Research trends and gaps in ant-mediated seed dispersal: a review with emphasis on Brazilian biomes

Tendências e lacunas na pesquisa sobre dispersão de sementes mediada por formigas: uma revisão com ênfase nos biomas brasileiros

Abstract

This scientometric review analyzes global research patterns on seed dispersal by ants (myrmecochory) from 2009 to 2019, with particular emphasis on geographic and methodological biases across Brazilian biomes. Our analysis reveals not only pronounced disparities in research distribution, with studies heavily concentrated in Atlantic Forest and Cerrado ecosystems, while Amazonia, Pantanal, and Pampa remain critically understudied, but also fundamental methodological limitations that constrain our understanding of the ecological role of myrmecochory. More importantly, we provide the first comprehensive biome-level comparison of research effort and methodological approaches, identifying critical gaps in distinguishing seed dispersal from predation events and assessing post-dispersal seed viability. Rather than merely documenting these imbalances, we offer a strategic framework for prioritizing research in underrepresented biomes and implementing standardized methodologies. Our findings have direct implications for conservation planning and restoration ecology, particularly in threatened tropical ecosystems where ant-mediated dispersal could significantly enhance recovery programs. Addressing these geographic and methodological biases is therefore essential for advancing myrmecochory research and improving its applied relevance.

Keywords:
ant-plant interaction; myrmecochory; research biases; ecological restoration; conservation planning; scientometric review

Resumo

Esta revisão cienciométrica analisa a pesquisa global sobre dispersão de sementes por formigas (mirmecocoria) de 2009 a 2019, com ênfase particular nos padrões geográficos e abordagens metodológicas nos biomas brasileiros. Nossa análise revela disparidades pronunciadas na distribuição da pesquisa, com estudos fortemente concentrados nos ecossistemas da Mata Atlântica e do Cerrado, enquanto a Amazônia, o Pantanal e o Pampa permanecem criticamente pouco estudados. Mais importante ainda, identificamos limitações metodológicas fundamentais que limitam nossa compreensão do papel ecológico da mirmecocoria, particularmente a falha em distinguir entre eventos de dispersão de sementes e predação, e a avaliação insuficiente da viabilidade de sementes pós-dispersão. Além de meramente mapear lacunas de conhecimento, nosso estudo fornece uma estrutura estratégica para priorizar a pesquisa em biomas sub-representados e implementar metodologias padronizadas. Essas percepções são particularmente importantes para orientar os esforços de restauração em ecossistemas tropicais ameaçados, onde a dispersão mediada por formigas pode aprimorar significativamente programas de recuperação. Abordar esses vieses geográficos e metodológicos é, portanto, essencial para o avanço dos estudos sobre mirmecocoria e para fortalecer sua relevância aplicada.

Palavras-chave:
interação formiga-planta; mirmecocoria; vieses de pesquisa; restauração ecológica; planejamento de conservação; revisão cienciométrica

1. Introduction

In an ecosystem, many interactions occur simultaneously between animals and plants, which can be positive or negative (Del-Claro and Torezan-Silingardi, 2021; Pereira et al., 2021; Couto et al., 2022; Alencar et al., 2023). Among these interactions, seed dispersal stands out as a fundamental process for the maintenance of ecosystem functioning (Corlett, 2021). Seed dispersal enables the movement of plant genes across space and, in some cases, through time, influencing reproduction, population genetic structure, and several other components of plant ecology (Lengyel et al., 2009). This mechanism benefits both the mother plant and the propagule by reducing intraspecific competition and mitigating density- and distance-dependent effects from pathogens and herbivores. It also contributes to floristic heterogeneity and supports the maintenance of forest areas (Lengyel et al., 2009; Corlett, 2021).

The structural traits, chemical composition, and nutritional properties of seeds and/or fruits influence how they are dispersed, forming what are classically known as dispersal syndromes, as defined by van der Pijl (1982). Among these syndromes, six major categories are recognized: autochory, zoochory, anemochory, anthropochory, hydrochory, and barochory (Oliveira et al., 2022; Torres et al., 2022). Zoochory, the dispersal of seeds by animals, is particularly prominent and is considered the dominant dispersal strategy in several pristine ecosystems (Jordano, 2000). In South American forests, zoochory is estimated to represent about 70-90% of Angiosperm seed dispersal (Almeida-Neto et al., 2008; Oliveira et al., 2022). Zoochoric fruits typically exhibit traits that facilitate animal-mediated dispersal, such as fleshy pulp, elaiosome-bearing seeds, mucilage production, or structures that enable adhesion (Almeida-Neto et al., 2008; Hernández-Brito et al., 2021; Oliveira et al., 2022). Within zoochory, there are subdivisions based on the taxonomic group of the dispersal agent, including saurochory (reptiles), mastozoochory (mammals), ornithochory (birds), ichthyochory (fish), and myrmecochory (ants) (Almeida-Neto et al., 2008; Silveira and Weiss, 2014; Gomes et al., 2021; Leal et al., 2011).

