Adult transport in the ant Cataglyphis iberica: a means to maintain a uniform colonial odour in a species with multiple nests

Abstract. . Societies of the ant Cataglyphis iberica (Hymenoptera, Formicidae) comprise several satellite queenless nests around a queenright nest. During spring, at the end of hibernation, the incidence of adult transport between these nests is high, but it decreases during summer. Quantitative analyses of the contents of postpharyngeal glands of transporter and transportee ants reveal the amounts of hydrocarbons to be lower in the latter, indicating that these ants are generally younger. Moreover, the more diverse composition of the transportees' secretion may reflect their individual makeup. Transporters, in contrast, maintained a uniform colony odour uirough trophallactic exchanges before entering hibernation. The odour disparity between the transportees and the general colony odour may stimulate the transporters to bear them to the nest containing the queen where they can obtain the colony odour via trophallaxis. The intense traffic between satellite nests in the spring may thus be the means by which a uniform colonial odour is regained after hibernation. Adult transport seems also to be necessary since the young transportee ants are not able to orient themselves and find the other nests.     

Prey availability and eclosion-help of callow workers in the formicine ant Camponotus mus

Summary In the ant Camponotus mus the eclosion of imagines from the cocoons was shown to be controlled by nurse workers, which broke the cocoons and assisted the callows to energe. Short-term colony prey deprivation modified the eclosion-help behavior resulting in a delay in cocoon opening times. During long-term prey deprivation, no new cocoons were spun; the starved larvae grew and pupated when prey was again available. These findings as a colony strategy adapted to the changing condition of prey availabitity in a temperate habitat are discussed.     

Social hybridogenesis in the clonal ant Cataglyphis hispanica

With a few rare exceptions, the vast majority of animals reproduce sexually. Some species have, however, evolved alternative modes of reproduction by shifting from classical bisexuality to unorthodox reproductive systems, like parthenogenesis, gynogenesis, or hybridogenesis. Under hybridogenesis, both the maternal and paternal genomes are expressed in somatic tissues, whereas the germline is purely maternal. Recently, a form of hybridogenesis at the level of the society has been reported in some ants, where purebred females develop into reproductive queens and interlineage hybrids into sterile workers. Here, we report a unique case of social hybridogenesis in the desert ant Cataglyphis hispanica. Workers are produced exclusively from interbreeding between two distinct genetic lineages, whereas male and female sexuals are produced by asexual reproduction through parthenogenesis. As a consequence, all workers are pure hybridogens, and only maternal genes are perpetuated from one generation to the next. Thus, queens of C. hispanica use sexual reproduction for colony growth, whereas they reproduce asexually through parthenogenesis for germline production.     

Microvillar orientation in the photoreceptors of the ant Cataglyphis bicolor

Polarization sensitivity in arthropod photoreceptors is crucially dependent on the arrangement of the microvilli within the rhabdom. Here, we present an electron-microscopical study in which the degree of microvillar alignment and changes in the cross-sectional areas of the rhabdoms along their length were studied in the compound eye of the desert ant, Cataglyphis bicolor. Serial cross-sections through the retina were taken and the orientation of the microvilli was determined in the photoreceptors of individually identified ommatidia. The reconstructions of microvillar alignment were made in the three anatomically and functionally distinct regions of the Cataglyphis compound eye: the dorsal rim area (DRA), the dorsal area (DA), and the ventral area (VA). The following morphological findings are consistent with polarization sensitivities measured previously by intracellular recordings. (1) The microvilli of the DRA photoreceptors are aligned in parallel along the entire length of the cell from the distal tip of the rhabdom down to its proximal end, near the basement membrane. The microvilli of the retinular cells R1 and R5 are always parallel to each other and perfectly perpendicular, with only minor deviation, to the microvillar orientation of the remaining receptor cells. (2) In the DA and VA regions of the eye, the microvillar tufts of the small receptors R1, R3, R5, R7, and R9 change their direction repetitively every 1-4 7m for up to 90°. In contrast, the large receptor cells R2, R4, R6, and R8 maintain their microvillar orientation rigidly. (3) In the DRA ommatidia, the cross-sectional areas of the rhabdomeres do not change along the length of the rhabdom, but substantial changes occur in the DA and VA ommatidia.     

Ontogeny of hydrocarbon profiles in the ant Aphaenogaster senilis and effects of social isolation

In ants, cuticular hydrocarbons are used for nestmate recognition; they are stored in the postpharyngeal gland and shared among the individuals. Newly emerged ants have a very small quantity of hydrocarbons. We studied the ontogeny of the hydrocarbon profile in Aphaenogaster senilis. The total quantities of both cuticular and postpharyngeal gland (PPG) hydrocarbons increased with age from 0 to 20 days after emergence and then stabilised. These quantities are correlated with the development of the ovary. Under individual social isolation, cuticular hydrocarbons increased as normal, but the total quantity of PPG hydrocarbons never increased from the initial low level. This effect of social isolation on the PPG hydrocarbon level indicates the importance of hydrocarbon transfer between nestmates through the PPG and lends support to the gestalt model of nestmate recognition. To cite this article: K. Ichinose, A. Lenoir, C. R. Biologies 332 (2009).     

