The Importance of Maize Management on Dung Beetle Communities in Atlantic Forest Fragments

Dung beetle community structures changes due to the effects of destruction, fragmentation, isolation and decrease in tropical forest area, and therefore are considered ecological indicators. In order to assess the influence of type of maize cultivated and associated maize management on dung beetle communities in Atlantic Forest fragments surrounded by conventional and transgenic maize were evaluated 40 Atlantic Forest fragments of different sizes, 20 surrounded by GM maize and 20 surrounded by conventional maize, in February 2013 and 2014 in Southern Brazil. After applying a sampling protocol in each fragment (10 pitfall traps baited with human feces or carrion exposed for 48 h), a total of 3454 individuals from 44 species were captured: 1142 individuals from 38 species in GM maize surrounded fragments, and 2312 from 42 species in conventional maize surrounded fragments. Differences in dung beetle communities were found between GM and conventional maize communities. As expected for fragmented areas, the covariance analysis showed a greater species richness in larger fragments under both conditions; however species richness was greater in fragments surrounded by conventional maize. Dung beetle structure in the forest fragments was explained by environmental variables, fragment area, spatial distance and also type of maize (transgenic or conventional) associated with maize management techniques. In Southern Brazil’s scenario, the use of GM maize combined with associated agricultural management may be accelerating the loss of diversity in Atlantic Forest areas, and consequently, important ecosystem services provided by dung beetles may be lost.     

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

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

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.

     

Invasion and retreat: shifting assemblages of dung beetles amidst changing agricultural landscapes in central Peru

This study examines the effects of deforestation, and the habitat value of coffee and regenerated forest for tropical dung beetles, a functionally significant insect group. Pitfall trapping was conducted at 22 sites in a montane region of central Peru during April and November/December of 2002. Sites included primary and secondary forest, shade-coffee, regenerated forest and open farms (mainly with banana, yuca, and corn). Ordination techniques indicated that beetle assemblages in forests, regenerated forest and coffee were relatively similar. However, assemblage compositions in forested areas differed even at similar altitudes under the influence of biogeographical factors, and the assemblages at disturbed sites (farms/coffee) were influenced by beetle dispersal from adjacent forests. During dry months, when beetle activity is low, communities at all habitat types tended to converge because fewer unique species were recorded in forests at that time and habitat/season generalists were dominant. Preliminary results also indicate that beetles in shady crops such as bananas responded to plant growth: as the banana canopy closed-in, producing more shade, open-habitat specialists retreated and forest/shade specialists invaded the sites. Chronosequence data at two of the sites demonstrate the rapid and dramatic changes in species richness and assemblage composition caused by deforestation. As forests become increasingly fragmented, and open farms continue to expand, dung beetles will become more restricted to the remaining fragments and reserves. In the mosaic landscape studied here, shade crops, like coffee, act as habitat and corridors for many dung beetle species. Small farm size and the consequent magnitude of edge effects, likely contributed to beetle movement between habitat types and determined the apparent generalist nature of many of the dung beetle species in this study.     

Dung beetles in continuous forest, forest fragments and in an agricultural mosaic habitat island at Los Tuxtlas, Mexico

With the aim of determining what kind of landscape mosaics might sustainmaximum diversity and minimum species loss, dung beetles were sampled withbaited pitfall traps to compare species richness and species composition in atract of continuous forest, forest fragments and a habitat island consisting ofa mosaic of forest and arboreal crops in Los Tuxtlas, southern Mexico. Wecaptured 7332 dung beetles representing 33 species. Similar numbers of specieswere captured in the three habitats. However, 56% of individuals were capturedin the continuous forest, 29% in the mosaic habitat and 15% in the forestfragments. Eight species (Canthon femoralis,Copris laeviceps, Canthidium centrale,Onthophagus batesi, Deltochilumpseudoparile, O. rhinolophus, Canthonviridis vazquezae and Dichotomius satanus) accounted for 90% of thecaptures, but their relative dominance varied among habitats. A clear trend wasevident in the number of dung beetles captured in the dung processing guilds(rollers/tunnelers) as well as in the diurnal and nocturnal guilds, withcaptures decreasing from the continuous forest to the mosaic habitat to theforest fragments. A similar trend was detected in detection rates for medium andsmall size dung producing mammals. Species richness of forest fragments andmosaic habitat did not differ from that found in the continuous forest, butthese habitats differ significantly in species richness from isolated shaded andunshaded plantations, linear strips of vegetation, the forest–pasture edge andpastures according to rarefaction analysis. The co-occurrence of the continuousforest, the mosaic habitat and the cluster of forest fragments in closeproximity seems to be preserving a diverse assemblage of dung beetle species inthe local landscape.     

