Intraspecific comb usurpation in the social wasp Polistes fuscatus

Summary Incidents of usurpation were observed in colonies of Polistes fuscatus nesting on farm buildings (1977–79) and in nestboxes (1980–84) in Johnson County, Iowa, USA. Most usurpations (84.8%) occurred in the latter half of the preworker phase of the colony cycle, which coincided with periods of high predation of combs by vertebrates. Usurpers were probably displaced single foundresses which did not join neighbors or refound colonies after comb loss. Most (89–100%) usurpers of known relatedness to the foundresses they replaced were cousins or less related to them. Usurpation was a significant source of nest loss (19.6%) among single foundresses, but was rare (2.2%) in multiple-foundress colonies and colonies with workers (3.5%). Usurpers often destroyed younger brood (eggs and larvae in instars 1–3) in host colonies, while older larvae and pupae were usually spared. Brood destruction was more pronounced in more advanced host combs. Usurper survivorship after workers eclosed was lower than than of queenright single foundresses (61.5% vs 87.0%). Reproductive success by usurpers was less than that of queenright single foundresses, but greater than that of foundresses which initiated colonies late in the preworker colony cycle.     

Nestmate discrimination in social wasps (Polistes metricus,Hymenoptera: Vespidae)

Summary Laboratory-overwintered paper wasp (Polistes metricus) females preferentially associate on new nests with former nestmates (presumably their sisters) rather than with non-nestmates in the absence of cues associated with their natal nests or nest sites. Females isolated from conspecifics and nests for 74–99 days retain the ability to discriminate nestmates from nonnestmates.     

Control of reproduction in social insect colonies: individual and collective relatedness preferences in the paper wasp, Polistes annularis

Social insect colonies often have one or a few queens. How these queens maintain their reproductive monopoly, when other colony members could gain by sharing in the reproduction, is not generally known. DNA microsatellite genotyping is used to determine reproductive interests of various classes of colony members in the paper wasp, Polistes annularis. The relatedness estimates show that the best outcome for most individuals is to be the reproductive egg-layer. For workers, this depends on the sex of offspring: they should prefer to lay their own male eggs, but are indifferent if the queen lays the female eggs. The next-best choice is usually to support the current queen. As a rule, subordinates and workers should prefer the current queen to reproduce over other candidates (though subordinates have no strong preference for the queen over other subordinates, and workers may prefer other workers as a source of male eggs). This result supports the theory that reproductive monopoly stems from the collective preferences of non-reproductives, who suppress each other in favor of the queen. However, we reject the general hypothesis of collective worker control in this species because its predictions about who should succeed after the death of the present queen are not upheld. The first successor is a subordinate foundress even though workers should generally prefer a worker successor. If all foundresses have died, an older worker succeeds as queen, in spite of a collective worker preference for a young worker. The results support the previous suggestion that age serves as a conventional cue serving to reduce conflict over queen succession. Received: 3 May 1996 / Accepted after revision: 22 September 1996     

Cuticular hydrocarbons and reproductive status in the social wasp Polistes dominulus

Most species of social insect are characterized by a reproductive division of labor among morphologically specialized individuals. In contrast, there exist many species where all individuals are morphologically identical and dominance relationships determine which individuals mate and/or reproduce. In newly founded multiple-foundress associations of the social wasp Polistes dominulus, foundresses establish dominance hierarchies where the top-ranked (alpha) female monopolizes egg laying. The possibility that chemical cues are used for recognition of egg-laying individuals has not been explored in this wasp. Using non-destructive techniques, we examined the relationship between ovarian activity and the proportions of cuticular hydrocarbons of three female types (dominant and subordinate foundresses and workers) in 11 colonies. Immediately after nest foundation, no differences were found between alpha and subordinate females. In contrast, at worker emergence, alpha females were statistically distinguishable from both subordinates and workers. We experimentally removed the alpha female in 5 of the original nests and reanalyzed hydrocarbon proportions of the new dominant individual. Replacement individuals were all found to acquire a cuticular signature characteristic of the alpha female. This suggests that cuticular hydrocarbons are used as cues of ovarian activity in P. dominulus, and we discuss our results in terms of a switch from behavioral dominance to chemical signaling in this wasp.     

Physical variability among nest foundresses in the polygynous social wasp,Polistes annularis

Summary An individual's rank in a dominance hierarchy is often based on size or weight, especially in insects. Foundresses of the social wasp, Polistes annularis, vary greatly in size as measured by wing length, dry weight, fat weight, and residue weight after fat has been extracted. Females that emerged from the same nest are much more similar in size to each other than they are to females that emerged from other nests. Within nests however queens are usually larger than their subordinates. Queens that emerged from one nest may be smaller than subordinates that emerged from another nest. We found no evidence of a group of females that are forced into being subordinates because of inadequate feeding as larvae. Females are also probably not attempting to begin new nests with females as different in size as possible from themselves since means and variances of winglength of females on new nests do not differ from those of all females emerging from that natal nest.     

