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.     

Patterns of alimentary resource use by dung beetles in introduced Brazilian pastures: Cattle versus sheep dung

Dung beetles (Coleoptera: Scarabaeidae) perform essential ecological functions in pastures, such as dung removal, nutrient recycling and parasite control. However, the patterns of alimentary use by dung beetles in introduced Brazilian pastures are poorly known. Here, we compared dung beetle species richness, abundance and species composition in cattle and sheep dung, and identified the dung beetle species preference by each dung type. In January 2019, dung beetles were sampled with pitfall traps baited with cattle and sheep dung in 12 introduced pastures (Urochloa spp.), in Aquidauana, Mato Grosso do Sul, Brazil. A total of 592 individuals belonging to 14 species of dung beetles were collected. Of the 14 species sampled, nine were recorded in both dung types, five were found exclusively in sheep dung and no species was exclusive to cattle dung. Species richness and abundance were higher in sheep dung. Species composition was different between the dung types. Dichotomius bos (Blanchard), Genieridium bidens (Balthasar), Onthophagus aeneus Blanchard and Trichillum externepunctatum Preudhomme de Borre were associated with sheep dung. Our results provide evidence that sheep dung is more attractive to dung beetles with a distinct community species between the two dung types, although the studied pastures have never been used before for sheep breeding. Thus, our data shows that the introduction of a new alimentary resource (e.g. sheep dung) can be an important strategy to help to obtain a more diverse dung beetle assemblage in introduced Brazilian pastures.     

Diversity, distinctiveness and conservation status of the Mediterranean coastal dung beetle assemblage in the Regional Natural Park of the Camargue (France)

The Mediterranean region as a whole has the highest dung beetle species richness within Europe. Natural coastal habitats in this region are among those which have suffered severe human disturbance. We studied dung beetle diversity and distinctiveness within one of the most important coastal protected areas in the west Euro‐Mediterranean region (the regional Park of Camargue, southern France) and made comparisons of dung beetle assemblages with other nearby Mediterranean localities, as well as with other coastal protected area (Doñana National Park, Spain). Our finding showed that: (1) The species richness of coastal habitats in the Camargue is low and only grasslands showed a similar level of species richness and abundance to inland habitats of other Mediterranean localities. The unique habitats of the coastal area (beaches, dunes and marshes) are largely colonized by species widely distributed in the hinterland. (2) In spite of their low general distinctiveness, dune and marsh edges are characterized by the occurrence of two rare, vulnerable, specialized and large roller dung beetle species of the genus Scarabaeus. As with other Mediterranean localities, current findings suggest a recent decline of Scarabaeus populations and the general loss of coastal dung beetle communities in Camargue. (3) The comparison of dung beetle assemblages between the Camargue and Doñana shows that, in spite of the low local dung beetle species richness in the Camargue, the regional dung beetle diversity is similar between both protected areas. Unique historical and geographical factors can explain the convergence in regional diversity as well as the striking divergence in the composition of dung beetle assemblages between both territories.     

Altitudinal variation of dung beetle (Scarabaeidae: Scarabaeinae) assemblages in the Colombian Andes

Aim We describe the changes in species richness, rarity and composition with altitude, and explore whether the differences in Scarabaeinae dung beetle composition along five altitudinal transects of the same mountain range are related to altitude or if there are interregional differences in these altitudinal gradients. Location Field work was carried out on the eastern slope of the eastern Cordillera, Colombian Andes, between Tamá Peak to the north, in the Tamá National Park (07°23′ N, 72°23′ W) and the San Miguel River (00°28′ N, 77°17′ W) to the south. Methods Sampling was carried out between February 1997 and November 1999 in five regions spanning elevation gradients. In each gradient, six sites were chosen at 250 m intervals between 1000 and 2250 m a.s.l. Results We found a curvilinear relationship between altitude and mean species richness, with a peak in richness at middle elevations. However, the diversity of dung beetle assemblages does not seem to be related to the interregional differences in environmental conditions. The number of geographically restricted species is negatively and significantly related to altitude, with geographically restricted species more frequent at low altitude sites. Ordination delimited the two main groups according to altitude: one with all the highest sites (1750–2250 m a.s.l.) and a second group with the remaining sites (< 1750 m a.s.l.). Analysis of species co‐occurrence shows that these dung beetle assemblages seem to be spatially structured when all sites have the same probability of being chosen. In contrast, the spatial structure of species assemblages seems to be random when the probability of choosing any site is proportional to its altitude. Main conclusions The altitude of sites is the main factor that influences the diversity of these dung beetle assemblages. The peak in species richness at middle elevations, the higher number of geographically restricted species at lower altitudinal levels, and the compositional differences along these mountain gradients seem to result from the mixing at these altitudes of dung beetle assemblages that have different environmental adaptations and, probably, different origins. The relevance of altitude in these assemblages is related to the limited role of these Neotropical high altitude environments as centres of refuge and vicariance for a monophyletic group of warm‐adapted species, for which the vertical colonization of these high mountain environments by lineages distributed at lower altitudes would have been very difficult.     

