Reproduction in foundress associations of the social wasp, Polistes carolina: conventions, competition, and skew

Who reproduces in colonies of social insects is determined bysome combination of direct competition and more peaceful convention.We studied these two alternatives in foundresses of the paperwasp, Polistes carolina, by examining two different contexts:what determines who becomes the dominant reproductive and whatdetermines the amount of reproduction obtained by subordinates.The dominant queen on most nests was the foundress to arrivefirst, rather than the largest foundress, expected to be bestat fighting. This suggests that dominance is initially determinedby convention, although the persistence of some aggressiveconflict throughout the foundress period suggests that thisconvention is not absolute. Attempts to explain the divisionof reproduction using several skew theories were generallyunsuccessful. Skew was not correlated with relatedness, size differences, colony productivity, and challenges by the subordinate.P. carolina showed high constraints against solitary nesting,with a minority of females attempting to nest alone, and nonesucceeding. In this situation, most skew theories predict thatgroup stability will be independent of relatedness, yet nearlyall collected subordinates were full sisters to the queen.Reproductive partitioning in early P. carolina colonies may have more to do with enhancing worker production than with conflictover direct fitness.     

Viability of diploid males in the parasitic wasp,Diadromus pulchellus (Hym.: Ichneumonidae)

In the HymenopteraDiadromus pulchellus diploid males have been observed in samples collected in the wild and bred in the laboratory. Using a yellow body color mutant strain, a protocol of crosses, involving inbred individuals, allows the routine production of such diploid males. These males result from the development of fertilized ova and emerge normally. Their preimaginal viability is similar to those of other individuals, and their imaginal viability does not differ from that of haploid males. Diploid males present normal external morphology and neither mosaicism nor intersexuality was observed. However, they are bigger than haploid males and have head size and wing length similar to those of females. The significance of diploid male viability in hymenopteran populations is discussed.     

Worker interests and male production in Polistes gallicus, a Mediterranean social wasp

The resolution of social conflict in colonies may accord with the interests of the most numerous party. In social insect colonies with single once-mated queens, workers are more closely related to the workers' sons than they are to the queens' sons. Therefore, they should prefer workers to produce males, against the queen's interests. Workers are capable of producing males as they arise from unfertilized eggs. We found Polistes gallicus to have colonies of single, once-mated queens, as determined by microsatellite genotyping of the workers, so worker interests predict worker male production. In colonies lacking queens, workers produced the males, but not in colonies with original queens. Thus worker interests were expressed only when the queen was gone. The high fraction of missing queens and early end to the colony cycle relative to climate so early in the season is surprising and may indicate a forceful elimination of the queen.     

Cooperation between non-relatives in a primitively eusocial paper wasp,Polistes dominula

In cooperatively breeding vertebrates, the existence of individuals that help to raise the offspring of non-relatives is well established, but unrelated helpers are less well known in the social insects. Eusocial insect groups overwhelmingly consist of close relatives, so populations where unrelated helpers are common are intriguing. Here, we focus on Polistes dominula—the best-studied primitively eusocial wasp, and a species in which nesting with non-relatives is not only present but frequent. We address two major questions: why individuals should choose to nest with non-relatives, and why such individuals participate in the costly rearing of unrelated offspring. Polistes dominula foundresses produce more offspring of their own as subordinates than when they nest independently, providing a potential explanation for co-founding by non-relatives. There is some evidence that unrelated subordinates tailor their behaviour towards direct fitness, while the role of recognition errors in generating unrelated co-foundresses is less clear. Remarkably, the remote but potentially highly rewarding chance of inheriting the dominant position appears to strongly influence behaviour, suggesting that primitively eusocial insects may have much more in common with their social vertebrate counterparts than has commonly been thought.     

