Competition between Honey-Bees (

Honeydew, the sugary exudate of the scale insect Ultracoelostoma brittini, is an important food source in black beech (Nothofagus solandri var. solandri) forests in the South Island of New Zealand. Two of the most prominent foragers of honeydew are honey bees (Apis mellifera) and wasps (Vespula germanica and V. vulgaris). Observations in the field and using a captive bee hive were used to investigate competition between bees and wasps feeding on honeydew. In laboratory trials, interference competition was often strong, and many cases of aggression were noted. In the forest, there was invariably enough room on the trees for bees and wasps to feed while rarely encountering one another. Over the whole year, environmental variables (especially low temperatures and rain), were found to constrain honey bee foraging to a greater degree than competition with wasps. Because the competition that did occur was primarily exploitation competition, reciprocal effects were likely to be felt. At Coopers Creek, bees may be reducing wasp densities, compared with the situation in Nelson—Marlborough where commercial hives are scarce. It may be possible to reduce wasp densities locally by increasing the number of bee hives in an area.     

Prey Diets and Population-Densities of the Wasps

Prey collected by Vespula vulgaris and V. germanica were sampled by intercepting foragers returning to nests at two sites in scrubland-pasture near Hamilton. About 12% of returning foragers carried animal prey and 5% carried wood pulp. The remaining 83% carried no external load. The most common prey item for both species was Diptera, followed by Lepidoptera and Araneae (spiders). Even in similar habitats the two species collected different prey, with V. germanica collecting more Diptera and V. vulgaris more Lepidoptera. The prey changed slightly over time at both sites, V. germanica and V. vulgaris collected an estimated 1800 g and 600 g of prey nest-1 season-1 respectively. The wasps collected an estimated 75 000 prey loads ha-1 season-1 at Mystery Creek and 12 000 at Ruakura. This represents biomasses of 50 and 470 g ha-1, respectively, which are at least an order of magnitude lower than estimates of prey biomass for honeydew beech forest in parts of the South Island. The distribution of wasp nests was patchy however, and the biomass of prey collected exceeded 10 kg ha-1 in some places.     

Volatiles from green‐lipped mussel as a lead to vespid wasp attractants

Vespid wasps (Vespula vulgaris L. and V. germanica Fab. Hymenoptera; Vespidae) are highly abundant in 1 million ha of New Zealand's indigenous beech forests (Nothofagus spp.) and have had detrimental effects on the New Zealand native fauna. This hyperabundance is due in part to the vast supply of carbohydrate‐rich honeydew produced by scale insects Ultracoelostoma spp. native to New Zealand. Current control methods include the use of wet cat food as a protein source with insecticide as a lure‐and‐kill‐based system, but there are problems with fresh baits degrading rapidly, and a more durable formulation would enable the expansion and longevity of wasp control. Four crude protein baits were tested for vespid attraction. Green‐lipped mussels had the highest vespid catch of the crude baits tested, and aged and fresh mussels were equally attractive. From headspace analysis of the green‐lipped mussel volatiles, a series of butanoate esters, 3‐octanone and 1‐octen‐3‐ol were identified as possible attractants. These compounds were tested individually and in various blend combinations for the attraction of Vespula wasps in matagouri vegetation at the edge of beech forests. We found synergistic effects between single attractive compounds when tested in various combinations, and the multicomponent lures were more attractive to these wasps than heptyl and octyl butanoate, previously identified attractants for vespid species. The new multicomponent lures could form the basis for a new generation of attractants for social wasps that can provide sustained control methods for invasive vespid wasps.     

Preference by Vespula germanica (Hymenoptera: Vespidae) for processed meats: implications for toxic baiting

