Leaf‐cutting ants use relative humidity and temperature but not CO2 levels as cues for the selection of an underground dumpsite

1. Leaf‐cutting ants remove copious amounts of colony waste, a potential pathogen source for workers and reared symbiotic fungus, to above‐ground heaps or deep underground chambers. However, the dumpsite may also contain information about plants initially harvested and disposed of because of unsuitability for the fungus. 2. The underground environment presents climatic gradients across the soil profile and it is an open question whether leaf‐cutting ants use microclimatic cues to choose suitable sites for waste disposal, as displayed for other in‐nest tasks. 3. Climatic preferences in leaf‐cutting ants were investigated for the deposition of colony waste. In the laboratory, deposition of waste particles by workers of Atta laevigata was quantified by offering them, in different experiments, a binary choice of temperatures (range, 15–30 °C), levels of air humidity (range 10–98%), and CO₂ concentrations (range, atmospheric values to 10%). 4. Leaf‐cutting ants used temperature and air humidity, but not CO₂ levels, as cues for the deposition of their waste. They consistently preferred a dry (≤ 33% air humidity) environment. Less consistent, temperature preferences varied depending on colony (15–25 °C for one colony and 25–30 °C for the other). Although workers showed clear preferences for high levels of CO₂ for themselves, they were CO₂‐indifferent for waste deposition. 5. It is argued that the observed climatic preferences for underground waste disposal might aid nest hygiene by providing unsuitable dry conditions for pathogen growth, with thermal preferences that do not hinder worker activities for further waste management and inspection of discarded plants.     

Simple rules based on pile slope are used in the self organization of sand pile formation by <Emphasis Type="Italic">Pheidole oxyops</Emphasis> ants

We tested the hypothesis that slope influences where worker ants deposit excavated soil on piles near the nest entrance. We predicted that ants will deposit their load near the top of a pile where the slope changes from upward to downward, to prevent material rolling back towards the entrance. We tested this hypothesis by studying five natural colonies of Pheidole oxyops ants at a field site at S?o Sim?o, Brazil. At this site, each colony was dumping sandy soil excavated from its underground nest in a crescent-shaped pile c. 13 cm from and perpendicular to the nest entrance. Each nest was given an experimental sand pile of symmetrical curved cross section on a plywood platform that could be tilted 15 degrees up or down. From videos, the locations where individual ants dumped their soil loads were measured in relation to the inner (position = 0) and outer (position = 1) edges of the pile. When the platform was tilted down the ants deposited their loads significantly closer to the inner edge (0.458 ± 0.007) than when not tilted (0.530 ± 0.006). When the platform was tilted up the ants deposited their loads significantly further from the inner edge (0.626 ± 0.006) than when not tilted (0.522 ± 0.006). These results support the hypothesis that ants use pile slope in deciding where to dump their load. A similar rule is probably used in other ant species that place excavated soil in steep piles near the nest entrance. Received 5 February 2007; revised 10 June 2007; accepted 9 October 2007.     

Waste of leaf-cutting ants: disposal,nest structure,and abiotic soil factors around internal waste chambers

Leaf-cutting ants produce large quantities of waste that harbor bacteria and fungi that are harmful to the colony. To be protected from these pathogens, the workers of Atta species present a sophisticated organization to manage harmful material, which can be deposited outside the nest or in internal chambers. However, little is known about the behavior of Acromyrmex species in handling and disposal of waste. Due to some observations, we assume that the same species of Acromyrmex can deposit waste outside the nest and into internal chambers and raise the following question: what determines the occurrence of internal waste chambers in Acromyrmex? To address this question, we verified whether nest depth influences the waste-chamber occurrence. We also verified the nest structure and the abiotic factors of soil beside each waste-chamber: pH and water content of the soil. For this, eight nests were excavated for Acromyrmex balzani and Acromyrmex rugosus rugosus. We verified that not only can the same leaf-cutting ant species deposit debris both outside and inside the nest but also the same nest can present internal chambers and external waste deposit. The soil beside the waste chamber always presented an acidic pH, while the humidity varied widely. Our results showed that the nest depth was highly correlated with the depth of the waste chamber (p = 0.0003) and probably has some influence on waste disposal. The characteristics of the nest and the role of depth in the choice of waste chamber location are discussed.     