Factors affecting the acceptance of alien conspecifics on nests of the primitively eusocial wasp,Ropalidia marginata (Hymenoptera: Vespidae)

In the primitively eusocial wasp, Ropalidia marginata,individual females are known to drift from one newly founded nest to another. In the laboratory, young (<6- to 8-day-old) alien wasps are accepted onto unrelated colonies, while older (>6- to 8-day-old) wasps are not. Here we have investigated the factors that could influence the acceptance of foreign conspecifics onto unrelated nests. Individually marked wasps of different ages, isolated immediately after eclosion from the natal nest and from each other, were introduced onto unrelated recipient nests. Considered separately, both age and ovarian condition seemed to influence the probability of acceptance as well as the levels of aggression and tolerance received by the introduced wasps. However, partial correlation analysis and multiple regression analysis indicated that only age had a direct influence and that the ovarian condition acts only through age, a variable with which it is highly correlated. The observed acceptance of young aliens and rejection of old aliens are less likely to be due to the perception of older wasps as a reproductive threat rather than some age-related factor, other than ovarian condition, for example, the relative ease with which younger wasps can be molded into desired roles.     

The influence of temperature on nest site choice and reproductive strategy in a temperate zone Polistes wasp

Abstract. 1. Field experiments with Polistes fuscutus showed that more nests were initiated, co-foundress groups were larger, broods developed faster, and workers eclosed earlier in sun-warmed sites than in cool ones.
2. These results suggest that colonies nesting in sun-warmed shelters gained a fitness advantage over those in cooler shelters. More rapid brood development means that such colonies reach the ergonomic stage more quickly, which in turn means a higher expectancy of survival and greater productivity at the end of the season.
3. We argue that thermal properties constitute an important component of Polistes nest site quality in temperate regions and that these properties influence joining and usurping strategies as well as nest site choice in spring Polistes females. Other factors being equal, a joining or usurping female should opt for a colony in a warm site over one in a cool site, leading to the larger foundress groups we found in warm shelters.
4. We hypothesize that the selective advantage of philopatry in temperate zone Polistes populations is that it brings spring females to nest sites of proven quality.     

Prey diversity of Polistes rothneyi koreanus in different landscapes using DNA barcoding

For the present study, we investigated the prey of Polistes rothneyi koreanus, which is the most common social wasp in South Korea, and the relationship between prey diversity and vegetation cover around their nests. Prey was collected over two 6‐hr sampling from seven nests between July and mid‐August 2015 in Daegu, Gyeongsan and Gunwi in South Korea. To analyse the prey spectrum, we identified species using DNA barcodes; to analyse vegetation cover, we used the normalized difference vegetation index in a 200 m radius around the nests. A total of 338 prey samples were collected, and eight orders, 24 families, and 65 species were identified, demonstrating a much broader prey spectrum than those previously recorded for Polistes spp. Lepidoptera were the most prevalent, with 158 samples and 47 species. Nest 7, located in a rural area, had the highest numbers of samples and species per worker (5.2 and 1.9, respectively). Lepidoptera accounted for over half of the prey spectrum for all nests, and the families Noctuidae and Geometridae accounted for 60% of all Lepidoptera. Tenodera sinensis (Mantodea) and Gabala argentata (Noctuidae) were the most ubiquitous species, collected at five locations. Six species and some genera of prey are designated as pests in South Korea, indicating that P. r. koreanus also has a beneficial role in pest control. A higher vegetation cover was associated with significantly higher prey species diversity (R² = .4597, p < .1) and abundance (R² = .5986, p < .05), indicating that vegetation cover is an important factor for maintaining colonies. Therefore, the recent increase in green spaces in South Korean cities is probably a major contributor to the increased density of social wasps.     

Application of microsatellite primers for the social wasp Polistes to another social wasp genus, Parapolybia, to estimate genetic relationships among nestmates

We tested 14 pairs of microsatellite primers that had been developed for the genus Polistes to assess their applicability for determining genetic structure in a colony of the genus Parapolybia. At least one of the 14 Polistes primer sets was useful for Parapolybia indica, and the locus identified by this primer set was highly polymorphic. Using this primer set, we estimated the average relatedness among female nestmates of Parapolybia indica to be 0.82 ± 0.05 (mean ± SE), indicating that in this species single mating is normal, although some cases of multiple mating was found.     

