The comparative biology of two sympatric paper wasps, the native Polistes fuscatus and the invasive Polistes dominulus (Hymenoptera, Vespidae)

Summary: Polistes dominulus (Christ), an old world paper wasp, was introduced accidentally into the eastern coast of the United States in the late 1970s and has been rapidly spreading westward, displacing the native P. fuscatus (F.). The biology of naturally nesting P. fuscatus and P. dominulus was compared at a field site in Rochester, Michigan. The basic methodology consisted of simultaneously videotaping spatially proximate, matched single-foundress colonies of P. fuscatus and P. dominulus (13 matched sets, 176.8 h of videography). In addition, extensive surveys and censuses were taken of colonies to record colony productivity, dates of nest initiation and first worker emergence, usurpation and parasitism.¶There was no evidence that P. dominulus is negatively impacting P. fuscatus through direct, agonistic encounters. However, P. dominulus is 4-5 times more productive than P. fuscatus, suggesting that P. dominulus is replacing P. fuscatus via exploitative competition. P. dominulus appears to have a number of advantages over P. fuscatus, including earlier production of workers, higher per capita foraging rates by queens and workers, higher queen survivorship, and lack of conspecific pressures. Nest site and/or prey availability may be limiting factors in the competition between the two species.     

The comparative biology of two sympatric paper wasps in Michigan,the native <Emphasis Type="Italic">Polistes fuscatus</Emphasis> and the invasive <Emphasis Type="Italic">Polistes dominulus</Emphasis> (Hymenoptera,Vespidae)

Summary The biology of the invasive Polistes dominulus and the native P. fuscatus was compared at a field site in Rochester, Michigan over a two-year period. Colonies nesting semi-naturally in plywood nestboxes were studied using videography, extensive surveys, and colony-specific marking of gynes.Both single- and multiple-foundress colonies of P. dominulus were significantly more productive than comparable colonies of P. fuscatus. The disparity in productivity was significantly more pronounced in single-foundress colonies than in multiple-foundress colonies. P. dominulus had significantly shorter larval and pupal development times than P. fuscatus, which allowed P. dominulus to produce its first workers about a week earlier than P. fuscatus. P. dominulus had a number of additional advantages over P. fuscatus that contributed to its productivity including (1) significantly less parasitism by Strepsiptera, (2) significantly greater probability of renesting after predation by raccoons, (3) significantly lower usurpation pressures, and (4) possibly longer foraging days . The recovery of colony-marked foundresses indicated that gynes of P. dominulus suffered significantly greater mortality than gynes of P. fuscatus during winter diapause and that foundresses of both species were equally, strongly philopatric.P. dominulus is likely replacing P. fuscatus in many areas of southeastern Michigan via indirect or exploitative competition. The two species may be competing for nest sites.Received 7 July 2003; revised 10 October 2003; accepted 3 November 2003.     

The cost of flight: a role in the <Emphasis Type="Italic">Polistes dominulus</Emphasis> invasion

Polistes dominulus is a primitively eusocial paper wasp from Mediterranean Europe that is invasive to North America. In Eastern North America, P. dominulus is in competition with P. fuscatus. One reason for the success of P. dominulus is that their colonies produce more reproductive offspring than P. fuscatus colonies. A partial explanation for this difference is that P. dominulus foundresses make more foraging trips in the pre-worker period, which likely helps them to rear workers more quickly. In comparing the species, we found that P. dominulus had a lower absolute flight metabolic rate, but that P. fuscatus had a lower mass-specific flight metabolic rate. In addition, in P. fuscatus, wingloading correlated with flight metabolic rate, but that this was not the case in P. dominulus. This suggests that P. fuscatus is not able to transport large loads inexpensively. Therefore, the lower overall cost of transport of P. dominulus may provide an advantage by allowing the foundresses to make more relatively efficient foraging trips. In addition, we compared time in flight by P. dominulus and P. fuscatus over a range of temperatures and found that while P. fuscatus flew well over a broad range of temperatures, P. dominulus had a relatively narrow range of optimal temperatures for flight (30–33°C). These differences may help explain both the success and the limitations of the P. dominulus invasion.     