Trophallaxis Mediates Uniformity of Colony Odor in Cataglyphis iberica Ants (Hymenoptera, Formicidae)

We studied the effect of nestmate separation on trophallaxis in the polydomous ant Cataglyphis iberica. After dividing three colonies into two equivalent subgroups, one queenright and one queenless, we quantified the frequency of trophallaxis within each subgroup, between the workers from the two subgroups (mixed trophallaxis), and trophallaxis involving the queen. Observations of trophallaxis were conducted over four periods of time: for 2 weeks before the separation of the two subgroups, 8 weeks during separation, immediately after reunification, and 3 weeks following reunification. Subgroups were identically fed on the eve of each day of observation. Group separation induced an increase in mixed frequencies of trophallaxis just after reunification, after which trophallaxis returned to the initial level observed before separation. Previous results showed that group separation in C. iberica induces hydrocarbon profile divergence and that reunification restores this chemical modification. The current results seem to indicate that increased trophallaxis permits a uniform odor to be reestablished among previously separated ants. Trophallaxis involving the queen is infrequent and does not seem to be crucial in the process of odor exchange. Our data confirm that trophallaxis plays a key role in establishing the Gestalt colony odor, particularly among naturally separated satellite nests in a polydomous species like C. iberica.     

Local resource competition and sex ratio in the ant Cataglyphis cursor

The local resource competition (LRC) hypothesis predicts thatwherever philopatric offspring compete for resources with theirmothers, offspring sex ratios should be biased in favor of thedispersing sex. In ants, LRC is typically found in polygynous(multiple queen) species where foundation of new nests occursby budding, which results in a strong population structure anda male-biased population-wide sex ratio. However, under polygyny,the effect of LRC on sex allocation is often blurred by theeffect of lowered relatedness asymmetries among colony members.Moreover, environmental factors, such as the availability ofresources, have also been shown to deeply influence sex ratioin ants. We investigated sex allocation in the monogynous (singlequeen) ant Cataglyphis cursor, a species where colonies reproduceby budding and both male and female sexuals are produced throughparthenogenesis, so that between-colony variations in relatednessasymmetries should be reduced. Our results show that sex allocationin C. cursor is highly male biased both at the colony and populationlevels. Genetic analyses indicate a significant isolation-by-distancein the study population, consistent with limited dispersal offemales. As expected from asexual reproduction, only weak variationsin relatedness asymmetry of workers toward sexual offspringoccur across colonies, and they are not associated with colonysex ratio. Inconsistent with the predictions of the resourceavailability hypothesis, the male bias significantly increaseswith colony size, and investment in males, but not in females,is positively correlated with total investment in sexuals. Overall,our results are consistent with the predictions of the LRC hypothesisto account for sex ratio variation in this species.     

Characterization of nuclear DNA microsatellite markers in the ant Cataglyphis cursor

The ant Cataglyphis cursor is exceptional in that unmated workers are potentially able to lay both male and female eggs. We characterized eight pairs of primers for microsatellite loci, developed from genomic DNA for this species. Variability was tested with DNA from 19 workers and all eight loci were highly polymorphic, displaying 5–10 alleles and a high level of heterozygosity. Cross‐species amplifications indicate that these microsatellites might be useful in genetic studies of other species belonging to the genus Cataglyphis.     

Hydrocarbon site of synthesis and circulation in the desert ant Cataglyphis niger

Chemical analyses revealed that in Cataglyphis niger both the hemolymph and the crop contain the same hydrocarbons that are found in the postpharyngeal gland (PPG). On the cuticle, on the other hand, alkanes, and in particular nonacosane, were more abundant than in the PPG. Studies of their biosynthesis in vivo, using intact ants, revealed the presence of newly synthesized hydrocarbons in both the PPG and the crop. In decapitated ants (in the absence of the PPG), however, the crop did not contain any newly synthesized hydrocarbons, indicating the PPG as the major source of crop hydrocarbons. The fat body, as demonstrated by in vitro studies, is the major tissue that biosynthesizes hydrocarbons. The PPG failed to do so, but showed good de novo biosynthesis of other lipid constituents. The large amount of hydrocarbons in the crop suggests that the alimentary canal may serve as an outlet for the overflow of PPG hydrocarbons, or as a route for the directed clearance of hydrocarbons from the PPG.These results confirm and enlarge the model proposed for hydrocarbon circulation in C. niger. They are synthesized by the fat body, released to the hemolymph and transported to the cuticle and the PPG. The PPG hydrocarbons are applied to the cuticle by self-grooming, but can also be cleared via the alimentary canal. Partial emptying of the PPG may facilitate the admixing of recognition cues that the ant may acquire from nestmates by trophallaxis. The reason for the dissimilarity in hydrocarbon composition between the PPG and the cuticle is not yet clear; it may be due to secretions from additional glands, or reflect deviant hydrocarbon transport mechanisms between the PPG and the cuticle.     

