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Original Article • Braz. J. Biol. 85 • 2025 • https://doi.org/10.1590/1519-6984.294155 linkcopy

Open-access External and internal architecture of nests of fungus-growing ants of the genera Cyphomyrmex, Mycetarotes, Mycocepurus, Myrmicocrypta, and Trachymyrmex (Hymenoptera: Formicidae)

Arquitetura externa e interna dos ninhos de formigas cultivadoras de fungo dos gêneros Cyphomyrmex, Mycetarotes, Mycocepurus, Myrmicocrypta e Trachymyrmex (Hymenoptera: Formicidae)

AuthorshipSCIMAGO INSTITUTIONS RANKINGS

Abstract

Studying the nest structures of fungus-growing ants of the genera Cyphomyrmex, Mycetarotes, Mycocepurus, Myrmicocrypta, and Trachymyrmex is essential to understanding their ecological role. Structural variations affect adaptation of these ants and their impact on agriculture and forest ecosystems. The internal and external structures of 49 fungus-growing ant nests were marked over three months, with four, seven, 31, four, and three nests, respectively, of the genera Cyphomyrmex, Mycetarotes, Mycocepurus, Myrmicocrypta, and Trachymyrmex (Hymenoptera: Formicidae: Attini) were described. The study were studied in the João Dias da Silveira Campus (Bela Vista) of the São Paulo State University (UNESP) in Rio Claro, São Paulo state, Brazil (22°23'73” S and 47°32'53” W). Forty-nine nests were marked over three months, with four, seven, 31, four, and three nests, respectively, of the genera Cyphomyrmex, Mycetarotes, Mycocepurus, Myrmicocrypta, and Trachymyrmex Four spherical chambers were observed in Cyphomyrmex nests, with height, width, length, and depth, respectively, of 2.8 cm, 2.5 cm, and 2.1 cm, and 15 cm to 30 cm. A single spherical chamber at a depth from 10 to 25 cm was observed in a Mycetarotes sp. nest and in each of the 31 nests of Mycocepurus sp. and Myrmicocrypta sp., with height, width, length, and depth, respectively, of 3.6 cm, 4.5 cm, and 4.1 cm, and 10 to 110 cm; and 5 cm, 5.9 cm, 6.3 cm, and 10 cm to 72 cm. The number of fungus chambers ranged from one to four per Trachymyrmex sp. nest, at depths between 15 cm and 205 cm and height, width, and length, respectively, of 3.9 cm, 7.2 cm, and 7.9 cm. Variations in the internal and external structures of Attini ant nests indicate that they should be considered to understand the ecology of these insects

Keywords:
ant ecology; forest ecosystems; nest architecture

Resumo

O estudo das estruturas dos ninhos de formigas cultivadoras de fungo dos gêneros Cyphomyrmex, Mycetarotes, Mycocepurus, Myrmicocrypta e Trachymyrmex é essencial para compreender seu papel ecológico. As variações estruturais afetam a adaptação dessas formigas e seu impacto na agricultura e nos ecossistemas florestais. As estruturas internas e externas de 49 ninhos de formigas cultivadoras de fungo foram marcadas ao longo de três meses, sendo descritos, respectivamente, quatro, sete, 31, quatro e três ninhos dos gêneros Cyphomyrmex, Mycetarotes, Mycocepurus, Myrmicocrypta e Trachymyrmex (Hymenoptera: Formicidae: Attini). O estudo foi realizado no Campus João Dias da Silveira (Bela Vista) da Universidade Estadual Paulista (UNESP), em Rio Claro, estado de São Paulo, Brasil (22°23'73” S e 47°32'53” W). Nos ninhos de Cyphomyrmex, foram observadas quatro câmaras esféricas, com altura, largura, comprimento e profundidade, respectivamente, de 2,8 cm, 2,5 cm, 2,1 cm e de 15 cm a 30 cm. Em um ninho de Mycetarotes sp. e em cada um dos 31 ninhos de Mycocepurus sp. e Myrmicocrypta sp., foi observada uma única câmara esférica a uma profundidade de 10 a 25 cm, com altura, largura, comprimento e profundidade, respectivamente, de 3,6 cm, 4,5 cm, 4,1 cm e de 10 cm a 110 cm; e 5 cm, 5,9 cm, 6,3 cm e de 10 cm a 72 cm. O número de câmaras de fungo variou de uma a quatro por ninho de Trachymyrmex sp., com profundidades entre 15 cm e 205 cm e altura, largura e comprimento, respectivamente, de 3,9 cm, 7,2 cm e 7,9 cm. As variações nas estruturas internas e externas dos ninhos das formigas Attini indicam que essas características devem ser consideradas para compreender a ecologia desses insetos.

