Nourishment affects colony demographics in the paper wasp Polistes metricus

Abstract  1. Colony survivorship and numbers of nest cells, pupae, and adult females were monitored throughout the nesting season for a cohort of 78 colonies of the paper wasp Polistes metricus Say. Thirty-nine colonies received a twice-weekly nourishment supplement of honey during pre-emergence and early emergence phases of the colony cycle; 39 colonies were unsupplemented controls.
2. Colony survivorship was unaffected by the supplemental nourishment. Loss of colonies to predation differed among three sites but was unaffected by supplementation.
3. Honey-supplemented colonies constructed more nest cells than did control colonies but this effect was not expressed until after supplementation had ceased.
4. Honey-supplemented colonies produced more pupae than did control colonies but the number of adult females at nests did not differ between supplemented and control colonies. Because honey-supplemented colonies had more offspring but fewer of them remained as workers at the nest, honey supplementation led to a lower frequency of workers and corresponding higher frequency of reproductives than in control colonies.
5. In a second year of study, colony survivorship and numbers of nest cells, pupae, and adult females were monitored from late pre-emergence until the end of the nesting season for a cohort of 32 colonies of Polistes metricus . In 16 colonies, trophallactic saliva was taken from final-instar larvae on nine dates in the late pre-emergence and early emergence periods; 16 colonies served as controls.
6. Saliva-diminished colonies had lower survivorship, fewer nest cells, fewer pupae, and fewer adult females at the nest than did control colonies.
7. These results show that variation in nourishment in the early to mid phases of the colony cycle can have significant effects on the subsequent colony demographics of Polistes metricus paper wasps.     

Behaviour in usurpers and late joiners ofPolistes biglumis bimaculatus (Hymenoptera,Vespidae)

Summary During the late pre-emergence phase, a foundress of the paper waspPolistes biglumis bimaculatus may be expelled by a conspecific female from her own nest (usurpation) or, less frequently, joined by another female of the same species (late association). The behaviour of femalePolistes biglumis bimaculatus, when usurping a conspecific colony or joining another foundress, is compared with that of foundresses on non-usurped colonies. The most conspicous difference is the intense abdomen stroking behaviour the usurper performs over the comb surface on the first days after usurpation. As observed in otherPolistes species, once usurpers and joiners arrive on a strange nest they will destroy most of the immature brood of the previous nest owner. Although host workers are not aggressive towards the intruder females, reproductive success of usurpers and joiners is low compared with that of legitimate foundresses. The same behaviours observed on usurped colonies are found in the obligate social parasites ofPolistes. These behaviours are therefore discussed in the context of the evolution of intra- and inter-specific parasitism.     

Three viruses found in the braconid parasitoid Microplitis croceipes and their implications in biological control programs

Productivity and longevity decreased in a laboratory colony of the parasitoid wasp Microplitis croceipes (Cresson) (Hymenoptera: Braconidae). Using light microscopy, it was determined that the colony was free of microsporidia. However, samples of the colony examined for pathogens by electron microscopy revealed three types of viruses: a nonpathogenic polydnavirus which is produced by all female wasps; a nonoccluded baculovirus which is pathogenic to late-stage pupae and adults; and a picorna-like virus which is present in larvae, pupae, and adults. The nonoccluded baculovirus was eliminated from the laboratory colony of M. croceipes by selection of progeny from wasps which had oviposited within 2 to 3 days after emergence from the cocoons and which had lived for at least 14 days post-emergence. Upon death, the wasps were examined by negative stain electron microscopy and only progeny from baculovirus-free wasps were retained. Parasitoid colonies should be systematically examined for pathogenic viruses that may reduce their productivity and efficacy as biological control agents. In addition, exotic parasitoids and predators should be evaluated for viruses and other pathogens while in quarantine.     

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.     

