The infrabuccal pellet piles of fungus-growing ants

Fungus-growing ants (Attini) live in an obligate mutualism with the fungi they cultivate for food. Because of the obligate nature of this relationship, the success of the ants is directly dependent on their ability to grow healthy fungus gardens. Attine ants have evolved complex disease management strategies to reduce their gardens exposure to potential parasitic microbes, to prevent the establishment of infection in their gardens, and to remove infected garden sections. The infrabuccal pocket, a filtering device located in the oral cavity of all ants, is an integral part of the mechanisms that leaf-cutter ants use to prevent the invasion and spread of general microbial parasites and the specific fungal-garden parasite Escovopsis. Fungus-growing ants carefully groom their garden, collecting general debris and pathogenic spores of Escovopsis in their infrabuccal pocket, the contents of which are later expelled in dump chambers inside the nest or externally. In this study we examined how a phylogenetically diverse collection of attine ants treat their infrabuccal pellets. Unlike leaf-cutters that deposit their infrabuccal pellets directly in refuse piles, ants of the more basal attine lineages stack their infrabuccal pellets in piles located close to their gardens, and a separate caste of workers is devoted to the construction, management, and eventual disposal of these piles.     

<Emphasis Type="Italic">Gnamptogenys hartmani</Emphasis> Wheeler (Ponerinae: Ectatommini): an agro-predator of<Emphasis Type="Italic"> Trachymyrmex</Emphasis> and<Emphasis Type="Italic"> Sericomyrmex</Emphasis> fungus-growing ants

The fungus gardens of fungus-growing ants are a potentially valuable resource for exploitation by natural enemies, but few of these antagonistic interactions have been studied. Here we describe key aspects of the behavioral ecology of Gnamptogenys hartmani (Ponerinae: Ectatommini), a specialized agro-predator of Trachymyrmex and Sericomyrmex fungus-growing ants in Panama. Raiding columns of G. hartmani attack and usurp nests with remarkably little effort: a few intruding workers are sufficient to cause panic among the attine ants and make them abscond from the nest. Both G. hartmani larvae and adults consume the fungus and the host brood, after which the colony migrates to a new fungus-growing ant nest discovered by scouting workers. The morphology of the G. hartmani larval mouthparts is similar to that of Gnamptogenys species with a non-fungal diet. However, we suggest that the presence of long spinules on the larval mandibles in the genus Gnamptogenys, comparable to those found in attine larvae, may have pre-adapted G. hartmani to fungus eating. G. hartmani workers do not actively maintain or modify fungus gardens, which makes them less efficient exploiters than Megalomyrmex, the only other agro-predatory ant species known so far.     

Vector navigation in desert ants, Cataglyphis fortis: celestial compass cues are essential for the proper use of distance information

Foraging desert ants navigate primarily by path integration. They continually update homing direction and distance by employing a celestial compass and an odometer. Here we address the question of whether information about travel distance is correctly used in the absence of directional information. By using linear channels that were partly covered to exclude celestial compass cues, we were able to test the distance component of the path-integration process while suppressing the directional information. Our results suggest that the path integrator cannot process the distance information accumulated by the odometer while ants are deprived of celestial compass information. Hence, during path integration directional cues are a prerequisite for the proper use of travel-distance information by ants.     

Ant-lepidopteran associations along African forest edges

Working along forest edges, we aimed to determine how some caterpillars can co-exist with territorially dominant arboreal ants (TDAAs) in tropical Africa. We recorded caterpillars from 22 lepidopteran species living in the presence of five TDAA species. Among the defoliator and/or nectarivorous caterpillars that live on tree foliage, the Pyralidae and Nymphalidae use their silk to protect themselves from ant attacks. The Notodontidae and lycaenid Polyommatinae and Theclinae live in direct contact with ants; the Theclinae even reward ants with abundant secretions from their Newcomer gland. Lichen feeders (lycaenid; Poritiinae), protected by long bristles, also live among ants. Some lycaenid Miletinae caterpillars feed on ant-attended membracids, including in the shelters where the ants attend them; Lachnocnema caterpillars use their forelegs to obtain trophallaxis from their host ants. Caterpillars from other species live inside weaver ant nests. Those of the genus Euliphyra (Miletinae) feed on ant prey and brood and can obtain trophallaxis, while those from an Eberidae species only prey on host ant eggs. Eublemma albifascia (Erebidae) caterpillars use their thoracic legs to obtain trophallaxis and trophic eggs from ants. Through transfer bioassays of last instars, we noted that herbivorous caterpillars living in contact with ants were always accepted by alien conspecific ants; this is likely due to an intrinsic appeasing odor. Yet, caterpillars living in ant shelters or ant nests probably acquire cues from their host colonies because they were considered aliens and killed. We conclude that co-evolution with ants occurred similarly in the Heterocera and Rhopalocera.     

