Beetle Species-Area Relationships and Extinction Rates in Protected Areas.
ABSTRACT: The species-area relationship (SAR, i.e., the increase in species richness with area) is one of the most general ecological patterns. SARs can be used to calculate expected extinction rates following area (habitat) loss. Here, using data from Italian reserves, extinction rates were calculated for beetle groups with different feeding habits: Carabidae (terrestrial predators), Hydradephaga (aquatic predators), coprophagous Scarabaeoidea (dung feeders), phytophagous Scarabaeoidea (herbivores), and Tenebrionidae (detritivores). The importance of other factors besides area (namely latitude and elevation) was investigated. Reserve area was recovered as an important predictor of species richness in all cases. For Carabidae, Hydradephaga, and Tenebrionidae, elevation exerted a negative influence, whereas latitude had a negative influence on coprophagous Scarabaeoidea and Tenebrionidae, as a consequence of current and historical biogeographical factors. Extinction rates were higher for dung beetles, due to their dependence on large grazing areas, and Tenebrionidae, due to their low dispersal capabilities. The lower extinction rates predicted for Carabidae, phytophagous Scarabaeoidea, and Hydradephaga can be explained by their higher dispersal power. If other variables besides area are considered, extinction rates became more similar among groups. Extinction rates by area loss are always relatively low. Thus, in reserves with few species, many local extinctions might be unnoticed.
Project description:Extant terrestrial biodiversity arguably is driven by the evolutionary success of angiosperm plants, but the evolutionary mechanisms and timescales of angiosperm-dependent radiations remain poorly understood. The Scarabaeoidea is a diverse lineage of predominantly plant- and dung-feeding beetles. Here, we present a phylogenetic analysis of Scarabaeoidea based on four DNA markers for a taxonomically comprehensive set of specimens and link it to recently described fossil evidence. The phylogeny strongly supports multiple origins of coprophagy, phytophagy and anthophagy. The ingroup-based fossil calibration of the tree widely confirmed a Jurassic origin of the Scarabaeoidea crown group. The crown groups of phytophagous lineages began to radiate first (Pleurostict scarabs: 108 Ma; Glaphyridae between 101 Ma), followed by the later diversification of coprophagous lineages (crown-group age Scarabaeinae: 76 Ma; Aphodiinae: 50 Ma). Pollen feeding arose even later, at maximally 62 Ma in the oldest anthophagous lineage. The clear time lag between the origins of herbivores and coprophages suggests an evolutionary path driven by the angiosperms that first favoured the herbivore fauna (mammals and insects) followed by the secondary radiation of the dung feeders. This finding makes it less likely that extant dung beetle lineages initially fed on dinosaur excrements, as often hypothesized.
Project description:At the basis of a trophic web, coprophagous animals like dung beetles (Scarabaeoidea) utilize resources that may have advantages (easy gain and handling) as well as drawbacks (formerly processed food). Several studies have characterized the nutrients, e.g. C/N ratios and organic matter content, for specific types of dung. However, a comparative approach across dung types and feeding guilds of dung producers, and relationships between dung nutrients and preferences by coprophages, have been missing. Hence, we analyzed water content, C/N ratio, amino acid, neutral lipid fatty acid, free fatty acid and sterol composition and concentrations in dung from 23 vertebrates, including carnivore, omnivore and herbivore species. Our analyses revealed significant differences among the three vertebrate feeding guilds for most nutritional parameters. Although formerly processed, dung grants sufficient amounts of essential nutrients for insects. We tested whether nutrients can explain the dung beetles' preferences in a field experiment, using 12 representative dung types in baits that were installed in 27 forests and 27 grasslands. Although consistent preferences for specific dung types were pronounced, the nutritional composition did not predict the variation in attractiveness of these dung diets, suggesting a primary role of dung volatiles irrespective of food quality.
