INTRODUCTION

Dung beetles are well-known taxonomically and ecologically (Barragán et al., 2021). They belong to the subfamily Scabaeinae, comprising approximately 20 tribes, nine subtribes, 270 genera, 7,000 living species, and 23 fossil species (Tello et al., 2021, 2023; Daniel & Davis, 2024). In Argentina, about 247 species have been reported (Cupello et al., 2023). Regarding their ecology, these insects feed on the dung of mammals, mainly (native and exotic), and decaying organic matter (fungi, fruits, meat, among others), burying a variable number of masses or balls as provisions for their larvae or feeding (Simmons & Ridsdill-Smith, 2011). This remarkable characteristic of food relocation evolved as a response to intra- and interspecific competition for an ephemeral resource (Arellano et al., 2023). Due to this burial process, dung beetles are associated with important ecological functions, including the degradation of organic matter, secondary seed dispersal, and bioturbation, among others (Braga et al., 2013; Noriega et al., 2021; Barragán et al., 2022).

Dung beetles serve as effective bioindicators for assesing anthropogenic impacts on biodiversity since they are sensitive to changes in microclimate conditions, availability of organic matter, and physicochemical properties of the soil, which are key factors for their development (Nichols et al., 2007; Otavo et al., 2013). Exploring the diversity and distribution of dung beetles is necessary to understand the mechanisms involved in the loss (Gómez-Cifuentes et al., 2017), replacement (Guerra Alonso et al., 2022), and repopulation of species (Bobadilla et al., 2024).

There are numerous reviews in the literature on the distribution of these species at global (Davis & Scholtz, 2001) and continental scales (Cupello et al., 2023). Additionally, studies have focused on forest and savanna diversity (Nichols et al., 2007, Reis et al., 2024) and on the effects of cattle ranching on their diversity and functioning (Arellano et al., 2023); however, many of these studies have been carried out mainly in tropical countries. According to the study of Arellano et al. (2023), three countries in the Neotropics have the highest number of studies with dung beetles (Brazil, Mexico, and Colombia). The authors conclude that the lack of funding in the rest of the countries creates a critical information gap, which translates into a poor understanding of the diversity and distribution of these insects within each country, particularly in those located in the subtropical and temperate regions of the American continent.

In Argentina, studies focused on dung beetles seem to be relatively low. In this review, we compiled an in-depth list of all studies conducted in the Argentine territory, including those that report species stored in national collections. Based on the above, we examine the temporal and spatial trends in Argentine dung beetle research and identify knowledge gaps to propose future research priorities.

MATERIALS AND METHODS

We used two databases for the literature search: Google Scholar and Web of Science. In both, we included keywords in English and Spanish, as Argentina is a Spanish-speaking country, and some works may have been published in Spanish or included abstracts and keywords in this language. The following terms were used to search for information: (Dung beetles OR Scarabaeidae OR Scarabaeinae AND Argentina) and (estercoleros OR coprófagos OR copronecrofagos OR peloteros AND Argentina) in Spanish. In each search, we exclude the countries with the highest number of studies in the Neotropics (Arellano et al., 2023): "-Colombia", "-México", "-Mexico", "-Brasil", and "-Brazil". In addition, we conducted a specific search in all issues published by the Journal of the Sociedad Entomologica Argentina, as well as in Google and Google Scholar, to find as many studies as possible published by Dr. Antonio Martínez, one of the most notable entomologists and probably the leading specialist in the Scarabaeidae family in Argentina. It is worth clarifying that we considered in this review some articles published in journals not indexed in these platforms, but with peer review (Table S1 – DATA STATEMENT).

From the papers collected, we decided to extract the following information:

(i) Temporal variation: refers to the number of studies published by year (Table S1 – DATA STATEMENT).

(ii) Spatial variation: refers to the ecoregions and provinces in which these studies were performed based on the Biodiversity Information System (https://sib.gob.ar) and the Official Portal of the Argentine State (https://www.argentina.gob.ar/pais), respectively. In some cases, a study included two or more ecoregions or provinces. We used Google Earth Pro to find the specific location reported in each study (such as geographic coordinates or names); we also used a KMZ file with the ecoregions of Argentina for greater precision.

