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Overview

Brief Summary

Overview

Atta cephalotes is the most widely distributed species of leaf cutting ant. It occurs  from southernmost Mexico to Ecuador and Brazil, and from the Lesser Antilles as far north as Bardbados (Holldobler & Wilson, 1990). An apparently disjunct population occurs in the Atlantic Coastal Forests of Brazil (Solomon, 2007). The species is widely regarded as a significant pest on crops including citrus, coffee, cocoa and ornamentals (Cherrett, 1986; Cherrett & Peregrine, 1976). The ecological success and ubiquity of A. cephalotes is owed in large part to its broad niche tolerance with respect to both nesting habitats and diet breadth (Solomon, 2007). The species has evolved to specialize in forest gaps, and consequently thrives in disturbed habitats such as farms and plantations (Cherrett & Peregrine, 1976). Niche modeling suggests that the current range of A. cephalotes is limited by dispersal constraints rather than suitable habitat (Solomon, 2007), and could cause significant ecological and agricultural devastation should it become introduced elsewhere (Hlldobler & Wilson, 1990). Atta cephalotes is highly variable, both among castes of the same colony and across populations. Workers vary enormously in shape, color, sculpture and pubescence (Borgmeier, 1959). 

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Comprehensive Description

Taxonomic History

Taxonomic history

Olivier, 1792: 500 (q.m.); Wheeler, 1949 PDF: 677 (l.).
Combination in Atta: Fabricius, 1804: 421; in Oecodoma: Latreille, 1818a PDF: 224.
Senior synonym of Atta migratoria: Retzius, 1783 PDF: 76; of Atta grossa: Latreille, 1802b: 224; of Atta fervens: Smith, 1858a PDF: 180; of Atta visitatrix: Emery, 1892c PDF: 162; of Atta integrior, Atta isthmicola, Atta lutea, Atta oaxaquensis, Atta opaca, Atta polita: Borgmeier, 1959b: 340.
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Biology

In Costa Rica (Jack Longino)
This is the most common species of Atta in Costa Rica and it is found throughout the country at mid to low elevations. It occurs in wet or dry forest habitats and prefers clearings and forest edges. In the lowlands it is most common in agricultural areas and young second growth forest but can also occur in mature closed-canopy forest. At higher elevations it is increasingly restricted to open, highly insolated areas. For example, in the Monteverde area large nests of A. cephalotes occur in pastures around the community, at about 1400m elevation, but not in adjacent forest. Residents observe that nests are becoming more abundant and occurring at ever higher elevations, perhaps related to increased development and climate change.

Atta cephalotes has subterranean refuse dumps where spent fungus substrate is deposited. This is in contrast to A. colombica, which has conspicuous refuse dumps on the surface.

Nuptial flights occur during predawn hours and are synchronized across colonies in a population. Massive emergences occur sporadically during the year. On mornings following nuptial flights trails and clearings are sprinkled with the large queens and males, most of them being eaten by ants and other predators. 

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Atta cephalotes is the most widely distributed species of leaf cutting ant. It occurs from southernmost Mexico to Ecuador and Brazil, and from the Lesser Antilles as far north as Barbados (Hölldobler & Wilson, 1990). An apparently disjunct population occurs in the Atlantic Coastal Forests of Brazil (Solomon, 2007). The species is widely regarded as a significant pest on crops including citrus, coffee, cocoa and ornamentals (Cherrett, 1986; Cherrett & Peregrine, 1976). The ecological success and ubiquity of A. cephalotes is owed in large part to its broad niche tolerance with respect to both nesting habitats and diet breadth (Solomon, 2007). The species has evolved to specialize in forest gaps, and consequently thrives in disturbed habitats such as farms and plantations (Cherrett & Peregrine, 1976). Niche modeling suggests that the current range of A. cephalotes is limited by dispersal constraints rather than suitable habitat (Solomon, 2007), and could cause significant ecological and agricultural devastation should it become introduced elsewhere (Hölldobler & Wilson, 1990; Mikheyev, 2008). A pest risk assessment of unprocessed Eucalyptus concluded the introduction of South American Atta species into the United States was a low probability (Kliejunas et al., 2001).

