Ambush Predation by the Ponerine Ant Ectatomma ruidum Roger (Formicidae) on a Sweat Bee Lasioglossum umbripenne (Halictidae), in Panama

An individual ambush predation is used by huntresses of the ponerine ant Ectatomma ruidum to capture halictid bees (Lasioglossum umbripenne) in the Panamanian mountains. Workers, which use this strategy and originated from a colony (A) situated within a nest aggregation of halictid bees, capture almost four times more prey than colony (B) with a foraging area which does not include this type of bee's nest. Forty-eight percent of the prey of colony (A) are halictid bees, demonstrating the local importance of this predatory strategy in E. ruidum. A close examination of the behavioral sequence of predation shows that ambush is successful in only 4.8% of cases, a very low success rate in comparison to other predatory strategies described in this species. Nevertheless, due to the high repetition (16.1 ± 5.9 times on average) of very short waiting phases (6.3 ± 1.9 s), the true success rate (i.e., according to the number of individual trips) can reach up to 80%. A review of ambush predation among ant species allows us to distinguish approaches between jumping, long stealth approaches, and true ambushes. Implications of learning and expectation processes are also discussed.     

Polyethism within hunters of the ponerine ant,Ectatomma ruidum Roger (Formicidae,Ponerinae)

Summary During laboratory experiments, two categories of prey-foraging workers were found inEctatomma ruidum: stingers and transporters. When numerous live drosophila were offered to the ants, one group of hunters specialized in killing the prey and another in transporting simultaneously the dead drosophila to the nest. Sometimes, there was a transfer of prey by a stinger towards a transporter, after an active soliciting of the transporter by antennation or by using the forelegs. We found high positive correlations between the colony size and the number of ants in each subcaste. A negative correlation existed between the colony size and the proportion of hunters. However, the proportion of workers in the two behavioral subcastes of hunters was stable in spite of differences in colony size. The phylogenetic interest of this type of cooperative predation is discussed.     

Queen dimorphism and reproductive capacity in the ponerine ant, Ectatomma ruidum Roger

We report here the first case of queen dimorphism in a ponerine ant species. A total of 550 colonies of the Neotropical ponerine ant Ectatomma ruidum, from two natural populations in southeastern Mexico, were investigated for polymorphism and reproductive function within the queen caste. The distribution of different morphological traits (head and thorax widths, and scutum and alitrunk lengths) was shown to be bimodal. Thorax width and head width were significantly isometric for both macro- and microgynes and thorax proportions did not change with the category of the queen. Microgynes appear as an isometric reduction of the normal queens. On average, macrogynes were found to be approximately 20% larger in size and twice as heavy as microgynes (and up to three times heavier in terms of dry weight). Their wing surface was 72% greater. There was no difference between the two morphs in terms of their capacity to be inseminated and all dealate females present in natural colonies could be considered as true potentially reproductive queens whatever the class to which they belonged. Ovary size, number of ovarioles per ovary, and number of mature oocytes produced were significantly higher for macrogynes than for microgynes, but small queens were able to function as active egg-layers. However, the number of egg-laying individuals was significantly lower for this form, although egg-laying activity appeared to be independent of queen weight. Despite its markedly reduced reproductive capacity compared to macrogynes, the distribution of the microgyne form of E. ruidum in both studied populations was very broad, reaching one-third of all alate and dealate females and extending to one-third of all colonies. In 72.2% of the colonies where alate and/or dealate microgynes were present, both forms co-occurred. Fat content analysis results supported the idea that semi-claustral independent colony founding, typical for this species, would essentially be achieved by macrogynes, while microgynes would be adopted by established colonies. We argue that queen dimorphism in E. ruidum could represent an attractive alternative dispersal strategy for female sexuals. The small females may constitute an important reserve of potential reproductives at a very low energetic cost to the colony, the production of a macrogyne being about 7.5 times more costly than a microgyne. The significantly greater `wing surface/body weight' ratio of microgynes also suggests their greater capacity for dispersion. The combination of such a capacity for microgynes, along with their ability to contribute in the production of both female morphs, including the reproductively efficient macrogynes, would contribute to insuring genetic reassortment at the population level and could explain, in part, the ecological success of E. ruidum in Neotropical zones. Received: 4 January 1999 / Accepted: 12 May 1999     

Behavioural Variability and Development of Cataglyphis cursor Ant Workers (Hymenoptera,Formicidae)1)

Variability and development of behaviour among Cataglyphis cursor ant workers was analyzed, after identifying each member of the studied nests by individual marking. Behavioural differences between workers of the same nest were considerable, even when they were assigned to the same behavioural class defined with multifactorial analysis. Workers' behaviour also changed considerably over short-time intervals, for example from one day to another. Behavioural development was very variable. Only ? of the workers followed the classical age polyethism, while the remainder showed stable behaviour or irregular variations from the expected scheme.     

