From forest to pasture: an evaluation of the influence of environment and biogeography on the structure of beetle (Scarabaeinae) assemblages along three altitudinal gradients in the Neotropical region

The objective of this study is to evaluate the effect of environmental (associated with the expansion of cattle ranching) and biogeographical factors on the diversity of dung beetle (Scarabaeinae) assemblages along three altitudinal gradients in the Neotropical region. One gradient is located in the Mexican Transition Zone, on the Cofre de Perote mountain, the other two are in the northern Andes (the Chiles Volcano and the Río Cusiana Basin). For the three gradients, the number of species and of individuals was similar in both forest and pasture, while species composition was different between habitats. On this mountain, species turnover in pastures was characterized by the addition of new species as altitude increased. In the northern Andes, species diversity was always greater in the forest than in the pasture, and species turnover between habitats was notably influenced by species loss with increasing altitude. As such the pasture fauna of the northern Andes was an impoverished derivative of the fauna present in the forests at the same altitude characterized by species of Neotropical affinity with a limited capacity for colonizing open, sunnier habitats. The opposite occurs in the areas used by cattle on the Cofre de Perote. This habitat has its own fauna, which is mainly comprised of Holarctic and Afrotropical species adapted to the prevailing environmental conditions of areas lacking arboreal vegetation. These results suggest that the impact on beetle communities caused by human activities can differ depending on the geographic position of each mountain and, particularly, the biogeographical history of the species assemblage that lives there.     

Diversity and biogeographical makeup of the dung beetle communities inhabiting two mountains in the Mexican Transition Zone

The role of horizontal and vertical colonization on the diversity and integration of the dung beetle fauna of two mountains in the Mexican Transition Zone (Los Tuxtlas and La Chinantla) are analyzed and compared. On each mountain standardized sampling was done using pitfall traps baited with dung and carrion along elevation gradients. On both mountains diversity decreased linearly with increasing elevation. The decrease in the number of genera and species was not different between mountains, but the cumulative total number for both taxonomic levels was significantly higher on La Chinantla. There, three well-defined groups were identified for which species turnover was mainly a result of species gain. On Los Tuxtlas there was no evident grouping pattern, and species turnover was determined by species loss. For both mountains the dominant biogeographic distribution pattern was Neotropical; however, at the higher elevations of La Chinantla, a clear replacement by lineages of Holarctic affinity was observed. We suggest that for La Chinantla—a mountain that is geographically connected to the Sierra Madre Oriental mountain range and is of ancient geological origin—the processes of horizontal and vertical colonization have had relatively different weights in terms of their effect on the pattern of diversity and the biogeographic integration of the beetle community, while on Los Tuxtlas, the limited role of horizontal colonization appears to be a consequence of its isolation and more recent geological origin. We discuss the potential use of these models for studying the effects of climate change on elevation gradients.     

Does the Trans‐mexican Volcanic Belt represent a natural biogeographical unit? An analysis of the distributional patterns of Coleoptera

Aim We analysed the geographical distribution of beetle species of the families Buprestidae, Cerambycidae, Dryophthoridae, Melolonthidae, Passalidae and Staphylinidae from the Trans‐mexican Volcanic Belt (TVB) through a track analysis and a parsimony analysis of endemicity (PAE), in order to test its naturalness and determine its affinities. Location The area analysed corresponds to the TVB, which is a biogeographical province of the Mexican Transition Zone. Methods The panbiogeographical analysis was based on the comparison of the individual tracks of 299 species of Buprestidae, Cerambycidae, Dryophthoridae, Melolonthidae, Passalidae and Staphylinidae (Coleoptera). The TVB was divided into 1^o × 1^o grid cells and we also included in the analysis the remaining Mexican biogeographical provinces. Parsimony analysis of endemicity with progressive character elimination (PAE‐PCE) was applied to classify areas by their shared taxa according to the most parsimonious cladograms. The nested sets of areas were represented as generalized tracks. Results Three generalized tracks were obtained: (1) grid cells 9C, 9D, 10D, 10E, Sierra Madre Oriental, Chiapas, Mexican Gulf and the Sierra Madre del Sur; (2) grid cells 3B, 3C, 4B, 4C, 5C, 6C, 7C, Sierra Madre Occidental, Sierra Madre del Sur, Balsas Basin and the Mexican Pacific Coast, and (3) grid cells 8D, 9C, 9D, 10D, 10E, Yucatán Peninsula, Chiapas, Sierra Madre Oriental and the Mexican Gulf. Main conclusions We conclude that the TVB does not represent a natural biogeographical unit because it shows different relationships with other biogeographical provinces, being clearly transitional between the Nearctic and Neotropical provinces. Some parts of the TVB are related to Neotropical provinces (Chiapas, Mexican Gulf and Mexican Pacific Coast) and others to the remaining provinces of the Mexican Transition Zone (Sierra Madre Oriental, Sierra Madre del Sur, Sierra Madre Occidental and Balsas Basin).     

