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.

     

Landscape diversity of pasture dung beetle communities in the central region of mainland Japan and implications for conservation management

We studied the diversity of dung beetle communities in Japanese pastures to identify the factors that maintain or enhance the diversity of dung beetles at a landscape scale. We surveyed dung beetles from 17 pastures located in the northeastern part of Tochigi Prefecture, which is in the center of mainland Japan. From 1999 to 2001, surveys were conducted during the 6-month grazing period (May to October) by using dung baited basket traps. We also collected information about the environmental conditions and pasture management practices. Twenty-five dung beetle species belonging to Geotrupinae, Scarabaeinae, and Aphodiinae (including 13 tunneler and 12 dweller species) were recorded. The abundance of dweller species decreased with increasing elevation, possibly because of the effect of rainfall, whereas the species richness of tunneler species was affected by cattle disturbance and soil condition. Beetle species richness significantly increased with the number of years that the pastures had been grazed. Ivermectin administration did not appear to have any adverse effect on dung beetle abundance, species richness, or species diversity. The dung beetle datasets of the current study (including specific tunneler and dweller beetle groups) supported the widely documented positive relationship between local abundance and species distribution ranges. The within pasture, within area, and between area hierarchical additive partitioning of regional total diversity indicated that landscape-scale management should be implemented to conserve the regional diversity of the dung beetle communities inhabiting Japanese pastures.     

Experimental mixtures of dung fauna affect dung decomposition through complex effects of species interactions

Dung fauna plays an important role in dung decomposition, a key ecosystem process in nutrient cycling in grazed grasslands. The diversity of a three‐species community (dung beetles, dung flies and epigeic earthworms) was systematically manipulated to produce different relative abundance distributions (evenness levels) and the resulting communities were introduced to standardised dung pats in laboratory experiments. Dung decomposition was modelled using an analysis that disentangled species identity effects and the interactions among species that contribute to the diversity effect. This revealed that the net diversity effect was composed of positive (fly–earthworm), negative (beetle–earthworm) and neutral (fly–beetle) effects of species interaction. These pairwise interactions resulted in complex, but systematically varying and predictable effects on dung decomposition as the relative abundances of species changed. Decomposition was consistently greater in communities with higher decomposer biomass. The utility of the adopted analytical approach was emphasised by comparison with an ANOVA that found that dung decomposition did not differ among evenness levels. Thus, the averaging of decomposition across different community structures within evenness levels masked the different effects of species interactions. These results highlight methodological insights into the quantification of diversity–function relationships.     

How do sheep affect plant communities and arthropod populations in temperate grasslands?

Grasslands being used in sheep farming systems are managed under a variety of agricultural production, recreational and conservational objectives. Although sheep grazing is rarely considered the best method for delivering conservation objectives in seminatural temperate grasslands, the literature does not provide unequivocal evidence on the impact of sheep grazing on pasture biodiversity. Our aim was therefore to review evidence of the impacts of stocking rate, grazing period and soil fertility on plant communities and arthropod populations in both mesotrophic grasslands typical of agriculturally improved areas and in native plant communities. We therefore conducted a literature search of articles published up to the end of the year 2010 using ‘sheep’ and ‘grazing’ as keywords, together with variables describing grassland management, plant community structure or arthropod taxa. The filtering process led to the selection of 48 articles, with 42 included in the stocking rate dataset, 9 in the grazing period dataset and 10 in the soil fertility dataset. The meta-analysis did not reveal any significant trends for plant species richness or plant community evenness along a wide stocking rate gradient. However, we found frequent shifts in functional groups or plant species abundance that could be explained by the functional properties of the plants in the community. The meta-analysis confirmed that increasing soil fertility decreased plant species richness. Despite the very limited dataset, plant species richness was significantly greater in autumn-grazed pastures than in ungrazed areas, which suggests that choosing an appropriate grazing period would be a promising option for preserving biodiversity in sheep farming systems. Qualitative review indicated that low grazing intensity had positive effects on Orthoptera, Hemiptera (especially phytophagous Auchenorrhyncha) and, despite a diverse range of feeding strategies, for the species richness of Coleoptera. Lepidoptera, which were favoured by more abundant flowering plants, also benefited from low grazing intensities. Spider abundance and species richness were higher in ungrazed than in grazed pastures. In contrast, there are insufficient published studies to draw any firm conclusions on the benefits of late grazing or stopping fertilization on insect diversity, and no grounds for including any of this information in decision support tools at this stage.     