Circulating hemocytes from larvae of the paper wasp Polistes dominulus (Hymenoptera, Vespidae)

Circulating hemocytes from larval stages of the paper wasp Polistes dominulus were characterized by light and transmission electron microscopy. Three types were identified: prohemocytes, plasmatocytes and granulocytes. The first two are agranular cells while the latter present typical cytoplasmic inclusions called granules. Plasmatocytes differ from prohemocytes being larger, showing lower nucleus/cytoplasm ratio and they possess many phagolysosomes. The substantial uniformity of most subcellular features and the presence of "intermediate forms" support the "single-cell theory" i.e., there is only one cell line that originates from the prohemocyte and leads to the granular cell passing through the plasmatocyte. This hypothesis seems to be confirmed by functional tests. Indeed, most part of cells adheres to the glass and is able to phagocytize fluorescent microspheres.     

Control of male production in the swarm‐founding wasp,Polybioides tabidus

In social insect colonies, male production may involve conflicts over the sex ratio, worker vs. queen reproduction, and each queen's contribution to the males when there are multiple queens. We examined male production in the swarm‐founding, multiple‐queened wasp, Polybioides tabidus, for which previous work suggested worker control of the sex ratios. We found that queens produced the males in accord with the collective worker preference. We also found that diploid males were produced, but only in association with haploid males. Simulations show they should have been produced in other colonies as well and their absence indicates that they were killed in some of these other colonies. The pattern of their removal indicates that P. tabidus cannot distinguish diploid from haploid males, and that haploid males would have been removed from these colonies too. This provides evidence that the workers are able to manipulate male production when collective preferences dictate.     

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.     

Colony cycle in the south-eastern coastal populations of Ropalidia plebeiana, the only Ropalidia wasp occurring in temperate Australia

Of the 22 species of Ropalidia occurring in Australia, only R. plebeiana is distributed in temperate areas. This wasp is abundant in the south‐eastern coastal regions of Australia, where it forms huge nest aggregations. Based on our observations in late autumn and early spring, as well as information in the literature, we outline the colony cycle of R. plebeiana in the south‐eastern coastal area of New South Wales. The nesting period lasts for approximately 9.5 months, which is unusually long for temperate polistine wasps. Most foundresses seem to return to their natal nests in early spring (mid‐August to early September) to start their own colonies by reusing pre‐existing nest combs, and the pre‐emergence stage (before the emergence of the first adult brood) lasts for approximately 3.5 months. Reproductives (gynes and males) start to emerge as adults in early March, and the reproductive‐producing period lasts until the end of May. Both the pre‐emergence stage and the reproductive‐producing period are much longer than in other temperate polistine wasps, which is discussed in terms of fluctuating temperature conditions in the area and the formation of nest aggregations, a peculiar habit for temperate populations of R. plebeiana.     

Inbreeding in a natural population of Euodynerus foraminatus (Hymenoptera: Vespidae), a solitary wasp with single-locus complementary sex determination

The solitary wasp Euodynerus foraminatus has single-locus complementary sex determination (sl-CSD), which is normally incompatible with inbreeding because it increases the production of sterile or inviable diploid males. Previous field observations of E. foraminatus have suggested that high levels of sibling mating are present in this species. However, conclusions about inbreeding and its genetic consequences could be flawed if based solely upon behavioural observations. Through microsatellite DNA genotyping of 102 E. foraminatus females in southwest Michigan, we estimate that between 55% and 77% of the matings in this population take place between siblings, but the frequency of diploid males is lower than expected. Our data suggest that a mixture of inbreeding and outbreeding persists in E. foraminatus despite the presence of sl-CSD.     

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).     