The ladybird<Emphasis Type="Italic"> Thalassa saginata</Emphasis>, an obligatory myrmecophile of<Emphasis Type="Italic"> Dolichoderus bidens</Emphasis> ant colonies

The larvae and pupae of the ladybird Thalassa saginata develop inside colonies of the dolichoderine ant Dolichoderus bidens. This association is the first specific and obligatory relationship recorded between ants and ladybirds. The ants provide shelter and protection to the larvae but the diet of the latter remains unclear. The integration of T. saginata larvae into the ant colonies is achieved by mimicking the cuticular patterns of the ants brood. Moreover, the larvae secrete substances from their hairs and anal gland that are likely to enhance their attractiveness.     

Rapid modification in the olfactory signal of ants following a change in reproductive status

In insect societies, the presence and condition of egg-layers can be assessed with pheromones. Exocrine secretions are expected to vary in time in order to give up-to-date information on an individuals reproductive physiology. In the queenless monogynous ant Streblognathus peetersi, we allowed a previously infertile high-ranking worker to accede to the alpha rank, thus triggering the onset of her oogenesis (15 replicates). We then studied her interactions with an established egg-layer from the same colony after different durations, ranging from 20 h to several days. Even though her eggs are only ready to be laid after 30 days, the new alpha was recognised within 1–2 days. Detection occurred at a distance of a few millimetres, suggesting the involvement of a pheromone with low volatility, such as cuticular hydrocarbons. When the new alpha had differentiated for >48 h, she was attacked by the established egg-layer. In all cases, low-ranking workers eventually immobilised one of the two alphas: the new alpha was the target if she had differentiated only recently, suggesting that police workers select the dominant worker with the less fertile odour. Using the behaviour of ants as our measure, we demonstrate that a dominants olfactory signal changes rapidly with a modification in her social status, and it occurs well before the onset of egg-laying.     

Circadian consequences of social organization in the ant species<Emphasis Type="Italic"> Camponotus compressus</Emphasis>

The locomotor activity rhythm of different castes of the ant species Camponotus compressus was monitored individually under laboratory light/dark (LD) cycles, and under continuous darkness (DD). The colony of this ant species comprises two sexual castes, the queens and the males, and three worker castes, namely the major, media, and minor workers. The virgin males and virgin queens display rhythmic activity patterns, but the mated queens were arrhythmic while laying eggs, with the rhythmicity resuming soon after egg-laying. Under the LD regime, major workers showed nocturnal patterns, while about 75% of the media workers displayed nocturnal patterns and about 25% showed diurnal patterns. Under the DD regime, most major workers exhibited circadian rhythm of activity with a single steady state, whereas media workers displayed two types of activity patterns, with activity patterns changing after 6–9 days in DD (turn-arounds). The pre-turn-around of the ants that showed nocturnal activity patterns during LD entrainment was <24 h after release into DD, which then became >24 h, after 6–9 days. On the other hand, the pre-turn-around of those ants that exhibited diurnal patterns during LD entrainment was first >24 h after release into DD, and then became <24 h, after 6–9 days. The activity of the minor workers neither entrained to LD cycles nor showed any sign of free-run in DD. It appears that the circadian clocks of the ant species C. compressus are flexible, and may perhaps depend upon the tasks assigned to them in the colony.     

Symmetry is in the eye of the ‘beeholder’: innate preference for bilateral symmetry in flower-naïve bumblebees

Bilateral symmetry has been considered as an indicator of phenotypic and genotypic quality supporting innate preferences for highly symmetric partners. Insect pollinators preferentially visit flowers of a particular symmetry type, thus leading to the suggestion that they have innate preferences for symmetrical flowers or flower models. Here we show that flower-naïve bumblebees (Bombus terrestris), with no experience of symmetric or asymmetric patterns and whose visual experience was accurately controlled, have innate preferences for bilateral symmetry. The presence of color cues did not influence the bees original preference. Our results thus show that bilateral symmetry is innately preferred in the context of food search, a fact that supports the selection of symmetry in flower displays. Furthermore, such innate preferences indicate that the nervous system of naïve animals may be primed to respond to relevant sensory cues in the environment.     