Monitoring the ecosystem service provided by dung beetles offers benefits over commonly used biodiversity metrics and a traditional trapping method

Surveying terrestrial invertebrates often requires lethal techniques that can also kill non-target vertebrates. Removing the desirable components of biodiversity is at odds with the philosophy of ecological restoration and biodiversity conservation more generally. Moreover, commonly used metrics generated by such survey approaches (e.g. abundance and species richness) are only indirectly related to the ecosystem services (e.g. pollination) that are often of primary interest. We examined the relationship between rates of dung removal (a direct measure of an ecosystem service) and dung beetle abundance and species richness in a temperate region of New South Wales, Australia, and examined if dung removal in revegetated riparian areas of different ages were trending monotonically toward rates in areas with mature native vegetation. Pellets of pig manure and conventional traps were left at study sites for 48 h to examine the relationships between rates of dung removal and dung beetle abundance and richness. Regressions of abundance and richness with average percent dung removal were positive and significant, demonstrating the potential of the method as a non-lethal proxy. While the dung removal method cannot determine the species responsible, percentage dung removal was more time-efficient, costing 4 min per sample, while abundance and richness cost 13 and 17 min, respectively. Despite variability among replicates of the same habitat type, the trajectory across the restoration gradient showed an increase from sites recently revegetated toward those with mature woody vegetation. We interpreted these results as a positive response of dung beetle activity and an indication of recovery of this ecosystem service. We argue that responses that can be collected efficiently such as dung removal should be used if restorationists have limited resources for data collection and analysis; non-specialists are involved; knowledge of ecosystem function is required, and animal ethics constrain options.     

Changes of dung beetle communities from rainforests towards agroforestry systems and annual cultures in Sulawesi (Indonesia)

Little is known about how tropical land-use systems contribute to the conservation of functionally important insect groups, including dung beetles. In a study at the margin of Lore Lindu National Park (a biodiversity hotspot in Central Sulawesi, Indonesia) dung-beetle communities were sampled in natural forest, young secondary forest, agroforestry systems (cacao plantations with shade trees) and annual cultures (maize fields), each with four replicates (n = 16 sites). At each site we used 10 pitfall traps, baited with cattle dung, along a 100 m transect for six 3-day periods. The number of trapped specimens and species richness at the natural forest sites was higher than in all land-use systems, which did not significantly differ. Each land-use system contained, on average, 75% of the species richness of the natural forest, thereby indicating their importance for conservation. However, a two-dimensional scaling plot based on NESS indices (m = 6) indicated distinct dung beetle communities for both forest types, while agroforestry systems and annual cultures exhibited a pronounced overlap. Mean body size of dung beetles was not significantly influenced by land-use intensity. Five of the six most abundant dung beetle species were recorded in all habitats, whereas the abundance of five other species was significantly related to habitat type. Mean local abundance and number of occupied sites were closely correlated, further indicating little habitat specialisation. The low dung beetle diversity (total of 18 recorded species) may be due to the absence of larger mammals in Sulawesi during historical times, even though Sulawesi is the largest island of Wallacea. In conclusion, the dung beetle fauna of the lower montane forest zone in Central Sulawesi appears to be relatively robust to man-made habitat changes and the majority of species did not exhibit strong habitat preferences.     

