Polistes Dominulus Research Articles

Using social parasitism to test reproductive skew models in a primitively eusocial wasp.

Remarkable variation exists in the distribution of reproduction (skew) among members of cooperatively breeding groups, both within and between species. Reproductive skew theory has provided an important framework for understanding this variation. In the primitively eusocial Hymenoptera, two models have been routinely tested: concessions models, which assume complete control of reproduction by a dominant individual, and tug-of-war models, which assume on-going competition among group members over reproduction. Current data provide little support for either model, but uncertainty about the ability of individuals to detect genetic relatedness and difficulties in identifying traits conferring competitive ability mean that the relative importance of concessions versus tug-of-war remains unresolved. Here, we suggest that the use of social parasitism to generate meaningful variation in key social variables represents a valuable opportunity to explore the mechanisms underpinning reproductive skew within the social Hymenoptera. We present a direct test of concessions and tug-of-war models in the paper wasp Polistes dominulus by exploiting pronounced changes in relatedness and power structures that occur following replacement of the dominant by a congeneric social parasite. Comparisons of skew in parasitized and unparasitized colonies are consistent with a tug-of-war over reproduction within P. dominulus groups, but provide no evidence for reproductive concessions.

Experimental male size manipulation in Polistes dominula paper wasps: being the right size

In previous studies on Polistes dominula paper wasps, male size affects both mating success and territorial strategy. Larger males occupy exclusive territories at leks, i.e. landmarks where they achieve most copulations (Residents), while smaller males search for females in sub-optimal locations across territories as Transients. We here observed spatial and mating behaviour of a selected size-bimodal population of males, caught at leks and released in semi-natural conditions. Residents, mainly Large males, were more successful in sexual interactions with virgin females than Transients. Residents also engaged in ‘mate guarding’, reported here for the first time. On the other hand, some Small males achieved copulations inside their own territories, at foraging sites or by grasping females and falling on the ground. Both tactics of mate access – patrolling a territory for perched females or intercepting females – were successful, the former more than the latter, whereas size itself was irrelevant to sexual performance and mating success, regardless of our size-bimodal sample. Moreover, we carried out morphological analysis of male reproductive apparatus. Males with larger accessory glands, directly involved in reproductive success in many insects, mated more than males with smaller accessory glands, regardless of male body size.

Expression of Th1, Th2, lymphocyte trafficking and activation markers on CD4+ T-cells of Hymenoptera allergic subjects and after venom immunotherapy

Systemic reactions to Hymenoptera stings can be fatal and represent a reduction in the quality of life. The immune mechanisms involved in venom allergic subjects are barely known. Nevertheless, a shift towards a Th1-type response with an increase in IFNγ levels has been observed after venom immunotherapy (VIT). There is currently no information available about the expression of markers on CD4+ T-cells or their involvement in venom allergy, nor following VIT. For this, we have studied the expression of Th1 and Th2-cell markers, homing receptors and activation markers on CD4+ T-cells of subjects who presented systemic allergic reactions, mainly to Polistes dominulus, and after receiving a 4-month conventional VIT protocol. The markers studied were: CD26 (Th1), CD30 (Th2), CXCR4, CXCR3 (Th1), CCR4 (Th2), CD154 (CD40L), CD152 (CTLA-A), and ICOS. We also determined the IL-4 (Th2) and IFNγ (Th1) intracellular cytokine levels in T-cells and carried out a basophil activation test (BAT). Comparing venom allergic subjects with non-allergic healthy controls, we have found up-regulation of CD26, CXCR4, CXCR3, CD154 and ICOS. Conversely, a down-regulation of CD30, CD154 and CD152 occurred upon immune intervention, whereas the remaining markers were not affected. Equally, VIT has been shown to be effective, as evidenced by the decrease of basophil degranulation and increase of IFNγ levels in T-cells after the fourth month of treatment. These new findings highlight the possible application of these surface molecules as markers to distinguish between symptomatic and asymptomatic subjects sensitized to Hymenoptera venom, as well as revealing information about the immune changes associated with VIT.

Novel insights into the ontogeny of nestmate recognition in Polistes social wasps.

