Queen acceptance and the complexity of nestmate discrimination in the Argentine ant

In most social insect species, individuals recognize and behave aggressively towards non-nestmate conspecifics to maintain colony integrity. However, introduced populations of the invasive Argentine ant, Linepithema humile, exhibit pronounced variation in intraspecific aggression denoting diversity in nestmate recognition behavior, which possibly shapes their social structure and the varying levels of unicoloniality observed among these populations. One approach to better understand differential aggression behaviors towards conspecifics and recognition cue perception and response in L. humile is to examine variation in nestmate discrimination capability among genetically distinct colonies under different social contexts. Consequently, we investigated the dynamics of queen and worker recognition in southeastern US L. humile queenless and queenright colonies by measuring rates of non-nestmate worker and queen adoption and intercolony genetic similarity. Aggression levels between colony pairs differed and were associated with non-nestmate worker, but not queen adoption. Adoption of queens and workers was a function of host colony origin, while colony queen number affected adoption of queens, but not workers, with queens more readily accepted by queenless hosts. Fecundity of adopted non-nestmate queens was comparable to that of rejected non-nestmate and host colony queens, suggesting that queen fecundity did not affect adoption decisions. Genetic similarity between colonies ranged from 30 to 77% alleles shared, with more genetically similar colonies showing lower levels of intraspecific aggression. Non-nestmate queens and workers that were more genetically similar to host colony workers were more likely to be adopted. We provide the first evidence for the role of L. humile colony queen number on queen discrimination and suggest an effect of resident queens on worker conspecific acceptance thresholds. Our findings indicate a role for genetically based cues in L. humile nestmate recognition. However, subtle discrimination capability seems to be influenced by the social context, as demonstrated by more frequent recognition errors in queenless colonies.     

Mixed messages: fertility signaling interferes with nestmate recognition in the monogynous ant Camponotus floridanus

Chemical communication is crucial for the organization of social insect colonies. However, with the heavy use of one communication modality, problems may arise such as the interference of different types of information. This study investigated how information about fertility and colony membership is integrated in the ant Camponotus floridanus. We introduced into mature, queenright colonies (a) the nestmate queen, (b) a nestmate worker, (c) a foreign, high-fertility queen, (d) a foreign, low-fertility queen, and (e) a foreign worker. As expected, workers did not attack their nestmate queen or a nestmate worker but responded aggressively to foreign workers and foreign, low-fertility queens. Surprisingly, workers did not attack foreign, high-fertility queens. Chemical analysis demonstrated that the cuticular hydrocarbon profile of C. floridanus encodes information about fertility status in queens and workers and colony membership in workers. We suggest that ants respond to this information in the cuticular hydrocarbon profile: individuals with strong fertility signals are accepted regardless of their colony membership, but individuals without strong fertility signals are tolerated only if their cuticular hydrocarbon profile matches that of colony members. Learning how social insects respond to multiple types of information presented together is critical to our understanding of the recognition systems that permit the complex organization of social insect colonies.     

Queen primer pheromone affects conspecific fire ant (<Emphasis Type="Italic">Solenopsis invicta</Emphasis>) aggression

Monogyne fire ant, Solenopsis invicta, colony workers are territorial and are aggressive toward members of other fire ant colonies. In contrast, polygyne colony workers are not aggressive toward non-nestmates, presumably due to broader exposure to heritable and environmentally derived nestmate recognition cues (broad template). Workers from both monogyne and polygyne fire ant colonies execute newly mated queens after mating flights. We discovered that monogyne and polygyne queens have a remarkable effect on conspecific recognition. After removal of their colony queen, monogyne worker aggression toward non-nestmate conspecifics quickly drops to merely investigative levels; however, heterospecific recognition/aggression remains high. Queenless monogyne or polygyne worker groups were also not aggressive toward newly mated queens. Queenless worker groups of both forms that adopted a monogyne-derived newly mated queen became aggressive toward non-nestmate workers and newly mated queens. We propose that the powerful effect of fire ant queens on conspecific nestmate recognition is caused by a queen-produced recognition primer pheromone that increases the sensitivity of workers to subtle quantitative differences in nestmate recognition cues. This primer pheromone prevents the adoption of newly mated queens (regulation of reproductive competition) in S. invicta and when absent allows queenless workers to adopt a new queen readily. This extraordinary discovery has broad implications regarding monogyne and polygyne colony and population dynamics.     

