Interspecific Trail Following and Commensalism Between the Ponerine Ant Gnamptogenys menadensis and the Formicine Ant Polyrhachis rufipes

Gnamptogenys menadensis (subfamily Ponerinae) foragers use chemical trails to home to their nests. Although prey capture and retrieval are generally performed solitarily, trails seem to enhance foraging to areas rich in prey or to sugar sources. Trail laying and following are most conspicuous during nest migration. These trails are laid down by tapping the sting onto the substrate. In laboratory tests, only extracts from the Dufour's gland were readily followed. Workers of Polyrhachis rufipes (Formicinae) use the trails of G. menadensis to gain access to otherwise nonavailable sugar sources. When they encounter Gnamptogenys foragers, P. rufipes workers show a typical aggressive antennal boxing, to which Gnamptogenys reacts with a submissive behavior. This is the first report of commensalism between a ponerine and a formicine ant.     

Task Allocation Depends on Matriline in the Ponerine Ant Gnamptogenys striatula Mayr

Matriline and the predominant social tasks performed by workers are correlated in the functionally polygynous ponerine ant Gnamptogenys striatula. This result favors the idea that polygyny might have been secondarily selected and maintained in ants because it provided more genetic variability and, thus, more potential variation in the regulation of the division of labor within mutualistic societies. As in previous studies on ants, nepotism could not be demonstrated. Because of the relatively small number of individuals per colony, these ponerine ants constitute a valuable model for exploring how polyethism is determined in insect societies.     

Male Behavior and Regulation of Worker Mating in a Ponerine Ant, Pachycondyla (Bothroponera) sp. (Hymenoptera: Formicidae)

Male behavior and regulation of worker mating were observed in Indonesian Pachycondyla sp. The colonies had some mated workers but only one gamergate (mated and egg-laying workers). Males frequently tried to mate with workers in their natal nests irrespective of worker age and dominance rank, however, workers never accepted mounting of nestmate males. Worker mating occurred only under the gamergateless condition with alien males.     

Production of trophic eggs by virgin workers in the ponerine ant Gnamptogenys menadensis

Abstract. Most colonies (thirty-five out of thirty-seven) of the ant Gnamptogenys menadensis (Mayr) (Hymenoptera: Formicidae: Ponerinae) lack queens. Mated workers produce reproductive eggs, whereas virgin workers can lay only smaller trophic eggs (350 ovipositions observed). These two egg types are morphologically distinct (e.g. in the pattern of oogenesis and ultrastructure of membranes and micropyle) and relate to different ovarian characteristics (ovariole length, number of yolky oocytes and yellow bodies). When reproductives are removed, a small number of virgin workers switch to producing reproductive eggs, although only 3% of these develop into larvae. Once workers are mated, up to 50% of their eggs develop further. Trophic eggs are generally absent in social insects lacking physical castes, and we review adaptive explanations of its occurrence in G. menadensis .     

Sting,Carry and Stock: How Corpse Availability Can Regulate De-Centralized Task Allocation in a Ponerine Ant Colony

We develop a model to produce plausible patterns of task partitioning in the ponerine ant Ectatomma ruidum based on the availability of living prey and prey corpses. The model is based on the organizational capabilities of a “common stomach” through which the colony utilizes the availability of a natural (food) substance as a major communication channel to regulate the income and expenditure of the very same substance. This communication channel has also a central role in regulating task partitioning of collective hunting behavior in a supply&demand-driven manner. Our model shows that task partitioning of the collective hunting behavior in E. ruidum can be explained by regulation due to a common stomach system. The saturation of the common stomach provides accessible information to individual ants so that they can adjust their hunting behavior accordingly by engaging in or by abandoning from stinging or transporting tasks. The common stomach is able to establish and to keep stabilized an effective mix of workforce to exploit the prey population and to transport food into the nest. This system is also able to react to external perturbations in a de-centralized homeostatic way, such as to changes in the prey density or to accumulation of food in the nest. In case of stable conditions the system develops towards an equilibrium concerning colony size and prey density. Our model shows that organization of work through a common stomach system can allow Ectatomma ruidum to collectively forage for food in a robust, reactive and reliable way. The model is compared to previously published models that followed a different modeling approach. Based on our model analysis we also suggest a series of experiments for which our model gives plausible predictions. These predictions are used to formulate a set of testable hypotheses that should be investigated empirically in future experimentation.     

