10.1007/978-1-4020-6359-6

From: Subject: 10.1007/978-1-4020-6359-6_1983 Date: Tue, 7 Oct 2008 05:39:37 -0400 MIME-Version: 1.0 Content-Type: multipart/related; type="text/html"; boundary="----=_NextPart_000_0000_01C9283F.15CDA860" X-MimeOLE: Produced By Microsoft MimeOLE V6.00.2900.5579 This is a multi-part message in MIME format. ------=_NextPart_000_0000_01C9283F.15CDA860 Content-Type: text/html; charset="iso-8859-1" Content-Transfer-Encoding: quoted-printable Content-Location: http://springerlink.com/content/l844231n882375g7/fulltext.html 10.1007/978-1-4020-6359-6_1983

Encyclopedia of Entomology

Springer Science+Business Media B.V. 2008

10.1007/978-1-4020-6359-6_1983

John L. Capinera

Leaf-Cutting Ants (Formicidae: = Myrmicinae:=20 Attini)

Klaus Jaffe²

(2)	Universidad Sim=F3n Bol=EDvar, = Caracas,=20 Venezuela

Without = Abstract

All ant species in the tribe Attini (Formicidae: = Myrmicinae)=20 cultivate a symbiotic fungus (Basidomycete: Lepiotacae) in order to feed = their=20 brood. The most conspicuous members of this tribe are undoubtedly the=20 leaf-cutter ants from the genera Atta, Acromyrmex and=20 Trachymyrmex. These ants cut leaves, which they prepare by = removing=20 surface waxes that normally harbor fungicides, before feeding their = symbiotic=20 fungus with it. The fungus helps to detoxify the leaves by degrading the = insecticides which are normally found inside the leaves. The larvae feed = basically on the fungus, whereas workers feed also on sugary plant sap = flowing=20 from the leaves while they are being cut. These ants build subterranean = nests in=20 which they grow their fungus which also needs a symbiotic bacterium in = order to=20 prosper. The workers regulate nest conditions so as to maintain = humidity,=20 temperature and carbon dioxide concentrations between narrow ranges. = Workers=20 avoid contaminations from other fungi and bacteria thanks to antibiotic=20 secretions from their metapleural glands and the action of the symbiotic = bacterium. The filamentous bacterium (Actinomycete) of the genus=20 Streptomyces produces antibiotics specifically targeted to = suppress the=20 growth of the specialized garden-parasite Escovopsis. The = symbiotic=20 fungus is known only to occur in association with Attini ants. The lower = Attini,=20 which are not considered to be leaf-cutting ants, feed their fungus = flowers,=20 fruit pulp, dead insects, and animal excrements.

The Colony

The leaf-cutting ant colony is normally monogynic, that = is, it=20 contains a single queen. A few cases of polygynic colonies (2=966 = queens) have=20 been reported, but this certainly is not the norm for these ants. The = colony is=20 formed by different individuals and each type of individuals is called a = caste.=20 The leaf-cutting ant colony contains normally only female individuals of = at=20 least two different castes, the queen and the workers. Just before the = beginning=20 of the tropical rain season, the colony produces winged individuals, = fertile=20 males and females which wait inside the nest for their nuptial flight = after the=20 first rains. All these types of individuals or castes are found in their = different life stages such as eggs, larvae, nymphs and adults in a = single=20 colony. The architecture of the subterranean nest is characteristic for = each=20 species. Workers also have behavioral castes. Young workers are more = often=20 engaged in caring for the fungus whereas older workers forage or engage = in=20 dumping rubbish and dead ants. The different components of a typical = colony are:=20

The Queen

Most mature leaf-cutting ant colonies have a single = fertile queen,=20 i.e., they are monogynic. The queen is the fertile female individual,=20 inseminated during the nuptial flight by various males, in charge of = laying=20 eggs. The eggs might have been fertilized by a male spermatozoon or not. = In the=20 first case, they will produce females; in the second, they produce = males.=20 Depending on the amount of food the larvae of females receive (and may = be on=20 hormones provided by the colony), they will become either virgin queens, = ready=20 to start a nuptial flight and eventually a new colony, or sterile worker = ants.=20

The Workers

Workers are sterile female ants. A single Atta = colony has=20 workers of different sizes =96 they are polymorphic =96 whereas those = from=20 Acromyrmex and Trachymyrmex colonies are monomorphic, = i.e., they=20 are all approximately the same size. Polymorphism among workers from = Atta=20 species is continuous, that is, sizes of workers vary from just above a=20 millimeter to over 2 cm in length. The smallest workers are found mostly = inside=20 the nest, caring for the fungus and larvae, whereas the bigger ones = engage in=20 foraging, leaf cutting, and transport of leaf fragments to the nest. = Atta=20 colonies have a special soldier caste, formed by very large workers with = strong=20 mandibles, which make them have big heads with small brains.