Nest‐site selection by golden plover: why do shorebirds avoid nesting on slopes?

It is widely known that many upland-breeding shorebirds tend to nest on plateaus but to date no studies have put forward explanations for this phenomenon. We examined the effect of slope and habitat on the distribution of ground-nesting golden plover Pluvialis apricaria at two study sites in County Durham, U.K. Golden plovers showed strong selection for nesting on flat ground. Habitat significantly affected nest-site location on one study site (heather burnt within the past 2 years was favoured and older stands of heather were avoided) but not on the other. Fifty-nine per cent of all nests failed. We attributed 95% of all losses to predation. Seventy-five per cent of nests, in which the predator was identified, were taken by ground predators, mainly stoats Mustela erminea . Nests on flat ground had significantly higher rates of survival than those on slopes. Nest survival did not vary significantly with habitat type nor with vegetation height or density around the nest. Neither habitat type, vegetation height nor vegetation density around nests differed between nests on slopes and on flat ground. In addition, individuals that nested on flat ground tended to have less black on their underparts (a suggested indicator of dominance). We suggest that birds nesting on slopes are less efficient at avoiding nest predation than those nesting on the flat. The nature or degree of a bird's response to a predator may be related to the efficiency of individual anti-predator responses and/or to visibility from the nest. This study cannot differentiate between these two explanations. We encourage further work to investigate differences in behaviour between individuals nesting on slopes and those on flat ground.     

Solar Elevation Triggers Foraging Activity in a Thermophilic Ant

Nycthemeral rhythm is an important biological trait that allows animals to escape predation and competition and, conversely, to coincide with mutualists. Although laboratory studies have shown that the rhythm depends on both endogenous factors and cyclic environmental cues, the latter is often poorly understood, particularly in the wild. Because insects are mostly ectothermal organisms, their activity rhythm is often thought to depend directly on ground temperature. In Mediterranean habitats, Cataglyphis ants are well known for their unusual thermoresistance, allowing them to forage in summer at the central hours of the day when the ground reaches temperatures that are lethal to their competitors. However, we show that the rhythm of Cataglyphis floricola in south‐western Spain is governed by light cues rather than by temperature. First, variations in ant traffic at the nest entrance were better explained by solar elevation angle than by ground temperature on both seasonal and daily scales. Second, if ants waited for the ground to reach a threshold temperature to start their activity, we would expect similar temperatures regardless of the opening hour. However, we found a significant increase in ground temperature as opening hour got later in the day. Third, by using a simple experimental set‐up that increased the apparent solar elevation over the nest entrance, we provoked a delay of nest closure time. We discuss the relevance of these results with respect to the life history of Cataglyphis species and their possible consequences in relation to global warming.     

Do ants make direct comparisons?

Many individual decisions are informed by direct comparison of the alternatives. In collective decisions, however, only certain group members may have the opportunity to compare options. Emigrating ant colonies (Temnothorax albipennis) show sophisticated nest-site choice, selecting superior sites even when they are nine times further away than the alternative. How do they do this? We used radio-frequency identification-tagged ants to monitor individual behaviour. Here we show for the first time that switching between nests during the decision process can influence nest choice without requiring direct comparison of nests. Ants finding the poor nest were likely to switch and find the good nest, whereas ants finding the good nest were more likely to stay committed to that nest. When ants switched quickly between the two nests, colonies chose the good nest. Switching by ants that had the opportunity to compare nests had little effect on nest choice. We suggest a new mechanism of collective nest choice: individuals respond to nest quality by the decision either to commit or to seek alternatives. Previously proposed mechanisms, recruitment latency and nest comparison, can be explained as side effects of this simple rule. Colony-level comparison and choice can emerge, without direct comparison by individuals.     

