|Atta mexicana workers carrying a leaf section|
Fungus-growing ants (subtribe Attina) comprise all the known fungus-growing ant species participating in ant-fungus mutualism. They are known for cutting grasses and leaves, carrying them to their colonies' nests, and growing fungi on them on which they later feed.
Their farming habits typically have large effects on their surrounding ecosystem. Many species farm large areas surrounding their colonies and leave walking trails that compress the soil and no longer grow plants. Attine colonies commonly have millions of individuals, though some species only house a few hundred.
They are the sister group to the subtribe Dacetina. Leafcutter ants, including Atta and Acromyrmex, make up two of the genera. Their cultivars mostly come from the fungal tribe Leucocoprineae of family Agaricaceae.
Attine gut microbiota is often not diverse due to their primarily monotonous diets, leaving them at a higher risk than other beings for certain illnesses. They are especially at risk of death if their colony's fungus garden is affected by disease, as it is most often the only food source used for developing larvae. Many species of ants, including several Megalomyrmex, invade fungus-growing ant colonies and either steal from and destroy these fungus gardens, or they live in the nest and take food from the species.
Fungus-growing ants are only found in the Western Hemisphere. Some species stretch as far north as the pine barrens in New Jersey, USA (Trachymyrmex septentrionalis) and as far south as the cold deserts in Argentina (several species of Acromyrmex). This New World ant clade is thought to have originated about 60 million years ago in the South American rainforest. This is disputed, though, as they likely evolved in a drier habitat while still learning to domesticate their crops.
- 1 Evolution
- 2 Behavior
- 3 Caste system
- 4 Habitat
- 5 Farming
- 6 Genera
- 7 See also
- 8 References
- 9 External links
This section needs expansion. You can help by adding to it. (January 2019)
Early ancestors of attine ants were probably insect predators. They likely began foraging for leaf sections, but then converted their primary food source to the fungus these leaf cuts grew. Higher attines, such as Acromyrmex and Atta, are believed to have evolved in Central and North America about 20 million years ago (Mya), starting with Trachymyrmex cornetzi. While the fungal cultivars of the 'lower' attine ants can survive outside an ant colony, those of 'higher' attine ants are obligate mutualists, meaning they cannot exist without one another.
Generalized fungus farming in ants appears to have evolved about 55-60 Mya, but early 25 Mya ants seemed to have domesticated a single fungal lineage with gongylidia to feed colonies. This evolution of using gongylidia appears to have developed in the dry habitats of South America, away from the rainforests where fungus-farming evolved. About 10 million years later, leaf-cutting ants likely arose as active herbivores and began industrial-scaled farming. The fungus the ants grew, their cultivars eventually became reproductively isolated and co-evolved with the ants. These fungi gradually began decomposing more nutritious material like fresh plants.
Shortly after attine ants began keeping their fungus gardens in dense aggregations, their farms likely began suffering from a specialized genus of Escovopsis mycopathogens. The ants evolved cuticular cultures of Actinobacteria that suppress Escovopsis and possibly other bacteria. These cuticular cultures are both antibiotics and antifungals. The mature worker ants wear these cultures on their chest plates and sometimes on their surrounding thoraces and legs as a biofilm.
Typically, one queen lives per colony. Every year after the colony is about three years old, the queen lays eggs of female and male alates, the reproductive ants that will pass on the genes of the queens. Before leaving the nest, queens stuff some of the fungus' mycelia in her cibarium. These winged males and queens then take their nuptial flights to mate high in the air. In some areas, species flights are synchronized with all local colonies' virgin royalty flying at the same time on the same day, such as Atta sexdens and Atta texana.
Some species' queens mate with only one male, as in Seriomyrmex and Trachymyrmex, while some are known to mate with as many as eight or 10, such as Atta sexdens and many Acromyrmex spp. After mating, all males die, but their sperm stays alive and usable for a long time in the spermatheca, or sperm bank, of their mate, meaning that many ants technically father offspring years after their death.
After their mating flights, queens cast off their wings and begin their descent into the ground. After creating a narrow entrance and digging 20–30 cm (7.9–11.8 in) straight down, she creates a small 6 cm (2.4 in) chamber. In here, she spits on a small wad of fungus and starts her colony's garden. After about three days, fresh mycelia are growing out of the fungus wad and the queen has lain three to six eggs. In a month, the colony has eggs, larvae, and often pupae surrounding the ever-growing garden.
Until the first workers are grown, the queen is the sole worker. She grows the garden, fertilizing it with her fecal liquid, but does not eat from it. Instead, she gains energy from eating 90% of the eggs she lays, in addition to catabolizing her wing muscles and fat reserves.
Though the first larvae feed on the eggs of the queen, the first workers begin growing and eating from the garden. Workers feed malformed eggs to the hungry larvae while the garden is still fragile. After about a week of this underground growth, workers open the closed entrance and begin foraging, staying close to the nest. The fungus begins growing at a much faster rate [13 μm (0.00051 in)] an hour. From this point on, the only work the queen does is egg-laying.
Colonies grow slowly for the first two years of existence, but then accelerate for the next three years. After winged males and queens begin being produced at around five years, the growth of the colony begins to level out.
The founding of a nest by these queens is highly difficult, and successful cases are not likely. After three months, newly founded colonies of Atta capiguara and Atta sexdens are 0.09% and 2.53% likely to still exist, respectively. Some species have better odds, such as Atta cephalotes, which are 10% likely to survive a few months.
