9 urn:lsid:arphahub.com:pub:F9D0807C-3C52-5A33-8AB6-FB2607A29353 urn:lsid:zoobank.org:pub:E14246FD-79C0-4B14-A52F-3F6A3D7ECCA8 Subterranean Biology SB 1768-1448 1314-2615 Pensoft Publishers 10.3897/subtbiol.57.193389 193389 Research Article Mycetozoa Myxomycetes Protozoa Systematics Taxonomy Cenozoic Americas Brazil South America First records of myxomycetes (Amebozoa, Eumycetozoa) in cave in Central Brazil Moreira Izabel Cristina izacristina26@yahoo.com.br https://orcid.org/0000-0001-5302-7264 1 Cavalcanti Laise de Holanda https://orcid.org/0000-0002-6011-7142 2 Marley-Santos Roberth Lopes https://orcid.org/0009-0009-3749-3439 1 Leonardo-Silva Lucas https://orcid.org/0000-0001-6298-4293 1 de Lima Luciene Araújo https://orcid.org/0009-0004-7668-6755 1 Xavier-Santos Solange https://orcid.org/0000-0002-3397-0885 1 Laboratório de Micologia Básica, Aplicada e Divulgação Científica (FungiLab), Universidade Estadual de Goiás, Anápolis, Goiás, Brazil Laboratório de Micologia Básica, Aplicada e Divulgação Científica (FungiLab), Universidade Estadual de Goiás Anápolis Brazil https://ror.org/03ta25k06 Centro de Biociências - CB, Universidade Federal de Pernambuco (UFPE), Recife, Pernambuco, Brazil Centro de Biociências - CB, Universidade Federal de Pernambuco (UFPE) Recife Brazil https://ror.org/047908t24

Corresponding author: Izabel Cristina Moreira (izacristina26@yahoo.com.br)

Academic editor: Rosaura Mayén-Estrada

 2026 16 07 2026 57 91 103 6E90ED7E-8561-5451-B699-1E2F39E26421 55DCF944-32EE-425F-8E0B-920890548B33 28 03 2026 27 05 2026 Izabel Cristina Moreira, Laise de Holanda Cavalcanti, Roberth Lopes Marley-Santos, Lucas Leonardo-Silva, Luciene Araújo de Lima, Solange Xavier-Santos This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. https://zoobank.org/55DCF944-32EE-425F-8E0B-920890548B33

More than 26,000 caves occur in the extensive Brazilian territory, distributed across the country’s five geopolitical regions. Some of these caves have been studied for the biodiversity they harbor; however, many groups of organisms are neglected, as is the case with myxomycetes. Herein we present the second record of myxomycetes in Brazilian caves, based on specimens sampled in a cave in the Central-West region, in the state of Goiás. Leaf litter and decaying wood fragments from the photic, penumbra, and aphotic zones were incubated in humid chambers and monitored for 90 days. Of the 22 humid chambers prepared, 45% were positive. Three species were identified: Arcyria cinerea (Arcyriaceae), Cribraria elegans (Cribrariaceae), and Perichaena depressa (Trichiaceae), in addition to one unidentified specimen. This study, the first conducted in Cerrado caves, expands the known occurrence of myxomycetes in Brazilian subterranean environments, reporting Cribrariaceae and P. depressa for the first time in caves and A. cinerea for the first time in a Brazilian Cerrado cave.

 Cave-dwelling Cerrado Karst zones Slime molds Conselho Nacional de Desenvolvimento Científico e Tecnológico 501100003593 https://ror.org/03swz6y49 http://doi.org/10.13039/501100003593 Coordenação de Aperfeiçoamento de Pessoal de Nível Superior 501100002322 https://ror.org/00x0ma614 http://doi.org/10.13039/501100002322 Fundação de Amparo à Pesquisa do Estado de Goiás 501100005285 https://ror.org/00w8cq239 http://doi.org/10.13039/501100005285 Citation

Moreira IC, Cavalcanti LdeH, Marley-Santos RL, Leonardo-Silva L, de Lima LA, Xavier-Santos S (2026) First records of myxomycetes (Amebozoa, Eumycetozoa) in cave in Central Brazil. Subterranean Biology 57: 91–103. https://doi.org/10.3897/subtbiol.57.193389

 Introduction

Myxomycetes (Amebozoa, Eumycetozoa) have a wide distribution in terrestrial environments, occurring in different microhabitats rich in organic matter where they act as predators of microorganisms, contributing to regulate microbial communities and to decompose and cycle nutrients (Stephenson and Stempen 1994; Dussutour et al. 2010; Ferreira and Cavalcanti 2010; Hosokawa et al. 2019; Leonardo-Silva et al. 2021; Stephenson and Novozhilov 2021; Keller et al. 2022; Sá et al. 2022; Toschi et al. 2025).