The German yellowjacket, Vespula germanica (F.) (Hymenoptera: Vespidae), was introduced into Australia in 1959 and has established throughout southern Australia. In urban environments, V. germanica is frequently a nuisance pest at public gatherings and to homeowners. In native environments, it has the potential to pose a threat to native invertebrates. The current practice for controlling the wasps is nest destruction with pesticide. However, locating the nest(s) is not always practical or possible. Meat baits impregnated with an insecticide that foraging wasps cut and carry back to the nest offer a means of suppressing wasps where the nest sites are unknown. The success of meat baits depends on the attractiveness and acceptance of the meat to the wasp and the mode of action of the insecticide. Our objective was to determine wasp preference and acceptance of five processed meats: canned chicken or fish and freeze-dried chicken, fish, or kangaroo. We found that more wasps visited and took freeze-dried kangaroo and canned chicken than the other baits. Canned and freeze-dried fish were similarly preferred, and freeze-dried chicken was the least attractive and accepted by foraging wasps. Our findings demonstrate that wasps prefer some processed meats and hence take more loads back to the nest. By combining a suitable insecticide with a meat bait preferred by wasps, the likelihood of effective suppression of nuisance wasp populations should be increased.     

Foraging niche separation of social wasps in an invaded area: Implications for their management

Foraging niche separation may be a mechanism to promote coexistence of two competing species by concentrating intraspecific competition relative to interspecific competition. The present study investigated foraging behaviour and microhabitat use of two coexisting species of invasive social wasps, Vespula germanica and Vespula vulgaris, when foraging for two different food resources. Also, we tested the attractiveness of traps baited with a synthetic lure for those two species. We found that V. germanica wasps prefer to forage at ground level regardless of the resource, while V. vulgaris prefers protein resources at the shrubland level given a choice between a protein bait at ground or at shrubland level. However, when baited with the synthetic lure, the species caught was not affected by the height at which traps were placed. That is, in a no choice scenario, the traps were sufficiently attractive to lure both species of wasps to both microhabitats (ground and shrubland levels). Thus, our results support the existence of spatial niche differentiation at least in protein foraging and suggest that the synthetic lure evaluated could be used to trap both species of Vespula wasps present in Argentina. These results could help to improve management strategies of these social wasps in an invaded area.     

Competition for honeydew between two social wasps in South Island beech forests,New Zealand

Summary Honeydew is a sugary secretion of beech scale insects (Ultracoelostoma spp.). Two introduced species of wasps forage on the sugar droplets in New Zealand beech forests. We hypothesize that competition between them may explain whyVespula germanica has become locally extinct in beech (Nothofagus) forest invaded byV. vulgaris. Changes in behaviour of the two wasp species in response to changes in the honeydew resource were monitored. Foraging and rainfall separately and together reduced the honeydew standing crop. In response to the standing crop decreasing, more wasps were found on honeydew trees, they became less active, spent more time lapping the tree surface, and ingested drops of honeydew at a slower rate.V. vulgaris was more active, and obtained drops and energy at a higher rate thanV. germanica. These behavioural differences may lead to competitive advantages affecting queen size and possibly survival.     

Attraction of Vespula germanica (Hymenoptera: Vespidae) foragers by conspecific heads

The socialwasp Vespula germanica (F.) is a serious pest in many regions it has invaded. Control programs to reduce its populations are commonly based on the use of poison baits. These baits also attract nonpestiferous invertebrates and vertebrates. In this work we studied the attraction of V. germanica foragers by conspecific worker squashes, comparing the effect of head and abdomen squashes in wasps behavior. We found that head squashes attract V. germanica foragers, elicit landing and transportation to nests. Furthermore, the addition of squashed heads to a protein bait increased attraction. This could be an alternative to improve baiting programs.     

Orchids mimic green-leaf volatiles to attract prey-hunting wasps for pollination

An outstanding feature of orchids is the diversity of their pollination systems [1]. Most remarkable are those species that employ chemical deceit for the attraction of pollinators [2]. The orchid Epipactis helleborine is a typical wasp flower, exhibiting physiological and morphological adaptations for the attraction of pollinating social wasps [3]. As noted by Darwin [1], this species is almost entirely overlooked by other potential pollinators, despite a large nectar reward. Therefore, the mechanism for the attraction of pollinating social wasps was something of a mystery. By using a combination of behavioral experiments, electrophysiological investigations, and chemical analyses, we demonstrate for the first time that the flowers of E. helleborine and E. purpurata emit green-leaf volatiles (GLVs), which are attractive to foragers of the social wasps Vespula germanica and V. vulgaris. GLVs, emitted by damaged plant tissues, are known to guide parasitic wasps to their hosts [4]. Several E. helleborine GLVs that induced response in the antennae of wasps were also emitted by cabbage leaves infested with caterpillars (Pieris brassicae), which are common prey items for wasps [5]. This is the first example in which GLVs have been implicated in chemical mimicry for the attraction of pollinating insects.     