Myrmecochory is a relatively common type of seed dispersal, occurring in several regions and ecosystems worldwide and present in more than 11,000 species of angiosperms (approximately 4.5%) (Lengyel et al., 2009). This interaction is primarily mediated by the presence of a lipid-rich elaiosome attached to the seed coat (Beattie, 1985), although factors such as seed size, odor, and chemical cues may also influence ant-seed interactions. Elaiosomes are highly attractive to ants due to their protein and carbohydrate content (van der Pijl, 1982), functioning both as a food reward and a structure that facilitates seed transport. In primary dispersal, ants actively collect seeds bearing elaiosomes and transport them to their nests to consume this nutritive structure. Secondary dispersal occurs when ants relocate seeds previously dispersed by other vectors or attempt to predate upon seeds lacking elaiosomes. After handling, ants typically consume the elaiosome and may discard the intact seed in nutrient-rich nest chambers or midden piles (Beattie, 1985), potentially enhancing germination, protecting against predators, and placing seeds in favorable microsites (Dáttilo et al., 2009; Rapalino and Haydar, 2017). However, ants may also act as seed predators, and dispersal effectiveness may be reduced when seeds are deposited in unsuitable locations (Oliveira et al., 2023).

Despite its ecological relevance, the occurrence and importance of myrmecochory studies are not equally distributed among world ecosystems. In addition, many studies do not verify whether ants truly disperse seeds, whether seeds remain viable after handling, or whether deposition sites are ecologically advantageous. Therefore, mapping where, when, and how myrmecochory has been studied becomes essential, particularly in biodiverse regions. Although myrmecochory has been studied globally, the distribution of research effort across Brazilian biomes remains poorly understood. Here, we aim to detect current and geographical trends and gaps in myrmecochory research. We examined how published studies are distributed spatially (a) and temporally (b). In addition, we assessed which type of myrmecochory is most studied (c), whether there is empirical evidence of dispersal (d), whether native seeds were used (e), whether ants disperse viable seeds (f), and how far seeds are transported (g). These results provide a foundation for future studies to address knowledge gaps and inform strategies for vegetation recovery. Although this is a global scientometric review, our discussion places particular emphasis on Brazilian biomes in order to identify national research gaps and provide applied recommendations for conservation and restoration.

2. Material and Methods

2.1. Literature survey

We carried out a literature search in the Scopus (2025) database during 2020, because this database provides broad, multilingual journal coverage, advanced filtering by fields (title/abstract/keywords) and consistent indexing that supports reproducible searches. The search used the following terms in English, Portuguese, and Spanish: “myrmecochory”, “seed AND dispersal AND ants”, and their translated equivalents. The search was conducted in the fields “title”, “abstract”, and “keywords”. We restricted the survey to peer-reviewed articles published between 2009 and 2019, a decade-long interval commonly used in scientometric analyses to identify recent trends. This timeframe captures methodological developments in myrmecochory research, such as the use of artificial seeds and evaluations of secondary dispersal, while minimizing dilution by older studies based on less comparable approaches. Additionally, the period precedes the most severe constraints on Brazilian science funding, offering a relatively stable context for interpreting research productivity. We therefore consider this interval appropriate for identifying current geographic and methodological gaps in myrmecochory research. Although the survey encompasses global studies, the Discussion places particular emphasis on Brazilian biomes to highlight region-specific research gaps and applied implications.

2.2. Information compilation

All retrieved papers were screened for thematic relevance and redundancy. We included only empirical studies directly investigating seed dispersal by ants. Review articles, meta-analyses, data papers, scientometric articles, and studies that did not fit the topic were excluded. Non-peer-reviewed materials such as books, book chapters, conference abstracts, technical reports, and graduate dissertations were also excluded. Duplicate records were removed when the same data set appeared in more than one publication. Screening was performed by the first author, with support from the senior author. To ensure consistency in the screening process, a second coauthor independently validated a subset of the selected records, and any discrepancies were resolved through discussion. For transparency, we report a concise summary of the screening flow: 295 records were retrieved from Scopus, 144 were excluded based on the criteria above, and 151 studies met the inclusion criteria and were analyzed.