Comparison of intraspecific nest usurpation between two haplometrotic paper wasp species (Hymenoptera: Vespidae:Polistes)

Intraspecific nest usurpation by foundresses was studied in 2 haplometrotic (solitary founding) species of different subgenera,Polistes (P.) riparius andP. (Polistella) snelleni, in areas where they cohabited. The overall probability for a nest to be usurped by a foreign foundress during the season was about twice as large inP. snelleni as inP. riparius. In both, however, probability of usurpation was largest on late pre-emergence nests, or in late June and early July. InP. riparius, all the usurpers of known origins were those foundresses that had lost their pre-emergence nests to destruction probably by some vertebrates; inP. snelleni, some usurpers had the same history as above, while the others had lost many of all larvae to predation by unknown agents before worker emergence. Usurpers of both species destroyed eggs and younger larvae to much greater extents than older larvae or pupae, and they produced fewer numbers of reproductives in comparison with non-usurping foundresses. We concluded that usurpation behavior has been maintained despite its relatively low productivity because renesting would lead to even lower or no reproductive production.     

The Polistes war: weak immune function in the invasive P. dominulus relative to the native P. fuscatus

Invasive species are of growing ecological concern, in part because of conflicts arising with native congeners. The European paper wasp Polistes dominulus was first introduced to North America in the 1970s, and may be displacing at least one native species, P. fuscatus. Previous reports indicate that in native territories over half of P. dominulus colonies are infected by Strepsipteran parasites, which decrease host fitness. In North America, P. fuscatus are parasitized to a lesser degree (approximately one-third), but no infected colonies of invasive P. dominulus have been reported. Because immune function is an indicator of susceptibility to parasitism, we quantified activated levels of immune function by measuring the encapsulation response and phenoloxidase activity and then compared these levels between species. Counterintuitively, our results indicate that P. dominulus has lower levels of both mechanisms of immunity. Additionally, P. dominulus displayed less self-grooming activity than P. fuscatus. We briefly discuss possible immunological explanations for this invasion success, including the selective expression of low immunocompetence.     

A role for thermoregulation in the <Emphasis Type="Italic">Polistes dominulus</Emphasis> invasion: a comparison of the thermoregulatory abilities of the invasive wasp <Emphasis Type="Italic">P. dominulus</Emphasis> and the native wasp <Emphasis Type="Italic">P. fuscatus</Emphasis>

Social insects are excellent invaders that have had negative impacts on native species and humans. Many invasive species move from warmer to cooler climates. For these species, thermal adaptations may both be important for their ability to invade and to limit their invasion range. The invasion of Polistes dominulus into North America provides an example of a primitively eusocial invader from a warmer climate. We studied the differences in thermoregulation between P. dominulus and the native P. fuscatus. We found that, during flight, thorax temperature in P. fuscatus was less affected by ambient temperature than thorax temperature of P. dominulus. We also found that P. dominulus and P. fuscatus showed different patterns of warming after removal from a cold environment. Unlike P. dominulus, live P. fuscatus never fully cooled down in a cold environment. P. fuscatus also reached their relative minimum flight temperatures earlier than P. dominulus, but P. dominulus maintained higher elevated temperatures for longer. These differences in thermoregulatory ability suggest that the lower winter survival of P. dominulus could be offset by a greater thermal tolerance during flight, while the lower thermal tolerance of P. fuscatus in flight is offset somewhat by better thermoregulatory ability.     