Host nest usurpation and colony foundation in the European amazon ant,Polyergus rufescens Latr. (Hymenoptera: Formicidae)

Summary The socially parasitic mode of founding new colonies by queens of the European amazon antPolyergus rufescens was analysed in the laboratory. Newly-mated females of this obligatory slave-maker were individually introduced into queenright and queenless artificially established colonies of bothFormica cunicularia (the slave present in the natal dulotic nest) andF. rufibarbis (another potentialServiformica host). Particular attention was devoted to the behavioural patterns displayed by these young queens during the usurpation phases. Our observations, supported also by video-taping, show that the slave-making female, before laying her eggs, must penetrate the host colony, kill the resident queen, become accepted by the adult workers and appropriate the host brood. The parasite was almost always adopted in the colonies ofF. cunicularia, whereas in the presence ofF. rufibarbis it was generally killed in a short time. The failure in the attempt of usurping the colonies ofF. rufibarbis is discussed in relation to the host specificity typical of this slave-maker. Finally, egg-laying byPolyergus successful usurpers, the subsequent eclosion of the brood, and its complete social integration in the newly-established mixed colonies were also recorded.     

Laboratory studies on the biology ofSpalangia nigra [Hym.: Pteromalidae]

At 21 °C,Spalangia nigra Latreille (Hymenoptera: Pteromalidae) averaged 29.3 days between exposure and emergence of 1^st progeny from host house flies,Musca domestica L. (Diptera: Muscidae). At 27 °C, the average developmental time to 1^st emergence was reduced to 26.6 days, and a majority of adult wasps emerged from host house fly puparia between 29 and 40 days postoviposition. The sex ratio of progeny ranged from 1.4 to 1.8 female-to-male, but all progeny of virgin females were male. Male wasps lived from 6.8–15 and females 11–17.8 days at 27 °C; honey as a food source increased longevity. No significant differences in parasitism byS. nigra were associated with host house fly pupal densities ranging from 1 to 200 pupae per female-male pair of wasps, but average percent parasitism decreased at host densities greater than 50. House fly pupae exposed to parasitism at ages ranging from 4 to 96 h did not differ in subsequent production of adult flies.S. nigra did not demonstrate preference for house flies or stable flies,Stomoxys calcitrans (L.) (Diptera: Muscidae) as hosts. The results of these studies indicate thatS. nigra may contribute significantly to previously unexplained mortality of house flies and stable flies.      

Nest-founding behavior induced in the first descendants ofPolistes dominulus Christ (Hymenoptera: Vespidae) colonies

Summary First descendants' founding ability was tested inP. dominulus colonies by experimentally removing the original nests and queens 21–24 days after the first emergence. Foundation tests were carried out after three different periods had elapsed after the removal and the results were analyzed with regards to the effects of participation in social tasks, duration of stay within the colony and seasonal factors. 1.Immediately after the removal, the foundations that occurred were performed almost entirely by workers, regardless of the duration of their stay within the colony. The number of foundations depended on the season in which the females had emerged. 2.Two months and four months after the removal, foundations were made by only a small number of females (workers and non-workers). These females, which were therefore able to found a nest the year they were born, were mostly among the very first-born individuals in the colonies. 3.The following year after hibernation: females (workers and non-workers) which had stayed for only a short time within the colony (1–15 days) had the highest survival rates. The foundation rates among the surviving females depended only on seasonal factors, since the foundation rates were higher among the non-workers which had emerged later in the season. The results throw some light on the first descendants' nest-founding potential in colonies developing under natural conditions, where many workers in fact stay for only a short time at the nest.     

Life history studies of <Emphasis Type="Italic">Prorops nasuta</Emphasis>, a parasitoid of the coffee berry borer

Life history studies were conducted in the laboratory on the African parasitoid Prorops nasuta Waterston (Hymenoptera: Bethylidae), a parasitoid of the coffee berry borer, Hypothenemus hampei (Ferrari) (Coleoptera: Scolytidae). The female wasp enters an infested coffee berry, kills the adult borer and seals the entrance of the berry with the body of the borer, impeding the entry of other organisms into the berry. The preoviposition period ranges from 3 to 14 days (mean 5.42 ± 0.37 SE). During this time females feed on the immature stages and paralyse fully grown larvae and pupae of the CBB. P. nasuta is an idiobiont solitary ectoparasitoid. Eggs are laid externally on the last instar larvae and pupae. Mean development time (egg to adult) for males and females was 27.7 (±0.37 SE) and 30 (±0.12 SE) days, respectively. Median survival for wasps fed on final instar CBB larvae was 27.7 days, significantly longer than any other treatment, while for females without food it was 2.5 days. In culture, females produced an average of 4.3 (±0.39 SE) progeny during their lifetimes. Adults began emerging at 30.6 days (±0.28 SE) after cultures were started and peak production was reached at 36 days, declining thereafter. Males normally emerged from coffee beans 2–3 days before females. Males usually emerged from 07:00 to 09:00h and females from 10:00 to 14:00 h. The culture sex ratio (proportion of males) was 0.21. Virgin females produced only male offspring.     