Track analysis of the Mexican species of Buprestidae (Coleoptera): testing the complex nature of the Mexican Transition Zone

Aim  We analysed the geographical distributions of species of Buprestidae (Coleoptera) in Mexico by means of a panbiogeographical analysis, in order to identify their main distributional patterns and test the complex nature of the Mexican Transition Zone, located between the Nearctic and Neotropical regions.
Location  Mexico.
Methods  The geographical distributions of 228 species belonging to 33 genera of Buprestidae were analysed. Localities of the buprestid species were represented on maps and their individual tracks were drawn. Based on a comparison of the individual tracks, generalized tracks were detected and mapped. Nodes were identified as the areas where generalized tracks converged.
Results  Thirteen generalized tracks were obtained: one was restricted to the Mexican Transition Zone and five to the Neotropical region (Antillean and Mesoamerican dominions), a further two occurred in both the Nearctic region (Continental Nearctic dominion) and the Mexican Transition Zone, and a further five in both the Neotropical region (Mesoamerican dominion) and the Mexican Transition Zone. Seven nodes were identified at the intersections of the generalized tracks – in the Mesoamerican dominion (Mexican Pacific Coast, Mexican Gulf and Chiapas biogeographical provinces) and the Mexican Transition Zone (Trans-Mexican Volcanic Belt, Balsas Basin and Sierra Madre Oriental biogeographical provinces).
Main conclusions  We conclude that the geographical distribution of Buprestidae is mainly Neotropical, corresponding to the Mesoamerican dominion and the Antillean dominion of the Neotropical region, and the Mexican Transition Zone. Most of the generalized tracks and nodes correspond to the Mexican Transition Zone, thus confirming its complex nature. We suggest that the nodes we have identified could be particularly important areas to choose for conservation prioritization.     

Effect of landscape structure on the spatial distribution of Mediterranean dung beetle diversity

Aim  Landscape structure influences the distribution of animals, altering their movements and their ability to reach habitat patches. We analysed the spatial patterns of dung beetle species diversity in three differently structured natural landscapes in a Mediterranean protected area in the centre of the Iberian Peninsula.
Location  Cabañeros National Park, Central Spain.
Methods  Diversity components within (α) and among (β) the three main vegetation types in the reserve were compared by using a hierarchical nested design. These were forests, scrublands and grasslands embedded in three different landscapes, where each was the most dominant and structurally connected habitat.
Results  Species richness of grassland habitat did not vary across landscapes, but forest habitat showed lower species richness in the grassland-dominated landscape. Scrubland was the least species-rich habitat, but here again there was no significant variation across landscapes. However, in all cases, there was a significant influence of habitat context (configuration of habitat patches within landscape matrix) on similarity of species composition. These tended to be more similar to the dominant landscape matrix where they were embedded, rather than to the same habitat type in other landscapes. Additive partitioning of diversity showed higher than expected values of β in all landscapes, which indicated a structured response. Highest values of β in the grassland-dominated landscape suggest that this was the least connected landscape for dung beetles.
Main conclusions  Our results suggest that in homogeneous conditions of climate and trophic resources, landscape structure may well be more important than habitat type as a determinant of dung beetle distribution in the Mediterranean.     