A ten-year comparative study of the population dynamics and parasitoidism in the native paper wasp Polistes fuscatus and the invasive P. dominulus

The invasive success of Polistes dominulus in North America has been attributed to its greater productivity relative to native Polistes. Liberation from parasites and parasitoids are thought to be major factors contributing to the high productivity of P. dominulus. We analyzed historical records of colony relative abundance and productivity of P. dominulus and the sympatric, native Polistes fuscatus from 1995 to 2010 using historical data from our Michigan Polistes study site. We also analyzed evidence of parasitoids from 294 P. fuscatus and 507 P. dominulus archived combs from 2001 to 2010. Additionally, we examined field and laboratory colonies from outside of our study site for parasites and parasitoids in 2009 and 2010. We documented one parasite and three parasitoids exploiting Polistes in our Michigan study sites. Our historical records document that P. dominulus initially displaced P. fuscatus rapidly, then slowed, and finally the two populations stabilized. Furthermore, the historical pattern of decreasing displacement of P. fuscatus by P. dominulus corresponded temporally with a significant decline in the productivity and a significant increase in Dibrachys cavus infestation of P. dominulus. Our evidence indicates that the parasitoid, D. cavus, is a major factor in stabilizing the populations of the sympatric P. dominulus and P. fuscatus.     

Social parasitism of <Emphasis Type="Italic">Polistes dominulus</Emphasis> by <Emphasis Type="Italic">Polistes nimphus</Emphasis> (Hymenoptera,Vespidae)

Summary Interspecific facultative social parasitism is well known in ants and in bumble-bees, but it is rarer in wasps. This form of parasitism is traditionally considered to be an intermediate stage in the evolution of obligate interspecific parasitism, where the parasites are no longer able to nest alone. We report field and experimental observations of a newly discovered facultative parasitic relationship between two closely related free-living Polistes species: P. nimphus and P. dominulus. P. nimphus foundresses sometimes usurp the nests of the larger P. dominulus before worker emergence. The invading queen takes over the nest with abundant abdomen stroking on the nest surface and is accepted by workers if they emerge 6 or more days after usurpation. Morphometric comparisons show that the usurper species, though smaller than its victims, has morphological adaptations consisting of larger heads, mandibles and front femora relative to their body size that may give it an advantage during nest invasion. This strategy is likely to be taken only after the foundress loses her original nest because invading P. nimphus queens have lower reproductive success than they would have had on their own nest. Overall, we found that P. nimphus usurpers use strategies of invasion similar to those of two obligate parasites, suggesting that this may be an example of one of the pathways by which social parasitism evolved.Received 4 April 2003; revised 8 August 2003; accepted 14 September 2003.     

Rearing conditions influence quality signals but not individual identity signals in Polistes wasps

Given the diversity of animal signals, there has been recentinterest in categorizing signals into probable functions accordingto their properties. For example, models predict that signalsof quality should be costly and condition dependent, whereassignals of individual identity should be cheap and expressedindependently of condition. Here, we test these predictionsby comparing the condition dependence of signals of individualidentity and quality in Polistes wasps. Polistes fuscatus waspshave black and yellow patterns on the face and abdomen thatsignal individual identity, whereas Polistes dominulus waspshave black and yellow facial patterns that signal aspects ofquality related to dominance. We reared both species with andwithout supplemental food and examined the facial patterns ofthe resulting offspring. As predicted, food availability didnot influence the development of identity signals in P. fuscatus.In strong contrast, P. dominulus wasps reared with supplementalfood had facial patterns that signaled higher levels of qualitythan P. dominulus reared without supplemental food. Interestingly,the identity and quality signals have different condition dependence,despite being composed of similar pigments, suggesting thatsignal function has a stronger influence on signal propertiesthan pigmentation. Because body size is often correlated withquality signal elaboration, we also tested how food supplementationinfluenced offspring size. In both species, supplemented coloniesproduced smaller offspring than nonsupplemented colonies, suggestingthat queens may invest in producing fewer, larger offspringin stressful environments.     