(iii) Research topic: refers to the main line of research in which each study was developed. For this, we used two artificial intelligence (AI) sources (ChatGPT and Sider) to identify the main research areas and then classified each study into one of these areas (Table S2 -DATA STATEMENT). We want to clarify that the use of AI was supported by our criteria and experience working with dung beetles.

(iv) Data analyzed: refers to the variables explored, evaluated, and analyzed in the different studies to test the objectives and hypotheses of the work (Table S3 – DATA STATEMENT). We divided these variables into four large groups: community, ecological, physiological, and evolutionary metrics. In some cases, a single study analyzed two or more metrics.

(v) Species reported in Argentina: refers to the number of dung beetles captured or reported in each Argentine province. In some cases, one study reported two or more species in one or more provinces. In addition, we explored the species reported by geographic region: northeast, northeast, Pampas, Cuyo, and Patagonia (Table S4 – DATA STATEMENT). We would like to clarify that this is not a measure of species richness.

RESULTS

We found a total of 81 studies, with most written in English, while Spanish was the second language (80 % and 20 %, respectively) (Table S1 – DATA STATEMENT). Regarding gender balance, men were the first authors in 56 % of the studies, while women were the first authors in 44 % (Table S1 – DATA STATEMENT).

Temporal variation

The first study in English or Spanish with dung beetles in Argentina was published in 1944 in the Revista de la Sociedad Entomológica Argentina, while the last published paper, at the end of this review (June 2025), was published in 2025 in the journal Scientific Reports (Table S1 – DATA STATEMENT). Most studies were published in the 2010s (n= 29) (Fig. 1); 25 % of those papers were published in 2019 (Table S1 – DATA STATEMENT). It is important to note that there are likely more studies in other languages and grey literature, such as graduate and postgraduate theses, non-indexed journals, bulletins, reports, and other sources that were not consulted in this review, which may explain this result (Fig. 1).

Figure 1.
Number of studies with dung beetles in Argentina per decade.

Spatial variation

The ecoregion with the most studies published in Argentina was the Paranaense rainforest (n= 17), while the High Andes and Iberá Wetlands had only one study each. Furthermore, six ecoregions in Argentina have not been explored yet (Fig. 2). Regarding the studies with dung beetles per province, Misiones had the highest number of studies published (n= 19), while Jujuy, San Juan, Santa Cruz, San Luis and Neuquén had only one each; moreover, eight provinces had no studies with dung beetles (Fig. 2).

We want to clarify that 29 studies were excluded from this section because they did not report a specific geographic location.

Figure 2.
Number of studies with dung beetles in Argentina.

a. Ecoregions. b. Provinces. In some cases, a single study encompasses one or more ecoregions.

Research areas

Based on the AI complemented with our criteria and experience resulting from our extensive work with dung beetles, we categorized the 81 studies into seven primary research areas, each encompassing multiple sub-areas (Table S2 – DATA STATEMENT). Based on this, 28 % of the analyzed studies explored the behaviour and evolution of dung beetles. The main focuses of this research area were the past and present of different food relocation strategies, as well as reproductive and mating behaviour (Fig. 3). Furthermore, 27 % were on taxonomy, systematics, and phylogeny (mainly focused on the diagnosis, description, and classification of species), 21 % of the studies were on biodiversity and conservation (loss, fragmentation, and replacement of natural environments due to different land uses), and 11 % dealt with biogeography and species distribution (primarily associated with sub-areas such as paleontology and ichnology) (Fig. 3). The least explored research areas were the ecological function and physiology (6 %), while OR whereas the impact of livestock management (effect of antiparasitic agents, the intensity and management of grazing areas, and the impact of pollution on dung beetles) was explored in a single study (Fig. 3). We want to clarify that several studies related to Biodiversity and Conservation analyzed the effects of livestock on dung beetles. Still, none of them directly evaluated livestock management, as they only explored changes in the microclimate (an indirect variable), which is why they were not considered in the area of livestock management (Table S2 – DATA STATEMENT).

