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.     

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.     

Nesting strategies affect altitudinal distribution and habitat use in Alpine dung beetle communities

1. Dung beetles are key contributors to a suite of ecosystem services. Understanding the factors that dictate their distributions is a necessary step towards preventing negative impacts of biodiversity loss. 2. Alpine dung beetle communities were analysed along altitudinal gradients to assess how different components of the community, defined in terms of nesting strategy [dung‐ovipositing Aphodiidae (DOAs), soil‐ovipositing Aphodiidae (SOAs) and two paracoprid (PAR) groups, Geotrupidae and Scarabaeidae] and parameters relevant to dung removal rates (species richness, total biomass and functional diversity), are distributed, and to identify to which environmental factors they respond. 3. Species richness declined with altitude. There was no significant variation in functional diversity or total biomass in relation to altitude. There were significant variations when considered by nesting group: DOA species richness and biomass decreased, SOA biomass increased, and Geotrupidae biomass showed a non‐linear trend, as altitude increased. 4. Functional diversity and total species richness were positively related to vegetation cover. DOA species richness was highest in forest and scrub; SOA species richness was highest in grassland and PAR species richness was lowest in rocky areas. 5. Dung beetle species show different trends in species richness and biomass depending on nesting strategy. Management to promote the dung beetle community should include maintenance of a mosaic of habitat types. Given the likely importance of species richness and biomass to ecosystem functioning, and the complimentary effect of different dung beetle groups, such a strategy may protect and enhance the ecosystem services that Alpine dung beetles provide.     

Functional diversity predicts overyielding effect of species combination on primary productivity

The effect of biodiversity on ecosystem functioning has proven variable both within and among manipulative studies. Species richness is the most commonly used measure of biodiversity in such studies, but the range of species’ functional traits (functional diversity), not the number of species per se, likely underpins a key mechanistic link between species richness and ecosystem functioning. However, the majority of experiments that have examined the effect of functional diversity have manipulated functional group richness, an approach recognised to suffer numerous limitations. Continuous measures of functional diversity avoid many of these limitations, but the relationship between continuous functional diversity and the magnitude of ecosystem processes has been largely untested. Using one vs two‐species mixtures of rock pool macroalgae as a model, we conducted a field experiment to determine the effect of a continuous measure of functional diversity (functional attribute diversity, FAD, the degree of functional differentiation based on four functional traits) on the magnitude of net primary productivity and overyielding, based upon two alternative null‐models. The total magnitude of productivity was largely determined by the identity of species present, not FAD. However, FAD proved to be a good predictor of overyielding (variation in productivity after the dominant effects of species identity had been accounted for). Furthermore, despite differences in the mean magnitude of the effect of combining species, the positive relationship between FAD and overyielding was consistent according to both additive and substitutive null‐models. Our findings imply that whilst knowledge of species’ independent contributions remains indispensable in the prediction of biotic effects on ecosystem functioning within a trophic level, continuous measures of functional diversity should be used as a supplementary tool to predict the magnitude of overyielding, thereby refining predictions.     

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.     

Effect of dung beetle species richness and chemical perturbation on multiple ecosystem functions

1. The relationship between biodiversity and ecosystem functioning is typically positive but saturating, suggesting widespread functional redundancy within ecological communities. However, theory predicts that apparent redundancy can be reduced or removed when systems are perturbed, or when multifunctionality (the simultaneous delivery of multiple functions) is considered. 2. Manipulative experiments were used to test whether higher levels of dung beetle species richness enhanced individual functions and multifunctionality, and whether these relationships were influenced by perturbation (in this case, non‐target exposure to the veterinary anthelmintic ivermectin). The four ecosystem functions tested were dung removal, primary productivity, soil faunal feeding activity and reduction in soil bulk density. 3. For individual functions, perturbation had limited effects on functioning, with only dung removal significantly (negatively) affected. Species richness did not, on its own, explain significant variation in the delivery of individual functions. In the case of primary productivity, an interaction between richness and perturbation was found: species‐rich dung beetle assemblages enhanced forage growth in the unperturbed treatment, relative to the perturbed treatment. 4. Using a composite ‘multifunctionality index’ it was found that species‐rich dung beetle assemblages delivered marginally higher levels of multifunctionality in unperturbed conditions; however, this benefit was lost under perturbation. Using a relatively new and robust method of assessing diversity–multifunctionality relationships across a range of thresholds, no significant effect of species richness on multifunctionality was found.     

