Polymorphic microsatellite loci for eusocial wasps (Hymenoptera: Vespidae)

Hymenopterans are an important model for studying the evolution of cooperation in animal societies. Here, we characterize 19 microsatellite loci, isolated from the common wasp Vespula vulgaris, that can be used to study genetic variation in three genera (seven species) within the Vespidae. The number of alleles in V. vulgaris was moderate, varying from 2 to 14, with expected heterozygosity ranging between 0.04 and 0.93. Eleven loci amplified DNA in V. austriaca and Dolichovespula sylvestris, nine in V. germanica, eight in Vespa crabro and V. rufa, seven in D. media and only five loci could be used for D. norwegica.     

Polymorphic microsatellite loci for the Japanese paper wasp,Polistes chinensis antennalis (Hymenoptera: Vespidae)

We isolated 13 microsatellite markers for the Japanese paper wasp Polistes chinensis antennalis. Null alleles were ruled out in the loci because we could detect polymerase chain reaction products for 578 haploid males. Our results indicate that these markers are excellent tools for the analysis of pedigrees within colonies for species with a complicated pedigree structure due to worker reproduction (male production by workers in queen‐right colonies).     

Cryptically dimorphic caste differences in a Neotropical, swarm-founding paper wasp genus, Parachartergus (Hymenoptera: Vespidae)

Morphological differences in body shape between females of different reproductive conditions (in terms of insemination and ovarian development) were examined in two species of the Neotropical polistine genus Parachartergus: P. smithii and P. fraternus. The present study shows, for the first time, that non‐size‐based morphological divergence between queens and workers occurs in Parachartergus, an epiponine genus once believed to have little or no morphological caste differences. In the P. smithii colony examined, queens were significantly larger than workers in five of the eight body parts measured (head width, eye width, genal width, mesosomal length, wing length, first metasomal tergum width, and width and length of the second tergum), but the mean values of wing length and first and second tergum widths were not significantly different between them. The queen : worker size ratios tended to be greater anteriorly and smaller posteriorly, although the size ratio was greatest in second tergum length. Analysis of covariance (ancova ) with mesosomal length as covariate showed that queens had proportionally wider heads and narrower first terga than did workers. In the P. fraternus colony, size differences between queens and workers were not significant, and there was little or no difference in shape, but queens had significantly proportionally wider first terga than did workers.     

Adult mortality rates in young colonies of a swarm-founding social wasp (Polybia occidentalis)

The swarm founders are unusual among the social wasps in having socialized the dispersal stage of the life cycle. Colonies are initiated by groups of workers accompanied by smaller numbers of queens. Thus, swarm founders avoid the colony size bottleneck faced by the independent founders, whose colonies start with one or a few queens. Among the advantages of swarm founding is a reduction of the risk of colony failure due to attrition of the founding adults. Stochasticity in adult mortality is less likely to lead to outright extinction of a large founding group before new workers are produced (pre-emergence period). However, it is not known how important pre-emergence mortality is as a selective force on founding and dispersal behaviour in swarm founders, since colony-wide mortality rates have never been reported for a large-colony social wasp. Sixty-eight swarms of Polybia occidentalis were censused just before colony initiation and again 24 days later. Mortality over this period averaged 0.41±0.12 of the founding swarm population. Including mortality on the day of emigration and extrapolating to day 30, when the first adult offspring eclose, the original absconding swarm would be reduced by 0.52 of its initial size. Rates of loss during the first week, while the colonies engaged in nest construction, did not differ from rates over the full 24 days. Thus, colony founding in P. occidentalis is both costly and highly variable in terms of mortality of the founding adults.     

Worker biting interactions and task performance in a swarm-founding eusocial wasp (Polybia occidentalis, Hymenoptera: Vespidae)

Workers in many insect societies interact via body contact withtheir nest mates, and social biting and other forms of contactmay play a general role in regulating task performance. HereI present evidence that social biting affects task performancewithout direct reproductive conflict in Polybia occidentalis,a swarm-founding eusocial wasp. Polybia occidentalis workersengaged in social biting with nest mates. Most workers thatwere active on the nest surface participated in biting interactions,but individuals differed significantly in their rates of biting and of being bitten. Rates of being bitten corresponded withnonreproductive task performance: more biting was directedat foragers than nonforagers, and foraging rates were correlatedwith rates of being bitten. Furthermore, some on-nest workersinitiated foraging activity immediately after they were bitten.Together these patterns suggest that social biting influencesforaging rates by increasing workers' probabilities of leavingthe nest. Variation in biting rates did not correspond withdifferences in reproductive physiology: highly active bitersand recipients did not differ in body size or in ovary development.In P. occidentalis and in other eusocial insects with largeworker forces, biting and other types of social contact amongworkers may regulate task performance independently of directreproductive competition.     

