Unexpectedly diverse forest dung beetle communities in degraded rain forest landscapes in Madagascar

Tropical forests, which harbor high levels of biodiversity, are being lost at an alarming speed. Madagascar, a biodiversity hotspot, has lost more than half of its original forest cover. Most of the remaining forests are small fragments of primary and secondary forest with differing degrees of human impact. These forests, as well as coffee and fruit plantations, may be important in supporting the forest-dependent biodiversity in Madagascar but this has been little studied. In Madagascar, dung beetles, which offer important ecosystem services, are largely restricted to forests. We examined the ability of fragmented and degraded forests to support dung beetle diversity, compared to the large areas of primary forest in eastern Madagascar. We found a general trend of a reduction of species with a loss of forest connectivity. In contrast, a higher level of forest disturbance was associated with higher species diversity. In several sites of low-quality forest as many or more species were found as in less disturbed and primary forests. The average size of dung beetles was smaller in the lower quality localities than in the primary forests. These findings suggest that many forest dung beetles in Madagascar are better adapted to forest disturbance than earlier expected, although they require some level of connectivity to surrounding forest.     

Climate as a determinant of dung beetle response to native forest replacement by cattle pastures in South America

Anthropogenic disturbance in natural ecosystems reduces the number of species in biological communities and homogenizes their composition across different regions. Climate is one of the main abiotic determinants of species distributions and different factors were proposed as the main climatic drivers. Here we explored the role of regional climate on the local response of dung beetle assemblages to the replacement of native forest by cattle pastures in South America by simultaneously contrasting three climatic hypotheses: energy, seasonality and heterogeneity. We compiled a database by searching published studies comparing dung beetle richness and composition between both native forests and cattle pastures. We calculated the proportional difference in species richness and composition between habitat types. As explanatory variables, we used seven abiotic variables grouped into the three climatic hypotheses. Energy/Productivity: mean annual temperature (°C/year) and total annual precipitation (mm/year). Seasonality: annual thermal amplitude (°C/year), the average coefficient of variation of monthly precipitation and the coefficient of average monthly variation in temperature. Heterogeneity: coefficient of variation of mean annual temperature, coefficient of variation of mean annual precipitation. Using regression analyses and a model selection procedure, we found differences in species richness between native forests and cattle pastures were explained by the coefficient of variation of mean annual precipitation, whereas changes in species composition were explained by total annual precipitation and the coefficient of variation of mean annual precipitation. The response of dung beetle assemblages to livestock grazing in South American forests was associated with precipitation variation. The heterogeneity hypothesis better explained changes in species richness following forest replacement by cattle pastures, while both energy/productivity and heterogeneity hypotheses explained the changes in species composition.     

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.

     

Response of a dung beetle assemblage along a reforestation gradient in Restinga forest

Scarabaeinae dung beetles are indicator insects used in the evaluation of the ecological effects caused by changes in habitat structure and ecosystem integrity resulting from environmental degradation. We compared dung beetle diversity in conserved restinga forests (coastal tropical moist broadleaf forest) and in reforested areas of various ages during the rainy and dry seasons, on the coast of Paraíba State, Brazil. A total of 3,634 individuals comprising 14 species were collected. In the reforested areas there was a gradual increase in species abundance relative to the area’s age, but in the conserved restinga the abundance of individuals was 10–20 times higher than that recorded in areas of recent reforestation. The highest species richness was found in the conserved restinga and in the oldest reforested area (16 years old) during the rainy season. During the dry season, when environmental conditions do not seem to favor adult survival, most of the species were found in the conserved restinga forest. The dung beetle community structure was related to the increases in habitat heterogeneity in the successional processes of the reforested areas. Our results suggest that reforested areas act as a source of and refuge for dung beetle species.     

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.     

