved restinga forests (coastal tropical moist broadleaf forest) and in reforested areas of various ages during the rainy and dry seasons, on the coast of Paraíba State, Brazil. A total of 3,634 individuals comprising 14 species were collected. In the reforested areas there was a gradual increase in species abundance relative to the area’s age, but in the conserved restinga the abundance of individuals was 10–20 times higher than that recorded in areas of recent reforestation. The highest species richness was found in the conserved restinga and in the oldest reforested area (16 years old) during the rainy season. During the dry season, when environmental conditions do not seem to favor adult survival, most of the species were found in the conserved restinga forest. The dung beetle community structure was related to the increases in habitat heterogeneity in the successional processes of the reforested areas. Our results suggest that reforested areas act as a source of and refuge for dung beetle species.     

Beyond Coprinisphaera: fossil nests of dung beetles

Forty‐two remains of fossil nests of dung beetles (Coleoptera: Scarabaeidae: Scarabaeinae), which were recorded in four formations of the Cenozoic of South America, are described herein for the first time. Most of them are represented by nesting chambers containing a fossil brood ball (Coprinisphaera). However, in the most remarkable cases, parts of the burrows constructed by the parents and/or the vertical emergence burrows constructed by the offspring are preserved too. The preservation of the dung beetle nests is very unusual but more recurrent in younger formations probably because of the short‐term action of the diagenetic processes. Considering that the construction of brood balls always involves the construction of nests, the ichnotaxonomical proposal is that the remains of fossil nests should be considered as ‘structures associated with Coprinisphaera’ and added to the diagnosis to avoid the proliferation of names. The study of the fossil nests provides new palaeoetological inferences for dung beetles, such as the Nesting Pattern of the trace makers of Coprinisphaera tonnii and Coprinisphaera akatanka, and how phylogenetically related were they with the extant necrophagous species of the genera Coprophanaeus and Canthon, respectively. Additionally, the fossil evidence suggests that simple nests of dung beetles predate compound nests.     

Roles of environmental variables and land usage as drivers of dung beetle assemblage structure in mopane woodland

Colophospermum mopane woodland covers large areas of dry lowland savanna in southeastern Africa. Dominant land usage is conservation (45%) with the remainder mostly modified by farming. Dung beetle responses to environment (dung type, habitat, weather) and land usage (conservation, farming, mining) were examined at Phalaborwa (23.9431°S 31.1411°E) in the Phalaborwa‐Timbavati Mopaneveld, South Africa. Partitioning of gamma species richness and diversity showed lower alpha values in mine areas than in farm and conserved areas. However, between‐land usage differences in species richness, alpha diversity, abundance and biomass, showed lower significance than those between dung type and different weather. At two sampling scales, three multivariate techniques variously separated assemblages according to land usage, dung type and weather. Analysis of 21 mean samples separated clusters according to dung type (Canonical Correspondence Analysis, CCA) or mine assemblages, conserved plus farm assemblages on pig plus elephant, or cattle dung (NMDS, Factor Analysis) with shared variance of >80% and unique variance of 16–18% per cluster. In analysis of 188 samples (CCA), each overlapping dung type cluster was offset in ordinal space with congruent patterns of separation according to land usage and weather (drier days distant from moister days; conserved plus farm areas distant from early succession mine areas, which were distant from disturbed and later succession mine areas). Mining, dung types, and moist conditions were the strongest contributors to between‐assemblage differences. Compared with conserved areas, dung beetle diversity is appreciably altered by mining but only slightly altered by intensive game farming or livestock ranching with subsistence agriculture.     

Patterns of alimentary resource use by dung beetles in introduced Brazilian pastures: Cattle versus sheep dung

Dung beetles (Coleoptera: Scarabaeidae) perform essential ecological functions in pastures, such as dung removal, nutrient recycling and parasite control. However, the patterns of alimentary use by dung beetles in introduced Brazilian pastures are poorly known. Here, we compared dung beetle species richness, abundance and species composition in cattle and sheep dung, and identified the dung beetle species preference by each dung type. In January 2019, dung beetles were sampled with pitfall traps baited with cattle and sheep dung in 12 introduced pastures (Urochloa spp.), in Aquidauana, Mato Grosso do Sul, Brazil. A total of 592 individuals belonging to 14 species of dung beetles were collected. Of the 14 species sampled, nine were recorded in both dung types, five were found exclusively in sheep dung and no species was exclusive to cattle dung. Species richness and abundance were higher in sheep dung. Species composition was different between the dung types. Dichotomius bos (Blanchard), Genieridium bidens (Balthasar), Onthophagus aeneus Blanchard and Trichillum externepunctatum Preudhomme de Borre were associated with sheep dung. Our results provide evidence that sheep dung is more attractive to dung beetles with a distinct community species between the two dung types, although the studied pastures have never been used before for sheep breeding. Thus, our data shows that the introduction of a new alimentary resource (e.g. sheep dung) can be an important strategy to help to obtain a more diverse dung beetle assemblage in introduced Brazilian pastures.