Social prophylaxis through distant corpse removal in ants

Living in groups raises important issues concerning waste management and related sanitary risks. Social insects such as ants live at high densities with genetically related individuals within confined and humid nests, all these factors being highly favorable for the spread of pathogens. Therefore, in addition to individual immunity, a social prophylaxis takes place, namely, by the removal of risky items such as corpses and their rejection at a distance from the ant nest. In this study, we investigate how Myrmica rubra workers manage to reduce encounters between potentially hazardous corpses and nestmates. Using both field and laboratory experiments, we describe how the spatial distribution and the removal distance of waste items vary as a function of their associated sanitary risks (inert item vs. corpse). In the field, corpse-carrying ants walked in a rather linear way away from the nest entrance and had an equal probability of choosing any direction. Therefore, they did not aggregate corpses in dedicated areas but scattered them in the environment. In both field and laboratory experiments, ants carrying corpses dropped their load in more remote—and less frequented—areas than workers carrying inert items. However, for equidistant areas, ants did not avoid dropping corpses at a location where they perceived area marking as a cue of high occupancy level by nestmates. Our results suggest that ants use distance to the nest rather than other occupancy cues to limit sanitary risks associated with dead nestmates.     

Influence of interspecific competition on the recruitment behavior and liquid food transport in the tramp ant species Pheidole megacephala

This study was conducted on the reactions of Pheidole megacephala scouts when finding liquid food sources situated on territories marked by competing dominant ant species or on unmarked, control areas to see if the number of recruited nestmates is affected and if soldiers behave in ways adapted to the situation. We show that scouts recruit more nestmates, particularly soldiers, on marked rather than on unmarked areas. This recruitment allows P. megacephala to organize the defence and rapid depletion of these food sources prior to any contact with competitors. Soldiers can carry liquid foods both (1) in their crops like other Myrmicinae and (2), in a new finding concerning myrmicine ants, under their heads and thoraxes like certain poneromorph genera because the droplets adhere through surface tension strengths. Later, the liquids stored in the crop are distributed to nestmates through regurgitations during trophallaxis and the external droplets are distributed through social buckets, or the mode of liquid food transfer common in poneromorphs. Their flexibility to use or not use the latter technique, based on the situation, corroborates other reports that Pheidole soldiers have a relatively large behavioral repertoire.     

Morphology and structure of the tarsal glands of the stingless bee Melipona seminigra

Footprint secretions deposited at the nest entrance or on food sources are used for chemical communication by honey bees, bumble bees, and stingless bees. The question of the glandular origin of the substances involved, however, has not been unequivocally answered yet. We investigated the morphology and structure of tarsal glands within the fifth tarsomeres of the legs of workers of Melipona seminigra in order to clarify their possible role in the secretion of footprints. The tarsal gland is a sac-like fold forming a reservoir. Its glandular tissue is composed of a unicellular layer of specialized epidermal cells, which cover the thin cuticular intima forming the reservoir. We found that the tarsal glands lack any openings to the outside and therefore conclude that they are not involved in the secretion of footprint substances. The secretion produced accumulates within the glands reservoir and reaches as far as into the arolium. Thus it is likely that it serves to fill and unfold the arolium during walking to increase adhesion on smooth surfaces, as is known for honey bees and weaver ants.     

In-nest environment modulates nestmate recognition in the ant<Emphasis Type="Italic"> Camponotus fellah</Emphasis>

Multiple behavioral and chemical studies indicate that ant nestmate recognition cues are low-volatile substances, in particular hydrocarbons (HCs) located on the cuticular surface. We tested the hypothesis that in the ant Camponotus fellah, nest environment, in particular nest volatile odors, can modulate nestmate-recognition-mediated aggression. Workers were individually confined within their own nest in small cages having either a single mesh (SM = limited physical contact permitted) or a double mesh (DM = exposed to nest volatiles only) screen. Individual workers completely isolated outside their nest (CI) served as control. When reintroduced into a group of 50 nestmates, the CI workers were attacked as alien ants after only 2 weeks of separation, whereas the SM workers were treated as nestmates even after 2 months of separation. Aggression towards DM ants depended on the period of isolation. Only DM workers isolated for over 2 months were aggressed by their nestmates, which did not significantly differ from the CI nestmates. Cuticular HC analyses revealed that the profile of the non-isolated ants (NI) was clearly distinct from that of CI, SM and DM ants. Profile differences matched the aggressive response in the case of CI ants but were uncorrelated in the case of SM or DM ants. This suggests that keeping the ants within the nest environment affected nestmate recognition in additional ways than merely altering their HC profile. Nest environment thus appears to affect label–template mismatch by modulating aggressive behavior, as well as the direction at which cuticular HCs diverged during the separation period.     

