The capacity of ants for multiple alterations in the maze habit]

Ability was shown of ants Myrmica rubra to multiple reconstructions of the habit elaborated in symmetrical multialternative maze under motivation of care for the progeny (transportation of breed of the own species). Reconstruction consisted in the change of reinforcement location on the left or right aim spot. The ants showed the ability to carry out the series of eight reconstructions during one-two days. An improvement took place of the fulfillment of the last reconstructions in comparison with the first ones. Peculiarities of learning and reconstructions were found in two groups of animals differing by conditions of learning: at reinforcement on both ain spors or on one of them. The results obtained are considered as indices of high plasticity of the behaviour of ants of the studied species.     

Sucking pump activity in feeding behaviour regulation in carpenter ants

Modulation of liquid feeding-rate would allow insects to ingest more food in the same time when this was required. Ants can vary nectar intake rate by increasing sucking pump frequency according to colony requirements. We analysed electrical signals generated by sucking pump activity of ants during drinking solutions of different sucrose concentrations and under different carbohydrate-deprivation levels. Our aim was to define parameters that characterize the recordings and analyse their relationship with feeding behaviour.Signals showed that the initial and final frequencies of sucking pump activity, as well as the difference between them were higher in sugar-deprived ants. However, these parameters were not influenced by sucrose solution concentration, which affected the number of pump contractions and the volume per contraction. Unexpectedly, we found two different responses in feeding behaviour of starved and non-starved ants depending on concentration. Starved ants drank dilute solutions for the same length of time as non-starved ants but ingested higher volumes. While drinking the concentrated solutions, starved ants drank the same volume, but did so in a shorter time than the non-starved ones. Despite these differences, for each analysed concentration the total number of pump contractions remained constant independently of sugar-deprivation level. These results are discussed in the frame of feeding regulation and decision making in ant foraging behaviour.     

Prey-capture techniques and prey preferences of Chrysilla, Natta and Siler, ant-eating jumping spiders (Araneae, Salticidae) from Kenya and Sri Lanka

The predatory behaviour of Chrysilla lauta and Sliersemiglaucus from Sri Lanka, and four species of Natta from Kenya, was studied in the laboratory for the first time. These salticids eat ants, a prey most salticids avoid. These species' specialized behaviour for catching ants is described and compared to their behaviour for catching other insects. Three different types of tests of prey preference were carried out and, in each type, ants were taken in preference to other insects. Preference for ants, and prey-specific predatory behaviour did not depend on prior experience with ants. Results from this study are discussed in relation to recent findings on other ant-eating salticids.     

An empirical test of Lanchester's square law: mortality during battles of the fire ant Solenopsis invicta

Lanchester's models of attrition describe casualty rates during battles between groups as functions of the numbers of individuals and their fighting abilities. Originally developed to describe human warfare, Lanchester's square law has been hypothesized to apply broadly to social animals as well, with important consequences for their aggressive behaviour and social structure. According to the square law, the fighting ability of a group is proportional to the square of the number of individuals, but rises only linearly with fighting ability of individuals within the group. By analyzing mortality rates of fire ants (Solenopsis invicta) fighting in different numerical ratios, we provide the first quantitative test of Lanchester's model for a non-human animal. Casualty rates of fire ants were not consistent with the square law; instead, group fighting ability was an approximately linear function of group size. This implies that the relative numbers of casualties incurred by two fighting groups are not strongly affected by relative group sizes and that battles do not disproportionately favour group size over individual prowess.     

Three-dimensional orientation in desert ants: context-independent memorisation and recall of sloped path segments

Desert ants (Cataglyphis fortis) navigate by a combination of path integration and landmark-based route memories. Their ability to correct sloped path segments to their ground distances enables them to orientate accurately even in undulating terrain. In this study, we tested whether or not ants incorporate vertical components of an itinerary into their route memory in similar ways as they do with visual landmarks and horizontal changes of direction. In two separate experiments, we trained desert ants to walk over artificial hills and later tested their acceptance of slopes within novel contexts. In the first paradigm, ants had to traverse a hill only on their outbound run, but not on their homebound trip. In a follow-up experiment, we confronted ramp-trained animals with descents in a completely new temporal and spatial context. The animals transferred their newly acquired acceptance of slopes from the outbound to the homebound run as well as to novel foraging trips. Cataglyphis obviously dissociates the experience of sloped path segments from the original context in which they appeared, thus reducing their significance as a navigational aid.     