Effects of the progressive abandonment of grazing on dung beetle biodiversity: body size matters

Dung beetles are an ecologically important group of insect species inhabiting semi-natural grasslands in Europe. Despite their ecological importance, several dung beetle species are currently facing local threats mainly stemming from changes in land use practices, including the abandonment of pasturelands. The aim of this study was to analyze the impact of the progressive abandonment of grazing lands on dung beetle alpha diversity, abundance, biomass and body size classes. Areas representing a range of trophic resource availability were compared: (i) abandoned, (ii) low and (iii) moderate grazing intensity. A total of 136,884 specimens belonging to 56 species (3 Geotrupinae; 16 Scarabaeinae; 37 Aphodiinae) of dung beetle were collected. Pastureland abandonment was shown to lead to a decrease in alpha diversity and biomass density, a reduction of 22 and 78% respectively in abandoned areas. From a biomass standpoint, the effects of pastureland abandonment varied according to the body size classes of the dung beetles, with larger species proving more susceptible to local extinction. Such body-size dependent variations led us to propose the use of both biomass and abundance data in making assessments since they are differentially sensitive to trophic resource availability. Hence, from a conservationist standpoint, the obtained results suggest that traditional pastureland management with low-moderate grazing intensity should be maintained.     

Effects of livestock breed and grazing pressure on ground‐dwelling arthropods in Cantabrian heathlands

Abstract 1. The effects of different goat breeds and grazing pressures on epigeal arachnids (Lycosidae and Opiliones) and ground beetles (Carabidae) were assessed in a heath–gorse shrubland located in the north of the Iberian Peninsula. Three treatments (low grazing pressure with Cashmere breed, high grazing pressure with Cashmere breed, and high grazing pressure with local Celtiberic breed) with three replicates of each treatment were allocated randomly to nine plots. 2. Arthropod fauna abundance, diversity, and species richness were estimated using pitfall trapping over 3 years (2003, 2004, and 2005). Vegetation cover, composition, and height were assessed along transects and above‐ground plant biomass was sampled in quadrats. 3. Lower grazing pressure led to lower herbaceous cover and higher heather cover than high grazing pressure. Overall, it did not enhance arthropod diversity or abundance, but the abundance and species richness of carabids, and abundance of lycosids like Pardosa pullata, were higher under high stocking rates, whereas Opiliones were favoured by the low stocking rates. Although it has been assumed that traditional breeds enhance biodiversity, the present study found no significant differences for both arthropod fauna and vegetation between the goat breeds. 4. Arthropod–vegetation relationships were examined by constrained ordination. Fauna assemblages followed a gradient from closed canopy shrubland areas, in the lowest grazing pressure, to grassier open canopy areas resulting from higher grazing pressure, revealing the importance of grazing management as a driver of arthropod community structure of the heathland.     

松嫩草地牛粪中大型节肢动物种类组成及种群动态变化

研究并分析了松嫩草地牛粪中大型节肢动物种类组成及动态变化.结果表明,在草地牛粪中共调查到大型节肢动物7目20科43种,以鞘翅目、膜翅目和双翅目为主.依其食性不同,分为粪食性型(8科19种)、腐食性型(4科4种)、植食性型(4科6种)、捕食性型(5科11种)、杂食性型(2科5种)、尸食性型(1科1种)和寄生性型(1科1种).根据鞘翅目金龟总科中营粪食性型生活的粪甲虫对牛粪利用方式的不同,分为外生粪甲虫(6种)、内生粪甲虫(5种)、远生粪甲虫(2种)和偷窃寄生粪甲虫(1种).除了双顶嗡蜣螂,牛粪中大型节肢动物数量与鲜牛粪重量具有显著正相关关系(p＜0.01).粪食性型种类数和个体数量随鲜牛粪堆置时间的延长而迅速减少,7d后消失,其个体总数量与鲜牛粪堆积时间呈显著的负相关关系(p＜0.01),而与牛粪含水量呈显著正相关关系(p＜0.01),各种大型节肢动物因其食性不同在鲜牛粪中占据不同的生态位.鲜牛粪中大型节肢动物以粪食性型占绝对优势,而干牛粪中则以捕食性型和杂食性型为主.牛粪中大型节肢动物种群季节动态变化受温度、降水量与牛粪含水量影响,鲜牛粪中大型节肢动物优势种为直蜉金龟(69.25%)和肖秋家蝇与东方角蝇幼虫(25.12%).4、5、6月份和9月份,优势种为直蜉金龟,7月和8月份,优势种为蝇幼虫.     

