Load-size determination in the leaf-cutting ant, Atta cephalotes

I examined load-size determination by a highly polymorphic leaf-cuttingant, Atta cephalotes, cutting leaves of artificial trees (branchesplaced in the top of plastic tubes). I compared load size forants cutting thin leaves (starfruit, Averrhoa carambola) andthick leaves (grapefruit, Citrus parodist). At each source,larger ants cut larger fragments. Distance from the nest hadno effect on load size. The mass of fragments cut by an antof a given size was significantly greater when cutting grapefruitleaves. The leaf area cut, however, showed no significant differencebetween the two leaf types. Leaf area increased approximatelyin proportion to ant body mass to the 0.6 power. As a resultof their method of load-size determination, ants of a givensize cut heavier loads when cutting the thicker leaves. Thisdifference, however, was counteracted at the colony level byrecruitment of larger ants, which cut smaller area fragmentsrelative to their body mass, to cut at thicker leaf sources.     

Hitchhiking and the removal of microbial contaminants by the leaf‐cutting ant Atta colombica

1. The ecologically dominant leaf‐cutting ants exhibit one of the most complex forms of morphological caste‐based division of labour in order to efficiently conduct tasks, ranging from harvesting fresh leaf material to caring for the vulnerable fungal crop they farm as food. While much of their division of labour is well known, the role of the smallest workers on foraging trails is puzzling. Frequently these minim workers hitchhike on leaf fragments and it has been suggested that they may act to reduce the microbial contamination of leaf material before they enter the nest. Here we investigated this potentially important role of minims with field colonies of Atta colombica. 2. We experimentally increased the microbial load of leaf fragments and found that this resulted in minims hitchhiking on leaf fragments for longer. Furthermore, we show that leaves naturally have a significant microbial load and that the presence of hitchhikers reduces the microbial load of both experimentally manipulated and natural leaf fragments. 3. Intriguingly, the microbial load of leaves high in the canopy where ants were foraging was much lower than closer to the ground where the ants avoided cutting leaves. This suggests that the often perplexing foraging patterns of leaf‐cutting ants may in part be explained by the ants avoiding leaves that are more heavily contaminated with microbes. 4. The removal of microbial contaminants is therefore an important role of hitchhiking minim workers in natural colonies of Atta leaf‐cutting ants, although other tasks such as trail maintenance and defence also explain their occurrence on trails.     

Cutters,carriers and transport chains: Distance-dependent foraging strategies in the grass-cutting ant <Emphasis Type="Italic">Atta vollenweideri</Emphasis>

Summary Most studies on leaf-cutting ant foraging examined forest species that harvest dicot leaves. We investigated division of labor and task partitioning during foraging in the grass-cutting ant Atta vollenweideri. Workers of this species harvest grass fragments and transport them to the nest for distances up to 150 m along well-established trunk trails. We recorded the behavior of foraging ants while cutting and monitored the transport of individually-marked fragments from the cutting site until they reached the nest. A. vollenweideri foragers showed division of labor between cutting and carrying, with larger workers cutting the fragments, and smaller ones transporting them. This division was less marked when plants were located very close to the nest and no physical trail was present, i.e., the cutter often transported its own fragment back to the nest. On long foraging trails, the transport of fragments was a partitioned task, i.e., workers formed transport chains composed of 2 to 5 carriers. This sequential load transport occurred more often on long than on short trails. The first carriers in a transport chain covered only short distances before dropping their fragments, and they were observed to turn back and revisit the patch. The last carriers covered the longest distance. The probability of dropping the carried fragment on the trail was independent of both worker and fragment size, and there was no particular location on the trail for dropping, i.e., fragments were not cached. Transport time of fragments transported by a chain was longer than for those transported by single workers all the way to the nest, i.e., sequential transport did not save foraging time. Two hypotheses concerning the possible adaptive value of transport chains are discussed. The first one argues that sequential transport may lead to an increased material transport rate compared to individual transport. The second one considers sequential transport as a way to enhance the information flow among foragers, thus leading to a quicker build-up of workers at particular harvesting places. It is suggested that rather than increasing the gross transport rate of material, transport via chains may favor the transfer of information about the kind of resource being actually harvested.Received 19 December 2002; revised 14 March 2003; accepted 19 March 2003     

The predatory behavior of Pheidole megacephala

We studied the foraging and predatory behaviors of the invasive African myrmicine ant, Pheidole megacephala (F.) in its native range. Workers can singly capture a wide range of insects, including relatively large prey items. For still larger prey, they recruit at short range those nestmates situated within reach of an alarm pheromone and together spread-eagle the insect. These behaviors are complimented by a long-range recruitment (of nestmates remaining in the nest) based on prey size. P. megacephala scouts also use long-range recruitment when they detect the landmarks of termites and competing ant species, thus permitting them to avoid confronting these termites and ants solitarily.     

