Large-scale spatial ecology of dung beetles

Recent modelling work shows that the composition of local communities can be influenced by the configuration of the surrounding landscape, but many of these models assume that all community members display the same type of extinction‐colonization dynamics. I use Aphodius dung beetles to test the hypothesis that interspecific differences in habitat selection and dispersal capacity may translate into differences in spatial population dynamics, even among closely related species coexisting on the same resource. If this is true, then groups of species with different characteristics would show different responses to landscape configuration. I first divided the area of Finland into a grid, and used collection records to describe regional variation in the Aphodius fauna of open cattle pastures. I then sampled dung beetles on 131 cattle farms, to examine whether the subset of species found on each farm was related to the density of pastures in the surrounding grid square. Finally, I used historical records to analyze changes in dung beetle communities during the last century, when there was great loss of pasture. Overall, I found no relationship between landscape characteristics and the total proportion of the regional species pool that was found on each farm. However, the distribution of species among guilds with different habitat specificity did relate to the configuration of the landscape, and the pattern was most pronounced in a specialists species with limited dispersal. Associations between community structure and landscape configuration were superimposed on two much larger and stronger patterns: a large‐scale latitudinal gradient in regional species richness, and a decelerating gain of species to local communities with an increasing regional species pool. I conclude that ecological variation among community members is a crucial factor in the analysis of local community composition, and that local species richness should always be conditioned on regional richness.     

Habitat fragmentation and the functional efficiency of temperate dung beetles

To understand how current patterns of habitat loss and fragmentation will ultimately affect ecosystem functioning, we need to match experimental manipulations of community structure with real changes occurring in the landscapes of today. In this study, we examine the consequences of habitat fragmentation on a key function: the decomposition of dung by invertebrates. In a microcosm experiment, we use previous observations of dung beetle assemblage structure in fragmented and intact landscapes to create realistic differences in assemblages of small, dung‐dwelling species in the genus Aphodius. We ask whether such differences will affect ecosystem functioning, and how their effects compare to those of removing full functional groups: dung‐dwelling Aphodius, tunnelling Geotrupes stercorarius, and/or earthworms. As measured by changes in dung fresh weight, we observe an overriding impact of removing G. stercorarius, with the amount of dung remaining at any one time doubling if the species is excluded. Compared to this major effect, there seem to be less effects of removing Aphodius, ambiguous effects of excluding earthworms, and no detectable effects of relatively minor changes in Aphodius assemblages as induced by current levels of fragmentation. Overall, our results support the general notion that different species contribute highly unevenly to overall ecosystem functioning. Most importantly though, our findings suggest that the functional consequences of habitat loss will depend on taxon‐specific responses to landscape modification. Only by addressing these responses may we predict the actual consequences of habitat loss.     

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.     

Habitat and weather affect reproduction and size of the dung beetle Aphodius fossor

1. The influence of sun exposure and moisture on reproduction and size of a north temperate dung beetle Aphodius fossor was investigated. Adult beetles were added to cow pats that were either shaded or exposed and treated with different watering regimes. 2. After 26 days, more eggs and larvae were recovered from shaded and wetter pats than from exposed and drier pats. 3. At the end of the experiment, more beetles had developed in shaded and wetter pats, and these beetles were larger than those from exposed and drier pats. 4. There was no between‐treatment difference in the ratio between adults in final sampling and larvae in first sampling. 5. It was concluded that exposure and moisture can influence both reproduction and size of A. fossor. It is suggested that weather may be a factor regulating reproduction in Aphodius not only through heat‐ or drought‐induced mortality, as suggested earlier, but also by limiting the time available for egg laying. Further studies are needed to clarify this suggestion.     

