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.     

Use of habitat resources by scarab dung beetles in an savanna

In the Queen Elizabeth National Park, Uganda, we compared the scarab beetle assemblages in the dung of three wild ungulates (African buffalo, a ruminant foregut fermenter; hippopotamus, nonruminant foregut fermenter; and warthog, nonruminant hindgut fermenter). Dung was collected from two sandy-clay soils with different percentage of coarse sand. We aimed at investigating habitat resource selection by dung beetle species within a savanna natural contest with abundant and diverse food availability. Analyses were performed to detect differences for dung beetle assemblages in abundance, diversity, functional groups. Species richness in the three dung types and in the two soil types was similar. However, warthog dung and sandy-rich soil appeared the preferred habitat resources, in terms of abundance and biomass, while hippopotamus dung hosted the lowest values for these parameters. The analysis of functional groups revealed that slow-burying tunnellers held the major role, both in terms of abundance and biomass, and were mainly found in warthog dung.     

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.     

Experimental evidence for the effects of dung beetle functional group richness and composition on ecosystem function in a tropical forest

Much of the literature on the relationship between species richness or functional group richness and measures of ecosystem function focuses on a restricted set of ecosystem function measures and taxonomic groups. Few such studies have been carried out under realistic levels of diversity in the field, particularly in high diversity ecosystems such as tropical forests. We used exclusion experiments to study the effects of dung beetle functional group richness and composition on two interlinked and functionally important ecological processes, dung removal and secondary seed dispersal, in evergreen tropical forest in Sabah, Malaysian Borneo. Overall, both dung and seed removal increased with dung beetle functional group richness. However, levels of ecosystem functioning were idiosyncratic depending on the identity of the functional groups present, indicating an important role for functional group composition. There was no evidence for interference or competition among functional groups. We found strong evidence for overyielding and transgressive overyielding, suggesting complementarity or facilitation among functional groups. Not all mixtures showed transgressive overyielding, so that complementarity was restricted to particular functional group combinations. Beetles in a single functional group (large nocturnal tunnellers) had a disproportionate influence on measures of ecosystem function: in their absence dung removal is reduced by approximately 75%. However, a full complement of functional groups is required to maximize ecosystem functioning. This study highlights the importance of both functional group identity and species composition in determining the ecosystem consequences of extinctions or altered patterns in the relative abundance of species.     

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 in forests and pastures of El Salvador: a functional comparison

This study examines the functional attributes of Neotropical dung beetles, and, based on the analysis, evaluates the role of dung beetles in pastures and tropical dry forest in El Salvador. Dung beetle diversity was lower in pastures than in forests. However, the total biomass of beetles at similarly-sized dung baits was frequently higher in pastures. Diversity loss followed structured patterns: (1) carrion and fruit feeding beetles were well represented (species number) in the forests but were largely absent from pastures; (2) large ball rollers and small fast tunnellers were present in forests but were absent from pastures; (3) large fast tunnellers and small slow tunnellers were poorly represented in pastures compared to forests but the few species that survived in pastures attained extremely high populations; and (4) each functional group in the species-poor pasture assemblages was divided almost equally into day and night active species. Substrate and habitat generalists that were present in both the forests and pastures were attracted to fruit and carrion in forested habitat but not in open pastures. In open habitats, flies and other insects may be the principal decomposers of decaying fruit and carrion.     

Experimental mixtures of dung fauna affect dung decomposition through complex effects of species interactions

Dung fauna plays an important role in dung decomposition, a key ecosystem process in nutrient cycling in grazed grasslands. The diversity of a three‐species community (dung beetles, dung flies and epigeic earthworms) was systematically manipulated to produce different relative abundance distributions (evenness levels) and the resulting communities were introduced to standardised dung pats in laboratory experiments. Dung decomposition was modelled using an analysis that disentangled species identity effects and the interactions among species that contribute to the diversity effect. This revealed that the net diversity effect was composed of positive (fly–earthworm), negative (beetle–earthworm) and neutral (fly–beetle) effects of species interaction. These pairwise interactions resulted in complex, but systematically varying and predictable effects on dung decomposition as the relative abundances of species changed. Decomposition was consistently greater in communities with higher decomposer biomass. The utility of the adopted analytical approach was emphasised by comparison with an ANOVA that found that dung decomposition did not differ among evenness levels. Thus, the averaging of decomposition across different community structures within evenness levels masked the different effects of species interactions. These results highlight methodological insights into the quantification of diversity–function relationships.     