Attines have seven castes performing roughly 20-30 tasks, meaning the potential exists for development of more specialized castes performing individual tasks for Atta's future. For now, a reproductive caste, made of male drones and female queens, and a worker class, that vary greatly in size, are known. Queens have much larger ovaries than females in the working castes. Since their needs are constantly taken care of, queens rarely move from a single location, which is typically in a centralized fungal garden. Workers take their eggs and move them to other fungal gardens. Differences in size between worker castes begin to develop after a colony is well established.
Lower attines have very minor polymorphism within the minor workers, though higher attines commonly have very different sizes of worker ants. In the higher attines, though, head width varies eight-fold and dry weight 200-fold between different castes of workers. The size differences in workers is nearly nonexistent in newly founded colonies.
Due to the variety of tasks needed to be performed by a colony, the widths of workers heads are important and good measures of what jobs workers are likely to perform. Those with the heads about 0.8–1.0 mm (0.031–0.039 in) wide tend to work as gardeners, although many with heads 0.8–1.6 mm (0.031–0.063 in) wide participate in brood care.
Workers need heads only about 0.8 mm wide to do the work of caring for the very delicate hyphae of the fungus, which they care for by stroking with their antennae and moving with their mouths. These tiny workers are the smallest and most abundant and are called minim. Ants of 1.6 mm (0.063 in) appear to be the smallest workers that cut vegetation, but they cannot cut very hard or thick leaves. Most foragers have heads oaround 2.0–2.2 mm (0.079–0.087 in) wide.
Attines, particularly the workers that cut leaves and grass, have large mandibles powered by strong muscles. On average, 50% of worker ants' head mass and 25% of their full body mass is the mandibular muscles alone.
Though all castes defend their nests in the event of invasion, a true soldier caste, with individuals called majors, exists. They are larger than other workers, and use their large, sharp mandibles, powered by huge adductor muscles, to defend their colonies from large enemies, such as vertebrates. When a foraging area is threatened by conspecific or interspecific ant competitor, the majority of respondents are smaller workers from other castes, since they are more numerous, and therefore better suited for territorial combat.
Tasks are divided not only by size, but by the age of individuals workers, as well. Young workers of most subcastes tend to work inside the nest, but many older workers take on tasks outside. Minims, whichare too small to cut or carry leaf fragments, are commonly found at foraging sites. They often ride from the foraging site to the nest by climbing onto the fragments carried by other workers. Most likely, they are older workers that defend carriers from parasitic phorid flies that attempt to lay eggs on the backs of the foragers.
All size groups defend their colonies from invaders, but older workers have been found to attack and defend territories most often. At least three of four physical castes of A. sexdens change their behavior based on their age.
Lower attines mostly live in inconspicuous nests with 100-1000 individuals and relatively small fungus gardens in them. Higher attines, in contrast, live in colonies made of 5-10 million ants that live and work within hundreds of interconnected fungus-bearing chambers in huge subterranean nests. Some colonies are so large, they can be seen from satellite photos, measuring up to 600 m3 (21,000 cu ft).
The majority of fungi that are farmed by attine ants come from the family Lepiotaceae, mostly from the genera Leucoagaricus and Leucocoprinus, though variance occurs within the tribe. Some species in the genus Apterostigma have changed their food source to fungi in the family Tricholomataceae. Some species cultivate yeast, such as Cyphomyrmex rimosus.
Some fungi that have supposedly been vertically transmitted are believed to be millions of years old. It was previously assumed that the cultures are always transmitted vertically from colony to young queen, but some lower attines have been found to be growing recently domesticated Lepiotaceae. Some species transfer cultures laterally, such as Cyphomyrmex and occasionally some species of Acromyrmex, whether by joining a neighboring tribe, stealing, or invading another colony's garden.
Lower attines do not use leaves for the majority of the substrate for their gardens, and instead prefer dead vegetation, seeds, fruits, insect feces, and corpses.
The number of ants that are recruited to cut varies greatly based on the leaf quality available in addition to the species and location of the colony. Leaf quality is complex to measure because many variables exist, including "leaf tenderness, nutrient composition, and the presence and quantity of secondary plant chemicals" such as sugar.
Early studies found the pheromones used to mark foraging trails come from poison gland sacs. Studies suggest there are two purposes for marking the trails this way: worker recruitment and orientation cues. The trail recruitment pheromone methyl-4-methylpyrrole-2-carboxylate (MMPC), was the first whose chemical structure was identified. It is also the main trail recruitment phermone in all Atta species except Atta sexdens, which uses 3-ethyl-2,5-dimethylpyrazine.
MMPC is incredibly potent and effective at attracting ants. One milligram is theoretically powerful enough to create a path that A. texana and A. cephalotes would follow three times the Earth's circumference [74,703 miles (120,223 km)] and that 50% of A. vollenweideri foragers would follow 60 times around the Earth [1,494,060 miles (2,404,460 km)].
After following the pheromone trail to vegetation, ants climb onto leaves or grass and begin cutting off sections. To do this, they place one mandible, called the fixed mandible, onto a leaf and anchor it. Then they open the other, called the motile mandible, and place it on the leaf tissue. The ant keeps moves the motile jaw and pulls the fixed jaw behind it by closing them together until the fragment detaches. Which jaw is fixed and which is motile varies depending on the direction in which the ant chooses to cut a fragment.
The sizes of leaf fragments have been found in some studies to vary based on the size of ants due to the ants' anchoring of their hind legs while cutting, though other studies have not found correlations. This is likely because many factors affect how ants cut leaves, including neck flexibility, body axis location, and leg length. Load sizes that do not impact the running speed of the collecting ants are favored.