Dung‐associated arthropods influence foraging ecology and habitat selection in Black‐necked Cranes (Grus nigricollis) on the Qinghai–Tibet Plateau

Variation in grassland vegetation structure influences the habitat selection of insectivorous birds. This variation presents a trade‐off for insectivorous predators: Arthropod abundance increases with vegetation height and heterogeneity, but access to arthropod prey items decreases. In contrast, grazing by large herbivores reduces and homogenizes vegetation, decreasing total arthropod abundance and diversity. However, the presence of livestock dung may help counteract the overall reduction in invertebrates by increasing arthropods associated with dung. It is unclear, however, how the presence of arthropod prey in dung contributes to overall habitat selection for insectivorous birds or how dung‐associated arthropods affect trade‐offs between vegetation structure, arthropod abundance, and access to prey. To explore these relationships, we studied habitat selection of the Black‐necked Crane (Grus nigricollis), a large omnivorous bird that breeds on the Qinghai–Tibet Plateau. We assessed the relationships between habitat selection of cranes and vegetation structure, arthropod abundance, and the presence of yak dung. We found that Black‐necked Cranes disproportionately foraged in grassland patches with short sward height, low sward height heterogeneity, and high numbers of dry yak dung, despite these habitats having lower total arthropod abundance. Although total arthropod abundance is lower, these habitats are supplemented with dry yak dung, which are associated with coleopteran larvae, making dung pats an indicator of food resources for breeding Black‐necked Cranes. Coleopteran adults and larvae in yak dung appear to be an important factor influencing the habitat selection of Black‐necked Cranes and should be considered when assessing grassland foraging trade‐offs of insectivorous birds. This research provides new insights into the role of livestock dung in defining foraging habitats and resources for insectivorous predators.     

Does natural reforestation represent a potential threat to dung beetle diversity in the Alps?

Traditional agro-pastoral practices are in decline over much of the Alps, resulting in the complete elimination of livestock grazing in some areas. Natural reforestation following pastoral abandonment may represent a significant threat to alpine biodiversity, especially that associated with open habitats. This study presents the first assessment of the potential effects of natural reforestation on dung beetles by exploring the relationships between the beetle community (abundance, diversity, species turnover and assemblage structure) and the vegetation stages of ecological succession following pastoral abandonment. A hierarchical sampling design was used in the montane belt of the Sessera Valley (north-western Italian Alps). Dung beetles were sampled across 16 sampling sites set in four habitat types corresponding to four different successional stages (pasture, shrub, pioneer forest and beech forest) at two altitudinal levels. The two habitats at the extremes of the ecological succession, i.e. pasture and beech forest, had the greatest effect on the structure of local dung beetle assemblages. Overall, dung beetle abundance was greater in beech forest, whereas species richness, Shannon diversity and taxonomic diversity were significantly higher in pasture, hence suggesting this latter habitat can be considered as a key conservation habitat. Forests and pastures shared a lower number of species than the other pairs of habitats (i.e. species turnover between these two habitats was the highest). The two intermediate seral stages, i.e. shrub and pioneer forest, showed low dung beetle abundance and diversity values. Local dung beetle assemblages were also dependent on season and altitude; early-arriving species were typical of pastures of high elevation, whereas late-arriving species were typical of beech forests. It is likely that grazing in the Alps will continue to decrease in the future leading to replacement of open habitats by forest. This study suggests therefore that, at least in the montane belt, reforestation may have potentially profound and negative effects on dung beetle diversity. Maintaining traditional pastoral activities appears to be the most promising approach to preserve open habitats and adjacent beech forests, resulting in the conservation of species of both habitats.     

