Gut microbiota of dung beetles correspond to dietary specializations of adults and larvae

Vertebrate dung is central to the dung beetle life cycle, constituting food for adults and a protective and nutritive refuge for their offspring. Adult dung beetles have soft mandibles and feed primarily on nutritionally rich dung particles, while larvae have sclerotized mandibles and consume coarser dung particles with a higher C/N ratio. Here, using the dung beetles Euoniticellus intermedius and E. triangulatus, we show that these morphological adaptations in mandibular structure are also correlated with differences in basic gut structure and gut bacterial communities between dung beetle life stages. Metagenome functional predictions based on 16S rDNA characterization further indicated that larval gut communities are enriched in genes involved in cellulose degradation and nitrogen fixation compared to adult guts. Larval gut communities are more similar to female gut communities than they are to those of males, and bacteria present in maternally provisioned brood balls and maternal ‘gifts’ (secretions deposited in the brood ball along with the egg) are also more similar to larval gut communities than to those of males. Maternal secretions and maternally provisioned brood balls, as well as dung, were important factors shaping the larval gut community. Differences between gut microbiota in the adults and larvae are likely to contribute to differences in nutrient assimilation from ingested dung at different life history stages.     

A review of the systematic position of the North American species of the genusGlyptotendipes

The systematic position of the North American species of the genusGlyptotendipes is reviewed using morphological characters from all life history stages. The cladistic analysis revealed adult characters which support the monophyly of the three species groups (A, B and C) of PINDER and REISS (1983, 1986) for pupae and larvae. These species groups are given formal subgeneric status, with Group A becomingGlytotendipes, s.s., placingPhytotendipes Goetghebuer, 1937 in synonymy, Group B is namedCaulochironomus (type-species,Tendipes caulicola Kieffer, 1913) and Group C is namedTrichotendipes (type-species,Tendipes signatus Kieffer, 1911).     

Elephant dung decomposition and coprophilous fungi in two habitats of semi-arid Botswana

In order to understand the impact of habitat changes on ecosystem processes caused by increased populations of elephants, elephant dung decomposition was studied in semi-arid Botswana. Dung decomposition rates were studied with and without the presence of arthropods, using pairs of exposed dung and dung enclosed in nylon-mesh bags, respectively. Dung decomposition rates were lower in the absence of arthropods. The rates in the late wet season were higher in the scrubland than in the woodland. In the early dry season, immediately after the wet season, the rates were higher in the woodland than in the scrubland. The difference in decomposition rates between habitats was attributed to microclimatic conditions created by vegetation cover. With regard to fungal succession, Cladosporium cladosporioides and Eurotium brefeldianum occurred only in the late stages of dung decomposition whereas Talaromyces helicus, Cercophora coprophila and Sporormiella minima occurred in all the stages. Although there was no significant difference in Shannon–Weiner fungal species diversity index between habitats, seasons, dung ages and laboratory incubation periods, there were significant differences in fungal community composition between these parameters. Species richness was higher in the late wet season than in the early dry season, indicating the importance of moist conditions for a large diversity of fungal species.     

Acoustic estimation of wildlife abundance: methodology for vocal mammals in forested habitats

Habitat loss and hunting pressure threaten mammal populations worldwide, generating critical time constraints on trend assessment. This study introduces a new survey method that samples continuously and non‐invasively over long time periods, obtaining estimates of abundance from vocalization rates. We present feasibility assessment methods for acoustic surveys and develop equations for estimating population size. As an illustration, we demonstrate the feasibility of acoustic surveys for African forest elephants (Loxodonta africana cyclotis). Visual surveys and vocalizations from a forest clearing in the Central African Republic were used to establish that low‐frequency elephant calling rate is a useful index of elephant numbers (linear regression P < 0.001, r_adj.² = 0.58). The effective sampling area was 3.22 km² per acoustic sensor, a dramatic increase in coverage over dung survey transects. These results support the use of acoustic surveys for estimating elephant abundance over large remote areas and in diverse habitats, using a distributed network of acoustic sensors. The abundance estimation methods presented can be applied in surveys of any species for which an acoustic abundance index and detection function have been established. This acoustic survey technique provides an opportunity to improve management and conservation of many acoustically‐active taxa whose populations are currently under‐monitored.     