The importance of early experience in animals’ life is unquestionable, and imprinting-like phenomena may shape important aspects of behaviour. Early learning typically occurs during a sensitive period, which restricts crucial processes of information storage to a specific developmental phase. The characteristics of the sensitive period have been largely investigated in vertebrates, because of their complexity and plasticity, both in behaviour and neurophysiology, but early learning occurs also in invertebrates. In social insects, early learning appears to influence important social behaviours such as nestmate recognition. Yet, the mechanisms underlying recognition systems are not fully understood. It is currently believed that Polistes social wasps are able to discriminate nestmates from non-nestmates following the perception of olfactory cues present on the paper of their nest, which are learned during a strict sensitive period, immediately after emergence. Here, through differential odour experience experiments, we show that workers of Polistes dominula develop correct nestmate recognition abilities soon after emergence even in absence of what have been so far considered the necessary cues (the chemicals spread on nest paper). P. dominula workers were exposed for the first four days of adult life to paper fragments from their nest, or from a foreign conspecific nest or to a neutral condition. Wasps were then transferred to their original nests where recognition abilities were tested. Our results show that wasps do not alter their recognition ability if exposed only to nest material, or in absence of nest material, during the early phase of adult life. It thus appears that the nest paper is not used as a source of recognition cues to be learned in a specific time window, although we discuss possible alternative explanations. Our study provides a novel perspective for the study of the ontogeny of nestmate recognition in Polistes wasps and in other social insects.

Parasitic castration byXenos vesparumdepends on host gender

Host castration represents a mechanism used by parasites to exploit energy resources from their hosts by interfering with their reproductive development or to extend host lifespan by removing risks associated with reproductive activity. One of the most intriguing groups of parasitic castrators is represented by the insects belonging to the order Strepsiptera. The macroparasite Xenos vesparum can produce dramatic phenotypic alterations in its host, the paper wasp Polistes dominula. Parasitized female wasps have undeveloped ovaries and desert the colony without performing any social task. However, very little attention has been given to the parasitic impact of X. vesparum on the male phenotype. Here, we investigated the effects of this parasite on the sexual behaviour and the morpho-physiology of P. dominula males. We found that, differently from female wasps, parasitized males are not heavily affected by Xenos: they maintain their sexual behaviour and ability to discriminate between female castes. Furthermore, the structure of their reproductive apparatus is not compromised by the parasite. We think that our results, demonstrating that the definition of X. vesparum as a parasitoid does not apply to infected males of P. dominula, provide a new perspective to discuss and maybe reconsider the traditional view of strepsipteran parasites.

Erratum to: Examining the “evolution of increased competitive ability” hypothesis in response to parasites and pathogens in the invasive paper wasp Polistes dominula

Erratum to: Examining the “evolution of increased competitive ability” hypothesis in response to parasites and pathogens in the invasive paper wasp Polistes dominula

A test of the haploid susceptibility hypothesis using a species with naturally occurring variation in ploidy

The haploid susceptibility hypothesis (HSH) was proposed as an explanation for how behavioral roles in haplodiploid social systems evolved. It posits that haploid males are more susceptible to disease than diploid females due to decreased genetic variability at key disease resistance loci. The resulting decreased immunocompetence is hypothesized to have played a role in the evolution of social behavior by limiting the behavioral repertoire haploids perform. Here, we test this hypothesis in a study system that separates ploidy from behavioral sex roles: Polistes dominulus, a social wasp, has colonies with naturally occurring diploid males. We report results from two immune function assays—hemolymph phenoloxidase activity and encapsulation response—performed on haploid males, diploid males, and diploid females. Our data suggest that ploidy is not a significant contributor to immune function in P. dominulus; thus, our data do not support the HSH for the evolution of behavioral roles. Instead, our data indicate that time of emergence is the best predictor of immune function in Polistes. We speculate that seasonal trends result from seasonal differences in pathogens and parasites.

Dominant aggression as a deterrent signal in paper wasps

Low-level social aggression is a conspicuous feature of cooperative animal societies, but its precise function is usually unclear. One long-standing hypothesis is that aggressive displays by dominant individuals serve to reduce uncertainty about relative strength and deter subordinates from starting fights that they are unlikely to win. However, most formal theoretical models of this idea do not consider how the credibility of deterrent signals might change over time in social groups. We developed a simple model of dominant aggression as a deterrent signal, which takes into account how credibility changes over time and how selection should act on receiver memory. We then carried out an experimental test of the predictions of our model on a field population of the paper wasp, Polistes dominulus. The match between our theoretical and empirical results suggests that low-level social aggression can help to maintain the stability and productivity of cooperative associations in this species. Moreover, our work suggests that rates of aggression in animal societies and the robustness of social memories are likely to be intimately related.