El Niño impacts on human-modified tropical forests: Consequences for dung beetle diversity and associated ecological processes

Our knowledge of how tropical forest biodiversity and functioning respond to anthropogenic and climate-associated stressors is limited. Research exploring El Niño impacts are scarce or based on single post-disturbance assessments, and few studies assess forests previously affected by anthropogenic disturbance. Focusing on dung beetles and associated ecological functions, we assessed (a) the ecological effects of a strong El Niño, (b) if post-El Niño beetle responses were influenced by previous forest disturbance, and (c) how these responses compare between forests impacted only by drought and those affected by both drought and fires. We sampled 30 Amazonian forest plots distributed across a gradient of human disturbance in 2010, 2016, and 2017—approximately 5 years before, and 3–6 and 15–18 months after the 2015–16 El Niño. We found 14,451 beetles from 98 species and quantified the beetle-mediated dispersal of >8,600 seed mimics and the removal of c. 30 kg of dung. All dung beetle responses (species richness, abundance, biomass, compositional similarity to pre-El Niño condition, and rates of dung removal and seed dispersal) declined after the 2015–16 El Niño, but the greatest immediate losses (i.e., in 2016) were observed within fire-affected forests. Previous forest disturbance also influenced post-El Niño dung beetle species richness, abundance, and species composition. We demonstrate that dung beetles and their ecological functions are negatively affected by climate-associated disturbances in human-modified Amazonian forests and suggest that the interaction between local anthropogenic and climate-related stressors merits further investigation.     

Dung beetle diversity and functions suggest no major impacts of cattle grazing in the Brazilian Pantanal wetlands

1. Dung beetles perform relevant ecological functions in pastures, such as dung removal and parasite control. Livestock farming is the main economic activity in the Brazilian Pantanal. However, the impact of cattle grazing on the Pantanal's native dung beetle community, and functions performed by them, is still unknown. 2. This study evaluated the effects of cattle activity on dung beetle community attributes (richness, abundance, biomass, composition, and functional group) as well as their ecological functions (dung removal and soil bioturbation) in the Pantanal. In January/February 2016, dung beetles were sampled and their ecological functions measured in 16 sites of native grasslands in Aquidauana, Mato Grosso do Sul, Brazil, 10 areas regularly grazed by cattle and six control ungrazed areas (> 20 years of abandonment). 3. In all, 1169 individuals from 30 species of dung beetles were collected. Although abundance, species richness, and biomass did not differ between grasslands with and without cattle activity, species composition and functional groups differed among systems. Large roller beetles were absent from non‐cattle grasslands, and the abundance, richness, and biomass of medium roller beetles was higher in those systems. 4. Despite causing changes in species/functional group composition, the results of this study show that a density compensation of functional groups in cattle‐grazed natural grasslands seems to have conserved the ecological functions (dung removal and soil bioturbation), with no significant differences between systems. 5. Therefore, these results provide evidence that cattle breeding in natural grasslands of the Brazilian Pantanal can integrate livestock production with the conservation of the dung beetle community and its ecological functions.     

Negative impacts of human land use on dung beetle functional diversity

The loss of biodiversity caused by human activity is assumed to alter ecosystem functioning. However our understanding of the magnitude of the effect of these changes on functional diversity and their impact on the dynamics of ecological processes is still limited. We analyzed the functional diversity of copro-necrophagous beetles under different conditions of land use in three Mexican biosphere reserves. In Montes Azules pastures, forest fragments and continuous rainforest were analyzed, in Los Tuxtlas rainforest fragments of different sizes were analyzed and in Barranca de Metztitlán two types of xerophile scrub with different degrees of disturbance from grazing were analyzed. We assigned dung beetle species to functional groups based on food relocation, beetle size, daily activity period and food preferences, and as measures of functional diversity we used estimates based on multivariate methods. In Montes Azules functional richness was lower in the pastures than in continuous rainforest and rainforest fragments, but fragments and continuous forest include functionally redundant species. In small rainforest fragments (<5 ha) in Los Tuxtlas, dung beetle functional richness was lower than in large rainforest fragments (>20 ha). Functional evenness and functional dispersion did not vary among habitat types or fragment size in these reserves. In contrast, in Metztitlán, functional richness and functional dispersion were different among the vegetation types, but differences were not related to the degree of disturbance by grazing. More redundant species were found in submontane than in crassicaule scrub. For the first time, a decrease in the functional diversity in communities of copro-necrophagous beetles resulting from changes in land use is documented, the potential implications for ecosystem functioning are discussed and a series of variables that could improve the evaluation of functional diversity for this biological group is proposed.     

