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Original Article • Braz. J. Biol. 86 • 2026 • https://doi.org/10.1590/1519-6984.306398 linkcopy

Open-access From Plants to the Fungus Garden: Physical, Chemical, and Behavioral Factors Influencing Foraging in Leaf-Cutting Ants

Das plantas ao jardim de fungo: fatores físicos, químicos e comportamentais que influenciam o forrageamento em formigas cortadeiras

AuthorshipSCIMAGO INSTITUTIONS RANKINGS

Abstract

Leaf-cutting ants selectively forage plant material to cultivate their symbiotic fungus, Leucoagaricus gongylophorus, which constitutes the primary food source for their colonies. Plant selection involves complex interactions among plants, ants, and fungi. The orientation and selectivity of leaf-cutting ants are influenced by physical characteristics of plants, such as hardness and water content, as well as by secondary chemical compounds, including attractants, deterrents, and repellents. In addition, prior experience and learning appear to modify foraging behavior and the selection of more suitable substrates for the symbiotic fungus through a potential feedback mechanism. Consequently, the complexity of foraging behavior varies according to environmental stimuli, ant learning, and the nutritional demands of the colony. Understanding these mechanisms is essential for elucidating the ecological and evolutionary interactions between ants and fungi and for developing more sustainable management strategies for leaf-cutting ants. This mini-review summarizes the multiple factors that influence the foraging behavior of leaf-cutting ants.

Keywords:
ant-fungus interaction; foraging behavior; leaf-cutting ants; learning; plant selection; symbiosis

Resumo

As formigas cortadeiras forrageiam seletivamente material vegetal para cultivar seu fungo simbionte, Leucoagaricus gongylophorus, que constitui a principal fonte de alimento para suas colônias. A seleção de plantas envolve interações complexas entre plantas, formigas e fungos. A orientação e a seletividade das formigas cortadeiras são influenciadas por características físicas das plantas, como dureza e teor de água, bem como por compostos químicos secundários, incluindo atrativos, deterrentes e repelentes. Além disso, a experiência prévia e o aprendizado parecem modificar o comportamento de forrageamento e a seleção de substratos mais adequados para o fungo simbionte por meio de um possível mecanismo de retroalimentação. Consequentemente, a complexidade do comportamento de forrageamento varia de acordo com os estímulos ambientais, o aprendizado das formigas e as demandas nutricionais da colônia. Compreender esses mecanismos é essencial para elucidar as interações ecológicas e evolutivas entre formigas e fungos e para o desenvolvimento de estratégias mais sustentáveis de manejo de formigas cortadeiras. Esta mini-revisão sintetiza os múltiplos fatores que influenciam o comportamento de forrageamento das formigas cortadeiras.

Palavras-chave:
interação formiga-fungo; comportamento de forrageamento; formigas cortadeiras; aprendizado; seleção de plantas; simbiose

1. Introduction

Leaf-cutting ants of the genera Atta, Acromyrmex, and Amoimyrmex, comprising 53 species and 26 subspecies (Cristiano et al., 2020; Bolton, 2025), are major pests in forest plantations, agricultural crops, and pastures (Britto et al., 2016), while also playing important roles in nutrient cycling and seed dispersal (Schaefer et al., 2021; Oliveira et al., 2023). The coevolutionary trajectory between ants and their cultivated fungi has resulted in an obligate mutualism, with multiple factors influencing the foraging behavior of these insects (Barrera et al., 2022). This mini-review addresses the main factors that influence the foraging behavior of leaf-cutting ants.

