Distribution, Invasion History, and Ecology of Non-native Pine Bark Beetles (Coleoptera: Curculionidae: Scolytinae) in Southern South America.

The growth of international trade, coupled with an expansion of large-scale pine plantations in South America during the second half of the twentieth century, has significantly increased the opportunities for the invasion of forest insects. Bark beetles (Coleoptera: Curculionidae, Scolytinae) are a large and diverse group of insects, commonly recognized as one of the most important tree mortality agents in coniferous forests worldwide and an important group among invasive forest species. In this study, we combined data from field sampling with published records of established non-native pine bark beetles, to describe their distribution and invasion history in pine plantations across southern South America, reviewing the available information on their phenology and host range. We obtained records of established populations of six Eurasian species distributed in two major regions: the southwest region comprises plantations in Chile and the Argentine Patagonia, with four bark beetle species: Hylastes ater, Hylastes linearis, Hylurgus ligniperda, and Orthotomicus laricis; the northeastern zone includes northeastern Argentina, Uruguay, and southern Brazil, and includes three bark beetle species: Cyrtogenius luteus, H. ligniperda, and O. erosus. The establishment of non-native populations across the study area began in the 1950s, and from the 1980s onwards, there has been an exponential increase in introductions. We predict that several of these species will continue spreading across South America and that new species will continue arriving. We highlight the importance of international collaboration for early detection and management of non-native pine bark beetles.

Gustatory responsiveness in Vespula germanica workers: exploring the interplay between sensory perception and task specialization.

Workers' task specialization and division of labor are critical features of social insects' ecological success. It has been proposed that the division of labor relies on response threshold models: individuals varying their sensitivity (and responsiveness) to biologically relevant stimuli and performing a specific task when a stimulus exceeds an internal threshold. In this work, we study carbohydrate and protein responsiveness and their relation to worker task specialization in Vespula germanica, an invasive social wasp. The sucrose and peptone responsiveness of two different subcastes, preforagers and foragers, was determined by stimulating the antenna of the wasps with increasing concentrations of the solution and quantifying whether each concentration elicited a licking response. We studied responsiveness in five different ways: (1) response threshold, (2) concentration 50 (concentration to which at least 50% of wasps responded), (3) maximum response, (4) mean scores and (5) median scores. Our results suggest that V. germanica foragers are more sensitive to sucrose (lower thresholds) than preforager workers. However, we found no differences for peptone thresholds (i.e., a protein resource). Nonetheless, this is the first study to investigate response thresholds for protein resources. The intercaste variation in sucrose responsiveness shown in our work contributes to the existing knowledge about response threshold theory as a mechanism for task specialization observed in V. germanica.

Allee effects in an invasive social wasp: an experimental study in colonies of Vespula germanica.

Allee effects rely on the existence of mutually beneficial intraspecific interactions that increase individual fitness and per capita growth rate, as the number of individuals in a population or group increases. When the number of individuals falls below a given number, the success of a group or population drops. Social insects heavily rely on cooperation between individuals for various tasks such as foraging and breeding. In this study, we experimentally explored component Allee effects and the underlying mechanisms in colonies of the social wasp Vespula germanica. After the removal of workers, we counted the number of gynes produced, estimated the body mass index as a proxy of their quality, and registered the protein-food collected by foragers in colonies. Our research revealed a correlation between the decrease in worker population and a subsequent decrease in the production of gynes. However, the removal of workers did not impact the quality of the produced gynes or the quantity of protein-food collected by the colonies. These findings highlight the crucial role of the worker caste in the success of eusocial species and suggested an ability of workers to perform multiple tasks that enable colonies to respond to disturbances. Additionally, our study provides the first evidence of Allee effects at the colony level of V. germanica, with potential practical implications for managing this invasive species.

Climate, host and geography shape insect and fungal communities of trees.