Prey-capture techniques and prey preferences of Corythalia canosa and Pystira orbiculata, ant-eating jumping spiders (Araneae, Salticidae)

Corythalia canosa from Florida is an unusual salticid because it is known to eat ants. This species' specialized behaviour for catching ants is described in detail for the first time and compared to its behaviour for catching other insects. Pystira orbiculata from Queensland is shown to be another ant-eating salticid, although its behaviour for catching ants seems less specialized than that of C. canosa. Three different types of tests of prey preference were carried out. In each type of test C. canosa and P. orbiculata took ants in preference to other insects. Another species of salticid, Trite planiceps from New Zealand, failed to eat ants in these tests, although T. planiceps often attacked then released the ants. Corythalia canosa's and P. orbiculata's preference for ants, and their prey-specific predatory behaviour for catching ants, are shown not to depend on prior experience with ants.     

Quarantining behaviour in ants: are Myrmica aphid milkers able to detect and get rid of fungus-contaminated aphids?

Fungal infections are highly dangerous for social insects including ants. Close trophobiotic interaction between ants and aphids promotes infection transmission, as aphids can be a disease vector. The ability of ants to detect fungus-infected aphids and get rid of them is important to the prosperity of both symbiotic partners. However, the diversity of quarantining behaviour among ants is still poorly studied. Here, the behaviour of honeydew foragers of two ant species – Myrmica rubra L. and Myrmica scabrinodis Nylander (Hymenoptera: Formicidae, Myrmicinae, Myrmicini) – was studied in laboratory towards Schizaphis graminum (Rondani) (Hemiptera: Aphididae, Aphidini) aphids contaminated with the generalist fungal pathogen Beauveria bassiana (Balsamo-Crivelli) Vuillemin (Hypocreales). This fungus attacks a wide range of hosts including aphids and ants. The removal of conidia-contaminated aphids from the host plant was found not to be typical of the ants studied. Aphid milkers of M. rubra and M. scabrinodis usually displayed non-aggressive behaviour (tolerance, antennation, honeydew collection, grooming) towards the experimental aphids regardless of whether they were covered with conidia or not. Neither ant species, nor the number of milkers had significant effects on their behaviour towards ‘infected’ aphids. However, some individuals were found to demonstrate quarantining behaviour in full. They quickly detected and removed contaminated aphids, placing them at some distance from the plant. Moreover, in addition to the simple carrying of ‘infected’ aphids down, the more effective technique of dropping them from the plant was noted as well. Ants of the genus Myrmica appear to have a tendency to perform a certain sequence of actions to remove conidia-contaminated aphids from the plant. It is likely that in larger colonies or under conditions of increased risk of infection with entomopathogenic fungi, some Myrmica ants are able to deploy and actively use the behavioural pattern of quarantining behaviour to increase their viability.     

Tuned protection of aphids by ants against a predatory hoverfly

1. Aphid‐tending ants that feed on honeydew have evolved strategies against aphidophagous insects and tune their aggressive behaviour according to the level of danger for their trophobionts. Here we investigate how Lasius niger Linnaeus (Hymenoptera: Formicidae) ants react to different instars of Episyrphus balteatus De Geer (Diptera: Syrphidae) hoverflies which vary in their voracity and defensive abilities. 2. During pairwise encounters, early syrphid instars (eggs, L1, and L2 larvae) elicited lower aggression scores compared to third larval instars (L3), which was intensively bitten by ants. L3 tried to escape from ants by releasing a sticky and toxic secretion over biting ants that died or underwent severe morbidity. 3. In a standardised system including the host plant, aphid, tending ant, and hoverfly, the ability of ants to protect an Aphis fabae Scopoli (Hemiptera: Aphididae) colony was evaluated. Early E. balteatus instars placed onto the plant elicited no mobilisation of ants, which often removed the hoverfly successfully. Eggs and early instars appeared as the weak links for integrated pest management by hoverfly auxiliaries. 4. In contrast, L3 induced the number of ant patrollers to increase at a local scale without any further recruitment from inside the ant nest. L3 syrphids were quite efficient at gluing ants with defensive secretions and at resisting to removal attempts by ants. 5. While supporting the assumption that ants tune their defensive response to the aphidophagous predator, the present results also showed a lack of efficient protection of their trophobionts from the most voracious late syrphid instar.     