Project description:In this study, we newly sequenced five mitogenomes of representatives of phytophagous scarab beetles (Coleoptera: Scarabaeidae) by using next-generation sequencing technology. Two species have complete (or nearly complete) mitogenome sequences, namely Popillia mutans Newman (Coleoptera: Scarabaeidae) and Holotrichia oblita Faldermann (Coleoptera: Scarabaeidae). The remaining three species have the partial mitogenomes, and the missing genes are mainly located adjacent to the control region. The complete (or nearly complete) mitogenomes have the same genome structure as most of the existing Scarabaeidae mitogenomes. We conducted phylogenetic analyses together with 24 published mitogenomes of Scarabaeoidea. The results supported a basal split of coprophagous and phytophagous Scarabaeidae. The subfamily Sericinae was recovered as sister to all other phytophagous scarab beetles. All analyses supported a non-monophyletic Melolonthinae, which included two different non-sister clades. The Cetoniinae was recovered as sister to a clade including Rutelinae and Dynastinae. Although the Rutelinae was rendered paraphyletic by Dynastinae in the Bayesian trees inferred under the site-heterogeneous CAT-GTR or CAT-MTART model, discordant patterns were given in some of ML trees estimated using the homogeneous GTR model.
Project description:Dung beetles (Coleoptera: Scarabaeoidea) support numerous ecosystem functions in livestock-grazed pastures. Exposure to veterinary anthelmintic residues in livestock dung can have lethal and sublethal effects on dung beetles, and can reduce rates of dung removal, but the immediate and longer-term consequences for other dung beetle mediated functions have rarely been studied. We investigated the consequences of anthelmintic exposure on survival of the dung beetle Aphodius fossor and its delivery of four ecosystems functions that underpin pasture production: dung removal, soil fauna feeding activity, primary productivity, and reduction of soil compaction. We tested whether anthelmintic exposure had immediate or delayed effects on these functions individually and simultaneously (i.e., ecosystem multifunctionality). We found no evidence that ivermectin residues had a lethal effect on adult beetles. For dung removal, we found a significant interaction between the timing of exposure and functioning: while dung removal was impaired by concurrent exposure to high levels of ivermectin, functioning was unaffected when beetles that had been exposed previously to the same concentration of anthelmintic later interacted with untreated dung. Other ecosystem functions were not affected significantly by anthelmintic exposure, and there was no evidence to suggest any persistent impact of anthelmintic exposure on ecosystem multifunctionality. While anthelmintic residues remain a significant threat to dung beetle populations, for adult beetles, we found no evidence that residues have detrimental consequences for ecosystem functioning beyond the immediate point of exposure.
Project description:The dung beetle subfamily Scarabaeinae is a cosmopolitan group of insects that feed primarily on dung. We describe the first case of an obligate predatory dung beetle and contrast its behaviour and morphology with those of its coprophagous sympatric congeners. Deltochilum valgum Burmeister killed and consumed millipedes in lowland rainforest in Peru. Ancestral ball-rolling behaviour shared by other canthonine species is abandoned, and the head, hind tibiae and pygidium of D. valgum are modified for novel functions during millipede predation. Millipedes were killed by disarticulation, often through decapitation, using the clypeus as a lever. Beetles killed millipedes much larger than themselves. In pitfall traps, D. valgum was attracted exclusively to millipedes, and preferred injured over uninjured millipedes. Morphological similarities placing D. valgum in the same subgenus with non-predatory dung-feeding species suggest a major and potentially rapid behavioural shift from coprophagy to predation. Ecological transitions enabling the exploitation of dramatically atypical niches, which may be more likely to occur when competition is intense, may help explain the evolution of novel ecological guilds and the diversification of exceptionally species-rich groups such as insects.