Palavras-chave:
ecologia de formigas; ecossistemas florestais; arquitetura de ninhos

1. Introduction

Ant nests are primarily located underground, as well as in rotten logs, plant parts, and under leaves (Weber, 1945, 1972; Pereira-da-Silva et al., 1981; Mayhé-Nunes, 1995a). Soil excavated from chambers and channels by ant workers, along with plant debris such as straw, plant leaves, and/or twigs, forms the external appearance of Attini ant nests, with only one entrance hole in species of more primitive genera (Weber, 1945, 1972; Pereira-da-Silva et al., 1981; Mayhé-Nunes, 1995a).

Some species of leaf-cutting ants of the genus Acromyrmex and all of the genus Atta deposit soil in mounds above the nest underground, facilitating their identification in the field. An external mound of soil was also reported for species of primitive genera such as Mycetarotes, Mycocepurus, Sericomyrmex, and Trachymyrmex (Mayhé-Nunes, 1995a; Leal, 1998). Species of these genera supposedly use a wide variety of organic matter as substrate for their fungus gardens, including arthropod feces, wood pellets, insect cadavers, seeds, flower parts, dry leaves, and other plant debris (Mann, 1916; Weber, 1941, 1945, 1947, 1966, 1968, 1969; Hölldobler and Wilson, 1990; Mueller et al., 2005; Santos and Sousa Souto, 2023; Lutinski et al., 2023; Oliveira et al., 2024).

Chambers with fungus culture, workers, brood, and queen; and others with soil or waste (exhausted plant residues, depleted fungus, and ant cadavers), as well as channels or tunnels connecting one chamber to another and to the external area, characterize leaf-cutting ant nests of the genera Acromyrmex and Atta (Mariconi, 1970).

Waste chambers are found in nests of different leaf-cutting ant species, including Acromyrmex balzani, Acromyrmex fracticornis, Acromyrmex landolti, Acromyrmex lobicornis, and Acromyrmex striatus, and Atta colombica, Atta laevigata, Atta sexdens, and Atta texana (Wheeler, 1907; Gonçalves, 1961; Haines, 1978; Fowler, 1979; Von Ihering, 1984; Waller and Moser, 1990; Fargi-Brener, 2000). Waste inside colonies can increase the proliferation of pathogenic bacteria and fungi attracting other animals, while its deposition outside the colony reduces the energy expenditure of chamber excavation and the humidity for the symbiotic fungus (Haines, 1978). Deposition of waste externaly is positively correlated with cleaning behavior and elimination of pathogenic microorganisms (Diehl-Fleig and Araujo, 1996). However, waste deposited externally by ants can attract other organisms (Weber, 1972), such as Apterostigma sp., Cyphomyrmex rimosus, Odontomachus bauri, Pheidole fallax, and Pheidole radoszkowskii, which collected seeds and fruit pieces in the waste of Atta cephalotes (Roberts and Heithaus, 1986).

The structure of Atta nests is the greatest complexity among the Attini, with more than 7,000 chambers and depths of up to 8 meters for Atta laevigata (Moreira et al., 2004a), with internal variations in the shape and location of waste and fungus chambers relative to the external area. This shape of A. sexdens nests is semi-ellipsoidal with arm-like extensions in the waste chambers, both under the mound of loose soil (Pretto, 1996), while those of A. laevigata and A. bisphaerica are spherical (Moreira et al., 2004a, b). Variations in the structure and external shape of nests are used to identify species and subspecies of Acromyrmex, although similar nests are reported for different species of this genus (Gonçalves, 1967; Pacheco and Berti-Filho, 1987).

The structural complexity of nests of primitive Attini, such as those of Cyphomyrmex, Mycetarotes, and Mycetophylax, with few and superficial chambers not exceeding 20 cm deep, differs from those of Atta and Acromyrmex (Mayhé-Nunes, 1995a). Chambers of Mycetosoritis and Mycocepurus are found at 30 to 120 cm deep, and those of Trachymyrmex spp. and Trachymyrmex fuscus at 2.7 to 59 cm and 111 to 208 cm deep, respectively, generally with three chambers and a single entrance (Araújo and Della Lucia, 1997). The objective of this study was to describe the external and internal architecture of ants of fungus-growing ant of the genera Cyphomyrmex, Mycetarotes, Mycocepurus, Myrmicocrypta, and Trachymyrmex (Hymenoptera: Formicidae) in Brazil.