Aggression and resource sharing among foundresses in the social wasp Polistes dominulus: testing transactional theories of conflict

Tug-of-war models of within-group conflict predict that the frequency of conflict will be positively related to the degree of reproductive sharing within the group; in contrast, a negative relationship supports transactional models, in which reproductive payments among group members limit the degree of within-group selfishness. We tested predictions of the tug-of-war and transactional models by examining cofoundress interactions during the founding (preworker) phase of colony development in 30 naturally nesting colonies of the paper wasp Polistes dominulus. We found that the mean rate of foundress aggression and the mean probability of food sharing were significantly negatively associated, which supports the prediction of the transactional, not the tug-of-war model. Further, cofoundress aggression significantly increased over the founding phase (independently of temperature), while the fraction of aggression initiated by the dominant (alpha) foundress significantly decreased over this period. We show that both of these results are predicted by the transactional model of within-group conflict. Interestingly, the alpha’s rate of aggression was significantly positively temperature dependent, while the beta’s was not. This indicates that the alpha’s aggression level may often be near her physiological maximum, while the beta’s aggression is limi- ted by other factors, contradicting the prediction of the tug-of-war model. Moreover, the alpha’s aggression was significantly positively temperature dependent only in the second half of the founding period, as predicted by the transactional model since this is when there is least reproductive sharing. Finally, our results indicate that the alpha’s level of aggression depends on the resources controlled by the beta. Received: 18 January 2000 / Revised: 19 June 2000 / Accepted: 24 June 2000     

Solitary nesting and reproductive success in the paper wasp Polistes aurifer

Female Polistes paper wasps are capable of independent nesting, yet many populations demonstrate a mixture of solitary and cooperative nest foundation. Previous studies of Polistes have found survival and/or productivity advantages of cooperative nest foundation compared to solitary nesting, and reproductive skew models have been designed to predict the dynamics of such flexible cooperation. In this paper, we examine the success of different nesting strategies in a previously unstudied population of Polistes aurifer in southern California. The colony cycle of this population is less synchronous than that of other temperate species, and the frequency of solitary nesting averages 86.2%. Our data suggest that this low rate of cooperative nest founding is adaptive, as demonstrated by the lack of survival or productivity advantages for cooperative foundress associations. Due to foundress turnover and nest foundation later in the season, many nests produce only one set of offspring. This results in a loss of the eusocial nature of some nests in the population. Data from a small sample of multifoundress nests show significant positive reproductive skew, despite concession model predictions that skew should be low in populations with low ecological constraints on independent nesting. This lack of support for the concessions skew model reflects a diminished incentive for cooperation.Communicated by L. Keller     

Role of ovaries and ecdysteroids in dominance hierarchy establishment among foundresses of the primitively social wasp,Polistes gallicus

Summary The contribution of ovaries and ovarian ecdysteroids to dominance and nest initiation was tested in foundresses of Polistes gallicus at the end of hibernation. Removal of the ovaries resulted in a significant decrease of the ecdysteroid titre in haemolymph. Dominance in ovariectomized foundresses was found to be closely correlated to the size of corpora allata. The operated wasps started nest building, but they did not show any egg-laying behaviour. We conclude that both juvenile hormone and ovarian ecdysteroids contribute to degree of dominance, and that factors other than ovary development may stimulate building.     

Recognition of former nestmates during colony founding by the social wasp Polistes fuscatus (Hymenoptera: Vespidae)

Summary In the laboratory overwintered females of Polistes fuscatus preferentially associated with former nestmates during the founding of a colony. They did not associate with non-nestmates or only with nestmates with whom they had overwintered, but affiliated with all former nestmates, regardless of where the former nestmates overwintered.     

Stable relatedness structure of the large-colony swarm-founding wasp Polybia paulista

In social-insect colonies, cooperation among nestmates is generally stabilized by their high genetic similarity. Thus, fitness gained through cooperation drops quickly as the number of reproductive females (queens) increases. In this respect, wasps of the tribe Epiponini have attracted special attention, because the colonies have tens, or even hundreds of queens. It has been empirically or genetically confirmed that relatedness within nestmates can be elevated by a mechanism known as cyclical monogyny, under which new queens are produced only after the number of old queens is reduced to one. Another likely factor that can increase relatedness within nestmates under polygyny is comb partitioning by queens. If queens concentrate their egg laying on one or a subset of the available combs, then workers may be able to rear closer relatives by focusing their work on the comb where they emerged. Using microsatellite markers, we tested the hypotheses of cyclical monogyny and comb partitioning by queens increasing relatedness within nestmates under polygyny in the large-colony epiponine wasp, Polybia paulista. There were no significant differences between relatedness within combs and between combs, and thus we ruled out the possibility that each queen partitions reproduction between combs. However, as cyclical monogyny predicts, a lower effective number of queens contributed to queen production than to worker production. Cyclical monogyny explained well the observed smaller effective number of queens for new queens than that for workers, but failed to explain the stable relatedness values throughout colony cycles.Communicated by L. Keller     

Timing matters when assessing dominance and chemical signatures in the paper wasp Polistes dominulus

A recent study (Izzo et al., Behav Ecol Sociobiol 64: 857–864, 2010) reported that cuticular hydrocarbons (CHCs) correlate with fertility, not dominance, in the paper wasp Polistes dominulus thus contradicting the results of recent investigations which concluded that social dominance is the main determinant for CHC signatures in this species. We suggest here that different forms of dominance in the pre-nesting and post-nesting phases caused the apparently contradictory results. Thus the assumption that dominance behaviour in the pre-nesting stage is synonymous with dominance after colony foundation is incorrect. We provide standardised definitions for forms of “dominance” observed in the P. dominulus life cycle to avoid apparent discrepancies in the future among studies dealing with the same topics in different annual stages.     

