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.     

Local dynamics of worker activity of the invasive Vespula germanica and V. vulgaris (Hymenoptera: Vespidae) wasps in Argentina

1. The abundance of insects depends essentially on the reproductive success of individuals. In social insects, however, the abundance of sterile workers outside a nest depends on colony size but is also determined by ontogeny, nest demands, and local environmental factors. For invasive social wasps, the drivers of worker abundance are important because they determine the impact that these species have on the native systems, people, and their goods. 2. The aim of the present study was to understand the relative importance of endogenous and exogenous factors on the abundance of workers of populations of Vespula spp., by analysing 12 years of trap captures in NW Patagonia. This is the first attempt to model the activity levels of invasive Vespula spp. wasps over time in Argentina. 3. It was shown that between years, the worker activity of both vespids presents fluctuations, and that of V. germanica is determined by the spring mean temperatures. Within the flight season, V. germanica worker activity is affected by the relative abundance of workers in that year, whereas for V. vulgaris, activity it is affected by the relative abundance of both species that year. We found no relationship between individual weather variables and activity within a flight season for both wasps. 4. The patterns observed for Argentina are similar to those observed in all invaded temperate areas where Vespula spp. are established. This study provides useful information to understand the driving factors that affect Vespula spp. worker activity in Argentina. This could be a necessary step to develop plans to manage these invasive social insects.     

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.     

Long‐term spatial memory in Vespula germanica social wasps: the influence of past experience on foraging behavior

Social insects exhibit complex learning and memory mechanisms while foraging. Vespula germanica (Fab.) (Hymenoptera: Vespidae) is an invasive social wasp that frequently forages on undepleted food sources, making several flights between the resource and the nest. Previous studies have shown that during this relocating behavior, wasps learn to associate food with a certain site, and can recall this association 1 h later. In this work, we evaluated whether this wasp species is capable of retrieving an established association after 24 h. For this purpose, we trained free flying individuals to collect proteinaceous food from an experimental plate (feeder) located in an experimental array. A total of 150 individuals were allowed 2, 4, or 8 visits. After the training phase, the array was removed and set up again 24 h later, but this time a second baited plate was placed opposite to the first. After 24 h we recorded the rate of wasps that returned to the experimental area and those which collected food from the previously learned feeding station or the nonlearned one. During the testing phase, we observed that a low rate of wasps trained with 2 collecting visits returned to the experimental area (22%), whereas the rate of returning wasps trained with 4 or 8 collecting visits was higher (51% and 41%, respectively). Moreover, wasps trained with 8 feeding visits collected food from the previously learned feeding station at a higher rate than those that did from the nonlearned one. In contrast, wasps trained 2 or 4 times chose both feeding stations at a similar rate. Thus, significantly more wasps returned to the previously learned feeding station after 8 repeated foraging flights but not after only 2 or 4 visits. This is the first report that demonstrates the existence of long‐term spatial memory in V. germanica wasps.     

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.     

Carbohydrate bait preferences of wasps (Vespula vulgaris and V. germanica) (Hymenoptera: Vespidae) in New Zealand

Abstract

Carbohydrates are both a source of food and a potential bait for control of common and German wasps. Carbohydrate preferences of wasps were determined in a series of paired trials by counting the number of wasps caught in traps baited with carbohydrate‐based baits versus a standard protein‐based bait. Factors influencing wasp attraction to carbohydrate‐based baits included bait type, season, weather, and location. A 30% sucrose solution was more attractive than honey, jams, gels, or dry (solid) sugars. The sucrose solution was more attractive in spring and autumn than in summer, probably because rainfall washed away natural carbohydrate supplies. There was a significant relationship between rainfall in the previous week and the proportion of wasps caught in traps baited with sucrose solution versus sardine cat‐food. A higher proportion of German wasps than common wasps was caught in sucrose‐baited traps. This may have been related to seasonal differences in abundance and/or behavioural differences between the two species. In some locations, more wasps were caught in traps baited with sucrose solution than in those baited with sardine cat‐food. The proportion of wasps caught in sucrose‐baited traps was higher in locations without beech scale honeydew than with beech scale honey‐dew. Proportionally more non‐target species (mainly honey bees) and fewer wasps were caught in traps baited with sucrose solution than in those baited with sardine cat‐food. All five bee repellents tested also repelled wasps. These results indicate the importance of determining the bait preferences of wasps (carbohydrate or protein) and the presence of non‐target species before attempting a poison‐baiting operation. They also emphasise the need for a wasp‐specific carbohydrate‐based bait.     

