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.     

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.     

The role of habitat and daily activity patterns in explaining the diversity of mountain Neotropical dung beetle assemblages

The interaction between land use and climate change is expected to strongly affect species distributions along high elevation landscapes. We aimed to test the effect of climatic variables on community metrics among five types of land use in a high elevation landscape. We described dung beetle spatial and temporal taxonomic and functional diversity patterns, and partitioned β‐diversity into turnover and nestedness components. The interaction between land use and daily period of activity mostly drives abundance, functional richness and functional diversity, but not dung beetle species richness. Unlike Neotropical lowlands, species richness and abundance in open environments are similar to those existing in forests. Temperature is an important predictor of abundance and functional divergence. There is a higher spatial component of the taxonomic β‐diversity, which is highly driven by species turnover. The temporal component of the taxonomic β‐diversity was strongly driven by nestedness, where night assemblages are sub‐sets, although not entirely, of diurnal assemblages. For functional diversity, the temporal β‐diversity was much higher than the spatial β‐diversity, but both were similarly represented by functional group turnover and nestedness. The composition of nocturnal and diurnal assemblages is clearly different, even more than the differences observed between habitats. However, taxonomic turnover is the dominant force between sampling sites while nestedness dominates the daily pattern. This means that forest habitats are unlikely to act as shelters for grassland species under a scenario of rising temperature.     

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.     

Analysis of trophic structure of two carnivore assemblages by means of guild identification

We evaluated the existence of trophic guild structure, considering seasonal and annual variation, in two terrestrial carnivore assemblages: one from Santa Cruz province (Argentinean Patagonia, composed by six carnivore species), and the other from Doñana National Park (SW Spain, composed by five carnivore species). To identify trophic guilds, we first studied seasonal and annual diets of predators, calculated trophic overlap among species pairs, and then constructed overlap matrices (similarity matrices). We determined guild membership objectively by entering the similarity matrices into the clustering technique unweighted pair-group method with arithmetic averaging. Carnivores from both assemblages were grouped, respectively, into four feeding guilds. Lagomorphs and rodents promoted the formation of two feeding guilds in both study sites, although the taxonomic composition of predator species that composed them was different. The ungulates-edentates feeding guild was only present at Santa Cruz, whereas the birds and reptiles feeding guild was only present at Doñana. Invertebrates and fruits were the base for the formation of a guild composed by species of the same taxonomic origin both in Santa Cruz and Doñana. Guild structure of Santa Cruz and Doñana assemblages did not exhibit seasonal or annual variation, although the specific guild composition changed over the two studied periods for both assemblages. This structure probably responded to discontinuities in resource spectra in Santa Cruz and fluctuations in rabbit abundance in Doñana. Our results support the hypothesis that establishes that guilds are originated by opportunistic convergence of species on abundant and energetically rewarding resources.     

Population genetics after fragmentation: the case of the endangered Spanish imperial eagle (Aquila adalberti)

The highly endangered Spanish imperial eagle, Aquila adalberti, has suffered from both population decline and fragmentation during the last century. Here we describe the current genetic status of the population using an extensive sampling of its current distribution range and both mitochondrial control region sequences and nuclear microsatellite markers. Results were evaluated in comparison to those obtained for the Eastern imperial eagle, Aquila heliaca, its nearest extant relative. Mitochondrial haplotype diversity was lower in the Spanish than in the Eastern species whereas microsatellite allelic richness and expected heterozygosity did not differ. Both allelic richness and expected heterozygosity were lower in the small Parque Nacional de Doñana breeding nucleus compared to the remaining nuclei. A signal for a recent genetic bottleneck was not detected in the current Spanish imperial eagle population. We obtained low but significant pairwise F_ST values that were congruent with a model of isolation by distance. F_ST and exact tests showed differentiation among the peripheral and small Parque Nacional de Doñana population and the remaining breeding subgroups. The centrally located Montes de Toledo population did not differ from the surrounding Centro, Extremadura and Sierra Morena populations whereas the latter were significantly differentiated. On the other hand, a Bayesian approach identified two groups, Parque Nacional de Doñana and the rest of breeding nuclei. Recent migration rates into and from Parque Nacional de Doñana and the rest of breeding nuclei were detected by assignment methods and estimated as 2.4 and 5.7 individuals per generation, respectively, by a Bayesian approach. We discuss how management strategies should aim at the maintenance of current genetic variability levels and the avoidance of inbreeding depression through the connection of the different nuclei.     