From each eligible study, we compiled 11 data metrics: (1) year of publication; (2) geographical coordinates; (3) phytophysiognomy or biome of the study area; (4) country; (5) type of dispersal investigated (primary or secondary); (6) type of seed used (native, exotic, or artificial); (7) distance of seed removal or dispersal; (8) focal event (removal, dispersal, or predation); (9) whether the event was verified; (10) whether a germination test was conducted; and (11) how the germination test, when present, was performed.

When coordinates were provided, we assigned the study to a biome using official Brazilian biome limits (IBGE, 2025). When coordinates were absent, biome classification was inferred from the vegetation descriptions reported in the original article. Although biome or phytophysiognomy was one of the recorded metrics, this information was consistently available only for studies conducted in Brazil. Most international articles lacked sufficient detail to allow reliable biome assignment, and attempting to force a global classification would likely generate artifactual patterns. For this reason, biome-level analyses were restricted to Brazilian studies, where vegetation descriptions are more standardized and where evaluating the spatial distribution of research effort is particularly relevant given the country’s large area and high representation in our data set. Descriptive analyses based on simple frequencies were conducted, and results were visualized in R (R Core Team, 2023) using the ggplot2 package (Wickham, 2016).

3. Results

A total of 295 peer-reviewed articles were initially retrieved, of which 151 met the inclusion criteria. The 144 excluded studies were primarily review articles or papers not focused directly on seed dispersal by ants.

We observed a temporal oscillation in the annual number of publications on myrmecochory, with no clear trend across the 11-year period. The highest number of publications occurred in 2018 (18 articles), whereas 2014 had the lowest output (8 articles) (Figure 1). On average, 15.1 articles were published per year during the study period.

Figure 1
Annual number of peer-reviewed articles on myrmecochory retrieved from the Scopus database.

Brazil was the country with the highest number of publications (27 articles; 17.9% of all studies), followed by the United States (22; 14.6%), Australia (16; 10.6%), Spain (15; 9.9%), Japan (13; 8.6%), and China (12; 7.9%) (Figure 2). Seventeen countries (11.3%) had only one published study on this topic between 2009 and 2019.

Figure 2
Number of peer-reviewed articles on myrmecochory published per country between 2009 and 2019. Dark gray areas indicate countries with no studies indexed in Scopus; warmer colors represent higher publication numbers.

In Brazil, studies were unevenly distributed among biomes. The Atlantic Forest had the highest number of publications (11 studies), followed by the Cerrado (10). However, additional records occurred outside these two dominant biomes, including three studies in the Caatinga, one study in the Cerrado-Atlantic Forest transition (CAFT), one study in the Cerrado-Amazon transition (CAT), and one study that could not be assigned to a specific biome due to insufficient geographic information (“Not specified”). No studies were recorded for the Amazon, Pantanal, or Pampa biomes sensu stricto, despite the presence of records in their ecotonal zones (Figure 3).

Figure 3
Number of peer-reviewed articles on myrmecochory conducted in different Brazilian biomes. CAFT = Cerrado-Atlantic Forest transition; CAT = Cerrado-Amazon transition.

To contextualize this geographic bias, we compared the number of studies per biome with their total area and cumulative deforestation between 2009 and 2019 (MapBiomas, 2025 – Table 1). This comparison revealed that research effort is not aligned with environmental vulnerability: although the Caatinga does contain studies (n = 3), research effort in this biome remains very low relative to its extent and deforestation levels. Research gaps are particularly concerning in the Amazon, Pantanal, and Pampa, which had no studies, and in the Cerrado, which was only modestly represented (10 studies). These biomes also experienced the highest proportional deforestation during the study period (Pampa 13.9%, Caatinga 7.0%, Cerrado 6.0%). This overlap reveals an acute mismatch between ecological threat and research effort, indicating that the ecosystems under the greatest anthropogenic pressure are precisely those for which empirical data on ant-mediated seed dispersal are most lacking. Overall, these results indicate that myrmecochory research in Brazil remains geographically concentrated and does not track the environmental urgency of the biomes most impacted by land-cover change.

Table 1
Area of Brazilian biomes, accumulated deforestation (2009-2019), and number of myrmecochory studies identified in this review. Deforestation values were converted from hectares to km² based on the data set provided (MapBiomas, 2025).