Recruitment and Other Communication Behavior in the Ponerine Ant Ectatomma ruidum

The ponerine ant Ectatomma ruidum, though previously reported to possess only rudimentary recruitment ability, was found to lay chemical trails for mass recruitment to rich or difficult food sources. The pheromone originates from the Dufour's gland, a new source of trail pheromones in the primitive ant subfamily Ponerinae. During nest emigrations, E. ruidum practices stereotyped social carrying in the myrmicine mode. The discovery of this form of social carrying and of a recruitment pheromone in the Dufour's gland secretions support the hypothesis that the subfamily Myrmicinae is derived from an ectatommine ancestor. Other communication behaviors exhibited by E. ruidum include exchange of liquid food carried between the mandibles, chemical alarm communication, nest entrance marking, and an additional social carrying posture previously unknown in ants.     

Multiply Mating Males in Gnamptogenys striatula Mayr (Hymenoptera, Formicidae)

Multiply mating ant males are rare, because in most ant species, multiple mating opportunities for males are scarce. Gnamptogenys striatula is an exception to this rule, as intranidal mating offers protection and a local female-biased sex ratio. In a behavioral experiment, we confirm the males’ ability to inseminate several receptive workers, with four females being the observed maximum. D. Allard is a Research Assistant for the Fund of Scientific Research—Flanders.     

Behavioral and Chemical Correlates of Long-Term Queen Adoption in the Facultative Polygynous Ant Ectatomma tuberculatum

In ants, queen adoption is a common way of achieving secondary polygyny but the mechanisms involved are little known. Here we studied the process of long-term adoptions of alien queens in the facultative polygynous ant Ectatomma tuberculatum. In eight out of 10 successful adoption experiments, all the introduced queens showed similar behavior and fecundity as the resident queens even after 2 months, indicating complete integration into the colony. Chemical analysis revealed that the cuticular hydrocarbon profiles of resident and introduced queens were clearly distinct from those of workers and that they did not change after adoption. We propose that queen-specific cuticular hydrocarbon profile may represent a reliable signal of queen’s fertility and discuss about the evolution of high level of queen acceptance in E. tuberculatum.     

Functional Monogyny and Dominance Hierarchy in the Queenless Ponerine Ant Pachycondyla (=Bothroponera) sp. in West Java,Indonesia (Hymenoptera,Formicidae, Ponerinae)

Social organization of the queenless ponerine ant Pachycondyla (Bothroponera) sp. was studied in Bogor, West Java, Indonesia. Colonies had no morphologically distinct queens. Worker number per colony was 11 on average. Most colonies contained multiple mated workers (gamergates), however, only one of them laid eggs, whereas additional gamergates were sterile or ovipositing at a low rate, indicating that this ant species shows functional monogyny. Some virgin workers also developed their oocytes and actually laid eggs. Ritualized dominant — subordinate behavior was frequently observed and a linear dominance hierarchy was established among workers. The topranking individual was a fertile gamergate. Sterile gamergates positioned in lower ranks and they often engaged in foraging. Among virgin workers, callows and young workers monopolized higher ranks.     

Tandem Recruitment and Trail Laying in the Ponerine Ant Diacamma rugosum: Signal Analysis1

During nest emigration, Diacamma rugosum scouts recruit female nestmates to the new nest site by tandem running. ♂♂, brood and prey objects are carried. The tandem leaders stimulate their nestmates to follow them by pushing or, more rarely, pulling them with the mandibles. The signals releasing the following behaviour are mainly chemical. Only 15–30% of motivated followers react positively to an odourless paper or wax dummy. The ♀♀ follow cut-off thoraces, legs and gasters plugged at the end or without hind gut and sting apparatus, just as well as a leading ant. The ants even follow a dummy consisting of body parts if they are not in direct mechanical contact with it. The leading effect of the body surface is not reduced by intensive cleaning with water or by extraction in solvents of different polarity. Only maceration in potash lye eliminates the leading effectiveness of the cuticula. None of the secretions of the tested abdominal glands release following behavior. Only the hind gut fluid possesses leading signal qualities. No distribution of faeces over the bodies of the leading ants can be observed before or during tandem running. The faecal pheromone responsible for this leading effect does not seep into the superficial layer of a leader dummy made from bee wax but it can be washed off with water. These results show that the actual signal releasing tandem following is an unspecific odour substance of the cuticula surface. The presence of the hind gut fluid is not necessary for the tandem following response. It may be an additional signal. During nest emigration the hind gut fluid is scattered patchily in the form of small droplets by the nest finders on their way to the new nest. No increased deposition of faecal droplets could be observed on the way to repeatedly visited rich food sources. The hind gut fluid serves as a long lasting trail substance and has an exclusively orienting effect. The secretions of the other abdominal glands are not trail pheromones. The nest entrance is marked colony-specifically.     

