Cuticular hydrocarbons rather than peptides are responsible for nestmate recognition in Polistes dominulus

A colony of social insects is like a fortress where access is allowed only to colony members. The epicuticular mixture of hydrocarbons has been widely reported to be involved in nestmate recognition in insects. However, recent studies have shown that polar compounds (mainly peptides) are also present, mixed with hydrocarbons, on the cuticle of various insects, including the paper wasps of the genus Polistes. As these polar compounds are variable among Polistes species and are perceived by the wasps, this cuticular fraction could also be involved in nestmate recognition. Through MALDI-TOF (Matrix-Assisted Laser Desorption Ionization Time of Flight) mass spectrometry analysis, we assessed, for the first time, the intercolonial variability of the cuticular polar fraction of Polistes dominulus in order to evaluate its reliability as source of nestmate recognition cues. We then tested through behavioral assays the importance of the 2 isolated fractions (apolar and polar) in nestmate recognition by presenting them separately to colonies of P. dominulus. Our results showed that the cuticular polar compounds are not colony specific and they are not used by paper wasps to discriminate nestmates from non-colony members. On the contrary, we confirmed that the isolated cuticular hydrocarbons are the chemical mediators prompting nestmate recognition in paper wasps.     

Thermoregulation in the primitively eusocial paper wasp, Polistes dominulus

Regulation of wing muscle temperature is important for sustaining flight in many insects, and has been well studied in honeybees. It has been much less well studied in wasps and has never been demonstrated in Polistes paper wasps. We measured thorax, head, and abdomen temperatures of inactive Polistes dominulus workers as they warmed after transfer from 8 to ~25°C ambient temperature, after removal from hibernacula, and after periods of flight in a variable temperature room. Thorax temperature (T _th) of non-flying live wasps increased more rapidly than that of dead wasps, and T _th of some live wasps reached more than 2°C above ambient temperature (T _a), indicating endothermy. Wasps removed from hibernacula had body region temperatures significantly above ambient. The T _th of flying wasps was 2.5°C above ambient at T _a = 21°C, and at or even below ambient at T _a = 40°C. At 40°C head and abdomen temperatures were both more than 2°C below T _a, indicating evaporative cooling. We conclude that P. dominulus individuals demonstrate clear, albeit limited, thermoregulatory capacity.     

Wolbachia in the invasive European paper wasp Polistes dominulus

The European paper wasp Polistes dominulus has been expanding its North American range since its introduction in the 1970s. We screened P. dominulus from Italy and the northeastern U.S. for the presence of the intracellular reproductive symbiont Wolbachia. Infection rates among females varied from 16% to 87% among U.S. sites and from 33% to 71% in Italy. We also found infected haploid and diploid males, indicating that this is not a male-killing Wolbachia infection. Our data show that infected individuals from New York, Massachusetts, and Italy carry the same Wolbachia strain, and that some mtDNA haplotypes include both infected and uninfected individuals. We discuss possible implications of Wolbachia infection in this invasive social hymenopteran. Received 28 October 2005; revised 25 January 2006; accepted 17 February 2006.     

Diploid males and their triploid offspring in the paper wasp Polistes dominulus

Although the hymenopteran sex-determining mechanism generally results in haploid males and diploid females, diploid males can be produced via homozygosity at the sex-determining locus. Diploid males have low fitness because they are effectively sterile or produce presumably sterile triploid offspring. Previously, triploid females were observed in three species of North American Polistes paper wasps, and this was interpreted as indirect evidence of diploid males. Here we report what is, to our knowledge, the first direct evidence: four of five early male-producing Polistes dominulus nests from three populations contained diploid males. Because haploid males were also found, however, the adaptive value of early males cannot be ignored. Using genetic and morphological data from triploid females, we also present evidence that both diploid males and triploid females remain undetected throughout the colony cycle. Consequently, diploid male production may result in a delayed fitness cost for two generations. This phenomenon is particularly relevant for introduced populations with few alleles at the sex-determining locus, but cannot be ignored in native populations without supporting genetic data. Future research using paper wasp populations to test theories of social evolution should explicitly consider the potential impacts of diploid males.     