How much Dillenia indica seed predation occurs from Asian elephant dung?

Elephants are thought to be effective seed dispersers, but research on whether elephant dung effectively protects seeds from seed predation is lacking. Quantifying rates of seed predation from elephant dung will facilitate comparisons between elephants and alternative dispersers, helping us understand the functional role of megaherbivores in ecosystems. We conducted an experiment to quantify the predation of Dillenia indica seeds from elephant dung in Buxa Reserve, India from December 2012 to April 2013. Using dung boluses from the same dung pile, we compared the number of seeds in boluses that are a) opened immediately upon detection (control boluses), b) made available only to small seed predators (<3 mm wide) for 1–4 months, and c) made available to all seed predators and secondary dispersers for 1–4 months. Using a model built on this experiment, we estimated that seed predation by small seed predators (most likely ants and termites) destroys between 82.9% and 96.4% of seeds in elephant dung between the time of defecation and the median germination date for D. indica. Exposure to larger seed predators and secondary dispersers did not lead to a significant additional reduction in the number of seeds per dung bolus. Our findings suggest that post-dispersal seed predation by small insects (<3 mm) substantially reduces but does not eliminate the success of elephants as dispersers of D. indica in a tropical moist forest habitat.     

Small-scale dung survey reveals high forest elephant density and preference for mixed species forest in an intact protected area

Understanding the ecological factors influencing African forest elephant (Loxodonta cyclotis) abundance and distribution is crucial for their conservation management in Central Africa. Dung surveys have been conducted at the landscape scale and confirmed the overwhelming impact of anthropogenic activities on forest elephants. We present results from a small-scale survey in a pristine protected area without anthropogenic activities to elucidate the ecological factors influencing forest elephant density. We conducted a line transect dung survey in a small study area (110 km²) around Mbeli Bai, a natural forest clearing in the Nouabalé-Ndoki National Park, Republic of Congo, and compare results with a landscape survey conducted during the same period. We used habitat specific dung decay data collected on site to estimate elephant density using distance sampling. We fitted Generalized Additive Models to elephant dung encounter rate using explanatory variables collected during the transect survey and from geospatial data. The small-scale survey revealed a precise estimate of forest elephant density that was twice as high as the result from a landscape survey with higher density in mixed species forest for the small-scale survey. We could not find an impact of the proximity of forest clearings and proximity to rivers at the small scale. Fine-scale habitat features, e.g. degree of canopy and understorey closure, had little explanatory power for elephant dung encounter rate. Small-scale dung surveys are a useful method to reveal spatio-temporal variation in forest elephant density and distribution which can inform conservation practitioners in a timely manner. Combining monitoring methods at various spatial scales improves our knowledge and conservation efforts of forest elephants. Nouabalé-Ndoki National Park is a stronghold for forest elephants and of global importance for their conservation.

     

Evaluation of Metarhizium anisopliae for the control of Culicoides brevitarsis Kieffer (Diptera: Ceratopogonidae), the principal vector of bluetongue virus in Australia

Four isolates of the entomopathogenic fungus Metarhizium anisopliae were tested for their potential to control the biting midge Culicoides brevitarsis, the principal vector of bluetongue virus in Australia. Adult C. brevitarsis died three to eight days after walking on paper substrate treated with 0.7 g/m² conidia of any of the isolates, indicating that M. anisopliae has potential as a surface treatment or topical application control strategy. Incorporation of the fungus into freshly excreted cattle dung at rates of between 0.25 and 1 g conidia/kg reduced the emergence of adult midges by up to 98.5% compared to untreated dung indicating that M. anisopliae has the potential to control C. brevitarsis larvae in cattle dung. Three of the isolates produced similar mortality rates on adult and immature C. brevitarsis while the fourth isolate produced lower, but still significant, mortality rates on adult and immature stages.     