Intermediate disturbance promotes diversity and the conservation of dung beetles (Scarabaeoidea: Scarabaeidae and Aphodiidae) in the Eastern Cape,South Africa

Environmental fluctuations, such as changes in climate, agricultural management and anthropogenic land-use patterns can affect the diversity of organisms inhabiting an area. Losses of biodiversity alter ecosystems processes, eroding their capacity to deliver ecosystem services. Dung beetles are critical ecosystem service providers, making them an ideal ecological indicator to explore the effects of land-use change on biodiversity. Dung beetles were sampled across three land-use types, in the summers of 2015 and 2016 in the Eastern Cape province, South Africa. Game ranching is regarded as a relatively low-intensity land use type. It was compared with cattle ranching (medium intensity) and dairy farming (high intensity) to examine their effect on dung beetle assemblage metrics (abundance, species richness and true Shannon diversity index), guild diversity (as nesting guilds) and spatial turnover. The intermediate grazing intensity of cattle ranching supported a higher abundance and diversity of both whole dung beetle assemblage and the nesting guilds, followed by the game ranches and then dairy farms. Differences between the sampling years were dependent on the beetle nesting guild, and largely correlated with rainfall and temperature. Cattle and game ranches shared a higher number of species than either shared with dairy farms. Whittaker's Beta-diversity index showed the highest species turnover between game ranches and dairy farms. A mix of game and cattle ranching, minimising dairy farming or restricting it to already ecological degraded sites, appears the best alternative for maintenance of dung beetle diversity and their ecosystem services. The year-to-year trends of the data were in general consistent, confirming that dung beetles are reliable ecological indicators; but also suggest that climate change that affects rainfall will result in the reduction of the abundance and diversity of this key ecological group.     

Effects of plant diversity, plant productivity and habitat parameters on arthropod abundance in montane European grasslands

Arthropod abundance has been hypothesized to be correlated with plant diversity but the results of previous studies have been equivocal. In contrast, plant productivity, vegetation structure, abiotic site conditions, and the physical disturbance of habitats, are factors that interact with plant diversity, and that have been shown to influence arthropod abundance. We studied the combined effect of plant species diversity, productivity and site characteristics on arthropod abundance in 71 managed grasslands in central Germany using multivariate statistics. For each site we determined plant species cover, plant community biomass (productivity), macro- and micronutrients in the soil, and characterized the location of sites with respect to orographic parameters as well as the current and historic management regimes. Arthropods were sampled using a suction sampler and classified a priori into functional groups (FGs). We found that arthropod abundance was not correlated with plant species richness, effective diversity or Camargo's evenness, even when influences of environmental variables were taken into account. In contrast, plant community composition was highly correlated with arthropod abundances. Plant community productivity influenced arthropod abundance but explained only a small proportion of the variance. The abundances of the different arthropod FGs were influenced differentially by agricultural management, soil characteristics, vegetation structure and by interactions between different FGs of arthropods. Herbivores, carnivores and detritivores reacted differently to variation in environmental variables in a manner consistent with their feeding mode. Our results show that in natural grassland systems arthropod abundance is not a simple function of plant species richness, and they emphasize the important role of plant community composition for the abundance patterns of the arthropod assemblages.     

Evaluating long-term ivermectin use and the role of dung beetles in reducing short-term CH4 and CO2 emissions from livestock faeces: a mesocosm design under Mediterranean conditions

1. Greenhouse gas (GHG) emissions from livestock contribute significantly to global warming, and a reduction of this source of emissions is crucial in achieving the goal of mitigating global warming. 2. CO₂ and CH₄ emissions from dung pats were analysed by means of a mesocosm experiment in a Mediterranean ecosystem. The experiment consisted of a total of 30 mesocosms distributed across three treatments: a well-preserved, undisturbed dung beetle assemblage associated with organic livestock; a dung beetle assemblage that was impoverished as a result of the long-term use of veterinary medical products; and a control treatment without dung beetles. 3. Corrections related to insect respiration allow researchers to provide more precise measurements of CO₂ emissions from dung, especially in the initial and final phases of dung exposure, when the percentage of CO₂ emitted by dung beetles can become greater than the emissions from the dung pats themselves. 4. The effects of dung beetles on CO₂ and CH₄ emissions are much more accentuated in warm-temperate conditions than in northern temperate areas previously studied. Mediterranean assemblages remove and spread dung faster and more effectively than do northern dung beetle assemblages characterised by a lower functional richness and beetle abundance and biomass. 5. From a livestock management viewpoint, mesocosms representing areas with impoverished dung beetle assemblages, due to the long-term use of veterinary medical products, such as ivermectin, emitted 1.6- and 2.8-fold higher total CO₂ and CH₄, respectively, than mesocosms mimicking sites with untreated livestock.     