1.1. Origin of the ant-fungus symbiosis

The proliferation of fungi approximately 66 million years ago, associated with an asteroid impact at the end of the Cretaceous that temporarily disrupted photosynthesis and caused global mass extinctions, favored the emergence of interactions between ants and fungi (Schultz et al., 2024). An ancestral fungal population, cultivated as an obligate mutualist, originated in seasonally dry habitats and subsequently expanded throughout South America, likely becoming isolated from its free-living congeners inhabiting humid forests (Schultz et al., 2024). Fungus-growing ants arose in a single evolutionary event, giving rise to 247 species, primarily in the Neotropics (Schultz and Brady, 2008), all of which maintain obligate relationships with symbiotic fungi. These associations resulted in four distinct agricultural systems within the Agaricales (Mehdiabadi and Schultz, 2010). Lower agriculture is ancestral and involves the cultivation of Agaricaceae fungi (Branstetter et al., 2017). Yeast agriculture is characterized by the cultivation of yeast-like fungi (Branstetter et al., 2017). Coral-fungus agriculture occurs in ants that cultivate Pterulaceae fungi (Branstetter et al., 2017). Higher agriculture comprises 113 ant species that cultivate multinucleate or polyploid agaricaceous fungi, which produce nutritive structures known as gongylidia (Branstetter et al., 2017; Leal-Dutra et al., 2023). Leaf-cutting ants represent a highly specialized lineage within higher agriculture. They use fresh vegetation as a substrate to cultivate Leucoagaricus gongylophorus, enabling extensive ecological expansion and the formation of highly complex colonies, comparable to the most organized societies among non-human animals.

2. Physical Factors of Plants

Anatomical features, including physical characteristics of plants, are associated with acceptance or rejection and directly affect the selection of plant substrates by leaf-cutting ants (Cherrett, 1968; Rockwood, 1975; Littledyke and Cherrett, 1978; Pollard et al., 1983; Forti, 1985; Vitório, 1996; Garcia, 1997). Physical factors such as tissue hardness and the production of resins or latex influence plant material selection by leaf-cutting ants. For example, Atta cephalotes (Hymenoptera: Formicidae) preferentially selects softer, less dense, and more vigorous plant tissues, possibly due to their higher water or liquid content (Cherrett and Seaforth, 1970; Barrer and Cherrett, 1972; Littledyke and Cherrett, 1978; Waller, 1982; Rockwood, 1976; Nichols-Orians and Schultz, 1989).

3. Chemical Factors of Plants

The stimulus generated by a substrate that enables ants to recognize food can be classified into different categories: (a) attractive chemical compounds, which induce ant movement toward the material to be foraged, whereas repellent compounds inhibit its transport; (b) arrest compounds, which interrupt ant traffic and promote investigation of the material; and (c) loading, cutting, and incorporation of the substrate, which depend on the balance between repellent and arrest compounds present in the material to be foraged (Howard et al., 1988; Littledyke and Cherrett, 1978). Although the mechanisms underlying stimuli perception and odor learning remain poorly understood, leaf-cutting ants orient themselves against airflow during foraging to receive olfactory cues (Littledyke and Cherrett, 1978) and use learned information to select plant material (Roces, 1990). Ants preferentially select plants containing attractive or stimulatory compounds, such as a variety of plant-derived substances, particularly specific fatty acids and volatile organic compounds. These compounds may mask repellent or toxic substances that are harmful to the ants and/or their symbiotic fungus, as reported for sesame (Sesamum indicum) (Hebling-Beraldo et al., 1986; Bueno et al., 1995; Pagnocca et al., 1996; Costa et al., 1997; Sinhori et al., 1997), castor bean (Ricinus communis) (Fernandes et al., 1997), Canavalia ensiformis (Takahashi-Del-Bianco et al., 1997), and Virola spp. (Pagnocca et al., 1996). Consequently, plant selection may be more closely related to ant preference than to the suitability of the substrate for symbiotic fungal growth (Camargo et al., 2003a). The chemical composition of substrates alone may not fully explain plant selection, which involves complex behaviors related to both the choice of material to be collected and its subsequent modification by the symbiotic fungus (Camargo et al., 2003b). In addition, the nutritional demands of the colony contribute to the high variability in leaf-cutting ant preferences for natural materials and their chemical fractions; consequently, ants tend to explore new alternative substrate sources (Camargo et al., 2003b).