Wind and sky as compass cues in desert ant navigation

While integrating their foraging and homing paths, desert ants, Cataglyphis fortis, depend on external compass cues. Whereas recent research in bees and ants has focused nearly exclusively on the polarization compass, two other compass systems—the sun compass and the wind (anemo) compass—as well as the mutual interactions of all these compass systems have received little attention. In this study, we show that of the two visual compass systems, it is only the polarization compass that invariably outcompetes the wind compass, while the sun compass does so only under certain conditions. If the ants are experimentally deprived of their polarization compass system, but have access simultaneously to both their sun compass and their wind compass, they steer intermediate courses. The intermediate courses shift the more towards the wind compass course, the higher the elevation of the sun is in the sky.     

Is dauer pheromone of<Emphasis Type="Italic"> Caenorhabditis elegans</Emphasis> really a pheromone?

Animals respond to signals and cues in their environment. The difference between a signal (e.g. a pheromone) and a cue (e.g. a waste product) is that the information content of a signal is subject to natural selection, whereas that of a cue is not. The model free-living nematode Caenorhabditis elegans forms an alternative developmental morph (the dauer larva) in response to a so-called dauer pheromone, produced by all worms. We suggest that the production of dauer pheromone has no fitness advantage for an individual worm and therefore we propose that dauer pheromone is not a signal, but a cue. Thus, it should not be called a pheromone.     

Mating behaviour in a slave-making ant, Rossomyrmex minuchae (Hymenoptera, Formicidae)

The mating behaviour of the ant Rossomyrmex minuchae, a rare, protected slave-making species in Spain, seems to be significantly affected by its particular life history and patchy habitat. The mating behaviour of the entire genus Rossomyrmex is virtually unknown. We present here the results of a 3-year study of mating behaviour in R. minuchae.Behavioural observations and limited nest excavations revealed that R. minuchae does not produce sexuals every year, the number of sexuals is low, and the sex ratio tends to be female biased. Females typically exhibit two distinct activity periods. The first, the mating period, takes place in early afternoon: the ants call near the natal nest, mate and then return to their nest. The second, the dispersal period takes place in late afternoon: the mated females exit their nest and fly in search of a new, non-parasitized Proformica longiseta host nest. Males are highly active during the mating period, but will remain inactive in the dispersal period even if experimentally presented with virgin females. It appears that females are monogamous, while males are polygamous. When males are late arriving at the female calling site, the females will frequently congregate presumably calling in chorus.The low reproductive efficiency exhibited by R. minuchae, coupled with the postulated low genetic variation in the population, as sisters may mate with the same male, could result in a low survival rate and risk of eventual extinction. The observed decrease in nest density we observed during the 2004 season may be indicative of such a process.     

Acoustic mirror effect increases prey detection distance in trawling bats

Many different and phylogenetically distant species of bats forage for insects above water bodies and take insects from and close to the surface; the so-called trawling behaviour. Detection of surface-based prey by echolocation is facilitated by acoustically smooth backgrounds such as water surfaces that reflect sound impinging at an acute angle away from the bat and thereby render a prey object acoustically conspicuous. Previous measurements had shown that the echo amplitude of a target on a smooth surface is higher than that of the same target in mid-air, due to an acoustic mirror effect. In behavioural experiments with three pond bats (Myotis dasycneme), we tested the hypothesis that the maximum distances at which bats can detect prey are larger for prey on smooth surfaces than for the same prey in an airborne situation. We determined the moment of prey detection from a change in echolocation behaviour and measured the detection distance in 3D space from IR-video recordings using stereo-photogrammetry. The bats showed the predicted increase in detection distance for prey on smooth surfaces. The acoustic mirror effect therefore increases search efficiency and contributes to the acoustic advantages encountered by echolocating bats when foraging at low heights above smooth water surfaces. These acoustic advantages may have favoured the repeated evolution of trawling behaviour.     