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.     

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.

     

Forest complexity drives dung beetle assemblages along an edge-interior gradient in the southwest Amazon rainforest

1. The habitat heterogeneity hypothesis predicts that heterogeneous habitats may provide more niches and diverse ways of exploiting environmental resources, thereby allowing more species to coexist, persist and diversify. 2. We aimed to investigate how an edge-interior gradient related to forest complexity influences species composition, abundance and richness of dung beetles in the western Amazon rainforest. We expected dung beetle abundance and richness to increase along the forest edge-interior gradient, in accordance with the habitat heterogeneity hypothesis. We also expected strong changes in species composition driven by species turnover in the forest interior and nestedness along the forest edges. We sampled dung beetles using baited pitfall traps across an edge-interior gradient. We also assessed the variation in forest features along the edge-interior gradient to identify changes in forest complexity. 3. Both species richness and abundance of dung beetles increased along the forest edge-interior, following the gradient of forest complexity. The Sorensen dissimilarity of dung beetle assemblages was higher among sampling units placed near the forest edge, although neither turnover, nor nestedness was different between the extremes of the forest edge-interior gradient. There was a clear compositional change along the edge-interior gradient mostly driven by species turnover. Individual indicator value analysis revealed that species were strongly associated with the forest interior conditions. 4. The simplification of the Amazon rainforest near clearings causes compositional changes in dung beetle assemblages. These changes are characterised by species-poor and low-abundance assemblages and may impair dung beetle ecological functions and therefore forest recovery.     

Effects of forest fragmentation on European birds: implications of regional differences in species richness

Aim In this paper, we adopted a large‐scale approach to evaluate the effect of regional richness of forest birds on the number of bird species retained by forest fragments in several localities across Europe. Location We studied bird assemblages in fourteen forest archipelagos embedded in agricultural matrices from southern Norway to central Spain. Tree composition varied from oak and beech forests of the northern localities to oak and pine xerophitic woodlands of the southern ones. The number of fragments in each forest archipelago ranged from eighteen to 211. Methods We used the Gleason equation (s = a + z log A; where s and A are, respectively, the species richness and size of forest fragments and z the rate of species loss) to estimate the species richness for 1‐ and 15‐ha fragments in each archipelago. The regional richness of forest birds was estimated by modelling the geographical distribution of species richness in the European atlas of breeding birds. Results The latitudinal distribution of regional richness displayed a convex form, with the highest values being in central Europe. Along this gradient, the number of species retained by fragments and the rate of species loss was positively related to regional richness. In addition, the percentage of the regional pool of species sampled by fragments decreased in the southern localities. Main conclusions Relationships between regional richness of forest birds and richness in fragments seem to explain why fragments in central Europe shelter more species than their southern counterparts. The decreased ability of southern forest fragments to sample the regional richness of forest birds, could be explained as an effect of the low abundance of many species in the Mediterranean, which could depress their ability to prevent extinction in fragments by a rescue effect. Alternatively, high beta diversity in the Mediterranean could produce undersampling by fragments of the regional pool of species. These regional differences in the response of bird assemblages to forest fragmentation are used to discuss the usefulness of large‐scale, biogeographical approaches in the design of conservation guidelines.     

Species Richness and Distribution of Copepods and Cladocerans and their Relation to Hydroperiod and Other Environmental Variables in Doñana, South-west Spain

Twenty-five sites located in five wetland zones within Doñana, south-west Spain were studied for copepod and cladoceran species richness and composition in relation to habitat characteristics from January to March 2004. The γ-diversity of copepods and cladocerans combined varied between wetland zones, which differed significantly in conductivity, surface area and abundance of vegetation. However, there were no significant differences between zones in local species richness of either copepods or cladocerans or the two combined. Species richness was significantly higher in sites with intermediate hydroperiods (duration between 3 and 5 months) than in sites with shorter or longer hydroperiods. CCA analyses performed separately for copepods and cladocerans extracted conductivity, surface area, and vegetation cover as the main factors structuring species composition of both groups. Ours is one of the first studies of zooplankton in the natural marshes of the Doñana World Heritage Site, and we recorded five species new to the Doñana area.     

