Alarm behaviour in Eciton army ants

The effective communication of alarm can be critical for social animals so that they are able to deal with threats posed by predators and competitors. In the case of many of the most ecologically dominant, large‐colony ant species, these alarm responses are aggressive and coordinated by alarm pheromones, produced generally from the mandibular glands. In the present study, the alarm behaviour of two Neotropical army ant species is examined, the swarm raiding Eciton burchellii (Westwood) and the column raiding Eciton hamatum (Fabricius). Both species exhibit aggressive alarm responses in response to crushed heads, suggesting that the alarm pheromone is indeed produced by the mandibular glands in these ants. The most abundant component of the mandibular gland secretion, 4‐methyl‐3‐heptanone (10 µL on a rubber septum), stimulates a substantial alarm response, although this is less than the response to a single crushed head. This suggests that 4‐methyl‐3‐heptanone may be an alarm‐stimulating compound in Eciton. The alarm response of E. burchellii involves more workers than that of E. hamatum, although major workers play a much greater role in the response of the latter species. The differences in the alarm response of the two closely‐related species may relate to their foraging strategies, with E. burchellii relying more on quantity rather than the caste of ants responding and possibly using alarm pheromones for offensive as well as defensive functions.     

Filamentous fungi found on foundress queens of leaf‐cutting ants (Hymenoptera: Formicidae)

Queens of the leaf‐cutting ant species Atta laevigata and Atta capiguara were collected soon after their mating flight and maintained in the laboratory until death. Ant corpses showing signs of contamination by insect pathogenic fungi were selected for fungal identification. Filamentous fungi such as Beauveria bassiana and Paecilomyces lilacinus actively sporulated in the ant’s corpses. This is the first report of the latter fungus on reproductives of leaf‐cutting ants. The fact that queens may acquire filamentous fungi including saprophytic and potential insect pathogens after their mating event is especially interesting regarding the impacts of such microbes on the establishment of a new nest.     

Polyethism and nestmate recognition in the alarm reaction of Atta leaf-cutting ants

Abstract.  Ethological studies are conducted under laboratory conditions using workers of Atta sexdens sexdens and Atta opaciceps collected from field colonies to investigate the mechanisms involved in the alarm response and intra-specific recognition in leaf-cutting ants. Hexane extracts from the heads of gardeners and generalist workers elicit higher levels of alarm response in foragers from the same colony than do mandibular gland extracts from foragers and soldiers, indicating that gardeners, generalists and foragers are primarily responsible for the production of alarm pheromone. Foragers subjected to extracts from non-nestmates exhibit significantly greater alarm responses than are induced by similar extracts derived from nestmate workers, suggesting that the alarm pheromone may have a role in nestmate recognition.     

The use of alarm pheromones to enhance bait harvest by grass-cutting ants

The enhancement of bait for the control of grass-cutting ants was investigated using two species of grass-cutting ant, Atta bisphaerica (Forel) and Atta capiguara (Gon?alves) (Hymenoptera: Formicidae). Bait was applied in loose piles to obtain a direct relationship between ant attraction and bait harvest. Enhancement with alarm pheromone compounds significantly increased the attractiveness and harvest of bait under certain conditions. A large proportion of the ants attracted to the enhanced bait were minor workers. These ants rarely transport bait because of their small size, and so it may be possible to increase the effect of bait enhancement by using smaller bait granules. Foragers of A. capiguara were less inclined to transport citrus-pulp bait than were those of Atta laevigata (Fr. Smith), a species that also harvests dicotyledonous plants. This emphasizes the importance of developing a bait matrix that is more acceptable to grass-cutting species. Nevertheless, the results suggest that alarm pheromone compounds have significant potential to improve the efficacy of baits for the control of grass-cutting ants.     

Hitchhiking behaviour in leaf-cutter ants: An experimental evaluation of three hypotheses

