Changes in the dynamics of functional groups in communities of dung beetles in Atlantic forest fragments adjacent to transgenic maize crops

We investigated the composition and structure of dung beetle communities (Coleoptera, Scarabaeidae, Scarabaeinae) inhabiting areas of forest fragments next to either conventional or Bt-transgenic maize crops. The purpose of the study was to examine possible impacts of transgenic plants on non-target organisms associated with mammals through their food chain. In February 2011, we collected a total of 1502 beetles belonging to 33 species in Campos Novos, Santa Catarina state (SC), Brazil. Beetles were captured using 200 pitfall traps distributed among 20 forest fragments, 10 fragments in each site type (adjacent to conventional vs. Bt crops). In the fragments adjacent to conventional maize, 805 dung beetles from 27 species were collected. In the fragments adjacent to Bt-transgenic maize, 697 dung beetles from 27 species were caught. Dung beetle community composition was affected by fragment size and environmental complexity, and by distance between fragments. However, it did not explain the differences related to the two crop types, i.e., the functional group of dwellers was significantly over-represented in the fragments surrounded by transgenic maize. Hence, the dweller species Eurysternus francinae and Eurysternus parallelus were more frequent and abundant in fragments located near the transgenic maize, while the tunneler species Onthophagus tristis, Uroxys terminalis, Ontherus sulcator and the roller species Canthon lividus seminitens were more frequent and abundant in fragments surrounded by conventional maize. This observed impact of transgenic crops on functional group dynamics within dung beetle communities could potentially lead to impaired capacity for feces removal, seed dispersal, edaphic aeration, and incorporation of organic matter in the soil in these areas, as such ecosystem services are not performed by the dominant functional group (i.e., dwellers).     

Interspecific and intraspecific variation in diet preference in five Atlantic forest dung beetle species

1. Dung beetles are commonly assumed to be generalist feeders, but there has been limited work in identifying whether there is interspecific variation in feeding preference. Equally, there has been no work exploring whether generalist feeding behaviour in a species is a result of within‐species specialisation. 2. This study identified the individual and species‐level feeding preferences of five dung beetle species towards human, jaguar and pig dung using a choice experiment. 3. It was found that species varied in their preference for the dung types, but there was no evidence that within a species, individual beetles varied in their dung choice. These findings were similar to results from field experiments that are more typically used to assess feeding preferences in dung beetles. 4. The results suggest that individual specialism in feeding may not be common in dung beetles. However, there is variation in feeding preferences among species, which is often overlooked and can have implications for the ecosystem functions they provide. It is suggested that choice arenas can be used to assess feeding preferences in dung beetle species that are not abundant enough to reliably estimate dietary choice from field studies.     

Dung beetle-megafauna trophic networks in Singapore’s fragmented forests

We investigated trophic networks between dung beetles and megafauna species in five forest fragments in Singapore varying in size and isolation. We found that Singapore's dung beetle communities were attracted to extant and extinct dung types from different dietary groups. All forest fragment networks were similar and displayed high generalism and high nestedness.     

Co‐declining mammal–dung beetle faunas throughout the Atlantic Forest biome of South America

The millennial–scale evolutionary relationships between mammals and dung beetles have been eroded due to several drivers of contemporary biodiversity loss. Although some evidence of co‐decline has been shown for mammals and dung beetles at some Neotropical sites, a biome‐scale analysis for the entire Atlantic Forest of South America would strengthen our understanding of how relictual sets of mammal species can affect dung beetle co‐occurrences and co‐declines. We therefore collated hundreds of assemblages of both dung beetles and medium‐ to large‐bodied mammals throughout the world's longest tropical forest latitudinal gradient to examine to what extent mammal assemblages may exert a positive influence on dung beetle species composition and functional assembly, and whether this relationship is scale dependent. We also collated several climatic and other environmental variables to examine the degree to which they shape mammal–dung beetle relationships. The relationships between local mammal and dung beetle faunas were examined using regression models, variation partitioning, dissimilarity indices and ecological networks. We found a clear positive relationship between mammal and dung beetle species richness across this forest biome, indicating an ongoing process of mammal–dung beetle niche‐mediated co‐decline. We found a strong relationship between the species composition of both taxa, in which dung beetle species dissimilarity apparently track changes in mammalian dissimilarity, typically in 80% of all cases. Co‐variables such as phytomass and climatic variables also influenced mammal–dung beetle patterns of co‐decline along the Atlantic Forest. We conclude that dung beetle diversity and community assembly are shaped by the remaining co‐occurring mammal assemblages and their functional traits, and both groups were governed by environmental features. We emphasize that ecosystem‐wide effects of mammal population declines remain poorly understood both quantitatively and qualitatively, and curbing large vertebrate defaunation will ensure the persistence of co‐dependent species.     