The maintenance of extensively exploited pastures within the Alpine mountain belt: implications for dung beetle conservation (Coleoptera: Scarabaeoidea)

The abandonment of marginal areas and the decline of traditional small-scale cattle-breeding is bringing about substantial changes in many areas of the Alpine mountain belt. Extensively exploited pastures, which were colonized by typical ecological communities, are now being replaced by shrublands and forests. In this paper, we evaluate the effectiveness of the maintenance of small pastures for dung beetle (Scarabaeoidea: Geotrupidae, Aphodiidae, Scarabaeidae) conservation in a protected area of the north-western Italian Alps. From July until September 2007, we studied the dung beetle coenosis in the main habitats of the mountain belt: beech forest, scots pine forest, mountain pine forest and pasture. Twenty-three species were sampled. We compared abundance, species richness, α-diversity, evenness and species turnover between the four habitat assemblages and performed a correspondence analysis based on dominance patterns of the three families. Two main ensembles were identified: a pasture ensemble, highly diverse and dominated by Scarabaeidae, and a closed habitats ensemble, less structured and dominated by Aphodiidae and Geotrupidae. Combining habitat specificity and fidelity, the IndVal method showed that half of the species collected made a strong habitat selection toward pasture, while only C. granarius (Aphodiidae) was a reliable indicator of woody areas. We therefore conclude that the loss of extensively exploited pastures may bring about fundamental changes in dung insect communities. In protected areas, a management policy intended to preserve the traditional, sustainable human activities within the mountain belt is highly recommended for the conservation of these insects.     

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.     

The impact of overgrazing on dung beetle diversity in the Italian Maritime Alps

Dung beetles (Coleoptera: Scarabaeidae) are undoubtedly the most typical and ecologically relevant insects of grazed alpine habitats because they provide valuable ecological services such as biological pest control and soil fertilization. Despite the great ecological contribution of these insects to pasture ecosystem functioning, little is known about their direct or indirect relationships with pastoral activities. The main aim of the study was to assess whether dung beetle diversity was influenced by different intensities of cattle grazing. Dung beetle communities of two adjacent alpine valleys within the Maritime Alps Natural Park (north-western Italian Alps), representing overgrazed and ungrazed pastures, were studied by pitfall trapping. A hierarchical design (three levels: valleys, transects, and replicates) was established for additive partitioning of γ-diversity and Indicator Species Analysis. Evenness and Shannon diversity were significantly higher at the ungrazed than at the overgrazed site because abundances were much more evenly distributed at the former than at the latter site (where one species was dominant over all the others). Dung beetle abundance and species richness of the overgrazed graminaceous pasture vegetation types were in most cases significantly lower than those of the ungrazed nongraminaceous vegetation type. In the additive partitioning of γ -diversity analysis relative to the whole study area, the randomization procedure indicated that the contribution of β to γ-diversity was significantly different from that expected by chance, suggesting that one or more environmental factors has intervened to change the partition of total diversity in the system considered. The analysis of the preferences and fidelity of species (Indicator Species Analysis) showed that only one species chose overgrazed pastures; all the others positively selected the ungrazed site, or the only ungrazed pasture vegetation type (Rumicetum alpini Beger) occurring at the overgrazed site. Results conformed to evidences that overgrazing represents a serious threat to the conservation of alpine dung beetles. To conserve local dung beetle assemblages, especially in protected areas, cattle overgrazing should be avoided. This does not mean, however, that pastoral activities are incompatible with biodiversity conservation. The contemporaneous presence of wild ungulates and low intensity extensive pastoral activities may be useful to preserve both local dung beetle assemblages and alpine pasture ecosystems.     

