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.     

Dung beetles as secondary seed dispersers in a temperate grassland

The two-phase dispersal event in which dung beetles move seeds after endozoochory is often assumed to be advantageous for plant regeneration. Because seeds are expected to end up in favourable and safe germination sites, it is considered as an example of directed dispersal. However, literature so far is restricted to tropical rain forest ecosystems, while data for temperate regions are lacking. In this study, the effect of dung beetles on seedling establishment of endozoochorically dispersed seeds is evaluated for a temperate grassland ecosystem. We performed a field experiment in which cages excluded dung beetles from horse and cattle dung samples with mixed-in grass seeds. Seed germination from these samples was significantly higher than that from samples which were accessible to dung beetles. This indicates that the effect of dung beetles on short-term seedling establishment was negative, which contrasts with the patterns found for large-seeded species used in tropical studies. This is most likely attributed to the lack of roller species and the larger depth at which tunneling Geotrupes species bury seeds.     

Linking dung beetle-mediated functions to interactions in the Atlantic Forest: Sampling design matters

Interactions between dung beetles and vertebrate dung are intimately linked to a suite of ecosystem functions in tropical forests. We show that the trapping method and the type of dung used affect the suite of beetles captured, with the potential to influence the outcome of experiments linking functions to interactions.     

Diversity of dung and carrion beetles in a disturbed Mexican tropical montane cloud forest and on shade coffee plantations

This paper analyzes the diversity of dung and carrion beetles (Scarabaeinae and Silphidae) in four human-induced habitats of a disturbed tropical montane cloud forest: polyspecific shade coffee plantations, monospecific shade coffee plantations, tropical montane cloud forest fragments, and clear cuts. The four habitats had similar richness, species composition, and assemblage structure of dung and carrion beetles. Differences were found in abundance and biomass levels for the four dominant species in the landscape. Dung beetles were more abundant than carrion beetles, but the biomass was higher for the latter. Carrion beetles were seasonal, while dung beetles were clearly not. When forest fragments and shade coffee plantations were compared to other similar habitats in the region, the same general pattern was observed. However, forests with high disturbance and monospecific shade coffee plantations had lower species richness than forests with low and medium disturbance and polyspecific shade coffee plantations. Thus shade coffee plantations maintain connectivity between patches of cloud forest in a landscape that is strongly affected by human activities. Protecting landscape diversity appears to ensure high species richness.     

The activity of dung beetles increases foliar nutrient concentration in tropical seedlings

Dung beetles are extensively used as a focal taxon in tropical forests. Yet, information for most of their ecological functions comes from other systems. We present results from a field experiment in a tropical rain forest showing that dung beetle activity increases foliar phosphorus concentration in seedlings of the tree Brosimum lactescens. Our results open new lines of research to assess the multiple effects that dung beetles may have on rain forest plants.     

Low local but high beta diversity of tropical forest dung beetles in Madagascar

Aims We 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. Results Species 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 (R² 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 conclusion The 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.     

Effects of Season and Vegetation Type on Community Organization of Dung Beetles in a Tropical Dry Forest1

Dung beetles are important components of most terrestrial ecosystems. In tropical rain forests, dung beetle communities can be very rich in number of species and individuals, and they are known to be useful bioindicators of habitat disturbance. In contrast, very little is known about the organization of dung beetle communities in tropical dry forests. The aim of this study was to describe in detail the dung beetle community of a Mexican tropical dry forest and to assess the relative importance of rainfall seasonality and forest structure in affecting the temporal and spatial dynamics of this community. Dung beetles were captured with pitfall traps at the beginning of the rainy season, the middle of the rainy season, and the middle of the dry season, in two distinct forest types: deciduous forest (DF) and semideciduous forest (SDF) at the Estación de Biología Chamela. Both rainfall seasonality and forest structure affected the community organization of dung beetles. During both rainy periods, 14 species were captured, but only three during the dry season. Dung beetles captured during the dry season were only found in the SDF. When comparing the beginning and the middle of the rainy season, differences in abundance and guild structure were also observed between both periods and between forest types, but these differences were much less pronounced.     