Effects of Degradation and Replacement of Southern Brazilian Coastal Sandy Vegetation on the Dung Beetles (Coleoptera: Scarabaeidae)

In this paper we address the effects of anthropogenic disturbance and replacement of Brazilian Coastal sandy vegetation (restingas) on dung beetles communities. We sampled dung beetles in the four main vegetative physiognomies of Guriri Island, Espírito Santo State: forest restinga, restinga Clusia, disturbed restinga (from burning events), and pastures. We placed four sets of two pitfall traps (baited with horse and human dung) in four independent areas of each vegetation type, and collected 14,534 individuals of 13 dung beetle species. Neither log₁₀ of individuals nor log₁₀ of species richness were good predictors of restinga disturbance. However, a significant amount of variation in dung beetle abundance and richness could be explained by bait type. Ordination of these sites using hybrid multidimensional scaling revealed a gradient of habitat disturbance from undisturbed restinga samples to pasture. Dung beetle communities along this gradient demonstrated a complete turnover in species composition, from restinga‐specialists to invasive and generalists species respectively. This complete turnover signals the local extirpation of forest‐adapted species in disturbed and converted areas. Only a single dung beetle species in preserved restingas is protected by Brazilian law (Dichotomius schiffleri). Given the extent of the clearing of restinga habitat, the conservation status of dung beetles associated with restinga forest gives cause for concern.     

Space-for-time substitution reveals a hump-shaped distribution of dung beetles

Unraveling how climate change impacts the diversity and distribution patterns of organisms is a major concern in ecology, especially with climate-sensitive species, such as dung beetles. Often found in warmer weather conditions, beetles are used as bio-indicators of environmental conditions. By using an altitudinal gradient as a proxy for climate change (i.e., space-for-time substitution), we assessed how changes in climatic variables, such as temperature and precipitation, impact patterns of dung beetle diversity and distribution in the Peruvian Andes. We recorded dung beetle diversity using three different types of baits, feces, carrion, and fruits, distributed in 18 pitfall traps in five different altitudinal sites (from 900 to 2500 m, 400 m apart from each other) in the rainy and dry season. We found that (i) dung beetle richness and abundance were influenced by the climate gradient, (ii) seasonality influenced beetle richness, which was high in the wet season, but did not influence abundance, (iii) dung beetle richness and abundance fit to a hump-shaped distribution pattern along the altitudinal gradient, and (iv) species richness is the beta-diversity component that best describes the composition of dung beetle species along the altitudinal gradient. Our data show that the distribution and diversity of dung beetles are different at larger scales, with different patterns resulting from the response of species to both abiotic and biotic factors.     

Dung beetle assemblages on tropical land‐bridge islands: small island effect and vulnerable species

Aim Using dung beetles (Coleoptera: Scarabaeidae: Scarabaeinae) in a tropical land‐bridge island system, we test for the small island effect (SIE) in the species–area relationship and evaluate its effects on species richness and community composition. We also examine the determinants of species richness across island size and investigate the traits of dung beetle species in relation to their local extinction vulnerability following forest fragmentation. Location Lake Kenyir, a hydroelectric reservoir in north‐eastern Peninsular Malaysia. Methods We sampled dung beetles using human dung baited pitfall traps on 24 land‐bridge islands and three mainland sites. We used regression tree analyses to test for the SIE, as well as species traits related to local rarity, as an indication of extinction vulnerability. We employed generalized linear models (GLMs) to examine determinants for species richness at different scales and compared the results with those from conventional linear and breakpoint regressions. Community analyses included non‐metric multidimensional scaling, partial Mantel tests, nestedness analysis and abundance spectra. Results Regression tree analysis revealed an area threshold at 35.8 ha indicating an SIE. Tree basal area was the most important predictor of species richness on small islands (<35.8 ha). Results from GLMs supported these findings, with isolation and edge index also being important for small islands. The SIE also manifested in patterns of dung beetle community composition where communities on small islands (<35.8 ha) departed from those on the mainland and larger islands, and were highly variable with no significant nestedness, probably as a result of unexpected species occurrences on several small islands. The communities exhibited a low degree of spatial autocorrelation, suggesting that dispersal limitation plays a part in structuring dung beetle assemblages. Species with lower baseline density and an inability to forage on the forest edge were found to be rarer among sites and hence more prone to local extinction. Main conclusions We highlight the stochastic nature of dung beetle community composition on small islands and argue that this results in reduced ecosystem functionality. A better understanding of the minimum fragment size required for retaining functional ecological communities will be important for effective conservation management and the maintenance of tropical forest ecosystem stability.     