Intercontinental chemical variation in the invasive ant Wasmannia auropunctata (Roger) (Hymenoptera Formicidae): a key to the invasive success of a tramp species

Unicoloniality emerges as a feature that characterizes successful invasive species. Its underlying mechanism is reduced intraspecific aggression while keeping interspecific competitiveness. To that effect, we present here a comparative behavioural and chemical study of the invasive ant Wasmannia auropunctata in parts of its native and introduced ranges. We tested the hypothesis that introduced populations (New Caledonia archipelago) have reduced intraspecific aggression relative to native populations (e.g., Ilhéus area, Brazil) and that this correlates with reduced variability in cuticular hydrocarbons (CHCs). As predicted, there was high intraspecific aggression in the Brazilian populations, but no intraspecific aggression among the New Caledonian populations. However, New Caledonian worker W. auropunctata remained highly aggressive towards ants of other invasive species. The chemical data corresponded with the behaviour. While CHCs of ants from the regions of Brazil diverged, the profiles of ants from various localities in New Caledonia showed high uniformity. We suggest that in New Caledonia W. auropunctata appears to behave as a single supercolony, whereas in its native range it acts as a multicolonial species. The uniformity of recognition cues in the New Caledonia ants may reflect a process whereby recognition alleles became fixed in the population, but may also be the consequence of a single introduction event and subsequent aggressive invasion of the ecosystem. Chemical uniformity coupled with low intraspecific but high interspecific aggression, lend credence to the latter hypothesis.     

The asymmetric scent: ringtailed lemurs (<Emphasis Type="Italic">Lemur catta</Emphasis>) have distinct chemical signatures in left and right brachial glands

Distinctive cues are predicted to evolve when the benefits obtained by the recognition process overcome its costs. When individual recognition is particularly beneficial for both senders and receivers, the expression of strongly distinctive signals is predicted to evolve. On the other hand, it could be predicted that each individual should show a very stable individual signature. In the same perspective, a great stability of the individual signatures could be expected. Lemur catta is the first non-human primate in which olfactory individual recognition has been demonstrated on the basis of the specialized brachial gland secretions. In this paper, I performed gas chromatograph analyses of right and left gland samples collected in two different periods (breeding and non-breeding seasons) from seven males. The aim was to verify if a diversification in such cues, already demonstrated at the inter-individual level, also occurs at the intra-individual level between left and right glands. I verified, by discriminant analysis and chemical distance comparisons, that each gland of each lemur has its particular signature that is maintained through time. Moreover, such diversification resulted so marked to make the overall intra-individual chemical differences similar to/as strong as the inter-individual ones. Since in rodents several odors from different glands may be integrated in individual recognition, I suggest that bilateral diversification in L. catta scents may offer an enhanced distinctiveness that could provide benefits in mate choice and social relationships.     

Orientation and navigation during adult transport between nests in the ant Cataglypis iberica

  Cataglyphis iberica is a polydomous ant species in which adult transports between nests are frequently observed. When pairs of workers were captured and released at the same location, the transporters (Ts) fled directly towards their destination nest and reached it in most of the cases. The transportees (Te), on the other hand, fled in the opposite direction and only a third of them eventually reached their nest of departure. Additional experiments suggest that this result may be explained by the fact that the Ts ants have a memory of the compass direction of the nest they are heading to and that they adjust their course by using a sequence of memorised landmarks. As regards to the Te, the reversal of their direction of transport seems to be based essentially on celestial cues. Received: 20 October 1999 / Accepted in revised form: 10 May 2000     

Nestmate recognition in ants is possible without tactile interaction

Ants of the genus Camponotus are able to discriminate recognition cues of colony members (nestmates) from recognition cues of workers of a different colony (non-nestmates) from a distance of 1 cm. Free moving, individual Camponotus floridanus workers encountered differently treated dummies on a T-bar and their behavior was recorded. Aggressive behavior was scored as mandibular threat towards dummies. Dummies were treated with hexane extracts of postpharyngeal glands (PPGs) from nestmates or non-nestmates which contain long-chain hydrocarbons in ratios comparable to what is found on the cuticle. The cuticular hydrocarbon profile bears cues which are essential for nestmate recognition. Although workers were prevented from antennating the dummies, they showed significantly less aggressive behavior towards dummies treated with nestmate PPG extracts than towards dummies treated with non-nestmate PPG extracts. In an additional experiment, we show that cis-9-tricosene, an alkene naturally not found in C. floridanus' cuticular profile, is behaviorally active and can interfere with nestmate recognition when presented together with a nestmate PPG extract. Our study demonstrates for the first time that the complex multi-component recognition cues can be perceived and discriminated by ants at close range. We conclude that contact chemosensilla are not crucial for nestmate recognition since tactile interaction is not necessary.