Mating system and population structure in the desert ant Cataglyphis livida

We investigated population genetic structure, mating system, worker reproduction and thelytokous parthenogenesis in the desert ant Cataglyphis livida. Pedigree analyses at polymorphic microsatellite loci show that colonies are headed by a single queen, and that queens are mated with two to eight males. No inbreeding was found in the population sampled. Colonies are genetically differentiated and exhibit no isolation-by-distance pattern, consistent with independent foundation of new colonies. Workers do reproduce and lay haploid (arrhenotokous) eggs in queenless colonies; conversely, we found no evidence of worker reproduction in queenright nests. In contrast with C. cursor, where new queens are produced by thelytokous parthenogenesis, female sexuals and workers of C. livida arise from classical sexual reproduction. We discuss the parallels and contrasts between the mating system and population structure in C. livida and the other Cataglyphis species studied so far.     

Discrimination of inclined path segments by the desert ant Cataglyphis fortis

Desert ants (Cataglyphis fortis) navigate by means of path integration, and perform accurately even in undulating terrain. They are able to correctly calculate the ground distance between nest and feeder even if their foraging excursion leads them over corrugated surfaces. To compute the respective ground projection when walking over an inclined surface, ants must measure its slope with sufficient accuracy—but how they do so is still not understood. Using a new behavioural assay that included a negative reinforcement, we investigated how well different slopes are discriminated by the ants. Ants were trained to visit an elevated feeder, via a ramp of fixed inclination (five training inclinations were used: 0°, 15°, 30°, 45°, 60°). The ants discriminated a steeper test slope that differed from the training slope by 12.5°. This discrimination performance was found to be constant for training slopes between 0° and 45°. Ants trained on a 60° slope, however, did not discriminate all steeper slopes, up to a vertical ascent, from the 60° inclination. The consequences of this discrimination accuracy for errors in the path integration process are discussed.     

Cooperative prey-retrieving in the ant Cataglyphis floricola: an unusual short-distance recruitment

Cataglyphis ants are mostly scavengers adapted to forage individually in arid environments. Although they are widely thought to have lost the capacity of recruitment, we provide evidence that C. floricola foragers that find a large prey near their nest are able to solicit the help of nestmates to carry it cooperatively. After discovering a non-transportable prey, these ants readily return to their nest and stimulate the exit of several recruits. This rudimentary form of recruitment, which is absent in the sympatric species C. rosenhaueri, is only employed when the prey is sufficiently close to the nest entrance (<1 m) and does not allow the food location to be communicated. Instead, C. floricola recruits search for the prey in all directions until they discover it and transport it cooperatively to their nest.     

Chemical integration of Thorictus myrmecophilous beetles into Cataglyphis ant nests

Thorictus beetles of the Dermestidae are obligate myrmecophiles. To understand how these beetles are integrated into and tolerated by their host colonies, the cuticular hydrocarbon profiles of different species of the Thorictus castaneus group that are generally associated with Cataglyphis were examined. The beetles are characterized by small amounts of cuticular hydrocarbons, which render them partly chemically “insignificant”. They also have the same cuticular hydrocarbon profiles as their hosts and thus likely use chemical mimicry to evade worker hostility but, like slaves in slave-maker species, they maintain some partial chemical identity. Thorictus martinezi from Burkina Faso were immediately adopted by conspecific colonies of their host, Cataglyphis sp. aff. bicolor, but were never adopted by colonies of other species (i.e. Cataglyphis viatica and Formica selysi). Thorictus buigasi from Morocco also mimicked the chemical profile of its host, C. viatica, but, in contrast to T. martinezi, individuals were adopted by colonies of Cataglyphis velox from Spain. This result can be explained by the similarity between the hydrocarbon profiles of C. viatica and C. velox, which may facilitate adoptions. T. buigasi beetles remained in Formica selysi colonies for some time but were ultimately rejected, probably due to their very different hydrocarbon profiles. In contrast, they were sometimes adopted by Camponotus herculeanus colonies and eventually chemically matched their new hosts, probably by passive camouflage. These data suggest that Thorictus of castaneus group myrmecophily is the result of coevolution with Cataglyphis hosts and that the mimicry is plastic, such that beetles can live with different hosts if the hosts show very limited CHC differences.     