<Emphasis Type="Italic">Ophiocordyceps pulvinata</Emphasis> sp. nov., a pathogen of ants with a reduced stroma

Ophiocordyceps pulvinata, a pathogen of ants, is formally described as a new species. Genus level designation of this species is difficult due to several apparently conflicting morphological and ecological characters. Affinity with Ophiocordyceps is suggested by the dark color stroma and ascospore morphology. However, the species was included in a book of entomopathogenic fungi of Japan as Torrubiella sp. due to the production of perithecia on an astipitate stroma. Phylogenetic analyses of molecular data support a close relationship with O. unilateralis, a finding consistent with morphological characteristics of the color, asci and ascospores and ecological traits of host affiliation. Thus, O. pulvinata represents another example of the loss of stipe for the hypocrealean arthropod pathogenic fungi and highlights the utility of asci and ascospore morphology as taxonomically informative characters of closely related taxa.     

从泰国进口椰色果中截获南方凹头臭蚁

依据形态学特征,对截获自泰国进口椰色果上的凹头臭蚁进行种类鉴定。经鉴定,该批凹头臭蚁为南方凹头臭蚁Philidris notiala ZhouZheng,经中国检验检疫科学研究院动植物检疫信息资源共享服务平台查询,系全国口岸首次截获。南方凹头臭蚁由周善义等人于1998年描述,此前仅知分布于中国广西的北流市,本次从来自泰国的椰色果上检出,是其在泰国分布的首次报道,是泰国新记录种。通过记述南方凹头臭蚁工蚁的形态特征,首次描述了该种雌蚁形态特征,给出了其与近缘种的区别,列出了世界凹头臭蚁属名录及分布信息,拍摄了工蚁和雌蚁成虫整体图,为该种的检疫鉴定、分布调查及防治提供参考。     

Adult and early‐stage characters of Brassolini contain conflicting phylogenetic signal (Lepidoptera,Nymphalidae)

This study examines the contribution of early‐stages and adult characters to the reconstruction of the phylogeny of Brassolini butterflies. Parsimony analyses used both equal weights and implied weights, and a series of analyses were performed. First, we analysed adult and early‐stages partitions independently and in combination for a subset of 27 species; in these cases the matrices were mostly complete. Whereas the adult partition alone produced a topology that was well resolved and congruent with previous studies, the early‐stages partition produced a poorly resolved tree under equal weights. Furthermore, implied weights produced a well‐resolved early‐stages topology that differed significantly from the adult topology. When both partitions were combined for 27 species, implied weights yielded a topology that resembled the adult tree except for the positions of Bia and Penetes, but statistical node support was generally lower. This suggests that stochastic noise increased when early‐stage characters were added to the adult partition, but the combined partitions topology was not statistically different from that based on adult characters alone. Second, given that preserved early stages are not as readily available as adults, we analysed a matrix including 45 species in which early‐stage data were missing for 18 species, and compared the topology to that produced by the adult partition alone. Results were similar to the analyses including fewer species; the combined partitions tree was similar to that from the adult partition except for the position of Bia and Penetes. We compare our findings to other genus‐level phylogenetic studies within Lepidoptera that have also used early‐stages and adult characters.     