Trail Communication During Foraging and Recruitment in the Subterranean Termite Reticulitermes santonensis De Feytaud (Isoptera, Rhinotermitidae)

The search for food in the French subterranean termite Reticulitermes santonensis De Feytaud is organized in part by chemical trails laid with the secretion of their abdominal sternal gland. Trail-laying and -following behavior of R. santonensis was investigated in bioassays. During foraging for food termites walk slowly (on average, 2.3 mm/s) and lay a dotted trail by dabbing the abdomen at intervals on the ground. When food is discovered they return at a quick pace (on average, 8.9 mm/s) to the nest, laying a trail for recruiting nestmates to the food source. While laying this recruitment trail the workers drag the abdomen continuously on the ground. The recruitment trail is highly attractive: it is followed within a few seconds, by more nestmates, and at a quicker pace (on average, 6.4 mm/s) than foraging trails (on average, 2.9 mm/s). The difference between foraging and recruitment trails in R. santonensis could be attributed to different quantities of trail pheromone. A caste-specific difference in trail pheromone thresholds, with workers of R. santonensis being more sensitive to trails than soldiers, was also documented: soldiers respond only to trails with a high concentration of trail pheromone.     

Foraging grass-cutting ants (Atta vollenweideri) maintain stability by balancing their loads with controlled head movements

Grass-cutting ants (Atta vollenweideri) carry leaf fragments several times heavier and longer than the workers themselves over considerable distances back to their nest. Workers transport fragments in an upright, slightly backwards-tilted position. To investigate how they maintain stability and control the carried fragment’s position, we measured head and fragment positions from video recordings. Load-transporting ants often fell over, demonstrating the biomechanical difficulty of this behavior. Long fragments were carried at a significantly steeper angle than short fragments of the same mass. Workers did not hold fragments differently between the mandibles, but performed controlled up and down head movements at the neck joint. By attaching additional mass at the fragment’s tip to load-carrying ants, we demonstrated that they are able to adjust the fragment angle. When we forced ants to transport loads across inclines, workers walking uphill carried fragments at a significantly steeper angle, and downhill at a shallower angle than ants walking horizontally. However, we observed similar head movements in unladen workers, indicating a generalized reaction to slopes that may have other functions in addition to maintaining stability. Our results underline the importance of proximate, biomechanical factors for the understanding of the foraging process in leaf-cutting ants.     

Stingless bees (<Emphasis Type="Italic">Scaptotrigona pectoralis</Emphasis>) learn foreign trail pheromones and use them to find food

Foragers of several species of stingless bees (Hymenoptera, Apidae and Meliponini) deposit pheromone marks in the vegetation to guide nestmates to new food sources. These pheromones are produced in the labial glands and are nest and species specific. Thus, an important question is how recruited foragers recognize their nestmates’ pheromone in the field. We tested whether naïve workers learn a specific trail pheromone composition while being recruited by nestmates inside the hive in the species Scaptotrigona pectoralis. We installed artificial scent trails branching off from trails deposited by recruiting foragers and registered whether newly recruited bees follow these trails. The artificial trails were baited with trail pheromones of workers collected from foreign S. pectoralis colonies. When the same foreign trail pheromone was presented inside the experimental hives while recruitment took place a significant higher number of bees followed the artificial trails than in experiments without intranidal presentation. Our results demonstrate that recruits of S. pectoralis can learn the composition of specific trail pheromone bouquets inside the nest and subsequently follow this pheromone in the field. We, therefore, suggest that trail pheromone recognition in S. pectoralis is based on a flexible learning process rather than being a genetically fixed behaviour.     