In leaf-cutter ants, small workers often ride or “hitchhike” on leaf fragments carried back to the nest by larger foragers. There are several possible explanations for this unusual behaviour, the main ones being defence against phorid flies, defence against fungal contaminants, and leaf sap obtention. Here we tested these three hypotheses using standardized paired assays with laboratory colonies of Atta sexdens and field colonies of Atta laevigata. For both ant species, the proportion of fragments with hitchhikers increased significantly in response to the presence of fungal contaminants and to leaf sap. The proportion of fragments with hitchhikers also increased in the presence of phorid flies (Neodohrniphora erthali), but only for A. sexdens. In addition, hitchhiker position varied with the treatment applied. In the presence of phorids, hitchhikers frequently moved around the fragment; in the presence of leaf sap, hitchhikers frequently stood at the edge of the fragments, whereas when fragments were inoculated with fungal contaminants, hitchhikers were more often in the centre of the fragment, where the contamination was greatest. Our results strongly suggest that hitchhiking behaviour in Atta has multiple functions. Such behaviour probably evolved as a mechanism of defence, both against potential contaminants of the symbiotic fungus and against ant parasitoids. The obtention of leaf sap by minims seems to be a secondary and probably opportunistic function of this behaviour. Received 7 November 2005; revised 16 March 2006; accepted 6 April 2006.     

Drought stress drives intraspecific choice of food plants by Atta leaf‐cutting ants

Leaf‐cutting ants (LCA) are polyphagous and dominant herbivores throughout the Neotropics that carefully select plant individuals or plant parts to feed their symbiotic fungus. Although many species‐specific leaf traits have been identified as criteria for the choice of food plants, the factors driving intraspecific herbivory patterns in LCA are less well studied. Herein, we evaluate whether or not drought‐stressed native plants are a preferred food source using free‐living colonies of two leaf‐cutting ants, Atta sexdens L. (Hymenoptera: Formicidae: Attini), in combination with five plant species, Ocotea glomerata Nees (Lauraceae), Lecythis lurida S. A. Mori (Lecythidaceae), Miconia prasina DC (Melastomataceae), Tovomita brevistaminea Engl. (Clusiaceae), and Tapirira guianensis Aubl. (Anacardiaceae), and Atta cephalotes L., in combination with two plant species, O. glomerata and Licania tomentosa Benth. (Chrysobalanaceae). In dual‐choice bioassays, ants removed about three times more leaf area from drought‐stressed plants compared to control plants. Both leaf‐cutting ant species consistently preferred drought‐stressed plants for all species tested, except T. guianensis. The mean acceptability index – expressing the preference for one of two options on a scale of 0 to 1 – of drought‐stressed plants ranged from 0.65 to 0.86 across plant species, and the preference did not differ significantly among the tested plant species. Our results suggest that selection of drought‐stressed individuals is a general feature of food plant choice by leaf‐cutting ants irrespective of ant or plant species. As human‐modified forest assemblages across the Neotropics are increasingly prone to drought stress, the documented preference of Atta for drought‐stressed plants may have tangible ecological implications.     

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.     

Temporal Response of Foragers and Guards of Two Stingless Bee Species to Cephalic Compounds of the Robber Bee <Emphasis Type="Italic">Lestrimelitta niitkib</Emphasis> (Ayala) (Hymenoptera,Apidae)

Lestrimelitta spp. are stingless bees that steal food and nesting materials from other highly social bees to survive. Though most of their victim species respond, either aggressively or submissively, to cephalic components of Lestrimelitta, little is known about if such response changes at some point during extended periods of exposure. Moreover, potential synergistic effects due to a mixture of victim’s alarm/defense pheromones and Lestrimelitta mandibular pheromones, like in an actual attack, have not been examined so far. In this paper, we investigated the response of two species of non-robber stingless bees, Scaptotrigona mexicana (Guérin) and Tetragonisca angustula (Latreille), to (a) cephalic compounds from crushed heads of nestmates, (b) cephalic compounds of Lestrimelitta niitkib (Ayala), and (c) a mixture of (a) and (b). We found that even though T. angustula did not react to nestmates’ crushed head, its response towards L. niitkib cephalic compounds was stronger and lasted longer than that of S. mexicana. Interestingly, the addition of crushed heads of the non-robber species to L. niitkib crushed heads caused no significant increase in the alarm response of both species. It may be that the absence of an alarm pheromone in T. angustula made this species more receptive to extraneous odors, which is not the case for S. mexicana; however, more species must be studied to elucidate any pattern regarding the absence/presence of alarm pheromones and the corresponding response to intruders’ pheromones.     