Variations in dung beetle assemblages across a gradient of hunting in a tropical forest

Populations of large mammals are severely depleted by hunting in tropical forests, with direct effects on plant regeneration. But indirect consequences on commensal taxa depending on them for food resources, like coprophagous beetles, are less documented. Cascading effects of species loss across Scarabaeinae are expected, with likely significant negative implications for ecosystem functions. We examined dung beetle assemblages using pitfall traps at three rain forest sites in French Guiana ranging from intact mammalian fauna (Nouragues) to moderate (Kaw) and heavy (Matoury) defaunation. The site with the most depauperate mammalian fauna showed significantly lower dung beetle species richness than the two other two sites, which were not different from each other. Mean abundance and biomass per trap were not different across sites whereas community composition strongly differed among sites. A positive correlation was observed between body size and the individual contribution to dissimilarity between Nouragues and Kaw. The species contributing the most to dissimilarity were large. By contrast, one medium-sized species, dominant in Matoury, contributed the most to dissimilarity between Matoury and other sites. Diurnal genera of large tunnellers showed a higher diversity and abundance in Nouragues compared to other sites, whereas a nocturnal genus showed no differences. Large rollers were more abundant in Kaw compared to other sites. None of the groups of small beetles but one were affected by defaunation. Our results suggest that loss of large mammal populations affects dung beetle assemblage structure and causes decreasing abundance or disappearance of large tunnellers species that have a major impact on several dung beetle-mediated ecological processes.     

Large herbivores affect forest ecosystem functions by altering the structure of dung beetle communities

Dramatic increases in populations of large mammalian herbivores have become a major ecological issue, particularly in the northern hemisphere, due to their substantial impacts on both animal and plant communities through processes such as grazing, browsing, and trampling. However, little is known about the consequences of these population explosions on ecosystem functions. Here, we experimentally investigated how the population density of sika deer (Cervus nippon) in temperate deciduous forest areas in Japan affected the decomposition of mammal dung by dung beetles, which is a key process in forest ecosystems. We measured a range of environmental variables (e.g., vegetation cover, soil hardness) and the dung decomposition rate, measured as the amount of deer dung decomposed during one week, and sampled dung beetles at 16 study sites with three different deer densities (high/intermediate/low). We then used structural equation modeling to investigate the relationships between deer density, environmental variables, the biomass of dung beetles (classified into small or large species), and the dung decomposition rate. We found that the biomass of small species increased with increasing deer density, whereas that of large species was not related to deer density. Furthermore, the dung decomposition rate was positively related to the biomass of small species but unrelated to that of large species. Overall, our results showed that an increase in deer density affects the decomposition rate of mammal dung by changing the structure of dung beetle communities (i.e., increasing the number of small dung beetles). Such an understanding of how increases in large herbivore populations affect ecosystem functions is important for accurately evaluating the ecological consequences of their overabundance and ultimately managing their populations appropriately.     

Experimental warming alters the relative survival and emigration of two dung beetle species from an Australian dung pat community

To preserve insect‐mediated ecosystem services under ongoing climate change, it is necessary to first understand the impact that warming will have on the insects that provide or mediate these services. Furthermore, because responses of a species may be modified by interactions with competitors, it is informative to examine warming effects on organisms and service provision under competition. Dung beetles provide numerous services to agriculture by burying the manure of other animals. To understand the potential impacts of climate warming on ecosystem service provision, we exposed two dung beetle species (Sisyphus rubrus and Euoniticellus fulvus (Coleoptera: Scarabaeidae)), occurring together in the same experimental pats, to warming and measured reproduction (dung ball production and burial, brood production, and egg laying), pat departure behaviour and survival of both species. These two species are likely competitors in pastures in northern New South Wales. To simulate climate warming, we used custom‐built chambers to add offsets (+0, +2 or +4°C) to field recorded, diurnally fluctuating baseline temperatures. There was no direct effect of increased temperature on any measured trait in either species. We did find however that the relative survival of the two species depended on temperature; S. rubrus had a higher probability (resulting in greater odds) of surviving than E. fulvus in the +0 and +4°C offset chambers, but not in the +2°C offset chambers. Likewise, the relative likelihood of the different species leaving a dung pat was temperature dependent; in the +2°C offset chambers, E. fulvus were more likely to leave than S. rubrus, but not in the +0 and +4°C offsets chambers. Our results highlight that it may be important for future studies to consider warming effects on relative survival and emigration because such effects could potentially lead to changes in dung beetle species composition.     