The Function of Hitchhiking Behavior in the Leaf‐cutting Ant Atta cephalotes1

In some leaf‐cutting ant species, minim workers ride on the fragments of leaves as they are carried back to the nest from the cutting site. There is convincing evidence that these “hitchhikers” can protect the leaf carriers from attack by phorid (Diptera: Phoridae) parasitoids, but we consider the possibility of other functions for the hitchhiking behavior. It has been hypothesized that the hitchhikers (1) feed on leaf sap from the edges of the cut leaves; (2) ride back to the nest to save energy; (3) get caught on the fragments as they are cut, and hitchhike because they cannot (or will not) get off; and (4) begin the process of preparing the leaf to enter the fungal gardens in the nest, perhaps by removing microbial contaminants. We observed hitchhikers of Atta cephalotes in 14 nests at the La Selva Biological Station in Costa Rica. There was no difference in the proportion of leaf carriers with hitchhikers between day and night. Because the nests we observed were largely nocturnal, more than 90 percent of the hitchhiking occurred at night. The phorid parasitoids are usually considered to be diurnal, so the preponderance of nocturnal hitchhiking suggests other functions in addition to parasitoid defense. Hitchhikers spent more time in the defensive head‐up posture during the day, but spent more time in the head‐down posture at night. The head‐down posture may indicate cleaning or other leaf preparation. The hitchhikers were never observed feeding on sap. Hitchhikers frequently got onto and off of the fragments, and so they were not “marooned.” Few hitchhikers rode all the way back to the nest and were often moving on the leaf fragment; these observations make the energy conservation hypothesis less likely, although we cannot reject it. We conclude that parasitoid defense is an important function of hitchhiking but also that there are probably other functions when parasitoids are absent. Based on available data, the most likely possibility is preparation of the leaf fragment before it enters the nest.     

Foraging by Atta sexdens (Formicidae: Attini): seasonal patterns, caste and efficiency

Abstract. 1. Atta sexdens changes diel periods of foraging, the size of its foraging territory, the numbers and lengths of foraging trails, and its rate of foraging with respect to seasonality in subtropical Paraguay. Leaf loads are significantly larger in warmer months than loads carried in cooler months.
2. Foragers segregate into three labour groups in the field: a small subset climb trees, cut large quantities of vegetation, and drop them to the ground; the second subset of foragers searches out these leaf caches, cut diem into smaller pieces, and carry and deposit the leaf fragments on the foraging trail; the third subset of workers retrieves leaf fragments on the trail and carry them to the nest.
3. Pitfall trapping shows a large degree of patchiness in activity, with media workers dominating the foraging population, more so closer to vegetation which is being harvested.
4. The recovery efficiency of the multi-staged foraging behaviour is estimated to be only 49%, with the recovery of leaf caches near 50%. The impact of A. sexdens may, thus, be twice as great as previous estimates on their herbivory.     

Predatory abilities favour the success of the invasive ant Pheidole megacephala in an introduced area

Abstract:  The invasive African big-headed ant, Pheidole megacephala , is a dominant species in the many areas it has invaded. We examined whether one potential reason for its ecological success might be its predatory efficiency. We compared the density of termite nests in an area of Mexico invaded by P. megacephala with an adjacent area where P. megacephala is not present. We also compared the success of P. megacephala in preying on termites with that of 13 native ant species. We found that termite nest density was significantly lower in areas invaded by P. megacephala (0.33 vs. 1.05 nests per 30 m transect). In field trials, we established that P. megacephala workers were significantly more successful at capturing termite workers from termite nest fragments than even the most successful native ant species, Dorymyrmex pyramicus . For both P. megacephala and D. pyramicus , single scouts could trigger the mass recruitment of nestmates, but P. megacephala was able to recruit greater numbers of nestmates. Combined with their aggressiveness towards other ant species, their highly efficient predatory capacities help explain the ecological success of P. megacephala and demonstrate how it can be a major threat to invertebrate biodiversity in the areas it invades.     