Nest surface hydrocarbons facilitate nestmate recognition for the social wasp,Polistes metricus Say (Hymenoptera: Vespidae)

Nestmate recognition was observed inPolistes metricus Say workers only if the workers were exposed to their nest surface hydrocarbons after eclosion. If the newly emerged workers were never exposed to the nest hydrocarbons as adults, they showed no discrimination between nestmates and nonnestmates. Furthermore, the newly emerged workers were accepted more readily by their experienced nestmates than by experienced nonnestmates only if the newly emerged wasps had been exposed to the nest surface hydrocarbons. This reciprocal recognition implies that the nest recognition cues are nest surface hydrocarbons that are learned and that may be acquired byP. metricus workers as adults on the natal nest.     

Genetic relatedness in early associations of Polistes dominulus: from related to unrelated helpers

Indirect benefits obtained through the reproduction of relatives are fundamental in the formation and maintenance of groups. Here, we examine the hypothesis that females of the temperate paper wasp Polistes dominulus preferentially form groups with close relatives. Genetic relatedness data were obtained for 180 groups of females collected at the early stages of the nesting cycle of a large population of P. dominulus in two sites in southwestern Spain. Average within-group relatedness values ranged from 0.189 to 0.491. Foundresses on early nests were significantly more closely related than females in winter aggregations or in stable groups (just before workers emerged). Within-group relatedness values were independent of group size. The vast majority of worker-producing nests ( c . 85%) had one or more females that were unrelated (or distantly related) to the remaining members of the group. These results provide further support to the hypothesis that indirect fitness benefits alone are unlikely to explain why P. dominulus foundresses form cooperative associations.     

Social behavior of early emerging males of a Japanese paper wasp,Polistes chinensis antennalis (Hymenoptera: Vespidae)

Summary Five cases of the early emergence of males in a Japanese paper wasp,Polistes chinensis antennalis, in which male emerged together with the first group of workers, were described. In one case of the five where the queen disappeared before the emergence of male, worker(s) produced female offspring. The frequency of the nests where the early emergence of male was observed was 16.7% (5/30 nests). In two colonies, worker(s) and/or queen chased off males. But in an orphan nest where worker(s) produced female offspring, the dominance order among workers which was similar to that of colony without male was observed. The significance of the early emergence of male in the social evolution of wasps was discussed.     

Mate‐searching behavior in the invasive German wasp,Vespula germanica,in Patagonia

In order to increase the probability of reproduction, social insects can adopt various mate‐finding strategies, such as increasing densities of males at specific locations, and/or visual and chemical cues that attract the opposite sex. In field and laboratory studies we investigated strategies used by the invasive eusocial wasp Vespula germanica (Fabricius) (Hymenoptera: Vespidae). In tethered flight assays, we established contrasting flight patterns in females and males that may partly explain how related individuals distribute spatially during the mating period. We also determined experimentally, in the field and in the laboratory via olfactometer assays, that gynes produce airborne pheromonal cues that attract drones and are important during mate location. Our field trials also suggest that visual cues play a role in mate location. We conclude that in addition to aspects of the social biology of the species, an efficient mate‐location strategy can partly explain the invasion success of the species. Tools to mitigate the damage caused by yellowjackets may be developed by focusing on reproductive castes, in addition to workers.     

A nocturnal Provespa wasp species as the probable pollinator of epiphytic orchid Coelogyne fimbriata

Many vespid wasps visit flowers to forage nectar. These hymenopterans sometimes contribute to flower pollination. However, none of the nocturnal wasp species is a known pollinator. We collected individuals of light‐attracted Provespa nocturna workers in a montane rainforest on Peninsular Malaysia: some wasps collected bore orchid pollinia on their thoraxes. Among 114 trapped individuals, four bore pollinaria and nine bore only viscidia, suggesting that pollinia had been successfully transported. Molecular barcoding of the pollinia (based on their ITS sequences) assigned the orchid to a species in Coelogyne fimbriata complex. These findings and our other analyses suggest that this nocturnal wasp contributes to pollination of an epiphytic nectarless orchid that probably releases olfactory attractants. This discovery sheds light on the importance of mutualistic relationships between the nocturnal social wasps and epiphytic orchids in Southeast Asian tropical rainforest canopies.