Non-native pests, climate change, and their interactions are likely to alter relationships between trees and tree-associated organisms with consequences for forest health. To understand and predict such changes, factors structuring tree-associated communities need to be determined. Here, we analysed the data consisting of records of insects and fungi collected from dormant twigs from 155 tree species at 51 botanical gardens or arboreta in 32 countries. Generalized dissimilarity models revealed similar relative importance of studied climatic, host-related and geographic factors on differences in tree-associated communities. Mean annual temperature, phylogenetic distance between hosts and geographic distance between locations were the major drivers of dissimilarities. The increasing importance of high temperatures on differences in studied communities indicate that climate change could affect tree-associated organisms directly and indirectly through host range shifts. Insect and fungal communities were more similar between closely related vs. distant hosts suggesting that host range shifts may facilitate the emergence of new pests. Moreover, dissimilarities among tree-associated communities increased with geographic distance indicating that human-mediated transport may serve as a pathway of the introductions of new pests. The results of this study highlight the need to limit the establishment of tree pests and increase the resilience of forest ecosystems to changes in climate.

Oviposition substrate location by the invasive woodwasp Sirex noctilio: the combined effect of chemical cues emitted by its obligate symbiont Amylostereum areolatum and different host-tree species.

BACKGROUND: Sirex noctilio is an invasive forest wasp that affects pines and can result in severe economic losses. The use of semiochemicals offers an opportunity to develop sensitive and specific capturing systems to mitigatenegative impacts. Previous research showed that female S. noctilio would use volatiles emitted by its fungal symbiont, Amylostereum areolatum, but little is known about how these modulate behaviour when combined with pine-wood emissions. Our aim was to understand the relevance of fungal volatiles grown on artificial media and wood from two hosts trees, Pinus contorta and Pinus ponderosa, on behavioural and electroantennographic responses of wasp females. Because background odours can modify an insect's response towards resource-indicating semiochemicals, we propose that the behaviour towards the symbiont (resource) will be modulated by host pine emissions (background odours). RESULTS: Olfactometric assays showed that both host species with fungus were attractive when contrasted against air (P. contorta versus Air, χ2 = 12.19, P < 0.001; P. ponderosa versus Air, χ2 = 20.60, P < 0.001) and suggest a clear hierarchy in terms of female preferences towards the tested stimuli, with response highest towards the fungus grown on P. contorta (olfactory preference index: 5.5). Electrophysiological analyses indicate that females detect 62 volatile compounds from the tested sources. CONCLUSION: Results indicate a strong synergy between symbiont and host semiochemicals, suggesting that the pine species could play a fundamental role in the interaction. Further understanding of the chemical basis of this, could guide the development of specific and attractive lures, in order to maximize attraction of wasps in surveillance programmes. © 2023 Society of Chemical Industry.

Spatiotemporal outbreak dynamics of bark and wood-boring insects.

Bark and wood-boring insects (BWI) can cause important disturbances in forest ecosystems, and their impact depends on their spatiotemporal dynamics. Populations are usually at stable, low densities but can be disrupted by stochastic perturbations that trigger a transition to an epidemic phase. For less aggressive species, outbreaks die out quickly once the perturbation is removed, while aggressive species exhibit density-dependent feedback mechanisms facilitating persistent landscape-scale outbreaks. The interactions of attributes of tree, stand, landscape- and regional-scale stressors, and insect life history and behavior determine system-specific dynamics. However, most of our knowledge is based on a few species of mainly Holarctic bark beetles. With global change, it is becoming increasingly important to improve our understanding of the frequency and severity of BWI outbreaks.

Long-term coexistence of two invasive vespid wasps in NW Patagonia (Argentina).