Newly released non‐native dung beetle species provide enhanced ecosystem services in New Zealand pastures

1. Whether the release of non‐native insect species benefits or harms ecosystem services has been the subject of debate. In New Zealand, the release of new non‐native dung beetle species was intended to enhance ecosystem services but concerns were raised over possible negative effects. 2. Field cage trials used three newly released dung beetle species to investigate two concerns: that soil disturbance from dung beetle activity increases soil losses in runoff after rainfall; and that dung burial increases survival of infective parasitic nematodes on pasture. 3. Three treatments – dung + beetles, dung‐only, and controls (without dung or beetles) – were applied on each of three soil types with different permeability: sandy loam, clay loam, and compacted clay. 4. Dung beetle activity resulted in significant reductions of 49% and 81% in mean surface runoff volume, depending on simulated rainfall intensity. Amounts of sediment in the runoff did not change under an extreme rainfall simulation, but in a less extreme rainfall simulation the presence of dung beetles resulted in a 97% reduction in mean sediment amount in runoff. 5. The numbers of infective third‐stage nematode larvae recovered from foliage varied considerably between soil types and through time; however, dung beetle activity reduced overall mean nematode numbers on grass around the dung pats by 71%. 6. This study adds to global evidence that dung beetles can improve agricultural ecosystem services by providing data on services that have rarely been investigated: reduced runoff/soil losses through increased soil porosity, and reductions in parasitic nematodes.     

Territorial ants depress plant growth through cascading non‐trophic effects in an alpine meadow

Understanding non‐trophic interactions is critical to mechanistically linking community structure and ecosystem functioning. Despite the widespread occurrence of territoriality across animal taxa and ecosystems, the cascading ecological consequences of non‐trophic interactions between territorial animals and intruders have been poorly studied. We experimentally investigated the non‐trophic interaction between territorial ants and members of a dung decomposer community (i.e. predatory arthropods, maggots and coprophagous beetles) in an alpine meadow. We further examined how this non‐trophic interaction cascaded to influence ecosystem properties including dung removal rate, soil nutrient status and aboveground plant biomass surrounding dung pats. Results indicated that territorial interference of ants on key decomposers cascaded to affect plant growth. Specifically, ants significantly decreased the abundance of coprophagous beetles at the time of their peak‐abundance and hence decreased dung removal rates and soil nitrogen concentrations, ultimately decreasing aboveground plant biomass. The strength of this non‐trophic cascading effect was comparable to those reported in studies addressing trophic cascades triggered by predator–prey interactions. Our findings suggest that the non‐trophic interactions and associated cascading effects stemming from territorial behavior should be incorporated into ecological network modeling and research addressing biodiversity–ecosystem functioning relationships.     

Heterotrophic succession within dung-inhabiting beetle communities in northern Spain

Successional patterns of beetles inhabiting dung pats were examined during May and July 1993 in a mountain area in northern Spain (Picos de Europa). Beetles belonging to six families were caught during the course of succession (30 d). Coprophagous beetles were more abundant in dung pats than predatory beetles (89 and 11 %, respectively). A trophic sequence was observed in relation to age of the dung, coprophagous beetles occurring earlier in the dung than predatory beetles. The pattern was observed on two occasions during the season, though succession proceeded somewhat faster in July than in May. These results suggest that food availability and microclimatic conditions in dung pats appear to determine the successional occurrence of beetle taxa. On the other hand, coprophagous species (Aphodius) were poorly segregated along the successional axis. Null models failed to support the hypothesis that successional overlap and differences in successional mean occurrence between species could be the result of competition. Successional patterns at the specific level probably reflect differences in behaviour, such as pat location, feeding, mating, egg-laying and larva requirements, rather than competitive replacement.     

Distribution and abundance of dung beetles in fragmented landscapes

Related species utilising similar resources are often assumed to show similar spatial population structures and dynamics. This paper reports substantial ecological variation within a set of Aphodius dung beetles occurring in the same patchily distributed resource, livestock dung in pastures. We show how variation in habitat and resource selectivity, in the rate of movements between pastures, and in the distribution of local population sizes all contribute to interspecific differences in spatial population structures. Local dung beetle assemblages are compared between two landscapes with different densities of pastures. In one of the landscapes, we contrast the abundances and regional distributions of Aphodius before and after 15 years of rapid habitat loss. Different species show very dissimilar responses to changes in the structure of the landscape. Our results suggest that generalist Aphodius species, and specialist species with high dispersal powers, occur as large "patchy" populations in the landscape. In contrast, a strict pasture specialist species with limited dispersal powers (A. pusillus) forms classical metapopulations. At the community level, interspecific differences in spatial population structures make the local community composition a function of the structure of the surrounding landscape.     