Ant-seed interactions: combined effects of ant and plant species on seed removal patterns

Seed dispersal by ants (i.e. myrmecochory) is usually considered as a mutualism: ants feed on nutritive bodies, called elaiosomes, before rejecting and dispersing seeds in their nest surroundings. While mechanisms of plant dispersal in the field are well documented, the behaviour of the ant partner was rarely investigated in details. Here, we compared in laboratory conditions the foraging behaviour of two ant species, the omnivorous Lasius niger and the insectivorous Myrmica rubra to which seeds of two European myrmecochorous plants (Chelidonium majus and Viola odorata) were given. Ant colonies were simultaneously presented three types of items: entire seeds with elaiosome (SE), seeds without elaiosome (S) and detached elaiosomes (E). The presence of elaiosomes on seeds did not attract workers from a distance since ants first contact equally each type of items. Although ants are mass-recruiting species, we never observed any recruitment nor trail-laying behaviour towards seeds. For ants having contacted seed items, their antennation, manipulation and seed retrieval behaviour strongly varied depending on the species of each partner. Antennation behaviour, followed by a loss of contact, was the most frequent ant-seed interaction and can be considered as a “hesitation” clue. For both plant species, insectivorous Myrmica ants removed items in larger number and at higher speed than Lasius. This fits with the hypothesis of a convergence between odours of elaiosomes and insect preys. For both ant species, the small Chelidonium seeds were retrieved in higher proportion than Viola ones, confirming the hypothesis that ants prefer the higher elaiosome/diaspore-ratio. Thus, in these crossed experiments, the ant-plant pair Myrmica/Chelidonium was the most effective as ants removed quickly almost all items after a few antennations. The presence of an elaiosome body increased the seed removal by ants excepting for Myrmica which retrieved all Chelidonium seeds, even those deprived of their elaiosome. After 24 h, all the retrieved seeds were rejected out of the nest to the refuse piles. In at least half of these rejected items, the elaiosome was discarded by ants. Species-specific patterns and behavioural differences in the dynamics of myrmecochory are discussed at the light of ant ecology. Received 10 September 2007; revised 5 February 2008; accepted 5 March 2008.     

Behavioral mechanisms of spatial competition between red wood ants (Formica aquilonia) and ground beetles (Carabidae)

Behavioral aspects of spatial competition between red wood ants (Formica aquilonia) and six mass species of Carabidae were studied in field and laboratory experiments. We showed that red wood ants essentially influence spatial distribution of ground beetles on their common territories. Transplantation experiments suggest that in newly established ants' settlements stronger forms of interrelations arise than in old stable colony. To examine the ability of beetles to avoid collisions with ants we used two experimental techniques. In laboratory, we tested carabids ability to avoid a clash in a Y-shaped labyrinth containing an active tethered ant in one section. In field experiments we compared quantitative characteristics of movements (such as crookedness of individual trajectories, speed of movement, the time spent on stops) for beetles placed close to ants foraging routes and on ant-free plots. All beetles studied displayed a clear tendency to learn, that is, to modity their behavior in order to avoid collisions with ants. Species that exhibited best parameters of learning were closer to ants by their size and characteristic movement, namely, Pterostichus oblogopunctatus and P. magus. Beetles' stereotyped behavioral tactics can be considered universal for avoiding collisions with any subject (for instance, with an ant) of a certain size and speed of movements. A set of tactics in the labyrinth included: (1) attempts to round the ant; (2) turns away after touching the ant with antennae; (3) turns away without a contact; (4) avoidances of a dangerous section; (5) stops near the ant with the antennae hidden. Comparing pairwise difference between four species shows that beetles use species-specific preference for definite combinations of tactics. Effective learning allows carabids to penetrate into ant foraging territory and partly avoide interference competition. It seems that red wood ants are not inclined to learn to avoid collisions with competing carabid species. Instead, they recognize an "enemy's image" and selectively attack relatively small predatory carabids rather than herbivorous species. Experiments with dummy beetles suggest that ants react for several hierarchically organized key characteristics of competitors such as speed of movement, dark color, bilateral symmetry, protuberances (legs, antennae), and smell. Among "professional" groups in ant family, guards and hunters react to beetles aggressively, whereas aphid tenders ignore them. Sophisticated combination of flexible and innate behavioral patterns enables insects to share territories and interact in the mode of relatively mild, spatial, competition instead of predation. Eliciting sets of hierarchically organized features of competitors that govern adjustment of spatial distribution in insects' community enable us to integrate ideas of classic and cognitive ethology.     