The acceptance rate of young wasps by alien colonies depends on colony developmental stages in the swarm-founding wasp, Polybia paulista von ihering (Hymenoptera: Vespidae)

In social insects, newly emerged individuals learn the colony-specific chemical label from their natal comb shortly after their emergence. These labels help to identify each individual's colony of origin and are used as a recognition template against which individuals can discriminate nestmates from non-nestmates. Our previous studies with Polybia paulista von Ihering support this general pattern, and the acceptance rate of young female and male wasps decreased as a function of their age. Our study also showed in P. paulista that more than 90% of newly emerged female wasps might be accepted by conspecific unrelated colonies. However, it has not been investigated whether the acceptance rate of newly emerged female wasps depends on colony developmental stage of recipient colonies. We introduced newly emerged female wasps of P. paulista into different colony developmental stags of recipient colonies, i.e., worker-producing and male-producing colonies. We found that the acceptance rate of newly emerged female wasps by alien colonies was pretty lower by male-producing colonies than worker-producing colonies. This is the first study to show that the acceptance rate of young female wasps depends on stages of recipient colonies.     

Mating structure and genetic relatedness among gynes in the primitively eusocial wasp Polistes snelleni (Hymenoptera: Vespidae)

Mating structure and genetic relatedness among gynes (potential reproductive females) of the paper wasp Polistes snelleni were investigated using DNA microsatellite markers. All colonies had one foundress inseminated by a single male, and no sign of inbreeding was detected. The mean genetic relatedness among gynes was 0.734 ± 0.028 (SE), which is not significantly different from the expected value of 0.75 for full sisters.     

Polymorphic microsatellite loci in the parasitic wasp Cotesia glomerata (Hymenoptera: Braconidae)

Five polymorphic microsatellite loci were isolated in a gregarious endoparasitoid, Cotesia glomerata, using an enrichment protocol. We detected two to 15 alleles per locus in a sample of 50 female wasps collected from a field population near Zürich, Switzerland. The observed heterozygosities ranged from 0.21 to 0.92. These markers are currently being used in our studies of sex determination in this species.     

Worker Age, Ovary Development, and Temporal Polyethism in the Swarm-Founding Wasp Polybia occidentalis (Hymenoptera: Vespidae)

I measured oocyte sizes of Polybia occidentalis workers to assess whether ovary development corresponded with adult age or with individual variation in behavior. Workers exhibited temporal polyethism by first working inside the nest, then performing on-nest tasks, and later foraging and defending the nest. Individuals varied in their ages at first performance of each of these sets of behavioral acts. Ovary development corresponded strongly with adult age. Workers' ovaries declined quickly after an initial phase of partial development. The period of ovary decline overlapped with the age range at which most workers began performing tasks on the exterior nest surface. However, after accounting for age effects, individual behavioral differences (rate of temporal polyethism and performance of foraging and defense tasks) did not correspond with ovary development. These data suggest that ovary development had little or no effect on variation in task performance by P. occidentalis workers on the nest surface.     

Behavioral mediators of cyclical oligogyny in the Amazonian swarm-founding wasp Asteloeca ujhelyii (Vespidae, Polistinae, Epiponini)

Summary In the Epiponini, queen number may vary from many to few during the colonial cycle. This cyclical oligogyny may be important for the maintenance of high genetic relatedness found in several taxa of epiponines. Even though queen reduction in accord with the colony cycle has either been demonstrated or inferred for many taxa, the factors responsible for queen number reduction remain largely unknown. In this study, behavioral interactions (queen-queen and queen-worker) were analyzed in Asteloeca ujhelyii. Conflicts among queens leading to queen elimination were more intense during colony establishment, and queens behavioral performances were significantly correlated with their potential reproductive capacity. Frequency of workers testing behaviors toward queens did not relate as a function of the colony cycle. Thus, it seems that in Asteloeca ujhelyii cyclical oligogyny is indeed behaviorally mediated.Received 16 January 2003; revised 1 June 2003; accepted 3 July 2003.     

Age and Morphological Changes in the Epiponini Wasp Polybia paulista Von Ihering (Hymenoptera: Vespidae)

The division of labor is a central theme in the study of social insects. In bees and wasps, this activity is regulated by age polyethism. Important physiological and morphological changes have been widely studied in the polyethism of honeybee workers. In contrast, this is a relatively unexplored subject in social vespids. Our goal was to determine if there are detectable morphological changes in the body of the Epiponini wasp Polybia paulista Von Ihering or in certain glands in relation to age polyethism. We observed changes in the body weight, the salivary gland, and the mandibular gland that were associated with age, and our results suggest that social relationships and task performance are important to these changes. This contrasts with observations in Polistes and is different from the Apis mellifera Linnaeus age polyethism model.     