Central place foraging by an ant (Cataglyphis bicolor Fab.): a model of searching

A computer model of the searching activity of Cataglyphis bicolor (Fab.), based on quantitative data obtained in the field in Greece, was investigated. This ant searches for individual items of food (mainly dead arthropods) in the open on flat ground, on which its paths can be followed and mapped. The model was based on an analogy with molecular diffusion from a centre, suggested by the fact that the average distance of ants from the nest increased in proportion to the square root of time, up to a distance more than half the radius of a circle enclosing 90% of the search area. Actual behaviour was well described by a model in which individual ants went out for the first time to search in a direction determined at random. Subsequently they went out in a straight line in a direction and to a distance related to that of the previous find before commencing a semi-random search, on a path consisting of straight portions branching at random angles, with restriction on turns in a backward direction. Individual real ants search in restricted areas, commonly sectors centred on the nest. This behaviour was reproduced in the model as were quantitative aspects of the nest behaviour. Communication between individuals was not required in the model.     

Experiments with artificial nests provide evidence for ant community stratification and nest site limitation in a tropical forest

Ants are dominant in tropical forests and many species nest in hollow cavities. The manner in which species are vertically stratified in these complex habitats is not known, with lack of nest sites being proposed to limit ant populations. Here, we assess ant community stratification and nest site limitation in a lowland rainforest in New Guinea using experimental addition of artificial bamboo nests of two cavity sizes (small: ~12 mm large: ~32 mm diameter) placed at ground level, in the understorey, and in the canopy. We also conducted a pilot experiment to test the utility of nest translocation. Nests were checked for occupancy after 10 weeks and half of the occupied nests were then translocated between forest plots, while keeping same vertical position. Occupancy of small nests was much higher in the understorey and canopy than at ground level (~75% vs. ~25%). Translocation was successful, as a majority of nests was inhabited by the same species before and after translocation and there was no impact of translocation to a different plot compared to the control, except for a reduction in colony size at ground level. Our experiment demonstrates a vertical stratification in community composition of ants nesting in hollow dead cavities and shows that these ants are more nest site limited in the higher strata than at ground level. Use of small artificial cavities has great potential for future experimental studies, especially for those focused on arboreal ants, as occupancy is high and translocation does not negatively affect their colony size. Abstract in Tok Pisin is available with online material.     

Nest Etiquette—Where Ants Go When Nature Calls

Sanitary behaviour is an important, but seldom studied, aspect of social living. Social insects have developed several strategies for dealing with waste and faecal matter, including dumping waste outside the nest and forming specialised waste-storage chambers. In some cases waste material and faeces are put to use, either as a construction material or as a long-lasting signal, suggesting that faeces and waste may not always be dangerous. Here we examine a previously undescribed behaviour in ants – the formation of well-defined faecal patches. Lasius niger ants were housed in plaster nests and provided with coloured sucrose solution. After two months, 1–4 well defined dark patches, the colour of the sucrose solution, formed within each of the plaster nests. These patches never contained other waste material such as uneaten food items, or nestmate corpses. Such waste was collected in waste piles outside the nest. The coloured patches were thus distinct from previously described ‘kitchen middens’ in ants, and are best described as ‘toilets’. Why faeces is not removed with other waste materials is unclear. The presence of the toilets inside the nest suggests that they may not be an important source of pathogens, and may have a beneficial role.     

Distance estimation in the third dimension in desert ants

Desert ants of the genus Cataglyphis perform large-scale foraging excursions from which they return to their nest by path integration. They do so by integrating courses steered and the distances travelled into a continually updated home vector. While it is known that the angular orientation is based on skylight cues, it still is largely enigmatic how the ants measure distances travelled. We extended the ants' task into the third dimension by training them to walk within an array of uphill and downhill channels, and later testing them on flat terrain, or vice versa. In these tests the ants indicated homing distances that did not correspond to the distances actually travelled, but to the ground distances; that is, to the sum of the horizontal projections of the uphill and downhill segments of the ants' paths. These results suggest a much more sophisticated mechanism of distance estimation than hitherto thought. The ants must be able to measure the slopes of undulating terrain and to integrate this information into their "odometer" for the distance estimation process.     

