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.

     

Essential oils as vespid wasp repellents: Implications for their use as a management strategy

Recently, plant‐based repellents have been proposed as a potential alternative to classic pesticides against pest wasps, in certain scenarios. Here, the repellent effect of Dysphania multifida essential oil and one of its main terpenoid components, α‐terpinene, were tested under field conditions with natural populations of wasps in Patagonia Argentina. D. multifida essential oil (paico), as well as α‐terpinene, repelled V. germanica wasps in the field. A strong avoidance of food baits treated with the essential oil or α‐terpinene was observed in choice and no‐choice tests. In no‐choice tests, the time it took wasps to arrive at the bait was significantly greater in treated baits than in control baits. Also, the total number of arriving wasps in 30 min was significantly greater in untreated baits in comparison with treated baits, under similar environmental conditions and wasp density. As the wasps’ flight season progressed, wasp density and motivation for proteinaceous food sources increased. This was evidenced by a greater total number of wasps in untreated baits with time. On the contrary, the number of wasps in treated baits remained low throughout the peak season. Both the paico essential oil and the α‐terpinene act as powerful repellents for V. germanica wasps, generating an avoidance response to treated food sources. Thus, these compounds have potential to be used as repellents to prevent wasps’ approaches and foraging, when applied in close proximity to a food source.     

Resource choice of social wasps: influence of presence, size and species of resident wasps

Summary: The role of visual cues provided by resident wasps on resource choice by yellowjacket and paper wasp foragers was investigated. Large spring queen yellowjackets and small early season yellowjacket foragers (Vespula germanica, Vespula maculifrons, and Vespula vidua) were extracted in hexane to remove odors and posed as though feeding at petri dish feeders bearing daisy-like flower models, equipped with microcapillary feeding tubes, and containing 1:3 honey:water solution. An array of five feeders was presented to foragers at a suburban and a woodland site in Saratoga Springs, New York. The visual cues provided by resident wasps influenced resource choice by approaching social wasp foragers. Vespula germanica, an introduced yellowjacket species that tends to dominate at rich resources, was the only wasp visiting the suburban feeders. Foragers of this species preferentially fed on feeders and flowers with posed wasps and fed most often next to large wasps. Polistes fuscatus foragers at the woodland site similarly preferred to feed on occupied feeders and flowers. Vespula maculifrons and V. consobrina preferentially visited unoccupied feeders. Individual V. maculifrons, V. consobrina and V. vidua foragers that landed on occupied feeders all preferentially visited unoccupied flowers on those feeders. Vespula vidua and V. flavopilosa foragers did not demonstrate a feeder preference based on the presence/absence of posed wasps. Vespula consobrina foragers that visited occupied feeders preferred those occupied by extracted V. maculifrons queens and workers; no other wasps showed species based landing preferences.     

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.     

Social Learning in Vespula Germanica Wasps: Do They Use Collective Foraging Strategies?

Vespula germanica is a social wasp that has become established outside its native range in many regions of the world, becoming a major pest in the invaded areas. In the present work we analyze social communication processes used by V. germanica when exploiting un-depleted food sources. For this purpose, we investigated the arrival pattern of wasps at a protein bait and evaluated whether a forager recruited conspecifics in three different situations: foragers were able to return to the nest (full communication), foragers were removed on arrival (communication impeded), or only one forager was allowed to return to the nest (local enhancement restricted). Results demonstrated the existence of recruitment in V. germanica, given that very different patterns of wasp arrivals and a higher frequency of wasp visits to the resource were observed when communication flow between experienced and naive foragers was allowed. Our findings showed that recruitment takes place at a distance from the food source, in addition to local enhancement. When both local enhancement and distant recruitment were occurring simultaneously, the pattern of wasp arrival was exponential. When recruitment occurred only distant from the feeder, the arrival pattern was linear, but the number of wasps arriving was twice as many as when neither communication nor local enhancement was allowed. Moreover, when return to the nest was impeded, wasp arrival at the bait was regular and constant, indicating that naive wasps forage individually and are not spatially aggregated. In conclusion, this is the first study to demonstrate recruitment in V. germanica at a distance from the food source by modelling wasps’ arrival to a protein-based resource. In addition, the existence of correlations when communication was allowed and reflected in tandem arrivals indicates that we were not in the presence of random processes.     