Asymmetrical competition between Neotropical dung beetles and its consequences for assemblage structure

Abstract.  1. This study combines the results of laboratory experiments using representative assemblage components and pitfall trapping over a large geographical area to examine the hypothesis that ongoing interspecific competition structures Neotropical dung beetle assemblages.
2. From Guatemala to Panama assemblages of large to medium-sized, fast-tunnelling dung beetles include a single large, nocturnal dichotomiine species, Dichotomius annae (Kohlmann & Solís, 1997). In competition experiments, this species out-competed the medium-sized coprine species, Copris lugubris Boheman and Phanaeus demon Laporte-Castelnau, for dung and nesting space, in spite of earlier colonisation by the diurnal species, P. demon .
3. Differences in the abundance of D. annae at Central American sites did not affect total fast-tunnelling dung beetle assemblage richness over the rainy season. However, D. annae rank order was directly related to the probability of interspecific encounters (Hurlbert's Δ ₁) among species. These trends were also observed when species lists from published and unpublished studies of other large allopatric dichotomiine species, with a more northerly distribution, were included in the analyses.
4. The results obtained suggest that where large dichotomiine species are abundant, their efficient pre-emption of a considerable proportion of available resources drives all, or most, other fast-tunnelling species to a lower population density, thereby decreasing assemblage diversity.     

Wide corridors with much environmental heterogeneity best conserve high dung beetle and ant diversity

Landscape ecological networks (ENs) consist of landscape-scale conservation corridors that connect areas of high natural value within a production mosaic with protected areas (PAs). In South Africa, ENs have been implemented on a large spatial scale to offset the negative impacts of plantation forestry on indigenous grasslands. We focus on corridor width as a factor for conserving dung beetle and ant diversity within an EN. We also investigate the importance of natural environmental heterogeneity (elevation, vegetation type) and habitat quality (soil hardness, invasive alien plant density). We sampled dung beetles and ants in 30 corridors of different sizes, and at ten sites in a nearby PA. In addition, we also analysed dung beetles according to their feeding guild. Tunnelling dung beetle species richness increased with corridor width. Rolling dung beetle species richness was higher in the PA than in the corridors of the EN. The dung beetle assemblage within the EN differed from that within the PA. Corridors of various widths differed in ant composition but not in species richness. Furthermore, the PA and the EN differed in environmental variables, which contributed to differences in dung beetle species richness and assemblage composition. Within the EN, environmental heterogeneity across the landscape was more important than corridor width for driving species diversity of both dung beetles and ants. When planning future ENs, wide corridors (>280 m) that encompass as much natural heterogeneity across the landscape as possible will best conserve the range of local insect species.     

Does an aridity and trophic resource gradient drive patterns of dung beetle food selection across the Botswana Kalahari?

1. Aridity gradients are paralleled by both reductions in resources and decreased species richness of animals. Across the aridity gradient of the Botswana Kalahari, a reduction in mammal species richness leads to reduced density and diversity of dung types, accompanied by reduced dung beetle species richness. We investigated whether this gradient also drives changes in dung beetle food type association and specialisation owing to a loss of some dung types to the arid southwest. 2. Dung beetles were sampled from three study sites in each of the six study areas using 2 × 10 grids of pitfall traps baited with dung (pig, elephant, cattle, and sheep) or carrion (chicken livers). 3. Canonical correspondence analysis (CCA) showed that distributions of dung beetle species between bait types deviated significantly from random associations. 4. Central Kalahari assemblages were more specialist than those at the mesic and arid extremes of the gradient. 5. Patterns of selection and specialisation to bait types differed between mesic northeast and arid southwest study areas. There were specialist faunas on carrion and more generalist faunas on ruminant herbivore dung (cattle and sheep) in each region. However, specialist species associated with elephant dung in the northeast were replaced by a more generalist fauna in the southwest with an opposite trend on pig dung. 6. Reduced species richness and high species turnover from the mesic northeast to the arid southwest is paralleled by a shift in patterns of food association that may reflect changes in the diversity of food types, particularly the absence of elephant dung from the southwest.     