Colony Budding and its Effects on Food Allocation in the Highly Polygynous Ant, Monomorium pharaonis

To advance our understanding of the causes and the consequences of budding (colony multiplication by fragmentation of main nests), we investigated nest movement in the facultatively polydomous Pharaoh ant, Monomorium pharaonis. Demographic data revealed that Pharaoh ants are highly polygynous and have a relatively low worker to queen ratio of 12.86. Budding experiments demonstrated that the number of available bud nests has a significant effect on colony fragmentation and increasing the number of bud nests resulted in smaller colony fragments. The overall distribution among bud nests was uneven, even though there was no evidence that the different life stages and castes partitioned unevenly among the bud nests and the analysis of individual colonies revealed no evidence of an uneven split in any of the colonies. This demonstrates that Pharaoh ants have the ability to exert social control over colony size and caste proportions during budding, which may contribute to their success as an invasive ant. The intensity of nest disturbance had a significant effect on whether or not the ants migrated into bud nests. Major disturbance resulted in the ants abandoning the source nest and migrating to bud nests and minor disturbance did not stimulate the ants to abandon the source nest. The results of the successive budding experiment which allowed the ants the opportunity to bud into progressively smaller nest fragments demonstrate that Pharaoh ants maintain a preferred minimum group size of 469 ± 28 individuals. Food allocation experiments utilizing protein marking revealed that nest fragmentation in Pharaoh ants has no negative impact on intracolony food distribution. Overall, our results suggest that nest units in the Pharaoh ant behave like cooperative, rather than competitive, entities. Such cooperation is most likely facilitated by the fact that individuals in all bud nests are genetically related, remain in close proximity to each other, and may continue to exchange individuals after budding.     

Cellular Interactions during the Mating Process in Chlamydomonas eugametos

A method to determine the mating competence of Chlamydomonas eugametos was developed. The contribution of each mating type in the pair formation was investigated using asymmetric gamete mixtures. It was established that pair formation is not mediated by a pheromonal attraction mechanism between partner gametes, but depends on collision chances. On the other hand, it was demonstrated that during transient contacts between partner gametes the flagellar agglutinability of both partners is stimulated, evidently to prepare a successful mating. The plus mating type was generally less agglutinable than the minus mating type and was a rate-limiting factor in the mating process.     

Dominance and Mating in a Communal Polygynandrous Bird: Cooperation or Indifference towards Mating Competitors?

The effect of dominance on the mating behaviour of both females and males was investigated for the communally breeding pukeko (Porphyrio porphyrio melanotus). Pukeko are polygynandrous with most groups consisting of three to 7 breeding males and one or two breeding females that lay eggs in a single nest. Linear dominance hierarchies are well established within breeding groups. Alpha females were involved in more copulations than beta females, but this difference did not appear to be a result of dominance status. There was also no evidence of egg tossing or egg destruction by breeding females. There was no positive correspondence between dominance rank of males and their frequency of sexual activity or copulations during the prelaying or laying periods. Dominants did interrupt matings by subordinates, but both qualitative and quantitative analyses of behaviour found no evidence of mate guarding. Furthermore dominants were no more likely to interrupt a copulation involving a subordinate male than they were to passively observe it or not react at all. This led us to suggest that males tended to behave indifferently toward mating competitors. As a result, multiple paternity is predicted to be high in pukeko groups. We also question the standard interpretation that males are behaving cooperatively in communal groups rather than coactively.     

Egg Cannibalism in a Facultative Polygynous Ant: Conflict for Reproduction or Strategy to Survive?

Policing, i.e. all behaviours that prevent a nestmate from reproducing, is currently observed in social insects. It is presumed to have evolved to regulate potential conflicts generated by genetic asymmetries or to enhance colony efficiency by avoiding surplus reproductives and brood. In the ant, Ectatomma tuberculatum, individual queen fecundity was similar in monogynous and polygynous colonies issued from a Mexican population. Egg cannibalism, however, occurred in the polygynous colonies. The stealing and destruction of reproductive queen‐laid eggs involved only nestmate queens, even if they were highly related. No queen appeared to monopolize reproduction in the polygynous colonies. But, the observed value of relatedness among workers differed from the expected value, suggesting an unequal sharing of reproduction between queens. We discussed whether the cannibalism of queen‐laid eggs in E. tuberculatum results from a competition for reproduction among queens or if this phenomenon is related to constraints on nutritional resources.     