Evidence of associative blocking during foraging in the social wasp Vespula germanica

While foraging, Vespula germanica usually return to abundant food sites. During this relocation behavior, these wasps learn to identify contextual cues associated with food position. We analyzed associative blocking in this species, that is, how an association with a conditioned stimulus (CS1) blocks subsequent learning when a novel stimulus (CS2) is added on a second foraging visit. Three groups of wasps (A, B, and C; total 74 individual wasps) were observed while collecting meat during one or two consecutive visits. In group A, an environmental cue (CS1) was paired with food placed at a specific site, and on the second visit, a second cue (CS2) was added while food remained in the same position. In a subsequent testing phase, CS1 was removed and the food source displaced nearby. We then recorded the number of hovers performed over the empty dish (previously baited). Group A wasps appeared to ignore the addition of CS2 on their second visit because they performed fewer hovers over the learned site. For group A, the duration of the decision-making process to finally fly toward the baited dish was shorter than when CS1 and CS2 were presented together on their first visit (group B). This is the first study to demonstrate the occurrence of associative blocking in vespids, confirming that a prior foraging experience influences subsequent food relocation in V. germanica. Our findings reveal that first learning episodes block further associations with novel contextual cues, contributing to understanding of complex cognitive processes involved in V. germanica´s foraging behavior.     

Factors affecting use of mirex‐poisoned protein baits for control of European wasp (Paravespula germanica) in New Zealand

A bait of canned fish poisoned with 0.5 % or 1 % mirex substantially reduced populations of the European wasp, Paravespula germanica (Fab.), in a part‐cleared, part‐forested resort area in the Marlborough Sounds. In separate baiting regimes, bait was taken for as long as 9 weeks when 0.5% formulation was offered from 5 stations, but when 1 % formulation was offered from 10 and from 15 stations the time was reduced to 5 weeks and 4 weeks respectively. Synthetic and extracted materials tested as baits were unattractive to wasps, but fish baits attractive when fresh could be freeze‐dried without loss of acceptability. Marked wasps were caught up to 1200 m from their nests, and in the control programme 64 ppm mirex was recovered from dry brood comb of a defunct nest 700 m from the nearest bait source. The importance to bait control programmes of data on flight distance and foraging habits, and of the winter survival of colonies with prolonged queen production, is discussed. Some observations on reproductive biology are made in an appendix to the paper.     

Interactions between ants attendingAphis fabae ssp.cirsiiacanthoidis on thistles and foraging parasitoid wasps

We describe the behavioral interactions between honeydew-collecting workers of the ants Lasius nigerand Myrmica ruginodisand females of three species of aphidiid wasps (Lysiphlebus cardui, Lysiphlebus testaceipes, Trioxys angelicae)foraging for their aphid host, Aphis fabaessp. cirsiiacanthoidis,on thistles. Using field and laboratory experiments, we show that the ant-parasitoid interactions are species specific. Workers of both ant species generally attacked and killed females of T. angelicae,but they ignored those of L. cardui.This pattern was not altered when we anesthetized the wasps slightly with carbon dioxide to reduce their mobility. Prior contacts between L. carduiand either conspecific L. nigerfrom a different nest or workers of a different ant species (M. ruginodis)did not influence L. niger'snonaggressive behavior. The number of aphids parasitized by L. testaceipeswas significantly reduced in aphid colonies attended by L. niger,although this parasitoid was rarely attacked by ants. In encounters between these species of ants and wasps, ant aggression is consistent with differences in wasp behavior. We suggest that, in addition, chemical cues located in the cuticula may enable L. carduito avoid detection by honeydew-collecting ants.     