An experimental analysis of pleometrotic advantage in the desert seed-harvester antMessor pergandei (Hymenoptera; Formicidae)

Summary Starting colonies of the desert seed-harvester antMessor pergandei are clumped in the field and face severe intraspecific competition through brood raiding. Single foundress laboratory colonies ofM. pergandei are more likely to succeed at brood raiding with conspecific colonies if they are given additional workers and mature pupae several days prior to brood raiding. Per foundress fecundity remains constant across laboratory starting colonies established with 1, 3 and 5 foundresses. These results suggest that the selective advantage of cooperative colony foundation (pleometrosis) in this and similar species may derive directly from the ability of multiple foundresses to produce a larger brood raiding force.     

Social relations among females in pre- and postemergence colonies of a subtropical paper wasp,Parapolybia varia (Hymenoptera: Vespidae)

In southern Taiwan, most colonies (13 out of 14) ofParapolybia varia were founded by multiple females. Several females were present at the founding and increased up to 25 about one week after it. Observations on 2 pre-emergence colonies (in a total of 21 h) and those on 4 postemergence colonies (in 62 h) revealed the following facts: In the pre-emergence period, the top-ranking female dominated all others like a despot. Some females attending from the founding also dominated other females who joined later. Interactions among females gradually decreased in frequency after the founding and became milder. The top-dominant rarely left the nest and monopolized most of ovipositions by physically disturbing the subordinates' attempts to oviposit. Differential oophagy was not observed even when the subordinates succeeded in laying (6 cases). After the emergence of workers, the subordinate associates gradually disappeared and the colonies became monogynous (one egg-layer or queen per colony). Monogyny was also confirmed in October on 3 big colonies which had already produced reproductive forms. Eighty-six percent of females (n=50) taken from nests of early stage containing nothing or eggs alone were inseminated and 66% of them had well developed ovaries, while 90% of those (n=20) from more developed nests containing mature larvae and/or pupae were inseminated and only 25% had developed ovaries. This suggests that the ovaries of the subordinates had possibly, though not completely, degenerated through dominance interactions. Social biology ofParapolybia wasps in Taiwan. II. Based on part of a thesis presented to Hokkaido University in partial fulfillment of the Doctoral degree.     

Oviposition avoidance of parasitized aphid colonies by the syrphid predator Episyrphus balteatus mediated by different cues

Oviposition decisions made by members of a guild of natural enemies can have evolved to avoid intraguild predation, potentially avoiding the disruption of the extraguild prey control. We have studied the oviposition preference of the aphidophagous predator Episyrphus balteatus De Geer (Diptera: Syrphidae) within colonies of Myzus persicae Sulzer (Hemiptera: Aphididae) in the presence of two developmental stages of the aphid parasitoid Aphidius colemani Viereck (Hymenoptera: Aphidiidae). Results from a greenhouse choice experiment showed that E. balteatus females lay significantly fewer eggs in colonies with mummified aphids than in unparasitized colonies. Colonies of parasitized, but not yet mummified did not contain significantly fewer eggs than colonies with unparasitized aphids. In three no-choice experiments, we assessed stimuli coming from aphid honeydew, from the aphids themselves and also from extracts of the aphid bodies, and all of these stimuli mediate the discrimination of mummified aphids from healthy aphids. To a lesser extent these stimuli also contribute to the discrimination against aphids that are parasitized but not yet mummified. These results suggest that the effects of these two species could be complementary for the control of M. persicae, since the species that acts as an intraguild predator, E. balteatus, avoids ovipositing on aphid colonies parasitized by the intraguild prey, A. colemani.     