The Role of Species Traits in Mediating Functional Recovery during Matrix Restoration

Reversing anthropogenic impacts on habitat structure is frequently successful through restoration, but the mechanisms linking habitat change, community reassembly and recovery of ecosystem functioning remain unknown. We test for the influence of edge effects and matrix habitat restoration on the reassembly of dung beetle communities and consequent recovery of dung removal rates across tropical forest edges. Using path modelling, we disentangle the relative importance of community-weighted trait means and functional trait dispersion from total biomass effects on rates of dung removal. Community trait composition and biomass of dung beetle communities responded divergently to edge effects and matrix habitat restoration, yielding opposing effects on dung removal. However, functional dispersion—used in this study as a measure of niche complementarity—did not explain a significant amount of variation in dung removal rates across habitat edges. Instead, we demonstrate that the path to functional recovery of these altered ecosystems depends on the trait-mean composition of reassembling communities, over and above purely biomass-dependent processes that would be expected under neutral theory. These results suggest that any ability to manage functional recovery of ecosystems during habitat restoration will demand knowledge of species'' roles in ecosystem processes.     

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.     

Effect of functional group richness and species richness in manipulated productivity–diversity studies: a glasshouse pot experiment

Species and functional group (grasses, legumes, creeping nonlegume forbs, rosette nonlegume forbs) richness of species assemblages composed of 16 species from four functional plant groups were manipulated to evaluate the productivity-diversity relationships in a greenhouse pot experiment. Pots were filled with sand, and supplied at two levels of nutrients. The plants were grown in monocultures, two, four, eight and 16 species mixtures. Individual two, four, and eight species mixtures differed in the richness of functional groups. Although the two characteristics of biodiversity, i.e. species and functional group richness, were necessarily correlated, it was shown that it is possible to separate their effect statistically, and also test for their common effect without pronounced loss of test power. There was a pronounced increase of average aboveground biomass and a mild increase in belowground biomass with biodiversity. The effect of functional group richness was more pronounced than the effect of the number of species. By using the method of Loreau and Hector (Nature 411 (2001) 72), selection and complementarity effects were statistically separated, and the overyielding index was calculated as a ratio of the productivity of a mixture to the productivity of its most productive component (to demonstrate transgressive overyielding). Positive values of complementarity and transgressive overyielding were both found, particularly in some rich communities and under high nutrient levels. Complementarity significantly increased only with functional group richness and mainly under high nutrients in the belowground biomass. Some species, when grown in monocultures, had decreased productivity under higher nutrients, and thus were more productive in mixtures than in monocultures. It seems that those species suffered from too high nutrient levels when grown in monocultures, but not in the presence of other species, which were able to use the nutrients in high concentrations and effectively decrease the nutrient levels. As a consequence, mixtures of high diversity were always more productive under high nutrients. The difference in species proportions between high and low nutrients, characterized by chord distance, increased with species richness. The relative change in productivity decreased with the number of functional groups. This suggests that species richness might lead to stabilization of aggregate characteristics (like total productivity) under changing environmental conditions by changing the proportions of individual species.     

Partitioning the effects of algal species identity and richness on benthic marine primary production

Influential research in terrestrial habitats indicates that several ecosystem processes are related to plant biodiversity, yet these links remain poorly studied in marine ecosystems. We conducted one field and one mesocosm experiment to quantify the relative effects of macroalgal species identity and richness on primary production in coral reef macroalgal communities off the north coast of Jamaica. We measured production as the net accumulation of algal biomass in the absence of consumers and as photosynthetic rate using oxygen probes in sealed aquaria. We used two recently developed techniques to attribute deviations in expected relative yield to components associated with species identity or diversity and then to further partition diversity effects into mechanistic components based on dominance, trait-dependent complementarity, and trait-independent complementarity. Our results indicate that algal identity had far greater effects on absolute net growth and photosynthesis than richness. The most diverse mixture of macroalgae did not outperform the most productive monoculture or the average monoculture in either measure of primary production (i.e. we did not find evidence of either transgressive or non-transgressive overyielding). Trait-independent complementarity effects were positive but dominance and trait-dependent complementarity were both negative and became stronger when richness was increased. Thus the potentially positive influence of species interactions and niche partitioning on production were negated by dominance and other negative selection effects. These results demonstrate that the counteracting influence of component effects can diminish the net richness effects on production. This could explain frequently observed weak net richness effects in other aquatic and terrestrial systems and suggests that life history tradeoffs greatly reduce the potential for ecologically relevant plant biodiversity effects on ecosystem properties.     