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Distribution

Native range. Southernmost Mexico to Ecuador and Brazil, and from the Lesser Antilles as far north as Barbados.
Introduced range. No records of introductions.

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Southernmost Mexico to Ecuador and Brazil, and from the Lesser Antilles as far north as Bardbados. An apparently disjunct population occurs in the Atlantic Coastal Forests of Brazil.

Occasionally intercepted at US Ports of Entry.

 

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Physical Description

Diagnostic Description

Taxonomic Treatment

Borgmeier, T., 1939:
  Como localidade do typo desta espécie deve ser considerada a Guyana Hollandeza (Surinam). Linneu baseou a sua diagnose (1758) sobre material colleccionado por Rolander, sobre o qual diz H. Baillon (1891, Dictionnaire de Botanique, vol. 3, Paris, p. 742): «Voyageur danois à la Guyane. Hornemann a fait connaitre (1812) le journal de son Voyage à Surinam».
  Considero como forma typica abundante material de operarios, 1 macho e 1 fêmea de Lelydorp, localidade que fica a uma distancia de 25 km de Paramaribo (Stahei e Geijskes leg. Abril 1938). Também possuo material colleccionado porBünzli em Paramaribo, e por N. A. Weber na Guyana Hollandeza e em Trinidad.
  Segundo Santschi (1929, p. 92) a cabeça do operá- rio maior apresenta-se muito dividida pelo sulco occipital, o que não se verifica no material de Surinam. A espécie parece variar, bastante e precisava ser revista.
  Sobre a nidificação desta espécie , Stahel e Geijskes publicaram ha pouco (1939, Rev. Ent. X, fasc. 1) importante trabalho.
 

Forel, A., 1905:
 - Saint-Esteban.
 

Forel, A., 1908:
 Costa-Rica, San Jose (Biolley, Alfaro). Ile de Coco (Alfaro), evidemment importee.
 
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Identification

Atta cephalotes vs. Atta sexdens
Atta cephalotes can be distinguished by the wooly hairs on the heads of major workers and the shinier integument caused by the lack of hexagonal microsculpture, and the lack of small teeth or dents on the head anterior to the posterolateral spines. The pronotal spines of A. cephalotes tend to be more curved than those of its congeners, but this is a highly variable character.

Atta cephalotes vs. Atta native to the United States
The majors of A. cephalotes are also distinguished from its two congeners native to the United States (A. mexicana and A. texana) by the presence of wooly hairs on the cephalic dorsum. The petiolar spiracles are visible from above in the two native species, but cannot be seen from above in A. cephalotes (Borgmeier, 1959).

 

Atta cephalotes vs. Acromyrmex
Atta cephalotes is distinguished from its close relatives in the genus Acromyrmex by the presence of only three long spines on the mesosomal dorsum, and the absence of tubercles on the mesosomal and gastric dorsum.

Diagnosis among introduced and commonly intercepted species
Antenna 11-segmented. Antennal club indistinct. Antennal insertions at least partly covered by frontal lobes. Antennal scrobe lacking. Antennal scapes not conspicuously short; easily extended beyond eye level. Posterolateral corners of head spinose; lacking pair of small teeth anterior to posterolateral spines. Head smooth and shiny, not covered by hexagonal microsculpture. Head of soldier appearing wooly, covered in abundant long fine overlapping hairs. Mandibles triangular. Waist 2-segmented. Petiole with a distinct and upright node. Postpetiole attached to lower surface of gaster. Dorsum of promesonotum with 2 pairs of spines or teeth. Propodeum armed with spines. First gastral tergite lacking numerous tubercles. Yellow to reddish brown.