Changes in arthropod diversity along a land use driven gradient of shrub cover in savanna rangelands: identification of suitable indicators

Shrub encroachment linked to heavy grazing has dramatically changed savanna landscapes, and is a major form of rangeland degradation. Our understanding of how shrub encroachment affects arthropod communities is poor, however. Here, we investigate the effects of shrub encroachment on abundance and diversity of ground-dwelling (wingless) arthropods at varying levels of shrub cover in the southern Kalahari. We also ascertain if invertebrate assemblage composition changes with habitat structure and identify which aspects of habitat structure (e.g., grass cover, herbaceous plant cover, shrub density) correlate most strongly with these changes. Ant, scorpion and dung beetle abundance increased with shrub cover, whereas grasshoppers and solifuges declined. Spider and beetle abundance exhibited hump-shaped relationships with shrub cover. RTU richness within orders either mirrored abundances, or exhibited no trend. Shrub density was the habitat component most correlated with similarities between invertebrate assemblages. Ground-dwelling arthropods showed clear shifts in species assemblage composition at a similarity level of 65% according to shrub density. Changes in indicator species showed that within the Tenebrionidae (darkling beetles), certain species respond positively to shrub thickening, replacing other species within the Family. Small-bodied, wingless Scarabaeidae (dung beetles) tended to increase with increased shrub density and three species emerged as significant indicators of more thickened habitats, although this might be a response to greater dung availability, rather than habitat structure itself. We conclude that because ground-dwelling invertebrates showed such clear responses in species assemblage composition, they present excellent candidates for use as indicator species in further studies into bush encroachment.     

Correlations of stocking with the cryptogamic soil crust of a semi-arid rangeland in southwest Queensland.

Abstract The soil crust community from a sub-tropical grassland in southwest Queensland was found to include 34 taxa with cyanobacteria, other algae, lichens, fungi, mosses and liverworts represented. Cyanobacteria and liverworts were the major components of the cryptogamic cover. This is a significant component of the biodiversity of the region. Changes in the structure of this community were significantly correlated with distance from a linear water supply (bore drain) and with dung density. It was concluded that hoof impact by grazing stock had measurably affected the cryptogamic community even under a moderate stocking policy. This research suggests that management for sustainable use of low-nutrient rangelands should include consideration of soil crust condition.     

Dung Beetle Assemblages and Seasonality in Primary Forest and Forest Fragments on Agricultural Landscapes in Budongo, Uganda

Very little is known about the diversity of arthropods in the fast-disappearing fragments of natural forests in sub-Saharan Africa. This study investigated: (1) the influence of forest fragment characteristics on dung beetle species richness, composition, abundance, and diversity; and (2) the relationship between dung beetle assemblages and rainfall pattern. Beetles were sampled through 12 mo using dung baited pitfall traps. A total of 18,073 dung beetles belonging to three subfamilies and 45 species were captured. The subfamily Scarabaeinae was the most abundant (99%) and species rich (89%). Fast-burying tunnellers (paracoprids) were the most dominant functional group. Catharsius sesostris, Copris nepos , and Heliocopris punctiventris were the three most abundant species, and had the highest contributions to dissimilarities between forests. With few exceptions, dung beetle abundance, species richness, and diversity were generally higher in larger forest fragments (100–150 ha) than in smaller ones (10–50 ha) and the nature reserve (1042 ha). Forest fragment size had a highly significant positive relationship with beetle abundance, but only when the nature reserve is excluded in the analysis. Dung beetle abundance and species richness showed direct weak relationships with litter depth (positive) and groundcover (negative) but not tree density, tree species richness, and fragment isolation distance. Dung beetle abundance and species richness were strongly correlated with monthly changes in rainfall. Results of this study indicate that forest fragments on agricultural lands in the Budongo landscape, especially medium-sized (100–150 ha) ones, represent important conservation areas for dung beetles.     