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.     

Recruitment Behavior in the Ant Genus Polyrhachis (Hymenoptera, Formicidae)

We present a detailed behavioral analysis of the signals involved in recruitment of 11 syntopic Polyrhachis species from West Malaysia. We found a considerable variety of recruitment techniques, including social carrying behavior, tandem running, group recruitment, and a technique which we call leader-independent trail communication. The latter mode superficially resembles chemical mass communication (sensu Wilson, 1962). All these recruitment techniques involve mechanical invitation behavior inside the nest, comprising back-and-forth jerking or pulling movement often combined with a sideways waggling. However, not in all cases of leader-independent trail communication is a mechanical invitation behavior obligatory. The trail pheromone of all investigated Polyrhachis species originates from the hindgut. Only in the tandem running P. proxima do additional secretions from the poison gland appear to be involved in tandem calling.     

Coordination of Raiding and Emigration in the Ponerine Army Ant Leptogenys distinguenda (Hymenoptera: Formicidae: Ponerinae): A Signal Analysis

Several glandular sources of trail pheromones have been discovered in army ants in general. Nevertheless, at present the understanding of the highly coordinated behavior of these ants is far from complete. The importance of trail pheromone communication for the coordination of raids and emigrations in the ponerine army ant Leptogenys distinguenda was examined, and its ecological function is discussed. The secretions of at least two glands organize the swarming activities of L. distinguenda. The pygidial gland is the source of an orientation pheromone holding the group of raiding workers together. The same pheromone guides emigrations to new nest sites. In addition, the poison sac contains two further components: one with a weak orientation effect and another which produces strong, but short-term attraction and excitement. The latter component is important in prey recruitment and characterizes raid trails. This highly volatile recruitment pheromone allows the extreme swarm dynamic characteristic of this species. Emigration trails lack the poison gland secretion. Due to their different chemical compositions, the ants are thus able to distinguish between raid and emigration trails. Nest emigration is not induced chemically, but mechanically, by the jerking movements of stimulating workers.     

Social Organization of Cataglyphis cursor Ant Colonies (Hymenoptera,Formicidae): Inter-, and Intraspecific Comparisons

The social organization of several colonies of the ant Cataglyphis cursor with different compositions has been studied by means of individual marking of workers and later treatment of data with correspondence analysis. There are always three main poles or groups of tasks: inactivity, unspecific tasks in the nest and outside activities. From them, three groups of workers are separated: inactives, which stay immobile in the nest a great part of their time; interiors, which perform various tasks in the nest; and foragers, that are engaged in activities outside the nest. Several factors, such as the presence of the queen, the colony size or the season of the year influence the social structure of C. cursor colonies and originate important inter-, and intracolonial variations.     

Polygyny and Monoandry in the Ant Formica japonica (Hymenoptera: Formicidae)

Queen number, mating frequency and nest kin-structure of the ant Formica japonica were studied in the field and the laboratory. Nest excavation in the study site, the east slope of Mt. Fuji, Gotenba, Japan, revealed that F. japonica is weakly polygynous all year round and the queen number increases after the nuptial flight season, suggesting the adoption of newly mated queens by established nests. Dissection and laboratory rearing demonstrated that nearly all queens in polygynous nests had mated and were fertile with mature oocytes in their ovaries. Multilocus DNA fingerprinting was used to examine kin relationships among ants found in the same nests. The fingerprint band patterns were apparently governed by a simple genetic rule and suggested monoandry (single mating per queen). The mean band sharing score of DNA fingerprints among full sisters was 0.90, and the mean value between queens and their daughters was 0.75. Comparison of DNA fingerprints of adult and pupal workers with pupal gynes suggested that multiple queens in a nest may contribute unequally to gyne (new queen) production.     

Individual and Cooperative Food Transport of the Red Imported Fire Ant (Hymenoptera: Formicidae): Laboratory Observations

The red imported fire ant, Solenopsis invicta Buren, is a significant urban and agricultural pest with a global distribution. However, the food transport patterns of S. invicta are poorly understood. In the present study, food of different sizes were provided to S. invicta colonies and transport for a short distance was observed under laboratory conditions. Individual transport of small food (1 × 1 mm deli meat) and cooperative transport of food of larger sizes (2 × 2 mm, 4 × 4 mm, and 8 × 8 mm deli meat) was observed. The majority of transporters were small workers. During cooperative transport, ants were either around or on the food, pulling/pushing the food item in multiple directions, leading to a slow process with frequent deadlocks. Individual transport required significantly shorter time in the preparation and transportation stages, but food transport was more likely to deviate from the nest. This observation suggests that the cooperative transport of S. invicta is closer to uncoordinated.     