Assessment of prey overlap between a native (Polistes humilis) and an introduced (Vespula germanica) social wasp using morphology and phylogenetic analyses of 16S rDNA

Abstract In newly invaded communities, interspecific competition is thought to play an important role in determining the success of the invader and its impact on the native community. In southern Australia, the native Polistes humilis was the predominant social wasp prior to the arrival of the exotic Vespula germanica (Hymenoptera: Vespidae). Both species forage for similar resources (water, pulp, carbohydrate and protein prey), and concerns have arisen about potential competition between them. The aim of this study was to identify the protein foods that these wasps feed on. As many prey items are masticated by these wasps to the degree that they cannot be identified using conventional means, morphological identification was complemented by sequencing fragments of the mitochondrial 16S rRNA gene. GenBank searches using blast and phylogenetic analyses were used to identify prey items to at least order level. The results were used to construct complete prey inventories for the two species. These indicate that while P. humilis is restricted to feeding on lepidopteran larvae, V. germanica collects a variety of prey of invertebrate and vertebrate origin. Calculated values of prey overlap between the two species are used to discuss the implications of V. germanica impacting on P. humilis. Results obtained are compared to those gained by solely 'conventional' methods, and the advantages of using DNA-based taxonomy in ecological studies are emphasized.     

Foraging behavior related to habitat characteristics in the invasive wasp Vespula germanica

When foraging on carrion resources, the wasp Vespula germanica usually makes repeated visits to the feeding site until depleting the resource. In the present study we analyzed how environmental cues affect wasps' behavior when re‐locating a protein food source. We studied this behavior in two different natural habitats: closed and open habitats. As closed habitats have more references to orient wasps to the feeding site than open habitats, we hypothesized that they would return to the foraging site more frequently in closed habitats than in open ones. We tested this hypothesis by studying wasp behavior in three different natural habitat conditions: (i) closed habitats, (ii) open habitats, and (iii) open habitats artificially modified by adding five sticks with flagging. Experiments consisted of training individual wasps to feed from a certain array, and at the testing phase we removed food and displaced the array by 60 cm. Therefore, we recorded wasps’choices when returning to the training area, by counting both the wasps’first approaches and the number of visits to the original feeding site and the displaced array. Wasps' behavior while re‐locating a protein food source was different if foraging at open or closed habitats. Wasps more frequently revisited a previous feeding location when foraging in closed habitats than when foraging in open ones. Furthermore, wasps more frequently visited the displaced array than the original feeding site in all three treatments. Nevertheless, when wasps were trained in closed habitats, they returned to the original feeding site more frequently than if trained in open ones. Interestingly, when five sticks with flagging were added in open habitats, wasps responded similarly as in closed habitats without these references. The results show that foraging behavior in V. germanica seems to be different in closed and open habitats, probably associated with the existence of references that guide foragers when re‐locating undepleted resources.     

Hierarchical genetic structure of the introduced wasp Vespula germanica in Australia

The wasp Vespula germanica is a highly successful invasive pest. This study examined the population genetic structure of V. germanica in its introduced range in Australia. We sampled 1320 workers and 376 males from 141 nests obtained from three widely separated geographical areas on the Australian mainland and one on the island of Tasmania. The genotypes of all wasps were assayed at three polymorphic DNA microsatellite markers. Our analyses uncovered significant allelic differentiation among all four V. germanica populations. Pairwise estimates of genetic divergence between populations agreed with the results of a model-based clustering algorithm which indicated that the Tasmanian population was particularly distinct from the other populations. Within-population analyses revealed that genetic similarity declined with spatial distance, indicating that wasps from nests separated by more than approximately 25 km belonged to separate mating pools. We suggest that the observed genetic patterns resulted from frequent bottlenecks experienced by the V. germanica populations during their colonization of Australia.     