Dung beetle community composition affects dung turnover in subtropical US grasslands

1. An important service in many ecosystems is the turnover and degradation of dung deposited by cattle. Dung beetles are the primary group of insects responsible for dung turnover, and factors affecting their abundance and distribution thus impact dung degradation. Lands lost to grazing due to dung buildup and pasture contamination total millions of acres per year in US pastures.
2. We evaluated the structural differences in dung beetle assemblages in natural grasslands versus a managed agroecosystem in subtropical southeastern Florida (USA). We measured the direct effect of dung longevity when dung beetle fauna normally inhabiting dung pats were excluded.
3. Our results indicate dung beetle abundance, functional diversity, and species richness have a substantial impact on the rate of dung turnover in subtropical pastoral lands with ~70% of dung removed from the soil surface after three months. Functional diversity and evenness did not have a significant positive effect on dung removal in managed, versus natural grasslands demonstrating a strong relationship between dung beetle assemblage composition and delivery of a key ecological process, dung degradation.
4. We suggest the importance of trees, which provide a thermal refuge for beetles, should be dispersed within matrixes of open pasture areas and within proximity to adjacent closed‐canopy hammocks to facilitate the exchange of dung beetles between habitats and therefore maintain the provisioning of dung degradation services by dung beetle assemblages.

     

Ecosystem functioning in relation to species identity,density, and biomass in two tunneller dung beetles

1. Species abundance, biomass, and identity are the main factors that influence ecosystem functioning. Previous studies have shown that community attributes and species identity help to maintain natural ecosystem functioning. 2. This study examined how species identity, biomass, and abundance in dung pats (i.e. density) of dung beetles affect multiple ecological functions: dung removal, seed dispersal, and germination. Specifically, two species of tunnellers were targeted: Onthophagus illyricus (Scopoli, 1763) and Copris lunaris (Linnaeus, 1758). In accordance with their natural abundance, densities ranging from 10 to 80 individuals were considered for O. illyricus, and those from two to eight were considered for C. lunaris, spanning the total biomass per treatment from 0.22 to 1.76 g. 3. Results showed that, even at higher abundance, O. illyricus is not as efficient as C. lunaris. These results show that species identity, biomass, and density are crucial factors for maintaining ecosystem functioning. The combined effect of species identity and density/biomass facilitated dung removal and seed dispersal. Conversely, species identity is the only relevant factor for germination. Moreover, relationships among functions depend on the species investigated: C. lunaris showed a positive correlation between dung removal and seed dispersal, whereas O. illyricus showed a positive correlation between germination and dung removal. 4. In conclusion, optimal ecosystem functioning depends on multiple factors, such as density and species identity, and thus also on body size, nesting strategies and ecological functions investigated. Moreover, the loss of larger and efficient species cannot be compensated by higher abundances of small species.     

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.     

Establishment of pasture species on cattle dung: the role of endozoochorous seeds

Abstract. The process of colonization of cattle dung patches and its relation to endozoochorous dispersal is analysed in a Mediterranean pasture. Dung pats dropped in spring and winter were marked, and the vegetation developing on them during the first two years of colonization was followed. The influence of endozoochorous seeds on dung colonization is assessed through the study of: (1) the vegetation on spring and winter dung pats and the seed contents of both, which is abundant in the former and negligible in the latter; and (2) the vegetation on spring dung pats compared with that of the surrounding pasture. The origin of plants growing on dung (either seeds in the manure or in the soil seed bank) was traced by studying dung colonization under controlled conditions in pots with manure put on seed-free sand, and on pasture turf. The results indicate that endozoochorous seeds germinate in manure; they are the main source of recovery in gaps generated by dung pats. The micro-succession involved is independent of the type of pasture. A small-scale spatial pattern results in which gaps of old dung are dominated by endozoochorous species. Thus, dung patches enhance the similarity between grazed communities, and the variation within communities.     