Cuticular hydrocarbon dynamics in young adult Polistes dominulus (Hymenoptera: Vespidae) and the role of linear hydrocarbons in nestmate recognition systems

In social insects, cuticular hydrocarbons (CHCs) play an important role in nestmate discrimination processes, but young individuals are usually not discriminated. We studied CHC changes in young workers of the social wasp Polistes dominulus. A quantitative estimation demonstrated that total quantities of CHCs increased after emergence, with branched alkanes increasing drastically when compared with other classes of hydrocarbons. The relative quantity of longer-chain compounds increased with respect to shorter ones; unsaturated compounds decreased. These changes might reduce the capacity of the cuticle to acquire compounds of environmental origin. We then tested whether individuals acquire hydrocarbons from the environment, and whether this capability equally characterises newly emerged and mature wasps. We exposed wasps of two age classes (adults younger or older than 24 h) to four linear hydrocarbons in turn, and observed how nestmates reacted to their re-introduction into the natal colony. Exposed young wasps elicited significantly more aggressive responses than control sisters; but treated wasps older than 24 h were generally accepted by nestmates. Chemical assays showed that exposed young wasps readily absorbed hydrocarbons; older ones did not incorporate hydrocarbons, suggesting that the chemical profiles of mature wasps are less prone to chemical shifts than those of newly emerged wasps.     

Prey capture and caste-specific payload capacities in the European paper wasp Polistes dominulus

Here, we present a detailed ethogram for the foraging behavior of the eusocial paper wasp, Polistes dominulus. The animal’s foraging process can be sub-divided into four main stages: (1) approach, (2) attack, (3) butchering, and (4) balling. Although considerable behavioral variation exists within each stage, an analysis of more than 20 individual foraging bouts reveals a single “common path” leading from prey approach to its transportation back to the nest; elucidating this path and separating it into distinct stages is useful when identifying targets for future ethological studies. Of particular interest here is the balling phase, i.e., the preparation of a bolus of flesh for transportation back to the colony. Using an experimental approach, we show that foundresses can carry significantly heavier payloads than workers, suggesting a foraging advantage during the initial stages of colony development. We also show that wasp body mass is significantly positively correlated with payload capacity in foundresses, a relationship not seen among workers or late reproductives. This correlation suggests a beneficial adaptation of foundresses for combating early season pressures associated with the foundation of a new colony.     

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.     

Cuticular lipids as trail pheromone in a social wasp

We investigated the origin and composition of the chemical trail of the common yellow jacket Vespula vulgaris L. (Vespidae) and found that an artificial trail made from an extract of cuticular lipids from V. vulgaris foragers was biologically as active as a trail laid naturally by the foragers. Chemical analysis of natural trail extracts and the behaviourally active cuticular extracts by coupled gas chromatography-mass spectrometry revealed that the majority of cuticular hydrocarbons were also present in the trail extract at similar ratios. Thus, bioassay data and chemical analysis provide strong evidence that these cuticular hydrocarbons act as a trail pheromone in V. vulgaris.     

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.     

The effect of juvenile hormone on temporal polyethism in the paper wasp Polistes dominulus

Juvenile hormone (JH) has an important role in the behavior of eusocial Hymenoptera. Previous work has shown that JH influences aggression and dominance behavior in primitive eusocial insects that lack discrete queen and worker castes (e.g. Bombus bees and Polistes wasps). In contrast, JH is one of the factors that mediates temporal polyethism among workers in advanced eusocial insects that have reproductive castes (e.g. Apis bees and Polybiawasps). Therefore, initial observations suggest that JH may have different roles in primitive and advanced eusocial taxa. Here, we use detailed behavioral observations of marked individuals to test whether JH influences temporal polyethism in the primitive eusocial wasp Polistes dominulus. First, we show that workers in P. dominulus have an age-related division of labor, as workers switch from nest work to foraging as they mature. Then, we show that application of JH accelerates the onset of foraging behavior.Workers treated with JH start foraging at a younger age than control workers. Therefore, JH mediates temporal polyethism in the primitively eusocial insect Polistes dominulus. Received 23 April 2008; revised 6 August 2008; accepted 11 August 2008     

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.     