Seasonal variation in the honeydew, invertebrate, fruit and nectar resource for bellbirds in a New Zealand mountain beech forest

To examine the seasonal availability of the major bellbird (Anthornis melanura) food sources in a mountain beech (Nothofagus solandrivar. cliffortioides) forest at Craigieburn, the invertebrate, honeydew, and mistletoe (Peraxilla tetrapetala and Alepis flavida) fruit and nectar resources were sampled over 12 months. The total available food varied 2.6-fold from a low in October (8798 kJ/ha) to a high in December (22,959 kJ/ha) with an annual mean of 15,782 kJ/ha. Invertebrates were available all year and represented 89% of the available food energy. Only 16% of the invertebrate resource was on beech foliage, and beech trunks with honeydew had 60% more invertebrate energy than trunks without honeydew. The energy value of honeydew at Craigieburn (0.9% of the total) was much lower than at lower altitude sites. The relative rankings of honeydew standing crops on 25 permanently marked trees were very constant. On an annual basis mistletoe nectar and fruit made up 6.3% and 4.9%, respectively, of total food energy, but P. tetrapetala nectar was 46% of available food in early January, and P. tetrapetala fruit was 25% of the total in March. Bellbirds spent less time foraging on invertebrates, and more time on the other foods, than energy values would predict. However, during the Peak of its short flowering season, P. tetrapetala nectar made up 46% of available energy but only 33% of bellbird foraging observations. At this site P. tetrapetala is pollen limited due to insufficient visits from pollinators. This may be because bellbirds require invertebrates for protein, or to feed to nestlings. Therefore the pollination mutualism is faltering, despite high investment in nectar by the plant.     

Influence of floral resources on sugar feeding and nutrient dynamics of a parasitoid in the field

Abstract.  1. To investigate how floral resources impact feeding and nutrient dynamics of parasitoids, the ichneumonid Diadegma insulare (Cresson) was monitored in cabbage fields with and without buckwheat Fagopyrum esculentum Moench borders. Biochemical analyses of field-collected wasps determined whether sugar feeding occurred, and whether it enhanced nutrient levels and increased parasitism of diamondback larvae Plutella xylostella (L.) from 2000 to 2003. The effects of experimental spatial scale on sugar feeding dynamics were also evaluated in 2001.
2. The majority of D. insulare foraging on cabbage had fed upon sugar, but floral borders did not consistently increase the proportion of sugar-fed wasps or their levels of glycogen and lipid over all 4 years. Floral borders increased sugar storage by females in 2001. More females fed on sugar when experimental plots were separated by at least 800 m compared with 67 m.
3. Feeding on buckwheat nectar vs. honeydew produced by the soybean aphid Aphis glycines Matsumura in surrounding fields was distinguished using the ratio of fructose to total sugars, which was higher in nectar- than in honeydew-fed parasitoids. Floral borders increased the probability that females collected from adjacent crops were nectar-fed in 2002, demonstrating that floral resources were utilised by foraging parasitoids. Diadegma insulare that fed upon any sugar source had higher levels of sugar and glycogen than unfed wasps, and feeding on honeydew appeared to enhance nutrient storage.
4. The prevalence of sugar or nectar feeding by female D. insulare was not correlated with parasitism rates over the 4 years from all plots, but nectar feeding was positively correlated with parasitism rates within floral plots.     

Sugar feeding by the aphid parasitoid Binodoxys communis: how does honeydew compare with other sugar sources?