Waste Management in the Leaf-Cutting Ant <Emphasis Type="Italic">Acromyrmex lobicornis</Emphasis>: Division of Labour,Aggressive Behaviour,and Location of External Refuse Dumps

In leaf-cutting ants, the handling of waste materials from the fungus culture increases the risk of infection. Consequently, ants should manage their waste in a way that minimizes the spread of diseases. We investigated whether in Acromyrmex lobicornis, waste-worker ants (a) also perform roles in foraging or mound maintenance, (b) are morphologically different than other ant workers, and (c) are aggressively discriminated by other worker ants from the same colony. In addition, we investigated whether the location of external waste piles minimizes the probability that wastes spread to the ant nest. In the field, we (a) marked with different colours waste-workers, foragers and mound-workers and monitored whether these ants interchanged their tasks; (b) measured head width, head length, hind femur length and total length of waste-workers; foragers and mound-workers; (c) forced field encounters between waste-workers and foragers, and (d) measured the cardinal orientation of the waste piles in relation to the colony mound. Waste-worker ants did not perform other function outside the nest; neither foragers nor mound-workers managed the waste. Moreover, waste-workers were smaller than foragers and mound-workers, and were attacked if they tried to enter their nest using foraging entrances. The location of external refuse dumps also appears to reduce contamination risks. Waste piles always were down-slope, and often followed the prevailing wind direction. The importance of behaviours such as the division of labour, aggressions against waste-workers and nest compartmentalization (i.e., the orientation of external waste piles) to minimize the spread of pathogens is discussed.     

Waste management in the leaf-cutting ant Atta colombica

Unlike most leaf-cutting ants, which have underground wastedumps, the leaf-cutting ant Atta colombica dumps waste in aheap outside the nest. Waste is hazardous, as it is contaminatedwith pathogens. We investigated the organization of the workforceinvolved in outside-nest tasks (foraging, waste disposal) andquantified task switching and heap location to test hypothesesthat these tasks are organized to minimize contact between the heap and foraging entrances and trails. Waste management isan important task: 11% of externally working ants were eithertransporting waste or manipulating waste on the heap, and theother 89% were foragers. There is strict division of laborbetween foragers and waste workers, with no task switching.Waste management also has division of labor and is undertakenby transporters that carry waste to the heap margins and heapworkers that manage the heap. Waste heaps are always locateddownhill from nest entrances. The distance to the waste heapis positively related to colony size and negatively relatedto slope. Foraging trails avoid the heap, with 92% of trailsgoing away from the heap. This avoidance behavior is costly,increasing foraging trail length by at least 6%. Waste managementin A. colombica is a sophisticated system that encompassesboth work and spatial organization. This organization is probablyadaptive in reducing disease transmission. Division of labor separates waste management from foraging, reducing the likelihoodof foragers becoming contaminated with waste. The downhilllocation of heaps reduces waste entering entrances during rain.The orientation of foraging trails reduces the possibilityof foragers becoming accidentally contaminated with waste.     

Marking as a decision factor in the choice of a new resting site in <Emphasis Type="Italic">Lasius niger</Emphasis>

Summary In ants, aggregation is mainly based on the attraction between individuals. We confirmed the existence of ground marking in Lasius niger and demonstrated its influence on the aggregative behaviour of ants: the place of the gathering is leading by the existence of a ground marking which favours clustering by increasing the rate of the process but does not change the aggregation level. This factor could play a role in the choice of a new nest site and in the spatial distribution of ants inside the nest. This marking, presuming to have a long duration, could represent an external memory which could play an important role in various situations such as the choice of new nest or the reorganisation of ants after a disturbance.Received 23 July 2003; revised 16 December 2003; accepted 19 December 2003.     