Intended and unintended receivers of the male pheromones of the burying beetles Nicrophorus humator and Nicrophorus vespilloides

Male Nicrophorus beetles (Coleoptera: Silphidae) attract females through volatiles that are emitted at species‐specific times of day. Not only beetles of the opposite sex but also conspecific males are attracted. Another observation is the co‐attraction of congeners, a phenomenon that was shown in particular for Nicrophorus vespilloides Herbst, the smallest Nicrophorus species in Central Europe. In the current study, we identified the Nicrophorus humator Gleditsch male pheromone as methyl 4‐methyloctanoate through gas chromatography/mass spectrometry analysis. In field experiments, we tested and compared the attractiveness of synthetic analogs of the male pheromones of N. humator and N. vespilloides in baited pitfall traps. An asymmetric cross‐attraction to the synthetic male pheromones was observed, which is best explained by the skewed competitive relationship of the two species, with regard to the restricted availability of breeding resources. Nicrophorus humator is attracted by both its own male pheromone and by the pheromone of the smaller N. vespilloides, whereas N. vespilloides is almost exclusively attracted by its own male pheromone. The observed attraction of conspecific males of either species to male pheromone baits can be explained by both competition for females and competition for breeding resources.     

Endophytic fungi increase the processing rate of leaves by leaf‐cutting ants (Atta)

1. Fungal endophytes are microfungi that reside asymptomatically inside of leaf tissues, increasing in density and diversity through time after leaves flush. Previous studies have suggested that the presence of fungal endophytes in the harvest material of leaf‐cutting ants (Atta colombica, Guérin‐Méneville) may negatively affect the ants and their fungal cultivar. 2. In the present study, it was tested whether the presence and diversity of fungal endophytes affected the amount of time necessary for leaf‐cutter ants to cut, process, and plant leaf material in their fungal garden. It was found that ants took 30–43% longer to cut, carry, clean, and plant leaf tissue with high relative to low endophyte abundance, and that the ants responded similarly to leaf tissue with high or low endophyte diversity. 3. It was further investigated whether the fungal cultivars' colonisation rate was greater on leaf material without fungal endophytes. No difference in the ants' cultivar colonisation rate on leaf tissue with high or low endophyte abundance was observed.     

Antennal olfactory sensitivity in response to task-related odours of three castes of the ant Atta mexicana (hymenoptera: formicidae)

Abstract.  The relationship between scent composition and antennal sensitivity in different castes of Atta mexicana is investigated under laboratory conditions. Extracts of dead ants are analysed by gas chromatography-mass spectrometry to identify the compounds presumably responsible for the specific undertaking behaviour. Oleic acid is identified as one compound that triggers undertaking behaviour. To determine differences in odour reception between workers of different castes (i.e. foragers, undertakers and soldiers), further antennal sensitivity to task-related odours is tested using electroantennographic techniques. Soldiers are the most specialized caste because of their low response to all odours, except odourants related to alarm pheromones. The behavioural specialization of soldiers and their reduced behavioural repertoire are discussed.     

An ambiguous function of an alarm pheromone in the collective displays of the Australian meat ant,Iridomyrmex purpureus

Alarm pheromones, which have been documented in many species of ants, are thought to elicit responses related to aggressive or defensive behaviour. The volatile odour 6-methyl-5-hepten-2-one is described as an alarm pheromone in several species of ants, including the Australian meat ant, Iridomyrmex purpureus. The alarm pheromone is released by displaying workers that aggregate in the characteristic collective display grounds, located mid-way between colonies or near contested food trees. Workers are typically more aggressive at the latter location, and the alarm pheromone may regulate the collective level of aggression. We investigated this possibility by exposing displaying workers to synthesised alarm pheromone 6-methyl-5-hepten-2-one in a field experiment, and measuring their aggressive behaviour. We found no evidence that exposure to synthesised alarm pheromone caused changes in the aggressive level of workers. Subsequent field experiments revealed that the pheromone functions as an attractant, thereby increasing the density of displaying workers. More densely populated workers also display more aggressively, indicating that the interaction rate of displaying workers may determine the level of aggression in collective displays. This underlying mechanism can explain why displaying ants are more aggressive at the more densely populated food-tree locations than those displaying at locations midway between two neighbouring colony nest sites.     