Reproductive plasticity in Polistes paper wasp workers and the evolutionary origins of sociality

Regulatory pathways in solitary species provide the raw materials for the evolution of sociality. Therefore, comparing the mechanisms that mediate reproductive plasticity in social species and their solitary ancestors can provide insight into the evolutionary origin of sociality. In many solitary insects, the effect of juvenile hormone (JH) on fertility is mediated through the fat body; individuals in good physical condition show a stronger fertility response to JH than individuals in poor physical condition. Here, we test whether a similar, condition-dependent JH response mediates fertility in workers of the primitively eusocial Polistes dominulus wasps. We test how body weight, JH, and adult nutrition influence worker ovarian development. Both JH-treatment and adult nutrition dramatically increased ovarian development. Body weight also influenced ovarian development, as large workers developed more eggs than smaller workers. Body weight and fat are strongly linked in P. dominulus workers, so these results suggest that the fat-dependent JH responsiveness common in solitary insects is conserved in social wasps. The simple, ancestral relationship between reproductive investment and physical condition may facilitate cooperation by allowing workers to adaptively allocate energy into reproduction based on their probability of successfully becoming a queen.     

Intra‐specific body size variation in Polistes paper wasps as a response to social parasite pressure

1. Like avian brood parasites, obligate insect social parasites exploit the parental care of a host species to rear their brood, causing an evident loss of host reproductive success. This fitness cost imposes selective pressure on the host to reduce the parasite effect. A possible outcome of an evolutionary arms race is the selection of host morphological counter‐adaptations to resist parasite attacks. 2. We studied host–parasite pairs of Polistes wasps in which the fighting equipment of the parasite's body allows it to enter the host colony. 3. We searched for host morphological traits related to fighting ability that could be considered counter‐adaptations. As a host–parasite co‐evolutionary arms race can only occur where the two lineages co‐exist, we compared morphological traits of hosts belonging to populations with or without parasite pressure. We report that host foundresses belonging to populations under strong parasite pressure have a larger body size than those belonging to populations without parasite pressure. 4. Behavioural experiments carried out to test if an increase in host body size is useful to oppose parasite usurpation show that large body size foundresses exhibit a greater ability of nest defence.     

Colony structure and reproductive sharing among queens in a tropical paper wasp, <Emphasis Type="Italic">Polistes olivaceus</Emphasis>

Reproductive partitioning among group members is a key feature in social Hymenoptera. We investigated the genetic colony structure of a tropical paper wasp Polistes olivaceus, with an emphasis on variation in the number of queens and reproductive sharing among queens. Among 22 P. olivaceus colonies, 6 were monogynous, 9 polygynous, and 7 were queenless. Adults and brood (eggs and larvae) were genotyped based on six polymorphic microsatellite loci. In each of the polygynous colonies, progenies were assigned to their mothers using maximum-likelihood methods. Nestmate queens were full sisters. The vast majority of reproduction appeared to be monopolized by the dominant queen (α), and the overall reproductive skews were 0.63 ± 0.04 (B index) and 0.97 ± 0.02 (S _c index). Although all nestmate queens had equal reproductive potential, the high magnitude of reproductive skew was enigmatic in this species. Although 9.55 ± 2.07 workers contained developed ovaries in 11 of 15 queen-right colonies, they were unrelated to the nestmate queens but related to each other as full sisters, suggesting that they were the remaining offspring of superseded queens. In 2 of the 11 colonies, we detected male eggs produced by reproductive workers. On average, 7.27 % of the total genotyped male eggs were derived from reproductive workers among the colonies. These results suggest three possibilities regarding the presence of reproductive workers in the P. olivaceus colonies: drifting between colonies, putative remaining offspring from superseded queens, and the offspring of unrelated females who joined the colonies and reproduced there. We found no worker-derived larvae or adult males, suggesting that male eggs were removed by nestmates at some point between oviposition and hatching.     

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     

The Mechanism of Queen Regulation of Foraging by Workers in Paper Wasps (Polistes fuscatus,Hymenoptera: Vespidae)

We examined how queens of the primitively eusocial wasp, Polistes fuscatus, stimulate foraging by workers in 10 small, post-worker-emergence field colonies. We experimentally increased colony needs, including needs of the brood, by removing a colony's most active foragers (thereby decreasing the colony's foraging rate), and found that the queen significantly increased both her level of activity and rate of aggressive interactions. Most aggressive interactions were directed at dominant workers. Removal of a colony's least active foragers, however, produced no such effect. Our results, together with those of Reeve & Gamboa (1983, 1987), indicate that queens are sensitive to brood needs, and that they behaviorally regulate worker foraging to match brood needs by increasing their level of activity and rate of aggressive interactions.     