3.1. Lignification and silicon content of plants affecting cutting behavior and worker mandibles

Atta sexdens rubropilosa preferentially forages dicotyledonous plants, whereas Atta bisphaerica primarily exploits grasses (Fowler et al., 1986), a pattern associated with the biomes in which these species occur (Barrera et al., 2022). Differences in leaf-cutting behavior and in the cultivation of the symbiotic fungus by workers are largely related to the morphological structure of the leaf blades of preferred plants. Grass leaves are highly vascularized and contain a large number of lignified cells, whereas dicotyledon leaves exhibit more widely spaced vascular bundles and fewer cells with lignified walls (Figures 1 and 2). Structural leaf parameters (Peeters, 2002) and leaf anatomy are positively correlated with plant responses to herbivory in tropical ecosystems (Corrêa et al., 2008). The lower digestibility associated with greater hardness and resistance to shear in grasses explains why grass-cutting ants process softer plant material than leaf-cutting ants that exploit dicotyledons (Silva et al., 2017). In grasses, epidermal cells with lignified walls constitute the first mechanical barrier, protecting leaves against herbivory and pathogen attack (McNaughton and Tarrants, 1983). Glandular and non-glandular trichomes, calcium crystals, and secondary metabolites also play important roles in plant defense against herbivores and pathogens (McNaughton and Tarrants, 1983). In addition, cells with lignified walls increase the degree of processing of plant fragments by leaf-cutting ants during fungus garden cultivation (Silva et al., 2017). Silicon concentration in leaf tissues represents an important defense mechanism against herbivory (Silva et al., 2017). Silicon levels are generally higher in grasses than in most dicotyledons and vary among plant species foraged by leaf-cutting ants. Higher concentrations occur in Paspalum spp. (≈10,500 mg kg−1), followed by Gmelina sp. (≈9,600 mg kg−1), whereas lower concentrations are found in woody and shrubby plants such as Ligustrum sp. (≈6,500 mg kg−1), Acalypha sp. (≈6,800 mg kg−1), and Eucalyptus sp. (≈7,000 mg kg−1), with intermediate values in Citrus sp. (≈7,900 mg kg−1) relative to grasses and dicotyledons (Silva et al., 2017). These physiological differences suggest that silicon accumulation may play a key role in determining the selection of foliar substrates by leaf-cutting ant colonies. Silicon deposition in plant tissues is an effective defensive strategy that reduces herbivory by both vertebrates and invertebrates (Vicari and Bazely, 1993). Deposited in the form of phytoliths, silicon increases tissue resistance to shear and exerts an abrasive effect on cutting structures such as mandibles (Figure 3), thereby functioning as a feeding deterrent (Massey and Hartley, 2006). This mechanical wear is ecologically significant in leaf-cutting ants: individuals exposed to cutting more highly silicified plants experience greater mandibular abrasion, which reduces foraging efficiency and limits the supply of plant material to the symbiotic fungus (Silva et al., 2017). Conversely, insects may incorporate inorganic elements into their mandibles, increasing hardness and durability and thereby reducing abrasive damage (Silva et al., 2017). In Atta sexdens rubropilosa, a positive correlation between zinc content and mandibular tooth hardness indicates mineral enrichment as a mechanical strategy (Schofield et al., 2002). However, this phenomenon has not yet been demonstrated in grass-cutting species, which regularly exploit plants with high silicon content, nor have alternative mechanisms that compensate for silicon-induced wear been clearly identified (Silva et al., 2017). Overall, variation in silicon content among plant species influences resource selection behavior by leaf-cutting ants and shapes the coevolutionary dynamics among plants, ants, and their mutualistic fungus. Understanding these interactions is fundamental for explaining foraging patterns, mandibular physiology, and plant defensive strategies in tropical ecosystems (Silva et al., 2017).