Around 1100 species of myxomycetes are currently recognized worldwide. Many of these species, such as Arcyria cinerea (Bull.) Pers. and Fuligo septica (L.) F.H. Wigg., exhibit high adaptability and the ability to occupy different ecological niches, being found in virtually all terrestrial ecosystems (Cavalcanti et al. 2014; Rojas and Valverde 2015; Xavier-Santos et al. 2016; Stephenson et al. 2021).

This ability to colonize a wide diversity of environments, together with their morphology that is easily detectable in the field, contributes to the broad geographic distribution of these species, recorded either through sporophores observed in the field or through development from collected material, including in habitats with particular environmental conditions, such as caves, which are characterized by low light, high humidity, and relative thermal stability.

Myxomycetes, along with bacteria, fungi, and protozoa, play important roles in decomposing organic matter in these locations, releasing nutrients for cave-dwelling invertebrates (Galán 2015). Records of myxomycetes have been made in caves in countries of the Iberian Peninsula (Spain), North America (USA, Mexico), and Central America (Puerto Rico), occurring in various substrates, including rocks, speleothems, leaf litter, bones, decaying wood, and soils enriched with bat guano (Landolt et al. 1992; Nieves-Rivera 2003; Galán and Nieto 2010, 2022; Galán et al. 2010, 2018, 2021a, 2021b, 2022; Galán 2011, 2015; Galán and Rivas 2019; Martínez-Rodríguez et al. 2023).

The occurrence of myxomycetes in caves in Brazil is very poorly known, only being recorded by Moreira et al. (2024), who reported the presence of Arcyria denudata (L.) Wettst., Hemitrichia calyculata (Speg.) M.L. Farr and Metatrichia vesparia (Batsch) Nann. -Bremek. ex G.W. Martin & Alexop. in caves in the karst region of Pains, in Minas Gerais, in the Southeast of the country. These species belong to the order Trichiales, and the last two were reported for the first time in subterranean environments. This scarcity of data highlights the need for research focused on the distribution of these organisms in cave environments in the country.

In this context, the present study aims to contribute to advance knowledge about this biodiversity by recording new occurrences and expanding the information available about the presence of myxomycetes in Brazilian caves.

 Method Study area

The municipality of Vila Propício is located in the Central-West region of Brazil, in the eastern mesoregion of the state of Goiás, within the Cerrado biome (15°27'25"S, 48°53'20"W), approximately 205 km from Goiânia, the state capital, and 194 km from Brasília, the country’s capital. The region has a pronounced relief, with altitudes between 760 and 840 m a.s.l. and a semi-humid tropical climate, with an average annual temperature between 22 and 26 °C, and annual rainfall of up to 1,500 mm (CECAV 2019; Miranda et al. 2024; IBGE 2026). From a geological point of view, the municipality is located within units belonging to the Araxá Group, predominantly composed of metasedimentary rocks, such as schists, quartzites, marbles, phyllites, and carbonate formations (Oliveira et al. 2024). These lithotypes contain recurring minerals such as calcite, dolomite, muscovite, and quartz (among others).

Among the 35 natural caves in the municipality registered with the National Center for Cave Research and Conservation (ICMBio/CECAV) is Lapa do Boqueirão, a cave developed in dolomitic rocks with an extension of approximately 300 m. The cave is located approximately 40 km from the urban perimeter of Vila Propício, in the microbasin of the Pai José Stream, within the Maria Cícera settlement, an area corresponding to the former Bom Jesus Brejão Farm. Access is via a trail of approximately 1 km, which begins in the backyard of one of the residences in the settlement. The route crosses sections of open and closed forest, leading to the bottom of a valley, where it follows the bed of a dry drainage channel. The presence of steep rock formations near the entrance requires a short climb to access the cave, from where it offers a wide view of the surrounding Cerrado vegetation due to its elevated position (Figs 1, 2).