Ecosystem services provided by dung beetles in Australia

Although there are nearly 500 species of native dung beetles in Australia, most are adapted to small, hard, dry, pelletised marsupial droppings and not to dealing with the large, moist deposits of cattle. In 1788, Governor Arthur Phillip arrived at Botany Bay with five cows, two bulls, 44 sheep and seven horses: this signalled major changes in Australia. Now there are about 27 million cattle, whose annual dung production has a dry matter content of about 42 million tonnes. Until CSIRO introduced exotic dung beetles in the 1960s, the dung of these herbivores sat on the soil surface, sometimes for years, locking up organic matter, smothering pasture and polluting waterways. CSIRO introduced 53 exotic dung beetle species, of which 43 were released to the Australian mainland between 1965 and 1985. Twenty-three of these have become established, many of which have reached the natural limits of their distribution. I consider the reason for the failure of the other 30 species to establish and briefly review previous contributions to examining the role of dung beetles in delivering ecosystem services, noting that much of the published literature concerns laboratory studies. New field data are then examined on the way in which introduced species are transforming dung communities and the ecosystem services they provide. The capacity of deep-tunnelling dung beetles to transform the soil profile is examined along with their effects on pasture production and the flow of nutrients from dung on pasture. The biocontrol capacity of dung beetle activity is considered in relation to the native bush fly, Musca vetustissima, the introduced buffalo fly, Haematobia irritans exigua, and dung-borne intestinal parasites (helminths and Cryptosporidium). The rationale for introducing additional species to Australia is considered.     

Evidence that sex‐specific signals may support mate finding and limit aggregation in the dung beetle Aphodius fossor

1. In temperate climates, dung is often colonised by several species of endocoprid (dwelling) dung beetles which use pats for feeding, shelter, and reproduction. 2. Endocoprid beetles aggregate even when offered patches (dung pats) of consistent age, size, and origin, suggesting that beetles themselves might influence the attractiveness of patches to members of their own species. Both pheromones, and physical changes to the structure of dung pats caused by colonising beetles have been suggested as mechanisms facilitating intraspecific aggregation, but neither of these hypotheses have been empirically tested. 3. Using a common European dung beetle (Aphodius fossor L.), we conducted a simple choice experiment designed to test whether (i) earlier colonisation by conspecifics could alter dung attractiveness and (ii) whether attraction was influenced by sex‐specific signals. 4. We found that female beetles are repelled by dung colonised by conspecific females and are attracted to dung colonised by conspecific males. Male beetles show no evidence of attraction or repellence for dung colonised by either sex. Neither in females nor males was uncolonised dung found to be significantly more or less attractive than predicted by non‐preference. 5. Our results suggest that for A. fossor male‐produced signals may support mate finding in patchy environments, and that female‐produced signals may serve to discourage subsequent colonisation by additional females.     

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.     

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.     

Quantifying Beetle-Mediated Effects on Gas Fluxes from Dung Pats

Agriculture is one of the largest contributors of the anthropogenic greenhouse gases (GHGs) responsible for global warming. Measurements of gas fluxes from dung pats suggest that dung is a source of GHGs, but whether these emissions are modified by arthropods has not been studied. A closed chamber system was used to measure the fluxes of carbon dioxide (CO₂), methane (CH₄) and nitrous oxide (N₂O) from dung pats with and without dung beetles on a grass sward. The presence of dung beetles significantly affected the fluxes of GHGs from dung pats. Most importantly, fresh dung pats emitted higher amounts of CO₂ and lower amounts of CH₄ per day in the presence than absence of beetles. Emissions of N₂O showed a distinct peak three weeks after the start of the experiment – a pattern detected only in the presence of beetles. When summed over the main grazing season (June–July), total emissions of CH₄ proved significantly lower, and total emissions of N₂O significantly higher in the presence than absence of beetles. While clearly conditional on the experimental conditions, the patterns observed here reveal a potential impact of dung beetles on gas fluxes realized at a small spatial scale, and thereby suggest that arthropods may have an overall effect on gas fluxes from agriculture. Dissecting the exact mechanisms behind these effects, mapping out the range of conditions under which they occur, and quantifying effect sizes under variable environmental conditions emerge as key priorities for further research.     