Deforestation and biodiversity: effects of bushland cultivation on dung beetles in semi-arid Tanzania

Dung beetles provide important ecosystem services in the habitats where they occur. The activity of dung beetles enhances soil nutrient cycling and increases the soil’s ability to absorb and hold water. Consequently, these beetles are particularly important in semi-arid environments. This study analyses the importance of remaining wooded habitat patches (bushland) for the survival of a diverse dung beetle fauna in an otherwise cultivated landscape in semi-arid Tanzania. Dung beetles were sampled by pitfall trapping in maize fields and bushland habitats. In total, 6037 dung beetles (Scarabaeidae: Scarabaeinae), representing 77 species from 25 genera, were collected. Many species, particularly amongst the ball-rollers, showed a clear preference for wooded patches, species richness being significantly higher in the bushland patches than in the cultivated sites. The number of trapped specimens in bushland was also considerably higher than that found in maize fields, although the differences were not significant. In conclusion, bushland fragments appear to have an important conservation value as to maintaining a high diversity and abundance of dung beetles, thereby enhancing the ecosystem services provided by these beneficial insects.     

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 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 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.     

Linking functional group richness and ecosystem functions of dung beetles: an experimental quantification

Dung beetles form an insect group that fulfils important functions in terrestrial ecosystems throughout the world. These include nutrient cycling through dung removal, soil bioturbation, plant growth, secondary seed dispersal and parasite control. We conducted field experiments at two sites in the northern hemisphere temperate region in which dung removal and secondary seed dispersal were assessed. Dung beetles were classified in three functional groups, depending on their size and dung manipulation method: dwellers, large and small tunnelers. Other soil inhabiting fauna were included as a fourth functional group. Dung removal and seed dispersal by each individual functional group and combinations thereof were estimated in exclusion experiments using different dung types. Dwellers were the most diverse and abundant group, but tunnelers were dominant in terms of biomass. All dung beetle functional groups had a clear preference for fresh dung. The ecosystem services in dung removal and secondary seed dispersal provided by dung beetles were significant and differed between functional groups. Although in absolute numbers more dwellers were found, large tunnelers were disproportionally important for dung burial and seed removal. In the absence of dung beetles, other soil inhabiting fauna, such as earthworms, partly took over the dung decomposing role of dung beetles while most dung was processed when all native functional groups were present. Our results, therefore, emphasize the need to conserve functionally complete dung ecosystems to maintain full ecosystem functioning.     

Dung beetle persistence in human-modified landscapes: Combining indicator species with anthropogenic land use and fragmentation-related effects