Variations in dung beetle assemblages across a gradient of hunting in a tropical forest

Populations of large mammals are severely depleted by hunting in tropical forests, with direct effects on plant regeneration. But indirect consequences on commensal taxa depending on them for food resources, like coprophagous beetles, are less documented. Cascading effects of species loss across Scarabaeinae are expected, with likely significant negative implications for ecosystem functions. We examined dung beetle assemblages using pitfall traps at three rain forest sites in French Guiana ranging from intact mammalian fauna (Nouragues) to moderate (Kaw) and heavy (Matoury) defaunation. The site with the most depauperate mammalian fauna showed significantly lower dung beetle species richness than the two other two sites, which were not different from each other. Mean abundance and biomass per trap were not different across sites whereas community composition strongly differed among sites. A positive correlation was observed between body size and the individual contribution to dissimilarity between Nouragues and Kaw. The species contributing the most to dissimilarity were large. By contrast, one medium-sized species, dominant in Matoury, contributed the most to dissimilarity between Matoury and other sites. Diurnal genera of large tunnellers showed a higher diversity and abundance in Nouragues compared to other sites, whereas a nocturnal genus showed no differences. Large rollers were more abundant in Kaw compared to other sites. None of the groups of small beetles but one were affected by defaunation. Our results suggest that loss of large mammal populations affects dung beetle assemblage structure and causes decreasing abundance or disappearance of large tunnellers species that have a major impact on several dung beetle-mediated ecological processes.     

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.     

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.     

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.     

Local and regional ecological morphology of dung beetle assemblages across four biogeographic regions

Aim Niche partitioning within species assemblages is thought to influence species packing and/or total niche space occupied. The evolution of dung beetles (Scarabaeinae) is likely to have been strongly influenced by inter‐specific competition, leading to niche partitioning. We consider whether local‐scale processes leave a signature in regional patterns of functional diversity in dung beetle assemblages, and investigate the correlation between total exploited ecomorphological space and density of species packing with increased species richness. We test whether ecomorphological space occupied by local assemblages reflects that of their regional species pool, and the extent to which ecomorphological space is convergent or divergent within functional groups across regional pools. Location Neotropics, Africa, Australia and Madagascar. Methods Dung beetle assemblages were collected in a standardized manner from four biogeographic regions. Ecomorphological similarity among the assemblages was assessed by multivariate analysis of 19 linear measurements for 300 species and three functional nesting types (roller, tunneller or dweller), firstly on a local level within the Neotropics and Afrotropics, and then between the regional species pools. Results Key body measurements, in particular the hind tibia, separated rollers and tunnellers into largely non‐overlapping entities along the first three axes of the shape analysis. Three Neotropical assemblages, which vary widely in species numbers, each harboured a similar amount of morphometric variation, resulting in increasingly dense species packing with greater species richness. Similar findings were obtained in two South African assemblages. Assemblages in the four biogeographic regions showed largely similar distributions of ecomorphological variation, including the separation of rollers and tunnellers, despite their distant phylogenetic relationships. Ecomorphological similarity among regions was particularly high in tunnellers, whilst the rollers exhibited greater regional differentiation. Main conclusions Local assemblages evidently represent the full diversity of functional groups available in the regional pool, even in species‐poor assemblages. There is a strong trend towards convergence in morphology separating tunnellers and rollers in phylogenetically independent lineages. The ecomorphological similarity of regional assemblages suggests that morphological convergence is the result of common selective forces active within the assemblages themselves. This lends support to the widely hypothesized effect of inter‐specific interactions and niche partitioning in determining assemblage composition and lineage evolution in the Scarabaeinae.     