Project description:In recent decades, pastoral abandonment has produced profound ecological changes in the Alps. In particular, the reduction in grazing has led to extensive shrub encroachment of semi-natural grasslands, which may represent a threat to open habitat biodiversity. To reverse shrub encroachment, we assessed short-term effects of two different pastoral practices on vegetation and dung beetles (Coleoptera, Scarabaeoidea). Strategic placement of mineral mix supplements (MMS) and arrangement of temporary night camp areas (TNCA) for cattle were carried out during summer 2011 in the Val Troncea Natural Park, north-western Italian Alps. In 2012, one year after treatment, a reduction in shrub cover and an increase in bare ground cover around MMS sites was detected. A more intense effect was detected within TNCA through increases in forage pastoral value, and in the cover and height of the herbaceous layer. Immediately after treatment, changes in dung beetle diversity (total abundance, species richness, Shannon diversity, taxonomic and functional diversity) showed a limited disturbance effect caused by high cattle density. In contrast, dung beetle diversity significantly increased one year later both at MMS and TNCA sites, with a stronger effect within TNCA. Multivariate Regression Trees and associated Indicator Value analyses showed that some ecologically relevant dung beetle species preferred areas deprived of shrub vegetation. Our main conclusions are: i) TNCA are more effective than MMS in terms of changes to vegetation and dung beetles, ii) dung beetles respond more quickly than vegetation to pastoral practices, and iii) the main driver of the rapid response by dung beetles is the removal of shrubs. The resulting increase in dung beetle abundance and diversity, which are largely responsible for grassland ecosystem functioning, may have a positive effect on meso-eutrophic grassland restoration. Shrub encroachment in the Alps may therefore be reversed, and restoration of grassland enhanced, by using appropriate pastoral practices.
Project description:The evolutionary success of beetles and numerous other terrestrial insects is generally attributed to co-radiation with flowering plants but most studies have focused on herbivorous or pollinating insects. Non-herbivores represent a significant proportion of beetle diversity yet potential factors that influence their diversification have been largely unexamined. In the present study, we examine the factors driving diversification within the Scarabaeidae, a speciose beetle family with a range of both herbivorous and non-herbivorous ecologies. In particular, it has been long debated whether the key event in the evolution of dung beetles (Scarabaeidae: Scarabaeinae) was an adaptation to feeding on dinosaur or mammalian dung. Here we present molecular evidence to show that the origin of dung beetles occurred in the middle of the Cretaceous, likely in association with dinosaur dung, but more surprisingly the timing is consistent with the rise of the angiosperms. We hypothesize that the switch in dinosaur diet to incorporate more nutritious and less fibrous angiosperm foliage provided a palatable dung source that ultimately created a new niche for diversification. Given the well-accepted mass extinction of non-avian dinosaurs at the Cretaceous-Paleogene boundary, we examine a potential co-extinction of dung beetles due to the loss of an important evolutionary resource, i.e., dinosaur dung. The biogeography of dung beetles is also examined to explore the previously proposed "out of Africa" hypothesis. Given the inferred age of Scarabaeinae as originating in the Lower Cretaceous, the major radiation of dung feeders prior to the Cenomanian, and the early divergence of both African and Gondwanan lineages, we hypothesise that that faunal exchange between Africa and Gondwanaland occurred during the earliest evolution of the Scarabaeinae. Therefore we propose that both Gondwanan vicariance and dispersal of African lineages is responsible for present day distribution of scarabaeine dung beetles and provide examples.
