Cuticular spectra in the slave-making ant Polyergus rufescens and the slave species Formica rufibarbis

Abstract. The relative proportions of cuticular components having the same retention times were compared between the slave-making ant Polyergus rufescens and the slave ant Formica rufibarbis living in monospecific or mixed colonies. The two species were found to present different spectra. The Formica workers, when enslaved by Polyergus , tend to lose their colony characteristics but they do not seem to adopt the characteristics of Polyergus.     

Cuticular spectra and inter-individual recognition in the slave-making ant Polyergus rufescens and the slave species Formica rufibarbis

Abstract. The relationship between behavioural tests and relative proportions of cuticular components were studied in the slave-making species Polyergus rufescens and the slave and Formica rufibarbis living in either monospecific or mixed colonies. A correlation between the relative proportions of the cuticular products and interindividual recognition exists in each of the two species Polyergus and Formica: Polyergus are fiercely aggressive towards individuals which have different cuticular spectra and originate from a geographically isolated nest. This seems to be true also in the case of Formica living in monospecific colonies. A similar correlation also exists between the two species, which have different cuticular spectra: encounters arranged between them show that free-living Formica are always fiercely aggressive towards Polyergus. The reason why no such correlation seems to exist, however, between Polyergus and Formica when the latter are enslaved and the two species coexist peacefully at the same nest is discussed.     

Regulation of cuticular and postpharyngeal hydrocarbons in the slave-making ant Polyergus rufescens: effect of Formica rufibarbis slaves

The purpose of this study was to compare cuticular hydrocarbon profiles of slave-making Polyergus rufescens ants reared alone or with their Formica rufibarbis slaves. Chemical analyses showed that due to the close contacts occurring when the Formica were tending the Polyergus, the synthesis of the cuticular hydrocarbons carried by the slaves was enhanced in the slave-makers. The postpharyngeal hydrocarbon levels increased during the first 15 days of life, whether or not the Polyergus were exposed to Formica. Our findings suggest that Polyergus is able to secrete all components of their cuticular hydrocarbon blend and that none are acquired through contact with Formica. In addition to presenting our experimental evidence, several hypotheses are proposed to explain the synthesis and regulation of hydrocarbon blends borne when these two species cohabitate within a single colony.     

Wingless reproductives in the slave‐making ant Polyergus samurai: specialized producers of males (Hymenoptera: Formicidae)

Three wingless reproductives of Polyergus samurai Yano were collected in the reproductive season. These wingless reproductives had developed ovaries and laid eggs immediately after collection. They lacked spermatheca and the eggs produced by wingless reproductives finally become adult males. These observations suggest that the wingless reproductives of P. samurai are potential producers of males.     

Integration raids in the Amazon ant Polyergus rufescens (Hymenoptera, Formicidae)

Summary. Groups of enslaved Formica fusca workers from mixed colonies of Polyergus rufescens with numerous slave workforce tend to split off and found small and almost homospecific nests around the main nest, with at least some of them connected with the latter with underground passages. Their inhabitants are able, at least temporarily, to adopt young F. fusca gynes. P. rufescens invades these satellite nests in a manner similar to the normal slave raids, and carries the slaves back to the main nest. The supposed evolutionary cause of this behaviour is to keep integrity of mixed colonies and prevent possible emancipation of slaves.Received 18 August 2004; revised 27 September 2004; accepted 11 October 2004.     

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.     

Decyl acetate synthesis in the ant,Formica schaufussi (Hymenoptera: Formicidae)

Decyl acetate occurs in high amounts in the Dufour's gland of Formica schaufussi. When worker ants were fed [¹⁴C]-acetate the acetate moiety of decyl acetate was labelled preferentially indicating that decyl acetate is synthesized via an esterification reaction.     

Population genetics of the black ant Formica lemani (Hymenoptera: Formicidae)

Colony kin structure and spatial population structure were studied in multiple populations of the ant Formica lemani , using allozymes and DNA microsatellites. Average genetic relatedness between nestmate workers varied little between populations ( r  = 0.51–0.76), indicating that the average colony kin structure was relatively simple. Worker genotypes could not be explained with a single breeding pair in all nests, however, and the distribution of relatedness estimates across nests was bimodal, suggesting that single- and multi-queen colonies co-occur. We studied spatial population structure in a successional boreal forest system, which is a mixture of different aged habitats. Newly clear-cut open habitat patches are quickly colonized by F. lemani , where it is able to persist for a limited number of generations. Newly-founded populations showed signs of a founder effect and spatial substructuring, whereas older populations were more homogenous. This suggests that new populations are founded by a limited number of colonizers arriving from more than one source. Genetic differentiation among local populations was minor, indicating strong migration between them. There were, however, indications of both isolation by distance and populations becoming more isolated as habitat patches grew older.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 97 , 247–258.     