Amelogenin-Collagen Interactions Regulate Calcium Phosphate Mineralization in Vitro

Collagen and amelogenin are two major extracellular organic matrix proteins of dentin and enamel, the mineralized tissues comprising a tooth crown. They both are present at the dentin-enamel boundary (DEB), a remarkably robust interface holding dentin and enamel together. It is believed that interactions of dentin and enamel protein assemblies regulate growth and structural organization of mineral crystals at the DEB, leading to a continuum at the molecular level between dentin and enamel organic and mineral phases. To gain insight into the mechanisms of the DEB formation and structural basis of its mechanical resiliency we have studied the interactions between collagen fibrils, amelogenin assemblies, and forming mineral in vitro, using electron microscopy. Our data indicate that collagen fibrils guide assembly of amelogenin into elongated chain or filament-like structures oriented along the long axes of the fibrils. We also show that the interactions between collagen fibrils and amelogenin-calcium phosphate mineral complexes lead to oriented deposition of elongated amorphous mineral particles along the fibril axes, triggering mineralization of the bulk of collagen fibril. The resulting structure was similar to the mineralized collagen fibrils found at the DEB, with arrays of smaller well organized crystals inside the collagen fibrils and bundles of larger crystals on the outside of the fibrils. These data suggest that interactions between collagen and amelogenin might play an important role in the formation of the DEB providing structural continuity between dentin and enamel.     

Genetic Interactions That Regulate Inflorescence Development in Arabidopsis

In Arabidopsis, floral meristems arise in continuous succession directly on the flanks of the inflorescence meristem. Thus, the pathways that regulate inflorescence and floral meristem identity must operate both simultaneously and in close spatial proximity. The TERMINAL FLOWER 1 (TFL1) gene of Arabidopsis is required for normal inflorescence meristem function, and the LEAFY (LFY), APETALA 1 (AP1), and APETALA 2 (AP2) genes are required for normal floral meristem function. We present evidence that inflorescence meristem identity is promoted by TFL1 and that floral meristem identity is promoted by parallel developmental pathways, one defined by LFY and the other defined by AP1/AP2. Our analysis suggests that the acquisition of meristem identity during inflorescence development is mediated by antagonistic interactions between TFL1 and LFY and between TFL1 and AP1/AP2. Based on this study, we propose a simple model for the genetic regulation of inflorescence development in Arabidopsis. This model is discussed in relation to the proposed interactions between the inflorescence and the floral meristem identity genes and in regard to other genes that are likely to be part of the genetic hierarchy regulating the establishment and maintenance of inflorescence and floral meristems.     

Queen Dominance May Reduce Worker Mushroom Body Size in a Social Bee

The mushroom body (MB) is an area of the insect brain involved in learning, memory, and sensory integration. Here, we used the sweat bee Megalopta genalis (Halictidae) to test for differences between queens and workers in the volume of the MB calyces. We used confocal microscopy to measure the volume of the whole brain, MB calyces, optic lobes, and antennal lobes of queens and workers. Queens had larger brains, larger MB calyces, and a larger MB calyces:whole brain ratio than workers, suggesting an effect of social dominance in brain development. This could result from social interactions leading to smaller worker MBs, or larger queen MBs. It could also result from other factors, such as differences in age or sensory experience. To test these explanations, we next compared queens and workers to other groups. We compared newly emerged bees, bees reared in isolation for 10 days, bees initiating new observation nests, and bees initiating new natural nests collected from the field to queens and workers. Queens did not differ from these other groups. We suggest that the effects of queen dominance over workers, rather than differences in age, experience, or reproductive status, are responsible for the queen–worker differences we observed. Worker MB development may be affected by queen aggression directly and/or manipulation of larval nutrition, which is provisioned by the queen. We found no consistent differences in the size of antennal lobes or optic lobes associated with differences in age, experience, reproductive status, or social caste.     

Trophallaxis and Aggression in the Ponerine Ant,Ponera coarctata: Implications for the Evolution of Liquid Food Exchange in the Hymenoptera

Trophallaxis (i.e. the exchange of alimentary liquid among colony members) plays a major role in the societies of many social Hymenoptera. Food is often not equally distributed among nestmates but instead is directed towards more dominant individuals by means of trophallaxis. Antagonistic behaviour can be associated with the exchange of food, or aggression may trigger the offering of food, even where social food exchange does not normally occur. In orphaned colonies of the ponerine ant, Ponera coarctata, workers interact aggressively at a high frequency. They establish hierarchies soon after the removal of the reproductive queen. One of the consequences of aggression among workers is trophallaxis, which also occurs regularly, although less frequently, in queenright colonies. The connection of trophallaxis and aggression in Ponera coarctata and in many other species of the Hymenoptera is discussed. This study and various other examples show that, besides the nutritional function of trophallaxis, the offering of food may often serve as an appeasement behaviour during aggressive interactions. We speculate that appeasing food offers may have provided the basis for the further evolution and elaboration of trophallaxis in many social Hymenoptera.