10.3897/subtbiol.57.193389.figure1 3AF54170-5787-5F41-BD47-E7BA7BEC5F78

Location of the study area. The map shows South America, with Brazil represented in dark gray and the distribution of the Cerrado biome in green. The circle indicates the location of Lapa do Boqueirão Cave. The inset map shows the location of the cave in the municipality of Vila Propício, Goiás State, Brazil.

https://binary.pensoft.net/fig/1717885 10.3897/subtbiol.57.193389.figure2 7AF68287-42E0-5DD8-B297-2486B1E2FADF

Images of Lapa do Boqueirão, municipality of Vila Propício, state of Goiás, Brazil. A, B aerial view, obtained by drone, of the main entrance to the cave B rock climbing to access the cave C view of the interior of the cave, aphotic zone, showing stalactites and stalagmites D, E secondary opening formed by the partial collapse of the rock wall.

https://binary.pensoft.net/fig/1717886

The cave features a wide opening with abundant limestone formations and stalactites. This initial section (photic zone) receives ample natural light and has a floor formed by sediments, rocky blocks, and debris resulting from erosion and weathering processes. A portal approximately 2 m high provides access to a gallery where the light gradually decreases until the aphotic zone. Accumulated branches and leaf litter were observed in the intermediate portion, indicating the occurrence of occasional currents, probably associated with increased water flow during rainy periods. More speleothem formations were observed in the innermost area in complete absence of light (aphotic zone), such as stalactites, stalagmites, columns, and curtains, in addition to plant debris transported into the cavity. The cave has a second opening formed by the collapse of part of the rocks that constituted its walls, which resulted in separating a small portion of the cavity; however, this opening does not present characteristics which allow it to be classified as a skylight. This opening is only accessible from inside the cave, as the external area is covered by dense vegetation and has no access trails (Fig. 2).

 Sampling, identification and preparation of samples

Leaf litter samples and decaying wood fragments were collected in three zones of the cave: (i) photic, or illuminated, zone, near the main entrance and the second opening; (ii) intermediate or penumbra zone; and (iii) aphotic zone, inside the cave, completely devoid of light. Three samples were collected from each zone and individually placed in properly labelled paper bags. These samples were used to make humid chambers using 9 cm diameter Petri dishes, lined with filter paper, upon which the samples were deposited. The interior of the chambers was kept humid by adding distilled water at least twice a week.

Once established, the humid chambers were monitored weekly for 90 days through observations under a stereomicroscope, aiming to detect myxomycete plasmodia, sclerotia and/or sporophores. Species identification was performed by macro and microscopic analysis of the sporophore using a Leica EZ4HD stereomicroscope and an Olympus CX31 optical microscope. Taxonomic determination was based on specialized literature, such as Barbosa et al. (2016), Cavalcanti et al. (2016, 2020), Poulain et al. (2011) and Martin and Alexopoulos (1969), while nomenclature followed the pattern proposed by Lado (2005–2026). The indication of character colors followed the color charts of Kornerup and Wanscher (1978). The global geographic distribution of the species was established based on information available in specialized literature and taxonomic databases, such as the Flora and Funga of Brazil platform. The analyzed specimens were deposited in the HUEG Fungarium of the Universidade Estadual de Goiás.

 Results and discussion

Of the 22 humid chambers prepared, 45% were considered positive, as they presented at least one phase of the myxomycete life cycle. Among these, 20% corresponded to samples from the aphotic zone and 80% from the photic zone. In turn, three species were identified among the specimens, distributed across three genera and three families, along with one unidentified specimen (Fig. 3): Arcyria cinerea (Arcyriaceae), associated with substrates from the aphotic zone; Cribraria elegans Berk. & Curt. (Cribrariaceae) and one unidentified specimen, from the photic zone; and Perichaena depressa Lib. (Trichiaceae), found in substrates from both zones.

10.3897/subtbiol.57.193389.figure3 95D81C84-6157-5D66-AAAC-56740AE42FDC

Sporophores of myxomycetes found. AArcyria cinereaB unidentified specimen CCribraria elegansD, EPerichaena depressa. Scale bar: 0.5 mm.

https://binary.pensoft.net/fig/1717887

Below, we present taxonomic information and comments on the recorded species.

Fungi Anguilliformes Arcyriaceae DA45F895-1666-54B0-952C-E45E62582434 Arcyria cinerea (Bull.) Pers. Fig. 3A Recorded as sporophore.

Sporophore stalked. Stalk: pale yellow, darker at base. Sporotheca: cylindric to subcylindric pale grey (1B1). Peridium: simple, partially fugacious except for the base, forming a small calyculus. Hypothallus: brown, membranous. Capillitium: abundant, elastic, firmly attached to calyculus, pale yellow to hyaline, highly branched, spinulose to smooth. Spores: globose to subglobose, pale yellow to hyaline, with minute spines to nearly smooth, 6.5–8.0 µm.