The importance of location and visual cues during foraging in the German wasp (Vespula germanica F.) (Hymenoptera: Vespidae)

Abstract

The way in which foraging wasps use cues for prey location and choice appears to depend on both the context and on the type of prey. Vespula germanica is an opportunistic, generalist prey forager, and individual wasp foragers often return to hunt at sites of previous hunting success. In this paper, we studied which cues are used by this wasp when relocating a food source. Particularly we analysed the response to a displaced visual cue versus a foraging location at which either honey or cat food had been previously presented. We conclude that location is used over a displaced visual cue for directing wasp hovering, although the landing response is directed differently according to bait type. When wasps are exploiting cat food, location also elicits landing, but if they are exploiting honey, a displaced visual cue elicits landing more frequently than location.     

Changes in abundance of Vespula germanica and V. vulgaris in England

1. Thirteen time series, varying from 17 to 27 years, of the abundance of Vespula germanica and V. vulgaris from lowland England are examined. The time series depend on either spring queens and workers taken in Malaise or suction traps, or collected colonies. 2. During the late 1970s and early 1980s, the abundance of V. germanica declined abruptly but that of V. vulgaris did not. 3. During the early 1980s, the 2‐year cycle of annual abundance of V. vulgaris changed to a nearly perfectly damped pattern of annual abundances. 4. The most likely factor causing these population changes was the increased use of pesticides acting directly by killing the wasps and indirectly by reducing their food resources. 5. The difference in response of the two species to increased pesticide use may be related to a difference in foraging ability.     

Competition for food between the exotic wasp <Emphasis Type="Italic">Vespula germanica</Emphasis> and the native ant assemblage of NW Patagonia: evidence of biotic resistance?

The success of a biological invasion may depend on the interactions between the invader and the native biota. However, little experimental evidence demonstrates whether local species can successfully compete with exotics. We experimentally determined the existence of competition for food between the exotic wasp Vespula germanica, one of the most recent Patagonian invaders, and the native ant assemblage. Both wasps and ants are generalist predators and scavengers, sharing habitat and food resources. We selected 30 sites within scrubland habitats where both ants and wasps were present. At each site, we placed containers with protein baits under three treatments: wasp exclusion, ant exclusion, and control (i.e., free access for wasps and ants). Ant exclusion increased the number of wasps (with regard to a control), but wasp exclusion did not affect ant abundance. This result suggests that native ants affect the foraging activity of exotic wasps but not vice versa. Aggressive behaviors and worker aggregation may explain the competitive advantage of ants. Ants bite wasp legs and massively aggregate on food sources, physically limiting the landing of wasps on baits. If the outcome of interactions at baits reported here influence wasp population-level parameters, this competitive interaction could be one of the factors explaining the low abundance of this exotic wasp in NW Patagonia in comparison with other invaded regions.     

Odour-mediated foraging by yellowjacket wasps (Hymenoptera: Vespidae): predation on leks of pheromone-calling Mediterranean fruit fly males (Diptera: Tephritidae)

Predation is probably the most important male mortality factor in insect species with courtship displays that render males performing them conspicuous targets of predators. Sexually active Mediterranean fruit fly males, Ceratitis capitata (Wied.), aggregate in leks, where they participate in agonistic encounters and engage in visual, acoustic and pheromone-calling displays to attract receptive females. The objective of this study was to assess: a) whether sexually displaying C. capitata males in leks inside host and non-host foliage are subject to predation by the most prominent predators yellow-jacket wasps, Vespula germanica (F.), and if so, b) whether olfactory, visual or auditive stimuli are used by foraging wasps in locating male C. capitata prey. Studies were carried out in a citrus orchard and surroundings on the island of Chios, Greece. Observations were conducted using perforated containers hung within mulberry, fig or citrus foliage. Living C. capitata flies of different sex and either mature or immature were placed inside. Our results show that the yellowjacket wasps have learned to associate the presence of sexually active medfly males aggregated in leks with their prey's pheromone (kairomone). Foraging wasps, flying through the crowns of host trees, responded to the odour source of C. capitata male pheromone by approaching from downwind. Even inside dense citrus tree foliage, wasps keyed in on aggregations of pheromone-calling males using olfactory stimuli. Stimuli of visual and acoustic male signalling were only used at close range, after having followed the pheromone plume close to its source. Visual cues played a greater role in directing wasp foraging under more open and exposed host foliage conditions. Odour-based foraging of wasps inside host foliage in the mid-morning hours, when medfly male lekking activities peak, shifted gradually to a more visual-based host fruit patrolling in the afternoons to capture ovipositing and feeding medfly females. On ripe fruit, particularly fig, V. germanica visual prey hunting also included the capture of feeding medfly males, other feeding Diptera, as well as medfly larvae extracted from wasp-made perforations in the fruit.     