In Patagonia (Argentina) two non-native vespid wasps became established in the last decades. Vespula germanica was first detected in 1980, while V. vulgaris arrived some 30 years later. Both species can have a strong negative impact on agro-industrial economic activities, the natural environment, and outdoor human activities. Biological invasions may be influenced negatively by the degree of interaction with the resident native community and alien species already present. The sequential arrival and coexistence of Vespula wasps in Argentina for several years allows us to understand key questions of invasion ecology. Additionally, recognizing the outcome of the invasion by vespids in Patagonia, a region lacking native social wasps, may help plan species-focused mitigation and control strategies. We explored the role of competition in terms of invasion success, and the strategies that promote coexistence. Two possible scenarios, using niche overlap indices and isocline equations, were proposed to determine competition coefficients. Using a simple mathematical modeling framework, based on field collected data, we show that food resources do not play a central role in competitive interaction. The competition coefficients obtained from the equations were different from those inferred from the overlap indices (0.53 and 0.54-0.076 and 0.197, respectively). Together, these findings suggest that no matter the arrival order, V. vulgaris, always reaches higher densities than V. germanica when both species invade new regions. Our work contributes to further our understanding on the worldwide invasion processes deployed by these two eusocial insects.

Integrated Pest Management Education: a Video-Game to Improve Management of Drosophila suzukii, Soft-Skin Fruit Pest.

The transition from the conventional global agriculture to an agroecological model poses a teaching and learning challenge to facilitate the wide variety of practices and the many biological processes involved. Games, such as board games, video-games, or mobile apps, are elements that may be used for teaching agroecology, environmental education, or animal conservation. Here, we present a video-game designed to contribute to integrated pest management education. The Spotted-Stop-It video-game is a single-player game which encourages participation, disseminates knowledge on the pest problem and its potential solutions (i.e., harvest frequency, sanitation, and management of alternative non-crop plants), and highlights the importance of good practices from an agroecological perspective at the farmer scale. In a farm-tech regional fair, we presented the game to its users and performed a simple retrospective survey. The survey results showed that most participants did not know about the fly prior to playing the game (34 ind., 68%), but were able to recognize the species among other flies after playing (23 ind., ~ 65%). Also, 21 individuals correctly responded about the effects of this pest on soft-skin fruits ("the fly lays eggs inside the fruit"). The training of future generations on new insect invasions and IPM practices with elements of their own environment may prove to be important to transmit concepts and practices at the service of sustainable crop protection.

Worldwide diversity of endophytic fungi and insects associated with dormant tree twigs.

International trade in plants and climate change are two of the main factors causing damaging tree pests (i.e. fungi and insects) to spread into new areas. To mitigate these risks, a large-scale assessment of tree-associated fungi and insects is needed. We present records of endophytic fungi and insects in twigs of 17 angiosperm and gymnosperm genera, from 51 locations in 32 countries worldwide. Endophytic fungi were characterized by high-throughput sequencing of 352 samples from 145 tree species in 28 countries. Insects were reared from 227 samples of 109 tree species in 18 countries and sorted into taxonomic orders and feeding guilds. Herbivorous insects were grouped into morphospecies and were identified using molecular and morphological approaches. This dataset reveals the diversity of tree-associated taxa, as it contains 12,721 fungal Amplicon Sequence Variants and 208 herbivorous insect morphospecies, sampled across broad geographic and climatic gradients and for many tree species. This dataset will facilitate applied and fundamental studies on the distribution of fungal endophytes and insects in trees.

Flexible foraging decisions made by workers of the social wasp Vespula germanica (Hymenoptera: Vespidae) in response to different resources: influence of ontogenetic shifts and colony feedback.

Decisions made by foraging animals conform a complex process based on the integration of information from multiple external environmental stimuli and internal physiological signals, which in turn are modulated by individual experience and a detection threshold of each individual. For social insects in which foraging is limited to given age subcastes, individual foraging decisions may also be affected by ontogenetic shifts and colony requirements. We studied the short-term changes in foraging preferences of the generalist wasp Vespula germanica, focusing on whether the individual response to different resources could be influenced by the ontogenetic shifts and/or by social interaction with nestmates. We carried both laboratory and field experiments to confront worker wasps to a short-term resource switch between either protein or carbohydrate-based foods. We tested the response of (1) Preforager workers (no foraging experience nor interaction with other wasps), (2) Forager workers (experience in foraging and no colony feedback), and (3) Wild forager workers (foraging naturally and exposed to free interactions with nestmates). We evaluated the maxilla-labium extension response (MaLER) for laboratory assays or the landing response for field assays. We observed that for wasps deprived of colony feedback (either preforagers or foragers), the protein-rich foods acceptance threshold increased (and thus a lower level of foraging on that item was observed) if they had foraged on carbohydrates previously, whereas carbohydrates were accepted in all assays. However, wasps immersed in a natural foraging context did accept protein foods regardless of their first foraging experience and reduced the carbohydrates collected when trained on protein foods. We provide evidence that short-term changes in foraging preferences depend on the type of resource foraged and on the social interactions, but not on ontogenetic shifts.