 Substrate.

leaf litter

 Material examined.

HUEG 22235.

 Geographic distribution.

Americas (Lado and Basanta 2008; Rincón-Marín et al. 2021; Salazar-Hernández et al. 2021; Stephenson et al. 2021); Africa (Ndiritu et al. 2009; Kryvomaz et al. 2020); Europe (Bortnikov et al. 2020; Cazabonne et al. 2021); Asia (Ranade et al. 2012; Nguyen et al. 2019; Stephenson 2021); Oceania (Stephenson 2021). In Brazil: There are records of occurrence in all biomes (Amazon, Atlantic Forest, Caatinga, Cerrado, Pampa and Pantanal) (Bezerra et al. 2007, 2010; Alves et al. 2010; Cavalcanti 2010; Agra et al. 2014; Bezerra et al. 2014; Cavalcanti et al. 2014).

Although the material is consistent with the morphological concept of A. cinerea, the name is treated here in a broad sense (sensu lato) due to the ongoing revision of the species complex.

 Fungi Cribrariales Cribrariaceae 150B9AC6-AD12-5FE9-B77D-559BE6EF3029 Cribraria elegans Berk. & Curt Fig. 3C Recorded as sporophore.

Sporophore stalked. Stalk: slender, blackish purple. Sporotheca: globose, dark purplish-brown, erect or nodding. Peridium: cup occupying 1/2 of the sporotheca, nodes of the peridial net expanded, flat, polygonal and scarcely thickened. Spores: pale yellow to hyaline globose, minutely warted, 5.6–6.5 μm.

 Substrate.

decaying wood fragments.

 Material examined.

HUEG 22236.

 Geographic distribution.

Americas (Lado and Basanta 2008); Europe (Bochynek 2015; Bortnikov et al. 2020). In Brazil: found only in the Cerrado biome (Putzke 1996; Maimoni-Rodella 2002).

Many of the sporophores found formed aggregates on the wood fragment. Although it has a cosmopolitan distribution, in Brazil its occurrence is restricted to the Cerrado biome. In this study, it was observed in substrates associated with the photic zone of the cave, representing the first record of the species and of the Cribrariaceae family in a cave environment.

 Fungi Anguilliformes Trichiaceae AEE3E1CE-C0A7-505A-A398-0F3270358F84 Perichaena depressa Lib. Fig. 3D, E Recorded as sporophore.

Sporophore: sessile, short plasmodiocarps. Sporotheca: strongly flattened, reddish brown or blackened brown (6F4). Peridium: double, external layer cartilaginous, firm, opaque, thickened with granules, inner layer membranous, translucent, brilliant, light yellow (3A4); dehiscence circumscissile along the line formed just by the inner layer of the peridium. Capillitium: tubular, abundant, filaments branched, minutely warted or, more frequently, with regularly spaced constrictions. Spores: subglobose, pale yellow, warted, 8.3–9.5 µm diam.

 Substrate.

leaf litter and decaying wood fragments.

 Material examined.

HUEG 22237; HUEG 22238, HUEG 22239, HUEG 22240.

 Geographic distribution.

Americas (Lado and Basanta 2008; Rincón-Marín et al. 2021; Salazar-Hernández et al. 2021; Stephenson et al. 2021); Africa (Ndiritu et al. 2009; Kryvomaz et al. 2020); Europe (Bortnikov et al. 2020; Cazabonne et al. 2021); Asia (Ranade et al. 2012; Nguyen et al. 2019; Stephenson 2021); Oceania (Stephenson 2021). In Brazil: found in all biomes (Amazon, Atlantic Forest, Caatinga, Cerrado, Pampa, and Pantanal) (Bezerra et al. 2007, 2010; Alves et al. 2010; Cavalcanti 2010; Cavalcanti et al. 2016; Parentes and Cavalcanti 2023).

This species was the most frequent in the sampling. It should be noted that the sporotheca of some samples presented a light gray coloration and a powdery appearance, giving the surface an appearance similar to a thin layer of sand; while the dehiscence and micromorphological characteristics were consistent with the species description.

Unidentified specimen

Fig. 3B

Recorded as sporophore. Sporophore stalked. Stalk: short, black, thick. Sporotheca: oval to subglobose, black (26F1). Peridium: fugacious.

Substrate. leaf litter.

Material examined. HUEG 22241.

The material found was aborted, preventing its identification; however, a preliminary assessment suggests that it may belong to the family Stemonitaceae.

