Dung Beetles: The Unsung Heroes of Ecosystem Health
Dung beetles are coprophagous insects that process vertebrate dung for feeding and nesting, and in doing so they drive multiple ecosystem functions including nutrient cycling, soil aeration, seed dispersal, and parasite suppression. This article explains the ecological roles of dung beetles, the traits that determine their functional contributions, and the practical implications for land managers and livestock producers. The content draws on peer-reviewed research published through the National Center for Biotechnology Information, PubMed, and related scientific journals.
At a Glance
Dung beetles are classified into three functional groups based on how they use dung. Tunnelers, also called paracoprids, dig beneath the dung pat and transport dung into underground chambers. Dwellers, or endocoprids, live and breed within the dung pat itself. Rollers, or telecoprids, shape dung into balls and roll them away from the source before burying them. Each strategy moves dung to a different location and depth, which influences the ecological outcomes.
| Functional Group | Nesting Strategy | Primary Ecosystem Contributions | Sensitivity to Disturbance |
|---|---|---|---|
| Tunnelers (paracoprids) | Bury dung in underground chambers beneath the pat | Nutrient transport to root zone, soil aeration, seed burial | Large tunnelers decline sharply with land use intensification |
| Dwellers (endocoprids) | Live and breed within the dung pat | Surface nutrient cycling, dung breakdown | Less affected by canopy removal but sensitive to dung contamination |
| Rollers (telecoprids) | Shape dung into balls and roll away before burying | Seed dispersal away from source, dung removal from surface | Small diurnal rollers may increase with land use intensity |
The functional ecology of dung beetles is best understood through a trait-based framework. A 2023 review in The Journal of Animal Ecology identified 66 dung beetle traits that function as response traits, effect traits, or both, across six categories: morphology, feeding, reproduction, physiology, activity, and movement. The review documented 136 trait-response relationships and 77 trait-effect relationships, showing that no single trait category explains how beetles respond to environmental stressors or affect ecological processes. Dung relocation behavior during nesting and feeding connects multiple trait categories, which highlights why species identity matters as much as species richness. See the full framework at PubMed.
The same review confirmed that dung beetles influence nutrient cycling, bioturbation, plant growth, seed dispersal, other dung-based organisms, and parasite transmission, with some cases of pollination and predation. These functions are interconnected. A field experiment using nitrogen-15 isotope tracing in alpine pastures demonstrated that dung beetles influenced at least seven ecological functions simultaneously, including dung removal, transport of dung-derived nitrogen into soil, microbial ammonification and nitrification, plant uptake of nitrogen, herbage growth, and changes in botanical composition. Tunnelers and dwellers performed most functions with similar efficiency, with differences based on spatial and temporal patterns. See the isotope tracing study at PubMed.
Nutrient Cycling and Soil Fertility
Dung beetles accelerate the decomposition of manure by breaking up dung pats and incorporating organic matter into the soil. This process moves nutrients from the surface into the root zone where plants can access them. The nitrogen-15 tracing experiment showed that dung beetles transported dung-derived nitrogen into soil up to 20 centimeters deep and that this nitrogen remained available in the system for up to one year after dung application. Plant uptake of dung-derived nitrogen increased in the presence of beetles, and herbage growth responded accordingly. See the full findings at PubMed.
The trait-based framework review confirmed that dung beetle effects on nutrient cycling are trait-mediated. Body size, dung relocation behavior, and nesting strategy determine how much dung is moved and to what depth. Large tunnelers that bury dung deep in the soil create different nutrient distribution patterns than dwellers that process dung near the surface. Land managers who want to maximize nutrient redistribution in pastures should recognize that a mix of functional groups provides more complete nutrient cycling than any single species.
Soil aeration is a related benefit. When tunnelers excavate burrows and move dung underground, they create channels that improve water infiltration and gas exchange. The same burrowing activity that transports dung also loosens compacted soil. The trait-based review identified bioturbation as one of the ecosystem processes influenced by dung beetles, with effects mediated by nesting behavior and body size. See PubMed.
Seed Dispersal and Plant Establishment
Dung beetles contribute to seed dispersal through two mechanisms. Primary dispersal occurs when beetles move dung containing seeds away from the original dung pat. Secondary dispersal occurs when seeds are buried along with the dung and later moved upward through beetle activity. A study in the Journal of Tropical Ecology documented upward movement of buried seeds by dung beetles, an additional ecological role that promotes seedling establishment. See the study at Cambridge Core.