Dung beetle activity affects rain forest seed bank dynamics and seedling establishment

Dung beetles relocate vertebrate feces under the soil surface, and this behavior has many ecological consequences. In tropical forests, for example, seeds defecated by mammals that are subsequently buried by dung beetles are less likely to suffer predation. While the effects of dung beetles on the fate of defecated seeds have been relatively well studied, their effect on seeds already buried in the soil has not. To contribute to fill this gap, we designed a study with three objectives: (a) Describe the vertical re‐distribution of soil seeds that occurs due to dung beetle activity; (b) Determine if beetle activity favors establishment of seedlings from the soil seed bank; and (c) Determine if the effect of dung beetles is stronger in sites of recurrent mammal defecation. We carried out three complementary field experiments, one with artificial seeds (plastic beads) of three sizes buried at known depths, one with two species of seeds buried at those same depths, and one with the natural soil seed bank in sites of single vs. recurrent defecation. Buried beads were moved by dung beetles along the vertical axis, both upwards (9.5%) and downwards (11.5%); smaller beads were more frequently moved downwards while the contrary occurred for larger beads. Dung beetle activity caused an increase in seedling establishment, both from experimentally buried seeds and from the natural seed bank. Defecation recurrence had no effect on seedling establishment. We conclude that dung beetle activity affects seed bank dynamics with important consequences for seedling establishment in tropical forests.Abstract in Spanish is available with online material.     

The effects of rainforest canopy loss on arboreal dung beetles in Borneo: implications for the measurement of biodiversity in derived tropical ecosystems

We examine the effect of selective timber extraction, and corresponding forest canopy loss, on arboreal dung beetles in the tropical rainforests of Sabah, Malaysian Borneo. Changes in vertical distribution of beetles are measured through differences in the abundance of beetles in ground-based pitfall traps in primary, logged and plantation forest. Previous research has demonstrated that arboreal dung beetles are not generally collected in pitfall traps in primary forest, but are present in large numbers above the ground in canopy vegetation: the presence of arboreal beetles in pitfalls in areas of reduced or modified canopy cover may therefore reflect a response to the absence or modification of their usual habitat, and the proliferation of these beetles nearer to the ground. In this paper, statistically significant differences are found in the abundance of beetles in ground pitfall traps from logged forest compared to primary forest. Results show that virtually no arboreal dung beetles are recorded in primary forest traps, with an increased abundance of arboreal dung beetles in traps from logged and plantation forest, with 1.72% of the total number of arboreal beetles recorded from primary forest, 22.32% from logged forest, and 75.96% from plantation forest. The presence of arboreal dung beetles in plantations demonstrates that arboreal dung beetles can survive outside their normal habitat, and we relate these observations to adaptations to upper rainforest canopy conditions, and proliferation of these microclimatic conditions in man-made habitats. Results are also discussed in terms of their relevance to the measurement of species richness and diversity in logged and other derived ecosystems, where mixing of the ground-based and arboreal faunas occurs.     

Effects of dung and seed size on secondary dispersal,seed predation,and seedling establishment of rain forest trees

Seeds dispersed by tropical, arboreal mammals are usually deposited singly and without dung or in clumps of fecal material. After dispersal through defecation by mammals, most seeds are secondarily dispersed by dung beetles or consumed by rodents. These post-dispersal, plant-animal interactions are likely to interact themselves, as seeds buried by dung beetles are less likely to be found by rodents than unburied seeds. In a series of three experiments with seeds of 15 species in central Amazonia (Brazil), we determined (1) how presence and amount of dung associated with seeds influences long-term seed fate and seedling establishment, (2) how deeply dung beetles bury seeds and how burial depth affects seedling establishment, and (3) how seed size affects the interaction between seeds, dung beetles, and rodents. Our overall goal was to understand how post-dispersal plant-animal interactions determine the link between primary seed dispersal and seedling establishment. On average, 43% of seeds surrounded by dung were buried by dung beetles, compared to 0% of seeds not surrounded by dung (n=2,156). Seeds in dung, however, tended to be more prone than bare seeds to predation by rodents. Of seeds in dung, probability of burial was negatively related to seed size and positively related to amount of dung. Burial of seeds decreased the probability of seed predation by rodents three-fold, and increased the probability of seedling establishment two-fold. Mean burial depth was 4 cm (0.5–20 cm) and was not related to seed size, contrary to previous studies. Probability of seedling establishment was negatively correlated with burial depth and not related to seed size at 5 or 10 cm depths. These results illustrate a complex web of interactions among dung beetles, rodents, and dispersed seeds. These interactions affect the probability of seedling establishment and are themselves strongly tied to how seeds are deposited by primary dispersers. More generally, our results emphasize the importance of looking beyond a single type of plant-animal interaction (e.g., seed dispersal or seed predation) to incorporate potential effects of interacting interactions.     