Overwintering colonies of German (Vespula germanica) and common wasps (Vespula vulgaris) (Hymenoptera: Vespidae) in New Zealand

Abstract

Nests of both common (Vespula vulgaris) and German wasps (V. germanica) sometimes overwinter in New Zealand. Three overwintering common wasp colonies were found in low-altitude honeydew beech forest; about 2% of the colonies initiated there in 1988 survived the winter. Wasp traffic rates from nests in Nelson city and a nation wide survey of wasp abundance, showed that more German wasp than common wasp workers were on the wing in winter and spring. German wasp colonies in Nelson city were more likely to overwinter than were common wasp colonies. Of the active German wasp colonies recorded in Nelson in January and February 1989, only two (4%) had previously overwintered, but these two nests accounted for 38% of all German wasp workers estimated to leave nests in the area. Had other overwintering colonies not been poisoned, overwintering colonies might have accounted for up to 11 % of nests and produced up to 64% of German wasp workers on the wing in January and February 1989.

Overwintering common wasp colonies did not produce queens or drones in their first year or second spring, but all 10 overwintering German wasp colonies examined produced sexuals at both times. German wasp queens produced in winter and spring may influence the number of colonies successfully initiated and affect the population dynamics of German wasps in New Zealand.     

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.     

Colony structure and nest characteristics of European wasps, Vespula germanica (F.) (Hymenoptera: Vespidae), in Victoria, Australia

Abstract European wasps, Vespula germanica , are common across southern and south-eastern Australia and have a negative impact upon urban areas, primary industries, and natural ecosystems. Aspects of colony structure and nest characteristics are examined for nests located in the ground and collected from urban and rural sites in Victoria during two summer−autumn field seasons (1996 and 2001). On average, nests were located 28 cm beneath the surface (range 5−58 cm). The average number of combs in the nest and the total area of the nests increased from early February to late May. In addition, the proportion of cells used to produce workers and different life stages (larvae, pupae) also varied throughout the season; being consistent with studies from New Zealand. No differences in colony structure or nest character­istics were detected between urban and rural nests.     

Dispersal behavior of yellowjacket (Vespula germanica) queens

Understanding the factors that affect animal dispersal behavior is important from both fundamental and applied perspectives. Dispersal can have clear evolutionary and ecological consequences, but for nonnative insect pests, dispersal capacity can also help to explain invasion success. Vespula germanica is a social wasp that, in the last century, has successfully invaded several regions of the world, showing one of the highest spread rates reported for a nonnative insect. In contrast with nonsocial wasps, in social species, queens are responsible for population redistribution and spread, as workers are sterile. For V. germanica, it has been observed that queen flight is limited to 2 distinct periods: early autumn, when new queens leave the nest to mate and find sheltered places in which to hibernate, and spring when new colonies are founded. Our aim was to study the flight behavior of V. germanica queens by focusing on the different periods in which dispersal occurs, characterizing as well the potential contribution of queen flight (i.e., distance) to the observed geographical spread. Our results suggest that the distances flown by nonoverwintered queens is greater than that flown by overwintered individuals, suggesting that the main queen dispersal events would occur before queens enter hibernation. This could relate to a behavioral trait of the queens to avoid the inbreeding with related drones. Additionally, given the short distances flown and remarkable geographical spread observed, we provide evidence showing that queen dispersal by flight is likely to contribute proportionately less to population spread than human‐aided factors.     

