Distribution,dispersal and spread of the invasive social wasp (Vespula germanica) in Argentina

We studied the distribution and spread of the invasive social wasp Vespula germanica in Argentina, focusing on the contribution of queen dispersal to territorial expansion. Vespula germanica is native to Eurasia and has invaded several regions of the world, including Southern Argentina. Flight potential of field‐collected queens was measured using flight mills. Also, by means of an extensive survey we estimated the rate of spread by analysing the relationship between years since arrival and distance from the introduction locality. The mean distance flown by wasp queens in flight mills was 404.7 ± 140.8 m (mean ± SE, n = 59), while the rate of spread of V. germanica was estimated at 37.2 ± 2.1 km year^?1 (mean ± SE, n = 67), although faster towards the south. The observed spread rate of V. germanica wasps in Argentina confirms the invasive potential shown by several Hymenoptera species worldwide. Still, a stratified geographical expansion pattern does not match observed queen dispersal abilities, suggesting that human‐aided transport of hibernating queens is the central driver of the current distribution of these wasps. We suggest that despite several life‐history traits known for social insects that contribute to successful invasion, wasp spread must still rely strongly on human mediated pathways. This observation sheds light on those factors that are crucial for managing invasions of this and related pestiferous wasps.     

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 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.     

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.     

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.     

Fitness and microbial networks of the common wasp,Vespula vulgaris (Hymenoptera: Vespidae), in its native and introduced ranges

1. Variation in microbial communities between populations is increasingly hypothesised to affect animal fitness and performance, including for invasive species. Pathogenic species may be lost during the introduction process, enhancing invader fitness and abundance. 2. This study assessed fitness, immune gene expression, and microbial network complexity of invasive common wasps, Vespula vulgaris. Microbial networks were assayed using 16S and 18S sequencing and gene expression arrays in the native (Belgium) and introduced range (New Zealand). The immune gene expression of the wasp Down syndrome cell adhesion molecule (Dscam) gene homologue was examined. Dscam expression can be induced by viruses, Gram‐positive and Gram‐negative bacteria, and parasites. 3. Individual nest fitness was higher in the native range of Belgium than in the introduced New Zealand range. Microbial communities of wasps in the introduced range were more diverse with more complex networks, although some microorganisms were range‐specific. Microbial networks in the introduced range showed higher clustering coefficients, number of connected paths, network centralisation, number of neighbours and network density. 4. Larvae, workers, virgin and foundress queens had higher expression of Dscam in the New Zealand samples. These immune gene expression patterns were associated with higher pathogen pressure and lower relative fitness. 5. Epidemiological theory predicts that a high density of pathogen and microbial hosts should result in a high rate of disease infection, prevalence, and highly connected microbial networks. The results of this study support these predictions. Wasps displayed lower relative fitness and more highly connected microbial networks in New Zealand than in Belgium.     

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.     

The spread of the social wasp,Vespula germanica,in Australia

Abstract

Between 1977 and 1989 the social wasp Vespula germanicacolonised mainland Australia and became established in New South Wales, Victoria, and South Australia. Accidental transport between towns is the main mechanism of spread. In many towns wasp nests have not been detected until they became widespread; therefore, eradication is difficult Successful eradication has, however, been achieved in many other towns.

Nesting places vary considerably among localities but relatively little among years at anyone locality. Peak wasp abundance occurs between January and April. Control of V. germanica nests costs Australia an estimated $600,000 (Aust. $) annually.     

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.     

Long-term impact of exotic ants on the native ants of Madeira

Abstract.  1. The earliest exotic records for two notorious invasive ants, the big-headed ant ( Pheidole megacephala ) and the Argentine ant ( Linepithema humile ), both come from the Atlantic islands of Madeira, where the two species underwent population explosions in the 1850s and 1890s respectively. Researchers have long assumed that these invaders spread across all of Madeira and exterminated most or all native ants, despite no research actually documenting such impact.
2. Re-examination of first-hand nineteenth century accounts suggest that P. megacephala and L. humile may never have spread beyond coastal lowland areas, representing < 10% of Madeira's land area. In 2002, native ants dominated most of Madeira; P. megacephala and L. humile were restricted to ≈ 0.3% and ≈ 6% of Madeira's land area respectively.
3. Of the 10 native ant species known from Madeira, only one ( Temnothorax wollastoni ) was not present in 1999–2002 surveys. Although exotic ants may have exterminated T. wollastoni , it seems likely that this species still survives.
4. Thus, even after 150 or more years of residence, P. megacephala and L. humile have come to occupy only a small part of Madeira, and appear to have had little impact.
5. Most of Madeira may be too cool for P. megacephala and perhaps too moist for L. humile to dominate. Also, Madeira's vast natural areas may generally lack weedy vegetation that can support high densities of plant-feeding Hemiptera critical for the ecological dominance of invasive ants. Finally, a dominant native ant, Lasius grandis , inhabiting ≈ 84% of Madeira, may actively exclude P. megacephala and L. humile .     