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

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

Co‐declining mammal–dung beetle faunas throughout the Atlantic Forest biome of South America

The millennial–scale evolutionary relationships between mammals and dung beetles have been eroded due to several drivers of contemporary biodiversity loss. Although some evidence of co‐decline has been shown for mammals and dung beetles at some Neotropical sites, a biome‐scale analysis for the entire Atlantic Forest of South America would strengthen our understanding of how relictual sets of mammal species can affect dung beetle co‐occurrences and co‐declines. We therefore collated hundreds of assemblages of both dung beetles and medium‐ to large‐bodied mammals throughout the world's longest tropical forest latitudinal gradient to examine to what extent mammal assemblages may exert a positive influence on dung beetle species composition and functional assembly, and whether this relationship is scale dependent. We also collated several climatic and other environmental variables to examine the degree to which they shape mammal–dung beetle relationships. The relationships between local mammal and dung beetle faunas were examined using regression models, variation partitioning, dissimilarity indices and ecological networks. We found a clear positive relationship between mammal and dung beetle species richness across this forest biome, indicating an ongoing process of mammal–dung beetle niche‐mediated co‐decline. We found a strong relationship between the species composition of both taxa, in which dung beetle species dissimilarity apparently track changes in mammalian dissimilarity, typically in 80% of all cases. Co‐variables such as phytomass and climatic variables also influenced mammal–dung beetle patterns of co‐decline along the Atlantic Forest. We conclude that dung beetle diversity and community assembly are shaped by the remaining co‐occurring mammal assemblages and their functional traits, and both groups were governed by environmental features. We emphasize that ecosystem‐wide effects of mammal population declines remain poorly understood both quantitatively and qualitatively, and curbing large vertebrate defaunation will ensure the persistence of co‐dependent species.     

Ecosystem functioning in relation to species identity,density, and biomass in two tunneller dung beetles

1. Species abundance, biomass, and identity are the main factors that influence ecosystem functioning. Previous studies have shown that community attributes and species identity help to maintain natural ecosystem functioning. 2. This study examined how species identity, biomass, and abundance in dung pats (i.e. density) of dung beetles affect multiple ecological functions: dung removal, seed dispersal, and germination. Specifically, two species of tunnellers were targeted: Onthophagus illyricus (Scopoli, 1763) and Copris lunaris (Linnaeus, 1758). In accordance with their natural abundance, densities ranging from 10 to 80 individuals were considered for O. illyricus, and those from two to eight were considered for C. lunaris, spanning the total biomass per treatment from 0.22 to 1.76 g. 3. Results showed that, even at higher abundance, O. illyricus is not as efficient as C. lunaris. These results show that species identity, biomass, and density are crucial factors for maintaining ecosystem functioning. The combined effect of species identity and density/biomass facilitated dung removal and seed dispersal. Conversely, species identity is the only relevant factor for germination. Moreover, relationships among functions depend on the species investigated: C. lunaris showed a positive correlation between dung removal and seed dispersal, whereas O. illyricus showed a positive correlation between germination and dung removal. 4. In conclusion, optimal ecosystem functioning depends on multiple factors, such as density and species identity, and thus also on body size, nesting strategies and ecological functions investigated. Moreover, the loss of larger and efficient species cannot be compensated by higher abundances of small species.     