The functional significance of seed burial depends on depth. Seeds buried too deep may fail to emerge, while seeds left on the surface are exposed to predators and desiccation. Dung beetles place seeds at intermediate depths that can be favorable for germination, depending on seed size and beetle body size. The trait-based framework review confirmed that seed dispersal is one of the ecosystem processes affected by dung beetle traits, particularly dung relocation behavior. See PubMed.
A study of cattle ranching duration in tropical dry forest landscapes in Mexico found that seed removal was mainly carried out by four beetle species, with the exotic species Digitonthophagus gazella being the most important. This finding shows that a small number of species can dominate seed removal functions and that species composition matters for ecosystem function. See the study at PubMed.
Parasite Control and Livestock Health
Dung beetles can reduce populations of gastrointestinal nematodes that affect livestock. The mechanisms include direct damage to nematode eggs and larvae during ingestion, increased aeration and desiccation of the dung pat, and movement of dung deeper into the soil where larvae cannot complete their development. A 2025 study in Veterinary Parasitology investigated the impact of the tunnel beetle Copris incertus on gastrointestinal nematodes in New Zealand pastures. The study applied treatments with different beetle densities across spring, summer, and autumn. In spring and autumn, the highest beetle abundances increased dung removal from the soil surface. However, the study found no evidence that Copris incertus reduced nematode larval numbers on pasture, regardless of beetle densities. In spring, increased dung removal was associated with more buried dung balls and an increase in infective third-stage larvae in the herbage in the high beetle treatment. See the study at ScienceDirect.
This finding matters for livestock producers. Dung beetles are often promoted as a biological control for internal parasites, but the evidence is mixed. The New Zealand study shows that beetle activity can redistribute dung and larvae in ways that do not consistently reduce larval exposure. Producers should not rely on dung beetles alone for parasite control. Integrated parasite management that includes pasture rotation, targeted deworming, and fecal monitoring remains necessary.
A related study examined how dung burial by coprophagous beetles modifies the emergence of infectious strongyle nematode larvae across different soil depths. The study is indexed at Europe PMC. The title indicates that burial depth is a key variable in determining whether nematode larvae can return to the soil surface and infect grazing animals.
Soil Structure and Aeration
The burrowing activity of tunnelers creates macropores that improve soil structure. These channels increase water infiltration, reduce surface runoff, and promote root penetration. The trait-based framework review identified bioturbation as a core ecosystem process influenced by dung beetles, with effects mediated by body size and nesting behavior. See PubMed.
Soil aeration benefits pasture productivity by improving oxygen availability in the root zone and supporting microbial activity. The nitrogen-15 tracing experiment showed that dung beetles facilitated microbial ammonification and nitrification processes in the soil, which are oxygen-dependent transformations. See PubMed.
Land managers can assess soil structure improvements by comparing infiltration rates and soil compaction measurements in areas with active dung beetle populations versus areas where beetles are absent. These measurements provide direct evidence of the physical effects of beetle activity.
Dung Beetle Diversity and Community Composition
Dung beetle diversity varies across landscapes and is influenced by climate, habitat structure, and land use history. A macroecological study of dung beetles in Italy found that the fauna includes about 170 species and subspecies, one of the richest in Europe. Scarabaeinae species followed a latitudinal gradient, supporting a possible role of southern areas as Pleistocene refuges for thermophilic beetles. Aphodiines were more associated with cold and humid climates and did not show a distinct latitudinal pattern. Species richness was influenced by area, with the Sardinian fauna strongly impoverished because of isolation. See the study at PubMed.
Habitat structure strongly influences dung beetle communities. A study in tropical dry forests of the Yucatán Peninsula assessed dung beetle communities across a forest recovery chronosequence from 1 to 100 years. The study collected 6,605 individuals from 23 species and 13 genera using 90 pitfall traps. Species richness, biomass, and abundance were significantly associated with forest structural and diversity metrics. Forest Shannon entropy, inverse Simpson concentration, and aboveground biomass emerged as strong predictors of community attributes. Functional group responses varied: small diurnal rollers increased with land-use intensity, while large nocturnal rollers, large diurnal tunnelers, and small nocturnal tunnelers declined sharply from mature forests to early successional stages and agricultural areas. Species richness peaked in early to intermediate successional stages of 5 to 20 years, while dominant species diversity was highest in mixed-use forests under moderate disturbance. Forest attributes including diameter at breast height, aboveground biomass, canopy openness, and litter depth jointly explained 48.7 percent of the variation in dung beetle assemblage structure. See the study at PubMed.