Nest-departure behaviour of gynes and drones in the invasive yellowjacket Vespula germanica (Hymenoptera: Vespidae).

Inbreeding costs can be high in haplodiploid hymenopterans due to their particular mechanism of sex determination (i.e., single-locus complementary sex-determination system, sl-CSD), as it can lead to the production of sterile males. Therefore, mechanisms contributing to reduced inbred matings can be beneficial. In this sense, asynchronous nest departure of sibling drones and gynes could reduce kin encounters in social hymenopterans. Using six observation colonies, we determined under field conditions the nest departure behaviour of sibling reproductives of the social wasp Vespula germanica (Hymenoptera: Vespidae). We determined that sexuals leave the nests definitively and detected asynchronous departure not fixed to a particular caste at a seasonal scale in some colonies, as gynes or drones delayed their departure as a function of the departure of the opposite sex, depending on the colony. At a higher temporal resolution (i.e., within a day), we discovered that drones consistently began to leave nests 1 h before gynes and this difference was driven by those individuals that left on the same day as did the opposite-sex kin. Even though other mechanisms such as polyandry and differential dispersal could also be important at reducing inbred matings in the species, the observed departure patterns (i.e., in some colonies actually leave together with the opposite caste, while in others temporal segregation seems to occur) from nests could be complementary to the former and be important at reducing the negative effects of inbreeding in this invasive species.

Drone aggregation behavior in the social wasp Vespula germanica (Hymenoptera: Vespidae): Effect of kinship and density.

Inbreeding can have negative consequences on population viability because of the reduced fitness of the progeny. In general, most species have developed mechanisms to minimize inbreeding such as dispersal and kin avoidance behavior. In the eusocial Hymenoptera, related individuals typically share a common nest and have relatively short mating periods, this could lead to inbreeding, and because of their single-locus complementary sex determination system, it may generate diploid males that could result in infertile triploid progeny representing a cost for the colony. Vespula germanica, is an eusocial wasp that has invaded many parts of the world, despite likely facing a reduced genetic pool during the arrival phases. We ask whether male wasp display specific aggregation behavior that favors genetic diversity, key to reduce inbreeding. Through a set of laboratory experiments, we investigated the effects of drone nestmateship and density on the aggregation behavior of V. germanica drones. We show that drones avoid aggregating with their nestmates at all densities while non-nestmates are avoided only at high densities. This suggests that lek genetic diversity and density could be regulated through drone behavior and in the long run minimize inbreeding favoring invasion success.

Drivers of global Scolytinae invasion patterns.

Biological invasions are affected by characteristics of invading species, strength of pathway connectivity among world regions and habitat characteristics of invaded regions. These factors may interact in complex ways to drive geographical variation in numbers of invasions among world regions. Understanding the role of these drivers provides information that is crucial to the development of effective biosecurity policies. Here we assemble for the first time a global database of historical invasions of Scolytinae species and explore factors explaining geographical variation in numbers of species invading different regions. This insect group includes several pest species with massive economic and ecological impacts and these beetles are known to be accidentally moved with wood packaging in global trade. Candidate explanatory characteristics included in this analysis are cumulative trade among world regions, size of source species pools, forest area, and climatic similarity of the invaded region with source regions. Species capable of sib-mating comprised the highest proportion on nonnative Scolytines, and these species colonized a higher number of regions than outbreeders. The size of source species pools offered little power in explaining variation in numbers of invasions among world regions nor did climate or forest area. In contrast, cumulative trade had a strong and consistent positive relationship with numbers of Scolytinae species moving from one region to another, and this effect was highest for bark beetles, followed by ambrosia beetles, and was low for seed and twig feeders. We conclude that global variation in Scolytine invasions is primarily driven by variation in trade levels among world regions. Results stress the importance of global trade as the primary driver of historical Scolytinae invasions and we anticipate other hitchhiking species would exhibit similar patterns. One implication of these results is that invasions between certain world regions may be historically low because of past low levels of trade but future economic shifts could result in large numbers of new invasions as a result of increased trade among previously isolated portions of the world. With changing global flow of goods among world regions, it is crucial that biosecurity efforts keep pace to minimize future invasions and their impacts.