Project description:The overabundance of large herbivores is now recognized as a serious ecological problem. However, the resulting ecological consequences remain poorly understood. The ecological effects of an increase in sika deer, Cervus nippon Temminck (Cervidae), on three insect groups of beetles was investigated: ground beetles (Carabidae), carrion beetles (Silphidae), and dung beetles (Scarabaeidae and Geotrupidae) on Nakanoshima Island, Hokkaido, northern Japan. We collected beetles on Nakanoshima Island (experimental site) and lakeshore areas (control site) and compared the species richness, abundance, diversity index, and community composition of beetles between the sites. Results showed that although both species diversity and abundance of carabid beetles were significantly higher at the lakeshore site, those of dung and carrion beetles were higher at the island site. It was additionally observed that abundance of larger carabid beetles was higher at the lakeshore site, whereas that of small-sized carabid beetles did not differ between the lakeshore and island sites. For dung beetles, abundance of smaller species was higher at the island site, whereas that of large species did not differ between the lakeshore and island sites. Abundance of two body sizes (small and large) of carrion beetles were both higher at the island site. Overall, the findings of this study demonstrated that an increase in deer population altered the insect assemblages at an island scale, suggesting further changes in ecosystem functions and services in this region.
Project description:BACKGROUND: Coleoptera is the most diverse order of insects (>300,000 described species), but its richness diminishes at increasing latitudes (e.g., ca. 7400 species recorded in Canada), particularly of phytophagous and detritivorous species. However, incomplete sampling of northern habitats and a lack of taxonomic study of some families limits our understanding of biodiversity patterns in the Coleoptera. We conducted an intensive biodiversity survey from 2006-2010 at Churchill, Manitoba, Canada in order to quantify beetle species diversity in this model region, and to prepare a barcode library of beetles for sub-arctic biodiversity and ecological research. We employed DNA barcoding to provide estimates of provisional species diversity, including for families currently lacking taxonomic expertise, and to examine the guild structure, habitat distribution, and biogeography of beetles in the Churchill region. RESULTS: We obtained DNA barcodes from 3203 specimens representing 302 species or provisional species (the latter quantitatively defined on the basis of Molecular Operational Taxonomic Units, MOTUs) in 31 families of Coleoptera. Of the 184 taxa identified to the level of a Linnaean species name, 170 (92.4%) corresponded to a single MOTU, four (2.2%) represented closely related sibling species pairs within a single MOTU, and ten (5.4%) were divided into two or more MOTUs suggestive of cryptic species. The most diverse families were the Dytiscidae (63 spp.), Staphylinidae (54 spp.), and Carabidae (52 spp.), although the accumulation curve for Staphylinidae suggests that considerable additional diversity remains to be sampled in this family. Most of the species present are predatory, with phytophagous, mycophagous, and saprophagous guilds being represented by fewer species. Most named species of Carabidae and Dytiscidae showed a significant bias toward open habitats (wet or dry). Forest habitats, particularly dry boreal forest, although limited in extent in the region, were undersampled. CONCLUSIONS: We present an updated species list for this region as well as a species-level DNA barcode reference library. This resource will facilitate future work, such as biomonitoring and the study of the ecology and distribution of larvae.
Project description:Fire has become an increasingly important disturbance event in south-western Amazonia. We conducted the first assessment of the ecological impacts of these wildfires in 2008, sampling forest structure and biodiversity along twelve 500 m transects in the Chico Mendes Extractive Reserve, Acre, Brazil. Six transects were placed in unburned forests and six were in forests that burned during a series of forest fires that occurred from August to October 2005. Normalized Burn Ratio (NBR) calculations, based on Landsat reflectance data, indicate that all transects were similar prior to the fires. We sampled understorey and canopy vegetation, birds using both mist nets and point counts, coprophagous dung beetles and the leaf-litter ant fauna. Fire had limited influence upon either faunal or floral species richness or community structure responses, and stems <10 cm DBH were the only group to show highly significant (p = 0.001) community turnover in burned forests. Mean aboveground live biomass was statistically indistinguishable in the unburned and burned plots, although there was a significant increase in the total abundance of dead stems in burned plots. Comparisons with previous studies suggest that wildfires had much less effect upon forest structure and biodiversity in these south-western Amazonian forests than in central and eastern Amazonia, where most fire research has been undertaken to date. We discuss potential reasons for the apparent greater resilience of our study plots to wildfire, examining the role of fire intensity, bamboo dominance, background rates of disturbance, landscape and soil conditions.