The influence of landscape structure on ants and dung beetles diversity in a Mediterranean savanna—Forest ecosystem

Ants and dung beetles are focal indicators of change in several ecological processes and successional vegetation stages in Mediterranean landscapes. Despite relatively good knowledge of local species distributions, there are few data on their distributions at different scales. In the present study, the influence of multiscale landscape structure was examined using both ants and dung beetles to identify species that can serve as indicators and detectors of changes in vegetation structure. Multiscale analysis is necessary to explore the different roles of indicator and detector species for use as tools in studies focused on monitoring ecological changes. The study area was the Cabañeros National Park, in the center of the Iberian Peninsula. This site was selected because it is a good setting to evaluate the effects of typical vegetation mosaics on Mediterranean species at different scales. In this study, dung beetles and ants were trapped for one year. A multiscale analysis was designed using three different vegetation habitats (forest, scrubland and grassland) and landscape matrices (woodland, scrubland and grassland). Among dung beetles, 23 indicator species (IndVal values higher than 70%) were found at different scales of analysis. In addition, 20 dung beetle species were characterized as detector species (IndVal values between 45 and 70%) for the three different levels analyzed. Similar to the dung beetles, the ants had different species assemblages at various habitats and landscape levels; however, no indicator ant species were found in this study. All species with significant IndVal values (n = 8) were identified as detector species. Thus, ant and dung beetle assemblages were influenced in different ways by vegetation structure. Both groups showed strong individual species responses to different Mediterranean landscape conditions and vegetation types. Further, both insect groups showed a great number of detector species, which can be useful in ecosystem management because they have varying degrees of preference and sensitivity for different ecological states (such as successional vegetation stages). The diverse indicator and detector species identified in this work could be useful tools for the detection of landscape structure changes at both levels habitat patches and landscape matrix. However, a generalisation of the results at landscape scale should be taken with precaution, but they encourage to study more regions and similar landscapes. The use of more than one indicator group in the analysis confirms the importance of selecting groups with different sensitivities at varying spatial scales as well as different ecosystem functions. This strategy allows the establishment of a clear baseline with which to analyze future direct and indirect impacts of management in Mediterranean protected areas.     

Do pygidial secretions of dung beetles have the potential to repel urban pest ants?

Various organisms emit malodorous secretions against competitors, and the potential use of these secretions in pest management should be investigated. For example, some ant species feed on similar resources as dung beetles, which might have led to counter chemical defences in dung beetles. We tested whether pygidial secretions of the dung beetle Canthon smaragdulus (Fabricius) (Coleoptera: Scarabaeidae, Scarabaeinae) alter the locomotor behaviour of the exotic urban pest ant Tapinoma melanocephalum (Fabricius) (Hymenoptera: Formicidae), specifically whether these secretions repel those ants. We also tested whether the disturbance in the locomotor behaviour of T. melanocephalum increases with the amount of pygidial secretion. We found that individual T. melanocephalum displayed changes in their locomotor behaviour when exposed to pygidial secretions of coupled dung beetles, single males, and single females. Additionally, the pygidial secretions from male and female dung beetles could repel ants. The change in the locomotor behaviour of T. melanocephalum increased with the amount of pygidial secretion. Our results suggest that the pygidial secretions of dung beetles have potential as a biological repellent of T. melanocephalum. Hence, pygidial secretions from dung beetles may be used in the future for the development of urban pest management strategies.     

Evolution of the Scarabaeini (Scarabaeidae: Scarabaeinae)

Abstract. A phylogenetic analysis of the Scarabaeini, based on 244 morphological characters, including 154 multistate and three biological characters, is presented. Tree topologies generated from unweighted data and some weighted algorithms are similar and support only two clades in the tribe representing the genera Scarabaeus L. and Pachylomerus Bertoloni. The only supported subordinate groups treated in this paper as subgenera are Kheper Kirby stat. nov. , Pachysoma MacLeay, Scarabaeolus Balthasar and Sceliages Westwood stat. nov. Drepanopodus Janssens syn. nov. is synonymised with Scarabaeus and six additional names, Madateuchus Paulian, Mnematidium Ritsema, Mnematium MacLeay, Neateuchus Gillet, Neomnematium Janssens and Neopachysoma Ferreira, remain synonyms. A single origin of flightlessness is supported with the subgenus Pachysoma, the most derived lineage in this clade. Rolling dung balls backwards is the ancestral behaviour and predominant mode of food relocation in Scarabaeini, although tunnelling, forward pushing, and carrying also are utilized by some lineages. Pushing food has evolved independently in Sceliages species and S. galenus (Westwood) and a novel mode of forward food relocation by dragging evolved in the subgenus Pachysoma. Feeding on wet dung is the plesiomorphic condition and maintained by the majority of species in the tribe. The most unusual feeding behaviours in the tribe are represented by the obligate millipede-feeding species of Sceliages and the dry dung pellet and/or detritus feeders of Pachysoma.     