Influence of maternal age and host deprivation on egg production and parasitization byMicroplitis croceipes [Hym.: Braconidae]

The effect of maternal age and hostHeliothis virescens (F.) deprivation on egg production and allocation byMicroplitis croceipes (Cresson) was determined in the laboratory on 0.5, 1.0, 3.0, 6.0, 9.0, and 12.0 days old parasitoids. In host-deprived parasitoids, newly-emerged (0.5-day old) females produced significantly fewer eggs than older ones; egg production peaked 3 to 6 days after emergence and declined thereafter. However, the number of eggs found in undeprived parasitoids at various ages suggested a cyclical pattern of egg production. The rate of parasitization by host-deprivedM. croceipes was higher in 1.0 to 6.0 days old than in newly-emerged or older females and was correlated strongly with egg production (r=0.93). The first observation on premature larval hatching inM. croceipes was documented.      

Use of learned visual cues during habitat location by Brachymeria intermedia

Like many other parasitoids, Brachymeria intermedia (Hymenoptera: Chalcididae) uses olfactory cues to find its host, but in addition it appears to use learned visual cues to focus its searching efforts at the macrohabitat level. In laboratory experiments, females were held in cages where Lymantria dispar pupae were hidden either in a vertical, tree-like structure or on the floor. After four days females had learned to search for hosts in the structure in which they previously had found pupae. Such training was reversible. During similar tests in a semi-natural situation, in which pupae were hung from a tree trunk or were hidden under leaf litter, females also tailored their subsequent searching to favour the macrohabitat where hosts previously had been found. This parasitoid most likely uses a combination of visual and olfactory cues during host searching.     

Early queen infection shapes developmental dynamics and induces long-term disease protection in incipient ant colonies

Infections early in life can have enduring effects on an organism's development and immunity. In this study, we show that this equally applies to developing ‘superorganisms’––incipient social insect colonies. When we exposed newly mated Lasius niger ant queens to a low pathogen dose, their colonies grew more slowly than controls before winter, but reached similar sizes afterwards. Independent of exposure, queen hibernation survival improved when the ratio of pupae to workers was small. Queens that reared fewer pupae before worker emergence exhibited lower pathogen levels, indicating that high brood rearing efforts interfere with the ability of the queen's immune system to suppress pathogen proliferation. Early-life queen pathogen exposure also improved the immunocompetence of her worker offspring, as demonstrated by challenging the workers to the same pathogen a year later. Transgenerational transfer of the queen's pathogen experience to her workforce can hence durably reduce the disease susceptibility of the whole superorganism.     

Pupation site and emergence time influence the mating success of bagworm females,Oiketicus kirbyi

In commercial oil palm plantations in Costa Rica, we tested the hypotheses that pupation site and emergence time affect the mating success of protogynous female bagworms,Oiketicus kirbyi (Guilding) (Lepidoptera: Psychidae). Greater proportions of female than male pupae on upper leaves of oil palms and greater proportions of mated females in the upper rather than lower crown strata support the hypothesis that selection of pupation site by female larvae influences the mating success of adults. Increasing captures of males with increasing trap height further suggest that enhanced mating success of females in tree tops may be attributed either to most effective dissemination of sex pheromone on higher sites, or to males foraging predominantly in the upper strata of oil palms. As the majority of females pupated in the middle rather than upper crown of oil palms, selection of pupation site by females may be affected by additional as yet unknown factors. Emergence of females significntly preceded emergence of males. Increasing proportions of mated females throughout the emergence seasons probably resulted from an increased ‘availability’ of males. In tropical rainforests with local variations inO. kirbyi developmental time and stage, protogyny may represent an evolutionary strategy that furthers outbreeding.     