Body temperature and thermoregulation in two species of yellowjackets,Vespula germanica and V. maculifrons

In spite of the abundance and broad distribution of social wasps, little information exists concerning thermoregulation by individuals. We measured body temperatures of the yellowjackets Vespula germanica and V. maculifrons and examined their thermoregulatory mechanisms. V. germanica demonstrated thermoregulation via a decreasing gradient between thorax temperature and ambient temperature as ambient temperature increased. V. maculifrons exhibited a constant gradient at lower ambient temperatures but thorax temperature was constant at high ambient temperatures. Head temperature exhibited similar patterns in both species. In spite of low thermal conductances, a simple heat budget model predicts substantial heat loads in warm conditions in the absence of thermoregulation. Both species regurgitated when heated on the head. A smaller volume of regurgitant was produced at lower head temperatures and a larger volume at higher head temperatures. Small regurgitations resulted in stabilization of head temperature, while large ones resulted in 4°C decreases in head temperature. Regurgitation was rare when wasps were heated upon the thorax. Abdomen temperature was 3–4°C above ambient temperature, and approached ambient temperature under the hottest conditions. No evidence was found for shunting of hot hemolymph from thorax to abdomen as a cooling mechanism. The frequency of regurgitation in workers returning to the nest increased with ambient temperature. Regurgitation may be an important thermoregulatory strategy during heat stress, but is probably not the only mechanism used in yellowjackets.Abbreviations M _b body mass - M _th thorax mass - T _a ambient temperature - T _ab abdomen temperature - T _b body temperature - T _h head temperature - T _th thorax temperature - C _t thermal conductance     

Meat ants as dominant members of Australian ant communities: an experimental test of their influence on the foraging success and forager abundance of other species

Meat ants (Iridomyrmex purpureus and allies) are perceived to be dominant members of Australian ant communities because of their great abundance, high rates of activity, and extreme aggressiveness. Here we describe the first experimental test of their influence on other ant species, and one of the first experimental studies of the influence of a dominant species on any diverse ant community. The study was conducted at a 0.4 ha savanna woodland site in the seasonal tropics of northern Australia, where the northern meat ant (I. sanguineus) represented 41% of pitfall catches and 73% of all ants at tuna baits, despite a total of 74 species being recorded. Meat ants were fenced out of experimental plots in order to test their influence on the foraging success of other species, as measured by access to tuna baits. The numbers of all other ants and ant species at baits in exclusion plots were approximately double those in controls (controlling for both the fences and for meat ant abundance), and returned rapidly to control levels when fences were removed after 7 weeks. Individual species differend markedly in their response to the fencing treatment, with species of Camponotus and Monomorium showing the strongest responses. Fencing had no effect on pitfall catches of species other than the meat ant, indicating that the effect of meat ants at baits was directly due to interference with foraging workers, and not regulation of general forager abundance. Such interference by meat ants has important implications for the sizes and densities of colonies of other ant species, and ultimately on overall ant community structure.     

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.     

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.     