Often, ants stridulate while cutting vegetation by raising and lowering their gasters in a way that makes a cuticular file on the first gastric tergite and a scraper on the postpetiole rub together. This makes a noise, audible by people with great hearing sitting very close to them and visible using laser-Doppler vibrometry. It also causes the mandibles to move like a vibratome and cut smoother through tender leaf tissue.
The behavior of the foragers that bring the material back to the nest varies greatly among species. In some species, especially those that harvest close to their nests, the harvesters bring the litter back to their colony themselves. Species such as A. colombica have one or more cache sites along a trail for foragers to grab litter. Other species, such as A. vollenweideri, that carry leaves as far as 150 m (490 ft), have two to five carriers per leaf. The first carrier takes the segment a short distance toward the nest and then drops it. Another picks it up and drops it, and this repeats until the last carrier brings it the greatest distance until reaching the nest. Data does not show that this behavior maximizes load transportation, so scientists have explained this behavior in other ways, though the data are still inconclusive. One theory is that this type of task partitioning increases the efficiency of individual workers as they become specialists. Another is that the chains accelerate communication between ants about the quality and species of the plants being cut, recruits more workers, and reinforces territorial claims by reinforcing the scent markings.
First, foragers bring in to and drop leaf fragments on the nest's chamber floor. Workers that are usually slightly smaller clip these pieces into segments that are about 1–2 mm (0.039–0.079 in) across. Smaller ants then crush these fragments and mold them into damp pellets by adding fecal droplets and kneading them. They add the pellets into a larger pile of other prill.
Smaller workers then pluck loose strands of fungus from dense patches and plant them on the surface of the freshly made pile. The smallest workers, the minim, move around and keep up the garden by delicately prodding the piles with their antennae, licking the surfaces, and plucking out the spores and hyphae of unwanted mold species.
Higher attine fungi grow gongylidia, which form clusters called staphylae. The staphylae are rich in carbohydrates and lipids. Though workers can also eat the hyphae of the fungi, which is richer in protein, they prefer staphylae and appear to live longer while eating them.
Cellulose has been found to poorly degraded and assimilated by fungus, if at all, meaning that the ants that eat the fungus do not get much energy from the cellulose in plants. Xylan, starch, maltose, sucrose, laminarin, and glycoside apparently play the important roles in ant nutrition. It is not known yet how ants can digest laminarin, but myrmecologists E.O. Wilson and Bert Hölldobler hypothesize that fungal enzymes may occur in the ants' guts, as evidenced by the enzymes found in larval extract.
In a laboratory experiment, only 5% of workers' energy needs were met by fungal staphylae, and the ants also feed on tree sap as they collect greens. Larvae seem to grow on all or nearly all fungus, whereas queens obtain their energy from the eggs nonqueen females lay and workers feed to them.
The actinomycete bacterium Pseudonocardia is acquired by pupae from the workers that care for them two days after pupae eclose for metamorphosis. Within 14 days, the ants are covered in the bacteria, where they are stored in crypts and cavities found in the exoskeletons. The bacteria produce small molecules that can prevent the growth of a specialized fungus garden pathogen.
Impact of farming
The scale of the farming done by fungus-farming ants can be compared to human's industrialized farming. A colony can "[defoliate] a mature eucalyptus tree overnight". The cutting of leaves to grow fungus to feed millions of ants per colony has a large ecological impact in the subtropical areas in which they reside.
- Acanthognathus Mayr, 1887
- Acromyrmex Mayr, 1865
- Allomerus Mayr, 1878
- Apterostigma Mayr, 1865
- Atta Fabricius, 1804
- †Attaichnus Laza, 1982
- Basiceros Schulz, 1906
- Blepharidatta Wheeler, 1915
- Cephalotes Latreille, 1802
- Chimaeridris Wilson, 1989
- Colobostruma Wheeler, 1927
- Cyatta Sosa-Calvo et al., 2013
- Cyphomyrmex Mayr, 1862
- Daceton Perty, 1833
- Diaphoromyrma Fernández, Delabie & Nascimento, 2009
- Epopostruma Forel, 1895
- Eurhopalothrix Brown & Kempf, 1961
- Ishakidris Bolton, 1984
- Kalathomyrmex Klingenberg & Brandão, 2009
- Lachnomyrmex Wheeler, 1910
- Lenomyrmex Fernández & Palacio, 1999
- Mesostruma Brown, 1948
- Microdaceton Santschi, 1913
- Mycetagroicus Brandão & Mayhé-Nunes, 2001
- Mycetarotes Emery, 1913
- Mycetophylax Emery, 1913
- Mycetosoritis Wheeler, 1907
- Mycocepurus Forel, 1893
- Myrmicocrypta Smith, 1860
- Ochetomyrmex Mayr, 1878
- Octostruma Forel, 1912
- Orectognathus Smith, 1853
- Paramycetophylax Kusnezov, 1956
- Phalacromyrmex Kempf, 1960
- Pheidole Westwood, 1839
- Pilotrochus Brown, 1978
- Procryptocerus Emery, 1887
- Protalaridris Brown, 1980
- Pseudoatta Gallardo, 1916
- Rhopalothrix Mayr, 1870
- Sericomyrmex Mayr, 1865
- Strumigenys Smith, 1860
- Talaridris Weber, 1941
- Trachymyrmex Forel, 1893
- Tranopelta Mayr, 1866
- Wasmannia Forel, 1893
- Xerolitor Sosa-Calvo et al., 2018
- Bolton, B. (2015). "Attini". AntCat. Retrieved 18 August 2015.