Do these eggs smell funny to you?: an experimental study of egg discrimination by hosts of the social parasite Polyergus breviceps (Hymenoptera: Formicidae)

Social parasites exploit the behaviours of other social species. Infiltration of host systems involves a variety of mechanisms depending on the conditions within the host society and the needs of the social parasite. For many species of socially parasitic ants, colony establishment entails the usurpation of colonies of other species. This frequently involves the eviction or death of the host colony queen and the subsequent adoption of the invading queen. The social parasite queen achieves host worker acceptance by either manipulating the nest-mate recognition processes of the host or undergoing chemical modification. Little is known, however, about how host workers respond to social parasite eggs or whether host species defend against brood parasitism during parasite invasions. Host species are believed to adopt social parasite offspring because the recent common ancestry between many social parasites and their hosts may grant the sharing of certain characteristics such as chemical cues. Use of multiple host species, however, suggests other processes are needed for the social bond between host and parasite young to form. This study reports the findings of adoption bioassays in which eggs from a slave-maker ant, Polyergus breviceps, were offered to workers of two of its host species from unparasitised or newly parasitised nests to determine whether P. breviceps eggs generally elicit rearing behaviours from multiple host species. Comparisons of parasite egg survival until adulthood with conspecific egg survival reveal that workers of both host species, free-living or newly enslaved, do not typically accept slave-maker eggs. Both host species thus have sufficient discriminatory power to reject social parasite eggs although our hydrocarbon analysis indicates parasite eggs may be adapted to their local host species. Combined these results suggest that host rearing of P. breviceps eggs may reflect an evolutionary equilibrium that is maintained by probability and cost of recognition errors.Communicated by L. Sundström     

Multiple mating and facultative polygyny in the Panamanian leafcutter ant Acromyrmex echinatior

Queen mating frequency of the facultatively polygynous ant Acromyrmex echinatior was investigated by analysing genetic variation at an (AG)_n repeat microsatellite locus in workers and sexuals of 20 colonies from a single Panamanian population. Thirteen colonies were found to be monogynous, 5 colonies contained multiple queens, whereas the queen number of 2 colonies remained unresolved. Microsatellite genotypes indicated that 12 out of 13 queens were inseminated by multiple males (polyandry). The mean queen mating frequency was 2.53 and the mean genetically effective paternity frequency was 2.23. These values range among the highest found in ants, and the results are in keeping with the high mating frequencies reported for other species of leafcutter ants. Consistent skew in the proportional representation of different patrilines within colonies was found, and this remained constant in two consecutive samples of offspring. Dissections showed that all examined queens from multiple-queen colonies were mated egg-layers. The mean relatedness value among nestmate workers in polygynous colonies was lower than that for monogynous colonies. No diploid males were detected in a sample of 70 genotyped males. Worker production of males was detected in one queenless colony. We discuss our findings in relation to known patterns of multiple maternity and paternity in other eusocial Hymenoptera. Received: 2 September 1998 / Received in revised form: 3 February 1999 / Accepted: 7 February 1999     

Sneak in or repel your enemy: Dufour''s gland repellent as a strategy for successful usurpation in the slave-maker Polyergus rufescens