The influence of past experience on wasp choice related to foraging behavior

Memory has been little studied in social wasps. Vespula germanica （Fab.） （Hymenoptera： Vespidae） frequently revisits nondepleted food sources, making several trips between the resource and the nest. In this study, we analyzed this relocating behavior in order to evaluate whether this species is capable of remembering an established association after 1 h. To this end, we trained wasps to feed from a certain array. Then it was removed, setting it up again 1 h later, but this time 2 baited feeders were put in place, one at the original feeding site and the other opposite the first. We recorded the proportion of returning foragers, and their choice of feeder, after either 1 or 4 feeding trials. After 1 h, 78% of wasps trained with 4 feeding trials and 65% trained with 1, returned to the experimental area. Furthermore, during the testing phase, wasps trained with 4 feeding trials collected food from the previously learned feeder significantly more frequently than from the nonlearned one （P 〈 0.05）. In contrast, wasps that had been trained only once chose both feeders equally. Thus, memory retrieval could be observed 1 h after wasps had collected food on 4 consecutive occasions, but not after only 1. To our knowledge, this is the first study showing that V. germanica is capable of remembering an association 1 h after the last associative event, demonstrating that 1 h does not impair memory retention if4 feeding experiences have occurred.     

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     

Olfactory Learning of Fruit Odors in the Eastern Yellow Jacket, Vespula maculifrons (Hymenoptera: Vespidae)

Food-seaching workers of eastern yellow jackets, Vespula maculifrons, are attracted by the natural odors of a wide variety of succulent fruits; particularly effective was pear. The only part of a fruit that repelled was the leathery epicarp of oranges. After rewarding with sugar water, odors of six fruits, including the pulpy mesocarp of oranges and, in addition, the leaves of catmint Nepeta cataria, all become highly attractive. To learn the distinctive odors of any one of three fruits (pear, apple, quince), nondiscrimination training with a rewarded fruit was sufficient for the subsequent olfactory preference of the training fruit over the control fruit. In the other cases [banana, hawthorn (Crataegus crus-galli), grape] simultaneous discrimination training with a rewarded and an unrewarded fruit was necessary and effective for obtaining differential responses to the odors of the training fruits. As far as current evidence goes, olfactory learning plays similar roles in the fruit foraging of this wasp and in the nectar foraging of the honey bee (Apis mellifera).     

Foraging and spatial dispersion in protein-scavenging workers ofVespula germanica andV. maculifrons (Hymenoptera: Vespidae)

A series of experiments on the discovery of meat bait by scavenging workers ofV. germanica andV. maculifrons showed that workers oriented to meats by visual cues and olfactory cues. Scavening workers ofV. germanica andV. maculifrons responded positively to the sight of conspecifics at meat baits; i.e., meat with (confined) wasps received more visits than meat without wasps. This attraction to conspecific workers on meat baits provides evidence of local enhancement, in that the presence of conspecifics at meat bait directs other wasps to a particular spatial locality. Local enhancement by yellowjackets foraging for meat may contribute to the clumped distribution of worker populations noted in studies of workers captured in meat-baited traps. We hypothesize that local enhancement of foraging is a ubiquitous behavior in species of theV. vulgaris species group and may be one attribute explaining their ecological success.     

The potential distribution of German wasps (Vespula germanica) in Argentina

Abstract

In the early 1980s, the German wasp (Vespula germanica) arrived in Argentina and became established in northern Patagonia. We use an ecoclimatic computer‐based program (CLIMEX) to predict the potential spread of wasps to other areas of the country. The model suggests that much of Argentina does not appear to be very suitable for the species’ establishment. However, more suitable areas than those currently colonised by the wasp occur in the centre of the country. Based on CLIMEX results, we suggest that, if the wasps ever break through the barrier of unfavourable habitat that surrounds their current distribution, they will continue dispersing to those areas. Notwithstanding this, factors other than climate may explain why German wasps are firmly established in areas otherwise considered to be inhospitable for the species.     

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.     

Distribution of social wasps (Hymenoptera: Vespidae) in New Zealand in 1987

Abstract

More than 50 000 social wasps (Hymenoptera: Vespidae) were collected between March and August 1987 from throughout New Zealand. The most widespread species is the German wasp (Vespula germanica). The common wasp (V. vulgaris) has colonised about half the country and appears to be still spreading. It tends to appear in urban areas first, presumably because it is transported there with people and/or their belongings. The Asian paper wasp (Polistes chinensis) and Australian paper wasp (P. humilis), are confined to the northern half of the North Island, but the former species is spreading south.     