Redescription translated from Borgmeier, 1959b: 340  
The most complete description of the soldiers of this important kind of gave De Geer (1773). On the reproductives did Mayr (1865) and Emery (1890, 1913), important information, and the drawings of male genitalia in Emery (1913 8) are absolutely accurate. Among the new authors are concerned Goncalves (1942) and Borg Meier (1950) with the taxonomy of this kind is an actual description of all searches in the box recent literature in vain.

 

Type locality: Paramaribo, Suriname.

 

Workers. - Maxima (Paramaribo): 14 mm length with his head bowed. Head width 6, 1 mm. Occiput gently bulged in the middle. Occipital zaehne short, often reduced to tubercles. Occipital lobes thickened and rounded, the front on each side with long hair of woolly Bueschel. Forehead with 3 ocelli, the posterior small, sometimes absent. Eyes strongly convex. Thorax narrower than the head of many (width front 2.7 mm). Pronotum on each side at the lower edge with a curved spine. Mesonotum with 2 pairs of spines, the anterior with a broad base, often rounded up and the back short and sharp. Epinotaldornen straight, directed backwards and upwards. Petiole and postpetiole side recently with one tooth. Gaster: width 3.5 mm. Legs slender; hind tibiae 5 mm. Shiny little head forward. Occiput polished, naked, matt body. Face, thorax, pedicel and gaster with long woolly hair, sparse hair, protruding, hairy legs sticking out. Rusty brown color. - Media (Paramaribo): length 7 mm. Head width 2, 6 mm. Occiput bulged wide. Kapfflach in profile. Occipital spines slightly shorter than the eye. Front Mesonotal spines thin and long, curved forward, the back short and pointed. Epinotal spines too thin, slightly shorter than the mesonotal spines. Forehead with shiny ribbon on the center. Thorax shiny little. Gaster dull. Kapf, thorax and gaster with long hair and short protruding fitting Pubescenz. - Minima (Paramaribo) L. about 2 mm. Head without mandibles as long as breil Occipital spines very short or obsolete. Mesonotal thorns short, often missing the rear. Epinotal spines longer. Color yellow. Tegument something shiny. Prominent scattered hairs.

 

Females (Paramaribo). - Length 22 mm. Head width 5, 25ri1m. Occipital edge almost straight. Lies behind corners without head gear, at most with weak tubercles. Sides of the head behind the eyes convex, strongly convergent forward. Ocelli large. Eyes convex. Mandibles glossy, striped, teeth with blunt Apikalzahn and 8-9. Epinotum reinforced. Thorax length 10 mm. Gaster Br 8.5 mm. Tibia III 5.4 mm. Wing about 28 mm at the outer edge strongly browned. Matt. Head, thorax and gaster with fine golden pubescence fitting. Legs fitting coat or diagonally. Yellow rust stain.

 

Males (Paramaribo). - Length about 18 mm. Head (Borgm. 1950, Figure 47): width 2, 8 mm. Occipital zaehne obsolete. Front plate with teeth. Eyes strongly convex. Mandibles narrow, inner margin concave, irregularly toothed, pointed Apikalzahn. Scape too thin, 2, 6 mm, on the apical half slightly thickened. Wing about 25 mm, outer margin strongly browned. Genitalia (Fig. 10): Not Squamula dorsally fused in the middle, longer than wide; stipes articulated to the Squamula, not fused with it, abgestuzt apical hair long. Voisellen stabfoermig, in profile gently curved, apically obliquely truncated. Sagittae built complex, each side with an upright sharpened keel, which extends to the base, where it terminates in a spike, the keels slightly diverging apically, basally more strongly, laterally outside of it there is a curved hook. is occupied at its end and the back edge with tiny sharp teeth, on the ventral side of the Sagitta's in the middle line, two closely spaced keels, which are in profile convex and serrated saw-are (the teeth are kopfwaerts directed and curved), which two keels separated by a furrow that widens and forms the apical foramen, located above the foramen on each side a broad, flat, triangular teeth. Subgenital bulged apically. Head and thorax woolly, 1st Gaster short segment pubescent. Color yellow rust or rust-brown.