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.

Brewer's Yeast, Saccharomyces cerevisiae, Enhances Attraction of Two Invasive Yellowjackets (Hymenoptera: Vespidae) to Dried Fruit and Fruit Powder.

The German yellowjacket, Vespula germanica F., and common yellowjacket, Vespula vulgaris L. (Hymenoptera: Vespidae), are pests of significant economic, environmental, and medical importance in many countries. There is a need for the development and improvement of attractive baits that can be deployed in traps to capture and kill these wasps in areas where they are a problem. Yellowjackets are known to feed on fermenting fruit, but this resource is seldom considered as a bait due to its ephemeral nature and its potential attractiveness to nontarget species. We analyzed the headspace volatiles of dried fruit and fruit powder baits with and without Brewer's yeast, Saccharomyces cerevisiae, using gas chromatography-mass spectrometry, and we field tested these baits for their attractiveness to yellowjackets in Argentina. The addition of yeast to dried fruit and fruit powder changed the volatile compositions, increasing the number of alcohols and acids and decreasing the number of aldehydes. Dried fruit and fruit powder baits on their own were hardly attractive to yellowjackets, but the addition of yeast improved their attractiveness by 9- to 50-fold and surpassed the attractiveness of a commercial heptyl butyrate-based wasp lure. We suggest that further research be done to test additional varieties and species of yeasts. A dried fruit or fruit powder bait in combination with yeast could become a useful tool in the management of yellowjackets.

Predicting North American Scolytinae invasions in the Southern Hemisphere.

Scolytinae species are recognized as one of the most important tree mortality agents in coniferous forests worldwide, and many are known invaders because they are easily transported in wood products. Nonnative trees planted in novel habitats often exhibit exceptional growth, in part because they escape herbivore (such as Scolytinae) pressure from their native range. Increasing accidental introductions of forest pest species as a consequence of international trade, however, is expected to diminish enemy release of nonnative forest trees. In this context, there is need to characterize patterns of forest herbivore species invasion risks at global scales. In this study, we analyze the establishment potential of 64 North American Scolytinae species in the Southern Hemisphere. We use climate-based ecological niche models (MaxEnt) to spatially define the potential distribution of these Scolytinae species in regions of the Southern Hemisphere were pines are planted. Our model predicts that all of the pine-growing regions of the Southern Hemisphere are capable of supporting some species of North American Scolytinae, but there are certain "hotspot" regions, southeastern Argentina, Bolivia, Chile, Peru and southwestern Australia, that appear to be suitable for a particularly large number of species. The species with the highest predicted risk of establishment were Dendroctonus valens, Xyleborus intrusus, Hylastes tenuis, Ips grandicollis, Gnathotrichus sulcatus, and Ips calligraphus. Given that global commerce is anticipated to continue to increase, we can expect that more Scolytinae species will continue to establish outside their range. Our results provide information useful for identifying a global list of potential invasive species in pine plantations, and may assist in the design of comprehensive strategies aimed at reducing pest establishment in Southern Hemisphere forest plantations.

Foraging Behavior Interactions Between Two non-Native Social Wasps, Vespula germanica and V. vulgaris (Hymenoptera: Vespidae): Implications for Invasion Success?