Summary. The dulotic queen ant, Polyergus rufescens, must first penetrate a host colony and kill the resident queen in order to successfully founding a new colony. Successful usurpation by a newly mated queen predictably depends on a dual strategy. Although, it can sneak in by being “chemically insignificant” with respect to cuticular hydrocarbons, it may also need to deter prospective host-worker aggressors. Chemical analysis of Dufour's gland secretion of P. rufescens queens and workers by GS/MS revealed that queen secretion is typified by esters of butanoic acid and acetic acid, of which decyl butanoate comprises over 80%. Butanoates and acetates are also present in the workers' secretion, but these are of higher molecular weight, and octadecyl butanoate represents the major compound. Using synthetic mixtures of queen and worker Dufour's gland, we tested the hypothesis that these secretions modify the aggressive behavior of the host species Formica cunicularia>. The queen-like synthetic mixture significantly reduced aggression of the host workers towards alien conspecifics, but neither pentane nor the worker-like synthetic mixture showed this effect. Although Dufour's gland content of >Polyergus queens was suggested to function as an appeasement pheromone (Topoff et al. 1988; Mori et al. 2000), we hypothesized that it may in fact act as a repellent. In order to test this hypothesis we exposed starved F. cunicularia workers to a droplet of honey on a glass slide applied with one of the following compounds: decyl butanoate (queen major compound), octadecyl butanoate (worker main compound), limonene (a reported ant repellent), and pentane (solvent control). Of these, the workers were repelled only by the decyl butanoate and did not approach the honey. We conclude that during usurpation the queen actively repels aggressive workers by emitting Dufour's gland repellent, comprising the alternative tactic in the usurpation dual strategy. This represents another chemical weapon in the diverse arsenal used by parasites to overcome the host's resistance. Received 7 April 2000; accepted 17 May 2000     

Nestmate recognition in the ant Cataglyphis niger : do queens matter?

This study compares two basic models for the origin and maintenance of colony gestalt odor in the polygynous ant species Cataglyphis niger. In the first model, queens are centers of de novo biosynthesis and distribution of recognition odors (“queen-centered” model); in the second, colony odors are primarily synthesized and distributed by workers (“worker-centered” model). We tested the behavioral patterns that are predicted from each model, and verified by biochemical means the distributional directionality of these signals. Encounters between nestmates originating from split colonies were as amicable as between nestmates from non-split colonies; queenless ants were as aggressive as their queenright nestmates, and both were equally aggressed by alien ants. These results indicate that queens have little impact on the recognition system of this species, and lend credence to the worker-centered model. The queen-centered model predicts that unique queen substances should be produced in appreciable quantities and that, in this respect, queens should be more metabolically active than workers. Analysis of the chemical composition of postpharyngeal glands (PPGs) or cuticular extracts of queens and workers revealed high similarity. Quantitatively, queens possessed significantly greater amounts of hydrocarbons in the PPG than workers, but the amount on the thoracic epicuticle was the same. Queens, however, possess a lower hydrocarbon biosynthesis capability than workers. The biochemical evidence thus refutes the queen-centered model and supports a worker-centered model. To elucidate the directionality of cue distribution, we investigated exchange of hydrocarbons between the castes in dyadic or group encounters in which selective participants were prelabeled. Queens tended to receive more and give less PPG content, whereas transfer to the epicuticle was low and similar in all encounters, as predicted from the worker-centered hypothesis. In the group encounters, workers transferred, in most cases, more hydrocarbons to the queen than to a worker. This slight preference for the queen is presumably amplified in a whole colony and can explain their copious PPG content. We hypothesize that preferential transfer to the queen may reflect selection to maintain her individual odor as close to the average colony odor as possible. Received: 4 November 1997 / Accepted after revision: 5 February 1998     

Hierarchical perception of fertility signals and nestmate recognition cues in two dolichoderine ants

In social insects, queens are likely to “honestly” inform their nestmates of their presence and fertility status through pheromonal communication. Cuticular hydrocarbons (CHCs) have been reported to be effective nestmate discriminators and strongly suspected to act as fertility signals, at least in some species. The use of the same chemical bouquet (i.e., the CHC profile) to convey two fundamentally different information seems puzzling. However, a recent threshold model proposes a hierarchy in the discriminating processes, i.e., fertility signals can only be perceived if nestmate recognition has been reached (Le Conte and Hefetz, Annu Rev Entomol 53:523–542, 2008). Here, we developed a simple behavioral bioassay based on chemical recruitment toward a queen placed outside the nest in two dolichoderine ants (Linepithema humile and Tapinoma erraticum), which allowed us to investigate the interplay between fertility signaling and colonial recognition. Using queen corpses of various origins (nestmates or aliens) and physiological states (fertile or infertile; mated or unmated), we demonstrated that nestmate recognition cues clearly override fertility signals under our experimental conditions. Indeed, while nestmate infertile queens were largely ignored by the workers, nestmate fertile queens (mated or not) induced worker recruitment, whereas alien fertile queens did not and were aggressed by the workers.     