- Hölldobler, Bert; Holldobler, Foundation Professor of Biology Bert; Wilson, Honorary Curator in Entomology and University Research Professor Emeritus Edward O.; Wilson, Edward O. (2009). The Superorganism: The Beauty, Elegance, and Strangeness of Insect Societies. W. W. Norton & Company. ISBN 9780393067040.
- Branstetter, M. G.; Ješovnik, A.; Sosa-Calvo, J.; Lloyd, M. W.; Faircloth, B. C.; Brady, S. G.; Schultz, T. R. (2017-04-12). "Dry habitats were crucibles of domestication in the evolution of agriculture in ants". Proceedings of the Royal Society B: Biological Sciences. 284 (1852): 20170095. doi:10.1098/rspb.2017.0095. PMC 5394666. PMID 28404776.
- Weber, N.A. (1966). "Fungus-Growing Ants". Science. 153 (3736): 587–604. Bibcode:1966Sci...153..587W. doi:10.1126/science.153.3736.587. PMID 17757227.
- Brady, Seán G.; Schultz, Ted R. (2008-04-08). "Major evolutionary transitions in ant agriculture". Proceedings of the National Academy of Sciences. 105 (14): 5435–5440. Bibcode:2008PNAS..105.5435S. doi:10.1073/pnas.0711024105. ISSN 1091-6490. PMC 2291119. PMID 18362345.
- Branstetter, Michael G.; Ješovnik, Ana; Sosa-Calvo, Jeffrey; Lloyd, Michael W.; Faircloth, Brant C.; Brady, Seán G.; Schultz, Ted R. (2017-04-12). "Dry habitats were crucibles of domestication in the evolution of agriculture in ants". Proceedings of the Royal Society B: Biological Sciences. 284 (1852): 20170095. doi:10.1098/rspb.2017.0095. ISSN 0962-8452. PMC 5394666. PMID 28404776.
- Gerardo, Nicole; Mueller, Ulrich G. (2002-11-26). "Fungus-farming insects: Multiple origins and diverse evolutionary histories". Proceedings of the National Academy of Sciences. 99 (24): 15247–15249. Bibcode:2002PNAS...9915247M. doi:10.1073/pnas.242594799. ISSN 1091-6490. PMC 137700. PMID 12438688.
- Kooij, P. W.; Aanen, D. K.; Schiøtt, M.; Boomsma, J. J. (November 2015). "Evolutionarily advanced ant farmers rear polyploid fungal crops". Journal of Evolutionary Biology. 28 (11): 1911–1924. doi:10.1111/jeb.12718. ISSN 1420-9101. PMC 5014177. PMID 26265100.
- Currie, Cameron R.; Poulsen, Michael; Mendenhall, John; Boomsma, Jacobus J.; Billen, Johan (2006-01-06). "Coevolved crypts and exocrine glands support mutualistic bacteria in fungus-growing ants". Science. 311 (5757): 81–83. Bibcode:2006Sci...311...81C. CiteSeerX 10.1.1.186.9613. doi:10.1126/science.1119744. ISSN 1095-9203. PMID 16400148.
- Schultz, Ted R.; Rehner, Stephen A.; Mueller, Ulrich G. (1998-09-25). "The Evolution of Agriculture in Ants". Science. 281 (5385): 2034–2038. Bibcode:1998Sci...281.2034M. doi:10.1126/science.281.5385.2034. ISSN 1095-9203. PMID 9748164.
- Boomsma, Jacobus J.; Zhang, Guojie; Schultz, Ted R.; Brady, Seán G.; Wcislo, William T.; Nash, David R.; Rabeling, Christian; Dikow, Rebecca B.; Deng, Yuan (2016-07-20). "Reciprocal genomic evolution in the ant–fungus agricultural symbiosis". Nature Communications. 7: 12233. Bibcode:2016NatCo...712233N. doi:10.1038/ncomms12233. ISSN 2041-1723. PMC 4961791. PMID 27436133.
- Shik, Jonathan Z.; Gomez, Ernesto B.; Kooij, Pepijn W.; Santos, Juan C.; Wcislo, William T.; Boomsma, Jacobus J. (September 6, 2016). "Nutrition mediates the expression of cultivar-farmer conflict in a fungus-growing ant". Proceedings of the National Academy of Sciences of the United States of America. 113 (36): 10121–10126. doi:10.1073/pnas.1606128113. ISSN 1091-6490. PMC 5018747. PMID 27551065.
- Villesen, Palle; Murakami, Takahiro; Schultz, Ted R.; Boomsma, Jacobus J. (2002-08-07). "Identifying the transition between single and multiple mating of queens in fungus-growing ants". Proceedings. Biological Sciences. 269 (1500): 1541–1548. doi:10.1098/rspb.2002.2044. ISSN 0962-8452. PMC 1691065. PMID 12184823.
- Licht, Henrik H. De Fine; Boomsma, Jacobus J. (2010). "Forage collection, substrate preparation, and diet composition in fungus-growing ants". Ecological Entomology. 35 (3): 259–269. doi:10.1111/j.1365-2311.2010.01193.x. ISSN 1365-2311.