Sperm ultrastructure of five species of the Neotropical ant genus Pseudomyrmex (Hymenoptera: Formicidae)

The seminal vesicles of adult males of five species of Pseudomyrmex were prepared for light and transmission electron microscopy. The Pseudomyrmex spermatozoa are long and slender with similar morphology. The head region has an acrosome and a nucleus. In all the studied species, two morphologically distinct types of acrosomal vesicles were observed, a long structure, as observed in all known ants, and a pear‐shaped one, never before observed in ants. The nucleus is elongated and both condensed and loose chromatin are present. The flagellum has an axoneme, a centriolar adjunct, two mitochondrial derivatives and two accessory bodies. The centriolar, the mitochondrial derivatives and the accessory bodies are similar to observations in most ant species that have been studied. The axoneme presents an uncommon 9 + 9 + 1 microtubule arrangement and the central microtubule has 13 protofilaments. The acrosomal dimorphism and the different levels of chromatin organization are exclusive characteristics of Pseudomyrmex. Furthermore, the 9 + 9 + 1 microtubule arrangement is different from all Hymenoptera, as well as from most insects, which present a 9 + 9 + 2 arrangement. These new morphological characters that are specific to Pseudomyrmex, are valuable synapomorphies of the genus and can be used in taxonomic characterization of the Pseudomyrmecinae subfamily and in phylogenetic analyses in Formicidae family.     

Elongated hindguts in desert‐living dung beetles (Scarabaeidae: Scarabaeinae) feeding on dry dung pellets or plant litter

Most adult dung beetles (Scarabaeidae: Scarabaeinae) feed on fresh, wet dung of larger herbivorous or omnivorous mammals. As refractory plant fragments are selected out before ingestion, the food is presumed easily digestible. However, members of the desert‐living scarabaeine genus Pachysoma (probably evolved from an ancestor closely related to the wet‐dung feeding genus Scarabaeus) select dry dung pellets and/or plant litter. Thus, they ingest a much higher proportion of structural plant material, which nevertheless appears to be digested rather efficiently. This study investigates morphological modifications of the gut for this digestion in adults of eight Pachysoma species, both pellet and litter feeders. To ascertain hypothesized ancestral conditions, four fresh‐dung feeding Scarabaeus species were also examined. The latter have the usual dung beetle gut consisting of a long, simple midgut, followed by an equally simple, much shorter hindgut of the same width. Lengths of midguts (M) and hindguts (H) divided by body length (B) for comparison between species of different size are: 4.9–6.3 (M/B) and 0.7–0.8 (H/B), which is normal for dung feeders. In Pachysoma, lengths are 6.3–6.5 (M/B) and 1.0–1.4 (H/B) in pellet feeders, and 4.4–5.0 (M/B) and 2.0–2.5 (H/B) for litter feeders. Hindguts are still morphologically undifferentiated and of midgut width, but clearly longer, particularly in litter feeders. Presumably, plant fragments in the food are digested, at least partly, in the hindgut. If so, the morphological adaptation is unusual: simple elongation rather than the expansion of part of the hindgut, as found in several other plant‐ or detritus‐feeding scarabaeids. J. Morphol. 2013. © 2013 Wiley Periodicals, Inc.     