Dung beetles in pasture landscapes of Central America: proliferation of synanthropogenic species and decline of forest specialists

This study examines the spread of synanthropogenic dung beetles (species favoured by human activities) in pasture landscapes in Central America, and evaluates the role of forest fragments and regenerating patches of native vegetation in maintaining beetle diversity. Pitfall trapping was carried out at nine locations in El Salvador and seven in Atlantic Nicaragua that included both pasture and remnant or regenerating native vegetation. More dung beetle species occurred in forest fragments than in pastures. Community composition differed considerably between forest fragments from El Salvador and Nicaragua with many species restricted to either the Caribbean or Pacific regions. In contrast, dung beetle community composition and structure were largely the same in the pastures of El Salvador and Nicaragua, regardless of region or original habitat-type, and were similar to published results from pastures in Mexico and elsewhere on the Isthmus. Very small patches of native shrubs and tree stands (<2.5 ha) maintained no forest specialists in Nicaragua, whereas, in El Salvador, some forest specialists occurred even in the smallest stands of trees (ca. 0.25 ha). The study indicates that the expansion of cattle pastures has caused a regional decline in dung beetle diversity. Forest fragments and small isolated patches of native trees and shrubs maintain some of the diversity of the original landscape but their conservation value for dung beetles will depend on the biogeographical history of the sites.     

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.     

Importance of habitat heterogeneity in remnant patches for conserving dung beetles

Landscape heterogeneity affects the spatial distribution of species. This makes it an important consideration for conservation planning, particularly when designing sustainable production landscapes. We determine whether conserving landscape elements within a transformed landscape is adequate for conserving dung beetle biodiversity. Dung beetles are excellent indicators for landscape biodiversity studies as they are ecologically sensitive. Here we measure dung beetle alpha-diversity, as well as beta-diversity within landscape elements and across different landscape elements. In doing so, we assess the value of landscape elements, as well as variation within landscape elements, in determining the spatial distribution of dung beetles across a production landscape. The study was conducted in the commercial timber production area of the KwaZulu-Natal Midlands, South Africa. In this system, the different landscape elements are a mosaic of natural indigenous forests, grasslands and alien pine plantation blocks. Our results show that the only response for dung beetle alpha-diversity was higher species richness in grasslands and pine blocks compared to natural forests. The highest beta-diversity for a landscape element was the grassland, for elevational category was low elevational areas and grassland type was the Midlands Mistbelt Grassland. The compositional diversity (beta-diversity between elements) was significantly different for all pairwise variations between landscape elements, the elevational categories and grassland types. Natural forests embedded in the two different grassland types had greater differences in compositional diversity than those embedded in natural (grassland) or transformed (pine blocks) matrices. This highlights the need to conserve a range of similar remnant patches of natural vegetation regionally, in addition to conserving broad landscape elements (i.e. grasslands or natural forests) as conservation targets. Furthermore, our results are encouraging for the potential benefits from the ecosystem service provided by dung beetles across the whole landscape, even in the transformed elements.     

Landscape attributes shape dung beetle diversity at multiple spatial scales in agricultural drylands

Land-use change is one of the main drivers of biodiversity loss worldwide, but its negative effects can vary depending on the spatial scale analyzed. Considering the continuous expansion of agricultural demand for land, it is urgent to identify the drivers that shape biological communities in order to balance agricultural production and biodiversity conservation in human-modified landscapes. We used a patch-landscape design and a multimodel inference approach to assess the effects of landscape composition and configuration at two spatial scales (patch and landscape) on the structure of dung beetle assemblages. We performed our study in the Caatinga, the largest dry forest in South America. We sampled 3,526 dung beetles belonging to 19 species and 11 genera. At patch scale, our findings highlight the positive relationship of forest cover and landscape heterogeneity with dung beetle diversity, which are the major drivers of beetle assemblages. Edge density, in turn, is a major driver at the landscape scale and has a negative effect on beetle diversity. Our results support the hypothesis that landscapes combining natural vegetation remnants and heterogeneous agricultural landscapes are the most effective at conserving the biodiversity of dung beetles in the Caatinga landscapes. Directing efforts to better understand the dynamics of dung beetles in agricultural lands can be helpful for policymakers and scientists to design agri-environment schemes and apply conservation strategies in tropical dry forests.     