Experimental evidence that workers recognize reproductives through cuticular hydrocarbons in the ant Odontomachus brunneus

Eusociality is characterized by a reproductive division of labor, wherein workers respond to the presence of reproductive individuals by refraining from reproduction themselves and restricting the reproductive efforts of others. Our understanding of how eusociality is maintained therefore depends on characterizing the mechanism by which workers detect the presence of a reproductive. Variations in cuticular hydrocarbons correspond to changes in reproductive ability in ants, and experimental studies are beginning to reveal the function of hydrocarbons as signals. In this study, we compare the cuticular hydrocarbon profiles of dominant and reproductive workers and queens of the ant Odontomachus brunneus with profiles of non-reproductive workers. Using split/reunification tests we document the existence of worker policing in both queenless and queenright colonies; supernumerary reproductives were treated aggressively by nestmates. Finally, we induce aggression and replicate queen-like submissive nestmate responses by supplementing the hydrocarbon profile of workers with (Z)-9-nonacosene, a compound that was significantly more abundant on the cuticles of reproductives. In three bioassays, we compare this manipulation to various control manipulations of the hydrocarbon profile and demonstrate that workers gauge the reproductive activity of nestmates through changes in their cuticular hydrocarbon profiles.     

The kin recognition system of carpenter ants (Camponotus spp.)

Summary The phenotypic cues that provide for rejection of non-nestmates by workers of the ant genus Camponotus could derive from any or all of four sources: (1) environmental odors; (2) the individual's own genetically-determined recognition pheromones or discriminators; (3) a gestalt or mixture of transferable discriminators, produced by each nestmate and absorbed by all; and/or (4) the discriminators of the queen applied to all nestmates. To test these hypotheses, four series of small experimental colonies were created: inter- and intraspecific mixed colonies containing queens, queenless worker groups, and pairs of worker groups between which a single queen was repeatedly switched. Intraspecific mixed colonies and queenless groups were further divided into groups receiving different diets. Aggression of workers in 165 experimental colonies was assayed in a total of 4064 neutral arena tests. Workers adopted into inter- and intraspecific mixed colonies with queens were highly aggressive to unfamiliar kin from pure colonies, independent of diet and of the proportion of different kin groups in the colony. However, queenless workers exhibited less aggression to unfamiliar kin than to non-kin, demonstrating the existence of worker discriminators. Diet differences slightly enhanced aggression among unfamiliar queenless kin. Non-kin sharing a switched queen were as unaggressive to one another as were sisters. The ability to adopt queenless workers between colonies gradually declined over 1–2 wks following their emergence from pupae. We propose a hierarchy of importance of cue sources in determining nestmate discrimination in small Camponotus colonies: Queen discriminators> worker discriminators>environmental cues. A flow-diagram model of social insect kin recognition, based on the phenotype matching concept of Holmes and Sherman (1983), is discussed.     

Co-evolution-driven cuticular hydrocarbon variation between the slave-making ant Rossomyrmex minuchae and its host Proformica longiseta (Hymenoptera: Formicidae)

Summary. Proformica longiseta exists as two populations in the Sierra Nevada Mountains in Spain, only one of which is parasitized by the slave-maker ant Rossomyrmex minuchae. To investigate the possible effect of co-evolutionary pressures on cuticular hydrocarbon (CHC) profiles (the presumed nestmate recognition cues), we performed a comparative analysis of the CHC of R. minuchae and P. longiseta colonies from both the allopatric and sympatric populations; the latter includes samples of enslaved as well as free-living workers. Discriminant analyses based on these chemical profiles showed two clear profile groups: the first comprised R. minuchae and both enslaved and free-living P. longiseta from the sympatric population; and the second the allopatric P. longiseta workers. As expected, the profiles of the two sympatric P. longiseta groups (enslaved and free-living) were distinct; but, interestingly, those of the enslaved P. longiseta and its parasite R. minuchae were also distinguishable. This indicates that despite their cohabitation each species maintains its own chemical identity. Profile similarity between the sympatric free-living P. longiseta and its parasite may explain the lower than expected aggression observed during raids. We further speculate that in view of the differences between the sympatric and allopatric population of P. longiseta, co-evolutionary pressures have driven changes in the profile of the former to better match that of its parasite R. minuchae. Such an adjustment may have enabled nests of the sympatric P. longiseta to endure multiple raids by the parasite (due to the reduced aggression) and thus to continue to reproduce despite the damage inflicted by the raids.     