Responses of an avian community to rain forest degradation

Cumulative impacts of logging and road building in a previously undisturbed tract of tropical rain forest in French Guiana were assessed by random sampling of the bird community and 1-km² plot counts of diurnal raptors. Similar surveys were carried out during road construction and 15 years later within 5km of the road and were also compared with the undisturbed bird community of a nearby primary forest. The main disturbance was the change in forest structure brought about by logging and secondarily the road opening, the roadside second growth and the depletion of large vertebrates by hunting pressure. The responses of different bird guilds were highly divergent according to their natural habitat requirements, their diet and their vulnerability to hunting. They resulted in a moderate decrease in overall species richness and equitability. All species pooled, the abundance of 118 species was lowered, that of 45 species did not change appreciably and 89 were favored or even appeared. The guilds most affected were those with large body sizes, from terrestrial foragers to canopy frugivores, mostly by hunting, and the open understorey specialists, notably large insectivores and mixed flock members, because of changes in forest structure after logging. The guilds favored by logging and road opening were many of the hummingbirds, upper canopy frugivores and omnivores, and gap, edge or low secondary growth specialists. Recommendations to minimize the negative consequences of human exploitation in rain forests include reducing the width of deforested roadsides to no more than 10–15m on either side, implementing much more careful logging practices and strongly limiting hunting pressure in newly opened areas.     

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.     

Gut microbial community response to herbicide exposure in a ground beetle

Gut microbiota plays a key role in physiological processes of insects, including nutritional metabolism, development, immunity and detoxification. Environmental stressors such as herbicides, used to optimize and improve crop yields, may interfere with the mutualistic relationships causing negative consequences for the host health. Dinitroaniline herbicides, for example pendimethalin, are used worldwide in pre-emergence application to control grass and some broadleaf weeds. They target microtubules to arrest cell division and inhibit the development of roots and shoots. Effects of a pendimethalin-based herbicide were assessed on the gut microbial community of Pterostichus melas italicus Dejean, 1828 (Coleoptera, Carabidae). The exposure effect was tested in vivo by using a recommended field rate (4 L per ha, 330 gL⁻¹ of active ingredient) and evaluating the variability of responses in 21 days, corresponding to the half-life of pendimethalin. The 16S rRNA sequencing data showed that the gut lumen was dominated by Proteobacteria, Firmicutes, Fusobacteria, Tenericutes and Bacteroidetes. The exposure interfered with the bacterial community richness and diversity associated with the gut from 2 days after the treatment. The differential abundance analyses highlighted a shift involving Lactobacillaceae, Streptomycetaceae, Neisseriaceae, Ruminococcaceae and Enterobacteriaceae. An increase in species such as Enterobacter sp., Pseudomonas sp., Pantoea sp and Paracoccus sp. involved in the herbicide degradation was also recorded after 21 days of exposure. Phylogenetic investigation of communities by reconstruction of unobserved states (PICRUSt) analysis indicated that the exposure has effects on the most predicted functional categories of gut microbiota related to metabolic function including carbohydrate, amino acid and lipid metabolism. These results demonstrate that pendimethalin can impact microbial communities associated with generalist predators inhabiting croplands leading to severe implications for the species’ ecological role. Understanding the effects of herbicides such as pendimethalin on ground beetles may help to protect beneficial soil insects that have a crucial role in the ecosystem services.     

Edge effects on dung beetle assemblages in an Andean mosaic of forest and coffee plantations

In landscapes dominated by agriculture, conspicuous edges often occur between landscape elements. However, there is disagreement about the existence and intensity of edge effects, and information about species‐specific responses remains scarce. Studying such edge effects can help elucidate functional landscape connectivity and contribute to agricultural management. We, therefore, assessed whether sun‐grown coffee represents a barrier to dung beetles in an Andean agricultural landscape. We also evaluated whether the response to edge effects differs among species. We found that diversity and abundance tend to decrease from forest to sun‐grown coffee and that there are sharp increases in species turnover at the forest–coffee edge. We detected several different species‐specific responses to the forest–coffee edge, suggesting differences in the mobility of the species (or spillover) and in the degree of penetration that takes place from forest patches to sun‐grown coffee plantations. This study demonstrates that the sun‐grown coffee matrix constitutes a barrier to forest species and suggests that the forest–coffee ecotone is more complex than expected. Our results support the notion that the conservation value of native forest patches in agricultural scenarios depends on the functional connectivity of forest units in the landscape to maximize the opportunities species have to disperse through the agricultural matrix.     