Host-ant trail following by myrmecophilous larvae of Liphyrinae (Lepidoptera,Lycaenidae)

In this study we report a case of ant-trail following by lycaenid caterpillars. Euliphyra mirifica and E. leucyana caterpillars are involved in a commensal association with the weaver ant Oecophylla longinoda. The host nests are made with leaves which over the course of time dry out or are broken open by storms, forcing the ants to migrate and build a new nest elsewhere. Euliphyra caterpillars are stimulated by recruitment behaviour which triggers the migration of their host. They then follow the host trails leading to the new nesting site. Laboratory experiments showed that these caterpillars are able to follow host trails under varied conditions: (1) fresh trails actually used by workers, (2) fresh trails in the absence of workers, (3) heterocolonial, 2-month-old trails, and (4) fresh trails washed with water (to simulate the effect of tropical rains). They can also bridge trail gaps of more than 1 cm. Under natural conditions, the trails are frequently situated along thin twigs. The forward progress of the ants in such a situation is not impeded by the presence of large Euliphyra larvae. Workers just climb over the caterpillars, even on larger trails where there is enough room to pass alongside them. This suggests that an allomone is secreted on the dorsal part of the caterpillars. When crawling along heterocolonial trails, the caterpillars are not attacked, even if about 21% of the workers from the new colony spread their mandibles when encountering them. They are then adopted and are admitted to the nest of the new host colony of O. longinoda.     

The ‘truck‐driver’ effect in leaf‐cutting ants: how individual load influences the walking speed of nest‐mates

The foraging behaviour of social insects is highly flexible because it depends on the interplay between individual and collective decisions. In ants that use foraging trails, high ant flow may entail traffic problems if different workers vary widely in their walking speed. Slow ants carrying extra‐large loads in the leaf‐cutting ant Atta cephalotes L. (Hymenoptera: Formicidae) are characterized as ‘highly‐laden’ ants, and their effect on delaying other laden ants is analyzed. Highly‐laden ants carry loads that are 100% larger and show a 50% greater load‐carrying capacity (i.e. load size/body size) than ‘ordinary‐laden’ ants. Field manipulations reveal that these slow ants carrying extra‐large loads can reduce the walking speed of the laden ants behind them by up to 50%. Moreover, the percentage of highly‐laden ants decreases at high ant flow. Because the delaying effect of highly‐laden ants on nest‐mates is enhanced at high traffic levels, these results suggest that load size might be adjusted to reduce the negative effect on the rate of foraging input to the colony. Several causes have been proposed to explain why leaf‐cutting ants cut and carry leaf fragments of sizes below their individual capacities. The avoidance of delay in laden nest‐mates is suggested as another novel factor related to traffic flow that also might affect load size selection The results of the presennt study illustrate how leaf‐cutting ants are able to reduce their individual carrying performance to maximize the overall colony performance.     

An Overlooked Mandibular-Rubbing Behavior Used during Recruitment by the African Weaver Ant,Oecophylla longinoda

In Oecophylla, an ant genus comprising two territorially dominant arboreal species, workers are known to (1) use anal spots to mark their territories, (2) drag their gaster along the substrate to deposit short-range recruitment trails, and (3) drag the extruded rectal gland along the substrate to deposit the trails used in long-range recruitment. Here we study an overlooked but important marking behavior in which O. longinoda workers first rub the underside of their mandibles onto the substrate, and then—in a surprising posture—tilt their head and also rub the upper side of their mandibles. We demonstrate that this behavior is used to recruit nestmates. Its frequency varies with the rate at which a new territory, a sugary food source, a prey item, or an alien ant are discovered. Microscopy analyses showed that both the upper side and the underside of the mandibles possess pores linked to secretory glands. So, by rubbing their mandibles onto the substrate, the workers probably spread a secretion from these glands that is involved in nestmate recruitment.     

The dynamics of collective exploration and trail-formation in Monomorium pharaonis: experiments and model

Abstract. When exploring a chemically unmarked area devoid of food sources, workers of the pest ant Monomorium pharaonis L. (Formicidae, Myrmicinae) leave scent marks on the ground and after 30–60min a network of diverging exploratory trails begins to emerge.
Exploratory activity is affected by the nutritional state of the colony and a period of food deprivation induces a dramatic increase in the number of workers leaving the nest. A mathematical model based on a logistic growth equation is proposed to describe the exploratory recruitment observed. When travelling along exploratory trails the proportion of ants displaying trail-laying behaviour is higher for outbound than for nestbound workers. Outbound ants also show a greater propensity than nestbound ants to follow the scent marks of their nestmates. The chemical used to mark a novel area does not appear to be colony-specific and thus does not have a territorial function sensu stricto. The adaptive value of the collective exploratory behaviour observed in this study is discussed in relation to the common features of other pest ant species described in the literature.     