Data Analyzed

Based on the studies found, we categorized the analyzed metrics into four broad areas: community (33 studies), ecological (45 studies), physiological (8 studies), and evolutionary (10 studies) (Table S3 – DATA STATEMENT). Fifteen studies analyzed metrics from two broad areas simultaneously: six included community and physiological metrics, five included community and ecological metrics, and one included community and evolutionary metrics. Furthermore, two studies analyzed ecological and evolutionary metrics simultaneously, and one also analyzed ecological and physiological metrics (Table S3 – DATA STATEMENT).

Figure 3
Number of studies on dung beetles in Argentina by research topic.

Among the explored community metrics, the variables analyzed were associated with species taxonomic diversity in 46 studies, species arrangement and similarity in 16, functional trait diversity in 17, and dung beetle biomass in 11 (Fig. 4a). Species richness and abundance were the most frequently analyzed response variables in all the studies explored, being assessed in 20 % of all reviewed papers (Table S3 – DATA STATEMENT). Furthermore, in 12 studies, species richness and abundance were correlated (or at least analyzed in parallel) with response variables associated with beta diversity (multivariate analyses and similarity indices), in six studies with functional diversity (groups and multidimensional index), and six studies with biomass, primarily the fresh weight of dung beetles (Table S3 – DATA STATEMENT).

Regarding the ecological metrics, 23 studies explored the description and diagnosis of species (mostly in relation to taxonomic revisions), 22 studies assessed the behavior of dung beetles through field observations and laboratory experiments (nesting, food relocation, bioacoustics, and biological interactions), and eight analyzed the ecological functions of dung beetles (Fig. 4b); besides, in eight studies, two response variables were simultaneously evaluated (Table S3 – DATA STATEMENT). Among studies examining ecological functions, three assessed nutrient cycling changes, three evaluated soil removal, and two investigated seed removal (Fig. 4b). Furthermore, one of these studies evaluated soil removal together with nutrient cycling, and another evaluated soil removal with seed removal (Table S3 – DATA STATEMENT). On the other hand, two of these studies were conducted through field experiments with native beetles found in the evaluated environments and under natural temperature and humidity conditions, whereas, four studies were conducted with native soils and beetles collected in the field, but under controlled temperature and humidity conditions (Table S3 – DATA STATEMENT).

Regarding physiological metrics, six studies evaluated changes associated with physiology, with body temperature and thermal amplitude being the most frequently assessed metrics. Moreover, endothermy, thermoregulation, thermal niche, thermal limits, and metabolic rate were the other variables explored in these studies (Fig. 4c). We found two works related with chemoecological metrics, in the first one, the authors explored several chemical components associated with different trophic resources to assess their attraction potential and trophic preference for dung beetles; while, in the second one, the authors analyzed some chemical components of the body of the Canthon chalybaeus Blanchard as an attack mechanism to prey on snails of the species Bulimulus apodemetes (D’Orbigny) (Fig. 4c, Table S3 – DATA STATEMENT).

Regarding the evolutionary metrics, seven studies analyzed the qualitative and quantitative structure of fossilized nesting balls and chambers, where the phytoliths were the most explored response variable (Fig. 4d). On the other hand, three studies focused on phylogenetic variables were found; two of them analyzed the phylogenetic relationships of the species based on morphological and molecular traits of adult beetles, while the other one, explored the phylogenetic relationships associated with the macro- and micromorphological structure of the brood chambers found within the nesting balls (Table S3 – DATA STATEMENT).

Figure 4.
Number of studies with dung beetles in Argentina per metrics.

a. Community metrics. b. Ecological metrics. c. Physiological metrics. d. Evolutionary metrics.