Production and storage of mealworm beetle as prey for predatory stinkbug

Arthropod predators can be produced using alternative prey, but the availability of a constant supply of prey at low cost is necessary to support large-scale production. The use of stored prey can be advantageous for maintaining a constant food supply, but its quality may decline in storage. Thus, we tested the effect of using stored pupae of the alternative prey Tenebrio molitor L. at low temperature (mean of 3.6°C) on the performance of predatory stinkbug Podisus nigrispinus (Dallas). The treatment conditions we tested were: pupae stored for exactly 30 days before being used as prey (30-d pupae); pupae stored for either 20 or 35 days prior to the initiation of their use as prey, with continued storage during nymphal and adult stages (20-d+ or 35-d+ pupae); and 2-day-old newly moulted pupae as the control treatment. The duration of the nymphal stage of females and the weight of males were lower for nymphs fed 20-d+ and 35-d+ pupae, respectively. The pupal weight consumed and the rate of eggs and nymphs produced per gram of consumed pupae were also lower for adult females fed 35-d+ pupae compared to females that were fed 2-day-old pupae. These measures, however, were similar for females fed 30-d and 20-d+ pupae. The greatest consumption of pupae was observed for 20-d+ and 35-d+ treatments during nymphal and adult stages, respectively. The use of stored T. molitor pupae supported the development and reproduction of P. nigrispinus, regardless of the period of storage but with variable results among treatments. Therefore, among any of the tested treatments those pupae that were stored for exactly 30 days produced the highest fitness for P. nigrispinus.     

Acceptance and suitability of four lepidopteran stemborers for the development of the pupal parasitoid Xanthopimpla stemmator (Hymenoptera: Ichneumonidae)

Xanthopimpla stemmator (Thunberg), a solitary endoparasitoid of lepidopteran stemborer pupae, was recently imported into East Africa as a candidate biological control agent of gramineous stemborers. Suitability of Busseola fusca Fuller, Chilo partellus (Swinhoe), Eldana saccharina (Walker) and Sesamia calamistis Hampson, for the development of X. stemmator was studied in the laboratory. One- to 6-day-old laboratory reared pupae of the four stemborer species were exposed to naïve X. stemmator females. All host pupae and ages were acceptable for oviposition. The parasitoids inflicted multiple probe wounds on 67.8% of pupae exposed. B. fusca, C. partellus and S. calamistis were equally suitable with 56.4, 59.4 and 52.3%, respectively, of probed pupae leading to emergence of adult parasitoids. E. saccharina was less suitable with only 22.6% of probed pupae producing parasitoids. Emergence of parasitoids did not differ significantly across the six pupal ages for B. fusca and S. calamistis, but varied for C. partellus and E. saccharina. No parasitoids emerged from 6-day-old E. saccharina pupae. Realized fecundity of females reared on the four stemborer pupae showed that fewer progeny were produced by females emerging from E. saccharina than females reared on the other three stemborer species. Eldana saccharina may be a poor host for X. stemmator in Kenya, but this parasitoid is a potential candidate for biological control of B. fusca, C. partellus and S. calamistis.     

Development ofBrachymeria ovata [hymenoptera: chalcididae] in three noctuid hosts reared on artificial diet and insect-resistant and susceptible soybean genotypes

The development of the pupal parasitoidBrachymeria ovata (Say) was studied in 3 lepidopterous hosts reared on artificial diet and insect-susceptible and insectresistant soybena genotypes.Pseudoplusia includens (Walker),Anticarsia gemmatalis Hubner andHeliothis zea (Boddie) pupae from larvae reared on soybean leaves were less suitable forB. ovata development than pupae from larvae reared on artificial diet.B. ovata emergence rates, size and sometimes development period were adversely affected on plant-reared hosts. Dissection of hosts 6 days after parasitisation byB. ovata showed a higher proportion of diet-reared host pupae contained large parasitoid larvae than plant-reared hosts. Suitability differences were also detected among hosts reared on different soybean genotypes but these differences did not parallel closely the response of the lepidopterous hosts to soybean genotypes.      

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.     

Effects of host stage on the development of the facultative autoparasitoid Encarsia tricolor (Hymenoptera: Aphelinidae)

Encarsia tricolor is a facultative autoparasitoid of the glasshouse whitefly, Trialeurodes vaporariorum, with a potential in biological control. The rate of development, number of mature oocytes at emergence, number of ovarioles and size of the emerged adults were studied. Five nymphal instars (N1, N2, N3, N4, and pharate adult) of T. vaporariorum were used as hosts for the females. Female larvae and pupae of E. tricolor and Encarsia formosa were used as hosts for the males. Females developed faster when the egg was laid on N3 (18.0 days from egg to adult) and slower on N1 (22.3 days). Females were bigger when developing from N1 and N3 than from N4 and pharate adult. On emergence the mean number of mature oocytes was always small (0.8–2.6). Males developed faster and were smaller than females, and developed faster and were larger on larvae of E. formosa.