Vespula vulgaris is an invasive scavenging social wasp that has very recently arrived in Patagonia (Argentina), a territory previously invaded - 35 yrs earlier - by another wasp, Vespula germanica Although V. vulgaris wasps possess features that could be instrumental in overcoming obstacles through several invasion stages, the presence of preestablished populations of V. germanica could affect their success. We studied the potential role played by V. germanica on the subsequent invasion process of V. vulgaris wasps in Patagonia by focusing on the foraging interaction between both species. This is because food searching and exploitation are likely to overlap strongly among Vespula wasps. We carried out choice tests where two types of baits were presented in a pairwise manner. We found experimental evidence supporting the hypothesis that V. germanica and V. vulgaris have an asymmetrical response to baits with stimuli simulating the presence of each other. V. germanica avoided baits with either visual or olfactory cues indicating the V. vulgaris presence. However, V. vulgaris showed no preference between baits with or lacking V. germanica stimuli. These results suggest that the presence of an established population of V. germanica may not contribute to added biotic resistance to V. vulgaris invasion.

Innate Host Habitat Preference in the Parasitoid Diachasmimorpha longicaudata: Functional Significance and Modifications through Learning.

Parasitoids searching for polyphagous herbivores can find their hosts in a variety of habitats. Under this scenario, chemical cues from the host habitat (not related to the host) represent poor indicators of host location. Hence, it is unlikely that naïve females show a strong response to host habitat cues, which would become important only if the parasitoids learn to associate such cues to the host presence. This concept does not consider that habitats can vary in profitability or host nutritional quality, which according to the optimal foraging theory and the preference-performance hypothesis (respectively) could shape the way in which parasitoids make use of chemical cues from the host habitat. We assessed innate preference in the fruit fly parasitoid Diachasmimorpha longicaudata among chemical cues from four host habitats (apple, fig, orange and peach) using a Y-tube olfactometer. Contrary to what was predicted, we found a hierarchic pattern of preference. The parasitism rate realized on these fruit species and the weight of the host correlates positively, to some extent, with the preference pattern, whereas preference did not correlate with survival and fecundity of the progeny. As expected for a parasitoid foraging for generalist hosts, habitat preference changed markedly depending on their previous experience and the abundance of hosts. These findings suggest that the pattern of preference for host habitats is attributable to differences in encounter rate and host quality. Host habitat preference seems to be, however, quite plastic and easily modified according to the information obtained during foraging.

Spatial Pattern of Attacks of the Invasive Woodwasp Sirex noctilio, at Landscape and Stand Scales.

Invasive insect pests are responsible for important damage to native and plantation forests, when population outbreaks occur. Understanding the spatial pattern of attacks by forest pest populations is essential to improve our understanding of insect population dynamics and for predicting attack risk by invasives or planning pest management strategies. The woodwasp Sirex noctilio is an invasive woodwasp that has become probably the most important pest of pine plantations in the Southern Hemisphere. Our aim was to study the spatial dynamics of S. noctilio populations in Southern Argentina. Specifically we describe: (1) the spatial patterns of S. noctilio outbreaks and their relation with environmental factors at a landscape scale; and (2) characterize the spatial pattern of attacked trees at the stand scale. We surveyed the spatial distribution of S. noctilio outbreaks in three pine plantation landscapes, and we assessed potential associations with topographic variables, habitat characteristics, and distance to other outbreaks. We also looked at the spatial distribution of attacked trees in 20 stands with different levels of infestation, and assessed the relationship of attacks with stand composition and management. We found that the spatial pattern of pine stands with S. noctilio outbreaks at the landscape scale is influenced mainly by the host species present, slope aspect, and distance to other outbreaks. At a stand scale, there is strong aggregation of attacked trees in stands with intermediate infestation levels, and the degree of attacks is influenced by host species and plantation management. We conclude that the pattern of S. noctilio damage at different spatial scales is influenced by a combination of both inherent population dynamics and the underlying patterns of environmental factors. Our results have important implications for the understanding and management of invasive insect outbreaks in forest systems.