A reassessment of the relations in Malaysia between ants (Crematogaster) on trees (Leptospermum and Dacrydium) and epiphytes of the genus Dischidia (Asclepiadaceae) including 'ant-plants'

Three species of epiphytic Dischidia have been investigated in terms of their relationship to ants on trees. Two species, D. parvifolia and D. astephana , are associated with ants and trees in montane areas. A clear association has been found between ants of the genus Crematogaster and the tree Leptospermum flavescens. This relationship is complex and probably both organisms benefit from the association. The ants live in tunnels in the wood of the major branches and the trunk, and the entire tree is occupied by one ant colony. Trees occupied by ants are maintained by the ants substantially clear of epiphytes other than the two species of Dischidia. The potential benefits to the tree and to the ants of this association are noted. The roots of D. astephana and D. parvifolia penetrate into the cavities of these ant nests and presumably gain nutrients from waste in the ant nests. Both Dischidia species are effectively scavenging upon the waste material from the ant-tree association. The leathery dome-shaped leaves of D. astephana are not vital to the development of the scavenging habit as D. parvifolia has lens-shaped leaves, but may offer some advantage to D. astephana by the uptake of nutrients from waste deposited by the ants under the dome-shaped leaves by interception of stem flow and by uptake of gaseous waste. Ants do not nest under these leaves. Seeds of these species of Dischidia are taken by ants into the central woody area of the ant nest where they germinate. Both Leptospermum and Dischidia can be visualized as showing adaptations to a nutient-deficient tropical montane environment. These adaptations are discussed as is the need for reassessment in this genus of the term 'ant-plant', and the need for wider recognition of the 1ant-tree' relationship between Crematogaster and Leptospermum.     

Path integration controls nest-plume following in desert ants

The desert ant Cataglyphis fortis is equipped with sophisticated navigational skills for returning to its nest after foraging. The ant's primary means for long-distance navigation is path integration, which provides a continuous readout of the ant's approximate distance and direction from the nest. The nest is pinpointed with the aid of visual and olfactory landmarks. Similar landmark cues help ants locate familiar food sites. Ants on their outward trip will position themselves so that they can move upwind using odor cues to find food. Here we show that homing ants also move upwind along nest-derived odor plumes to approach their nest. The ants only respond to odor plumes if the state of their path integrator tells them that they are near the nest. This influence of path integration is important because we could experimentally provoke ants to follow odor plumes from a foreign, conspecific nest and enter that nest. We identified CO(2) as one nest-plume component that can by itself induce plume following in homing ants. Taken together, the results suggest that path-integration information enables ants to avoid entering the wrong nest, where they would inevitably be killed by resident ants.     

Sand Pile Formation in <Emphasis Type="Italic">Dorymyrmex</Emphasis> Ants

We studied circular sand pile formation by two colonies of Brazilian Dorymyrmex ants, in which workers dumped sand excavated from their underground nest around the nest entrance hole. In most cases a worker dumped its load just beyond the ridge of the pile. Each dumped piece either stayed where it was deposited (81.9% in colony A and 73.0% in colony B) or rolled down the outer slope of the sand pile away from the entrance (17.9% in colony A and 27.0% in colony B). Ants almost never dumped in a way that resulted in the load rolling back to the entrance. When one side of the sand pile was experimentally removed, ants preferentially dumped soil on the now flat side, thereby restoring the original circular shape.     

Ants estimate area using Buffon's needle

We show for the first time, to our knowledge, that ants can measure the size of potential nest sites. Nest size assessment is by individual scouts. Such scouts always make more than one visit to a potential nest before initiating an emigration of their nest mates and they deploy individual-specific trails within the potential new nest on their first visit. We test three alternative hypotheses for the way in which scouts might measure nests. Experiments indicated that individual scouts use the intersection frequency between their own paths to assess nest areas. These results are consistent with ants using a 'Buffon's needle algorithm' to assess nest areas.     