Host-plant selection,diet diversity,and optimal foraging in a tropical leafcutting ant

Summary A month-long study was conducted on the comparative foraging behavior of 20 colonies of the leafcutting ant, Atta cephalotes L. in Santa Rosa National Park, Guanacaste Province, Costa Rica. The study was conducted during the middle of the wet season, when trees had mature foliage and the ants were maximally selective among species of potential host plants. The colonies always gathered leaves from more than a single tree species but on average one species constituted almost half the diet with the remaining species being of geometrically decreasing importance. Colonies exhibited greater diversity in their choice of leaves and lower constancy of foraging when the average quality of resource trees was lower, as predicted by elementary optimal foraging theory. Furthermore, the ants were more selective of the species they attacked at greater distances from the nest. However, the ants sometimes did not attack apparently palatable species, and often did not attack nearby individuals of species they were exploiting at greater distances.A classical explanation for why leafcutting ants exploit distant host trees when apparently equally good trees are nearer, is that the ants are pursuing a strategy of conserving resources to avoid long-term overgrazing pressure on nearby trees. We prefer a simpler hypothesis: (1) Trees of exploited species exhibit individual variation in the acceptability of their leaves to the ants. (2) The abundance of a species will generally increase with area and radial distance from the nest, so the probability that at least one tree of the species will be acceptable to the ants also increases with distance. (3) The ants forage using a system of trunk-trails cleared of leaf litter, which significantly reduces their travel time to previously discovered, high-quality resource trees (by a factor of 4- to 10-fold). (4) Foragers are unware of the total pool of resources available to the colony. Therefore once scouts have chanced upon a tree which is acceptable, the colony will concentrate on harvesting from that tree rather than searching for additional sources of leaves distant from the established trail.     

Influence of leaf epicuticular waxes from cerrado species on substrate selection byAtta sexdens rubropilosa

The influence of leaf epicuticular waxes of nine woody species from cerrado (a savanna ecosystem of Brazil) on the foraging behavior of the leaf-cutting antAtta sexdens rubropilosa Forel, 1908 (Formicidae, Attini) was investigated. Some waxes showed neither inhibitory nor attractive effect over a series of tests, while a second group caused a delayed rejection. Waxes of two species were shown to be strongly deterrent to the ants. Since different waxes caused different responses, it is concluded that their chemical composition, rather than hydrophobicity or mechanical constraints, accounts for the observed effects. Plant species with leaves rejected by the ants did not necessarily have repellent waxes. On the other hand, no species with acceptable leaves yielded waxes with deterrent effects.     

An Attraction Pheromone from Heads of Worker Vespula germanica Wasps

Vespula germanica is a social wasp that efficiently exploits food resources. Odor cues, derived from substances located in wasp heads, have been proved to be central in conspecific attraction in this species. However, it remains unknown whether this attraction is related to foraging or defense responses. In this study we analyze conspecific attraction under two different contexts: at the nest entrance (defense) and under foraging conditions. We also test wasp response with two dosages of head extract and crosschecked the attractiveness of extracts obtained from different populations. We found no evidence of alarm response to head extracts either at the nest entrance or under foraging conditions. Moreover, no differences in attractiveness were found to both doses tested. Head extracts attract similarly in the same or a different population suggesting that conspecific attraction is not restricted to colony nestmates.     

Parallel foraging cycles for different resources in leaf‐cutting ants: a clue to the mechanisms of rhythmic activity

1. Leaf‐cutting ants display regular diel cycles of foraging, but the regulatory mechanisms underlying these cycles are not well known. There are, however, some indications in the literature that accumulation of leaf tissue inside a nest dampens recruitment of foragers, thereby providing a negative feedback that can lead to periodic foraging. We investigated two foraging cycles occurring simultaneously in an Atta colombica colony, one involving leaf harvesting and the other exploiting an ephemeral crop of ripe fruit. 2. Leaf harvesting followed a typical diel pattern of a 10–12 h foraging bout followed by a period of inactivity, while fruit harvesting occurred continuously, but with a regular pre‐dawn dip in activity that marked a 24 h cycle. 3. Although the results of the present study are drawn from a single field colony, the difference found is consistent with a mechanism of negative feedback regulation acting in parallel on two resources that differ in their rates of distribution and processing, creating cycles of formation and depletion of material caches. 4. This hypothesis should provoke further interest from students of ant behaviour and some simple manipulative experiments that would begin to test it are outlined. Any role of resource caches in regulating foraging by Atta colonies may have similarities to the logistics of warehouse inventories in human economic activity.     