The conversion of natural habitats to agriculture has consistent negative effects on dung beetle communities. A study in the Brazilian Cerrado compared dung beetle assemblages in native savanna, organic agriculture, and conventional agriculture. The amount of savanna formations was the most important landscape type supporting dung beetles and their ecosystem services. Organic farming increased habitat permeability to dung beetle species, while conventional farming negatively affected them. Species richness mediated functional diversity and ecosystem services locally. See the study at Europe PMC.
Another Cerrado study found that complete conversion of native savanna to exotic pasture had a negative effect on dung beetle assemblages. The same dung types differed completely in dung beetle assemblages in native savannas and exotic pasture regarding richness, abundance, species composition, and dominance patterns. Environmental filters in exotic pastures, including low canopy cover and simplified herbaceous complexity, likely restricted the establishment of species from the Cerrado. The study recommended increasing herbaceous complexity and canopy cover along with diversifying livestock in exotic pastures to avoid the loss of dung beetle species and their associated ecosystem services. See the study at ScienceDirect.
Dung Type Preferences and Functional Traits
Dung beetles are not equally attracted to all dung types. A study in the journal Life examined dung beetle communities attracted to horse and cow dung from a functional diversity standpoint. The study examined 18 functional traits and found species-specific dung preferences for eight species, with two preferring horse dung and six preferring cow dung. Significant differences were found in the mouthpart traits of beetles attracted to horse dung versus cow dung. Specifically, zygum development and the percentage of the molar area and the conjunctive area differed between the two groups. The authors proposed that quantitative differences in mouthpart traits relate to the differential capacity of beetles to filtrate and concentrate small particles from dung. Larger and nesting beetles preferred cow dung, while smaller and non-nesting beetles preferred horse dung. See the study at MDPI.
This research has practical implications for livestock operations. Producers with cattle and horses on the same property may support different dung beetle communities. Diversifying livestock types can increase the range of dung resources available and support a more diverse beetle assemblage.
Threats to Dung Beetle Populations
Veterinary Pharmaceutical Residues
Ivermectin is the most common antiparasitic medication used in cattle in some regions, despite residues excreted in dung being highly toxic for non-target coprophagous insects. A study of the dung beetle Euoniticellus intermedius evaluated ivermectin tolerance in beetles from six sites along an elevational gradient. Large variation among sites was observed in median lethal concentration, with tolerance more than 200 times higher in the most tolerant site compared to the least tolerant site. At high concentrations, nearly zero eggs emerged in the most sensitive populations. No elevational pattern was detected in ivermectin tolerance, indicating that factors other than elevation drive susceptibility. The study confirmed that ivermectin tolerance had associated costs in the most tolerant population, revealed by a trade-off between offspring number and size that was exaggerated with increasing ivermectin concentration. See the study at ScienceDirect.
A related study examined whether ivermectin exposure affects thermal tolerance, thermoregulation, and energetic status in Euoniticellus intermedius. Ivermectin unexpectedly increased heat tolerance and improved cold tolerance at intermediate elevation, indicating a hormetic response. Contrastingly, ivermectin reduced cold tolerance at high elevation, revealing a physiological cost of colonizing high elevations. Lipid content was higher at high ivermectin concentrations and highest at low elevations. Ivermectin did not affect thermoregulatory capacity. See the study at ScienceDirect.
A study in the Journal of Insect Conservation examined how veterinary medical products disrupt ecological processes. The title indicates that functional identity, not functional diversity, accounts for ecological process disruption caused by veterinary medical products. See the study at Springer. This finding suggests that the loss of specific functional groups, particularly large tunnelers, has greater ecological consequences than the loss of overall species richness.
Microplastic Contamination
Microplastics are an emerging environmental hazard in agricultural environments. A study tested whether artificial contamination of cow dung with thermoplastic polyurethane microplastics affects juvenile development and maternal behavior in the bull-headed dung beetle Onthophagus taurus. Larvae exposed to dung containing 0.5 milligrams per gram or more of thermoplastic polyurethane microplastics experienced high mortality, while exposure to 0.1 milligrams per gram did not significantly increase mortality risk relative to controls. Adult females were equally likely to provision offspring with contaminated dung as with uncontaminated control dung, suggesting that females cannot differentiate between highly toxic microplastic-contaminated and uncontaminated resources. See the study at ScienceDirect.