Fire? They don't give a dung! The resilience of dung beetles to fire in a tropical savanna

1. Disturbance is a strong driver of community assembly and fire has long been recognised as one of the main disturbances of terrestrial ecosystems. This study tested the resilience of dung beetles to fire events in campos rupestres, which is a tropical savanna ecosystem that evolved under a frequent fire regime, by assessing the resistance and recovery of their communities. 2. Dung beetles were sampled before and after a fire event and the effect of fire on dung beetle richness, abundance, mean community biomass and composition was tested. The effects of time since last fire and fire frequency on the community were also tested. 3. No effect of fire occurrence, time since last fire and fire frequency on any community variable was found. 4. Some non‐mutually exclusive mechanisms promoting the resistance and recovery of dung beetles in campos rupestres could be acting in synergy. One potential mechanism is the mismatched seasonality between fire events and dung beetle occurrence, as fires occur during the dry season and dung beetles are present above ground during the rainy season. Furthermore, dung beetles are insects that remain buried during most of their lifetime, which could protect individuals from being burned. Another potential mechanism is the replacement of species in burned areas by the movement of individuals from unburned areas, attracted by resources and/or by metacommunity dynamics. 5. It is concluded that in this ‘fire‐dependent’ ecosystem, dung beetle communities are resilient to fire and seem not to be structured by this disturbance.     

Do riparian reserves support dung beetle biodiversity and ecosystem services in oil palm-dominated tropical landscapes?

Agricultural expansion and intensification are major threats to global biodiversity, ecological functions, and ecosystem services. The rapid expansion of oil palm in forested tropical landscapes is of particular concern given their high biodiversity. Identifying management approaches that maintain native species and associated ecological processes within oil palm plantations is therefore a priority. Riparian reserves are strips of forest retained alongside rivers in cultivated areas, primarily for their positive hydrological impact. However, they can also support a range of forest‐dependent species or ecosystem services. We surveyed communities of dung beetles and measured dung removal activity in an oil palm‐dominated landscape in Sabah, Malaysian Borneo. The species richness, diversity, and functional group richness of dung beetles in riparian reserves were significantly higher than in oil palm, but lower than in adjacent logged forests. The community composition of the riparian reserves was more similar to logged forest than oil palm. Despite the pronounced differences in biodiversity, we did not find significant differences in dung removal rates among land uses. We also found no evidence that riparian reserves enhance dung removal rates within surrounding oil palm. These results contrast previous studies showing positive relationships between dung beetle species richness and dung removal in tropical forests. We found weak but significant positive relationships between riparian reserve width and dung beetle diversity, and between reserve vegetation complexity and dung beetle abundance, suggesting that these features may increase the conservation value of riparian reserves. Synthesis and applications: The similarity between riparian reserves and logged forest demonstrates that retaining riparian reserves increases biodiversity within oil palm landscapes. However, the lack of correlation between dung beetle community characteristics and dung removal highlights the need for further research into spatial variation in biodiversity–ecosystem function relationships and how the results of such studies are affected by methodological choices.     

Dung beetle assemblages in forests and pastures of El Salvador: a functional comparison

This study examines the functional attributes of Neotropical dung beetles, and, based on the analysis, evaluates the role of dung beetles in pastures and tropical dry forest in El Salvador. Dung beetle diversity was lower in pastures than in forests. However, the total biomass of beetles at similarly-sized dung baits was frequently higher in pastures. Diversity loss followed structured patterns: (1) carrion and fruit feeding beetles were well represented (species number) in the forests but were largely absent from pastures; (2) large ball rollers and small fast tunnellers were present in forests but were absent from pastures; (3) large fast tunnellers and small slow tunnellers were poorly represented in pastures compared to forests but the few species that survived in pastures attained extremely high populations; and (4) each functional group in the species-poor pasture assemblages was divided almost equally into day and night active species. Substrate and habitat generalists that were present in both the forests and pastures were attracted to fruit and carrion in forested habitat but not in open pastures. In open habitats, flies and other insects may be the principal decomposers of decaying fruit and carrion.     

Landscape attributes shape dung beetle diversity at multiple spatial scales in agricultural drylands

Land-use change is one of the main drivers of biodiversity loss worldwide, but its negative effects can vary depending on the spatial scale analyzed. Considering the continuous expansion of agricultural demand for land, it is urgent to identify the drivers that shape biological communities in order to balance agricultural production and biodiversity conservation in human-modified landscapes. We used a patch-landscape design and a multimodel inference approach to assess the effects of landscape composition and configuration at two spatial scales (patch and landscape) on the structure of dung beetle assemblages. We performed our study in the Caatinga, the largest dry forest in South America. We sampled 3,526 dung beetles belonging to 19 species and 11 genera. At patch scale, our findings highlight the positive relationship of forest cover and landscape heterogeneity with dung beetle diversity, which are the major drivers of beetle assemblages. Edge density, in turn, is a major driver at the landscape scale and has a negative effect on beetle diversity. Our results support the hypothesis that landscapes combining natural vegetation remnants and heterogeneous agricultural landscapes are the most effective at conserving the biodiversity of dung beetles in the Caatinga landscapes. Directing efforts to better understand the dynamics of dung beetles in agricultural lands can be helpful for policymakers and scientists to design agri-environment schemes and apply conservation strategies in tropical dry forests.     