Choice of food resource by bees and ants (Hymenoptera: Apoidea,Formicidae) on coconut tree inflorescences (Cocos nucifera,Arecaceae)

Coconut trees are mostly anemophilous; however, because bees and ants forage on coconut tree inflorescences for floral food, entomophilous pollination can also occur. The aim of this study was to determine the food resource preference of bees and ants while they collect pollen, nectar and, for ants, occasionally prey on coconut tree inflorescences, as well as to evaluate their impact on self-pollination. The number of ant visits to first female and then male flowers is significantly higher than that of bees. For Apis mellifera (L.) and Pseudomyrmex gracilis (Fabricius) 14% of the sequences were favorable to direct self-pollination. The probability of visits for all of the sequences was similar for both bees and ants and there was no difference in resource choice. For these reasons, neither can be considered a more effective pollinator of the coconut tree.     

The worldwide transfer of ants: geographical distribution and ecological invasions

Aim This is the first comprehensive account of the biogeography of ants transferred and at least temporarily established outside their native habitat. Location Using museum and literature records, I established the distributions of transferred ant species. Methods I used taxonomic and functional groups to assess how geographical spread as a transferred species is affected by taxonomy and life history. Results 147 ant species in forty-nine genera have been recorded outside of their native habitat. The proportion of transferred ants is similar to the number of genera and species in each subfamily. The species-rich subfamily Myrmicinae contains nearly 50% of all transferred species, while many of the species-poor subfamilies have absolutely no transferred species. A disproportionate high number of transferred ants originate from the Neotropical and Oriental biogeographic regions. The Pacific Islands are the recipients of the most transferred ant species. Most transferred ants belong to the CRYPTIC, OPPORTUNIST, and GENERALIZED MYRMICINE functional groups, while there are no recorded transfers of army ants or leaf-cutting ants. Both invasive and human commensal ‘tramp’ ant species are nonrandom subsets of transferred ants. Main conclusions ‘Tramp’ species and invasive species tend to have widespread geographical distributions, and share life history characteristics including queen number, nest structure, and foraging behaviour. Combining observations of functional groups and biogeography may lead to a better understanding of the factors contributing to the spread of transferred species.     

How a lack of choice can force ants to climb up waxy plant stems

The aim of this study was to analyse the ability of generalist ants to climb waxy stems and to understand whether the avoidance of such stems is solely based on their reduced ability to adhere to these surfaces. We performed observations of the locomotory behaviour and recorded the traversed distances of Lasius niger ants on flower stems of three plant species covered with epicuticular wax crystals: Anethum graveolens, Dahlia pinnata, and Tagetes patula. Using conventional and cryo-scanning electron microscopy, the surface micromorphology of plant stems and structure of ant attachment organs were examined. The present study demonstrates that the reason ants do not climb wax-covered stems is not because they are incapable of walking on such vertical surfaces, as previously assumed, but rather because they have a choice of other substrates. Presumably, additional locomotory efforts are needed to master climbing on “greasy” stems, and ants need a certain time to adapt to a new quality of the stem surface, and after this period, they can walk on these waxy stems.     