Colony cycle of an Australian swarm-founding paper wasp,Ropalidia romandi (Hymenoptera: Vespidae)

Summary Colonies of an Australian swarm founding polistine wasp,Ropalidia romandi, on the Atherton Tableland, Queensland, manage perennial nests by practicing seasonal colony activity. In summer nests, most cells were occupied by immatures of all developmental stages. The proportion of cells that did not contain immatures increased towards winter, when most nest cells did not contain immatures and half of these winter nests had rather large amounts of nectar deposited in empty cells. While the visits to tree flowers byR. romandi were not frequent in summer, nearly 60 % of insects collected on tree flowers in winter wereR. romandi. With such seasonal switching in the modes of colony reproductive activity and of nectar foraging behavior,R. romandi responds to its physical and biotic environment. In this way,R. romandi may be able to continue colonies throughout the year, with much reduced level of colony activity in winter, in areas with seasons that are unfavorable for reproductive activity of a colony.     

Isolation and characterization of 10 microsatellite loci in the paper wasp Polistes rothneyi (Hymenoptera; Vespidae)

We developed 10 polymorphic microsatellite loci for the Japanese paper wasp Polistes rothneyi using the magnetic particle method. Eight of the 10 loci tested were highly polymorphic, having four to eight alleles in P. rothneyi with an expected heterozygosity of 0.60–0.75. They also appeared to be applicable to other related species such as Polistes jokahamae. These loci can be used to study parameters concerning genetic relatedness such as worker reproduction and kin structure.     

Chemical analysis of the swarming trail pheromone of the social wasp Polybia sericea (Hymenoptera: Vespidae)

The behaviour of those polistine wasps which found their nests by swarming, suggests that these species use trail pheromones for leading the swarm to the location chosen for the new nest. Apart from a recent report regarding the ropalidine Polybioides tabidus, where the pheromone is thought to originate from the Dufour gland, nothing is known about the chemistry of such pheromones. Polybia sericea is the only species for which the source of the trail pheromone, the Richards' gland, is known with certainty. The chemistry of the Richards' gland secretion of this species has been investigated in the present work and shown to be a complex mixture where the major compounds are alkyl and aromatic aldehydes, fatty acids, 3-phenylpropanoic acid, ketones, a macrolactone, a pyranone compound and nerolidol.     

Foundress behaviors and interactions in polygynous colonies of the haplometrotic Japanese paper wasp Polistes snelleni (Hymenoptera: Vespidae)

Only two out of 959 pre‐emergence colonies of the paper wasp Polistes snelleni de Saussure surveyed between 1989 and 1996 in Sapporo, northern Japan, were found to be two‐foundress colonies, and the others were single‐foundress colonies. The two foundresses in one of the two colonies showed neither aggressive dominance behavior nor clear division of labor between them during a total of approximately 30 h in the first half of the pre‐emergence stage. Although both of the foundresses foraged for pulp and laid eggs, only one foundress foraged for prey and delivered it to the other.     

Numerical and structural chromosome variation in the swarm-founding wasp Metapolybia decorata Gribodo 1896 (Hymenoptera, Vespidae)

The Neotropical Polistinae wasps are diverse in taxonomy, social behavior, and nesting founding characteristics. Although some species in this group have been used as models for studies on wasp’s biology, they are poorly known in terms of cytogenetics. Here we reported an intraspecific numerical-structural chromosome variation in the swarm-founding wasp Metapolybia decorata from the Brazilian Atlantic Rainforest using conventional and molecular cytogenetic techniques. The observed structural chromosome change involved a telomeric fusion that resulted in a chromosome number range of 2n = 34–36. The origin and geographic distribution of the variant chromosome forms as well as their frequency and maintenance in the studied populations are discussed. In addition, we reported a novel and geographically restricted deletion in the fused chromosomes indicating that the species is undergoing a continued process of karyotype evolution leading to fused chromosome stabilization by elimination of inactive centromeric sequences. Evidence of differences in the telomeric sequences of this wasp was also found by in situ hybridization using the motif (T₂AG₂)₇ as probe.     

Nest thermoregulation in Polybia scutellaris (White) (Hymenoptera: Vespidae)

Polybia scutellaris (White) builds large nests characterized by numerous spiny projections on the surface. In order to determine whether or not the nest temperature is maintained because of homeothermic conditions of the nest individuals or otherwise, we investigated the thermal conditions within the nests built by P. scutellaris. We measured the temperature within active and abandoned nests. The temperature in the active nest was almost stable at 27°C during data collection, whereas the temperature in the abandoned nest varied with changes in ambient temperature. These results suggest that nest temperature was maintained by the thermogenesis of the individuals of the colony. This is the first report of nest incubation caused by thermogenesis of species of Polybia wasps.     

Polymorphic microsatellite loci for primitively eusocial wasp Ropalidia marginata

We report here development and characterization of 48 novel microsatellite markers for Ropalidia marginata, a tropical, primitively eusocial polistine wasp from peninsular India. Thirty‐two microsatellites showed polymorphism in a wild population of R. marginata (N = 38) collected from Bangalore, India. These markers will facilitate answering some interesting questions in ecology and evolutionary biology of this wasp, such as population structure, serial polygyny, intra‐colony genetic relatedness and the pattern of queen succession.