Available kin recognition cues may explain why wasp behavior reflects relatedness to nest mates

Relatedness is predicted to be a key determinant of cooperative behavior, but kin discrimination within social insect colonies is surprisingly rare. A lack of reliable cuticular hydrocarbon (CHC) cues is thought to be responsible, but here we show that in a high-profile paper wasp model, kin recognition cues are available for some individuals that found nests with nonrelatives. Thus, unrelated Polistes dominulus helpers could potentially recognize themselves as such. On this basis, we reanalyzed a behavioral data set to investigate whether foraging effort, defense contributions and aggression toward nest mates might thus reflect CHC profiles. Both foraging behavior and aggression varied with genetic relatedness, but genetic relatedness itself was a better predictor of this variation than differences in CHC profiles. We propose that wasps use specific components of the CHC profile, the identity of which is as yet unknown, to identify relatives among nest mates. Our data provide the first evidence of within-nest kin discrimination in primitively eusocial wasps but leave open the question of which cues are responsible.

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.

Duration of development of males in colonies of Polistes dominula (Christ) (Hymenoptera, Vespidae: Polistinae) in the south of Ukraine

The relation between the duration of ontogenesis of Polistes dominula males and the social structure of their colonies was studied in July–August 2012. Males developing in successful colonies with a foundress passed through the later-instar larval and pupal stages more quickly than those in orphaned colonies. Successful colonies more often produced males with weakly melanized mesopleura as compared with orphaned ones. Males from the two categories of colonies did not differ in the head and wing size. The influence of mechanical stress and nutrition during larval development and the specific signaling behavior of the foundresses on the male phenotypic diversity in P. dominula is discussed.

Do paper wasps negotiate over helping effort?

Recent theory and empirical studies of avian biparental systems suggest that animals resolve conflict over parental care via a process of behavioral negotiation or “rules for responding.” Less is known, however, about whether negotiation over helping effort occurs in cooperatively breeding animal societies or whether behavioral negotiation requires a relatively large brain. In this study, we tested whether negotiation over help occurs in a social insect, the paper wasp Polistes dominulus, by recording individual responses to both observed and experimentally induced foraging returns by other group members. In our experiments, we manipulated food delivery to the nest in 2 ways: 1) by catching departing foragers and giving them larval food to take back to the nest and 2) by giving larval food directly to wasps on the nest, which they then fed to larvae, so increasing food delivery independently of helper effort. We found no evidence from Experiment 1 that helpers adjusted their own foraging effort according to the foraging effort of other group members. However, when food was provided directly to the nest, wasps did respond by reducing their own foraging effort. One interpretation of this result is that paper wasp helpers adjust their helping effort according to the level of offspring need rather than the work rate of other helpers. Negotiation based on indicators of demand rather than work rate is a likely mechanism to resolve conflict over investment in teams where helpers cannot observe each other’s work rate directly, as is commonly the case in insect and vertebrate societies.

Preferential phenotypic association linked with cooperation in paper wasps

Animals can influence their social environment by preferentially associating with certain conspecifics. Such preferential association has gained increasing theoretical attention, as it may influence social evolution and population dynamics. However, relatively little empirical work has examined the occurrence of preferential association and its effects on cooperative group formation. Here, we test the factors associated with cooperative group formation in Polistes dominulus nest-founding queen wasps. P.dominulus are a good system to study preferential association, as foundresses can nest alone or in groups and group membership is flexible. We found that both social and environmental factors were associated with partner choice. First, facial patterns were associated with cooperation. Wasps with more similar facial patterns were more likely to cooperate than wasps with less similar facial patterns. This preferential phenotypic association fits the theoretical criteria for the evolution of tag-based cooperation. Season was also associated with cooperation; wasps on early-season nests were more likely to cooperate than wasps on late-season nests. High levels of aggression by nest owners during initial interactions were also correlated with lower probabilities of subsequent cooperation, suggesting that nest owners have some control over group membership. Other factors including body weight, weight similarity and nest productivity were not linked with cooperation. Overall, multiple factors influence cooperation in paper wasps, including facial pattern similarity. The occurrence of preferential phenotypic association in paper wasps is quite interesting and may influence the evolution of cooperation and population divergence in this group.