Ontogeny of nestmate recognition cues in the red carpenter ant (Camponotus floridanus)

Summary A combination of behavioral and chemical analyses was used to investigate the nature of nestmate recognition cues and the effects of worker age and social experience on these cues in the ant Camponotus floridanus. Five categories of workers were tested: foragers, 5-day old and 0-day old callows, 5-day old and 0-day old naive callows. Bioassays consisted of introductions of dead workers from these categories into their own colonies or into an alien colony after the following treatments: 1) killed by freezing, 2) solvent-washed, 3) solvent-washed and coated with a nestmate soak, 4) solvent-washed and coated with a non-nestmate soak. Soaks were obtained from individual ants immersed in hexane and were applied individually to washed workers from the same category. Soaks were analyzed by gas chromatography (GC) and compared by multivariate analyses. Freeze-killed workers from each category elicited more aggressive behavior in the alien colony than in its own. By comparing GC profiles, a worker from any category can be assigned to its colony of origin. Thus all studied worker categories are colony-specific. Solvent-washed ants did not induce more aggressive behaviors in the alien colony than in their own, but they induced significantly less aggressivity in an alien colony than non-treated dead ants from the same category. Washed ants indced more aggressive behaviors when coated with a soak from a different colony as opposed to a soak from the colony in which they were introduced. The combination of behavioral and chemical results lead to the following conclusions: 1) Information contained in soak derived from workers was sufficient to allow nestmate recognition. 2) Nestmate recognition cues, and consequently the recognition response displayed to their bearer, change with age. 3) Social experience is necessary to develop or acquire a colony-specific label. The role of age and social experience on nestmate recognition in social Hymenoptera is discussed.     

Food influence on colonial recognition and chemical signature between nestmates in the fungus-growing ant <Emphasis Type="Italic">Acromyrmex subterraneus subterraneus</Emphasis>

Summary. Neotropical Fungus-growing leaf-cutting ants (tribe Attini) live in obligatory symbiosis with a fungus, which they grow on fresh leaves harvested by workers. Colonial recognition is likely based on chemical cues provided by cuticular hydrocarbons that have been found to be partly influenced by environmental odor sources. The diet breadth of Acromyrmex subterraneus subterraneus enabled us to test the impact of different plant diets on colonial recognition. The intermediary of the fungus in the ants feeding habit adds a special angle to the question. From a queenright (QR) mother colony of A. s. subterraneus we formed several groups of queenless (QL) workers with fungus (approx. 700 ants). The QR colony and two of the QL-groups were fed with the same diet of fresh bramble leaves. Two other QL-groups were fed with privet leaves and two with rose flowers. After 4 months, QR workers were significantly more aggressive towards the QL-group fed with rose flowers or privet leaves than towards workers of the QL-groups fed with fresh bramble leaves. Rose-fed QL workers were aggressive towards privet-fed QL workers and vice versa, but never towards workers of their counterpart group that fed on the same diet. These results suggest that the absence of the queen or the separation time between groups played a minor role in shaping nestmate recognition cues as compared to the diet. The behavioral studies were supplemented by chemical analyses of cuticles, postpharyngeal glands (PPG) and plant-food extracts revealing profiles variations that were correlated with the dietary changes. However, although the plant extract contained several hydrocarbons there was no congruency between the plant profile and the respective diet-group ants. These results support the hypothesis that the diet influences indirectly the chemical profiles and consequently the recognition cues in A. s. subterraneus.     

