Antibiotics are routinely used to improve livestock health and growth. However, this practice may have unintended environmental impacts mediated by interactions among the wide range of micro-and macroorganisms found in agroecosystems. For example, antibiotics may alter microbial emissions of greenhouse gases by affecting livestock gut microbiota. Furthermore, antibiotics may affect the microbiota of non-target animals that rely on dung, such as dung beetles, and the ecosystem services they provide. To examine these interactions, we treated cattle with a commonly used broad-spectrum antibiotic and assessed downstream effects on microbiota in dung and dung beetles, greenhouse gas fluxes from dung, and beetle size, survival and reproduction. We found that antibiotic treatment restructured microbiota in dung beetles, which harboured a microbial community distinct from those in the dung they were consuming. The antibiotic effect on beetle microbiota was not associated with smaller size or lower numbers. Unexpectedly, antibiotic treatment raised methane fluxes from dung, possibly by altering the interactions between methanogenic archaea and bacteria in rumen and dung environments. Our findings that antibiotics restructure dung beetle microbiota and modify greenhouse gas emissions from dung indicate that antibiotic treatment may have unintended, cascading ecological effects that extend beyond the target animal.

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Abstract:The wet tropical forests in Madagascar have endemic dung beetles that have radiated for tens of millions of years using a limited range of resources produced by the species-poor mammalian fauna. Beetles were trapped in two wet-forest localities over 4 years (6407 trap nights, 18,869 individuals). More limited data for six other local communities were used to check the generality of the results. Local communities are relatively species poor (around 30 species) in comparison with wet-forest-inhabiting dung beetle communities elsewhere in the tropics (typically 50 or more species). The species belong to only two tribes, Canthonini and Helictopleurina (Oniticellini), which have evolved, exceptionally for dung beetle tribes, completely nocturnal versus diurnal diel activities, respectively. Patterns in the elevational occurrence, body size and resource use suggest that interspecific competition restricts the numbers of locally coexisting species exploiting the limited range of resources that are available. On the other hand, regional turnover in the species composition is exceptionally high due to a large number of species with small geographical ranges, yielding a very large total fauna of dung beetles in Madagascar (>250 species). Apart from exceptionally low local (alpha) diversity and high beta diversity, the Malagasy dung beetle communities are ecologically distinctive from comparable communities in other tropical regions in having high numerical dominance of the most abundant species, small average body size and low degree of resource specialization.

AimsWe have compared local (alpha) and regional (beta) species diversities of dung beetles in wet forests in the main tropical regions including Madagascar. Madagascar is exceptional in lacking native large herbivorous mammals which produce the key resource for dung beetles elsewhere.Location Central and South America, mainland Africa, Madagascar and Southeast Asia.Methods Trapping data on dung beetles and data on mammalian faunas were obtained from published and unpublished studies. We used our original data for Madagascar. ResultsSpecies richness of dung beetles and that of large-bodied (> 15 mm length) species in particular were highly significantly explained by the regional number of large-bodied (> 10 kg) mammals (R 2 from 50 to 80%). For a given pairwise spatial distance between two communities, beta diversity was significantly higher in Madagascar than elsewhere, explaining the very high total species richness in Madagascar in spite of low local diversity. Main conclusionThe presence and numbers of large herbivorous mammals greatly influence the species richness of dung beetles in tropical wet forests. The lack of native large herbivores rather than a limited species pool explains the low local diversity in Madagascar. Exceptionally high beta diversity in Madagascar suggests a pattern of old radiation involving extensive allopatric speciation.

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Paleotropical clades with largely disjunct distributions are ideal models for biogeographic reconstructions. The dung beetle genera Grebennikovius Mlambo, Scholtz & Deschodt, Epactoides Olsouffief and Ochicanthon Vaz-de-Mello are distributed in Tanzania, Madagascar and Réunion, and the Oriental region, respectively. We combine morphology and molecular dataset to reconstruct the phylogenetic relationships between these taxa. Our analyses corroborate previous hypotheses of monophyly of the group, which is here described as new tribe Epactoidini trib. nov. Grebennikovius is recovered as sister to Epactoides, while Ochicanthon emerges as sister to them both. The disjunct distribution of our focal clade is unusual within the subfamily Scarabaeinae. Bayesian divergence time estimates and ancestral range reconstructions indicate an African origin of the crown group of the tribe Epactoidini trib. nov. in the early mid Eocene, ca. 46 Ma. The divergence between Epactoides and its sister is dated to 32.3 Ma, while the crown age for the genus Ochicanthon is dated to 27 Ma. We investigate the factors that may have shaped the current distribution of the tribe Epactoidini trib. nov. The formation of the Gomphotherium landbridge, along with favourable environmental conditions would have allowed dry-intolerant organisms, such as Ochicanthon, to disperse out of Africa. Remarkable climatic stability of the Eastern Arc Mountains was critical for the retention of the monotypic genus Grebennikovius. We suggest two subsequent overwater dispersal events: the migration of the most recent common ancestor (MRCA) of Epactoides from Africa to Madagascar (32.3-29.5 Ma); the lately dispersal of the MRCA of the today's extinct Epactoides giganteus Rossini, Vaz-de-Mello & Montreuil to Réunion island from Madagascar (3.4 Ma). We suggest that the high potential of dispersal of Epactoidini trib. nov. dung beetles and the strict association to forest habitat might have triggered two major radiations, one in Madagascar and one in the Oriental Region.