Habitat Fragmentation and Land Use Change
Forest management practices affect dung beetle populations. A full-factorial forest experiment in Germany tested how gaps and deadwood influence dung removal. The experiment was established at 29 sites in three regions with treatments including Gap, Gap plus Deadwood, Deadwood, and Control. All gaps were experimentally created with a diameter of around 30 meters. Dung beetle diversity, biomass, and dung removal were each lower in gaps than in controls. Dung removal decreased from 61.9 percent in controls to 48.5 percent in gaps, irrespective of whether the gap had deadwood. This treatment effect was primarily driven by dung beetle biomass but not diversity. Dung removal was reduced to 56.9 percent in the deadwood treatment. Gaps reduced the abundance of a large-bodied key forest species. See the study at PubMed.
A related study in German production forests found that more heterogeneously managed forests did not have higher dung beetle species diversity or dung removal rates. Canopy openings did not increase species turnover but decreased species diversity. Along the climate gradient, dung beetle average biomass and dung removal decreased with increasing temperature. Canopy openings in combination with higher temperatures negatively impacted all abundant dung beetle species, especially the large species Anoplotrupes stercorosus, which comprised more than 90 percent of total dung beetle biomass. See the study at Europe PMC.
Dung Beetles as Bioindicators
Dung beetles are effective bioindicators of ecosystem health because they respond quickly to habitat change and land use intensity. Their sensitivity to vegetation structure, soil conditions, and management practices makes them useful for monitoring restoration success. A study in the Revista Mexicana de Biodiversidad examined the role of ecological restoration on the recovery of dung beetle diversity and function in a tropical rainforest. See the bibliographic record at Elsevier.
The chronosequence study in the Yucatán Peninsula demonstrated that dung beetle community attributes track forest recovery. Species richness peaked in early to intermediate successional stages of 5 to 20 years, while dominant species diversity was highest in mixed-use forests under moderate disturbance. Litter volume was positively correlated with species richness. See PubMed.
Land managers can use dung beetles as a monitoring tool by conducting standardized pitfall trapping across seasons and comparing species composition and functional group representation against reference sites. Changes in the abundance of large tunnelers or rollers can signal habitat degradation before other indicators respond.
Practical Assessment of Dung Beetle Activity
Land managers can assess dung beetle activity using standardized methods. Pitfall trapping with dung baits provides data on species presence, abundance, and functional group composition. Dung pat observation provides a simpler assessment of activity. Fresh dung pats can be marked and monitored for signs of beetle activity, including tunnels beneath the pat, dung removal, and the presence of beetles within the pat.
A practical assessment protocol includes the following steps. First, select representative pasture areas that differ in management history, livestock type, or habitat context. Second, mark fresh dung pats and record their condition at 24, 48, and 72 hours after deposition. Third, record the presence of tunnels, the percentage of dung removed, and the presence of adult beetles. Fourth, conduct pitfall trapping during the active season to identify species and functional groups. Fifth, compare results across sites and seasons to identify patterns.
Records should include the date, location, weather conditions, livestock type, dung age, and management history for each assessment. This information allows land managers to track changes over time and correlate beetle activity with management practices.
Common Failure Patterns in Dung Beetle Conservation
Several management practices consistently reduce dung beetle populations and their ecosystem functions. The use of persistent antiparasitic drugs, particularly ivermectin, can eliminate sensitive species and reduce dung processing. The study of Euoniticellus intermedius showed that ivermectin tolerance varies more than 200-fold among geographically close populations, meaning that some populations may persist while others collapse. See ScienceDirect.
Habitat simplification is another common failure pattern. The conversion of native vegetation to monoculture pasture removes the structural complexity that many dung beetle species require. The Cerrado studies showed that exotic pastures support different and generally impoverished beetle communities compared to native savanna. See ScienceDirect and Europe PMC.
Canopy removal in forests reduces dung beetle biomass and dung removal rates. The German forest experiment showed that gaps reduced dung removal from 61.9 percent to 48.5 percent, with effects driven primarily by loss of large-bodied species. See PubMed.
Microplastic contamination of dung is an emerging threat. Female beetles cannot distinguish between contaminated and uncontaminated dung, so they may provision offspring with lethal resources. See ScienceDirect.