Species‐level predation network uncovers high prey specificity in a Neotropical army ant community

Army ants are among the top arthropod predators and considered keystone species in tropical ecosystems. During daily mass raids with many thousand workers, army ants hunt live prey, likely exerting strong top‐down control on prey species. Many tropical sites exhibit a high army ant species diversity (>20 species), suggesting that sympatric species partition the available prey niches. However, whether and to what extent this is achieved has not been intensively studied yet. We therefore conducted a large‐scale diet survey of a community of surface‐raiding army ants at La Selva Biological Station in Costa Rica. We systematically collected 3,262 prey items from eleven army ant species (genera Eciton, Nomamyrmex and Neivamyrmex). Prey items were classified as ant prey or non‐ant prey. The prey nearly exclusively consisted of other ants (98%), and most booty was ant brood (87%). Using morphological characters and DNA barcoding, we identified a total of 1,103 ant prey specimens to the species level. One hundred twenty‐nine ant species were detected among the army ant prey, representing about 30% of the known local ant diversity. Using weighted bipartite network analyses, we show that prey specialization in army ants is unexpectedly high and prey niche overlap very small. Besides food niche differentiation, we uncovered a spatiotemporal niche differentiation in army ant raid activity. We discuss competition‐driven multidimensional niche differentiation and predator–prey arms races as possible mechanisms underlying prey specialization in army ants. By combining systematic prey sampling with species‐level prey identification and network analyses, our integrative approach can guide future research by portraying how predator–prey interactions in complex communities can be reliably studied, even in cases where morphological prey identification is infeasible.     

Making the cryptic visible – resolving the species complex of Aphodius fimetarius (Linnaeus) and Aphodius pedellus (de Geer) (Coleoptera: Scarabaeidae) by three complementary methods

Species in cryptic complexes are, per definition, difficult to identify using morphological characters. One such complex was recently detected in the dung beetle Aphodius fimetarius (Linnaeus) sensu lato, an abundant dung beetle with a wide distribution. While the two component taxa, Aphodius fimetarius sensu stricto and Aphodius pedellus (De Geer) exhibit distinctly different karyotypes, the validity of subtle morphological characters proposed to distinguish between them has been debated. Given the variability and minor interspecific differences in external characters, the large‐scale distribution of respective taxa has remained unknown, as have potential differences in ecology and habits. In this study, we ask how A. fimetarius and A. pedellus can best be distinguished, whether the use of different types of characters (karyotypes, DNA sequences and morphological traits) results in consistent species identification, where these species occur and whether they exhibit ecological differences. In total, we inspected a material of 4401 individuals from across the globe, of which 183 were examined for both mtDNA sequences and morphology, 154 for both morphology and karyotype, and 9 (including the recently proposed neotype of Aphodius fimetarius) for all three types of characters. As a marker gene, we sequenced a 590 bp region of the cytochrome c oxidase I gene for 183 individuals. Overall, DNA sequences offered a clear‐cut distinction between taxa: sequences of A. fimetarius and A. pedellus differed by an average pairwise distance of 8.2%, whereas variation within species was only 0.9% for A. fimetarius and 0.5% for A. pedellus. Morphological and chromosomal characters offered species identifications consistent with that of molecular characters: karyotypes identified as A. pedellus consistently fell within one of the molecular clades, whereas karyotypes identified as A. fimetarius fell within the other clade. Likewise, the majority of individuals identified by morphological characters were assigned to the same species by sequence‐based characters. Both taxa thus defined were found to be Holarctic in distribution, with major sympatry within Central and Southern Europe and mixed patterns of sympatry within the US. Northern areas of Europe, Asia and North America are dominated by A. pedellus alone. Within A. pedellus, patterns of sequence diversity were indicative of a recent population expansion. In the western US, the phenology of a population of A. fimetarius was observed to significantly differ from that of a sympatric population of A. pedellus, thereby revealing an ecological difference between the two cryptic taxa. Overall, we conclude that all types of characters offered a consistent classification of the two species. Thus, the laborious karyotyping techniques used to originally establish the presence of two cryptic taxa can now be substituted by characters more easily applied to large ecological samples. Using this approach of integrative taxonomy, we were able to establish the global distribution and species‐specific ecology of these ecologically important cryptic taxa. This published work has been registered in ZooBank, http://zoobank.org/urn:lsid:zoobank.org:pub:4033473E-8BF7-40F4-852D-916E4F858593 .     