Biome	Area (km2)	Deforestation 2009–2019 (km2)	Proportion (%)	Number of studies
Amazon	4,196,000	129,162	3.08	0
Caatinga	844,000	59,087	7.00	3
Cerrado	2,036,000	123,018	6.04	10
Atlantic Forest	1,110,000	35,988	3.24	11
Pampa	176,000	24,398	13.90	0
Pantanal	151,000	5,033	3.33	0

Worldwide, primary dispersal was the focus of 117 studies (77.5%), whereas 30 studies (19.9%) investigated secondary dispersal. Four studies (2.6%) addressed both processes. Only 27 studies (17.9%) explicitly verified whether seeds were dispersed or predated by ants. Most studies used native seeds (137; 90.7%), followed by exotic seeds (13; 8.6%), while only three studies (2.0%) used artificial seeds.

Seed viability was rarely evaluated: 126 studies (83.4%) did not perform germination tests on seeds removed by ants. Only 25 studies (16.6%) included germination assessments, and even fewer described the specific procedures adopted. Seed removal distance was reported in 48 studies (31.8%), which quantified how far ants transported seeds from the point of origin.

4. Discussion

Despite expectations that scientific output on myrmecochory would increase over the years, especially with advances in methodology and the use of artificial seeds (Henao-Gallego et al., 2012; Bieber et al., 2014), our results indicate a relatively stable number of publications in the last decade. This pattern contrasts with the general growth of Brazilian scientific production over recent decades (Barreto, 2006; Zorzetto et al., 2006; Leta et al., 2013), including the consolidation of research groups dedicated to ant biology and ecology (Brandão, 2020). Although Brazil has significantly advanced the knowledge of ant diversity (Andrade-Silva et al., 2022; Feitosa et al., 2022; Melo et al., 2021; Schmidt et al., 2022; Silva et al., 2022; Queiroz et al., 2023; Vicente et al., 2023; Lopes et al., 2021) and ant-plant interactions (Baccaro et al., 2015; Barbosa et al., 2015; Prado et al., 2016, 2022; Vicente and Izzo, 2017; Sozanski et al., 2020; Alencar et al., 2022), myrmecochory remains comparatively underrepresented. This demonstrates that the scientific community has not yet integrated ant-mediated seed dispersal as a mainstream research theme, despite its ecological and applied relevance. More recent contributions (e.g., Anjos et al., 2020; Alcolea et al., 2022) reinforce the importance of myrmecochory for ecosystem resilience and restoration, highlighting the need to expand research efforts in this field.

Although Brazil contributed the largest absolute number of myrmecochory studies in our sample, this scientific production is concentrated in only two biomes, which prevents a biome-level understanding of ant-seed interactions across the country. Given Brazil’s continental scale and its wide range of environmental conditions, interactions between ants and seeds are likely to vary substantially among biomes. The absence of studies in vast regions such as the Amazon, Pantanal, and Pampa therefore represents not only a numerical gap but also a loss of ecological resolution. Although our results show the presence of studies in Caatinga (3 studies) and in ecotonal regions (CAFT and CAT), these records remain sparse and do not compensate for the lack of studies within the Amazon, Pantanal, and Pampa biomes sensu stricto. In this sense, Brazil’s numerical leadership does not translate into comprehensive coverage, and expanding research to underrepresented biomes is essential for capturing the full functional variability of myrmecochory.

Brazil, the United States, Japan, and Australia led in the number of myrmecochory studies. Although these countries are megadiverse, biodiversity alone does not explain the research effort. For instance, the USA invests a larger proportion of its Gross Domestic Product in research (2.7%) compared to Brazil (1.3%), and Brazilian science has experienced recurrent funding cuts and political instability (Caires, 2019; Escobar, 2019; Jordão, 2019). These constraints often limit research to more accessible areas, contributing to geographic gaps, especially in biomes facing climate and land-use pressures (Carvalho et al., 2023). Acknowledging these structural limitations is essential, as they influence not only scientific output but also the representativeness of ecological studies. Future syntheses should incorporate analyses of how financial and infrastructural disparities shape research agendas, particularly in regions of the Global South.