Winged Sexual Individuals

The sexually active individuals are born with transparent = wings that=20 allow them to engage in the nuptial flight, at the beginning of the rain = season,=20 where they copulate. Males die shortly after a single copulation. Once = the=20 winged female is inseminated, she lands on the ground, sheds her wings, = looks=20 for an appropriate sandy spot, and starts digging her initial nest. = Often winged=20 individuals are found inside the nest. These are virgin males or a queen = that=20 failed to swarm or will do so in the future. Virgin females in addition = have=20 much smaller gasters compared to their physogastric mothers. Males have = much=20 smaller heads than females. Males show little activity and their main = role is=20 transferring their sperm to the female=92s spermatheca.

The Super-Organism

In a given leaf-cutter ant colony we generally find only a = single=20 queen and her daughters, the workers. Thus, a colony is a family unit. = The=20 coordination between the members of a colony-family is so tight that = some=20 authors consider the colony to be a super-organism. That is, each = individual=20 acts to optimize the adaptive value of the colony rather than itself. = The queen=20 dedicates herself to produce eggs, the gut of the larvae and that of = workers=20 works like a communal digestive system, the foragers work as the movable = extremities of the colony providing the required food, and the glandular = secretions of all individuals mingle so as to produce a characteristic = colony=20 odor. Thus, the colony has properties that are known to characterize an=20 organism. This is especially true when we look at the systems regulating = the=20 temperature and humidity of the nest.

The Uninvited Guests

A leaf-cutting ant colony hosts a number of other animals. = They=20 range from invertebrate ectoparasites to vertebrates such as snakes, = lizards and=20 birds. They normally feed on the brood and/or use the nest as shelter. = Reptiles=20 especially profit from the nice temperature in a leaf-cutting ant nest, = which is=20 suitable for their eggs. That is certainly the case with the snake=20 Elapomorphus lemniscatus (family Boidae) which is a frequent = visitor of=20 Acromyrmex nests. Once inside the nest, the reptile houses in a = fungus=20 chamber, eats the ant=92s larvae, and exits the nest only when searching = for a=20 mate. Beetles, spiders, isopods, flies, collembolans, mites and other = arthropods=20 are known to live inside the ants=92 nest. A large number of these = ectoparasites=20 live in the refuse chamber where they feed on dead ants and other refuse = of the=20 colony.

The Nest

Atta Nests

The nests of the ants from the genus Atta are among = the=20 largest and more complex known among insects. A typical nest may occupy = a=20 subterranean space of 20 m in diameter and 5 m in depth. It consists of=20 interconnected galleries leading to chambers where ants cultivate their=20 symbiotic fungi. Other galleries lead to the surface and are used to = regulate=20 the flux of fresh air so as to maintain the temperature, carbon dioxide=20 concentration and humidity constantly at the optimum inside the nest. A = large=20 chamber is used to dispose of the colony=92s waste which consists mainly = of dead=20 fungus and dead ants. This chamber produces heat and carbon dioxide that = allows=20 for a better regulation of the colony=92s climate. Other galleries lead = deep into=20 the soil for access to the ground water. Galleries leading to the = surface end=20 beneath an earth mound with a crater, allowing the exit and entrance of = workers.=20 These mounds work as chimneys, helping to regulating the flux of fresh = air and=20 stopping running water from flooding the nest after torrential rains. = Often=20 workers seal the nest entrances with dry leaves and earth, especially = when=20 fighting other colonies. Some underground galleries lead up to 200 m = from the=20 nest to the colony=92s foraging areas. Each Atta species builds = nests with=20 its own characteristics. For example, Atta colombica has no = refuse=20 chamber but throws its rubbish outside the nest. The form of the mounds = with=20 craters depends on the soil=92s texture. The depth of the nest is also = dependent=20 on soil conditions; for example, nests are shallower in more humid = areas.=20