- de Man, Tom J. B.; Stajich, Jason E.; Kubicek, Christian P.; Teiling, Clotilde; Chenthamara, Komal; Atanasova, Lea; Druzhinina, Irina S.; Levenkova, Natasha; Birnbaum, Stephanie S. L. (2016-03-29). "Small genome of the fungus Escovopsis weberi, a specialized disease agent of ant agriculture". Proceedings of the National Academy of Sciences of the United States of America. 113 (13): 3567–3572. Bibcode:2016PNAS..113.3567D. doi:10.1073/pnas.1518501113. ISSN 1091-6490. PMC 4822581. PMID 26976598.
- Gerardo, Nicole M; Jacobs, Sarah R; Currie, Cameron R; Mueller, Ulrich G (August 2006). "Ancient Host–Pathogen Associations Maintained by Specificity of Chemotaxis and Antibiosis". PLoS Biology. 4 (8): e235. doi:10.1371/journal.pbio.0040235. ISSN 1544-9173. PMC 1489191. PMID 16805647.
- Poulsen, Michael; Boomsma, Jacobus J.; Yek, Sze Huei (2012). "Towards a Better Understanding of the Evolution of Specialized Parasites of Fungus-Growing Ant Crops". Psyche: A Journal of Entomology. 2012: 1–10. doi:10.1155/2012/239392.
- Currie, C. R. (2001). "A community of ants, fungi, and bacteria: a multilateral approach to studying symbiosis" (PDF). Annual Review of Microbiology. 55: 357–380. doi:10.1146/annurev.micro.55.1.357. hdl:1808/835. ISSN 0066-4227. PMID 11544360.
- Malloch, David; Summerbell, Richard C.; Scott, James A.; Currie, Cameron R. (April 1999). "Fungus-growing ants use antibiotic-producing bacteria to control garden parasites". Nature. 398 (6729): 701–704. Bibcode:1999Natur.398..701C. doi:10.1038/19519. ISSN 1476-4687.
- Barke, Jörg; Seipke, Ryan F.; Grüschow, Sabine; Heavens, Darren; Drou, Nizar; Bibb, Mervyn J.; Goss, Rebecca JM; Yu, Douglas W.; Hutchings, Matthew I. (2010-08-26). "A mixed community of actinomycetes produce multiple antibiotics for the fungus farming ant Acromyrmex octospinosus". BMC Biology. 8 (1): 109. doi:10.1186/1741-7007-8-109. ISSN 1741-7007. PMC 2942817. PMID 20796277.
- Haeder, Susanne; Wirth, Rainer; Herz, Hubert; Spiteller, Dieter (2009-03-24). "Candicidin-producing Streptomyces support leaf-cutting ants to protect their fungus garden against the pathogenic fungus Escovopsis". Proceedings of the National Academy of Sciences of the United States of America. 106 (12): 4742–4746. Bibcode:2009PNAS..106.4742H. doi:10.1073/pnas.0812082106. ISSN 1091-6490. PMC 2660719. PMID 19270078.
- Mattoso, Thalles C.; Moreira, Denise D. O.; Samuels, Richard I. (2012-06-23). "Symbiotic bacteria on the cuticle of the leaf-cutting ant Acromyrmex subterraneus subterraneus protect workers from attack by entomopathogenic fungi". Biology Letters. 8 (3): 461–464. doi:10.1098/rsbl.2011.0963. ISSN 1744-957X. PMC 3367728. PMID 22130174.
- Hutchings, Matthew I.; Goss, Rebecca J. M.; Yu, Douglas W.; Hill, Lionel; Brearley, Charles; Barke, Jörg; Seipke, Ryan F. (2011-08-03). "A Single Streptomyces Symbiont Makes Multiple Antifungals to Support the Fungus Farming Ant Acromyrmex octospinosus". PLOS ONE. 6 (8): e22028. Bibcode:2011PLoSO...622028S. doi:10.1371/journal.pone.0022028. ISSN 1932-6203. PMC 3153929. PMID 21857911.
- Holmes, Neil A.; Innocent, Tabitha M.; Heine, Daniel; Bassam, Mahmoud Al; Worsley, Sarah F.; Trottmann, Felix; Patrick, Elaine H.; Yu, Douglas W.; Murrell, J. C. (2016-12-26). "Genome Analysis of Two Pseudonocardia Phylotypes Associated with Acromyrmex Leafcutter Ants Reveals Their Biosynthetic Potential". Frontiers in Microbiology. 7: 2073. doi:10.3389/fmicb.2016.02073. ISSN 1664-302X. PMC 5183585. PMID 28082956.
- Oh, Dong-Chan; Poulsen, Michael; Currie, Cameron R.; Clardy, Jon (July 2009). "Dentigerumycin: a bacterial mediator of an ant-fungus symbiosis". Nature Chemical Biology. 5 (6): 391–393. doi:10.1038/nchembio.159. ISSN 1552-4469. PMC 2748230. PMID 19330011.
- Seipke, Ryan F.; Grüschow, Sabine; Goss, Rebecca J. M.; Hutchings, Matthew I. (2012). Isolating antifungals from fungus-growing ant symbionts using a genome-guided chemistry approach. Methods in Enzymology. 517. pp. 47–70. doi:10.1016/B978-0-12-404634-4.00003-6. ISBN 9780124046344. ISSN 1557-7988. PMID 23084933.
- Autuori (1956). "La fondation des sociétés chez les fourmis champignonnistes du genre Atta (Hym. Formicidae)". L'Instinct dans le Comportement des Animaux et de l'homme. pp. 77–104.