Worker demography and behavior in a supercolonial ant colony: The case of the desert ant Cataglyphis niger

Insect societies headed by multiple queens (polygyny) raise evolutionary questions, such as how does genetic heterogeneity among colony members affect in-nest interactions; or, are all queens equally reproductive or equally treated by workers? Answering such questions requires intensive and continuous observations of in-nest behavior. Here, we addressed these questions in the polygyne supercolonial ant, Cataglyphis niger, using a barcoding system that enables the tracking of individual interactions, together with polymorphic DNA microsatellite markers that indicate the matriline and patriline of all individuals. Our findings that both queens and workers have low interrelatedness corroborate earlier reports regarding the supercolony structure of C. niger. Ovary inspection and worker genotyping revealed that all the queens contribute similarly to nest demography. Tracking positions of individual workers through time revealed that only a small proportion of them are constantly engaged in tending the queens and can be considered as retinue workers. However, genotyping these workers and the attended queens revealed no relationship to genetic relatedness, again typical of a true polygyne and supercolonial species. Unlike invasive supercolonial species, C. niger is native to Israel, enabling us to address questions regarding the driving forces, other than kin selection, that stabilize this society.     

Structural and chemical ontogeny of the postpharyngeal gland in the desert ant Cataglyphis niger

Abstract. The postpharyngeal gland of Cataglyphis niger (André, 1881) workers is characterized, at the ultrastructural level, by a well-developed smooth endoplasmic reticulum, Golgi apparatus, and numerous mitochondria and lamellar inclusions suggesting an involvement in lipid metabolism. In addition, the microvillar differentiation of the apical membrane and basal invaginations of the cell imply a transportation process.
The gland shows a clear structural as well as chemical age-dependent development. Epithelial thickness increases with age, reaching its maximum within the first week after emergence. Concurrently there is a progressive development of microvilli and appearance of the lamellar inclusions. The gland's secretion consists mainly of hydrocarbons, with a predominance of monomethyl and dimethyl alkanes. Congruency was found between the hydrocarbon composition of glandular exudates and those of the epicuticular washes. An age-dependent increase in the total amount of hydrocarbons was found to correlate with the ultrastructural development and gross morphology (thickness of epicuticular lining) of the gland.     

Phenotypic plasticity of immune defence linked with foraging activity in the ant Cataglyphis velox

Because immune defences are costly, life-history theories predict a modulation of immune investment according to its potential benefits. Social insects provide interesting models since infection risk may vary among individuals within a colony. In particular, the foraging workers, that have to leave the nest, suffer a higher infection risk and can contaminate their nest, which may favour high immune investments. However, evolutionary theories of aging predict that foragers should reduce their immune investment when they suffer high extrinsic mortality. To test these two predictions, we investigated the levels of phenoloxidase (PO) and prophenoloxidase, two important enzymes of the insect immune system, in workers of the ant Cataglyphis velox. We found a higher PO activity in foragers than in intra-nidal workers. This could result from an adaptive upregulation of the harmful PO (an enzyme potentially leading to autoimmune reactions) only when the risk of infection and wounding is high.     

Golgi studies of the first optic ganglion of the ant,Cataglyphis bicolor

The neurons of the first optic ganglion (the lamina) in the desert ant, Cataglyphis bicolor, have been studied with the light microscope after Golgi silver impregnation. The different types of retinal and laminal fibres and their configuration are compared with the results obtained in the bee. The first synaptic region in the visual system of the ant lies proximally to the fenestrated layer below the basement membrane and the layer containing the monopolar cell bodies. The synaptic region can be separated into three morphologically different zones: (1) The most distal layer where the short visual fibres end at two different levels. The short visual fibres and some laminal fibres (monopolar cell fibres) also show lateral elements in this region. (2) The second layer appears almost free of branches of retinal and laminal fibres. (3) The most proximal layer, which has a characteristically dense horizontal structure resulting from the lateral elements of long visual, centrifugal, monopolar and tangential fibres. Nine cell axons arising from each ommatidium leave the retina. Six of these are short visual fibres and end at two different levels in the lamina. Three different types of short visual fibres can be distinguished by their different terminal depths and lateral branching pattern. The remaining three fibres, the long visual fibres, terminate in the medulla. They can be distinguished from each other by their lateral elements in the lamina neuropile. The five morphologically different laminal fibre types (axons of the monopolar cells in the lamina) have different shapes and different arborizations at different levels. Tangential, centrifugal and incerta sedis-fibres, which originate either from cell bodies in the cell body layer at the periphery of the outer chiasma or more centrally, terminate in the synaptic region of the lamina. Consideration is given to the clearly demarkated arrangement and length of the branching pattern of retinal and laminal fibres at different levels of the synaptic region of the lamina. In addition, a hypothetical connectivity pattern is discussed.