The Nests of Acromyrmex and = Trachymyrmex

Nests of Acromyrmex species have variable = architecture,=20 depending on the species that builds it. Acromyrmex landolti = builds nests=20 with a single vertical gallery with nest chambers connected on the sides = of the=20 gallery. The vertical gallery can go as deep as 3=964 m. Acromyrmex=20 octospinosus builds more superficial subterranean nests with many = chambers=20 and galleries connected in all directions. Acromyrmex coronatus = sometimes=20 builds superficial nests under accumulated dry leaves. These nests are = often=20 found in caves of dead tree trunks. Nests of Trachymyrmex species = are=20 less elaborate. They are subterranean but normally close to the surface = and=20 rather small compared to those of Acromyrmex and Atta = (Figs. 28=20 and 29).=20

Leaf-Cutting Ants (Formicidae: = Myrmicinae:=20 Attini), Figure 28 Nest of Atta laevigata.

Leaf-Cutting Ants (Formicidae: = Myrmicinae:=20 Attini), Figure 29 Acromyrmex landolti (drawing by = Eduardo=20 Perez).

The Life Cycle

Reproduction

Leaf-cutting ants reproduce sexually, with copulation = occurring=20 during the nuptial flight. Normally a virgin queen mates with several = males.=20 Nuptial flight occurs at the beginning of the rainy season. After the = first=20 heavy rain, thousands of winged individuals start flying from all = affected nests=20 in an area. They fly normally in the early evening and concentrate in a = given=20 space several meters above the ground. As males die shortly after = mating, dead=20 bodies of males that have copulated rain to the ground, just under the = visible=20 swarm. Once successfully inseminated, the new queen flies to the ground, = sheds=20 her wings and starts looking for an appropriate site to excavate a nest. = Queens=20 also reproduce asexually; non-fertilized eggs will produce haploid males = (as are=20 all hymenoptera, these ants are haplodiploid).

The Birth of a Colony

Once the recently fertilized queen has found an adequate = site (often=20 next to a small tree), she starts digging using her mandibles to remove = earth.=20 She digs a small gallery about 15 cm long leading to a small chamber = about 7 cm=20 wide. After digging is completed, she seals the nest entrance. In the = chamber=20 she lays some unfertilized eggs upon which she deposits the mycelium of = the=20 fungus she has taken from her mother=92s nest, stored in a cavity in her = mouth. If=20 available, roots are also used as a substrate for the fungus. Then she = lays her=20 first fertilized eggs. The first larvae that emerge are fed with special = unfertile =93trophic=94 eggs produced by the queen. Eventually small = =93nanitic=94=20 workers emerge. During that period, the queen survives on her fat = reserves and=20 by metabolizing her wing muscles, now useless. Once the first nanitic = workers=20 emerge, they open the nest entrance and start foraging for leaves, = flowers,=20 insect droppings, seeds, etc., to feed the fungus. After this initial = phase,=20 larger workers emerge capable of cutting fresh leaves.

The Life Cycle of a Worker

Leaf-cutting ants have four phases in their life cycle: = egg, larvae,=20 nymph and adult. The white larvae feed mainly on the symbiotic fungus. = They have=20 very little mobility. After several molts, the larvae initiate its = metamorphosis=20 without spinning a cocoon and eventually emerges as an adult worker. The = amount=20 of food ingested during the larval stage will determine the final size = of the=20 worker, as adult workers stop growing. Young workers have a lightly = pigmented=20 soft cuticle, whereas older ones are darker. Adult workers have = different=20 nutritional requirements than larvae do; they feed mainly on plant sap = because=20 their protein requirement is less than that of the larvae, which mainly = feed on=20 the fungus.

The Life Cycle of a Colony

The colony starts with the birth of the first nanitic = workers.=20 During the first few years, the colony grows continuously in number of = workers=20 and in the space occupied by the nest. During that period, workers = forage every=20 day, except when it is raining copiously, and feed their fungus, which = occupies=20 ever more chambers which must be prepared by the workers. When the = colony=20 reaches maturity (4=966 years after being initiated by the queen for = Atta,=20 and less for Acromyrmex and Trachymyrmex), the colony = produces=20 winged individuals that accumulate in the nest until the initiation of = the next=20 rainy season, when they will leave the nest for their nuptial flight. We = know=20 little of how the leaf cutter ant colony produces winged females instead = of=20 workers, or how the queen decides to fertilize an egg with sperm, = producing=20 female offspring, or lay unfertilized eggs that will produce males. It = is likely=20 that these ants use hormones for this purpose, as bees do, but little is = known=20 for leaf-cutting ants. After a few nuptial flights, the colony is left = without=20 winged individuals and with few workers. The colony then has the whole = year to=20 again build its workforce, forage for food to grow its fungus, and = eventually=20 produce a new batch of individuals capable of sexual reproduction. This = cycle is=20 repeated until the old queen dies, which may take several decades.=20