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Figure 1
Micrographs of transverse sections of leaf blades. Acalypha sp. General view with a uniseriate epidermis and dorsiventral mesophyll. Note idioblasts with calcium oxalate crystals in the mesophyll (A). Midrib with epidermis, cortex with collenchyma and parenchyma, and the vascular system (B). Gmelina sp. general view with a uniseriate epidermis with glandular and non-glandular trichomes, and dorsiventral mesophyll. Note the extension of the vascular bundle sheath (C). Section of the midrib with a uniseriate epidermis with trichomes, cortex composed of collenchyma and parenchyma, and the vascular system (D). Eucalyptus sp. general view with a uniseriate epidermis, mesophyll with palisade parenchyma cells and oil cavities, and vascular bundles (E). Midrib with a uniseriate epidermis, cortex with collenchyma and parenchyma, and a vascular system with a well-defined cambial zone (F). BSE: vascular bundle sheath extension; CO: collenchyma; CR: crystal; CZ: cambial zone; NT: non-glandular trichome; OC: oil cavity; PH: phloem; PP: palisade parenchyma; SP: spongy parenchyma; GT: glandular trichome; VB: vascular bundle; XY: xylem. Scale bars: 12, 14, 16= 100 μm; 13, 15, 17= 150 μm.
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Figure 2
Micrographs of transverse sections of leaf blades. Citrus sp. general view with a uniseriate epidermis with idioblasts containing calcium crystals, dorsiventral mesophyll with oil cavities, idioblasts with raphides, and vascular bundles (A, B). Midrib showing a uniseriate epidermis, cortex with collenchyma and parenchyma, and a vascular system surrounded by a sheath of lignified fibers. Note the well-developed cambial zone (C). Ligustrum sp. general view with a uniseriate epidermis with sparse glandular trichomes and dorsiventral mesophyll (D). Midrib with a uniseriate epidermis, cortex with collenchyma and parenchyma, and a vascular system with a well-developed cambial zone (E). Paspalum sp. sections with common cells with lignified walls, bulliform cells and stomata in the epidermis; mesophyll with chlorenchyma arranged radially around the vascular bundles. Note the endodermis with large cells surrounding the vascular bundles and groups of fibers beneath the epidermis (F, G). BC: bulliform cells; CO: collenchyma; CP: chlorenchyma; CR: crystal; CZ: cambial zone; EN: endodermis; GT: glandular trichome; OC: oil cavity; FI: fibers; PA: parenchyma; PH: phloem; PP: palisade parenchyma; RP: raphides; SP: spongy parenchyma; VB: vascular bundle; XY: xylem. Scale bars: 18, 19, 21, 23, 24= 100 μm; 20, 22= 150 μm.
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Figure 3
Anteroposterior surface of the right mandible of gardeners of Atta bisphaerica (A) and Atta sexdens rubropilosa (B). Anteroposterior surface of the right mandible of generalist workers of Atta bisphaerica (C) and Atta sexdens rubropilosa (D). Anteroposterior surface of the right mandible of foragers of Atta bisphaerica (E) and Atta sexdens rubropilosa (F). Anteroposterior surface of the right mandible of soldiers of Atta bisphaerica (G) and Atta sexdens rubropilosa (H). Anteroposterior surface of the mandible of foragers of Atta sexdens rubropilosa at different ages with natural wear due to foraging activity. Adult of undetermined age (I) and of one day after emergence (J).