Respiratory patterns in nurses of the red wood ant Formica polyctena (Hymenoptera, Formicidae)

Summary Cycles of discontinuous gas exchange (DGCs) and abdominal ventilatory movements were studied in nurses of red wood ant, Formica polyctena, using an electrolytic respirometer and an infra-red (IR) gas analyser or flow-through respirometry. Both respirometry systems were combined with an IR actographic device based on IR-emitting and IR-sensor diodes. After recovering from handling and apparatus stress, lasting 1–3 h, completely motionless intact ants displayed regular DGC. After decapitation the ants displayed DGC whose frequency was somewhat lower than that of the intact individuals (7.17 ± 0.79 mHz and 10.43 ± 01.12 mHz, respectively). In headless ants, there occurred continuous slow movements of legs. Bursts of carbon dioxide in the intact and in the headless ants always coincided with a bout of telescoping movements (contractions) of abdominal segments, which was interpreted as active ventilation. During interburst periods, the headless ants exhibited telescoping movements characterised by rapid protraction, lasting 0.07–0.09 s, followed by a slow retraction of segments, suggesting passive ventilation. The intact ants were very sensitive to the flowing air and tended to be continuously active during flow-through respirometry. The decapitated ants, on the contrary, were insensitive to the air current.Received 26 March 2003; revised 30 July 2003; accepted 6 August 2003.     

Intraspecific variation of cuticular hydrocarbon composition in Formica japonica Motschoulsky (Hymenoptera: Formicidae)

Cuticular hydrocarbons and morphological features were compared among 80 Formica japonica colonies collected in Japan. Although a few morphological differences were found in workers among the colonies, four different types of cuticular hydrocarbon composition were observed. This was supported by a principal component analysis. We further compared the cuticular hydrocarbons among a total of approximately 400 F. japonica colonies, and categorized the hydrocarbon components into four types based on the result of discriminant analyses for the first 80 colonies. Type 1 was observed in colonies mainly collected in southern Honshu, Shikoku, and Kyushu. Types 2, 3, and 4 were from colonies with primary collections in Southern Honshu, central and Pacific coast northern Honshu, and the Sea of Japan coasts of northern Honshu and Hokkaido, respectively. The occurrence of four distinct types of CHC composition suggests that the colonies that produce them are separate species.     

Responses of Two Potential Host Species (Formica gnava and Formica occulta) to Pupae of the Obligatory Slave-Making Ant, Polyergus breviceps (Hymenoptera: Formicidae)

Queens of the slave-making ant, Polyergus breviceps, take over nests of adult Formica workers when establishing new colonies. Although naïve to slave-maker brood, the usurped Formica rear Polyergus offspring and nests containing both host and parasite species forms. Host worker acceptance of parasite brood has been attributed to the similarity of brood tending signals between these closely related taxa and/or the presence of an attractive pheromone in the slave-maker brood. By presenting single-species groups of Formica occulta and Formica gnava (two host species of P. breviceps) with a choice of Formica pupae of both species or with a choice of P. breviceps pupae from both types of mixed-species nests, it seems that neither close phylogenetic relatedness nor an attractive brood odor alone can account for the propensity of host workers to adopt slave-maker pupae. Significantly greater numbers of P. breviceps pupae were adopted by enslaved workers than by free-living workers, and within the enslaved groups and the free-living F. gnava group, greater numbers of P. breviceps pupae were adopted if they were from nests where the host species was conspecific to workers used in tests. When presented with F. gnava and F. occulta pupae, Formica workers adopted conspecific pupae almost exclusively and ignored or consumed pupae of the other host species. Taken together, these results imply that P. breviceps pupae have both a species-specific odor and a general brood-tending pheromone, upon which a host odor may be imposed. The disparate requirements of immatures at different stages of development for cue specificity or generality in maintaining nest exclusivity and maximizing inclusive fitness are discussed.     

Adoption of lycaenid Niphanda fusca (Lepidoptera: Lycaenidae) caterpillars by the host ant Camponotus japonicus (Hymenoptera: Formicidae)

Caterpillars of the parasitic lycaenid butterfly are often adopted by host ants. It has been proposed that this adoption occurs because the caterpillars mimic the cuticular hydrocarbons of the host ant. This study aimed to examine whether caterpillars of the Japanese lycaenid butterfly Niphanda fusca induce adoption by mimicking their host ant Camponotus japonicus. Behavioral observations conducted in the laboratory showed that most second‐instar caterpillars were not adopted, whereas most third‐instar caterpillars were successfully adopted by host workers. A chemical comparison detected no characteristic differences in the cuticular hydrocarbon profiles between second‐ and third‐instar caterpillars. However, morphological features of the caterpillars differed between the second and third instars; third‐instar caterpillars developed exocrine glands (ant organs) such as tentacle organs and a dorsal nectary organ. These results suggest that multiple chemical signatures, not only cuticular hydrocarbons, may be important for invasion of the host ant nest.     