 

Geographical distribution. - Mexico (Veracruz), Central America, Ecuador, Columbia, Peru, Bolivia (Chapare up), Venezuela, Guyana, Brazil (Amazonas, Para, Rondonia, Maranhao, Pernambuco, Baia (up I1heus, Belmonte and Mucuri).

 

Discussion. - Atta colombica cephalotes is very close, as evidenced by the male genitalia and the form det average worker, but colombica differs in the following important points: 1) the head of the soldiers is totally different, much narrower, incised on occipital deep edge, 2) the soldier is practically naked, pubescent only very fine fitting, and 3) in the middle are the workers Occipital spines always longer than the eye (at shorter cephalotes), 4) the male genitalia is squamate the top as long as wide, 5) are the Sagittalhaken less curved and serrated edge on the back clearly. The females of both species are hardly distinguishable (by colombica are slightly smaller). The dark brown female from Colombia, Bolivia and Costa Rica can easily be confused sexdens, etc., but are identified by the absence of Occipital zaehne and the darkened edge of the wing.

 

Variation. - Apart from the strong polymorphism workers vary enormously in shape and painting, sculpture and pubescence. Copies of Columbia, Bolivia, Peru and Costa Rica are dark brown and the soldiers have dull occipital (var opaca), while the females are covered with hair sticking out. Little Workers of Trinidad (3-4 mm) are bright yellow and very shiny all over her body (formerly referred to by me as lutea). The shape of the stipes varies (cf Borgm. In 1950, FIG 39, 43-46), also the head of conspecific males (ibid. Fig 47-51). The shape of the Sagitta is relatively constant (Fig. ibid. 35-37).

 

Synonymy. - The large variability of cephaloles it is not surprising that a number of synonyms exist in the literature. In the old synonyms that are far from clear, I do not discuss (d Emery, 1923).

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Diagnosis of worker among Antkey species. Antenna 11-segmented. Antennal club indistinct. Antennal insertions at least partly covered by frontal lobes. Antennal scrobe lacking. Antennal scapes not conspicuously short; easily extended beyond eye level. Posterolateral corners of head spinose; lacking pair of small teeth anterior to posterolateral spines. Head smooth and shiny, not covered by hexagonal microsculpture. Head of soldier appearing wooly, covered in abundant long fine overlapping hairs. Mandibles triangular. Waist 2-segmented. Petiole with a distinct and upright node. Postpetiole attached to lower surface of gaster. Dorsum of promesonotum with 2 pairs of spines or teeth. Propodeum armed with spines. First gastral tergite lacking numerous tubercles. Yellow to reddish brown.


Atta cephalotes is highly variable, both among castes of the same colony and across populations. Workers vary enormously in shape, color, sculpture and pubescence (Borgmeier, 1959). The species is most readily confused with A. sexdens, but can be distinguished by the wooly hairs on the heads of major workers and the shinier integument caused by the lack of hexagonal microsculpture, and the lack of small teeth or dents on the head anterior to the posterolateral spines. The pronotal spines of A. cephalotes tend to be more curved than those of its congeners, but this is a highly variable character.


The majors of A. cephalotes are also distinguished from its two congeners native to the United States (A. mexicana and A. texana) by the presence of wooly hairs on the cephalic dorsum. The petiolar spiracles are visible from above in the two native species, but cannot be seen from above in A. cephalotes (Borgmeier, 1959). Atta cephalotes is distinguished from its close relatives in the genus Acromyrmex by the presence of only three long spines on the mesosomal dorsum, and the absence of tubercles on the mesosomal and gastric dorsum. 

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Como localidade do typo desta espécie deve ser considerada a Guyana Hollandeza (Surinam). Linneu baseou a sua diagnose (1758) sobre material colleccionado por Rolander, sobre o qual diz H. Baillon (1891, Dictionnaire de Botanique, vol. 3, Paris, p. 742): «Voyageur danois à la Guyane. Hornemann a fait connaitre (1812) le journal de son Voyage à Surinam».