Facultative polygyny in Ectatomma tuberculatum (Formicidae, Ectatomminae)

Summary. Polygyny, the presence of several mated queens within the same colony, is widespread in insect societies. This phenomenon is commonly associated with ecological constraints such as limited nest sites. In habitats where solitary nest foundation is risky, monogynous colonies can reintegrate young daughter queens (secondary polygyny). We studied the reproductive structure (i.e. queen number) of the ectatommine ant Ectatomma tuberculatum from Bahia State, Brazil. This species was found to present facultative polygyny: out of a total of 130 colonies collected, 39.2% were monogynous, while 43.8% were polygynous. Polygynous colonies had significantly more workers than monogynous ones. Queen number in polygynous colonies ranged from 2 to 26, with an average of 4 ± 4 queens per colony. All nestmate queens were egg-layers with no apparent dominance hierarchy or agonistic behavior. Non-nestmate queens were adopted by monogynous colonies suggesting that polygyny is secondary, originating through queen adoption. This species is characterized by an open recognition system, which probably allows a switch from monogynous to polygynous colonies. The behavioral acts of queens showed that resident queens remained frequently immobile on or near the brood, contrarily to alien or adopted queens and gynes. In addition, monogynous queens showed no behavioral or physiological (i.e. by ovarian status) differences in comparison with polygynous ones. Secondary or facultative polygyny, probably associated with queen adoption, may have been favored in particular environmental conditions. Indeed, by increasing colony productivity (i.e. number of workers) and territory size (by budding and polydomy), polygyny could uphold E. tuberculatum as a dominant species in the mosaic of arboreal ants in Neotropical habitats.Received 7 April 2004; revised 11 November 2004; accepted 15 November 2004.     

Thelytokous Parthenogenesis in the Fungus-Gardening Ant Mycocepurus smithii (Hymenoptera: Formicidae)

The general prevalence of sexual reproduction over asexual reproduction among organisms testifies to the evolutionary benefits of recombination, such as accelerated adaptation to changing environments and elimination of deleterious mutations. Documented instances of asexual reproduction in groups otherwise dominated by sexual reproduction challenge evolutionary biologists to understand the special circumstances that might confer an advantage to asexual reproductive strategies. Here we report one such instance of asexual reproduction in the ants. We present evidence for obligate thelytoky in the asexual fungus-gardening ant, Mycocepurus smithii, in which queens produce female offspring from unfertilized eggs, workers are sterile, and males appear to be completely absent. Obligate thelytoky is implicated by reproductive physiology of queens, lack of males, absence of mating behavior, and natural history observations. An obligate thelytoky hypothesis is further supported by the absence of evidence indicating sexual reproduction or genetic recombination across the species'' extensive distribution range (Mexico-Argentina). Potential conflicting evidence for sexual reproduction in this species derives from three Mycocepurus males reported in the literature, previously regarded as possible males of M. smithii. However, we show here that these specimens represent males of the congeneric species M. obsoletus, and not males of M. smithii. Mycocepurus smithii is unique among ants and among eusocial Hymenoptera, in that males seem to be completely absent and only queens (and not workers) produce diploid offspring via thelytoky. Because colonies consisting only of females can be propagated consecutively in the laboratory, M. smithii could be an adequate study organism a) to test hypotheses of the population-genetic advantages and disadvantages of asexual reproduction in a social organism and b) inform kin conflict theory.For a Portuguese translation of the abstract, please see Abstract S1.     

Trail Pheromone of the Argentine Ant,Linepithema humile (Mayr) (Hymenoptera: Formicidae)

The Argentine ant (Linepithema humile) is recognized as one of the world''s most damaging invasive species. One reason for the ecological dominance of introduced Argentine ant populations is their ability to dominate food and habitat resources through the rapid mobilization and recruitment of thousands of workers. More than 30 years ago, studies showed that (Z)-9-hexadecenal strongly attracted Argentine ant workers in a multi-choice olfactometer, suggesting that (Z)-9-hexadecenal might be the trail pheromone, or a component of a trail pheromone mixture. Since then, numerous studies have considered (Z)-9-hexadecenal as the key component of the Argentine ant trails. Here, we report the first chemical analyses of the trails laid by living Argentine ants and find that (Z)-9-hexadecenal is not present in a detectible quantity. Instead, two iridoids, dolichodial and iridomyrmecin, appear to be the primary chemical constituents of the trails. Laboratory choice tests confirmed that Argentine ants were attracted to artificial trails comprised of these two chemicals significantly more often than control trails. Although (Z)-9-hexadecenal was not detected in natural trails, supplementation of artificial dolichodial+iridomyrmecin trails with an extremely low concentraion of (Z)-9-hexadecenal did increase the efficacy of the trail-following behavior. In stark contrast with previous dogma, our study suggests that dolichodial and iridomyrmecin are major components of the Argentine ant trail pheromone. (Z)-9-hexadecenal may act in an additive manner with these iridoids, but it does not occur in detectable quantities in Argentine ant recruitment trails.     