A sophisticated,modular communication contributes to ecological dominance in the invasive ant <Emphasis Type="Italic">Anoplolepis gracilipes</Emphasis>

The ecological success of ants is founded on cooperative behaviour and a well functioning communication. Particularly invasive ants are able to act highly cooperatively, out-compete other species, and become ecologically dominant. Since ant communication is to a large extent chemical, we investigated the pheromone functions involved in foraging and alarm behaviour of the invasive tropical formicine Anoplolepis gracilipes. Our results suggest that long-lasting orientation cues are located in hindguts, while Dufour glands contain short-term attractants that trigger an effective recruitment. Poison gland effects were intermediate between hindgut and Dufour gland in terms of orientation, attraction and longevity. In contrast to the other pheromone sources, mandibular glands have a repellent effect and are most likely involved in alarm behaviour. Taken together, the pheromone glands of A. gracilipes contain functionally distinct signals with considerable differences in persistence. In this respect, its communication is exceptional in formicine ants. A strikingly similar communication system was previously detected in Paratrechina longicornis, another opportunistic and invasive formicine ant. Based on these similarities and the differences compared to non-invasive formicine ants, we discuss the role of chemical signals for the coordination of efficient foraging. We conclude that a sophisticated communication system can contribute significantly to ecological dominance and invasive success, in concert with other well known traits.     

Social Insect Pheromones: Their Chemistry and Function

Exocrine secretions of social insects are often characterizedby extraordinarily complex mixtures of natural products. Thus,chemical communication in social insects must be interpretedin terms of signals generated by multicomponent systems, theindividual constituents of which can affect the informationalcontent of the message. Alarm pheromones have been identified chiefly in three subfamiliesof ants and their distribution appears to be chemosystematicallysignificant. Myrmicine genera emphasize 3-alkanones as alarmreleasers, whereas methyl ketones, primarily of terpenoidalorigin, are widely utilized as alarm pheromones in the subfamilyDolichoderinae. Formicine species may employ formic acidas analarm pheromone in addition to the compounds produced in themandibular and Dufour's glands. The mandibular gland pheromonesare chiefly acyclic monoterpene aldehydes (e.g., citronellal)which are relatively low boiling compounds. Higher boiling n-alkanesare produced in the Dufour's glands and may serve as more persistentreleasers of alarm behavior. Alarm pheromones as well as thecaste-specific pheromones of male bees and ants, probably alsoserve as defensive products. In many cases it is likely thatpheromones were originally utilized as defensive compounds andtheir communicative function is a secondary development.     

Alarm communication in <Emphasis Type="Italic">Ropalidia</Emphasis> social wasps

Summary Alarm pheromones, chemical substances produced by social insects to alert the colony to threat, are the principal means by which colony defence is co-ordinated. We present the results of a study on alarm behaviour in 5 swarming species of wasps belonging to the genus Ropalidia. These species show a remarkably efficient strategy of alarm communication, including visual display and attack synchronization. We show that pheromones released from the venom gland play an important role in alarm recruitment in species belonging to the Ropalidia flavopicta group, but not in Ropalidia sumatrae. We analysed the contents of the venom reservoirs content of four of the studied species by gas chromatography-mass spectrometry. Glands were found to contain a complex mixture of volatile compounds as well as spiroacetals of higher molecular weight. Interestingly, despite all species producing similar chemical compounds from the venom gland, these were found to elicit alarm behaviour in only those species that build nest envelopes, suggesting a link between chemical release of alarm behaviour and the evolution of nest architecture in Ropalidia wasps.Received 19 August 2003; revised 29 February 2004; accepted 10 March 2004     

Polymorphism: a weak influence on worker aggregation level in ants

Abstract.  1. Aggregation of individuals, a basic behaviour in social species, plays an essential role in many aspects of animal life (reproduction, defence, and alimentation). Understanding how this phenomenon is modulated is important to comprehend the social organisation of the group.
2. In social insects, aggregation is influenced by environmental (e.g. the light level) and social (e.g. polyethism in monomorphic ants) factors. Ants display a great variation of biological characteristics (e.g. queen number, polymorphism, division of labour, etc.) that are likely to influence the level of inter-attraction and so the aggregation.
3. The present research focused on one biological characteristic: the morphological castes (minors, majors), testing the hypothesis that minors will aggregate more than majors due to their greater need to fight against the loss of heat and to increase their self-protection.
4. Aggregation experiments were conducted on two highly polymorphic species, Atta sexdens rubropilosa and Solenopsis interrupta , using the two extreme morphological castes (majors and minors).
5. All castes exhibited a low level of aggregation: 40–50% of workers assembled for both species, the biggest cluster involving 20% of the total population. The lack of difference between morphological castes in the aggregation shows the weak influence of polymorphism on the interactions between ants.
6. It is concluded that the main factor modulating the aggregation behaviour is polyethism, i.e. the division of labour associated with the presence of an outside-the-nest experience: workers that only take care of the brood, without outside world experience (brood-tenders) assembling more than foraging workers (foragers).