The effects of land use change on native dung beetle diversity and function in Australia's Wet Tropics

The impacts of land use change on biodiversity and ecosystem functions are variable, particularly in fragmented tropical rainforest systems with high diversity. Dung beetles (Scarabaeinae) are an ideal group to investigate the relationship between land use change, diversity and ecosystem function as they are easily surveyed, sensitive to habitat modification and perform many ecosystem functions. Although this relationship has been investigated for dung beetles in some tropical regions, there has been no study assessing how native dung beetles in Australia's tropical rainforests respond to deforestation, and what the corresponding consequences are for dung removal (a key ecosystem function fulfilled by dung beetles). In this study we investigated the relationship between dung beetle community attributes (determined through trapping) and function (using dung removal experiments that allowed different dung beetle functional groups to access the dung) in rainforest and cleared pasture in a tropical landscape in Australia's Wet Tropics. Species richness, abundance and biomass were higher in rainforest compared to adjacent pasture, and species composition between these land use types differed significantly. However, average body size and evenness in body size were higher in pasture than in rainforest. Dung removal was higher in rainforest than in pasture when both functional groups or tunnelers only could access the dung. Increased dung removal in the rainforest was explained by higher biodiversity and dominance of a small number of species with distinct body sizes, as dung removal was best predicted by the evenness in body size of the community. Our findings suggest that functional traits (including body size and dung relocation behaviour) present in a dung beetle community are key drivers of dung removal. Overall, our results show that deforestation has reduced native dung beetle diversity in Australian tropical landscapes, which negatively impacts on the capacity for dung removal by dung beetles in this region.     

Dung beetle communities as biological indicators of riparian forest widths in southern Brazil

Riparian forests provide important habitat for many wildlife species and are sensitive to landscape change. Among terrestrial invertebrates, dung beetles have been used to investigate the effects of environmental disturbances on forest structure and diversity. Since many studies demonstrated a negative response of dung beetle communities to increasing forest fragmentation, and that most dung beetle species had a more pronounced occurrence during warmest seasons, three hypotheses were tested: (1) Scarabaeinae richness, abundance, diversity and evenness are lower in thinner riparian zone widths than in wider widths during the warmest seasons; (2) Scarabaeinae richness and abundance are positively influenced by leaf litter coverage and height and canopy cover; and (3) Scarabaeinae composition varies with the reduction in riparian vegetation and among annual seasons. We selected four fragments with different riparian zone widths in three secondary streams in southern Brazil. In each fragment, four sampling periods were carried out seasonally between spring 2010 and winter 2011. We collected dung beetles using pitfall traps with two types of bait. We collected 1289 specimens distributed among 29 species. In spring and summer, dung beetle richness was higher in fragments with the widest riparian zone than in those with a thinner riparian zone, and it did not vary between fragments in fall and winter seasons. Dung beetle abundance did not differ among fragments with different riparian zone widths, but it was higher in spring and summer than fall and winter. Richness and abundance were positively influenced by leaf litter. While dung beetle diversity was higher in fragments with wider riparian zone widths than in those with thinner widths, the evenness was similar among fragments. Dung beetle composition differed between the fragments with the widest and thinnest riparian zones, and it also varied among the seasons. Our results suggest that decreased riparian zones affect negatively to dung beetle community structure in southern Brazil. Fragments with thinner riparian zones had lower beetle richness in warmest seasons and an altered community composition. In this sense, the dung beetles are potentially good indicators of riparian forest fragmentation since some species were indicators of a particular riparian zone width. From a conservation perspective, our results demonstrate that the new Brazilian Forest Code will greatly jeopardize not only the terrestrial and aquatic biodiversity of these ecosystems, but also countless other ecological functions.     