Parasitoids commonly forage in agricultural settings where the predominant sugar source is homopteran honeydew. The aphidiine braconid, Binodoxys communis, is an Asian parasitoid currently being released against the soybean aphid, Aphis glycines, in North American soybean fields. We conducted a number of laboratory experiments evaluating the quality of A. glycines honeydew as a sugar source for this parasitoid. Wasps readily fed on droplets of A. glycines honeydew, honey and 50% sucrose solution, but the length of feeding bouts on honey was significantly longer than on the other foods. Parasitoids lived significantly longer when fed honey or sucrose than honeydew, while starved wasps had the shortest lifespan. At 21+/-1 degrees C and 25+/-5% R.H., male B. communis that were fed honey lived for a maximum of 14 days, while females lived up to 20 days. Honeydew-fed wasps of both sexes lived approximately 3 days on average, which was 2-3 times longer than when they were only allowed access to water. Anthrone tests of whole insects showed that total sugar and glycogen levels of honey or sucrose-fed individuals were consistently higher than those fed honeydew or water. The glycogen levels of honeydew-fed wasps increased significantly after one day of feeding. HPLC analyses revealed that B. communis readily assimilates A. glycines honeydew oligosaccharides such as erlose, while others (e.g., raffinose) did not degenerate. Raffinose was present in much higher amounts in honeydew-fed wasps than in wasps fed other diets, so this sugar could be used as a 'signature' sugar for this species. Honeydew-fed wasps also had significantly lower fructose/(fructose+glucose) ratios than those from other diet treatments. Although A. glycines honeydew might be the main carbohydrate source within a soybean field, other sugar sources such as floral nectar appear to be more optimal foods for B. communis from a physiological standpoint. We discuss the results from the perspective of classical biological control of the soybean aphid in North America.     

A proteomic study of the major allergens from yellow jacket venoms

The venoms of stinging insects belong to the most dangerous allergen sources and can cause fatal anaphylactic reactions. Reliable prediction of a patient's risk to anaphylactic reactions is vital, and diagnosis requires the knowledge of the relevant allergens. Recently, a new hyaluronidase -like glycoprotein from Vespula vulgaris (Ves v 2b) was identified. This led us to investigate hyaluronidases and also other major allergens from V. germanica and four additional Vespula species. By MALDI-Q-TOF-MS, the new hyaluronidase-like protein was shown to be the major component of the 43-kDa band in all Vespula species studied. LC-ESI-Q-TOF-MS/MS sequencing of Ves g 2a and Ves g 2b facilitated the cloning of their cDNA. Ves v 2b and Ves g 2b turned out to be essentially identical on protein level. Whereas the less abundant "a" form displayed enzymatic activity, the new "b" homologue did not. This is probably caused by amino acid exchanges in the active site, and it raises questions about the physiological role of this protein. Sequence comparisons by MS/MS of antigen 5 and phospholipases from V. vulgaris, germanica, maculifrons, pensylvanica, flavopilosa and squamosa revealed the latter as a taxonomic outlier and led to the discovery of several not previously reported amino acid differences.     

Comparing floral nectar and aphid honeydew diets on the longevity and nutrient levels of a parasitoid wasp

We compared the effects of floral nectar from buckwheat, Fagopyrum esculentum Moench, and honeydew produced by the soybean aphid, Aphis glycines Matsumura (Homoptera: Aphididae), on longevity, nutrient levels, and egg loads of the parasitoid Diadegma insulare Cresson (Hymenoptera: Ichneumonidae). Diadegma insulare lived for 2 days in control treatments of water or clean soybean leaves, for 6–7 days with honeydew, and in excess of 2 weeks with buckwheat nectar. Potential reasons for the superiority of buckwheat nectar over soybean aphid honeydew for extending the longevity of parasitoids include: (i) parasitoids ingest more sugars from floral sources, (ii) oligosaccharides in honeydew have a lower nutritional value than nectar sugars, and (iii) honeydew has antagonistic compounds. Overall sugar levels were lower in honeydew‐ vs. nectar‐fed female wasps, suggesting a lower feeding rate, but other explanations cannot be excluded. Diadegma insulare eclosed with high levels of lipids and glycogen, and low levels of gut and storage sugars. All carbohydrates increased over the life of both nectar‐ and honeydew‐fed wasps, but remained low or decreased in starved wasps. Lipid levels declined over the lifespan of female wasps, but females fed floral nectar showed the slowest rate of lipid decline. Diet did not affect egg load, probably because the females were not given hosts in the experiment.     