Control of Vespula germanica (Hym. Vespidae) populations using toxic baits: bait attractiveness and pesticide efficacy

Abstract:  Our aim was to test a set of baits and pesticides for the control of yellowjacket populations in NW Patagonia (Argentina), through field trials. We tested the attractiveness of protein (fresh and freeze-dried beef) and carbohydrate (corn syrup and honey) baits (alone and mixed) and three pesticides commonly employed to control terrestrial domestic arthropods (hydramethylnon 2%, permethrin 0.3% and chlorpyrifos 0.25%). Our results show that beef proved to be the most attractive bait tested, for Vespula germanica wasps in NW Patagonia. Honey and corn syrup alone or mixed with beef did not attract foraging wasps as did beef-only baits throughout the wasp season. Additionally, we show that the attractiveness of lyophilized beef baits is similar to that of fresh beef. The efficacy of the insecticides tested was limited. In fact, only baiting with hydramethylnon 2% reduced wasp populations (54%) and this occurred after 72 h since poisoning. However, given the potential of insects to develop resistance to a consistent use of a single product (e.g. fipronil 0.1%), the use of hydramethylnon may allow the alternating of insecticides in specific situations. The information provided here contributes to the existent knowledge on baits and insecticides for the control of yellowjackets using toxic baits.     

Comparative characteristics of nests of Vespula germanica (F.) and Vespula vulgaris (L.) (Hymenoptera: Vespinae) from Christchurch City,New Zealand

Abstract

From July 1987 to June 1989, 141 nests and 365 nests, respectively, of the adventive social vespines the German wasp, Vespula germanica (F.), and the common wasp V. vulgaris (L.), from the City of Christchurch, New Zealand, were evaluated for seven major characteristics.

There were few, if any, differences in nest sites, and from spring to early summer in nest traffic, nest size, and numbers of combs. However, Getman wasps showed no preference for direction of nest entrances, while common wasp nests were more numerous in areas most exposed to the morning sun. Some Getman wasp nests survived the winter and began producing new worker cells by late June, but all common wasp nests died by June. Because common wasp nests have been reported surviving the winter in beech forest, which produces honey dew, and honey dew is not available in Christchurch City, common wasp nest survival over winter may be more dependent upon carbohydrates and/or prey dependent upon carbohydrates than the survival of German wasp nests. The presence of large, expanding wasp nests from early spring must impose localised predation pressures virtually unknown in the Northern Hemisphere where overwintering nests are rare. Wasp population dynamics, and impacts of wasps on fauna, are likely to vary between different geographical areas of New Zealand, but as targets for biological control, the two wasp species can generally be considered to be quite similar.     

Morphology, echolocation and foraging behaviour in two sympatric sibling species of bat (Tylonycteris pachypus and Tylonycteris robustula) (Chiroptera: Vespertilionidae)

We studied the wing morphology, echolocation calls, foraging behaviour and flight speed of Tylonycteris pachypus and Tylonycteris robustula in Longzhou County, South China during the summer (June–August) of 2005. The wingspan, wing loading and aspect ratio of the two species were relatively low, and those of T. pachypus were lower compared with T. robustula . The echolocation calls of T. pachypus and T. robustula consist of a broadband frequency modulated (FM) sweep followed by a short narrowband FM sweep. The dominant frequency of calls of T. pachypus was 65.1 kHz, whereas that of T. robustula was 57.7 kHz. The call frequencies (including highest frequency of the call, lowest frequency of the call and frequency of the call that contained most energy) of T. pachypus were higher than those of T. robustula , and the pulse duration of the former was longer than that of the latter. The inter-pulse interval and bandwidth of the calls were not significantly different between the two species. Tylonycteris pachypus foraged in more complex environments than T. robustula , although the two species were both netted in edge habitats (around trees or houses), along pathways and in the tops of trees. Tylonycteris pachypus flew slower (straight level flight speed, 4.3 m s⁻¹) than T. robustula (straight level flight speed, 4.8 m s⁻¹). We discuss the relationship between wing morphology, echolocation calls, foraging behaviour and flight speed, and demonstrate resource partitioning between these two species in terms of morphological and behavioural factors.