Species reported in Argentina

The Paranaense rainforest was the ecoregion with the highest number of species reported in Argentina, followed by the Humid and Dry Chaco (Fig. 5). In contrast, the Puna had the lowest record with one species (Fig. 5). Regarding the provinces, Misiones had the highest number of records with a total of 127 species, followed by Chaco with 79 species (Fig. 5). In contrast, Santa Cruz had the lowest number of records with four species (Fig. 5). On the other hand, in Tierra del Fuego and Islas Malvinas no dung beetle species have been reported (Fig. 5). It is important to note that our results do not represent the actual species richness by province, only the species reported in the literature.

Figure 5
Number of species recorded in Argentina.

a. Ecoregions. b. Provinces.

Regarding geographic regions, the north had the highest number of species reported in Argentina, with 166 species in the northeast and 106 in the northwest; in contrast, the south (Patagonia) had the lowest species richness in Argentina (15 species) (Table S4 – DATA STATEMENT). Furthermore, the Pampa and Cuyo regions had intermediate levels, with 85 and 36 species respectively (Table S4 – DATA STATEMENT).

DISCUSSION

Temporal and spatial advances

Under the search parameters established in this review, the first study focused on dung beetles in Argentina was published 85 years ago by A. Martínez (1944), entitled "Insectos nuevos o poco conocidos (Coleoptera)," and was published in the Revista de la Sociedad Entomológica Argentina (Journal of the Argentine Entomological Society). However, it is worth noting that 45 years prior (1899), F. Judulien published a study in French entitled "Quelques notes sur plusieurs coprophages de Buenos Aires" in the Revista del Museo de la Plata, which probably means that it was the first paper published on this subject in the history of Argentina. On the other hand, it was at the beginning of the 21st century when the number of studies on dung beetles grew exponentially and continued to grow until the current decade. The number of studies doubled from 2000 to 2010. In only four and a half years of the current decade (2020-2025), more than half of the articles published in the previous decade have already been published. This growth is probably associated with the increase in research groups in the country (more researchers and students) and the rise in GDP in science and technology, which went from 0.17 % in 2002 to 0.30 % in 2023, with a peak investment of 0.35 % in 2014 (EPC (Economía-Política-Ciencia), 2025a). While the current outlook is somewhat discouraging due to the lack of investment, with GDP in science and technology declining in the last year (EPC, 2025b), we believe that the number of studies in the current decade will continue to increase.

Regarding spatial variation, we found that the highest number of dung beetle studies were published in the Paranaense rainforest ecoregion. Additionally, half or fewer of these studies were conducted in five ecoregions, while no studies have been conducted yet in 37 % of the Argentinian ecoregions. The lack of publications in the Yungas Forests, Fields, and Scrublands draws attention, as microclimatic conditions and mammal composition should not restrict the distribution of these species, which could explain the lack of studies in the southern provinces of the country. The provincial panorama is very similar to that observed along the ecoregions; the highest number of studies have been carried out in Misiones, while one to eight studies have been performed in 14 other provinces. On the other hand, of the eight provinces without studies on dung beetles, it is worth mentioning Tucumán and Santa Fe, as these provinces have large technological and scientific centers and diverse ecoregions, including the Yungas Forests, the Dry and Humid Chaco, the Espinal, and the Pampa.

Research trends: gaps and opportunities

The most extensively explored research area in Argentina has been the behaviour of dung beetles, with nesting and food relocation strategies being the most analyzed metrics. These variables have been a determining factor in the diversification of this group (Simmons & Ridsdill-Smith, 2011). On the other hand, it is worth highlighting those studies that explored novel ecological variables, such as bioacoustics in larvae (Sánchez & Barria, 2023), the predation of some adult beetles (Martín et al., 2021), and physiological variables as volatile substances in the excrement that influence the attraction and trophic preference of these insects (Wurmitzer et al., 2017); consequently, these studies help identify potential research lines to address knowledge gaps in these unexplored areas.