Orientation and Communication in the Neotropical Ant Odontomachus bauri Emery (Hymenoptera,Formicidae, Ponerinae)

The Neotropical species Odontomachus bauri employs canopy orientation during foraging and homing. An artificial canopy pattern above the ants is much more effective as an orientation cue than horizontal landmarks or chemical marks. However, both horizontal visual cues and chemical marks on the ground can serve in localizing the nest entrance. Successful O. bauri foragers recruit nestmates to leave the nest and search for food. However, the recruitment signals do not contain directional information. Antennation bouts and pheromones from the pygidial gland most likely serve as stimulating recruitment signals. Secretions from the mandibular and poison gland elicit alarm and attack behavior.     

Alkylpyrazines: alarm pheromone components of the little fire ant, Wasmannia auropunctata (Roger) (Hymenoptera, Formicidae)

The previous identification of 2,5-dimethyl-3-(3-methylbutyl)pyrazine as the mandibular alarm pheromone of the little fire ant Wasmannia auropunctata (Roger), has been found to be incorrect. Gas chromatography–mass spectrometry (GC/MS) of ant extracts suggested the correct structure to be the regioisomer 2,5-dimethyl-3-(2-methylbutyl)pyrazine, which was confirmed by comparison with the synthetic pyrazine. GC/MS analysis also revealed the presence of an additional disubstituted alkylpyrazine which was identified as 3-methyl-2-(2-methylbutyl)pyrazine. Headspace sampling of confined ants with SPME and Porapak Q followed by GC/MS analysis showed 2,5-dimethyl-3-(2-methylbutyl)pyrazine as the major volatile released by W. auropunctata workers while 3-methyl-2-(2-methylbutyl)pyrazine was only detected in trace amounts. In laboratory bioassays, W. auropunctata workers were attracted and arrested by both pyrazines, although the results were not always consistent. Synthetic pyrazines generally attracted as many W. auropunctata workers as were attracted to a single crushed ant. However, higher numbers of W. auropunctata were arrested by crushed ant treatments than by synthetic pyrazines in all bioassays but one.     

Raiding and Foraging Behavior of the Blood-Red Ant,Formica sanguinea Latr. (Hymenoptera,Formicidae)

In this paper we report the results of a detailed study on the behavioral ecology of slave raiding and foraging activity in the European blood-red ant, Formica sanguinea Latr. The field study was conducted over an unbroken period of 78 days, during which the activity of two dulotic colonies of this facultative slave-maker was observed for 10 h each day. It was possible to observe 26 raids distributed over 23 days, among which 18 were followed by the sacking of nests belonging to the species F. cunicularia, F. fusca, and Lasius emarginatus, whereas 8 failed. Simple, continuous, and simultaneous raids occurred. We recorded the timing, frequency, distance, and direction of slave raids, including the number of participants and the type of booty. Particular attention was devoted to the scouting behavior and raiding organization. Moreover, every day, we observed foraging and predatory behavior, during which adult insects (mainly ants), seeds, and berries were retrieved to the dulotic colonies. On the basis of our observations F. sanguinea seems to be a very efficient slave-maker and predatory species of the Raptiformica subgenus. Moreover, its dulotic behavior may be regarded as a continuation and an expansion of its foraging and predatory behavior, as predicted by Darwin's hypothesis for the origin and evolution of slavery in ants.     

A Diverse Ant Fauna from the Mid-Cretaceous of Myanmar (Hymenoptera: Formicidae)

A new collection of 24 wingless ant specimens from mid-Cretaceous Burmese amber (Albian-Cenomanian, 99 Ma) comprises nine new species belonging to the genus Sphecomyrmodes Engel and Grimaldi. Described taxa vary considerably with regard to total size, head and body proportion, cuticular sculpturing, and petiole structure while all species are unified by a distinct shared character. The assemblage represents the largest known diversification of closely related Cretaceous ants with respect to species number. These stem-group ants exhibit some characteristics previously known only from their extant counterparts along with presumed plesiomorphic morphology. Consequently, their morphology may inform hypotheses relating to basal relationships and general patterns of ant evolution. These and other uncovered Cretaceous species indicate that stem-group ants are not simply wasp-like, transitional formicids, but rather a group of considerable adaptive diversity, exhibiting innovations analogous to what crown-group ants would echo 100 million years later.