Effects of dung-pad conditions and density on coprophagous beetle assemblages in a Mediterranean rangeland

Dung beetles highly depend on the ephemeral microhabitat dung which is food resource and larval habitat at the same time. Environmental conditions surrounding a dung pad, such as vegetation structure, have an impact on dung beetle assemblages. We investigated the influence of dung conditions and surrounding habitat characteristics on Mediterranean dung beetle assemblages in a permanently grazed landscape in northern Sardinia. We sampled the dung beetle assemblages of donkey and horse dung in three different vegetation types and assessed species richness and abundance of dung beetles. Species richness was determined by dung and surrounding habitat conditions, whereas abundance was solely affected by dung conditions. However, species richness and abundance decreased with increasing dung density. The effect of dung density on species richness varied depending on vegetation type, with dry grassland exhibiting the highest number of dung beetles species at high dung density. Species composition in dung pads was influenced by abiotic factors with dwellers being negatively affected by increasing dung-pad temperature. Our results underline the importance of diverse vegetation, particularly with respect to the complexity of vegetation which interrelates with the microclimate. Furthermore, our findings illustrate the negative effect of high dung densities on dung beetle assemblages, suggesting that the degree of the intensity of use by grazing animals is important when considering measures for the conservation of dung beetles.     

Response of vertebrates to fenceline contrasts in grazing intensity in semi‐arid woodlands of eastern Australia

Abstract Changes in the abundance, species richness and assemblage composition of vertebrates due to grazing by domestic stock were investigated in the semi‐arid woodlands of eastern Australia. Analyses were based on the differences found at 10 fenceline contrast sites. Two species of amphibians, 22 species of reptiles and two species of small mammal were captured in pit traps during the surveys. Kangaroos (red and eastern grey), sheep, goats and 66 species of birds were recorded along line transects. Analyses revealed that abundance of diurnal reptiles and species richness of diurnal reptiles and birds were significantly lower on heavily grazed sites than they were on lightly grazed sites. At a local scale, the gecko, Gehyra variegata, was more abundant where grazing was heavier, while Diplodactylus conspicillatus, Diplodactylus steindachneri and Rhynchoedura ornata responded to variables indirectly related to grazing intensity (kangaroo density, sheep and goat dung mass and sheep density, respectively). Birds more commonly sighted on lightly grazed areas than heavily grazed areas were the apostlebird, brown treecreeper, crested bellbird, grey butcherbird, hooded robin, jacky winter, little woodswallow, Australian magpie‐lark, mulga parrot, splendid wren, white‐browed treecreeper and yellow‐rumped thornbill. Birds more commonly sighted on heavily grazed areas than on lightly grazed areas were the Australian raven and chestnut‐crowned babbler. Most variation in species composition between sites was due to spatial separation and no regional‐level indicator species of grazing were evident. A combination of direct grazing‐related changes (e.g. loss of ground cover) and indirect effects of the pastoral industry (e.g. introduction of artificial sources of water) lead to changes in fauna at different scales of analysis across regions.     

Regional and environmental effects on the species richness of mammal assemblages

Aim Variation in species richness has been related to (1) environmental conditions (water, energy and habitat characteristics) and (2) regional differences (contingent historical events and regional particularities that result in differences between regional faunas acting at broad extents). Whereas climatic factors have been widely studied, the effects of regional differences are less often quantified. This work aims to characterize global trends in the species richness of mammal assemblages with respect to both current and historical influences. Location All terrestrial biogeographical realms except Antarctica. Methods Species richness in checklists from 224 sites distributed worldwide were investigated by partitioning the variation between a general set of habitat/climate factors, biogeographical regions, and their overlaps. Additional analyses studied the specific overlaps of region, water and energy. Data were also divided according to area to determine if the strength of these effects varies according to the size of sites. Results Environmental effects explained 38% of richness variation across all sites, whereas environmentally independent regional effects explained 11% and the overlap between region and environment explained 13%. Results were similar when only larger sites (between 1000 km² and 10,000 km²) were considered. However, the importance of the overlap between region and all environmental variables was greater in smaller sites (between 100 km² and 1000 km²). In contrast, the specific importance of water and energy variables and their overlap with region was greater in larger sites. The strength of the independent effect of region remained almost invariant regardless of the size of the sites studied. Main conclusions The relationship between species richness and climate varies with scale and among regions. Although environmental variables are the strongest correlates of richness, the unique history and physiographic characteristics of a region produce differences between the richness of mammal assemblages and their response to environmental gradients. The importance of environmental variables varies with scale: climatic gradients are more important at coarse grain (larger sites), possibly as a result of their effects on species ranges, whereas habitat type is more important at the smaller sites, where the importance of ecological interactions increases. Therefore, regional differences and the scale at which richness is measured should be taken into account when evaluating species richness–energy hypotheses.     