Another main research area explored in Argentina has been the Taxonomy, Systematics, and Phylogeny of dung beetles, which represent 27 % of the papers compiled in this review. The main results included diagnoses, descriptions, and analyses of the evolutionary relationships of these organisms. Most studies have focused on the tribe Eucraniini, endemic to Argentina’s desert thorn ecosystems, whose species exhibit specialized ecological adaptations and unique behaviors (Ocampo, 2004). On the other hand, studies on the biogeography and species distribution, were carried out primarily in the south of the country, in relation to terrestrial mammals fossils and the evaluation of phytoliths of the ichnogenus Coprinisphaera Sauer, one of the most common trace fossils of South American Cenozoic paleosols (Cantil et al., 2013). The studies, conducted during the 2010s, have a significant value in understanding the evolutionary processes of dung beetles, as well as the grass-dominated ecosystems and the herbivores that inhabit them (Sánchez et al., 2010). In this research area, it is worth highlighting the studies conducted by Álvarez-Bohle et al. (2009) and Guerra Alonso et al. (2022), which report and update the records and distribution of Digitonthophagus gazella F., the only exotic dung beetle found in Argentina. This species has a broad thermal tolerance, with high dispersal and reproduction rates that allow it to rapidly colonize grassland ecosystems and non-forest livestock systems, making it the dominant species in a very short time as it displaces native species (Noriega et al., 2020); furthermore, if we take into account that in a scenario of direct competition between invasive and native exotic species, the latter may end up becoming locally extinct and that in the current context of accelerated climate change, these colonization processes by exotic species could allow their distribution to reach temperate ecoregions such as the Pampas, the increase in this type of studies is key to fill the current information gaps to understand this problem and to conserve native species (Cardoso et al., 2020; Noriega et al., 2020).

The Biodiversity and Conservation of dung beetles research area represents 21 % of the studies compiled in this review, with the particularity that all of them were conducted in the northeast of the country, except for the work of Brewer et al. (1981), which was carried out in the province of Córdoba. Regarding the variables analyzed in this research area, most studies have explored species richness, abundance, and composition, as well as their functional diversity based on traits, groups, and indices. In the same order, biomass was one of the most evaluated variables in the country, a variable closely related to the functioning of dung beetles, as it allows inferring the quantity, depth, and distance at which these insects bury excrement (Horgan, 2005; Gómez-Cifuentes et al., 2020). Another particularity observed in this area was that 80 % of the studies explored the degradation of native forests by different livestock systems; however, most of them indirectly evaluated the effect of livestock through environmental variables such as vegetation composition, canopy cover, microclimatic and soil properties, without considering livestock management variables, which is why in this review none of these studies was classified in this line of research. Based on the above, Arellano et al. (2023) had already detected this trend in studies with dung beetles associated with livestock systems in the Neotropics, where the authors suggest that it is necessary to include more information about the type of management, use of antiparasitic drugs, characteristics of pastures and soils, livestock species and pastures used in grazing areas to replicate these studies methodologically and thus be able to compare them on a large scale; that is why in this review we only include the work of Gómez-Cifuentes et al. (2020) in the livestock management research area because in that study the authors include a multidimensional analysis of the main variables associated with livestock management obtained from interviews with producers in different localities in the north of Misiones province.

In contrast, the ecological functions and physiology of dung beetles were poorly explored in the country. In the first case, the most evaluated metrics were nutrient removal, soil quality, and seed viability. In contrast, in the second case, the metrics evaluated were thermal amplitude, endothermy, thermoregulation, thermal niche, thermal limits, and metabolic rate. Previous studies have reported that dung beetle community attributes associated with alpha diversity are good indicators of changes in the structure of these communities caused by anthropogenic disturbance (Alvarado et al., 2019). However, many of these studies evaluate the number of abundant and generalist species, exploring little what happens with the rare and specialists, which are key to the functioning of ecosystems and probably respond to thermal limits associated with environmental conditions, which means that these two lines of research are the ones that best complement studies on biodiversity and conservation of dung beetles (Cardoso et al., 2020; Giménez-Gómez et al., 2025). However, it is worth clarifying that these research areas (functioning and physiology) were the most recently explored in the country (at the end of the 2010s and the beginning of the 2020s).