Two independent mechanisms of egg recognition in worker <Emphasis Type="Italic">Formica fusca</Emphasis> ants

Informational constraints can be an important limitation on the accuracy of recognition. One potential constraint is the use of recognition information from the same sources in multiple discriminatory contexts. Worker wood ants, Formica fusca, discriminate eggs based on their maternal sources of origin in two main contexts: recognition of eggs laid by nestmate versus non-nestmate queens and recognition of worker-laid versus queen-laid eggs. We manipulated the experience of F. fusca workers in laboratory colonies to both worker-laid and queen-laid eggs by transferring eggs between colonies in order to investigate whether these two contexts of egg discrimination are independent. Experience of non-nestmate queen-laid eggs significantly increased worker acceptance of both familiar (18% accepted) and unfamiliar (10%) queen-laid eggs compared to control workers without experience of eggs other than those laid by their own colony’s queen (2%). In contrast, worker acceptance of worker-laid eggs was not affected by variation in the egg experience of workers (14% in workers from control colonies exposed only to eggs from their own colony’s queen versus 19% and 17% in workers from colonies which had received eggs laid by either a non-nestmate queen or nestmate workers, respectively). Our results suggest that these two recognition contexts do not strongly constrain each other and are different in their ontogeny. In particular, worker-laid eggs are universally discriminated against by workers from colonies with a queen whatever the egg experience of the workers, while non-nestmate queen-laid eggs are strongly discriminated against only by workers without experience of eggs laid by more than one queen.     

Colony specificity of Dufour gland secretions in a functionally monogynous ant

Summary. Queens in colonies of the small myrmicine ant, Leptothorax gredleri Mayr 1855 (Hymenoptera, Formicidae) engage in dominance interactions and form social hierarchies, in which typically only the top-ranking queen lays eggs. Occasionally, queen antagonism escalates to violent mandible fighting, during which the sting is used to apply Dufour gland secretions onto the cuticle of the opponent. Contaminated queens often are attacked by nestmate workers. Here we show that the chemical composition of the Dufour gland is colony-specific and that workers can discriminate between secretions from their own and other colonies. Our findings suggest that Dufour gland secretions are involved in the establishment of hierarchies within a colony. When invading an alien colony the queen presumably employs the secretions during the expulsion of the resident queen. Apparently, Dufour gland secretions play a role in intraspecific queen competition similar to that in slave-making and inquiline formicoxenine ants, where they function as "propaganda substances" in an interspecific context. Received 7 July 1998; accepted 15 September 1998.     

Worker self-restraint and policing maintain the queen’s reproductive monopoly in a pseudomyrmecine ant

Division of reproductive labor in insect societies is often based on worker self-restraint and both queen and worker policing. Workers of many hitherto studied wasps, bees and ants do not lay eggs in the presence of a queen. However, it is presently unclear how far these observations in a few select clades can be generalized. We investigated if and how queens maintain a reproductive monopoly in colonies of the elongate twig ant, Pseudomyrmex gracilis, a member of the previously unstudied ant subfamily Pseudomyrmecinae. Colonies are usually headed by a single, singly mated queen (monogyny, monandry). Workers therefore would be more closely related to males produced by other workers (r?=?0.375) than to the sons of queens (r?=?0.25). Nevertheless, workers appear to refrain from laying male-destined eggs in the presence of the queen. In queenless conditions, workers form dominance hierarchies by antennal boxing, and only one or a few high-ranking individuals readily begin to lay eggs. When returned into a queenright colony, egg-laying workers are immediately bitten, stung and expelled or killed by other workers. While the composition of cuticular hydrocarbons clearly differed between castes, it less clearly reflected worker ovarian development. An association with worker ovarian development that would allow workers to monitor the reproductive status of nestmates could only be tentatively postulated for certain substances. Our study broadens our knowledge about reproductive conflict in social Hymenoptera and shows that worker sterility in the presence of a queen is more common in monogynous, monandrous ants than expected from relatedness alone.     

Chemical mimicry in the root aphid parasitoid Paralipsis eikoae Yasumatsu (Hymenoptera: Aphidiidae) of the aphid-attending ant Lasius sakagamii Yamauchi & Hayashida (Hymenoptera: Formicidae)