On status badges and quality signals in the paper wasp Polistes dominulus: body size, facial colour patterns and hierarchical rank

To establish a dominance order, social animals often rely on indicators of fighting to avoid costly aggressive encounters. In some species, individuals use colour patterns to signal their social status. Recent studies claimed that facial markings in the eusocial paper wasp Polistes dominulus are status badges that allow co-foundresses to form a linear hierarchy based on individual quality. Here, we evaluated facial patterns in natural populations of P. dominulus, in its native range, to observe whether the marks reflect overall wasp quality in different contexts. We used the same measures of clypeus patterns used by earlier studies, but did not find that they functioned as status badges. Our analyses showed no evidence that visual markers are related to: (i) size, (ii) probability of surviving winter, (iii) social rank in spring associations, or (iv) health status (assessed by the presence of strepsipteran endoparasites). Size, however, is important. Larger wasps are more likely to survive the winter and to acquire the dominant position in spring associations. Larvae infected with endoparasites become smaller adult wasps. These findings suggest that body size is a reliable quality indicator on which wasps build their social networks, and that clypeus patterning is not involved.     

Timing of insemination in terms of age of new reproductive females of a temperate-zone polistine wasp,Polistes snelleni (Hymenoptera: Vespidae)

Summary The present study investigated the age at which insemination takes place in Polistes snelleni, a species in which mating begins while new reproductives are still emerging from the nest. Insemination of new reproductive females began as early as 5 days of age, and most females were inseminated by 10 days of age.     

Costs and benefits of colony aggregation in the social wasp, Polistes annularis

Females of the social wasp, Polistes annularis, nest in largeaggregations onriverside cliffs in central Texas. Aggregationsoccur where there is an overhang under which wasps build theirnests. Colony aggregations appear to occur partly because individualfemales do notdisperse far from the natal site where most coloniesare successful. In most taxa greater reproductive success nearthe natal nest site is related to unique physical features orto advantages ofbeing in a group. In P. annularis, the physicalfactors and not the social factors appear to be most important.Advantages to remaining near the natal nest site include protectionprovided by the cliff overhang from rain and afternoon sun.Central colonies are as likely to suffer predation as are peripheralcolonies, which indicates an absence of selfish herd advantagesto being in the center of the group. A cost of being in a denseaggregation is often increased reproductive competition, andthis is the case for P. annularis. Colonies in the center ofthe aggregation suffer high usurpation rates by unrelated femalesand in 1 of 3 years produced fewer reproductives than edge colonies.     

Benefits of foundress associations in the paper wasp Polistes dominulus: increased productivity and survival, but no assurance of fitness returns

Successful Polistes dominulus nests can be started by one ormore nest founding queens (foundresses). Consequently, thereis much interest in the specific benefits that induce cooperationamong foundresses. Here, we experimentally demonstrate one majorbenefit of cooperation, namely that multiple foundresses increasecolony productivity. This increase is close to the value predictedby subtracting the productivity of undisturbed single-foundresscolonies from the productivity of undisturbed multiple-foundresscolonies. However, we found no evidence that an associatingfoundress' contribution to colony growth is preserved if shedisappears (assured fitness returns). Our correlational datasuggest that cooperation provides survival benefits, multiple-foundresscolonies are more likely to survive to produce offspring thanare single-foundress colonies, and individual foundresses inmultiple-foundress groups are less likely to disappear beforeworker emergence than foundresses nesting alone. Therefore,association provides substantial productivity and survival benefitsfor cooperating foundresses.