Ivermectin residues in cattle dung impair insect-mediated dung removal but not organic matter decomposition

1. Veterinary parasiticide residues in livestock dung have been repeatedly shown to negatively affect the abundance and diversity of dung-associated insects. While these losses are concerning from a conservation perspective, they can also translate to impairment of ecosystem functions in agricultural landscapes (e.g. nutrient cycling, primary productivity and greenhouse gas mitigation). 2. Most research focusing on decomposition-related ecosystem functioning has focused on dung beetles (Coleoptera: Scarabaeoidea) while other insects, particularly flies, have received comparatively less attention. 3. Using mesocosms, this study manipulated the insect groups colonising cow dung (beetles only, flies only, beetles and flies together, and an insect-free control). Half of the insects were exposed to 1 mg kg⁻¹ ivermectin in dung, while the other half were exposed to ivermectin-free dung. Dung decomposition (mass of organic matter lost) and dung removal (change in the dry mass of the dung pat attributed to both dung decomposition and burial) were measured. 4. Comparison of beetles and flies in ivermectin-free dung showed that beetles removed nearly twice as much dung as did flies. Comparison of dung removal across all treatments showed that ivermectin residues significantly reduced dung removal provided by beetles by 47% and dung removal provided by beetles and flies together by 32%. 5. Organic matter decomposition was not significantly affected by insect colonists or by the presence of ivermectin, indicating that organic matter decomposition can occur independently of insect activity and chemical perturbations.     

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.     

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.     

Elephant dung decomposition and coprophilous fungi in two habitats of semi-arid Botswana

In order to understand the impact of habitat changes on ecosystem processes caused by increased populations of elephants, elephant dung decomposition was studied in semi-arid Botswana. Dung decomposition rates were studied with and without the presence of arthropods, using pairs of exposed dung and dung enclosed in nylon-mesh bags, respectively. Dung decomposition rates were lower in the absence of arthropods. The rates in the late wet season were higher in the scrubland than in the woodland. In the early dry season, immediately after the wet season, the rates were higher in the woodland than in the scrubland. The difference in decomposition rates between habitats was attributed to microclimatic conditions created by vegetation cover. With regard to fungal succession, Cladosporium cladosporioides and Eurotium brefeldianum occurred only in the late stages of dung decomposition whereas Talaromyces helicus, Cercophora coprophila and Sporormiella minima occurred in all the stages. Although there was no significant difference in Shannon–Weiner fungal species diversity index between habitats, seasons, dung ages and laboratory incubation periods, there were significant differences in fungal community composition between these parameters. Species richness was higher in the late wet season than in the early dry season, indicating the importance of moist conditions for a large diversity of fungal species.     

Habitat heterogeneity, richness and structure of assemblages of dung beetles (Scarabaeidae: Scarabaeinae) in areas of cerrado in the Chapada dos Parecis, Mato Grosso state, Brazil

Ecological theory of habitat heterogeneity and limited niche-similarity assumes that more heterogeneous environments provide a greater amount and diversity of resources than simple environments, resulting in a greater diversity of species. This study aimed to evaluate the effect of the habitat heterogeneity on the richness of dung beetles and to examine the spatial patterns of assemblage structure in relation to patterns of habitat heterogeneity. Dung beetles were collected using pitfall traps without bait in 30 points distributed in an area of cerrado sensu lato, in the region of Tangará da Serra, MT, Brazil, including areas of cerrado sensu stricto, campo sujo, cerrad?o and gallery forest. A total of 1,291 dung beetles were collected, distributed in 16 genera and 29 species. Overall habitat heterogeneity exerted a negative effect on patterns of dung beetles richness. Higher levels of species richness were observed in areas of cerrado campo sujo, while the areas of gallery forest were the most species poor. Regarding assembly structure, it was found that the dung beetles were separated into two major groups, one formed by the presence of specialized species in forest areas and other composed of species that occurred predominantly in cerrado. In conclusion, it was found that habitat complexity influenced the distribution of dung beetles, but the level of turnover in species composition along the heterogeneity gradient was relatively weak.     

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

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

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.