The literature on myxomycetes in cave environments is still incipient, with a reduced number of taxa recorded in these ecosystems. There is only one study on the subject in Brazil to date, conducted in the karst region of Pains, in Minas Gerais, in which three species of the family Trichiaceae were recorded associated with decaying wood fragments located in skylight areas, where there is greater deposition of plant material and light incidence in relation to other sections of the cave environment (Moreira et al. 2024).

Similarly, myxomycete sporophores were found in caves of the Rio Camuy system in Puerto Rico on decaying wood and other organic substrates, such as leaf litter and bat bones (Nieves-Rivera 2003). These records indicate that the occurrence of these organisms in subterranean environments is associated with the entry of allochthonous organic matter, transported by rainwater, wind, or faunal action (Hoffmann et al. 1986; Landolt et al. 1992; Nieves-Rivera 2003), from which propagules can germinate and give rise to plasmodia inside the cavities. This development can also occur on mineral surfaces, including speleothems, where there are records of the formation of fruiting bodies, as observed for F. septica in the field (Galán 2015). Considering the inherent limitations of humid chambers, in which development may occur from propagules present in the substrate under favorable conditions, these records should be interpreted with caution, since detection in culture does not necessarily imply completion of the life cycle in the natural environment.

Sporophore formation in humid chambers prepared with leaf litter and decomposing wood fragments evidenced the presence of viable propagules associated with these materials. This finding suggests that such organic materials may act as potential vehicles for the introduction of myxomycete propagules into cave environments, as previously indicated by Hoffmann et al. (1986) and Nieves-Rivera (2003), even when fruiting is not directly observed under natural conditions. In this sense, humid chambers provide a useful complementary approach for detecting viable propagules associated with cave substrates, favoring the development of reproductive structures under controlled conditions and allowing the recording of species that may not be observed directly in the field.

The data from this study expand current knowledge of myxomycetes associated with cave environments by recording representatives of Cribrariaceae and the Perichaena genus in this type of environment for the first time, as well as A. cinerea for the first time in a Brazilian Cerrado cave. Until then, records in caves mainly focused on representatives of the orders Trichiales, Physarales, and Stemonitales, belonging to the families: Arcyriaceae (A. denudata), Trichiaceae (H. calyculata, M. vesparia), Didymiaceae (Didymium trachysporum G. Lister), Physaraceae (F. septica) and Stemonitaceae (Comatricha sp., Stemonitis cf. herbatica Peck), reported in caves in Brazil (Moreira et al. 2024), Porto Rico (Nieves-Rivera 2003), Spain (Galán 2015), the United States (Landolt et al. 1992) and Mexico (Hoffmann et al. 1986). This scenario reinforces the importance of new surveys in different regions and types of underground environments to broaden understanding of the distribution and diversity of this group.

The conservation of these environments is fundamental considering that the combination of monitoring, tourism management, scientific research, and environmental education is essential to guarantee preservation of communities of myxomycetes, fungi, and other microorganisms involved in decomposing organic matter and consequently in maintaining the ecological balance of underground environments. Studies conducted in Brazilian caves indicates that tourism and mining activities have the potential to alter habitats and compromise cave communities, making continuous monitoring of these environments and adopting preventive measures before significant impacts occur essential (Taylor et al. 2013; Cunha et al. 2020). In this sense, the proximity of the Vila Propício caves to areas of intense mining activity raises concerns about possible impacts on cave biodiversity, including myxomycetes.

 Conclusion

This study reports, for the first time, the occurrence of myxomycetes in a Cerrado cave, including the first record of Cribrariaceae and Perichaena depressa in this type of environment. These findings expand current knowledge of the group in subterranean systems and contribute to the advancement of research in cave environments, reinforcing the importance of further studies to better understand the distribution of myxomycetes under these conditions. The study also highlights the importance of complementary approaches and the need for field investigations to improve our understanding of the diversity and distribution of myxomycetes in subterranean environments.

 Acknowledgements

We thank the edital Pró-Programas n° 01/2024, termo de fomento n° 98-2024, processo sei n° 202400020010680, for funding the publication of this article; the Fundação de Amparo à Pesquisa do Estado de Goiás (FAPEG) for the junior postdoctoral fellowship awarded to ICM and for funding this research through the Taxonomy Support Program (PROTAX); the Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq) for the research productivity grant awarded to SX-S and the scientific initiation fellowship awarded to RMS; and the Coordenação de Aperfeiçoamento de Pessoal de Nível Superior (CAPES) for the junior postdoctoral fellowship awarded to LL-S.

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