Dung beetle persistence in human-modified landscapes: Combining indicator species with anthropogenic land use and fragmentation-related effects

Identifying and making use of ecological indicators becomes an essential task in the conservation of tropical systems, mainly in fragmented landscapes where land use intensification and habitat loss are confounding factors in the detection of species’ responses to human-caused disturbance. We aimed to analyze the importance of anthropogenic land use and fragmentation-related effects on dung beetle (Coleoptera: Scarabaeinae) persistence according to the interior–exterior non-linear gradient (forest + matrix) in a fragmented Atlantic Forest landscape used to sugar cane production and cattle ranching/farming. We offer scores for a comprehensive set of community-level attributes, from beetle abundance to taxonomic and ecological composition (i.e. species body size), including a list of indicator species of different forest habitats and adjacent matrix. Dung beetles were surveyed by traps across forest interiors (i.e. core forest areas) and edges of a primary forest, small fragments, sugar cane fields and pastures in a total of 60 sites. Indicator analyses were conducted across the landscape, using two well-established methods (IndVal and SIMPER). Our results suggest that (1) cross-habitat taxonomic distinctness is associated with the presence of indicator species, (2) some species benefit or are dependent of open habitats created by human-disturbances, such as forest edges (e.g. Canthon nigripennis) and matrices (e.g. Canthon aff. piluliformis, Dichotomius nisus and Trichilum externepunctatum), (3) although landscape habitats exhibit reduced beta diversity, dung beetle assemblages are spatially organized in response to the presence of both forest habitats and matrix and fragment area, (4) forest interior supports beetle assemblages biased toward large-bodied species, (5) accordingly forest interior, forest edges and matrix support taxonomically distinct assemblages, both contributing to the bulk of species richness at landscape level, (6) the response of dung beetles to the interior–exterior non-linear gradient (i.e. forest edge + matrix) reveals a similar pattern regardless of the nature of the matrix, and (7) there is no within-habitat variation in beetle abundance and species richness associated with distance from forest edge. Given that there is a high number of forest-dependent or forest-interior specialist species (e.g. Aphengium aff. sordidum, Ateuchus aff. alipioi, Dichotomius mormon, Ontherus aff. erosus and Onthophagus aff. clypeatus) dung beetle persistence in human-modified landscape is highly dependent on the presence of core areas, although edge-affected and matrix habitats may be complementary. This information is essential to permit a better prospect for dung beetle persistence in human-modified landscapes as they continue to move toward edge-dominated landscapes with intensively managed matrices.     

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.     

Edaphic and climatic history has driven current dung beetle species pool and assemblage structure across a transition zone in central South Africa

We investigate biogeographical, regional and sub‐regional‐scale responses of scarabaeine dung beetles to late Cenozoic changes in edaphic and climatic character that created a Savanna/Karoo transition zone in the Northern Cape, South Africa. Across a 50 200 km² study area, the Northern Cape species pool comprised six biogeographical groups defined from distribution across southern Africa. These species groups contributed in different proportions to five regional assemblages defined from structural differences across the transition zone. Towards transition zone peripheries, regional assemblage structure was more strongly correlated to sandiness dating from Miocene to Pliocene deposition (Kalahari), aridity dating from Pliocene to Pleistocene climatic change (Bushmanland Karoo), or cooler temperatures dating from Miocene to Pliocene uplift (Upper Karoo). Correlates of sub‐regional assemblages trended to intensification of dominant drivers towards regional peripheries. Drivers of central transition zone, regional assemblages (‘Gariep Karoo’, ‘Gariep Stony Karoo’) showed no dominance. Biogeographically, endemism dominates the Northern Cape transition zone: south‐west arid groups in Nama Karoo regions; Kalahari plus north‐east savanna groups in the Kalahari. Regionally, transition drives assemblage structure: unique variance, 60% in the Kalahari, 21–30% in four Nama Karoo regions; shared variance (overlap), 25–65% between Kalahari and warmer Karoo regions, 11–71% between mainly cooler Karoo regions.     

The effects of land use change on native dung beetle diversity and function in Australia's Wet Tropics

The impacts of land use change on biodiversity and ecosystem functions are variable, particularly in fragmented tropical rainforest systems with high diversity. Dung beetles (Scarabaeinae) are an ideal group to investigate the relationship between land use change, diversity and ecosystem function as they are easily surveyed, sensitive to habitat modification and perform many ecosystem functions. Although this relationship has been investigated for dung beetles in some tropical regions, there has been no study assessing how native dung beetles in Australia's tropical rainforests respond to deforestation, and what the corresponding consequences are for dung removal (a key ecosystem function fulfilled by dung beetles). In this study we investigated the relationship between dung beetle community attributes (determined through trapping) and function (using dung removal experiments that allowed different dung beetle functional groups to access the dung) in rainforest and cleared pasture in a tropical landscape in Australia's Wet Tropics. Species richness, abundance and biomass were higher in rainforest compared to adjacent pasture, and species composition between these land use types differed significantly. However, average body size and evenness in body size were higher in pasture than in rainforest. Dung removal was higher in rainforest than in pasture when both functional groups or tunnelers only could access the dung. Increased dung removal in the rainforest was explained by higher biodiversity and dominance of a small number of species with distinct body sizes, as dung removal was best predicted by the evenness in body size of the community. Our findings suggest that functional traits (including body size and dung relocation behaviour) present in a dung beetle community are key drivers of dung removal. Overall, our results show that deforestation has reduced native dung beetle diversity in Australian tropical landscapes, which negatively impacts on the capacity for dung removal by dung beetles in this region.     