Diversity and seasonal fluctuations in dung beetle (Coleoptera) community in Kurukshetra,India

Five pitfall traps baited with 150–200 g of fresh cattle dung were installed for 24 h at weekly intervals. A total of 991 dung beetles from 11 genera, 31 species and three subfamilies was obtained. The community was dominated by Oniticellus spinipes individuals by 32.3%. Ten species appeared only once during the collection period and species composition and dominance changed throughout the period. The overall pattern we detected in the organization of the dung beetle community is that the species richness, abundance and diversity rise in September and the 2^nd week of October. The dung beetle community was found to be affected by season.     

Effect of forest fragmentation on dung beetle communities and functional consequences for plant regeneration

I compared dung beetle communities and assessed some of their functional effects (dung removal, seed burial, seedling establishment) in continuous forest with those in 1-ha and 10-ha forest fragments in Central Amazonia. I followed the fate of seeds until seedling establishment for three native tree species, using clean seeds and seeds surrounded by dung. The 1-ha fragments had half the number of dung beetle species captured in continuous forest and in 10-ha fragments. The continuous forest sites and the 1-ha fragments had similar number of individuals, but in the 10-ha fragments dung beetles were twice as abundant. Mean beetle size increased with increasing forest area. Dung removal and seed burial rates were higher in continuous forest than in forest fragments. Seed predation rates were higher in the forest fragments. In all sites, the proportion of seedlings established from seeds surrounded by dung vs clean seeds was the same, and it was the same in continuous forest vs fragments. When comparing seeds that remained on the forest floor with seeds buried by dung beetles, a higher percentage of seedlings established from the latter. Conservation programs that aim to maintain the regeneration ability of forest fragments must incorporate all the important components involved in seedling establishment; in Central Amazonia these include dung beetles as secondary dispersers. It is important that studies start measuring directly not only the first-order effects of forest fragmentation on species, but also the higher-order functional effects.     

Short‐term assessment of dung beetle response to carbosulfan treatment against desert locust in Sudan

The beneficial role of dung beetles (Coleoptera: Scarabaeidae) is well known. Potential risks to these beetles from the widespread use of insecticides against the desert locust, a significant plant pest in Africa, the Near East and South West Asia, have not been studied previously. Short‐term responses of dung beetles to carbamate carbosulfan (Marshal®, ultra low‐volume formulation, 100 g active ingredient ha^?1) were assessed during desert locust control operations at five sites within two major biotopes: Acacia tortilis shrubland and cultivated wetland; on the Red Sea Coast of Sudan. The study took place during January–February 2004. At each site, fresh dung from Zebu cows was placed in areas targeted for desert locust control. Dung pats were placed in plots in two areas and left for 24 h, before and after insecticide application. Beetles were extracted by floatation. There was a significant decrease in abundance between the pre‐ and post‐spray period in treated areas for the Scarabaeinae species Onthophagus margaritifer (a dark colour morph). In contrast, it was found that Aphodius lucidus and beetles at the subfamily level of Aphodiinae increased in numbers after insecticide treatment. Mortality and sublethal impacts as well as a repellent effect of the insecticide may explain the decrease in Onthophagus margaritifer, while the increase in Aphodiinae beetles could be an indirect response to lower numbers of Scarabaeinae beetles in competing for the same resource. These organisms and the applied methodology may be useful for environmental monitoring of desert locust control, thus further studies are suggested. The assessment also revealed a marked difference between the two biotopes with high abundance and species richness of dung beetles in A. tortilis shrubland, while these measures were low in the cultivated wetland. Five new species of dung beetles for Sudan were found in this study.     

Changes to the N cycle following bark beetle outbreaks in two contrasting conifer forest types