Ontogeny of queen attraction to workers in the antCataglyphis cursor (Hymenoptera: Formicidae)

Summary The behaviour ofCataglyphis cursor workers towards queens at 15 days, one month or two months after worker emergence was tested. Workers reared entirely with their own maternal queen were tested with this queen or with an unfamiliar alien queen. Workers transferred within 48 h of emerging to a new definitive nest with an alien queen were tested with this queen or with the original maternal queen. The degree of attraction to each of these queens and the workers' behavioural repertoire were measured and analysed. The results showed the following: 1) The attractiveness of queens and the workers' queen recognition behaviour were linked. 2) Although unfamiliar alien queens hardly attract workers, familiar alien queens were as attractive as maternal queens, and induced the same strongly marked and unique worker response, indicating that workers learn queen attractant cues in the days immediately after emergence. 3) Agonistic reactions were observed, but workers continued to be attracted to their maternal queen even after developing an attraction response to an alien queen with which they had been reared. These results agree with the proposal that queens produce two kinds of pheromones, those that attract workers and those that mediate recognition of queens by workers. These results show the ability of workers to discriminate between queens. Workers are attracted to any queen, but recognize as nestmates only maternal or alien queens with which they have been maintained. 4) The differential in worker attraction and recognition from 15 days to 2 months and its modifications by post-imaginal experience illustrate worker behavioural ontogeny, which is a basis of social discrimination.     

Foraging behaviour of Atta cephalotes (leaf-cutting ants): an examination of two predictions for load selection

Two mechanisms have been proposed to explain how colony-level foraging performance of leaf-cutting ants can be maximized when workers harvest leaf fragments of a size that does not maximize their individual performance. Each mechanism predicts that ants will adjust the size of leaf fragments between starting a foraging bout and establishing full traffic between the nest and foraging site, but the two models predict shifts in opposite directions. I examined fragment sizes at the start of daily foraging in five field colonies of Atta cephalotes in Costa Rica and detected an obvious shift in only one case. More shifts were detected when the small and large ends of the worker body size range were considered separately, but the direction was inconsistent among colonies. I also examined the role of returning laden workers in recruitment of nestmates by intercepting all laden workers for the first 2 h of foraging, and measuring the effect on the arrival of recruits at the foraging site. In two cases, the flow of recruits was not diminished by the interception of returning workers. The results suggest that neither mechanism correctly and consistently accounts for load size selection by leaf-cutting ants. Copyright 2000 The Association for the Study of Animal Behaviour.     

Regulated food recruitment through individual behavior of scouts in the ant,Myrmica sabuleti (Hymenoptera: Formicidae)

By presenting different kinds of food sources to colonies ofM. sabuleti, we have demonstrated that this species regulates its foraging activity by altering the proportion of scouts that return to the nest to recruit nestmates after discovering a food source and by varying the number of nestmates recruited by a scout. These two parameters are related to the kind of food discovered. Our behavioral experiments showed that the probability that a scout returned to the nest decreased with a decreasing quantity of sucrose solution. In contrast, the number of returned scouts that elicited recruitment from the nest and the mean number of nestmates recruited by one of these scouts were independent of the quantity of the sucrose solution. Recruitment even occurred toward a 1- or 0.25-µl droplet of sucrose solution. When a scout discovered a large dead prey, a large drop of prey juice, a cluster of 30 dead fruit flies, or 1 isolated fruit fly, it always went back to the nest, but it elicited recruitment only when the food source was a large dead prey or a large drop of prey juice. No recruitment occurred when the food source was a single fruit fly and recruitment occurred only once in 30 trials when a cluster of 30 fruit flies was discovered.     

Forager polymorphism, size-matching, and load delivery in the leaf-cutting ant, Atta cephalotes

Abstract.