 

Considero como forma typica abundante material de operarios, 1 macho e 1 fêmea de Lelydorp, localidade que fica a uma distancia de 25 km de Paramaribo (Stahei e Geijskes leg. Abril 1938). Também possuo material colleccionado porBünzli em Paramaribo, e por N. A. Weber na Guyana Hollandeza e em Trinidad.

 

Segundo Santschi (1929, p. 92) a cabeça do operá- rio maior apresenta-se muito dividida pelo sulco occipital, o que não se verifica no material de Surinam. A espécie parece variar, bastante e precisava ser revista.

 

Sobre a nidificação desta espécie , Stahel e Geijskes publicaram ha pouco (1939, Rev. Ent. X, fasc. 1) importante trabalho.

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Borgmeier, T.

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- Saint-Esteban.

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Forel, A.

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Costa-Rica, San Jose (Biolley, Alfaro). Ile de Coco (Alfaro), evidemment importee.

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Forel, A.

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Look Alikes

Atta sexdens, Acromyrmex octospinosus​.

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Evolution and Systematics

Functional Adaptations

Functional adaptation

Smaller loads more efficient: leaf-cutting ant
 

Leaf-cutting ants optimize efficiency by matching load size to the outbound flow from the nest and extrapolating future flow of returning, laden ants.

   
  "When carrying resources from a collecting point to the nest, one would assume that animals would attempt to carry as much as possible to maximize their foraging efforts. However, among social insects that is not always the best strategy. Foragers carrying large loads might overwhelm the individuals processing the resources in the nest, causing a bottleneck. Additionally, a heavy load slows the carrier down. That may not be a significant cost for a solitary forager, but it may reduce the gains for a colony as a whole. Carrying loads well below maximum carrying performance actually reduces the burden on the resource processors and speeds up the forager, allowing for more foraging trips per unit time. But there is yet another factor whereby lighter loads can increase colony efficiency. Walking foragers, such as ants, often forage along well-defined trails and, depending on the number of foragers, this can potentially cause traffic problems. In that context, when one ant is slowed down by a heavy load it also slows down those ants following behind, regardless of load size. This phenomenon, called the 'truckdriver' effect – where a heavily laden truck slows down normally faster cars – describes another situation where carrying too much negatively impacts whole colony foraging efficiency." (Klok 2011:vi)

"At low traffic flow ants can cut and carry larger leaf fragments without the concern of slowing other ants, but when traffic flow increases they refrain from cutting larger fragments. It appears that individual ant foragers leaving the nest can estimate the outbound traffic flow and use this information to estimate the future flow of returning laden ants, thereby modulating the sizes of leaf cuttings made in order to avoid delays in overall colony foraging rates. This study shows remarkable flexibility in foraging behaviour and supports the idea that leaf-cutting ants make choices not only as individuals but also collectively." (Klok 2011:vi)
  Learn more about this functional adaptation.
  • Farji-Brener AG; Chinchilla FA; Rifkin S; Sanchez Cuervo AM; Triana E; Quiroga V; Giraldo P. 2011. The ‘truck-driver’ effect in leaf-cutting ants: how individual load influences the walking speed of nest-mates. Physiological Entomology. 36(2): 128-134.
  • Klok CJ. Heavy loads slow truck drivers and ants. The Journal of Experimental Biology. 214(doi: 10.1242/​jeb.049825): vi.
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Molecular Biology and Genetics

Molecular Biology

Barcode data: Atta cephalotes

The following is a representative barcode sequence, the centroid of all available sequences for this species.


No available public DNA sequences.