Management Recommendations for Land Managers
Land managers can support dung beetle populations through several practices. Reducing or eliminating the use of persistent antiparasitic drugs, particularly ivermectin, is the most direct intervention. When treatment is necessary, managers should consider the timing relative to dung beetle activity and use products with shorter environmental persistence.
Maintaining habitat heterogeneity supports diverse beetle communities. In pasture systems, this includes retaining native vegetation patches, maintaining riparian buffers, and avoiding complete conversion of natural habitats. The Cerrado study recommended promoting low-intensity agricultural practices such as organic farming combined with the conservation and restoration of natural savanna patches. See Europe PMC.
Diversifying livestock types can support a broader range of dung beetle species. The study of horse and cow dung preferences showed that different species are attracted to different dung types, and mouthpart traits determine the capacity to exploit specific dung resources. See MDPI.
Avoiding unnecessary soil disturbance preserves beetle burrows and overwintering sites. Tillage and heavy grazing can destroy underground nesting chambers and reduce beetle survival.
Limitations of Current Knowledge
Several important knowledge gaps remain. The New Zealand study of Copris incertus found no evidence that this species reduced nematode larval numbers on pasture, despite increased dung removal. This finding contrasts with the common assumption that dung beetles control parasites. See ScienceDirect. The relationship between dung beetle activity and parasite transmission is context dependent and requires further research.
The effects of microplastic contamination on dung beetles are not fully understood. The thermoplastic polyurethane study noted that environmentally relevant exposure levels remain unknown and that future work should quantify microplastic concentrations in the field, test the effects of microplastic composition and size distribution, and identify the mechanisms underlying toxicity. See ScienceDirect.
The trait-based framework review identified the need for consistent trait measurement protocols across studies. The review noted that trait approaches to functional ecology are well established for plants, whereas consistent frameworks for animal groups are less developed. See PubMed.
Professional Escalation Criteria
Land managers should seek professional advice when they observe specific warning signs. A complete absence of dung beetle activity in areas that previously supported beetles may indicate contamination or habitat degradation that requires investigation. Sudden declines in dung removal rates may signal the introduction of a persistent antiparasitic drug or an environmental contaminant. The presence of dung pats that remain intact for weeks without signs of beetle activity warrants assessment of management practices.
Veterinarians and agricultural advisors should be consulted when antiparasitic treatment protocols may be affecting dung beetle populations. The high variability in ivermectin tolerance among populations means that local conditions determine risk. See ScienceDirect.
Researchers and conservation agencies should be engaged when land use changes threaten dung beetle diversity. The chronosequence study in the Yucatán Peninsula showed that functional group responses vary by land use intensity, with large nocturnal rollers and large diurnal tunnelers declining sharply from mature forests to early successional stages. See PubMed.
Frequently Asked Questions
What ecosystem services do dung beetles provide?
Dung beetles provide nutrient cycling, soil aeration, seed dispersal, and parasite suppression. The trait-based framework review confirmed that dung beetles influence nutrient cycling, bioturbation, plant growth, seed dispersal, other dung-based organisms, and parasite transmission. See PubMed. The nitrogen-15 tracing experiment showed that dung beetles influenced at least seven ecological functions simultaneously. See PubMed.
How do dung beetles improve soil fertility?
Dung beetles move dung from the soil surface into underground chambers, transporting nutrients into the root zone. The nitrogen-15 tracing experiment showed that dung beetles transported dung-derived nitrogen into soil up to 20 centimeters deep and that this nitrogen remained available for up to one year. Plant uptake of dung-derived nitrogen increased in the presence of beetles. See PubMed.
Do dung beetles control livestock parasites?
The evidence is mixed. A 2025 study of Copris incertus in New Zealand found no evidence that this species reduced nematode larval numbers on pasture, regardless of beetle densities. In spring, increased dung removal was associated with an increase in infective larvae in the herbage in the high beetle treatment. See ScienceDirect. Dung beetles should not be relied upon as the sole method of parasite control.
How do dung beetles disperse seeds?
Dung beetles disperse seeds by moving dung containing seeds away from the original pat and by burying seeds along with dung. A study in the Journal of Tropical Ecology documented upward movement of buried seeds by dung beetles, promoting seedling establishment. See Cambridge Core.
What threatens dung beetle populations?