Coping with the cold: minimum temperatures and thermal tolerances dominate the ecology of mountain ants

1. Ants (Hymenoptera: Formicidae) are often cited as highly thermophilic and this has led to a range of studies investigating their thermal tolerances. It is unknown, however, if the geographic distribution of ant thermal tolerance conforms to the two major macropyhsiological rules that have been found in other taxa: Janzen's and Brett's rules. In addition, there is a paucity of data on how the lower thermal tolerances of ants are able to influence behaviour. 2. These two knowledge gaps were addressed here by sampling ants across a 1500 m elevational gradient in southern Africa and estimating the upper (CTmax) and lower (CTmin) thermal tolerances of 31 and 28 species, respectively. Ant abundances and soil temperatures were also recorded across the gradient over 6 years. 3. It was found that the average CTmin of the ants declined with elevation along with environmental temperatures. It was also found that the correlation between abundance and local temperature depended on the ant species' CTmin. The activity of species with a low CTmin was not constrained by temperature, whereas those with a high CTmin were limited by low temperatures. 4. For the first time, evidence is provided here that the thermal tolerances of ants are consistent with two major macrophysiological rules: Brett's rule and Janzen's rule. A mechanistic link between physiology, behaviour and the environment is also shown, which highlights that the ability of ants to deal with the cold may be a key, but often overlooked, factor allowing multiple ant species to succeed within an environment.     

Response of the foraging behaviour of red wood ants (Formica rufa group) to exclusion from trees

Abstract 1 Because of the large numbers within a colony and their aggressive nature, red wood ants (Formica rufa group) have a potential to greatly influence the cold‐temperate forest ecosystem. Wood ants are omnivorous and hunt in trees as well as on the forest floor. 2 A field experiment in a mixed forest in central Sweden was carried out to examine (i) the foraging behaviour of wood ants on the forest floor and (ii) the impact of increased numbers of wood ants on the soil fauna. The foraging behaviour of wood ants was manipulated by excluding the ants from their food resources in the tree canopy, with the intention to increase ant activity on the forest floor. To estimate this activity, the number of trees with foraging ants, the numbers of ants going to and from their nests and the prey carried by home‐running wood ants were determined during the summer period. Pitfall traps were placed in the soil to determine effects on mobile soil invertebrates. 3 When excluded from local trees, wood ants searched other trees further away from the nests rather than searching more intensively for prey on the forest floor. By contrast to the initial hypothesis, more soil‐living prey were caught by ants in the control plots than in the plots where the local trees were not accessible to the wood ants. The proportion of soil‐living to tree‐living prey tended to be greater in the control plots. 4 In the treated plots (no access to the trees), wood ants had a negative effect on the activity of Linyphiidae spiders. There was little effect of wood ants on other soil invertebrates. 5 This study suggests that the role of wood ants as top predators in the forest soil food‐web in central Sweden is limited.     

The response of grass-cutting ants to natural and synthetic versions of their alarm pheromone

The responses of the grass‐cutting ants Atta bisphaerica (Forel) and Atta capiguara (Gonçalves) to the main components of their alarm pheromones were examined in simple field bioassays. Both species react most strongly to 4‐methyl‐3‐heptanone, which causes the full range of alarm behaviour and a large increase in the number of individuals near the sources. In later experiments with A. capiguara, this increase was found to be due primarily to attraction, with some arrestment also occurring. The ant response to 4‐methyl‐3‐heptanone was compared with that to crushed heads and to that with whole ants with crushed heads. The pheromone 4‐methyl‐3‐heptanone by itself stimulates the same level of attraction as crushed heads, but results in far less alarm behaviour and arrests fewer ants. Whole ants with crushed heads attract a greater number of ants than the other sources and also cause more alarm behaviour. Bodies alone attract ants, but do not result in alarm behaviour. The main component in both species is the same, supporting the view that alarm pheromones lack species specificity. However, it appears that other components may also be important either as synergists of the main compound, or by stimulating behaviours that would not be observed in its absence.     