2. Material and Methods

2.1. Study genera and location

The internal and external structures of ant nests of the genera Cyphomyrmex, Mycetarotes, Mycocepurus, Myrmicocrypta, and Trachymyrmex, all fungus-growing Attini, were studied in the João Dias da Silveira Campus (Bela Vista) of the São Paulo State University (UNESP) in Rio Claro, São Paulo state, Brazil (22°23'73” S and 47°32'53” W).

2.2. Nest mapping

Attini workers were followed to the entrance holes of their nests, which were marked with stakes bearing an identification number. Some workers from each nest were collected after offering oats to them while they returned to their respective nests. Forty-nine nests were marked over three months, with four, seven, 31, four, and three nests, respectively, of the genera Cyphomyrmex, Mycetarotes, Mycocepurus, Myrmicocrypta, and Trachymyrmex in Rio Claro, São Paulo state, Brazil.

2.3. Identification

Ant owrkers collected were placed in jars with 80% alcohol, taken to the laboratory, and identified at the genus level using keys (Weber, 1972; Bolton, 1995; Mayhé-Nunes, 1995a). Specimens of these workers were sent to Dra. Ana Eugênia de Carvalho C. Farinha of the Biological Institute of São Paulo, Brazil for identification at species level and deposited in the Adolpho Hempel Entomological Collection of that institution.

2.4. External architecture of the nests

The external area of the nests was calculated by measuring the loose soil mound, using the greatest length and width. The distance between the soil mounds and the nest entrance was not recorded, as only observations were made. The external area of some nests was measured, and their entrance holes photographed.

2.5. Internal architecture of the nests

Trenches with a width of 0.70 m and with variable depth depending on the genus of Attini were manually excavated to describe the internal structure of their nests. Soil layers were removed after openning the trench towards the nest, and the excavation deepened as necessary. The channels were marked with the application of talcum powder using a manual formicide applicator, facilitating the location of the chambers of each nest.

The number, width, height, length, and depth of the chambers of each leaf-cutting ant nest were measured. The material in the chambers, including fungus and larvae, pupae and workers of the ants collected and transferred to artificial nests in the laboratory.

3. Results

3.1. External structure of the nests

The apparent area of loose soil in one of the four nests of Cyphomyrmex sp. was 49 cm2 with a height of 5 cm and an its external structure with small soil granules forming a low mound with an orifice at the top.

Small soil granules forming a small mound with an orifice at the top characterized the external structure of Mycetarotes nests (Figure 1A).

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Figure 1
External aspects of ant nests of the genera Mycetarotes (A), Mycocepurus (B), and Trachymyrmex (C and D) (Hymenoptera: Formicidae).

Entrance orifices surrounded by soil granules or dry leaves and small mounds of loose soil were observed in the nests of Mycocepurus sp. (Figure 1B).

An entrance orifice surrounded by soil granules was observed in a Myrmicocrypta sp. nest.

The apparent area of loose soil of Trachymyrmex sp. nests ranged from 300 to 374 cm2, and the diameter of their entrance orifices from 2.5 to 4 cm. One to three entrance orifices, surrounded by soil granules or dry leaves (Figure 1C and D), and small mounds of loose soil were observed on e nests of a species of this genus.

3.2. Internal structure of the nests

The chambers of the four Cyphomyrmex sp. nests (Table 1) were spherical, with a height, width, length, and depth of 2.8 cm, 2.5 cm, 2.1 cm, and 15 to 30 cm, respectively (Table 1). Part of the fungus garden was suspended from roots with the other part at the base of the chamber.

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Table 1
Number of chambers (N), chamber (C), and height (H), width (W), length (L), and depth (D), in centimeters, of the nest chambers of the ant species of the genera Cyphomyrmex sp., Myrmicocrypta sp., and Trachymyrmex sp. in the Rio Claro region, São Paulo state, Brazil.

A single spherical chamber, between 10 cm and 25 cm deep, with a height, width, and length of 3.6 cm, 4.5 cm, and 4.1 cm, respectively, and channels with a diameter of 0.5 cm, was observed in the nests of Mycetarotes sp. (Table 2 and Figure 2A).

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Table 2
Height (H), width (W), length (L), and depth (D.), in centimeters, of the nest chambers of ants of the genera Mycocepurus sp. and Mycetarotes sp. in the Rio Claro region, São Paulo state, Brazil.
thumbnail of Figure 2
Figure 2
Internal structure of ant nests of the genera Mycetarotes (A), Mycocepurus (B), and Trachymyrmex (C), and the fungus chamber of Trachymyrmex (D).

A single fungus chamber per nest (n= 31), spherical in shape, with a height, width, length, and depth of 3.6 cm, 4.5 cm, 4.1 cm, and 10 to 110 cm, respectively, and channels with a diameter of 0.5 cm, were observed in the nests of Mycocepurus sp. (Table 2 and Figure 2B).