Communication

Pheromones

The most common communication system between workers of = leaf-cutting=20 ants consists of volatile chemicals or odors called pheromones. These = ants=20 produce their pheromones through a series of exocrine glands located in=20 different parts of their body. Each of these glands produces a specific=20 secretion composed of a mix of chemicals which are used to communicate=20 information between workers. These multicomponent pheromones are = perceived=20 through the insect=92s antennae. Some pheromones are detected only upon = contact of=20 the antenna with the substrate containing the pheromone; these are = called=20 contact pheromones. For example, larval recognition in leaf-cutting ants = seems=20 to be achieved using contact pheromones that are located on the cuticle = of the=20 larvae.

Alarm Pheromones

Leaf-cutting ant workers produce an alarm pheromone in = their=20 mandibular glands. This pheromone is composed of highly volatile = compounds that=20 disperse rapidly in the air and alarm nestmates up to distances of 60 = cm. This=20 same pheromone helps workers and soldiers to orientate to the source or = location=20 that caused the alarm, thus helping to coordinate cooperation in = defense. This=20 coordination can be very fast and effective when fighting other ant = colonies or=20 when deterring an intruder. The compounds of this pheromone are absorbed = on the=20 cuticle of these insects, thanks to the cuticular hydrocarbons that = cover the=20 insect=92s surface. There, the alarm pheromone serves as an individual = recognition=20 signal, helping ants to differentiate between nestmates and intruders.=20

Recruitment Pheromone

Another well studied pheromone is the recruitment or trail = pheromone. This pheromone is produced in the poison gland located in the = gaster.=20 Workers lay trails using this pheromone when returning from a palatable = food=20 source. The less volatile compounds in the pheromone serve as = orientation cues,=20 whereas the more volatile ones regulate the amount of new workers = recruited to=20 the trail, which in turn will depend on the quality and quantity of food = to=20 which they are recruited. Some of the compounds of this pheromone, = isolated from=20 leaf-cutting ant species, are 4-metyl-2-pyrrol-carboxilate for Atta=20 laevigata, Atta cephalotes, Acromyrmex octospinosus, = and=20 3-ethyl-2,5-dimethyl-pyrazine for Atta sexdens rubropilosa and=20 Acromyrmex niger.

Territorial Pheromones

These are used to mark the colony=92s territory using a = secretion from=20 the Dufour=92s gland. Workers on territories marked by their colony are = more=20 aggressive and fight longer, whereas workers over territories of a = foreign=20 colony try to escape. During ant wars, many small workers swarm outside = their=20 nest to mark the surroundings of their nest entrance and the = battlefield,=20 whereas larger workers engage in combat. These combats can last weeks if = the=20 food source they are defending is especially scarce or attractive. This=20 pheromone helps achieve the harmonious cohabitation of neighboring = colonies of=20 similar size, which seems to be the norm in nature.

Individual Recognition

Leaf-cutting ant workers recognize their nestmates and = differentiate=20 them from foreign ants, even if they are of the same species. They = achieve this=20 by detecting odors absorbed on the cuticle. These odors mainly come from = the=20 mandibular alarm pheromone. This system allows them to recognize = individuals=20 even at a distance, and is also used to recognize nestmates from = different parts=20 of the nest.

Other Pheromones

Other pheromones are less well known. The queen is = recognized as=20 such by pheromones, and so are the larvae. Leaf fragments are marked = using the=20 Dufour=92s gland, probably to ascertain the colony=92s property and to = help detect=20 it when workers drop it from the trees. Other communication systems = remain to be=20 discovered.

Visual Communication

Although known from other ants, nothing is known for = leaf-cutter=20 ants.

Communication by Sound

Leaf-cutter ants have a stridulatory apparatus between the = gaster=20 and the second petiole. This apparatus consists of a stridulated = cuticular=20 surface on the dorsal part of the gaster and a cuticular tooth extruding = from=20 the petiole that serves as a bow producing vibrations when gliding over = the=20 stridulated surface. The ants produce rhythmic ultrasounds with this = apparatus=20 by vibrating and elevating their gaster. The sound is barely perceived = by some=20 humans. The ants use this sound to guide workers toward branches on = trees that=20 need leaf-cutters. They also stridulate when buried, guiding workers to = dig in=20 their direction. The sound is transmitted over solid substrate and is = perceived=20 thanks to special receptors at the end of their legs.