Outbreaks of Dendroctonus beetles are causing extensive mortality in conifer forests throughout North America. However, nitrogen (N) cycling impacts among forest types are not well known. We quantified beetle-induced changes in forest structure, soil temperature, and N cycling in Douglas-fir (Pseudotsuga menziesii) forests of Greater Yellowstone (WY, USA), and compared them to published lodgepole pine (Pinus contorta var. latifolia) data. Five undisturbed stands were compared to five beetle-killed stands (4–5 years post-outbreak). We hypothesized greater N cycling responses in Douglas-fir due to higher overall N stocks. Undisturbed Douglas-fir stands had greater litter N pools, soil N, and net N mineralization than lodgepole pine. Several responses to disturbance were similar between forest types, including a pulse of N-enriched litter, doubling of soil N availability, 30–50 % increase in understory cover, and 20 % increase in foliar N concentration of unattacked trees. However, the response of some ecosystem properties notably varied by host forest type. Soil temperature was unaffected in Douglas-fir, but lowered in lodgepole pine. Fresh foliar %N was uncorrelated with net N mineralization in Douglas-fir, but positively correlated in lodgepole pine. Though soil ammonium and nitrate, net N mineralization, and net nitrification all doubled, they remained low in both forest types (<6 μg N g soil^?1 NH₄ ⁺or NO₃ ^?; <25 μg N g soil^?1 year^?1 net N mineralization; <8 μg N g soil^?1 year^?1 net nitrification). Results suggest that beetle disturbance affected litter and soil N cycling similarly in each forest type, despite substantial differences in pre-disturbance biogeochemistry. In contrast, soil temperature and soil N–foliar N linkages differed between host forest types. This result suggests that disturbance type may be a better predictor of litter and soil N responses than forest type due to similar disturbance mechanisms and disturbance legacies across both host–beetle systems.     

Regional and local influence of grazing activity on the diversity of a semi-arid dung beetle community

This study analyses the effect of resource availability (i.e. sheep dung) on dung beetle communities in an arid region of Central Spain, both at regional and at local scales. A total of 18 sites within 600 km² were sampled for the regional analysis and 16 sites within the 30 km² of an Iberian municipality were sampled for the local analysis. Spatial and environmental characteristics of sampling sites were also compiled at both scales, including measures of grazing activity (livestock density at regional scale, and two counts of rabbit and sheep dung at local scale). At a regional scale, any environmental or spatial variable can help to explain the variation in abundance. However, species richness was related to summer precipitation and composition was related to elevation. At local scale, abundance is not significantly related to any of the environmental variables, but species richness was related to the local amount of sheep dung (27% of variance). The amount of dung in a 2‐km buffer around the site accounts for 27–32% of variance in abundance and 60–65% of variance in species richness. The presence of the flock with the highest sheep density explains 53% of abundance variability and 73% of species richness variance. A cluster analysis of localities identified two main groups, one characterized by a lower abundance and species richness that can be considered a nested subsample of the species‐rich group. The mean and maximum amount of sheep dung in the sites separated by less than 2 km are the only significant explanatory variables able to discriminate both groups. These results suggest that grazing intensity (and the associated increase in the amount of trophic resources) is a key factor in determining local variation in the diversity and composition of dung beetle assemblages. However, dung beetle assemblages are not spatially independent at the analysed resolution, and the amount of dung in the surroundings seems to be more important for locally collected species than the dung effectively found in the site. Although differences in the availability and quantity of trophic resources among nearby sites could be affecting the population dynamics and dispersion of dung beetles within a locality, sites with larger populations, and greater species numbers would not be able to exercise enough influence as to bring about a complete local faunistic homogenization.     

The role of local and regional processes in shaping dung beetle communities in tropical forest plantations in Borneo

We examine whether rain forest dung beetle species found in plantations in Sabah, northern Borneo, tend to be endemic or geographically widespread. In addition, linear regressions of abundance vs. distance from a major river in primary rain forest are calculated to see if species found in plantation forest show affinity to one specific biotope (riverine vs. interior forest) in their natural habitat. Results show that 14 of the 40 species recorded from plantations are endemic to Borneo. Only edge‐specialist endemic species are found in plantation forest, with no interior‐forest specialists recorded. Data suggest that endemic species that are adapted to more exposed conditions in primary rain forest, such as riverine species, can in some instances tolerate man‐made habitats. Twenty‐nine species (±SE 4.0) per transect are recorded from plantation transects, whereas 44.2 (±1.7 SE) are recorded in primary rain forest. As species richness is much lower in plantations than natural forest, implying loss of biodiversity, we conclude that measures of biogeographic distinctiveness, whereby endemic species confer higher values, may be misleading unless they take into account edge‐affinity. Local‐ as well as regional‐distributional data may therefore be needed to interpret correctly patterns of species assemblages in derived forest ecosystems.