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.     

An intercontinental comparison of dung beetle diversity between two mediterranean-climatic regions: local versus regional and historical influences

The species richness of biological communities is influenced by both local ecological, regional ecological, and historical factors. The relative importance of these factors may be deduced by comparison between communities in climatically and ecologically equivalent, but geographically and historically separate regions of the world. This claim is based on the hypothesis that community processes driven by similar local ecological factors lead to convergence in species richness whereas those driven by differing regional or historical factors lead to divergence. An intercontinental comparison between the winter rainfall regions of South Africa and the Iberian Peninsula showed that overall species richness of dung beetles was dissimilar at local, subregional and regional scales in Scarabaeidae s. str. but similar at all scales in Aphodiinae. Removal of species widespread in the summer rainfall region of Africa or the temperate region of Europe (regional component) resulted in dissimilarity in species richness of mediterranean endemics at all scales in both dung beetle taxa. However, the lines joining each set of species richness values were parallel which may indicate similarities in processes between different mediterranean climatic regions despite slight differences in latitudinal range. The dominant pattern of dissimilarity or non-convergence may be related primarily to intercontinental differences in regional biogeographical and evolutionary history (faunal dispersal, glaciation effects in relation to geographical barriers to dispersal, speciation history, long-term disturbance history). The limited pattern of similarity or convergence in overall species richness of Aphodiinae may be a chance result or primarily related to intercontinental similarities in local ecological factors.     

Multi-scale determinants of dung beetle assemblage structure across abiotic gradients of the Kalahari–Nama Karoo ecotone, South Africa

Aim Changing conditions across spatial gradients are primary determinants of biotic regions, local habitats, and distributional edges. We investigate how a climatic gradient and edaphic mosaic interact as multi‐scale drivers of spatial patterns in scarabaeine dung beetles. The patterns are tested for congruency with ecoregion and floral boundaries over the same gradient, as responses to physical factors often differ among higher taxa. Location Southern Africa and the Nama Karoo–Kalahari ecotone, Northern Cape, South Africa. Methods Data consisted of the climatic distributions of 104 species and their abundances at 223 sites in two ecoregions/floral biomes, four bioregions, and 13 vegetation units. Factor analyses determined the biogeographical composition of the species, and regional‐ to local‐scale patterns in species abundance structure. Hierarchical analysis of oblique factors determined the proportional contribution of spatial variance to patterns. One‐way anova was used to test for significant separation of patterns along factor axes. Stepwise multiple regression was used to determine correlations of five physical attributes with species richness, Shannon‐Wiener diversity, and factor loadings for the study sites. Results Four biogeographical influences overlap in the study region, although rank contribution declines from south‐west arid through north‐east savanna to widespread and south‐east highland taxa. Species abundance structure comprises five subregional patterns, two centred to the north‐east (Kalahari, Isolated Kalahari Dune) dominated by Kalahari influence, and three to the south‐west (Nama Karoo subdivisions: Bushmanland, ‘Upper’, ‘Stony Prieska’) dominated by south‐west arid influence. Kalahari deep sands are characterized especially by a warmer, moister climate, whereas the Nama Karoo mosaic of deep or stony soils is characterized especially by north‐west aridity (Bushmanland), south‐east cooler temperatures (‘Upper’), or excessively stony soils (‘Stony Prieska’). Four of the subregional patterns each comprised three localized patterns related primarily to relative stoniness, edge effects from geographical position, or incidence of rainfall. Species richness and diversity declined with decreasing rainfall and increasing stoniness. Main conclusions Climatic and edaphic factors are important multi‐scale determinants of spatial patterns in dung beetle assemblage structure, with edaphic factors becoming more important at local spatial scales. The patterns are roughly congruent with the Kalahari Savanna–Nama Karoo ecotone at the floral biome or ecoregion scale, but show limited coincidence with finer‐scale floral classification.     

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.     

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.