Within Brazil, the mismatch between biodiversity and research effort is evident in the uneven distribution of studies across biomes. Most investigations occurred in the Atlantic Forest and the Cerrado that is global biodiversity hotspots undergoing rapid environmental change (Françoso et al., 2015; Lima et al., 2017; Gonella et al., 2021; Francisco et al., 2023). These biomes receive more scientific attention due to their conservation urgency and the concentration of research institutions. The focus on seed dispersal processes in these regions (Arruda et al., 2018; Fraga et al., 2019; Klink et al., 2020; Marques and Grelle, 2021) reflects their sensitivity to fragmentation (Rocha-Santos et al., 2020). The presence of studies in the Caatinga and in transition zones (CAFT and CAT) suggests emerging, although still limited, research efforts outside the two major biomes. Although justified, this concentration reinforces a geographic bias that limits national-scale conservation planning. Understudied biomes such as the Amazon, Pantanal, and Pampa still present substantial knowledge gaps, despite increasing anthropogenic pressures. Addressing these gaps is essential for designing restoration strategies that consider ant-mediated seed dispersal as a potential tool.

The Amazon, despite its extraordinary biodiversity (Vicente et al., 2020; Alencar et al., 2023), remains scarcely represented in myrmecochory studies. Only one study in our data set was conducted in the Amazon-Cerrado transition (Paolucci et al., 2016). More recent work has shown that land-use change, such as pasture conversion, reduces both the diversity of seed-dispersing ants and the quality of dispersal services (Fontenele and Schmidt, 2021). This is concerning given the intensification of forest degradation and the weakening of environmental protections (Carvalho et al., 2019; Ribeiro et al., 2022). Similarly, no studies were recorded for the Pantanal or Pampa biomes, both experiencing increasing environmental pressures (Albuquerque et al., 2017; Marengo et al., 2021; Ziliotto et al., 2023). These omissions overlook the potential of ants to contribute to forest regeneration (Gallegos et al., 2014; Bianchi et al., 2017; Cristo et al., 2019). Recent evaluations of Amazonian research gaps (Carvalho et al., 2023) reinforce that these ecosystems require urgent scientific attention.

Our analysis also highlights methodological limitations that hinder a complete understanding of myrmecochory's ecological role. A recurring issue is the lack of clarity about whether seeds removed by ants were consumed or dispersed, an essential distinction for evaluating dispersal effectiveness (Rodgerson, 1998; Suárez-Esteban et al., 2018). Although most studies used native seeds, few assessed seed viability after removal (17%) or quantified dispersal distance (32%), both critical for determining whether ant-mediated dispersal enhances plant fitness (Christianini et al., 2007; Slusarenko et al., 2012). Ants typically disperse seeds over short distances, but they may compensate for this by acting in contexts where vertebrates are absent or functionally reduced (Christianini and Oliveira, 2013; Anjos et al., 2020; Alcolea et al., 2022). Thus, the absence of viability and distance assessments limits the applicability of results to restoration planning. To reconcile these points, we now explicitly clarify that myrmecochory has strong ecological and theoretical potential to contribute to restoration, but its practical implementation is currently constrained by these methodological gaps, which must be resolved through standardized protocols and empirical tests that quantify seed fate, viability, and establishment.

Recent developments in restoration ecology emphasize the need for precision forest restoration, which argues that restoration success depends on verifying the fate, viability, and effective establishment of each seed rather than relying solely on broad-scale interventions. This perspective (Castro et al., 2021) directly aligns with the methodological gaps revealed by our review, particularly the limited verification of dispersal vs. predation, the rarity of post-removal viability tests, and the inconsistent measurement of removal distances. Integrating these principles into studies of ant-mediated seed dispersal would enable the development of standardized protocols capable of determining whether myrmecochory can reliably contribute to restoration outcomes.

Artificial seeds and elaiosomes remain underutilized in myrmecochory research, despite their potential to provide controlled experimental conditions that help evaluate ant behavior and ecological filters (Raimundo et al., 2004; Luna and Dáttilo, 2018; Luna et al., 2021). Wider adoption of these approaches would allow comparative analyses across regions and offer insights into how environmental variation shapes seed removal.

Globally, most studies focused on primary dispersal, while secondary dispersal remains poorly investigated. Diplochory (sequential dispersal by two vectors) is common in tropical ecosystems (Vander Wall and Longland, 2004; Leal et al., 2015; Zhu et al., 2018) and likely depends on local biodiversity and faunal integrity. For example, the overhunting of frugivores in the Amazon disrupts vertebrate-mediated dispersal (Peres and Palacios, 2007), which may affect subsequent ant-mediated secondary dispersal. Integrating diplochory into restoration ecology is particularly relevant, as degraded areas often lose large vertebrates first, making ants potential “last-resort” dispersers. Recognizing this applied dimension increases the conservation value of myrmecochory research.