Identifying and making use of ecological indicators becomes an essential task in the conservation of tropical systems, mainly in fragmented landscapes where land use intensification and habitat loss are confounding factors in the detection of species’ responses to human-caused disturbance. We aimed to analyze the importance of anthropogenic land use and fragmentation-related effects on dung beetle (Coleoptera: Scarabaeinae) persistence according to the interior–exterior non-linear gradient (forest + matrix) in a fragmented Atlantic Forest landscape used to sugar cane production and cattle ranching/farming. We offer scores for a comprehensive set of community-level attributes, from beetle abundance to taxonomic and ecological composition (i.e. species body size), including a list of indicator species of different forest habitats and adjacent matrix. Dung beetles were surveyed by traps across forest interiors (i.e. core forest areas) and edges of a primary forest, small fragments, sugar cane fields and pastures in a total of 60 sites. Indicator analyses were conducted across the landscape, using two well-established methods (IndVal and SIMPER). Our results suggest that (1) cross-habitat taxonomic distinctness is associated with the presence of indicator species, (2) some species benefit or are dependent of open habitats created by human-disturbances, such as forest edges (e.g. Canthon nigripennis) and matrices (e.g. Canthon aff. piluliformis, Dichotomius nisus and Trichilum externepunctatum), (3) although landscape habitats exhibit reduced beta diversity, dung beetle assemblages are spatially organized in response to the presence of both forest habitats and matrix and fragment area, (4) forest interior supports beetle assemblages biased toward large-bodied species, (5) accordingly forest interior, forest edges and matrix support taxonomically distinct assemblages, both contributing to the bulk of species richness at landscape level, (6) the response of dung beetles to the interior–exterior non-linear gradient (i.e. forest edge + matrix) reveals a similar pattern regardless of the nature of the matrix, and (7) there is no within-habitat variation in beetle abundance and species richness associated with distance from forest edge. Given that there is a high number of forest-dependent or forest-interior specialist species (e.g. Aphengium aff. sordidum, Ateuchus aff. alipioi, Dichotomius mormon, Ontherus aff. erosus and Onthophagus aff. clypeatus) dung beetle persistence in human-modified landscape is highly dependent on the presence of core areas, although edge-affected and matrix habitats may be complementary. This information is essential to permit a better prospect for dung beetle persistence in human-modified landscapes as they continue to move toward edge-dominated landscapes with intensively managed matrices.     

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.     

Influence of land use on the taxonomic and functional diversity of dung beetles (Coleoptera: Scarabaeinae) in the southern Atlantic forest of Argentina

The degradation and replacement of native ecosystems affects both their taxonomic and functional biodiversity. However, native species may find a gradient of habitat suitability in different land uses within a region. The aim of this study was to evaluate the effect of land use on the taxonomic and functional diversity of dung beetle assemblages in the southern Atlantic forest of Argentina. Dung beetles were sampled in both the native forest (control) and different land uses (Pine and Yerba mate plantations and cattle pastures) during the 2014 summer, using pitfall traps baited with human feces and rotten meat. Samplings were taken from 20 different sites, with five replicates of each land use and the native forest (100 pitfall traps in total). A total of 1699 beetles of 27 species were captured. Canthon quinquemaculatus, Canthon conformis and Dichotomius sericeus were the most abundant species. Cattle pastures were the land use most negatively affected in their taxonomic and functional diversity, particularly large paracoprid dung beetles. Pine plantations maintained their taxonomic and functional diversity in relation to the native forest and Yerba mate plantations showed, in general, an intermediate situation. Microclimatic conditions (average temperature and humidity and maximum temperature) were correlated with functional diversity (the proportion of large paracoprid dung beetles decreased with increasing temperature) and are probably good predictors to explain the observed patterns of functional diversity of dung beetles. The development of sustainable production systems that preserve the native biodiversity requires the conservation of key components from the ecological niche of native species, especially microclimatic conditions.     

Vertical Stratification Responses of an Arboreal Dung Beetle Species to Tropical Forest Fragmentation in Malaysia

Little is known about how arboreal arthropods respond to forest disturbance. We assessed how rain forest fragmentation affected the vertical stratification of arboreal dung beetles in Peninsular Malaysia. An arboreal dung beetle species (Onthophagus sp.) moved closer to the forest floor in smaller fragments, suggesting a downward shift in its foraging possibly in response to fragmentation.     