- Saes, N. B.; Forti, L. C.; Pereira-da-Silva, V.; Fowler, H. G. (1986). "Population dynamics of leaf-cutting ants: A brief review". Cite journal requires
- Hölldobler, Bert; Wilson, Edward O. (1990). The Ants. Berlin Heidelberg: Springer-Verlag. ISBN 9783540520924.
- John R. B. Lighten; Roces, Flavio (February 1995). "Larger bites of leaf-cutting ants". Nature. 373 (6513): 392. doi:10.1038/373392a0. ISSN 1476-4687.
- Feener, Donald H.; Moss, Karen A. G. (1990). "Defense against Parasites by Hitchhikers in Leaf-Cutting Ants: A Quantitative Assessment". Behavioral Ecology and Sociobiology. 26 (1): 17–29. doi:10.1007/bf00174021. ISSN 0340-5443. JSTOR 4600370.
- Irenäus; Eibl‐Eibesfeldt, Eleonore (1967). "Das Parasitenabwehren der Minima-Arbeiterinnen der Blattschneider-Ameise (Atta cephalotes)". Zeitschrift für Tierpsychologie. 24 (3): 278–281. doi:10.1111/j.1439-0310.1967.tb00579.x. ISSN 1439-0310.
- Institute of Medicine (US) Forum on Microbial Threats (2012). The Social Biology of Microbial Communities: Workshop Summary. The National Academies Collection: Reports funded by National Institutes of Health. Washington (DC): National Academies Press (US). ISBN 9780309264327. PMID 24027805.
- Schultz, Ted R.; Meier, Rudolf (1995). "A phylogenetic analysis of the fungus-growing ants (Hymenoptera: Formicidae: Attini) based on morphological characters of the larvae". Systematic Entomology. 20 (4): 337–370. doi:10.1111/j.1365-3113.1995.tb00100.x. ISSN 1365-3113.
- Mueller, U. G.; Schultz, T. R.; Currie, C. R.; Adams, R. M.; Malloch, D. (June 2001). "The origin of the attine ant-fungus mutualism". The Quarterly Review of Biology. 76 (2): 169–197. doi:10.1086/393867. ISSN 0033-5770. PMID 11409051.
- Villesen, Palle; Mueller, Ulrich G.; Schultz, Ted R.; Adams, Rachelle M. M.; Bouck, Amy C. (October 2004). "Evolution of ant-cultivar specialization and cultivar switching in Apterostigma fungus-growing ants". Evolution; International Journal of Organic Evolution. 58 (10): 2252–2265. doi:10.1111/j.0014-3820.2004.tb01601.x. ISSN 0014-3820. PMID 15562688.
- Chapela, I. H.; Rehner, S. A.; Schultz, T. R.; Mueller, U. G. (1994-12-09). "Evolutionary history of the symbiosis between fungus-growing ants and their fungi". Science. 266 (5191): 1691–1694. doi:10.1126/science.266.5191.1691. ISSN 0036-8075. PMID 17775630.
- Schultz, Ted R.; Rehner, Stephen A.; Mueller, Ulrich G. (1998-09-25). "The Evolution of Agriculture in Ants". Science. 281 (5385): 2034–2038. doi:10.1126/science.281.5385.2034. ISSN 0036-8075. PMID 9748164.
- Bot, A. N.; Rehner, S. A.; Boomsma, J. J. (October 2001). "Partial incompatibility between ants and symbiotic fungi in two sympatric species of Acromyrmex leaf-cutting ants". Evolution; International Journal of Organic Evolution. 55 (10): 1980–1991. doi:10.1111/j.0014-3820.2001.tb01315.x. ISSN 0014-3820. PMID 11761059.
- Leal, I.R.; Oliveira, P.S. (2000-11-01). "Foraging ecology of attine ants in a Neotropical savanna: seasonal use of fungal substrate in the cerrado vegetation of Brazil". Insectes Sociaux. 47 (4): 376–382. doi:10.1007/PL00001734. ISSN 1420-9098.
- Howard, Jerome J. (1988). "Leafcutting and Diet Selection: Relative Influence of Leaf Chemistry and Physical Features". Ecology. 69 (1): 250–260. doi:10.2307/1943180. ISSN 0012-9658. JSTOR 1943180.
- Nichols‐Orians, Colin M.; Schultz, Jack C. (1990). "Interactions among leaf toughness, chemistry, and harvesting by attine ants". Ecological Entomology. 15 (3): 311–320. doi:10.1111/j.1365-2311.1990.tb00813.x. ISSN 1365-2311.
- Wirth, Rainer; Herz, Hubert; Ryel, Ronald J.; Beyschlag, Wolfram; Hölldobler, Bert (2003). Herbivory of Leaf-Cutting Ants: A Case Study on Atta colombica in the Tropical Rainforest of Panama. Ecological Studies. Berlin Heidelberg: Springer-Verlag. ISBN 9783540438960.
- Blum, Murray S.; Moser, John C. (1963-06-14). "Trail Marking Substance of the Texas Leaf-Cutting Ant: Source and Potency". Science. 140 (3572): 1228–31. doi:10.1126/science.140.3572.1228. ISSN 0036-8075. PMID 14014717.
- Jaffe, K.; Howse, P. E. (1979-08-01). "The mass recruitment system of the leaf cutting ant, Atta cephalotes (L.)". Animal Behaviour. 27: 930–939. doi:10.1016/0003-3472(79)90031-9. ISSN 0003-3472.