Trophalaxis and Antennal Contacts

Workers exchange liquids through their mouthparts. They do = this=20 after contacting the other ant with their antennae. We do not know what=20 information they are transmitting with these behaviors but they might = transmit=20 information about the quality of food.

Foraging

Leaf-cutting ants have especially sophisticated foraging = behaviors.=20 They use trunk trails that they maintain free of leaves and obstacles,=20 facilitating their movement from the nest to the foraging areas. From = this trunk=20 trail, smaller trails lead to the plants they harvest. Scout ants use = the same=20 trail system but eventually leave it to explore new terrain. When = finding a=20 palatable food source, they return to the nearest trunk trail, leaving = trail=20 pheromone behind. Foragers might cut a leaf fragment and carry it = through the=20 trail system to the nest. Alternatively, they might do it in stages; = some=20 workers cut the leaves and let them drop from the tree, or just carry = the leaves=20 to the nearest junction, where other workers collect the leaf fragments = and=20 carry them to the nest entrance, where still other workers remove the=20 superficial waxes from the leaves and carry the clean leaves inside the = nest,=20 where other workers cut them into fine pieces to feed the fungus. Very = small or=20 minima workers are often seen climbing on leaf fragments that are = carried by=20 larger workers. The minima help fend off flies from the family Phoridae = and=20 other parasites that attempt to attack the carrying worker or enter the = nest via=20 the leaf fragment. Older workers remove dry leaves, dead ants and dry = fungus=20 from the nest chambers and drop it in the refuse pile.

The Decision Making System

The colony has to coordinate its efforts in order to = forage=20 efficiently. Not all ants are required at the same place at the same = time.=20 Specifically, in the case of recruitment to food, leaf-cutting ants use = a=20 decision making system that is different from that used by all other = ants=20 studied and which resembles the system used by termites. It works as = follows: a=20 scout discovering food will return to the nest or the trunk trail while = laying a=20 pheromone trail. The concentration of pheromone on the trail will depend = on the=20 quality of the food. Workers encountering this trail, according to their = motivation for food, will follow the new trail. On their way back, they = will=20 reinforce the pheromone trail but in such a way that the total = concentration of=20 pheromone is roughly constant. They will not just add their pheromone = onto that=20 of others (as done by Solenopsis geminata, for example), but will = restrain from adding pheromone if enough of it is already on the trail. = In this=20 way, the concentration of pheromone on the trail will not depend on the = amount=20 of ants laying trails or returning from the food source but only on the = quality=20 of the food. This system allows leaf-cutters to recruit to various = different=20 food sites at the same time.

Orientation

Foraging scouts need to find their way back to the nest. = They=20 achieve this by integrating various keys or environmental signals when = foraging=20 and homing. We know that they can use polarized light, visual cues, = spatial=20 memory, olfactory cues and tactile cues. They might possibly also use=20 gravitational cues and the earth=92s magnetic field. Each species seems = to have=20 its own hierarchy in which these cues are used in actual orientation in = the=20 field.

Defense

Organisms need to defend themselves against predators and = parasites=20 in order to increase their odds for survival. Leaf-cutter ants possess = no=20 functional sting but they do secrete a toxic fluid from their anal = region which=20 they deposit onto enemy combatant ants. This secretion dries or = polymerizes=20 quickly, helping to immobilize the affected worker. It also seems to = contain=20 neurotoxic substances and volatiles that attract other ants. Large = workers and=20 soldiers also use their mandibles to cut wounds on molesting vertebrates = and=20 then add irritant secretions to the wounds. The mandibles can cut = appendages=20 from invertebrate enemies or even cut them into pieces. Another form of = defense=20 is using cryptic behavior, including cessation of all movement until the = danger=20 has passed. This behavior is often used by lone foraging scouts. The = thick, hard=20 cuticle and the spines over the thorax help the insect to avoid being = eaten by=20 predators such as lizards, birds and spiders. In addition, the secretion = of the=20 metapleural gland is spread all over the body of these insects, helping = to fend=20 off fungi, bacteria and other potential illness-causing agents. The main = predators of these ants are spiders, armadillos and anteaters, which = feed=20 copiously on them.