A single spherical chamber with a height, width, length, and depth of 5 cm, 5.9 cm, 6.3 cm, and 10 to 72 cm, respectively, was observed per nest of Myrmicocrypta sp. (Table 1).

Four fungus chambers with an ellipsoidal shape, flat base, and a height, width, length, and depth of 3.9 cm, 7.2 cm, 7.9 cm, and 15 to 205 cm, respectively, with the fungus garden at the base or suspended from the top, were observed in each of the three Trachymyrmex sp. nests (Table 1) (Figure 2C, D).

4. Discussion

4.1. External structure of the nests

Soil removed by the workers during the excavation of the chambers and channels in the ground or straw or twigs, along with one small entrance and exit form the external structure of the ant nests were similar to that reported for Acromyrmex spp. (Weber, 1972).

The area of loose soil of one of the four nests of Cyphomyrmex sp., with 49 cm2 and a height of 5 cm, and an external structure with small soil granules forming a small mound and an orifice at the top, is considered simple (Leal et al., 2011; Ramos-Lacau et al., 2012) compared to those of the genera Acromyrmex and Atta (Wirth et al., 2003; Silva Junior et al., 2010). However, similar structures have been reported, such as a single circular entrance holes without a well-formed mound of loose soil, for Cyphomyrmex lectus and Cyphomyrmex morschi (Klingenberg et al., 2007) and for species of other Attini ant genera including Mycocepurus Forel, Mycetarotes Emery, Sericomyrmex Mayr, and Trachymyrmex Forel (Mayhé-Nunes 1995a; Leal et al., 2011). However, this differs from that reported for Cyphomyrmex transversus and Cyphomyrmex cornutus, with distinct nesting habits (Adams and Longino, 2007; Ramos-Lacau et al., 2012). Additionally, large masses of aggregated soil suspended in lower arboreal parts formed nests of Cyphomyrmex cornutus on the Atlantic slopes of Costa Rica (Adams and Longino, 2007). The genus Cyphomyrmex includes species, such as Cyphomyrmex longiscapus, considered ideal models for studies of behavior, cultivation specificity, ecology, mating frequency, etc. (Schultz et al., 2002).

Small soil granules forming a low mound with a hole at the top are the external structure of Mycetarotes sp. nests. This is similar to that reported for nests of this ant, with a single shallow chamber, in secondary forest with clay soil in a trail used by humans (Mayhé-Nunes and Lanziotti, 2004). Entrance orifices surrounded by soil granules or dry leaves, and small mounds of loose soil in Mycocepurus sp. nests are similar to those observed for M. smithii with a single hole. However, the high density and proximity of nest holes can lead to the mistaken conclusion of multiple ones per nest of this ant (Fernández-Marín et al., 2005), confirming reports of intraspecific variation in nest architecture as scarce and with conflicting data that may be correct (Kempf, 1963). A single entrance hole was also reported in nests of M. smithii and Mycocepurus tardus in Costa Rica and Trinidad and for M. goeldii in Brazil and Guyana (Fernández-Marín et al., 2005).

An entrance hole surrounded by soil granules, observed in a Myrmicocrypta sp. nest, agrees to that of Myrmicocrypta camargoi (Sosa-Calvo and Schultz, 2010), with an entrance hole surrounded by soil granules and a single shallow spherical chamber below the soil surface.

The area of loose soil visible in Trachymyrmex sp. nests, from 300 cm2 to 374, with entrance holes between 2.5 and 4 cm in diameter, is similar to that for T. urichi (Weber, 1945) and T. fuscus, with a single entrance hole shaped like a semi-hardened mud straw (Araujo et al., 2002). Trachymyrmex spp. nests are, generally, reported with seven chambers and one or more entrance holes surrounded by soil, reflecting the size of the nests (Weber, 1956, 1966, 1969, 1972). The three-dimensional structure of T. septentrionalis nests, described with one to five vertically connected oval chambers a few centimeters below the soil surface (Tschinkel, 2003), agrees with that of T. fuscus, a species synonymous to T. urichi, with some vertically connected oval chambers a few centimeters below the soil surface (Araujo et al., 2002). Additionally, aspects of nesting are mentioned and used in taxonomic reviews and other general nesting studies (Fernández-Marín et al., 2004; Mayhé-Nunes and Brandão, 2004).