Recovery of mammal diversity in tropical forests: a functional approach to measuring restoration

Ecological restoration is increasingly applied in tropical forests to mitigate biodiversity loss and recover ecosystem functions. In restoration ecology, functional richness, rather than species richness, often determines community assembly, and measures of functional diversity provide a mechanistic link between diversity and ecological functioning of restored habitat. Vertebrate animals are important for ecosystem functioning. Here, we examine the functional diversity of small‐to‐medium sized mammals to evaluate the diversity and functional recovery of tropical rainforest. We assess how mammal species diversity and composition and functional diversity and composition, vary along a restoration chronosequence from degraded pasture to “old‐growth” tropical rainforest in the Wet Tropics of Australia. Species richness, diversity, evenness, and abundance did not vary, but total mammal biomass and mean species body mass increased with restoration age. Species composition in restoration forests converged on the composition of old‐growth rainforest and diverged from pasture with increasing restoration age. Functional metrics provided a clearer pattern of recovery than traditional species metrics, with most functional metrics significantly increasing with restoration age when taxonomic‐based metrics did not. Functional evenness and dispersion increased significantly with restoration age, suggesting that niche complementarity enhances species' abundances in restored sites. The change in community composition represented a functional shift from invasive, herbivorous, terrestrial habitat generalists and open environment specialists in pasture and young restoration sites, to predominantly endemic, folivorous, arboreal, and fossorial forest species in older restoration sites. This shift has positive implications for conservation and demonstrates the potential of tropical forest restoration to recover rainforest‐like, diverse faunal communities.     

Ecosystem functioning in stream assemblages from different regions: contrasting responses to variation in detritivore richness, evenness and density

1. The diversity of species traits in a biological assemblage varies not only with species richness, but also with species evenness and organism density, which together influence the concentration of traits within functional guilds. Potential trait diversity at local scales is also constrained by the regional species pool. Implications of such variation for spatio-temporal variability in biodiversity-ecosystem functioning relationships are likely to be complex, but are poorly understood. 2. In microcosm experiments conducted at laboratories in Sweden, Ireland and Romania, we investigated effects of species richness, evenness and density of stream-living detritivores on two related processes: detritivore leaf-processing efficiency (LPE) and growth. Assemblage composition varied among laboratories: one taxonomic order (Plecoptera) was studied in Sweden, whereas two orders, encompassing wider trait variation, were studied in Romania (Trichoptera and Plecoptera) and Ireland (Trichoptera and Isopoda). 3. Relationships between density and both LPE and growth ranged from negative to positive across the study species, highlighting the potential for density-dependent variation in process rates to alter ecosystem functioning, but indicating that such effects depend on species identity. 4. LPE varied with species diversity in the two more heterogeneous assemblages, but whereas LPE in the Romanian study was generally enhanced as richness increased, LPE in the Irish study increased only in less-even polycultures dominated by particular species. Transgressive overyielding was detected in the Irish experiment, indicating complementary resource use and/or facilitation (complementarity). These mechanisms could not be distinguished from the selection effect in the Romanian study. 5. Growth was elevated in Romanian species mixtures, reflecting positive complementarity, but lower than expected growth in some Swedish mixtures was associated with negative complementarity, indicating interspecific interference competition. 6. Our results emphasize the potential importance of detritivore diversity for stream ecosystem functioning, but both the effects of diversity on the studied processes, and the mechanisms underlying those effects, were specific to each assemblage and process. Such variability highlights challenges in generalizing impacts of diversity change for functional integrity in streams and other ecosystems in which the occurrence of important species traits fluctuates over relatively small spatio-temporal scales.     

Co-declining mammals and dung beetles: an impending ecological cascade

Biodiversity loss can precipitate extinction cascades and impair ecological processes. These 'downstream' effects will be exacerbated if functionally important taxa are tightly linked with species threatened by extinction or population decline. We review the current evidence that such a scenario is currently playing out in the linked declines of persistently hunted mammal populations and the dung beetles communities (Coleoptera: Scarabaeidae: Scarabaeinae) that depend on them for adult and larval food resources. Through a close evolutionary association, mammal assemblages have played a fundamental role in structuring extant dung beetle communities. Today many game mammal species' populations are severely depleted by subsistence or commercial hunting, especially in tropical forest systems. Multiple lines of evidence from temperate and tropical systems indicate that the regional-scale decline or extirpation of medium and large bodied mammal faunas can severely disrupt the diversity and abundance of dung beetle communities through alterations in the composition and availability of dung resources. These observed community disassemblies have significant short- and long-term implications for the maintenance of key ecosystem processes including nutrient recycling and secondary seed dispersal. Identifying the species- and community-level traits that buffer or exacerbate these species and functional responses is essential if we are to develop a better understanding of the cascading ecological consequences of hunting in tropical forests.