Ecology

Leaf-cutting ants are only found in the neotropics. They = might=20 have an important effect on plant growth, on occasion achieving pest = status.=20 Densities of up to 60 adult Atta nests per hectare, or hundreds = to=20 thousands of Acromyrmex nests per hectare, can cause serious harm = to=20 plantations and crops. On the other hand, the large accumulation of = organic=20 material inside the nest, once the queen and the colony die, makes it a=20 formidable substrate for plant growth. In nutrient poor savannas, old = collapsed=20 Atta nests are colonized by trees and shrubs, forming forested = islands.=20 It is the occasional status as agricultural and forestry pests that has = made=20 leaf-cutters most famous. Leaf-cutters with the highest economic = importance as=20 pests are all Atta species and some Acromyrmex species. An = adult=20 Atta laevigata colony, for example, may cut an average of 5 kg of = plant=20 material each day. They target especially young plant tissue, which can = be=20 especially damaging to the plants. Estimates of the economic damage = caused in=20 the Americas by these ants are close to a hundred billion US dollars per = year=20 (Figs. 30=20 and 31).=20

Leaf-Cutting Ants (Formicidae: = Myrmicinae:=20 Attini), Figure 30 Atta laevigata: (a) male,=20 (b) queen, (c) worker, (d) soldier (drawings by = Eduardo=20 Perez).

Leaf-Cutting Ants (Formicidae: = Myrmicinae:=20 Attini), Figure 31 Atta cephalotes transporting = vegetation=20 (photo by Scott Bauer, USDA).

Control of Leaf-Cutting Ants

There are three basic methods for reducing the populations = of these=20 pest ants: (i) Stopping the ants from accessing the plant. Asphalt or = other=20 sticky greases painted at the base of tree trunks, plastic or rubber = rings that=20 retard ants from reaching the plants, or water barriers that deter the = access of=20 these ants are useful ways of dealing with this pest. (ii) Direct = control,=20 killing the ants with insecticides. For large nests this might be = difficult and=20 expensive. Fogging and powder pumps may help to get the insecticide = inside a=20 nest. (iii) Indirect control, using poisoned but attractive baits, is = the most=20 efficient way known to manage large areas with this pest. Attractants = (such as=20 citrus pulp) mixed with slow acting insecticides, allow the foragers to = carry=20 the poison into the nest so that it is distributed to all chambers by = the=20 workers before it starts killing the insects.

References

Cameron R, Currie CR, Scott JA, = Summerbell RC,=20 Malloch D (1999) Fungus-growing ants use antibiotic-producing = bacteria to=20 control garden parasites. Nature 398:701=96704

Cede=F1o A (1984) La ecolog=EDa de los = bachacos.=20 Fondo Editorial Acta Cientifica Venezolana, Caracas

Chapela IH, Rehner SA, Schultz TR, = Mueller UG=20 (1994) Evolutionary history of the symbiosis between = fungus-growing ants=20 and their fungi. Science 266:1691=961694
3DPubMed=20

Herman HR (1982) Social insects, vol 4. = Academic=20 Press, New York, NY, 385 pp

Hernandez JV, Lopez H, Jaffe K (2001) = Nestmate=20 recognition signals of the ant Atta laevigata. J Insect = Physiol=20 48:287=96295

Jaffe K (1984) Negentropy and the = evolution of=20 chemical mass recruitment in ants. J Theor Biol 106:587=96604 =

Jaffe K (1987) The evolution of agonistic = communication systems in ants. Exp suppl 54:295=96311

Jaffe K, Howse PE (1979) The mass = recruitment=20 system of the leaf-cutting ant Atta cephalotes. Anim Behav=20 27:930=96939

Jaffe K, Villegas G, Colmenares O, Puche = H,=20 Zabala N, Alvarez M, Navarro JG, Pino E (1985) Two different = decision=20 making systems in ants. Behavior 92:9=9621

Mayhe A, Jaffe K (1998) On the = biogeography of=20 the Attini. Ecotropicos 11:45=9654

Vilela E, Jaffe K, Howse PE (1987) = Orientation=20 in leaf-cutting ants. Anim Behav 35:1443=961453

Weber N (1972) The gardening ants. = American=20 Philosophical Society, Philadelphia, 142 pp

Wheeler WM (1965) Ants: their structure, = development and behaviour, 4th edn. Columbia University Press, New = York,=20 NY, 663 pp

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Wilson EO (1971) The insect societies. = Belknap=20 Press, Harvard University, Cambridge, MA, 548 pp

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