- Ruth, J. M.; Brownlee, R. G.; Moser, J. C.; Silverstein, R. M.; Tumlinson, J. H. (December 1971). "Identification of the Trail Pheromone of a Leaf-cutting Ant, Atta texana". Nature. 234 (5328): 348–349. doi:10.1038/234348b0. ISSN 1476-4687. PMID 4944485.
- Cross, John H.; Byler, Russell C.; Ravid, Uzi; Silverstein, Robert M.; Robinson, Stephen W.; Baker, Paul M.; De Oliveira, João Sabino; Jutsum, Alan R.; Cherrett, J. Malcolm (1979-03-01). "The major component of the trail pheromone of the leaf-cutting ant,Atta sexdens rubropilosa forel". Journal of Chemical Ecology. 5 (2): 187–203. doi:10.1007/BF00988234. ISSN 1573-1561.
- Riley, R. G.; Silverstein, R. M.; Carroll, B.; Carroll, R. (April 1974). "Methyl 4-methylpyrrole-2-carboxylate: a volatile trail pheromone from the leaf-cutting ant, tatta cephalotes". Journal of Insect Physiology. 20 (4): 651–654. doi:10.1016/0022-1910(74)90186-3. ISSN 0022-1910. PMID 4833350.
- al, Kleineidam CJ , et (2007). "Perceptual differences in trail-following leaf-cutting ants relate to body size. - PubMed - NCBI". Journal of Insect Physiology. 53 (12): 1233–41. doi:10.1016/j.jinsphys.2007.06.015. PMID 17716686.
- Hölldobler, Bert; Wilson, Edward O. (2009). The Superorganism: The Beauty, Elegance, and Strangeness of Insect Societies. W. W. Norton & Company. ISBN 9780393067040.
- Nichols-Orians, Colin M.; Schultz, Jack C. (1989). "Leaf Toughness Affects Leaf Harvesting by the Leaf Cutter Ant, Atta cephalotes (L.) (Hymenoptera: Formicidae)". Biotropica. 21 (1): 80–83. doi:10.2307/2388446. ISSN 0006-3606. JSTOR 2388446.
- van Breda, J. M.; Stradling, D. J. (1994-12-01). "Mechanisms affecting load size determination inAtta cephalotes L. (Hymenoptera, Formicidae)". Insectes Sociaux. 41 (4): 423–435. doi:10.1007/BF01240645. ISSN 1420-9098.
- Burd, Martin (1995-09-01). "Variable load size-ant size matching in leaf-cutting ants,Atta colombica (Hymenoptera: Formicidae)". Journal of Insect Behavior. 8 (5): 715–722. doi:10.1007/BF01997240. ISSN 1572-8889.
- Burd, Martin (1996-10-01). "Server System and Queuing Models of Leaf Harvesting by Leaf-Cutting Ants". The American Naturalist. 148 (4): 613–629. doi:10.1086/285943. ISSN 0003-0147.
- Burd, Martin (1996-10-01). "Foraging Performance by Atta colombica, a Leaf-Cutting Ant". The American Naturalist. 148 (4): 597–612. doi:10.1086/285942. ISSN 0003-0147.
- Markl, H. (1965-09-17). "Stridulation in Leaf-Cutting Ants". Science. 149 (3690): 1392–1393. doi:10.1126/science.149.3690.1392. ISSN 0036-8075. PMID 17741924.
- Tautz, J.; Roces, F.; Hölldobler, B. (1995-01-06). "Use of a sound-based vibratome by leaf-cutting ants". Science. 267 (5194): 84–87. doi:10.1126/science.267.5194.84. ISSN 0036-8075. PMID 17840064.
- Röschard, Jacqueline; Roces, Flavio (2002-04-01). "The effect of load length, width and mass on transport rate in the grass-cutting ant Atta vollenweideri". Oecologia. 131 (2): 319–324. doi:10.1007/s00442-002-0882-z. ISSN 1432-1939. PMID 28547700.
- Röschard, J.; Roces, F. (2003-08-01). "Cutters, carriers and transport chains: Distance-dependent foraging strategies in the grass-cutting ant Atta vollenweideri". Insectes Sociaux. 50 (3): 237–244. doi:10.1007/s00040-003-0663-7. ISSN 1420-9098.
- Stephen P., Hubbell; Johnson, Leslie K.; Stanislav, Eileen; Wilson, Berry; Fowler, Harry (1980). "Foraging by Bucket-Brigade in Leaf-Cutter Ants". Biotropica. Association for Tropical Biology and Conservation. 12 (3): 210. doi:10.2307/2387973. JSTOR 2387973. Retrieved 2019-06-06.
- Fowler, Harold G.; Robinson, S. W. (1979). "Foraging by Atta sexdens (Formicidae: Attini): seasonal patterns, caste and efficiency". Ecological Entomology. 4 (3): 239–247. doi:10.1111/j.1365-2311.1979.tb00581.x. ISSN 1365-2311.
- Anderson, Carl; Ratnieks, Francis L. W. (November 1999). "Task Partitioning in Insect Societies. I. Effect of Colony Size on Queueing Delay and Colony Ergonomic Efficiency" (PDF). The American Naturalist. 154 (5): 521–535. doi:10.1086/303255. ISSN 1537-5323. PMID 10561125.
- Hart, Adam G.; Ratnieks, Francis L. W. (2001-08-01). "Leaf caching in the leafcutting ant Atta colombica: organizational shift, task partitioning and making the best of a bad job". Animal Behaviour. 62 (2): 227–234. doi:10.1006/anbe.2001.1743. ISSN 0003-3472.