• 1 This study examined the importance of forager polymorphism and division of labour among foragers of different size for the economics of load delivery in a leaf-cutting ant, Atta cephalotes (L.). I collected A.cephalotes foragers coming down trees carrying leaf fragments to evaluate the degree of match between forager mass and the density (mass per unit area) of leaves being cut, and to quantify how this match affects whether the mass of leaf fragments cut by the ants are within the range which maximizes the rate and efficiency of load delivery.
• 2 Foragers ranged 23-fold in mass (1.4–32.1 mg). On average, larger workers cut at denser leaf sources. Leaf fragment area increased with ant mass, but relative area (fragment area/ant mass) decreased with ant mass. The density of a leaf type had little or no effect on the area cut by ants of a given size. As a result, ants of a given mass cut heavier fragments from the denser leaves. The effect of leaf density, however, was partly counteracted at the colony level by recruitment of larger ants, which cut smaller area fragments relative to their body mass, to cut at denser leaf sources.
• 3 Despite a fairly high variance in the relationship between ant mass and fragment mass, overall 87% of the laden ants (74–100% for different trees) carried leaf fragments in the 1.5–6 times body mass range. Earlier studies indicate that loads in this range yield the highest biomass transport rate and transport efficiency. Thus, the variance falls within bounds such that it has little effect on load transport efficiency. Having a broad range in optimal load mass may be considered an adaptation to the expected variability in load masses.
• 4 If there were no correlation between ant mass and leaf density, mismatches between ant mass and load mass would be more common than observed. Thus, size-matching of larger workers to cut denser leaves increases the rate and ergonomic efficiency of load delivery.

     

How does starvation affect spatial organization within nests in <Emphasis Type="Italic">Lasius niger</Emphasis>?

Spatial distribution of ant workers within the nest is a key element of the colony social organization contributing to the efficiency of task performance and division of labour. Spatial distribution must be efficiently organized when ants are highly starved and have to get food rapidly. By studying ants’ behaviour within the nest during the beginning of food recruitment, this study demonstrates how the spatial organization is affected by starvation and improves the efficiency and the speed of recruitment as well as the allocation of food. (1) In starved nests, nestmates left the deep part of the nest and crowded near the nest entrance. This modification of the spatial distribution is a local phenomenon concerning only the individuals situated in the first chamber near the nest entrance. These starved individuals have a higher probability of leaving the nest after a contact with recruiters than nestmates situated deeper in the nest. This strongly suggests that nestmates situated near the nest entrance have a low response threshold to the signals emitted by recruiters. Their higher responsiveness speeds up their exit to the foraging area and hence may increase the efficiency of highly starved colonies in exploiting new food opportunities. (2) In starved nests, the trajectory covered by recruiters between contacts with nestmates was nearly twice as small. For recruiters, this represented a gain of time in the allocation of food. As the recruitment process follows snowball dynamics, this gain of time by starved recruiters might also speed up the exploitation of food. This study evidences how the spatial distribution of individuals as a function of their motivational state might have a regulatory function in the recruitment process, which should be generic for many social species.     

Allocation of Colony‐Level Foraging Effort in Vespula germanica in Response to Food Resource Quantity,Quality, and Associated Olfactory Cues

In social insects, selection takes place primarily at the level of the colony. Therefore, unlike solitary insects, social species are expected to forage at rates that maximize colony fitness rather than individual fitness. Workers can increase the net benefit of foraging by responding to increased resource availability, by responding more strongly to higher‐quality resources, and by decreasing the uncertainty with which nestmates find resources. Unlike many ants and social bees, no social wasp is known to utilize a nest‐based recruitment signal to inform nestmates of food location. On the other hand, wasps do learn the odor of food brought to the nest and use this cue to locate the food source outside the nest. Here, we quantify the effects of three food‐associated variables on the allocation of foraging effort in the yellowjacket Vespula germanica. We used an experimental approach to assess whether resource quantity, quality, or associated olfactory information affect the probability that a forager will leave the nest on a foraging trip. We addressed these questions by inserting a known amount of sucrose solution directly into nests and recording foraging effort (departure rate) over the subsequent hour‐long observation period. No differences were found in foraging effort because of the presence/absence of olfactory cues, but there was strong evidence that foraging effort increased in response to resource influx and resource quality. Thus, while olfactory cues are learned in the nest, only resource quality and the cue of increased amount of food in the nest factor into a forager's decision of whether or not to depart on a foraging trip. However, as prior work has shown, once a wasp forager leaves the nest, it uses the learned olfactory cues to aid in finding resources.