Ants (Formicidae) as food for birds in southern Africa: opportunism or survival?

Ants (Hymenoptera: Formicidae) are eaten by a number of bird species in southern Africa. Our database contained 545 species (excluding waterbirds and raptors), of which 179 species have been observed feeding on ants, or had ants in their stomachs. Ants are eaten by birds in all ecosystems, but the consumption of ants is disproportionately high in species that occur in arid ecosystems. The number of bird species eating ants increases in proportion to the number of bird species in any ecosystem, but it increases disproportionately with decreasing mean annual rainfall. There was a small, but not significant, difference in the proportion of ant-eating bird species between 502 resident species, of which 154 (30.6%) eat ants, and 47 nomadic species, of which 18 (38.2%) eat ants. Foraging behaviour and substrate, i.e. whether foraging in the air, from a perch or on the ground, significantly influenced the probability that ants would be included in the diet.     

Ants move to improve: colonies of Leptothorax albipennis emigrate whenever they find a superior nest site

A high-quality home can be a major factor determining fitness. However, when house hunting becomes necessary, animals might often face a speed-versus-accuracy trade-off and therefore be unable to survey their environment extensively for the optimum site. We found that the ant Leptothorax albipennis was able to correct errors made in such a hurried decision by continuing to survey even after a colony had settled in a nest site. Colonies moved from intact undisturbed nests to another nest site whenever the new nest site presented a sufficient improvement in nest quality. Thus, scout ants must be able to judge and compare the quality of the new, empty nest site with the one currently inhabited by the colony. Emigrations from intact nests were initiated by high numbers of ants recruited by tandem runs. This evidence may explain how a small number of scouts can motivate an entire colony to move when there is no immediate need to do so. Compared with their behaviour in emigrations from destroyed nests, the ants favoured even more strongly accuracy over speed, because they waited for a larger number of scouts to agree on one site before starting the emigration. They could do this without increased risk because the rest of the colony remained safely in the old nest.     

Impact of seed abundance on seed processing and dispersal by the red ant Myrmica rubra

1. Myrmecochory sensu stricto is an ant–plant mutualism in which non‐granivorous ants disperse plant diaspores after feeding on their nutrient‐rich seed appendage, the elaiosome. Phenological traits associated with the diaspore can influence the behaviour of ants and thus their ultimate efficiency as seed dispersers. 2. This study investigated how a contrasting availability of seeds (20 vs. 200 seeds) from the diplochorous Chelidonium majus (Papaveraceae, Linnaeus) plant species influences the behaviour of Myrmica rubra (Formicidae, Linnaeus) ants, from the retrieval of seeds until their dispersal outside the ant nest. 3. Regardless of seed abundance, the ants collected the first diaspores at similar rates. Then, seed retrieval sped up over time for large seed sources until satiation took place with only one‐third of the tested colonies wholly depleting abundant seed sources. 4. No active recruitment by trail‐laying ants was triggered, even to an abundant seed source 5. In both conditions of seed abundance, the majority of the diaspores retrieved inside the nest were discarded with the elaiosome removed and were dispersed at similar distances from the nest. 6. The paper concludes with a discussion of how the quantity of seeds released by a plant with a dual mode of dispersal can potentially influence the behaviour of ant dispersers and hence the dispersal efficiency derived from myrmecochory.     

The volume of reinforcement and the structure of rat behavior in a radial labyrinth

In rats choice reaction was elaborated in 12 passages radial labyrinth with different volumes of food only in six of twelve passages. Then the conditions were modified: the first group of rats in the beginning of the experiment was placed to another labyrinth sector, and in the second group the food volume was changed in passages. In new situations in animals of both groups the reaction efficiency temporarily decreased, simultaneously the correction of behaviour structure took place. The character of behavioural tactics (tendencies to visit passages with larger food volume at the beginning of the experiment, spatial-motor asymmetry etc.) is described in dynamics of adaptation to new conditions. Factors determining the rats behaviour structure are discussed.