- Anderson, C.; Boomsma, J.J.; Bartholdi, III, J.J. (2002-05-01). "Task partitioning in insect societies: bucket brigades". Insectes Sociaux. 49 (2): 171–180. doi:10.1007/s00040-002-8298-7. ISSN 1420-9098.
- Roces, Flavio (June 2002). "Individual complexity and self-organization in foraging by leaf-cutting ants". The Biological Bulletin. 202 (3): 306–313. doi:10.2307/1543483. ISSN 0006-3185. JSTOR 1543483. PMID 12087004.
- Bass, M.; Cherrett, J. M. (1995). "Fungal hyphae as a source of nutrients for the leaf-cutting ant Atta sexdens". Physiological Entomology. 20 (1): 1–6. doi:10.1111/j.1365-3032.1995.tb00793.x. ISSN 1365-3032.
- Meurant, Gerard (2012-12-02). "The mutualism between leaf-cutting ants and their fungus". Insect-Fungus Interactions. Academic Press. ISBN 9780080984537.
- Gomes De Siqueira, Célia; Bacci, Maurício; Pagnocca, Fernando Carlos; Bueno, Odair Correa; Hebling, Maria José Aparecida (December 1998). "Metabolism of Plant Polysaccharides by Leucoagaricus gongylophorus, the Symbiotic Fungus of the Leaf-Cutting Ant Atta sexdens L." Applied and Environmental Microbiology. 64 (12): 4820–4822. ISSN 0099-2240. PMC 90928. PMID 9835568.
- Abril, Adriana B.; Bucher, Enrique H. (2002). "Evidence that the fungus cultured by leaf-cutting ants does not metabolize cellulose". Ecology Letters. 5 (3): 325–328. doi:10.1046/j.1461-0248.2002.00327.x. ISSN 1461-0248.
- D'Ettorre, P.; Mora, P.; Dibangou, V.; Rouland, C.; Errard, C. (February 2002). "The role of the symbiotic fungus in the digestive metabolism of two species of fungus-growing ants". Journal of Comparative Physiology B. 172 (2): 169–176. doi:10.1007/s00360-001-0241-0. ISSN 0174-1578. PMID 11916111.
- Quinlan, R. J.; Cherrett, J. M. (1979). "The role of fungus in the diet of the leaf-cutting ant Atta cephalotes (L.)". Ecological Entomology. 4 (2): 151–160. doi:10.1111/j.1365-2311.1979.tb00570.x. ISSN 1365-2311.
- Boomsma, Jacobus J.; Schiøtt, Morten; Sørensen, Søren J.; Hansen, Lars H.; Zhukova, Mariya; Sapountzis, Panagiotis (2015-08-15). "Acromyrmex Leaf-Cutting Ants Have Simple Gut Microbiota with Nitrogen-Fixing Potential". Appl. Environ. Microbiol. 81 (16): 5527–5537. doi:10.1128/AEM.00961-15. ISSN 1098-5336. PMC 4510174. PMID 26048932.
- Anderson, Kirk E.; Russell, Jacob A.; Moreau, Corrie S.; Kautz, Stefanie; Sullam, Karen E.; Hu, Yi; Basinger, Ursula; Mott, Brendon M.; Buck, Norman (May 2012). "Highly similar microbial communities are shared among related and trophically similar ant species". Molecular Ecology. 21 (9): 2282–2296. doi:10.1111/j.1365-294X.2011.05464.x. ISSN 1365-294X. PMID 22276952.
- Bae, Jin-Woo; Lee, Won-Jae; Kim, Sung-Hee; Shin, Na-Ri; Kim, Joon-Yong; Choi, Jung-Hye; Kim, Yun-Ji; Nam, Young-Do; Yoon, Changmann (2014-09-01). "Insect Gut Bacterial Diversity Determined by Environmental Habitat, Diet, Developmental Stage, and Phylogeny of Host". Appl. Environ. Microbiol. 80 (17): 5254–5264. doi:10.1128/AEM.01226-14. ISSN 1098-5336. PMC 4136111. PMID 24928884.
- Colman, D. R.; Toolson, E. C.; Takacs-Vesbach, C. D. (October 2012). "Do diet and taxonomy influence insect gut bacterial communities?". Molecular Ecology. 21 (20): 5124–5137. doi:10.1111/j.1365-294X.2012.05752.x. ISSN 1365-294X. PMID 22978555.
- Sapountzis, Panagiotis; Nash, David R.; Schiøtt, Morten; Boomsma, Jacobus J. (2018). "The evolution of abdominal microbiomes in fungus-growing ants". Molecular Ecology. 0 (4): 879–899. doi:10.1111/mec.14931. ISSN 1365-294X. PMC 6446810. PMID 30411820.
- Boomsma, Jacobus J.; Rosendahl, Søren; Guldberg-Frøslev, Tobias; Rouland-Lefèvre, Corinne; Eggleton, Paul; Aanen, Duur K. (2002-11-12). "The evolution of fungus-growing termites and their mutualistic fungal symbionts". Proceedings of the National Academy of Sciences. 99 (23): 14887–14892. Bibcode:2002PNAS...9914887A. doi:10.1073/pnas.222313099. ISSN 1091-6490. PMC 137514. PMID 12386341.
- Hölldobler, Bert and Wilson, EO. (2009). The Superorganism: The Beauty, Elegance, and Strangeness of Insect Societies. W. W. Norton & Company. ISBN 9780393067040
- Media related to Attini at Wikimedia Commons