The main threats are veterinary pharmaceutical residues, particularly ivermectin, habitat fragmentation, land use change, and microplastic contamination. Ivermectin tolerance varies more than 200-fold among populations of Euoniticellus intermedius. See ScienceDirect. Microplastic contamination of dung at 0.5 milligrams per gram caused high larval mortality in Onthophagus taurus. See ScienceDirect.
How can farmers support dung beetle populations?
Farmers can reduce or eliminate persistent antiparasitic drugs, maintain native vegetation patches, diversify livestock types, and avoid unnecessary soil disturbance. The Cerrado study recommended promoting low-intensity agricultural practices such as organic farming combined with conservation of natural savanna patches. See Europe PMC.
Why do dung beetle communities differ between habitats?
Dung beetle communities respond to vegetation structure, soil properties, climate, and food resources. The chronosequence study in the Yucatán Peninsula found that forest attributes including diameter at breast height, aboveground biomass, canopy openness, and litter depth jointly explained 48.7 percent of the variation in dung beetle assemblage structure. See PubMed.
Are all dung beetles equally important for ecosystem function?
No. Functional identity matters more than functional diversity for some ecological processes. A study in the Journal of Insect Conservation found that functional identity, not functional diversity, accounts for ecological process disruption caused by veterinary medical products. See Springer. Large-bodied species often have disproportionate effects on dung removal and nutrient cycling.
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References and Further Reading
- NCBI Literature Resources. National Center for Biotechnology Information.
- PubMed. National Library of Medicine.
- A trait-based framework for dung beetle functional ecology.. The Journal of animal ecology, 2023.
- Developmental and Ecological Benefits of the Maternally Transmitted Microbiota in a Dung Beetle.. The American naturalist, 2016.
- Macroecology of Dung Beetles in Italy.. Insects, 2024.
- Gut microbiota diversity in a dung beetle (Catharsius molossus) across geographical variations and brood ball-mediated microbial transmission.. PloS one, 2024.
- Dung beetle assemblage changes along a chronosequence in a recovering tropical dry forest.. PloS one, 2025.
- Ecological functions provided by dung beetles are interlinked across space and time: evidence from (15) N isotope tracing.. Ecology, 2017.
- Duration of Cattle Ranching Affects Dung Beetle Diversity and Secondary Seed Removal in Tropical Dry Forest Landscapes.. Insects, 2024.
- Negative effects of forest gaps on dung removal in a full-factorial experiment.. The Journal of animal ecology, 2022.
- Ivermectin tolerance and associated costs in the dung beetle Euoniticellus intermedius.. 2026.
- The potential impact of microplastic contamination in cow manure on maternal dung burying behavior and larval survival in the dung beetle Onthophagus taurus.. 2026.
- Pollution constrains thermal adaptation but not thermoregulation in dung beetles.. 2026.
- How does dung burial by coprophagous beetles modify the emergence of infectious strongyle nematode larvae? An experimental test across different soil depths. 2026.
- The role of natural savanna in maintaining dung beetle diversity and their ecosystem services in agricultural landscapes in the Brazilian Cerrado. 2026.
- Dung beetles do not profit from enhanced spatial heterogeneity in temperate production forests: A forest manipulation experiment. 2026.
- Dung Beetle Assemblages Attracted to Cow and Horse Dung: The Importance of Mouthpart Traits, Body Size, and Nesting Behavior in the Community Assembly Process. Life, 2021.
- Conversion of Cerrado savannas into exotic pastures: The relative importance of vegetation and food resources for dung beetle assemblages. 2020.
- The influence of the dung beetle Copris incertus on dung removal and gastrointestinal nematode density on pasture.. Veterinary parasitology, 2025.
- Deep deterministic policy gradient algorithm based on dung beetle optimization and priority experience replay mechanism. Scientific Reports, 2025.
- Success or failure: The role of ecological restoration on the recovery of dung beetle diversity and function in a tropical rainforest. Revista Mexicana De Biodiversidad, 2018.
- Upward movement of buried seeds: Another ecological role of dung beetles promoting seedling establishment. Journal of Tropical Ecology, 2014.
- Dung beetles: functional identity, not functional diversity, accounts for ecological process disruption caused by the use of veterinary medical products. Journal of Insect Conservation, 2020.
- Ecological functions and ecosystem services provided by Scarabaeinae dung beetles. Biological Conservation, 2008.
This article is educational and does not replace institutional policy, professional advice, or applicable safety and regulatory requirements.