3.2. Energy obtained through cutting selected leaves

Ants generally feed on substances in the liquid phase (Figure 4), which are obtained during scraping and licking of solid particles and semisolid foods with the glossa mouthpart. These materials are temporarily stored in the infrabuccal cavity, a sac-like outgrowth of the hypopharynx lined with bristle hairs that filter food particles. In leaf-cutting ants, the crop also functions as a site for food storage and for the production of digestive enzymes such as amylase, invertase, maltase, and trehalase (Fowler et al., 1991). However, there is no evidence that these enzymes are synthesized in the crop or regurgitated from the midgut (Erthal et al., 2004). Larvae and adult ants generally differ in their feeding modes (Schneider, 2003). In Solenopsis invicta (Hymenoptera: Formicidae), a filter formed by hairs in the mouthparts, located between the preoral cavity and the pharynx, prevents the ingestion of particles larger than 0.88 μm in diameter, whereas larvae are capable of ingesting particles up to 45.8 μm (Glancey et al., 1981). Similarly, workers of Acromyrmex octospinosus (Hymenoptera: Formicidae) filter particles up to 10 μm in diameter (Quinlan and Cherrett, 1978a). Trophallaxis, defined as the regurgitation of liquids stored in the crop and their transfer to nestmates, represents a fundamental mechanism of nutrient exchange in social insects (Richard and Errard, 2009; Moreira et al., 2010, 2015). However, the occurrence of oral trophallaxis in leaf-cutting ants remains debated (Moreira et al., 2006, 2007a, b, 2010, 2015; Richard and Errard, 2009). The frequency of trophallaxis varies among ant species and reflects both phylogenetic position and feeding habits (Wilson, 1971). In Acromyrmex subterraneus subterraneus (Hymenoptera: Formicidae), trophallaxis occurs in 23%, 13%, and 3% of interactions involving liquid, semisolid, and solid diets, respectively, whereas in Atta sexdens rubropilosa (Hymenoptera: Formicidae) the corresponding values are 13%, 3%, and 0% (Moreira et al., 2015). In workers of Acromyrmex subterraneus subterraneus and Acromyrmex octospinosus (Hymenoptera: Formicidae), 63% and 67%, respectively, of foragers returning to the nest transfer collected liquids to nestmates via trophallaxis, a behavior that is rarely observed in non-foraging individuals (Richard and Errard, 2009). Liquid ingestion rates in nectar-feeding species also vary, being higher in workers of Camponotus rufipes (Formicinae) and Pachycondyla villosa (Ponerinae) than in Atta sexdens (Myrmicinae) and in predatory ants of the Rhytidoponera impressa complex (Ponerinae). These differences in feeding habits are associated with morphological traits, such as the greater liquid storage capacity observed in Camponotus species, which possess a more developed crop (Paul and Roces, 2003). Workers of Atta cephalotes (Hymenoptera: Formicidae) ingest water during dry periods or under laboratory conditions to compensate for water deficiency (Weber, 1972a). Unladen foragers have been associated with the transport, consumption, and assimilation of liquids, particularly carbohydrate-rich plant nectar (Wirth et al., 2003), although there is no evidence that this represents a common behavior under field conditions (Rytter and Shik, 2016). During the preparation of plant substrates for incorporation into the fungus garden, workers ingest liquid substances and then leave the nest to forage with a full midgut, consuming these reserves during foraging activity (Schilman and Roces, 2008; Rytter and Shik, 2016). Overall, the available evidence supports the hypothesis that oral trophallaxis occurs at low frequency or may be absent in this group of leaf-cutting ants.

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Figure 4
Liquid foraging by leaf-cutting ant workers: group of workers sucking solution (A); forager sucking a solution with rhodamine (B); suction in frontal and lateral views (C and D).

4. Symbiotic and Microbiological Factors

Plant selection during foraging is also closely linked to the nutritional requirements of the symbiotic fungus, as plant material serves as the substrate for its growth (Weber, 1972a). Glycogen represents the primary energy reserve of the symbiotic fungus Leucoagaricus gongylophorus (Fisher et al., 1994; Castillo-Alfonso et al., 2024), particularly during the early stages of gongylidia formation. Gongylidia are swellings in the central or terminal portions of hyphae, associated with staphylae and polysaccharides in forms readily assimilable by ants (Weber, 1972a; Quinlan and Cherrett, 1979) (Figure 5). Staphylae constitute important nutritional supplements, containing crude protein, carbohydrates, and lipids, including essential steroids, which represent approximately 24%, 2%, and 27% of the dry weight of cultivated fungus, respectively (Mueller et al., 2001; Mueller, 2002). The symbiotic fungus, which is the sole food source for larvae, provides a rich and nutritionally complete diet on which larvae depend exclusively (Weber, 1972b; Quinlan and Cherrett, 1979; Hölldobler and Wilson, 1990).

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Figure 5
Isolate of the symbiotic fungus of Acromyrmex subterraneus brunneus; culture medium (A); staphylae removed from the fungus under laboratory conditions (10× magnification) (B) and in the field (10× magnification) (C); (D) gongylidia of the laboratory isolate (20× magnification).