Recent advances in ecological big data and citizen science offer promising opportunities to reduce geographic and taxonomic gaps in myrmecochory research. Large, curated biodiversity repositories such as speciesLink and the Brazilian Biodiversity Information System (SiBBr), together with structured ecological data sets (Iamara-Nogueira et al., 2022; Silva et al., 2022; Boscolo et al., 2023; Lacerda, 2025), demonstrate how standardized and open-access databases can expand spatial coverage and enable integrative analyses across ecosystems. A parallel expansion of citizen-science initiatives in Brazil, including major platforms such as WikiAves and iNaturalist, has proven effective for filling distributional gaps; for example, citizen-generated records substantially increased the known area of occupancy and ecological representation of Amazona vinacea (Bovo et al., 2024). These developments echo broader arguments in ecology that big data and participatory monitoring enhance inference, especially in poorly sampled systems (Kremen et al., 2011; Dickinson et al., 2012; Hampton et al., 2013). Together, these initiatives outline a clear path forward: integrating curated biodiversity databases, national repositories and validated citizen-science observations can accelerate the discovery of ant-seed interactions in undersampled biomes, support macroecological assessments and guide restoration planning in regions where traditional field sampling remains logistically challenging.

Standardization is essential to enable comparisons across regions and biomes. Without consistent methodologies, it is difficult to integrate findings into conservation strategies or identify global patterns. Future research agendas should prioritize the development of standardized protocols. Addressing the geographic and methodological gaps revealed by our review is not only a scientific goal but a conservation necessity. These findings have direct implications for conservation planning and ecological restoration, particularly in threatened tropical ecosystems where ant-mediated seed dispersal may play a crucial role. Confronting these biases should be treated as both a scientific and conservation priority in the face of accelerating environmental change.

5. Conclusions

This review highlights the main advances and persistent gaps in myrmecochory research, particularly the uneven geographic distribution of studies both globally and within Brazil. Although Brazil published the highest number of publications in our data set, research remains concentrated in a few well studied biomes, leaving large regions with little to no empirical information.

We also identified key methodological shortcomings that constrain broader inferences about the ecological role of ant-mediated seed dispersal. These include the limited verification of whether seeds are truly dispersed or predated, the low frequency of seed viability assessments, and the underuse of standardized experimental tools such as artificial seeds and elaiosomes. Establishing consistent and comparable protocols across regions and biomes is essential for integrating results into conservation planning and for building a more comprehensive understanding of global myrmecochory patterns.

Bridging these geographic and methodological gaps is fundamental for advancing both basic and applied aspects of ant-mediated seed dispersal. Incorporating myrmecochory into restoration programs (particularly in ecologically significant and understudied biomes such as the Amazon, Pantanal, and Pampa) represents both a scientific opportunity and a conservation priority. Given the accelerating pace of environmental change and habitat fragmentation across Brazilian ecosystems, recognizing and leveraging the role of ants in seed dispersal is an urgent step toward more effective restoration and conservation strategies.

Acknowledgments

We thank Livia Pires do Prado, Liliane Stedile de Matos, and Thiago Junqueira Izzo for reviewing and providing valuable feedback on earlier versions of the manuscript, as well as the five referees whose comments greatly improved this updated version. We also thank Gustavo Rocha Costa for his support with data compilation, Thiago Ferreira for producing the heat map, and Ivone Silva for allowing us to use the laboratory during the study. WSB acknowledges support from the National Council for Scientific and Technological Development (CNPq) through a scientific initiation scholarship and a Desenvolvimento Tecnológico e Industrial (DTI) fellowship (Grant No. 350388/2024-5). REV thanks the Fundação de Amparo à Pesquisa do Estado de Mato Grosso (FAPEMAT; DCR 003/2016), CNPq (PDJ 150826/2024-9) for postdoctoral research fellowships and the Programa de Pós-graduação em Zoologia, Universidade Federal do Amazonas (POSGRAD/FAPEAM and PROAP/ CAPES).

Data Availability Statement

The data set analyzed or produced in this study can be requested from the corresponding author.

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Edited by

• Editor:
  Takako Matsumura Tundisi

Publication Dates

• Publication in this collection
  15 May 2026
• Date of issue
  2026

History

• Received
  27 Aug 2025
• Accepted
  13 Jan 2026