4.2. Internal structure of the nests

A spherical chamber with a height, width, length, and depth of 2.8 cm, 2.5 cm, 2.1 cm, and 15 to 30 cm, respectively, and the fungus garden partially suspended from roots and the remainder at the base in four nests of Cyphomyrmex sp., is similar to the structurally simple nests described for C. transversus, with a single chamber (Ramos-Lacau et al., 2012). However, variations in this parameter were reported for ants of this genus such as C. longiscapus as one of the most unusual among fungus-growing ants, featuring a large entrance hole opening and clay walls often surrounding the nest cavity in a swallow-like style, predominantly in vertical banks along permanent streams (Mueller and Wcislo, 1998).

A single spherical chamber per nest of Mycetarotes sp., at a depth of 10 cm to 25 cm, with a height, width, and length of 3.6 cm, 4.5 cm, and 4.1 cm, respectively, differs in diameter but agrees with that of M. parallelus, with a single spherical chamber (Luederwaldt, 1918) with a width, height, and depth of 4 to 7 cm, 3 to 4 cm, and 10 cm, respectively, in Viçosa, Minas Gerais state, Brazil (Mayhé-Nunes, 1995b). The reduced diameter of Mycocepurus sp. channels is similar to reports for M. smithii with a slight downward inclining directly to the chamber and connecting the external and internal areas of the nest of this latter ant, with the fungus garden at the bottom of the chamber or hanging from grass roots (Luederwaldt, 1918; Mayhé-Nunes, 1995a).

A single spherical fungal chamber in each of the 31 nests of Mycocepurus sp., with a height, width, length, and depth of 3.6 cm, 4.5 cm, 4.1 cm, and 10 to 110 cm, respectively, and channels with a diameter of 0.5 cm, confirms variations in these parameters among species of primitive Attini genera. This contrasts with the more complex nests of Atta, with greater number of chambers at different depths and up to 7 million workers per nest (Moreira and Forti, 1999; Moreira et al., 2004a).

The number of chambers in Mycocepurus sp. nests differs from that reported for M. goeldii, up to four chambers at the depths of 60 cm (Leal, 1998) to 120 cm (Luederwaldt, 1918, 1926). However, the width and height of the chambers of species in this genus were similar to the 10 cm and 20 cm, respectively, reported for both M. goeldii and M. tardus (Luederwaldt, 1918, 1926; Weber, 1972). The fungus garden of Mycocepurus sp., hanging from the top, on roots, or at the base of the chamber, is similar to that of M. goeldii in Brazil (Luederwaldt, 1918, 1926; Weber, 1972; Mayhé-Nunes, 1995a; Leal 1998).

Spherical chambers in Myrmicocrypta sp. nests (5 cm heigh, 5.9 cm wide, and 6.3 cm long) are similar to that of species of this genus in other locations, with chamber width, height, and depth of 6.5 cm, 5.3 cm, and 5.2 to 11 cm, respectively (Weber, 1937, 1941, 1945, 1968). However, up to two chambers at depths of 25 cm to 62 cm and a maximum volume of 358.2 cm3 of loose soil have been reported for nests of ants of the Myrmicocrypta genus in the Brazilian Cerrado (Leal, 1998).

The number of chambers in Trachymyrmex sp. nests, one to four, with height, width, length, and depth of 15 cm, 7.2 cm, 7.9 cm, and 15 cm to 205 cm, respectively, are similar to the four chambers at a maximum depth of 118 cm for species of this genus in the Cerrado biome (Leal, 1998). However, a total of six chambers, between 2.5 cm and 59 cm deep, connected to each other by small-diameter circular channels, usually not exceeding 1 cm in T. fuscus nests and often perpendicular to the soil surface, have been observed (Mayhé-Nunes, 1995a). One to five oval chambers a few centimeters deep and connected vertically to the soil surface characterize the internal structure of T. septentrionalis nests. Four to five chambers were reported for each of the four nests of T. holmgreni excavated in the Restinga (Albuquerque et al., 2018).

5. Conclusion

Variations in the external and internal structures of ant nests within the tribe Attini confirm the behavioral and adaptive diversity of these social insects.

A single chamber per Cyphomyrmex nest, superficially under the soil or beneath stones and roots, and Trachymyrmex nests, with characteristics of primitive and more derived species such as Acromyrmex and Atta, suggest that the second genus is an intermediate stage in the evolution of nest structure within the Attini tribe. This is significant to understanding nesting strategies of fungus-growing ants.

The ecology and excavation behavior of Attini ants highlight the importance of understanding the external and internal structures of their nests to comprehend their social and ecological interactions in natural and modified habitats.

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Publication Dates

  • Publication in this collection
    20 June 2025
  • Date of issue
    2025

History

  • Received
    11 Feb 2025
  • Accepted
    22 Apr 2025
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