Males do not like the working class: male sexual preference and recognition of functional castes in a primitively eusocial wasp

Discrimination among potential partners is a critical step in sexual selection to avoid wasting reproductive resources on an unsuitable mate. In the female-dominated hymenopteran societies males have often been regarded as ‘flying sperm containers’ spending all their time and energy in trying to acquire a mate. We investigated the male sexual preference for potential partners using as a model the primitively eusocial wasp Polistes dominula in which female caste is rather flexible and difficult to determine. By means of laboratory bioassays, we compared the males' behaviour towards females of different reproductive potential. Males were able to recognize female castes, strongly preferring reproductive females to workers, regardless of female age or health. The results show that in this species caste plays a key role in orienting male discrimination and preference, presumably through chemical cues, towards reproductive females both healthy and parasite-castrated. Overall, our study shows that social Hymenoptera males are not always ‘small mating machines’ eager to mate.

Actinomycetes with Antimicrobial Activity Isolated from Paper Wasp (Hymenoptera: Vespidae: Polistinae) Nests

Actinomycetes-a group of antimicrobial producing bacteria-have been successfully cultured and characterized from the nest material of diverse arthropods. Some are symbionts that produce antimicrobial chemicals found to protect nest brood and resources from pathogenic microbes. Others have no known fitness relationship with their associated insects, but have been found to produce antimicrobials in vitro. Consequently, insect nest material is being investigated as a new source of novel antimicrobial producing actinomycetes, which could be harnessed for therapeutic potential. To extend studies of actinomycete-insect associations beyond soil-substrate dwelling insects and wood boring excavators, we conducted a preliminary assessment of the actinomycetes within the nests of the paper wasp, Polistes dominulus (Christ). We found that actinomycetes were readily cultured from nest material across multiple invasive P. dominulus populations-including members of the genera Streptomyces, Micromonospora, and Actinoplanes. Thirty of these isolates were assayed for antimicrobial activity against the challenge bacteria Pseudomonas aeruginosa, Escherichia coli, Staphylococcus aureus, Serratia marcescens, and Bacillus subtilis. Sixty percent of isolates inhibited the growth of at least one challenge strain. This study provides the first assessment of bacteria associated with nests of P. dominulus, and the first record of antimicrobial producing actinomycetes isolated from social wasps. We provide a new system to explore nest associated actinomycetes from a ubiquitous and cosmopolitan group of insects.

A survey of DNA methylation across social insect species, life stages, and castes reveals abundant and caste-associated methylation in a primitively social wasp

DNA methylation plays an important role in the epigenetic control of developmental and behavioral plasticity, with connections to the generation of striking phenotypic differences between castes (larger, reproductive queens and smaller, non-reproductive workers) in honeybees and ants. Here, we provide the first comparative investigation of caste- and life stage-associated DNA methylation in several species of bees and vespid wasps displaying different levels of social organization. Our results reveal moderate levels of DNA methylation in most bees and wasps, with no clear relationship to the level of sociality. Strikingly, primitively social Polistes dominula paper wasps show unusually high overall DNA methylation and caste-related differences in site-specific methylation. These results suggest DNA methylation may play a role in the regulation of behavioral and physiological differences in primitively social species with more flexible caste differences.

Distance transform: a tool for the study of animal colour patterns

Summary The information in animal colour patterns plays a key role in many ecological interactions; quantification would help us to study them, but this is problematic. Comparing patterns using human judgement is subjective and inconsistent. Traditional shape analysis is unsuitable as patterns do not usually contain conserved landmarks. Alternative statistical approaches also have weaknesses, particularly as they are generally based on summary measures that discard most or all of the spatial information in a pattern. We present a method for quantifying the similarity of a pair of patterns based on the distance transform of a binary image. The method compares the whole pattern, pixel by pixel, while being robust to small spatial variations among images. We demonstrate the utility of the distance transform method using three ecological examples. We generate a measure of mimetic accuracy between hoverflies (Diptera: Syrphidae) and wasps (Hymenoptera) based on abdominal pattern and show that this correlates strongly with the perception of a model predator (humans). We calculate similarity values within a group of mimetic butterflies and compare this with proposed pairings of Mullerian comimics. Finally, we characterise variation in clypeal badges of a paper wasp (Polistes dominula) and compare this with previous measures of variation. While our results generally support the findings of existing studies that have used simpler ad hoc methods for measuring differences between patterns, our method is able to detect more subtle variation and hence reveal previously overlooked trends.