Dung beetles reported in Argentina

From the studies compiled in this review, a total of 236 species of dung beetles were reported in Argentina. Based on this, the Paranaense rainforests (Misiones) and the Humid and Dry Chaco were the geographic areas with the highest records of dung beetles. In the first case, the province of Misiones is likely associated with the presence of the Atlantic forest, one of the most diverse ecosystems on the planet (Di Bitetti et al., 2003); in the second case, the diversity in Chaco province probably is associated with the great diversity of ecological niches resulting from the biogeographic context of the Chaco province (e.g., hydrophilic and mesophilic forests, natural grasslands, Caranday palm groves, among others) (Ibarra-Polesel, Gómez-Cifuentes & Zurita manuscript in preparation). However, the records of dung beetles in the Chaco are likely also linked to its recent historical context, i.e., the intensity of land use, the loss and fragmentation of natural habitats, and the increasing demand for productive areas. For example, the study by Guerra Alonso et al. (2019) conducted in the province of Chaco showed that livestock systems that preserved canopy cover (silvopastoral) also preserved native dung beetle richness; this result is similar to that reported in other studies with dung beetles associated with neotropical humid forests, which showed that higher dung beetle richness is, generally, associated to environments with higher forest cover (Giraldo et al., 2011; Damborsky et al., 2015; Gómez-Cifuentes et al., 2019, 2022).

Moreover, the richness of dung beetles reported in Santiago del Estero stands out, a province that includes the semi-arid subregion of the Argentine dry Chaco ecoregion. Previous studies in Santiago del Estero have shown that the biogeographic history of dung beetles plays a fundamental role in understanding the response of these insects to disturbances (Guerra Alonso et al., 2020; Gómez-Cifuentes et al., 2023); i.e., these studies have observed that dung beetles present in the Argentine dry Chaco do not depend on the microclimatic regulation provided by trees as occurs in ecoregions with humid forests, but are probably also adapted to native and exotic grasslands (with and without livestock) thanks to the development of active thermoregulatory mechanisms which likely appeared as a response to extreme daily fluctuations in temperature. In contrast, the low richness of dung beetles in the southern ecoregions and provinces of the country likely responds to thermal constraints (temperature and humidity) and resource availability (low mammal diversity and abundance), which may be associated with biogeographic patterns in the Neotropics (Davis et al., 2002).

CONCLUSIONS

The study of dung beetles in Argentina is about to celebrate its first century. Since the 2000s, the field has grown exponentially, accounting for 79 % of the total studies conducted in the country. The Paranaense rainforest ecoregion and Misiones resulted in the geographical areas with the highest number of studies with dung beetles in Argentina; in contrast, the center and south of the country have been poorly explored. In general, the behaviour, diagnosis, and diversity of dung beetles in the country have been well-studied; however, there are research areas, such as physiology and ecological functions, presenting enormous potential for future research in the country. Moreover, the Paranaense rainforest (Misiones), as well as the Dry and Humid Chaco (Chaco and Santiago del Estero), were the geographic regions with the highest records of dung beetles (all of them in the northern portion of the country). It is necessary to explore more ecoregions of the country, such as the Fields and Scrublands and the Yungas Forest, among others, due to their unique microclimatic conditions and the fauna present, which may support many species of dung beetles. In addition, half of the provinces likely can host many more species, among which we highlight La Rioja, Salta, Catamarca, Tucumán, and Jujuy in the northwestern region; Buenos Aires, La Pampa, Entre Ríos, and Santa Fe in the Pampa region, and San Juan and San Luis in the Cuyo region.

Based on the above, we recommend that future researchers focus on less-explored ecoregions, provinces, and research areas. Additionally, it is necessary to conduct more inventories to understand the complete richness of dung beetles in Argentina. Finally, we recommend that studies on biodiversity and conservation in livestock systems add as much information as possible about livestock management following data collection formats such as the one proposed by Arellano et al. (2023).

Acknowledgments

To all Argentine and foreign students, researchers, and naturalists for their invaluable contribution to the knowledge of the Argentinean dung beetles in the last 85 years. To CONICET, universities, and national institutions for financial, logistical, and academic support. To dung beetles!

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Notes

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