Functional implications of temperature‐correlated colour polymorphism in an iridescent,scarabaeine dung beetle

Abstract 1. In many Coleoptera, iridescent colouration is generated by exoskeleton ultra‐structure, within which multilayer interference reflects only certain wavelengths. Published work indicates that the colour polymorphism shown by some iridescent beetles is under genetic control. However, the present study suggests environmental involvement in the polymorphic southern African dung beetle, Gymnopleurus humanus Macleay. 2. At 24 study sites across a 1000‐km latitudinal temperature gradient, population samples of G. humanus were dominated by blue individuals in the cooler south, by cupreous individuals in the warmer north, and by locally co‐occurring blue, green and cupreous individuals in intermediate situations. 3. Using digital reflectance spectrophotometry, we measured wavelength intensity values across the visible spectrum (400–800 nm) and used the 70 measured specimens to estimate maximum reflectance from a further 3912 beetles. Differences in mean reflectance values between 24 populations were strongly correlated with average annual temperatures at study sites. 4. Much stronger correlations between mean reflectance values and average temperatures of the cool dry season months suggest that the cross‐climatic patterns may be related to interaction between breeding seasonality and development under different cooler temperatures. 5. Published evidence suggests that inherent physical properties of cholesteric liquid crystals and their responses to different thermal conditions could, potentially, generate the different exocuticle ultra‐structure responsible for different reflected colour wavelengths. Furthermore, colour polymorphism could be advantageous across a gradient from cooler to warmer climate as a result of the different thermal properties of different colours. 6. Given the correlation with temperature, it is predicted that the prevailing reflected colour balance in southern populations would shift in response to global climatic change.     

The daily onset of flight in the crepuscular dung beetle Onitis alexis

The role of light intensity and temperature in determining the onset of flight in the crepuscular dung beetle Onitis alexis Klug (Coleoptera: Scarabaeidae) was examined. Flight under natural light was highly synchronized, with two-thirds of the beetles that flew flying over a period of 10–12 min. In six flights on different days, the mean time of onset varied by up to 13 min, but mean onset occurred at fixed light intensity. Absolute light intensity therefore appears to be a vital cue in determining flight onset. Mean onset remained at this intensity when dusk was advanced artificially by up to about 8 min. However, when dusk was brought further forward, mean flight occurred at lower intensities and onset of flight took place over a longer period. This is interpreted as an overlapping of the period of light intensities suitable for flight with the circadian rhythm that brings the beetles to the surface. No beetles flew when kept under constant bright light or in the dark during the dusk period. Under conditions of constant dusk, the beetles appeared uncoordinated and, although the median time of onset of flight was only 3 min later than in the control flight, onset of flight was spread out over a much longer period. At soil temperatures of 20–22 °C, over 90% of the beetles flew. This percentage decreased with decreasing temperature, and less than 10% flew at temperatures of 16–17 °C.

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L'incitation quotidienne au vol chez e bousier crépusculaire Onitis alexis Klug (Col. Scarabaeidae)

Résumé L'examen a porté sur le rôle de l'intensité lumineuse et de la température sur l'incitation au vol chez Onitis alexis. Le vol en lumière naturelle est fortement synchronisé, avec les deux tiers des adultes qui s'envolent en 10–12 min. Sur six vols pour différents jours, le moment moyen d'envol variait de 13 min, mais l'envol moyen avait lieu à une intensité lumineuse déterminée. La valeur absolue de l'intensité lumineuse paraît ainsi un signal crucial dans la détermination et l'incitation au vol. Le moment d'envol se maintient à cette intensité quand le crépuscule est artificiellement avancé jusqu'à 8 min environ. Cependant, quand le crépuscule est rendu encore plus précoce, l'envol moyen se produit à des intensités plus basses et est étalé. Ceci peut être interprété comme un chevauchement de la période des intensités lumineuses favorables à l'envol avec le rythme circadien qui conduit les bousiers à la surface. Aucun adulte ne vole quand il y a maintien de lumière constante ou d'obscurité à l'heure du crépuscule. En présence de crépuscule constant, les réponses sont hétérogènes, et, bien que le moment médian d'envol ne soit retardé que de 3 min, par rapport aux témoins, l'incitation à l'envol est étalée sur une période plus longue. Avec des températures au sol de 20–22 °C, plus de 90% des adultes s'envolent. Le pourcentage diminue avec la température, et moins de 10% s'envolent à 16–17 °C.