The effect of some ecological factors on the intestinal parasite loads of the Asian elephant (Elephas maximus) in southern India

Some ecological factors that might potentially influence intestinal parasite loads in the Asian elephant (Elephas maximus Linn.) were investigated in the Nilgiris, southern India. Fresh dung samples from identified animals were analysed, and the number of eggs/g of dung used as an index of parasite load. Comparisons across seasons and habitats revealed that parasite loads were significantly higher during the dry season than the wet season, but were not different between the dry-deciduous and dry-thorn forests in either season. After accounting for the effect of age on body condition, there was no correlation between body condition, assessed visually using morphological criteria, and parasite load in either season. Individuals of different elephant herds were not characterized by distinct parasite communities in either season. When intra-individual variation was examined, samples collected from the same individual within a day differed significantly in egg densities, while the temporal variation over several weeks or months (within a season) was much less. Egg densities within dung piles were uniform, enabling a simpler collection method henceforth.     

Out of South America: phylogeny of non‐biting midges in the genus Labrundinia suggests multiple dispersal events to Central and North America

Non‐biting midges of the genus Labrundinia (Chironomidae: Tanypodinae) are minute dipterans with immature stages living in a variety of unpolluted water bodies, from small streams and ponds to lakes and bays. Extensively recorded in ecological studies, the genus comprises 39 species, all except one described from areas outside the Palearctic region. Internal structure among Labrundinia species was postulated by S. S. Roback, who recognized four species groups based on morphological characters of immature stages. We examined phylogenetic relationships among known Labrundinia species using partial DNA sequences of the nuclear protein‐coding gene CAD and morphological characters. Both analyses with Bayesian inference and parsimony methods recovered the monophyly of Labrundinia, strongly supported by five morphological synapomorphies. Internal relationships within the genus partly supported Roback's species groups with the addition of later described species. Biogeographical inferences were obtained by applying Bayesian binary MCMC (BBM) analysis and favoured a scenario where Labrundinia had its initial diversification in the Neotropical region and that current presence in the Nearctic region and southern South America is due to subsequent dispersal.     

What do dung beetles eat?

Abstract.  1. Most adult coprophagous beetles feed on fresh dung of mammalian herbivores, confining ingestion to small particles with measured maximum diameters from 2–5 to 130 μm, according to body size and kind of beetle. This study explores benefits and costs of selective feeding in a 'typical' dung beetle with a maximum diameter of ingested particles (MDIP) of 20 μm.
2. Examined dung types (from Danish domestic sheep, cattle and horse, and African wild buffalo, white rhino and elephant) contained 76–89% water. Costs of a 20 μm MDIP were often low, since 69–87% of the total nitrogen in bulk dung other than that of elephant and rhino (40–58%) was available to selective feeders.
3. Nitrogen concentrations were high – and C/N ratios low – in most types of bulk dung compared with the average food of terrestrial detritivores or herbivores. Exceptions were elephant and rhino dung with low nitrogen concentrations and high C/N ratios.
4. Estimated C/N ratios of 13–39 in bulk dung (sheep–elephant) were decreased by selective feeding to 7.3–12.6 in the ingested material. In assimilated food, ratios are probably only 5–7, as most assimilable nitrogen and carbon may be of microbial origin. If so, the assimilable food contains a surplus of nitrogen relative to carbon.
5. The primary advantage of selective feeding, particularly in dung with a high C/N ratio, may be to concentrate assimilable carbon in the ingested food. Effects of changing the MDIP within 20–106 μm are modest, especially in dung with a low C/N ratio.