Linking Biodiversity,the Environment and Ecosystem Functioning: Ecological Functions of Dung Beetles Along a Tropical Elevational Gradient

Biodiversity loss and anthropogenic environmental changes are known to impact ecosystem functions and services. However, there are still some uncertainties such as confounding environmental factors other than community attributes that affect ecosystem functioning. Our goal was to understand what factors influence the performance of Scarabaeinae dung beetle functions, testing the hypothesis that both community attributes and environmental variables influence the performance. Toward this aim, we collected dung beetles along an elevational gradient (800–1400 m a.s.l.) in the Espinhaço mountain range (Brazil) and quantified dung beetle functions, that is, dung removal, soil excavation and secondary seed dispersal. We recorded data on environmental factors related to climate, soil and vegetation and evaluated their effects on dung beetle functions. Dung beetle ecological functions declined with elevation and the decrease was more pronounced than richness, indicating that there are other factors involved in functions performance besides diversity of beetles. Indeed, we found that the ecological functions measured were dependent on both dung beetle community attributes and environmental factors. Climate, soil and vegetation influenced dung beetle function performance as much as richness, abundance and body size. Dung beetle functional diversity did not explain any of the functions measured. Our study demonstrates that ecological functions are directly influenced by both community attributes and environmental variables and confirms the link between biodiversity, environment and ecosystem functioning.     

Copro-necrophagous beetle (Coleoptera: Scarabaeidae) diversity in an agroecosystem in Yucatan, Mexico

Scarabaeinae are sensitive to structural habitat changes caused by disturbance. We compared copronecrophagous beetle (Scarabaeinae) community structure in three differently managed zones within an agroeco-system of the northern Yucatan Peninsula, Mexico. We placed dung and carrion traps once a month from June 2004 through May 2005. The beetle community included 17 species from the genera Canthon, Canthidium, Deltochilum, Pseudocanthon, Malagoniella, Onthophagus, Phanaeus, Copris, Uroxys, Sisyphus and Ateuchus. The secondary vegetation had a higher beetle diversity than the other two zones. Species richness was highest in the Brosimum alicastrum plantation. The pasture had the lowest species diversity and richness, but exhibited the highest abundance of Scarabaeinae in the dry season. The two zones with extensive tree cover were the most diverse. Roller beetles were dominant over burrower species and small-sized species outnumbered large species. Our data show two important issues: beetle species in the pasture extended their activity to the beginning of the dry season, while abundances dropped in the other, unirrigated zones; and the possibility that the Scarabaeinae living in neotropical forests are opportunistic saprophages and have specialized habits for resources other than dung. The B. alicastrum plantation is beneficial to the entire ranch production system because it functions as a dispersion and development area for stenotopic species limited to tree cover.     

Effects of land usage on dung beetle assemblage structure: Kruger National Park versus adjacent farmland in South Africa

Little quantitative evidence exists regarding how effective protected areas are for preserving species. We compared dung beetle assemblages (Coleoptera: Scarabaeidae: Scarabaeinae) inside and outside of the Kruger National Park, which protects indigenous flora and fauna over a large area of savanna in the northeast lowlands of South Africa. Although it is contiguous with other reserves in South Africa, Zimbabwe and Mozambique, parts of its border abut onto farmland. Some effects of differing land usage either side of this border were studied at the South African Wildlife College (24.541° S 31.335° E) and the nearby farming village of Welverdiend using dung beetle assemblage structure (Coleoptera: Scarabaeidae: Scarabaeinae) as indicators. Samples were taken from gabbro-derived and granite-derived soils in open woody vegetation, both within the reserve and on adjoining farmland, using composite pig, elephant and cattle dung baits in the early rainy season (November 2009) and separate pig and elephant dung baits in the late rainy season (March 2010). Despite much higher large mammal density around Welverdiend, significantly greater species richness, abundance, and biomass of dung beetles were recorded in the reserve where mammal species diversity is greater and elephants produce much larger droppings than any mammal in the farmland. Assemblage structure also differed strongly between dung types, weather conditions on sample days, and season, but weakly between sampled soil types. These differences in assemblage structure were recorded over short distances as the sites in the reserve were only 3?C4?km from those in farmland at Welverdiend.     