Summary. An aphidiid wasp, Paralipsis eikoae, was associated with both Lasius niger and L. sakagamii attending the wormwood root aphid Sappaphis piri. An L. sakagamii worker was observed carrying a winged female P. eikoae to its nest with its mandible, but it did not kill the wasp. Once accepted by the ants, the wasp often mounted and rubbed against the worker ants and sometimes teased them to regurgitate food to itself. No workers in the colony attacked the wasp. Conspecific foreign workers, however, viciously attacked the wasp when encountered. Gas chromatography-mass spectrometry analyses showed that the accepted wasp had complex cuticular hydrocarbons that were very similar to those of its host ants, whereas the winged wasps collected outside the ant nest showed only a series of n-alkanes. Additionally, the accepted wasp had a hydrocarbon profile closer to that of its host ants than to the conspecific foreign ants. We believe the wasp mimics ant cuticular hydrocarbons to integrate into the ant nest, acquiring the hydrocarbons by mounting and rubbing against the ants. In contrast, the cuticular hydrocarbons of the emerged wasp contained larval and pupal hydrocarbons of L. sakagamii that were also similar to those of L. niger. Both ant species rejected adult workers of the other species but accepted their larvae and pupae. We suggest that the emerged P. eikoae mimics the cuticular hydrocarbons of these Lasius larvae and pupae, which allows P. eikoae to be accepted by both L. sakagamii and L. niger. Received 11 March 1998; accepted 22 July 1998.     

Interspecific aggression in colonies of the slave-making ant Harpagoxenus sublaevis

Colonies of the slave-making ant, Harpagoxenus sublaevis, may simultaneously contain workers of several Leptothorax slave species. We observed aggressive interactions among slave-makers, between slavemakers and slaves, and among slaves in 11 mixed colonies. The first two types of aggression appear to be correlated with reproductive competition for the production of males. Aggressive interactions among slaves, however, occurred mainly between slaves belonging to different species. In two colonies, in which one slave species clearly outnumbered the other, the majority attacked and finally expelled all nestmates belonging to the minority species. Our observations thus suggest that in Harpagoxenus colonies a homogeneous colony odor is not always achieved and that heterospecific slaves may occasionally be mistaken for alien ants. Gas chromatographic analyses of ants from mixed colonies similarly show that cuticular hydrocarbon profiles may differ strongly between heterospecific nestmate slaves.     

Prior experience with eggs laid by non-nestmate queens induces egg acceptance errors in ant workers

We studied the effect of prior experience to eggs laid by nestmate and non-nestmate queens on the acceptance of queen-laid eggs by worker wood ants, Formica fusca. We transferred eggs from a non-nestmate queen into colonies during early spring, when their own queen was recommencing egg laying. A few weeks later, workers from these “experienced” colonies accepted eggs of both familiar (44% acceptance) and unfamiliar (40%) non-nestmate queens much more than workers from control colonies (2%) that had only had previous contact with their own queen’s eggs. Thus, prior exposure to eggs laid by a non-nestmate queen induces much greater acceptance of all non-nestmate queen-laid eggs. Mechanistically, we hypothesize that exposure to eggs from several queens may increase acceptance by causing a highly permissive acceptance threshold of non-nestmate queen-laid eggs rather than by widening the template for acceptable queen-laid eggs. These novel results show that egg-discrimination behaviour in F. fusca is flexible and that workers respond to the diversity of eggs experienced in their colony.     

Almost royal: incomplete suppression of host worker ovarian development by a social parasite wasp

Insect social parasites, like other parasites, may benefit from inhibiting their host from reproducing (complete or partial parasitic castration) because they can then exploit more of the host’s resources for their own reproduction. In particular, social parasites that kill or expel the host queen need to prevent host workers from reproducing; this is a common worker response to the absence of their queen. Indeed, host workers would benefit from detecting the presence of the parasite and investing in direct and indirect fitness. Studying whether and how social parasites control host worker reproduction can provide information about the degree of integration of the parasite in the host colony and help identify factors regulating workers’ reproductive decisions in social insects. We investigated whether the paper wasp social parasite, Polistes sulcifer, suppresses Polistes dominula (host) worker reproduction as efficiently as the dominant host female does in queen-right colonies by comparing worker reproductive efforts in parasitized and non-parasitized (control) colonies. Our results show that 6 weeks after usurpation of their colony by the social parasite, parasitized workers (1) had more developed ovaries than control workers and (2) laid more eggs as soon as the opportunity arose. This reproductive readiness of parasitized workers was not apparent 2 weeks after colony usurpation. This suggests that P. dominula workers have evolved means to react to social parasitism, as occurs in some ants, and that the parasite has only limited control over host reproduction.