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.     

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.     

Effects of dung beetles on seedling emergence from herbaceous seeds in the dung of sika deer (<Emphasis Type="Italic">Cervus nippon</Emphasis>) in a temperate Japanese grassland ecosystem

The effects of dung form and condition and of dung beetles on the emergence of seedlings from herbaceous seeds in sika deer dung were examined in a temperate grassland ecosystem dominated by Zoysia japonica and Hydrocotyle maritima. I conducted field experiments to compare seedling emergence between dung exposed to dung beetles and intact dung using both dung pellets and pats during a typical rainy month (June) and the hottest, drier month (August), when large numbers of seeds of the dominant species were present in the dung. The exposed dung was immediately attacked and broken up by dung beetles, whereas dung protected from the beetles remained intact. In June, at least 12 herbaceous species, including Z. japonica, H. maritima, Mazus pumilus, and Plantago asiatica, emerged from the dung, versus at least six species in August. Decomposition rates of the pellets in June and decomposition scores of the pats in June and August were positively correlated with the number of emerging seedlings, suggesting that the acceleration of decomposition by dung beetles can positively affect seed germination. In this system of interactions among sika deer, herbaceous plants, and dung beetles, sika deer dung prevented seeds from germinating, and beetles had an indirect positive effect on seedling emergence by accelerating decomposition of the dung, although the extent of the effect may depend on the dung type, plant species, and environmental factors.     

Getting the most out of waste: how dung beetles boost the nitrogen content in their food

The life cycle of almost all dung beetles revolve around mammalian dung, the feed on dung, look for mating partners on dung and lay eggs in the dung. We know they feed on dung, but we still do not understand how exactly they filter‐feed on the dung and which particles size range they are ingesting. The aim on this study was to investigate the filter feeding by particle selection by adult dung beetles using Scarabaeus goryi and how that improves the nutrient quality of the ingested particles. We compared the particle sizes and nutrient content of the dung with the ingested material in the foregut, hindgut and the faeces of the dung beetle. Adult dung beetles do select smaller dung particles when feeding, we found the maximum particle size for the ingested particle to be around 1400 μm. The average particle size increased through the gut length. Dung beetles also selected particles with higher nitrogen content when feeding, the nitrogen content increased from about 1.5% in the dung to just over 5% in the foregut which then decreased to the level of the unprocessed dung in the dung beetle faeces. Carbon content did not increase from the unprocessed dung to the foregut but decreased through the gut length. Feeding by particle size selection by dung beetles helps in selecting particles with higher nitrogen content to compensate for the low levels found in dung.     

Impacts of Small‐scale Clearings due to Selective logging on Dung Beetle Communities

Conservation of biodiversity in production forests is crucial for mitigating biodiversity loss in the tropics. The major ecological impacts of selective logging are often the result of small clearings for skid trails, logging roads, log yards, and logging camps; however, their impacts on forest biodiversity have rarely been examined. The purpose of this study was to assess the impacts of these clearings on a forest‐dependent faunal group, dung beetles, and to identify the environmental factors responsible. Abundance and species richness of dung beetles decreased drastically in clearings, but directly increased in forests with the distance from roads/trails; abundance and species richness at 10 m from roads/trails were almost comparable with those detected in further interior forests. Similarly, species composition was significantly different between forests and clearings (except skid trails) but recovered within a short distance from roads/trails. Canopy openness was the most important environmental factor affecting the abundance, and species richness and composition of dung beetles; most dung beetle species were concentrated under closed forest canopy with less than 10 percent of canopy openness, whereas canopy openness ranged from 16 to 53 percent in clearings. Our study demonstrates that even small‐scale, unpaved clearings affect dung beetle communities through increased canopy openness. Although the effective distance was not very large, a considerable portion of logged areas can be affected when road networks are dense therefore minimizing the density of road networks and enhancing canopy recovery after logging are important for retaining biodiversity in tropical production forests.     

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.