Leaf-cutting ants avoid plants containing chemical substances harmful to their fungus through grooming and trophallaxis. By circulating secondary compounds among nest workers, ants can assess and regulate plant selection for foraging (Ridley et al., 1996). In addition, ants obtain information about plant chemical composition by ingesting liquids from plant tissues during foraging (Littledyke and Cherrett, 1976; Andrade, 1997; Forti and Andrade, 1999).

A medium-sized worker of Atta cephalotes (Hymenoptera: Formicidae), with a dry weight of approximately 1.0 mg, consumes about 62.2 μg of carbohydrates in 1 μL of plant liquid during the preparation of Ligustrum ovalifolium leaves, corresponding to an energy intake of 1.064 J (Quinlan and Cherrett, 1979). This amount may satisfy the worker’s basic energetic requirements for up to 24 hours. Plant-derived liquids are distributed to other adult ants not directly involved in leaf preparation through trophallaxis (Cherrett, 1980). Nevertheless, the contribution of the fungus garden to the nutritional and energetic supply of workers may be even greater, with carbohydrates derived from leaves or fungal mycelium accounting for up to 50% of their energetic requirements (Silva et al., 2003).

Citrus pulp is considered one of the most attractive substrates for leaf-cutting ants (Mudd et al., 1978). Whereas Atta cephalotes shown preference for citrus pulp, Acromyrmex octospinosus (Hymenoptera: Formicidae) preferentially carried barley, citrus pulp, lentil, and corn flakes, in this order. Moreover, fungal growth in artificial media containing these substrates was greater with barley than with citrus pulp for both species, indicating that ants do not necessarily select materials that improve fungal growth (Quinlan and Cherrett, 1978b).

The symbiotic fungus may also supplement essential nutrients that insects are unable to synthesize, such as sterols (Cherrett, 1980). The fungus produces ergosterol (Martin et al., 1969), which has been detected in the pharyngeal gland (formerly the postpharyngeal gland) of Acromyrmex octospinosus (Hymenoptera: Formicidae) (Peregrine et al., 1973). Preparation of the substrate for incorporation into the fungus garden in Acromyrmex colonies involves licking the leaf surface and chewing leaf margins, accompanied by liquid ingestion. During this process, workers perceive the chemical composition of the plant material, thereby performing a second selection step (Littledyke and Cherrett, 1976). Five behavioral responses of Acromyrmex octospinosus workers during plant foraging have been described (Knapp et al., 1990): (i) collection, in which all or most plant species are cut and transported to the nest; (ii) delayed rejection Type A, in which workers initially cut and transport leaves but later reject them following evaluation based on information retained by the colony; (iii) delayed rejection Type B, characterized by a low number of foragers on the trail, with foraging reduced or terminated when unfamiliar plant species are encountered; (iv) immediate rejection type A, in which ants do not collect plant material despite partially or completely cutting leaves, and leaf fragments are abandoned or occasionally transported to the refuse pile; and (v) immediate rejection type B, in which neither cutting nor transport of plant material occurs.

Finally, plant selection by leaf-cutting ants may be influenced by previous experiences of workers, as induction and habitat-related effects constitute additional important behavioral factors shaping foraging decisions (Fowler, 1982).

4.1. Behavioral and cognitive factors

The foraging preferences of Acromyrmex octospinosus (Hymenoptera: Formicidae) among discs of Dioscorea cayenensis cayenensis, Ipomoea batatas, and Manihot esculenta under laboratory conditions are not random and vary among days and individuals, possibly as a function of worker age or genetic composition. Such variation may be adaptive, increasing the sampling of plant resources available around the nest and reducing the risk associated with reliance on a single plant species as a substrate should it become unavailable (Therrien, 1988).

Dietary conditioning, whereby adults preferentially feed on plant species or varieties previously consumed, is also an important factor influencing insect behavior (Lara, 1979). For example, adults of the Colorado potato beetle, Leptinotarsa decemlineata (Coleoptera: Chrysomelidae), whose larvae developed on Solanum tuberosum and Solanum dulcamara, preferentially fed on these host plants when offered multiple Solanum species (Rossetto, 1973, cited in Lara, 1979). Experience acquired during early life stages may therefore induce environmental preferences in ants. Newly emerged workers of Camponotus vagus and Formica polyctena (Hymenoptera: Formicidae), when conditioned in nests containing thyme for 30 days, were able to recognize presence or absence of this food resource (Jaisson, 1980).