Ant Colonies Prefer Infected over Uninfected Nest Sites

During colony relocation, the selection of a new nest involves exploration and assessment of potential sites followed by colony movement on the basis of a collective decision making process. Hygiene and pathogen load of the potential nest sites are factors worker scouts might evaluate, given the high risk of epidemics in group-living animals. Choosing nest sites free of pathogens is hypothesized to be highly efficient in invasive ants as each of their introduced populations is often an open network of nests exchanging individuals (unicolonial) with frequent relocation into new nest sites and low genetic diversity, likely making these species particularly vulnerable to parasites and diseases. We investigated the nest site preference of the invasive pharaoh ant, Monomorium pharaonis, through binary choice tests between three nest types: nests containing dead nestmates overgrown with sporulating mycelium of the entomopathogenic fungus Metarhizium brunneum (infected nests), nests containing nestmates killed by freezing (uninfected nests), and empty nests. In contrast to the expectation pharaoh ant colonies preferentially (84%) moved into the infected nest when presented with the choice of an infected and an uninfected nest. The ants had an intermediate preference for empty nests. Pharaoh ants display an overall preference for infected nests during colony relocation. While we cannot rule out that the ants are actually manipulated by the pathogen, we propose that this preference might be an adaptive strategy by the host to “immunize” the colony against future exposure to the same pathogenic fungus.     

Les relations entre fourmis arboricoles et guêpes sociales sous les tropiques

Due to the brood that it contains, the nest of social wasps constitutes a major source of protein for eventual predators. In the tropics, ants are generally considered as the most efficacious among them. In the face of such predation, wasps have developed over the course of evolution a great number of strategies. In addition to guarding and defensive behaviours displayed by workers, social wasps have nests whose structure is a response to the predation pressure exerted by ants. Thus, the presence of a petiole attaching the nest to a natural support itself sometimes selected because of its limited accessibility constitutes a frequent example of architectural defence. The effectiveness of the petiole is often improved through the repeated application of repellent substances on its surface. In the same way, wasps that found their new societies through swarming construct nests protected by an envelope. In the American tropics, army ants which hunt on the ground and can rapidly take over a tree, regularly pillage wasp nests. Because no defensive strategy is efficacious against such predators, the choice of nest site becomes decisive. Paradoxically, it is by nesting in close proximity to an arboreal ant nest that certain wasp species have been able to find a way of responding to the threat exerted by other ants. These ants, which tolerate the presence of the wasp nest, are capable of efficaciously protecting the tree or the branch sheltering the nest.     

Ant behaviour and seed morphology: a missing link of myrmecochory

Seed dispersal by ants (myrmecochory) is mediated by the presence of a lipid-rich appendage (elaiosome) on the seed that induces a variety of ants to collect the diaspores. When seeds mature or fall onto the ground, these ant species transport them to their nest. After eating the elaiosome, the seed is discarded in nest galleries or outside, in the midden or farther away, where seeds can potentially germinate. The final location of seeds with their elaiosomes removed was evaluated to assess the importance of possible handles (structures that ants can grasp to carry) in transporting ants during re-dispersal experiments of seeds from nests of six species of ants. The results indicate that seeds remained within the nest because the ants were not able to transport them out of the nest. As a consequence of the elaiosome being removed, small ant species could not take Euphorbia characias seeds out of their nests. Only large ant species could remove E. characias seeds from their nests. Attaching an artificial handle to E. characias seeds allowed small ant species to redistribute the seeds from their nests. On the other hand, Rhamnus alaternus seeds that have a natural handle after the elaiosome removal were removed from the nests by both groups of ant species. If a seed has an element that acts as a handle, it will eventually get taken out of the nest. The ants’ size and their mandible gap can determine the outcome of the interaction (i.e. the pattern of the final seed shadow) and as a consequence, could influence the events that take place after the dispersal process.