Changes in the temporal pattern of antennal drumming behavior across the <Emphasis Type="Italic">Polistes fuscatus</Emphasis> colony cycle (Hymenoptera,Vespidae)

Social wasps in several genera exhibit a diverse array of conspicuous vibrational behavior patterns closely associated with larval feeding. Polistes, as the only genus in which these substrate-borne mechanical signals have been studied in some detail, is a useful system for understanding their functions. Most Polistes species examined perform antennal drumming (AD) in the context of feeding prey liquid to larvae. Two existing hypotheses on the function of AD propose that it is a behavioral releaser signal that regulates the release of larval saliva, but with opposite effects. One proposes that AD stimulates larvae to release their saliva for the drumming adult to imbibe, whereas the other proposes that AD inhibits saliva release. A recently proposed third hypothesis argues instead that AD has a modulatory effect on development: exposure to high levels of AD biases larvae toward a worker phenotype as adults. While the larval-saliva-release hypothesis for AD function has little support, predictions made by both the inhibition hypothesis and the mechanical switch hypothesis are yet to be tested within the broader ontogenetic framework of the Polistes colony cycle. We investigated the contexts, rates of performance, and actors associated with AD across 13 weeks of the P. fuscatus colony cycle. Mean colony-wide rates of AD were high during pre-emergence and early post-emergence stages, but dropped dramatically following the third week after the first workers emerged. This variation in the temporal pattern of AD was correlated neither with the rate at which larvae were fed liquid, the number of larvae on the nest, nor with the adult-to-larva ratio, but was solely a function of colony stage. In contrast, rates of feeding liquid to larvae varied only as a function of the number of larvae in the nest. Queens drummed and fed liquid to larvae at much higher rates than did workers. Queen AD and feed-liquid rates decreased after the third week of worker emergence. During the same period, total feed-liquid rates of workers became as high as levels of queens during pre-emergence. Colony-wide AD rates dropped dramatically because workers seldom drummed while feeding liquid to larvae. The mean duration of AD bursts for queens also decreased after the second week of worker emergence. These results fail to support the salivary inhibition hypothesis, but provide indirect support for the mechanical switch hypothesis on AD function.     

Selection for individual recognition and the evolution of polymorphic identity signals in Polistes paper wasps

Individual recognition (IR) requires individuals to uniquely identify their social partners based on phenotypic variation. Because IR is so specific, distinctive phenotypes that stand out from the crowd facilitate efficient recognition. Over time, the benefits of unique appearances are predicted to produce a correlation between IR and phenotypic variation. Here, we test whether there is an association between elevated phenotypic polymorphism and IR in paper wasps. Previous work has shown that Polistes fuscatus use variable colour patterns for IR. We test whether two less variable wasp species, Polistes dominulus and Polistes metricus, are capable of IR. As predicted, neither species is capable of IR, suggesting that highly variable colour patterns are confined to Polistes species with IR. This association suggests that elevated phenotypic variation in taxa with IR may be the result of selection for identity signals rather than neutral processes. Given that IR is widespread among social taxa, selection for identity signalling may be an underappreciated mechanism for the origin and maintenance of polymorphism.     

Cross‐amplification of polymorphic microsatellites reveals extra‐pair paternity and brood parasitism in Sturnus vulgaris

We tested the cross‐amplification of 37 microsatellites in a population of starlings (Sturnus vulgaris). Twenty‐three of them amplified and five exhibited a large number of alleles per locus and high heterozygosity (on average: 14.6 alleles/locus and H_E = 0.704). We assessed the occurrence of extra‐pair paternity (EPP) and intraspecific brood parasitism (IBP) in this population. The EPP rate was 16% to 18% offspring from 43% to 45% of nests. IBP was very variable between two successive years (14% to 27% chicks from 25% to 64% of clutches). These five polymorphic markers will be of potential use in studies of genetic diversity, population structure and reproductive strategy of this species.     