Relationship between land uses and diversity of dung beetles (Coleoptera: Scarabaeinae) in the southern Atlantic forest of Argentina: which are the key factors?

The loss of natural habitats is one of the main drivers of biodiversity decline. Anthropogenic land uses preserving biotic and abiotic conditions of the native ecosystem are more suitable to preserve the native biodiversity. In this study, we explored changes in species richness and composition in different land uses of the southern Atlantic forest, considering three independent factors: (1) canopy (presence–absence), (2) type of vegetation (native–exotic) and (3) livestock (presence–absence). We expected a gradient of response in the richness and composition of the native forest dung beetle community, from land uses preserving canopy and native vegetation to open land uses with exotic vegetation. Dung beetles were sampled in protected native forests and four land uses, using two potential food resources: human dung and carrion. The species richness and composition of each habitat, as well as differences in composition and the influence of factors over diversity, were then analyzed. As expected, our results showed that land uses preserving canopy and native vegetation maintain the dung beetle diversity of the native forest. Moreover, while the three factors analyzed influenced dung beetle diversity, canopy cover was the main driver of dung beetle diversity loss. The main conclusion of this study is that the conservation of canopy (either native or exotic) is determinant to preserve highly diverse dung beetle communities and subsequently, the ecological functions performed by this taxon. However, the ecophysiological mechanism behind the response of dung beetles to habitat disturbance is poorly understood.     

Differences in Coprophilous Beetle Communities Structure in Sierra de Minas (Uruguay): a Mosaic Landscape

Coprophilous beetles represent an abundant and rich group with critical importance in the functioning of terrestrial ecosystems. Most coprophagous beetles have a stenotopic distribution in relation to vegetation types. Because of this, they are usually very sensitive to environmental changes and are considered well suited as bioindicator organisms. The aim of this study was to analyze variations in coprophilous beetle assemblages in natural and anthropogenic habitats. Coprophilous beetle communities were sampled monthly for 1?year using pitfall traps baited with cow dung, in native xeric upland forests, 15-years-old plantations of Pinus elliottii and pastures in Sierra de Minas, Lavalleja, Uruguay. A total of 7,436 beetles were caught and identified to species or morphospecies level. The most abundant families were Aphodiidae, Scarabaeidae, and Staphylinidae. Differences in species richness, abundance, Shannon index, evenness, and dominance were detected between habitats. Abundances of most frequent families were significantly higher in both kinds of forests. Species richness and diversity of Aphodiidae and Staphylinidae were higher in forests, while Scarabaeidae showed the highest richness and diversity in pine plantations. Species composition significantly differed between habitats. Uroxys terminalis Waterhouse and Ataenius perforatus Harold typified the assemblages in native forests and pine plantations and also discriminated both communities because of their differential pattern of abundance between habitats. Typifying species in pastures were Onthophagus hirculus, Ateuchus robustus (Harold), and Ataenius platensis Blanchard. Habitat type had a strong effect on the coprophilous beetle community structure and composition.     

The influence of landscape structure on ants and dung beetles diversity in a Mediterranean savanna—Forest ecosystem

Ants and dung beetles are focal indicators of change in several ecological processes and successional vegetation stages in Mediterranean landscapes. Despite relatively good knowledge of local species distributions, there are few data on their distributions at different scales. In the present study, the influence of multiscale landscape structure was examined using both ants and dung beetles to identify species that can serve as indicators and detectors of changes in vegetation structure. Multiscale analysis is necessary to explore the different roles of indicator and detector species for use as tools in studies focused on monitoring ecological changes. The study area was the Cabañeros National Park, in the center of the Iberian Peninsula. This site was selected because it is a good setting to evaluate the effects of typical vegetation mosaics on Mediterranean species at different scales. In this study, dung beetles and ants were trapped for one year. A multiscale analysis was designed using three different vegetation habitats (forest, scrubland and grassland) and landscape matrices (woodland, scrubland and grassland). Among dung beetles, 23 indicator species (IndVal values higher than 70%) were found at different scales of analysis. In addition, 20 dung beetle species were characterized as detector species (IndVal values between 45 and 70%) for the three different levels analyzed. Similar to the dung beetles, the ants had different species assemblages at various habitats and landscape levels; however, no indicator ant species were found in this study. All species with significant IndVal values (n = 8) were identified as detector species. Thus, ant and dung beetle assemblages were influenced in different ways by vegetation structure. Both groups showed strong individual species responses to different Mediterranean landscape conditions and vegetation types. Further, both insect groups showed a great number of detector species, which can be useful in ecosystem management because they have varying degrees of preference and sensitivity for different ecological states (such as successional vegetation stages). The diverse indicator and detector species identified in this work could be useful tools for the detection of landscape structure changes at both levels habitat patches and landscape matrix. However, a generalisation of the results at landscape scale should be taken with precaution, but they encourage to study more regions and similar landscapes. The use of more than one indicator group in the analysis confirms the importance of selecting groups with different sensitivities at varying spatial scales as well as different ecosystem functions. This strategy allows the establishment of a clear baseline with which to analyze future direct and indirect impacts of management in Mediterranean protected areas.     