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.     

Species richness of herbivores on exotic host plants increases with time since introduction of the host

Aim Species richness of insect herbivores feeding on exotic plants increases with abundance as well as range size of the host in the area of introduction. The formation of these herbivore assemblages requires a certain amount of time, and the richness of insect faunas should also increase with the length of time an exotic plant has been present in the introduced range. Location Central Europe. Methods We analysed the variation in species richness of leaf‐chewing Lepidoptera larvae and sap‐sucking Auchenorrhyncha (Hemiptera) associated with 103 exotic woody plant species in Germany in relation to time since introduction, range size, growth form (trees versus shrubs), biogeographical origin (distance from Central Europe) and taxonomic isolation of the host plant (presence or absence of a native congener in the introduced area). Results Using simple correlation analyses we found for Lepidoptera and Auchenorrhyncha that species richness increased with time since introduction of the host plant. For the Lepidoptera the increase of species richness with time since introduction remained significant even after removing the effects of all other independent variables. Main conclusions Our results provide some evidence that assemblages of insects on exotic plants do not reach saturation within a time scale of few hundred years. This contrasts with previous findings for crop plants.     

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.     

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.     

Queen brood reared in worker cells by the social wasp,Vespula germanica (F.) (Hymenoptera: Vespidae)

Summary The rearing of queen brood from worker cells in the nests ofVespula germanica (F.) in Australia was found at all stages of the colony cycle from early summer to autumn. Worker cells used to rear queens were 17% wider at their open ends than cells used for worker production, and in all other respects their dimensions were greater. The volume of a worker cell used for queen nearing was 21% larger than cells used for rearing workers but half the volume of a normal queen cell. Queen pupae reared in worker cells were significantly smaller than those reared in queen cells of the same nest. The spatial distribution of queen pupae in worker comb tended to be random although some evidence of nonrandom clustering was noted.The significance of this phenomenon is discussed in relation to caste differentiation and queen-control of gyne initiation and production. It is concluded that the phenomenon is neither seasonal nor due to some intrinsic component of the colony's annual cycle, and that it is unlikely to be an emergency queen-rearing response. Development of queens in worker cells may have occurred during favourable feeding regimes, perhaps determined by the benign Australian environment.     

Evaluation of the toxicity of eggs,hatchlings and tadpoles of the introduced toad Bufo marinus (Anura: Bufonidae) to native Australian aquatic predators

Abstract The early life history stages of anurans in the Family Bufonidae often possess chemicals that are noxious or toxic to predators. Predators with no evolutionary history of exposure to bufomds may be particularly susceptible to these toxins. We conducted a series of laboratory experiments to investigate the toxic effects of eggs, hatchlings and tadpoles of the introduced toad, Bufo marinus (Linnaeus), on native Australian aquatic predators. There was considerable interspecific and intraspecific variation in these effects. Bufo marinus were highly toxic to some predator species, but were readily consumed by other species without apparent ill effect. Interspecific variation in toxic effects was not related to predator feeding mode or the number of B. marinus ingested by predators, and there was no clear pattern of distribution of vulnerability among species within higher taxa. Intraspecific variation in responses to toxins may result from individual variation in the resistance of predators to B. marinus toxins, or from individual variation in toxicity among B. marinus. Some native species adversely affected by B. marinus appeared unable to detect and avoid B. marinus toxins. This may result from a general inability to assess the toxicity of food items or from a lack of evolutionary exposure to B. marinus toxins.     

Extent of invasion of Tasmanian native vegetation by the exotic bumblebee Bombus terrestris (Apoidea: Apidae)

Abstract Observations of the large earth bumblebee, Bombus terrestris (L.), in native vegetation were collated to determine the extent to which this exotic species has invaded Tasmanian native vegetation during the first 9 years after its introduction. The range of B. terrestris now encompasses all of Tasmania's major vegetation types, altitudes from sea level to 1260m a.s.L, and the entire breadth of annual precipitation in the state from more than 3200 mm to less than 600 mm. Observations of workers carrying pollen, together with the presence of large numbers of bumblebees at many localities across this range indicate that colonies are frequently established in native vegetation. Evidence that colonies are often successful was obtained from repeated observations of the species during more than 1 year at particular sites. Unequivocal evidence of colonies was obtained from six National Parks, including four of the five in the Tasmanian Wilderness World Heritage Area (WHA). Indeed, the species has been present in the WHA for at least as long as it has in the city of Hobart, where it was first recorded. In southwestern Tasmania, evidence of colonies was obtained up to 40km from gardens, 61 km from small towns and 93 km from large towns. Hence, contrary to previous suggestions, the species is established in the most remote parts of Tasmania and is not dependent on introduced garden plants. Given their strong record of invasion, it is likely that B. terrestris will form feral populations on the mainland of Australia and in many other parts of the world if introduced. Because of their likely negative impacts on native animals and plants, and potential to enhance seed production in weeds, the spread of bumblebees should be avoided.