Direct and indirect effects of predatory wasps (Polistes sp.: Vespidae) on gregarious caterpillars (Hemileuca lucina: Saturniidae)

Summary We examined how predation by vespid wasps,Polistes dominulus andP. fuscatus, affected the behavior, growth rate and survivorship of aggregated caterpillars ofHemileuca lucina (Saturniidae). Although these larvae can exhibit a variety of defense and escape behaviors, in general larvae reacted to wasp attacks by clinging to the hostplant. Neighboring larvae in the aggregation responded by leaving the feeding site and moving to the interior or base of the plant. To determine wheter wasp attack affected the behavior and growth of the caterpillars that escaped, a field experiment was conducted with treatments of: 1) larvae exposed to wasps, 2) larvae protected from wasps, and 3) larvae protected from wasps but with the attack of wasps simulated (=harassment). Over just one instar, protected larvae gained significantly more weight than the harassed larvae, which in turn weighed significantly more than the larvae that escaped the wasps. The behavior of attacked and harassed larvae differed from that of the protected larvae; the disturbed larvae often fed in smaller groups and in shaded portions of the plant where only mature leaves were available. A laboratory experiment showed that at 35° C (daytime temperature) larvae had significantly higher relative growth rates and significantly shorter instar duration than larvae reared at 25° C. Our results suggest that wasps, in addition to killing caterpillars, indirectly affect larval fitness by slowing larval growth, at least in part by forcing larvae into cooler microhabitats where leaves are of lower quality.     

Proximate Determinants of Reproductive Skew in Polygyne Colonies of the Ant Formica fusca

Understanding the determinants of reproductive skew (the partitioning of reproduction among co‐breeding individuals) is one of the major questions in social evolution. In ants, multiple‐queen nests are common and reproductive skew among queens has been shown to vary tremendously both within and between species. Proximate determinants of skew may be related to both queen and worker behaviour. Queens may attempt to change their reproductive share through dominance interactions, egg eating and by changing individual fecundity. Conversely, workers are in a position to regulate the reproductive output of queens when rearing the brood. This paper investigates queen behaviour at the onset of egg laying and the effect of queen fecundity and worker behaviour on brood development and reproductive shares of multiple queens in the ant Formica fusca. The study was conducted in two‐queen laboratory colonies where the queens produced only worker offspring. The results show that in this species reproductive apportionment among queens is not based on dominance behaviour and aggression, but rather on differences in queen fecundity. We also show that, although the queen fecundity at the onset of brood rearing is a good indicator of her final reproductive output, changes in brood composition occur during brood development. Our results highlight the importance of queen fecundity as a major determinant of her reproductive success. They furthermore suggest that in highly derived polygyne species, such as the Formica ants, direct interactions as a means for gaining reproductive dominance have lost their importance.     

Development rate and brood production in haplo- and pleometrotic colonies of <Emphasis Type="Italic">Oecophylla smaragdina</Emphasis>

Pleometrosis (colony founding by multiple queens) may improve life history characteristics that are important for early colony survival. When queens unite their initial brood, the number of workers present when incipient colonies open may be higher than for single queen colonies. Further, the time until the first worker emerges may shorten. For territorial species and species that rob brood from neighbouring colonies, a faster production of more workers may improve the chance of surviving intraspecific competition. In this study, the time from the nuptial flight to the emergence of the first worker in incipient Oecophylla smaragdina Fabr. colonies founded by 1–5 queens was compared and the production of brood during the first 68 days after the nuptial flight was assessed. Compared to haplometrotic colonies, pleometrotic colonies produced 3.2 times more workers, their first worker emerged on average 4.3 days (8%) earlier and the queen’s per capita egg production almost doubled. Further, colony production was positively, correlated with the number of founding queens and time to worker emergence was negatively correlated. These results indicate that pleometrotic O. smaragdina colo-nies are competitively superior to haplometrotic colonies as they produce more workers faster and shorten the claustral phase, leading to increased queen fecundity.