Roles of environmental variables and land usage as drivers of dung beetle assemblage structure in mopane woodland

Colophospermum mopane woodland covers large areas of dry lowland savanna in southeastern Africa. Dominant land usage is conservation (45%) with the remainder mostly modified by farming. Dung beetle responses to environment (dung type, habitat, weather) and land usage (conservation, farming, mining) were examined at Phalaborwa (23.9431°S 31.1411°E) in the Phalaborwa‐Timbavati Mopaneveld, South Africa. Partitioning of gamma species richness and diversity showed lower alpha values in mine areas than in farm and conserved areas. However, between‐land usage differences in species richness, alpha diversity, abundance and biomass, showed lower significance than those between dung type and different weather. At two sampling scales, three multivariate techniques variously separated assemblages according to land usage, dung type and weather. Analysis of 21 mean samples separated clusters according to dung type (Canonical Correspondence Analysis, CCA) or mine assemblages, conserved plus farm assemblages on pig plus elephant, or cattle dung (NMDS, Factor Analysis) with shared variance of >80% and unique variance of 16–18% per cluster. In analysis of 188 samples (CCA), each overlapping dung type cluster was offset in ordinal space with congruent patterns of separation according to land usage and weather (drier days distant from moister days; conserved plus farm areas distant from early succession mine areas, which were distant from disturbed and later succession mine areas). Mining, dung types, and moist conditions were the strongest contributors to between‐assemblage differences. Compared with conserved areas, dung beetle diversity is appreciably altered by mining but only slightly altered by intensive game farming or livestock ranching with subsistence agriculture.     

Spatio-temporal nested patterns in macroinvertebrate assemblages across a pond network with a wide hydroperiod range

Nestedness has been widely used to measure the structure of biological communities and occurs when species-poor sites contain subsets of species-rich ones. Here, we examine nested patterns across the macroinvertebrate assemblages of 91 ponds in Doñana National Park, Spain, and explore temporal variation of nestedness and species richness in 19 temporary ponds over 2 years with differing rainfall. Macroinvertebrate assemblages were significantly nested; both pond spatial arrangement and environmental variation being important in driving nested patterns. Despite the nested structure observed, a number of taxa and ponds deviate from this pattern (termed idiosyncratic), by occurring more frequently than expected in species-poor sites, or having assemblages dominated by species largely absent from species-rich sites. Aquatic adults of winged insects, capable of dispersal, were more highly nested than non-dispersing taxa and life-history stages. Idiosyncratic taxa were found in ponds spanning a wide range of hydroperiods, although nestedness was higher in more permanent waterbodies. Monthly sampling demonstrated a gradual increase of species richness and nestedness from pond filling to April–May, when the most temporary ponds started to dry. Although the degree of nestedness of individual pond assemblages varied from month to month, the overall degree of nestedness in the two study years was practically identical despite marked differences in hydroperiod. Our results suggest that differential colonization and environmental variation are key processes driving the nested structure of Doñana ponds, that macroinvertebrate assemblages change in a predictable manner each year in response to cycles of pond wetting and drying, and that connectivity and environmental variability maintain biodiversity in pond networks.     

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.