Response of the dung beetle community to different climatic zones: Does the land use system matter?

The conversion of Brazilian savanna into exotic pastures leads to the loss of dung beetles and a decrease in their contribution to ecological functions. We hypothesized that the dung beetle communities from exotic pastures would show greater significant differences between climatic zones, when contrasted to communities from Brazilian savanna in the same region, since dung beetle assemblages in pastures are more simplified. We assessed which variables (purpose of production, type of management, percentage the habitat per buffer, soil penetration resistance, pasture area and herd size) affect more the dung beetle community in exotic pastures. We carried out this study in 48 areas of native Brazilian savannas and exotic pastures distributed across four bioclimatic zones: BZ1, hot with three dry months; BZ2, hot with 4–5 dry months; BZ3, sub-hot with 4–5 dry months and BZ4, meso-thermal with 4–5 dry months of Minas Gerais State, Brazil. In each BZ, six areas of Brazilian savannas and six areas of exotic pasture were selected. In the Brazilian savanna areas, the species richness, abundance and biomass of dung beetles did not differ between the bioclimatic zones, unlike the exotic pastures. The composition of the dung beetle community was different between land use systems and between bioclimatic zones; the interaction between the two factors was also significant. Our results provide evidence that dung beetle communities active in exotic pastures are more susceptible to climatic environmental variations than communities from more complex and stable habitats, such as savannas. Finally, the best model suggested that all the six variables combined explained about 91% of the total variability in species composition observed between sampling sites.     

Local soil characteristics determine the microbial communities under forest understorey plants along a latitudinal gradient

The soil microbial community is essential for maintaining ecosystem functioning and is intimately linked with the plant community. Yet, little is known on how soil microbial communities in the root zone vary at continental scales within plant species. Here we assess the effects of soil chemistry, large-scale environmental conditions (i.e. temperature, precipitation and nitrogen deposition) and forest land-use history on the soil microbial communities (measured by phospholipid fatty acids) in the root zone of four plant species (Geum urbanum, Milium effusum, Poa nemoralis and Stachys sylvatica) in forests along a 1700 km latitudinal gradient in Europe.Soil microbial communities differed significantly among plant species, and soil chemistry was the main determinant of the microbial community composition within each plant species. Influential soil chemical variables for microbial communities were plant species-specific; soil acidity, however, was often an important factor. Large-scale environmental conditions, together with soil chemistry, only explained the microbial community composition in M. effusum and P. nemoralis. Forest land-use history did not affect the soil microbial community composition.Our results underpin the dominant role of soil chemistry in shaping microbial community composition variation within plant species at the continental scale, and provide insights into the composition and functionality of soil microbial communities in forest ecosystems.     

Spatial distribution, resource utilisation and intraspecific competition in the dung beetle Aphodius ater

Competitive interactions in northern temperate dung beetles are poorly understood. This investigation therefore comprises a series of field and experimental work on a dung beetle species common in northern Europe, Aphodius ater, with special focus on intraspecific competitive interactions. The between-pat distribution of adult A. ater in relation to the age of sheep dung pats was studied in the field. The distribution of both sexes was contagious in the fresh pats but became more regular with increasing pat age. The successional occurrence of males and females did not differ, but immature females tended to occur in fresh pats while mature females were mainly found in older pats. With increasing age of pats, the egg load of females also increased. Egg-laying behaviour of the beetles was studied in laboratory experiments. The mean number of eggs laid per female per dung pat decreased with increasing beetle density. Thus, density-dependent processes seem to regulate resource utilisation with regard to breeding behaviour, resulting in equal exploitation of the available pats. Survival and weight of recently hatched beetles decreased with increasing initial density of eggs. Hence, in A. ater, competition between larvae for food within pats does occur. Received: 4 February 1998 / Accepted: 20 April 1998