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Statistics of barcoding coverage: Atta cephalotes

Barcode of Life Data Systems (BOLDS) Stats
Public Records: 1
Specimens with Barcodes: 1
Species With Barcodes: 1
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Relevance to Humans and Ecosystems

Benefits

The vibrating mandible of the leaf-cutter ant has been studied in detail to determine how the ants cut the leaves so efficiently (Holldobler & Wilson, 2009). The discovery of how the ant vibrates and uses its mandibles led to the human invention of new microsurgical instruments which operate similarly (Boudreau, 2010). It has also been hypothesized that leaf-cutter ants in general may be useful in controlling certain pest insect populations for some farmers in Mesoamerica and the Caribbean. However this has not yet been specifically tested on A. cephalotes (Varon, 2007).

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Risks

The species is widely regarded as a significant pest on crops including citrus, coffee, cocoa and ornamentals.

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Many humans consider Atta cephalotes to be a pest insect species due to the substantial damage they can do to native foliage and farmer's fields in order to supply their fungal crop with enough leaf material to continue growth. As a result human farmers are forced to either plant shade trees to provide A. cephalotes with another food source, or attempt to deter them by baiting another species of ants (Azteca sp.) to take up residence in the tree. This however, requires a colony of Azteca sp. in every tree, which may lead to uninteded problems for farmers (Jutsom, et al, 1981; Varon, 2007).

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Wikipedia

Atta cephalotes

Atta cephalotes is one of 41 species of leafcutter ant. This species is part of the Attini tribe (the fungus-growing ants). A single colony of ants can contain up to 5 million members, and each colony has one queen that can live more than 15 years. The colony comprises different castes, known as ‘task partitioning’, and each caste has a different job to do.

Biology and behaviour[edit]

Ants are split into different castes. Soldiers are the largest of the ants and they often stand guard at the entrance of the nest, or will even go on scouting missions to watch for predators. Workers, who are all female, are divided into two types, media and minima, that both have big jaws and sharp teeth. Media workers are the larger of the two, and they cut the leaves and bring them back to the nest. These ants are able to navigate to the source of leaves by following the pheromone scent of the other ants ahead, a process known as ‘tandem running’. Once the media workers have deposited the material into the nest, slightly smaller workers called ‘minima workers’ cut up the leaves into small pieces and then feed it to a fungus they cultivate. They also cover the leaf in antibacterial saliva which stops other types of fungus growing.

A special caste of workers manage the colony's rubbish dump. These ants are excluded from the rest of the colony. If any wander outside the dump, the other ants will kill or force them back. Rubbish workers are often contaminated with disease and toxins, and live only half as long as their peers.[2]

The smaller ‘minima’ workers feed the entire colony of ants. These minima workers also act as guards and follow the media workers or hitch a ride on the leaf they’re carrying to the source of the leaves and attack small parasitoids called phroid flies.

The queen is larger than the other females and is winged. Her role is to give birth to workers.

The females can be workers or soldiers, and cannot produce workers but instead produce males only.

Males are also winged, and their only role is to inseminate the virgin queen.

The leaves that the worker ants cut are not for their consumption, as it is toxic to ants. Instead, the minima ants feeding it to a fungus that they farm.

Habitat[edit]

Across the rainforest floor they occupy an area typically an area of approximately 20 feet. They live in nests that can be as deep as 7 metres that they have carefully positioned so that a breeze can rid the nest of the dangerous levels of CO2 given off by the fungus they farm and eat.

Evolution and history[edit]

Leaf-cutter ants are an extremely specialised species. It has evolved over 25 million years to have a symbiotic relationship with the fungus it farms.

Mating[edit]

Mating in Atta cephalotes requires flight, which is why the queen and the males are winged. In preparation, the queen will store some fungus in a pouch in her mouth to begin cultivating in her future colony. The process starts with the queen flying up off the ground. The male will then join her and inseminate her, at which point he is no longer needed and dies. The queen will reach the ground and very quickly lose her wings, and will then look for a suitable place to begin a new colony. Once an adequate place is found, she will start digging into the earth and release a bit of the fungus she had previously stored to begin the process of a new underground system.

References[edit]

www.arkive.org/leaf-cutter-ant San Francisco State University Department of Geography www.thewildones.org

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