Nest structure and colony cycle of the Allegheny mound ant,Formica exsectoides Forel (Hymenoptera: Formicidae)

Summary The nest structure and colony cycle of a population of Allegheny mound ants,Formica exsectoides, were examined in central Michigan. The dispersion pattern of mounds was random. Nest structure and presence of brood were primarily determined by excavation of twenty-three nests over three intervals from June through September. Additional excavations of five nests in 1990 and ten nests in 1991 provided further details on nest structure and colony cycle. Most galleries occurred within the mound and upper 30 cm of soil, but some activity reached depths of 100 to 270 cm. Depth of nests showed little correlation with external measurements of height and diameter. Immature stages were recovered from two strata: the upper 20 cm of nest and mound and the lowest nest depths. Alate sexual forms were found in or near the mound in July, and numerous dealate queens were collected in September from peripheral galleries near the soil surface.     

Host nest usurpation and colony foundation in the European amazon ant,Polyergus rufescens Latr. (Hymenoptera: Formicidae)

Summary The socially parasitic mode of founding new colonies by queens of the European amazon antPolyergus rufescens was analysed in the laboratory. Newly-mated females of this obligatory slave-maker were individually introduced into queenright and queenless artificially established colonies of bothFormica cunicularia (the slave present in the natal dulotic nest) andF. rufibarbis (another potentialServiformica host). Particular attention was devoted to the behavioural patterns displayed by these young queens during the usurpation phases. Our observations, supported also by video-taping, show that the slave-making female, before laying her eggs, must penetrate the host colony, kill the resident queen, become accepted by the adult workers and appropriate the host brood. The parasite was almost always adopted in the colonies ofF. cunicularia, whereas in the presence ofF. rufibarbis it was generally killed in a short time. The failure in the attempt of usurping the colonies ofF. rufibarbis is discussed in relation to the host specificity typical of this slave-maker. Finally, egg-laying byPolyergus successful usurpers, the subsequent eclosion of the brood, and its complete social integration in the newly-established mixed colonies were also recorded.     

Raid process, activity pattern and influence of abiotic conditions in the slave-making ant Rossomyrmex minuchae (Hymenoptera, Formicidae)

Summary: In the present work, we present the results of a study on the seasonal and daily activity, as well as on the raid process in Rossomyrmex minuchae. A comparative study of the annual and daily activity of the host species, Proformica longiseta, has also been made in parasitized and non-parasitized colonies. R. minuchae is an obligate slave-maker, in which the annual activity is strongly related to abiotic conditions, especially temperature and moisture of the soil. The raid process is comprised of three phases: scouting, assault of the host nest and robbing of the host brood. Recruitment of nestmates by mean of adult transport, concentration close to and invasion of the host nest, are the three tasks carried out by R. minuchae workers during the assault phase. Major differences between Rossomyrmex and Polyergus (the other obligate dulotic genus of the tribe Formicini) in terms of behaviour, both in nestmate recruitment and in the host-nest invasion, reflect different parasitic strategies in the two genera, supporting the assignment of these two genera to different phylogenetic groups.     

Structural and functional changes of the Dufour gland in gynes of the amazon ant Polyergus rufescens (Hymenoptera, Formicidae)

Colony foundation by the slave-making amazon ant Polyergus rufescens requires penetration of the young gyne soon after mating into a colony of the slave species. During this process, she uses decyl butyrate from her large Dufour gland as an effective appeasement allomone. The structural appearance and development of this gland shows a clear age-dependent evolution that is in line with its behavioural function. At the moment of eclosion, young females have a gland with an empty lumen but thick lining epithelium, of which the active secretory cells are characterised by a well-developed Golgi apparatus, abundant mitochondria and smooth endoplasmic reticulum. The basal plasmalemma shows deep invaginations that facilitate the uptake of precursors from the haemolymph, while intercellular contacts display conspicuous interdigitations in the apical cell part. During the first days of adult life, secretion starts to accumulate in the lumen. At the age of 3 weeks, the gland displays a large lumen filled with secretion and a thin epithelium that no longer displays secretory activity. By this time, the gynes are ready for penetration into a slave colony, being loaded with large quantities of the appeasement allomone that are necessary at this initial stage of the usurpation process. Accepted: 13 March 2001