Abundance of wasps and prey consumption of paper wasps (Hymenoptera, Vespidae : Polistinae) in Northland, New Zealand

Polistine and vespine wasps were captured in Malaise traps in two fire-modified shrubland habitats of varying canopy height and composition at Lake Ohia, Northland, New Zealand. Prey consumption rates were calculated for the Asian paper wasp (Polistes chinensis antennalis) occupying these two areas of shrubland and a home garden in Whangarei, Northland. The sites were systematically searched for nests and wasp prey determined by intercepting foragers returning to nests. The Asian paper wasp predominated in the Malaise trap samples from the low- growing habitat while the German wasp (Vespula germanica) was more common in the taller vegetation type. The Asian paper wasp was more abundant than the German wasp in the samples in February and early March. Only four Australian paper wasps (Polistes humilis) and no common wasps (Vespula vulgaris) were caught. Asian paper wasps collected an estimated 15 000 prey loads per ha per season from one of the shrubland areas, and 478 000 prey loads per ha per season from the second area. These convert to estimates of 31 and 957 g per ha per season of invertebrate biomass removed by paper wasps from each habitat, respectively. The estimate for the garden site was 79 g per ha per season. Wasp nest densities varied between 20 and 210 nests per hectare. The biomass estimates are similar to average figures calculated for vespine wasps in scrubland and pasture. Both Asian paper wasps and Australian paper wasps preyed mainly on lepidopteran larvae. The cabbage white butterfly (Pieris rapae) was the most commonly collected species. Noctuid species were also well represented. Both male and female Asian paper wasps collected nectar in late March and early April.     

Contrasting foraging related behaviours in two sympatric wasps (Vespula maculifrons and V. germanica)

ABSTRACT.

• 1 Using pinned freshly killed wasps (yellowjackets) to simulate feeding animals, it was found that V. germanica was socially facilitated, and concentrated its foraging activity on those feeders with the most pinned wasps (twenty). V. maculifrons foragers responded inversely, preferentially foraging at feeders with few or no pinned wasps.
• 2 Using hand-painted drawing pins (thumb tacks) as wasp models, similar results were found, indicating that visual, rather than olfactory, cues were responsible for the observed distribution.
• 3 Vespula maculifrons and V. germanica exhibit aggression while foraging at honey-baited feeders. Both species attack conspecifics at frequencies lower than would be expected under random encounter.

     

Comparison of three liquid lures for trapping social wasps (Hymenoptera: Vespidae)

Two citrus-based sodas and a known wasp attractant were compared in a field trial to assess their attractiveness to local nuisance wasp species. The wasps captured included Vespula germanica (F.), Vespula maculifrons (Buysson), Vespula vulgaris (L.), Vespula flavopilosa Jacobson, Vespula squamosa (Drury), Dolichovespula maculata (L.), Polistes fuscatus (L.), Polistes metricus Say, and Polistes dominulus (Christ). Wasps in the genus Vespula were present in significantly higher numbers in traps than Dolichovespula and Polistes. Both citrus soda products were superior to the isobutanol-acetic acid mixture as attractants for almost all of the wasp species.     

Honeydew composition and its effect on life-history parameters of hyperparasitoids

1. Diets that maximise life span often differ from diets that maximise reproduction. Animals have therefore evolved advanced foraging strategies to acquire optimal nutrition and maximise their fitness. The free-living adult females of parasitoid wasps (Hymenoptera) need to balance their search for hosts to reproduce and for carbohydrate resources to feed. 2. Honeydew, excreted by phloem-feeding insects, presents a widely available carbohydrate source in nature that can benefit natural enemies of honeydew-producing insects. However, the effects of variation in honeydew on organisms in the fourth trophic level, such as hyperparasitoids, are not yet understood. 3. This study examined how five different honeydew types influence longevity and fecundity of four hyperparasitoid taxa. Asaphes spp. (Pteromalidae) and Dendrocerus spp. (Megaspilidae) are secondary parasitoids of aphid parasitoids and are thus associated with honeydew-producing insects. Gelis agilis and Acrolyta nens (both Ichneumonidae) are secondary parasitoids of species that do not use honeydew-producing hosts. 4. Most honeydew types had a positive or neutral effect on life span and fecundity of hyperparasitoids compared with controls without honeydew, although negative effects were also found for both aphid hyperparasitoids. Honeydew produced by aphids feeding on sweet pepper plants was most beneficial for all hyperparasitoid taxa, which can partially be explained by the high amount of honeydew, but also by the composition of dietary sugars in these honeydew types. 5. The findings of this study underline the value of aphid honeydew as a carbohydrate resource for fourth-trophic-level organisms, not only those associated with honeydew-producing insects but also ‘interlopers’ without such a natural association.