This phenomenon has also been demonstrated in leaf-cutting ants, in which plant selection is influenced by prior experience (Fowler, 1982). Workers of the grass-cutting ant Acromyrmex landolti fracticornis (Hymenoptera: Formicidae) seems to be environmentally induced, probably through experience-based conditioning (Fowler, 1982). The selectivity of scout workers depends strongly on conditioning to specific food sources. In Atta colombica (Hymenoptera: Formicidae), workers cut plants and initiate recruitment more rapidly at familiar food sources than at unfamiliar ones (Howard et al., 1996).

Consequently, leaf-cutting ants are generally slow to accept unfamiliar food sources, given the wide variability in nutritional content and the diversity of secondary compounds among plant species, which complicate their recognition and evaluation (Mattson Junior, 1980). During their foraging lifespan, which has a half-life of approximately four months (Camargo et al., 2007), workers are exposed to a broad range of plant species and environmental stimuli, and the accumulated experience of each individual may reflect a complex pattern of herbivory (Howard et al., 1996). The collective organization of foraging and individual behavioral responses together generate extremely complex foraging patterns in leaf-cutting ants (Roces, 2001).

Responses of scout workers to relatively immutable plant characteristics, such as odors, depend on prior experience and on information regarding plant quality received from nestmates during foraging, as well as on the productivity of the fungus garden (Lopes et al., 2004). The type of material collected for fungal cultivation can, in turn, drive changes in plant selectivity by workers, suggesting that ants actively manipulate the status of the symbiotic fungus (Roces, 2002). Leaf-cutting ants do not directly process the collected plant fragments nor do they transmit information through a linear “working chain” within the nest. Instead, continuous feedback between the fungus and the gardener workers, and between gardeners and foragers, regulates foraging responses (Roces, 2002).

Individual worker preferences broaden the range of plant species exploited, generating colony-level variation in substrate selection that reflects the distribution of plant resources along foraging trails (Therrien, 1988). This pattern may arise from worker specialization in different tasks, ranging from foraging to the removal of dead individuals, as well as from internal colony differences, as observed in Acromyrmex versicolor (Hymenoptera: Formicidae) (Julian and Cahan, 1999). Individual behavioral variability thus increases overall colony flexibility (Schneirla, 1952, cited in Therrien, 1988) and enhances efficiency in the performance of specific tasks (Oster and Wilson, 1978).

5. Concluding Remarks

Physical, chemical, symbiotic, and behavioral factors interact dynamically to shape the foraging behavior of leaf-cutting ants, highlighting a coadaptive evolutionary process among plants, ants, and their symbiotic fungus. Plant selection is therefore closely linked to colony survival and to the stability of this mutualistic relationship. Experimental approaches that integrate chemical ecology, behavioral physiology, and molecular biology are likely to advance our understanding of the factors that govern the foraging behavior of leaf-cutting ants.

Acknowledgements

Luiz Carlos Forti was the recipient of a grant from the Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq) (Grant No. 301938/2017-2). Frederico Guilherme is grateful for financial support from the PELD through CNPq (process 23/2024, Grant No. 445997/2024-9) and FAPEG (process 23/2024, Grant No. 2025.1026.700.1634). Frederico Guilherme also acknowledges support from a CNPq Productivity Fellowship (PQ-C) (Process No. 302200/2025-8).

Data Availability Statement

No new data were generated or analyzed in this study. Data sharing is not applicable to this article.

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Edited by

  • Editor:
    Takako Matsumura Tundisi

Publication Dates

  • Publication in this collection
    03 Aug 2026
  • Date of issue
    2026

History

  • Received
    11 Mar 2026
  • Accepted
    11 June 2026
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This is an Open Access article distributed under the terms of the Creative Commons Attribution license (https://creativecommons.org/licenses/by/4.0/), which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited.
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