The impact of an exotic social wasp (Vespula germanica) on the native arthropod community of north-west Patagonia, Argentina: an experimental study

Abstract 1. Biological invasions are usually thought to have a negative impact on native communities. However, data supporting this idea are often based on comparative studies between invaded and non‐invaded areas, and are spatially and temporally limited. 2. The present study experimentally assessed the impact of an exotic wasp, Vespula germanica, on the native arthropod community of north‐west Patagonia during 3 years in an area of 80 ha. Vespula germanica is an exotic social vespid that invaded north‐west Patagonia 20 years ago. It has been suggested that its populations affect native arthropods because of its broad diet and also because Patagonia lacks natural enemies and potential competitors for these wasps. 3. Using wasp‐specific toxic baits, V. germanica abundance was reduced in five sites of native woodlands during 3 consecutive years. The abundance, species richness, and composition of arthropods between non‐poisoned (control) and poisoned sites was then compared, both before and after the wasps were poisoned. 4. Wasp abundance represented 6% of the total arthropod catches in non‐poisoned sites and was reduced, on average, by 50% in the treated areas. The abundance, species richness, and composition of the arthropod community (305 species, 24 600 individuals) did not differ between control areas and areas where the abundance of V. germanica was reduced. Significant differences in response variables were found only before wasp poisoning had begun and were related to variations among sites. 5. These results suggest that V. germanica is not affecting the local arthropod assemblages, contradicting past work in other regions. The low relative abundance of wasps in Patagonia, when compared with other invaded regions, might explain the findings. 6. The present study provides further evidence for the importance of large‐scale experimental work with before/after comparisons to fully understand the impact of invaders on natural communities.     

Interspecific displacement mechanisms by the invasive little fire ant <Emphasis Type="Italic">Wasmannia auropunctata</Emphasis>

Competition between invasive species and native ones in the new environment was found to be significant and to affect both animal and plant species. Invasive ants are notorious for displacing local ant species through competition. Competitive displacement of native species can occur through interference and or resource competition. However, for invasive ants, little is known about the relative importance of competitive displacement. We studied competitive interactions of the little fire ant, Wasmannia auropunctata, one of the most destructive invasive ant species, with two other ant species, Monomorium subopacum and Pheidole teneriffana. We compared the species’ foraging behavior and studied their aggressive interactions around food baits for the short (2 h) and long (21 days) term in the laboratory. Surprisingly we found that in short term experiments W. auropunctata had the poorest foraging abilities of the three species studied: it took the workers the longest to locate the bait and retrieve it; in addition they retrieved the lowest amount of food. When both W. auropunctata and M. subopacum were foraging the same bait, in the short term competition experiment, W. auropunctata workers did not defend the bait, and ceased foraging when encountered with competition. The long-term experiments revealed that W. auropunctata had the advantage in aggressive interactions over time; they eliminated seven of nine M. subopacum’s nests while consuming some of the workers and brood. According to our laboratory studies, W. auropunctata cannot be considered an extirpator species, unless it has a substantial numerical advantage, in contrast with previous assumptions. Otherwise it may behave as an insinuator species, i.e. the workers do not initiate aggression and by staying undetected they can continue foraging adjacent to dominant species.     

<Emphasis Type="Italic">Vespula germanica</Emphasis> Foragers Do Not Scent-Mark Carbohydrate Food Sites

We tested whether foragers of Vespula germanica leave a chemical attractant at a carbohydrate food site. Foragers were allowed to make 50 or 100 visits to a feeder filled with a rich, unscented fructose solution. We then gave them a choice between this and an identical but unvisited feeder. There was no difference between the number of individuals that chose the visited feeder and those that chose the unvisited one. Instead, wasps showed a tendency to choose a feeder based on its relative position on the feeding stand. We conclude that foragers of V. germanica do not scent-mark food sites. Instead, in the absence of food odor, they rely on local visual cues to relocate carbohydrate food sources.     

Variation in colour markings of German wasps Vespula germanica (F.) and common wasps Vespula vulgaris (L.) (Hymenoptera: Vespidae) in New Zealand

Abstract

German wasps (Vespula germanica F.) and common wasps (V. vulgaris L.) both show variations in colour markings in New Zealand. Overlap in the ranges of markings of the head and thorax is limited enough for ready distinction of queens and workers of the two species. Abdominal markings overlap considerably between the species. Colour markings of males were too similar to identify the species, which were completely separated by the shape of their genitalia. The frequency of occurrence of “intermediate” markings in V. germanica populations did not increase in areas where the two species overlap. This suggests there is little or no genetic exchange between the two species.