Temporal coexistence of dung-dweller and soil-digger dung beetles (Coleoptera, Scarabaeoidea) in contrasting Mediterranean habitats

The western part of the Mediterranean basin is a transitional biogeographical region for the distribution of the representatives of the main guilds of dung beetles; towards the south, Aphodiinae (dung-dwellers) become scarce, whereas northwards Scarabaeinae (soil-diggers) progressively disappear. The number of species in local dung beetle assemblages is enhanced by this double faunistic contribution. Annual dung beetle assemblages were sampled in two sub-Mediterranean sites, which differed by 600 m in elevation, in order to determine the phenological dynamics related to the way of using dung (dung-dwellers/Aphodiinae vs. soil-diggers/Scarabaeinae and Geotrupinae). Aphodiids were active all year round, although they were affected by summer drought and, at high elevation, by the length of the cold season. This reduced activity was related to an impoverishment of Aphodiinae and to reduced temporal segregation between species. In contrast, soil-diggers were not active all year round and showed different species assemblages in the two sites. An extension of the activity period of these beetles was observed due to the occurrence of cold resistant species at high elevation. Our results suggested that the occurrence of soil-diggers seemingly did not affect the seasonality of dung-dwellers; their local abundance showed no negative correlation and, most importantly, phenological differences between dung-dwellers were always significantly higher than the seasonal differences between dwellers and diggers.     

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.     

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.     

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 copronecrophagous beetle communities to habitat disturbance in two mountains of the Mexican Transition Zone: influence of historical and ecological factors

We analyze the influence of historical and ecological factors on the diversity and composition of communities of copronecrophagous beetles associated with conserved and disturbed habitats on two mountains of the Mexican Transition Zone: one mountain of xeric (Las Derrumbadas) and the other of temperate (El Pinal) climate. We collected a total of 1113 beetles of 14 species. While abundance was similar between mountains, the number of species found on the temperate mountain (589 individuals: 11 species) was higher than on the xeric mountain (524 individuals: 5 species). On the temperate mountain, the disturbed habitat was approximately twice as diverse as the conserved habitat, while the opposite pattern was observed on the xeric mountain. Analysis of species turnover between mountains indicated the presence of two relatively different communities and the magnitude of the species turnover related to habitat disturbance depended on the type of mountain, having a higher turnover between habitat types in the temperate mountain. The xeric mountain was dominated by species belonging to the Paleoamerican Plateau distribution pattern, while the Paleoamerican Montane pattern dominated on the temperate mountain, whose species increased their abundance in disturbed habitats together with species of Nearctic affinity. Our results suggest a negative effect of habitat disturbance on the xeric mountain. While on the temperate mountain, beetle diversity seems to increase with disturbance. These results reiterate the need to consider regional-scale historical and ecological processes in order to understand the effects of disturbance and permit the establishment of conservation strategies to adequately protect the organisms, as well as the functions they provide for natural and anthropic ecosystems alike.     

Scale-Dependence of Processes Structuring Dung Beetle Metacommunities Using Functional Diversity and Community Deconstruction Approaches

Community structure is driven by mechanisms linked to environmental, spatial and temporal processes, which have been successfully addressed using metacommunity framework. The relative importance of processes shaping community structure can be identified using several different approaches. Two approaches that are increasingly being used are functional diversity and community deconstruction. Functional diversity is measured using various indices that incorporate distinct community attributes. Community deconstruction is a way to disentangle species responses to ecological processes by grouping species with similar traits. We used these two approaches to determine whether they are improvements over traditional measures (e.g., species composition, abundance, biomass) for identification of the main processes driving dung beetle (Scarabaeinae) community structure in a fragmented mainland-island landscape in southern Brazilian Atlantic Forest. We sampled five sites in each of four large forest areas, two on the mainland and two on the island. Sampling was performed in 2012 and 2013. We collected abundance and biomass data from 100 sampling points distributed over 20 sampling sites. We studied environmental, spatial and temporal effects on dung beetle community across three spatial scales, i.e., between sites, between areas and mainland-island. The γ-diversity based on species abundance was mainly attributed to β-diversity as a consequence of the increase in mean α- and β-diversity between areas. Variation partitioning on abundance, biomass and functional diversity showed scale-dependence of processes structuring dung beetle metacommunities. We identified two major groups of responses among 17 functional groups. In general, environmental filters were important at both local and regional scales. Spatial factors were important at the intermediate scale. Our study supports the notion of scale-dependence of environmental, spatial and temporal processes in the distribution and functional organization of Scarabaeinae beetles. We conclude that functional diversity may be used as a complementary approach to traditional measures, and that community deconstruction allows sufficient disentangling of responses of different trait-based groups.