Behavioural evidence for the involvement of Dufour’s gland secretion in nestmate recognition in the social wasp Polistes dominulus (Hymenoptera: Vespidae)

The results of recent studies by our group have suggested that in social wasps of the genus Polistes the Dufour’s gland is involved in kin recognition. In fact, the same hydrocarbons occurring on the cuticle are found in the gland secretion, and in P. dominulus the composition of the glandular secretion is more similar in foundresses belonging to the same colonies than in heterocolonial foundresses. In this study, P. dominulus colonies were experimentally presented with previously treated dead conspecific females. These lures had been deprived of their epicuticular lipids and coated with epicuticular or Dufour’s gland secretion extracts from females hetero- or homocolonial with respect to the tested colonies. The behaviour of the colonies towards these lures indicates that, like the epicuticular lipids, the Dufour’s gland secretion is involved in nestmate recognition. Received: 14 August 1995/Accepted after revision: 5 November 1995     

Cuticular hydrocarbons correlate with fertility, not dominance, in a paper wasp, Polistes dominulus

Cuticular hydrocarbons (CHCs) are information-rich signals in social insects that coordinate behaviors within nests. However, in some taxa, the precise information conveyed by CHCs is poorly understood. In particular, there is a debate over whether CHCs convey information about their bearer's dominance or fertility. Distinguishing between dominance and fertility signaling is difficult because fertility and rank are frequently correlated within social insect colonies. This study disentangles those relationships by examining CHCs of Polistes dominulus paper wasps during the early nest-founding stage before dominance and fertility become correlated. First, we confirm that dominance and fertility are not associated in early spring foundresses. Then we show that CHCs are more strongly associated with fertility than dominance. There was no relationship between cuticular hydrocarbon profiles and a wasp’s ability to dominate rivals through aggression, suggesting that CHCs are unlikely to provide information about dominance. However, there was a significant correlation between ovarian development and the CHC profile, suggesting that CHCs could convey valuable information about their bearer's fertility. Furthermore, our data provide a potential mechanism for chemical signaling of fecundity, as there is a relationship between endogenous juvenile hormone titer (a gonadotropin), degree of ovarian development, and the CHC profile. Hormonal regulation of CHC profile expression offers a physiological mechanism to coordinate behavior, physical state, and signal expression.     

Influence of insect hormones on the establishment of dominance hierarchies among foundresses of the paper wasp,Polistes gallicus

Summary Laboratory studies on overwintered foundresses of Polistes gallicus show that differences in the endocrine activity are mainly responsible for achieving the dominance rank. Females that became dominant had either larger corpora allata or more developed ovaries than subordinate females. Body size did not contribute to dominance rank. Since a correlation exists between the length of terminal oocytes and the ecdysteroid titre in haemolymph as well as between the volume of corpora allata and the synthesis of juvenile hormone, dominant behaviour is thought to depend upon an elevated hormone titre in haemolymph. Injections of juvenile hormone (JHI) and 20-hydroxyecdysone, separately and simultaneously, significantly increased the probability that the treated female would be the dominant female of a test pair. After a hierarchy has been established, endocrine activity in subordinate foundresses is inhibited by the dominant foundress that then monopolizes reproduction.     

Social context influences cue-mediated recruitment in an invasive social wasp

Social insects often serve as model systems for communication and recruitment studies, and yet, it remains controversial whether social vespid wasps can reliably communicate resource information to nestmates. In this study, I present empirical evidence that foraging strategies depend on the initial assessment of resource size and potential competition by foraging yellowjackets. The context dependent foraging behavior of Vespula pensylvanica provides a potential explanation for the inconsistent reports of the existence of recruitment communication in vespid wasps. Furthermore, life history traits may influence yellowjacket foraging behavior; annual V. pensylvanica colonies, whose foragers routinely patrol near the nest, exhibited increased bait visitation in response to the return of successful foragers, whereas perennial colonies did not. These behavioral disparities provide insight into how foraging strategies and search patterns may shift with colony size and longevity. In experiments that investigate the effects of visual cues of conspecifics and bait dispersion, foraging decisions corresponded with expectations of yellowjackets integrating resource quantity and access into a perception of demand. When resource competition could be assessed as high, V. pensylvanica foragers quickly exploited the bait closest to their colony regardless of occupation by other wasps; however, foragers preferred visiting unoccupied baits in situations where competition could be perceived as low. Moreover, a meta-analysis revealed that context-dependent, cue-mediated recruitment was widespread in Vespidae, where such foraging behaviors changed with habitat and the potential for resource competition. Such plastic foraging strategies may contribute to the invasion success of some vespid wasps.