Using dung beetles to evaluate the effects of urbanization on Atlantic Forest biodiversity

We used dung beetles to evaluate the impact of urbanization on insect biodiversity in three Atlantic Forest fragments in Londrina, Paraná, Brazil. This study provides the first empirical evidence of the impact of urbanization on richness, abundance, composition and guild structure of dung beetle communities from the Brazilian Atlantic Forest. We evaluated the community aspects (abundance, richness, composition and food guilds) of dung beetles in fragments with different degrees of immersion in the urban matrix using pitfall traps with four alternative baits (rotten meat, rotten fish, pig dung and decaying banana). A total of 1 719 individuals were collected, belonging to 29 species from 11 genera and six Scarabaeinae tribes. The most urban‐immersed fragment showed a higher species dominance and the beetle community captured on dung presented the greatest evenness. The beetle communities were distinct with respect to the fragments and feeding habits. Except for the dung beetle assemblage in the most urbanized forest fragment, all others exhibited contrasting differences in species composition attracted to each bait type. Our results clearly show that the degree of urbanization affects Atlantic Forest dung beetle communities and that the preservation of forest fragments inside the cities, even small ones, can provide refuges for Scarabaeinae.     

Ecosystem services provided by dung beetles in Australia

Although there are nearly 500 species of native dung beetles in Australia, most are adapted to small, hard, dry, pelletised marsupial droppings and not to dealing with the large, moist deposits of cattle. In 1788, Governor Arthur Phillip arrived at Botany Bay with five cows, two bulls, 44 sheep and seven horses: this signalled major changes in Australia. Now there are about 27 million cattle, whose annual dung production has a dry matter content of about 42 million tonnes. Until CSIRO introduced exotic dung beetles in the 1960s, the dung of these herbivores sat on the soil surface, sometimes for years, locking up organic matter, smothering pasture and polluting waterways. CSIRO introduced 53 exotic dung beetle species, of which 43 were released to the Australian mainland between 1965 and 1985. Twenty-three of these have become established, many of which have reached the natural limits of their distribution. I consider the reason for the failure of the other 30 species to establish and briefly review previous contributions to examining the role of dung beetles in delivering ecosystem services, noting that much of the published literature concerns laboratory studies. New field data are then examined on the way in which introduced species are transforming dung communities and the ecosystem services they provide. The capacity of deep-tunnelling dung beetles to transform the soil profile is examined along with their effects on pasture production and the flow of nutrients from dung on pasture. The biocontrol capacity of dung beetle activity is considered in relation to the native bush fly, Musca vetustissima, the introduced buffalo fly, Haematobia irritans exigua, and dung-borne intestinal parasites (helminths and Cryptosporidium). The rationale for introducing additional species to Australia is considered.     

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.     

Tree retention in cattle ranching systems partially preserves dung beetle diversity and functional groups in the semideciduous Atlantic forest: The role of microclimate and soil conditions

The disturbance of natural environments affects, among others, the diversity of dung beetle assemblages, which could have serious consequences for the ecological processes regulated by these insects. The objective of this study was to evaluate and compare species diversity and functional groups of dung beetle assemblages both in the native forest and in three livestock systems that differed in their structure and composition of vegetation: a livestock system with native trees, a livestock system with exotic trees (Pinus taeda), and traditional open pastures, in the semideciduous Atlantic forest of Argentina, in an area previously covered by continuous forest and currently with a heterogeneous landscape of native forest and different land uses. Pitfall traps baited with cow dung were used in the natural forests and the livestock systems studied. A total of 2461 beetles belonging to 38 species were captured. Treed livestock systems showed the highest species richness (⁰D) and diversity (¹D and ²D). Twelve functional groups were identified. The native forest showed the highest functional group richness, while open pastures had the lowest. In general, livestock systems showed a low proportional abundance of telecoprid, diurnal and large beetles. Microclimate (average temperature and humidity) and soil conditions (soil composition: sandy or clayey) were closely associated with the species and functional group composition. Results confirm that cattle ranching with tree retention preserves dung beetle diversity, and suggest that cattle systems without canopy cover have higher impact (negative effects) than silvopastoral systems on both species and functional groups.