Juvenile hormone influences precontest assessment behaviour in Polistes dominulus paper wasps

Theory on animal contests often assumes that individuals can rapidly and accurately assess their own resource-holding potential (RHP). In empirical work, individuals behave as if they ‘know’ their own RHP, but little is known about how such assessment occurs. Here, we test how juvenile hormone (JH) influences precontest behaviour in P. dominulus paper wasps. Previous work has shown that P. dominulus have variable black facial patterns that are a signal of agonistic ability and that JH influences fighting ability. In this experiment, we tested how JH influences rival challenge behaviour in the absence of direct social competition. Wasps were given the opportunity to challenge or avoid a model wasp guarding a patch of food. Individuals treated with the JH analogue methoprene were more likely to challenge rivals than were control individuals, and they also had a shorter latency to challenge than controls. However, hormone treatment was only one of multiple factors that influenced behaviour, suggesting that assessment prior to contests is based on the interplay between individual physiology and social context. Contest decisions were also dependent on a wasp's own facial pattern and mass, as well as the facial pattern of its rival. Interestingly, these complex assessment decisions occurred in the absence of direct social cues that could be acquired during a contest. These results are consistent with JH playing a role in self-assessment of RHP. Hormones such as JH are influenced by the physical and social environment, so they provide diverse information about relative RHP and may be commonly involved in assessment behaviour.

Examining the “evolution of increased competitive ability” hypothesis in response to parasites and pathogens in the invasive paper wasp Polistes dominula

Successful invaders often become established in new ranges by outcompeting native species. The "evolution of increased competitive ability" hypothesis predicts that invasive species are subjected to less predation and parasitization than sympatric native species, and thus can allocate resources from defence and immunity to growth and fecundity, thereby achieving higher fitness. In this study, we examined whether American invasive Polistes dominula paper wasps have reduced immunocompetence. To explore this scenario, we tested their susceptibility towards parasites and pathogens at both the individual (immune defence) and colony levels, i.e. hygienic behaviour (removal of diseased individuals by nestmates). First, we examined the response to the specific coevolved parasite Xenos vesparum (lost after invasion) in terms of individual host susceptibility and hygienic behaviour. Second, we explored the response against general pathogens by quantifying the bacterial clearance in individual wasps after a challenge with Escherichia coli and hygienic behaviour after a challenge with the fungus Beauveria bassiana. Our results show that American invasive P. dominula have a higher response against X. vesparum at the colony level, but at the individual level their susceptibility is not significantly different from conspecifics of the native range. On the other hand, invasive P. dominula display lower response after a challenge with general pathogens at both the individual and colony levels. While supporting the hypothesis of a reduction of immunocompetence towards general pathogens in invasive species, these findings also suggest that the response against coevolved parasites might follow different evolutionary pathways which are not always easily predictable.

Clypeal patterning in the paper wasp Polistes dominulus: no evidence of adaptive value in the wild

Status signals function in a number of species to communicate competitive ability to conspecific rivals during competition for resources. In the paper wasp Polistes dominulus, variable black clypeal patterns are thought to be important in mediating competition among females. Results of previous behavioral experiments in the lab indicate that P. dominulus clypeal patterns provide information about an individual’s competitive ability to rivals during agonistic interactions. To date, however, there has been no detailed examination of the adaptive value of clypeal patterns in the wild. To address this, we looked for correlations between clypeal patterning and various fitness measures, including reproductive success, hierarchical rank, and survival, in a large, free-living population of P. dominulus in southern Spain. Reproductive success over the nesting season was not correlated with clypeal patterning. Furthermore, there was no relationship between a female’s clypeal patterning and the rank she achieved within the hierarchy or her survival during nest founding. Overall, we found no evidence that P. dominulus clypeal patterns are related to competitive ability or other aspects of quality in our population. This result is consistent with geographical variation in the adaptive value of clypeal patterns between P. dominulus populations; however, data on the relationship between patterning and fitness from other populations are required to test this hypothesis. Key wor ds: clypeal pattern, fitness, Polistes dominulus, quality, status signalling, wild. [Behav Ecol]