Dung beetles and their ecological functions in three agroforestry systems in the Lacandona rainforest of Mexico

There is growing interest in evaluating the impact that management intensity of agroecosystems has on animal communities and their ecological functions. Dung beetles are a highly used focal taxon for assessing the effects of anthropogenic disturbances and management practices on biodiversity. In the Lacandona rainforest region in southern Mexico, we quantified several metrics of the dung beetle community (number of species, number of individuals, total biomass, mean beetle size) and four of their ecological functions (dung removal, soil excavation, seed dispersal, seed exhumation) in conserved rainforest and three agroforestry systems with different management intensities: rustic cocoa, polyculture cocoa, and rubber monoculture. We also assessed the correlation between dung removal and the other functions, as well as the relationships between functions and community metrics. Land-use type affected the dung beetle communities as well as their functions, with negative effects on response variables in the most intensely managed agroecosystems (polyculture cocoa and rubber). Rustic cocoa had values similar to those of the conserved forest for all functions and community metrics, except the mean number of species per trap. Dung removal was correlated with the other ecological functions. The mean number of species per trap was significantly associated with all four functions. In our study region rustic cocoa plantations favor the maintenance of a high proportion of dung beetle species and maintain their ecological functions. Our findings corroborate that agroecosystems with less intense management may contribute to buffering the effects of landscape homogenization caused by more intensely managed agroecosystems, such as rubber plantations.     

荒漠草原马粪中粪金龟子组成及分解作用

2007年4-10月,采用野外和室内实验相结合的方法,研究了荒漠草原马粪中的粪金龟子群落组成和对马粪分解的影响.共捕获粪金龟子13528头,隶属于3科4属26种,优势种为符号蜉金龟(Aphodius comma),占总捕获量的88.5%,不同季节粪金龟子群落组成明显不同;在鲜马粪中活动的粪金龟子以粪居型蜉金龟为主,在鲜马粪堆置3d时种数和个体数达到最高,6d后全部离开;马粪经过136 d分解,残留率为52.5%,残留粪样中氮和磷释放较慢,钾释放较快;结合室内分解实验的结果,认为粪金龟子主要通过在马粪中活动对其分解产生影响,并与风蚀作用成为荒漠草原马粪块破碎的重要动因,主要发生在春季和秋季.     

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.     

Aggregated distribution of resources creates competition refuges for rainforest dung beetles

This study examines the effects of resource distribution on colonisation, intra- and interspecific aggregation, and the occurrence of low-density, competition refuges for tropical dung beetles. In field experiments from central Peru, using dung pats (resource patches) of different volumes, the numbers of interacting species and total beetle biomass at individual pats increased with increasing pat volume. In two of three separate experiments (including an experiment that also varied patch density), this represented a decrease in the biomass of beetles per unit volume (biomass-density) at larger patches. The numbers of interacting tunneller species and tunneller biomass-density were also related to the distance between pats (patch density) in one of two experiments with constant numbers of pats. Closely positioned pats had generally fewer interacting species and a lower biomass-density of beetles. For the most abundant Dichotomius species, interspecific associations increased as distances increased between dung pats. The numbers of interacting species and biomass-density declined linearly under the combined effects of increasing patch density and local patch abundance in 25 m² plots. In experimentally placed grids with large numbers of pats, colonisation of pats at the edge of the grids was generally higher than at the centre of the grids for tunnellers and Eurysternus spp. but not for ball rollers; however, at least tunnellers did not readjust to avoid patches with high densities of competitors. These results indicate that an aggregated distribution of dung and natural variability in patch size contribute to species coexistence by creating low-density refuges for weaker competitors.     

Disintegration of Dung Pats from Cattle Treated with the Ivermectin Anthelmintic Bolus,or the Biocontrol Agent Duddingtonia flagrans

An experiment was performed during the grazing seasons of 1998, 1999 and 2000 to study the influence of the antiparasitic drug ivermectin and the nematophagous fungus Duddingtonia flagrans on cattle dung disintegration. The faeces originated from groups of animals that were part of a separate grazing experiment where different control strategies for nematode parasite infections were investigated. Each group consisted of 10 first-season grazing cattle that were either untreated, treated with the ivermectin sustained-release bolus, or fed chlamydospores of D. flagrans. Faeces were collected monthly on 4 occasions and out of pooled faeces from each group, 4 artificial 1 kg dung pats were prepared and deposited on nylon mesh on an enclosed pasture and protected from birds. The position of the new set of pats was repeated throughout the 3 years of the study. Each year, the dung pats were weighed 4, 6, 8 and 10 weeks after deposition and immediately afterwards replaced to their initial positions.

Results showed that there was no difference in faecal pat disintegration between groups. However, the time-lag between deposition and complete disintegration of the faeces varied significantly between deposition occasions. Dung pats disappeared within 2 weeks (visual observation) when subjected to heavy rainfall early after deposition, whereas an extended dry period coincided with faeces still remaining 12 months after deposition.