Bet-hedging in parasitoids: when optimization is not the best strategy to cope with climatic extremes.

Bet-hedging occurs when unreliable environments select for genotypes exhibiting a lower variance in fitness at the cost of a lower mean fitness for each batch of progeny. This means that at the level of the genotype, the production of mostly non-optimal phenotypes may be favored when at least some phenotypes are successful. As extreme unreliable climatic events are increasing because of climate change, it is pertinent to investigate the potential of bet-hedging strategies that allow insects to cope with climate change. Evidence for bet-hedging is scarce in most insects, including parasitoids, but the unique lifestyle and biology of parasitoids leads to the expectation that bet-hedging may occur frequently. Here, we evaluate a range of parasitoid traits for which a bet-hedging strategy could be envisioned even if bet-hedging has not been identified as such yet. Under-identification of bet-hedging in nature could have resulted from a major focus of studies on parasitoid life history evolution and foraging behavior on optimality models, predicting how mean fitness can be maximized. Most environmental factors, however, vary unpredictably. Life history and behavioral adaptations are thus expected to be affected by environmental stochasticity. In this paper, we review different aspects of parasitoid behavior, physiology, and life histories and ask the question whether parasitoid traits could have evolved under selection by environmental stochasticity.

No evidence for competition over floral resources between winter-active parasitoids and pollinators in agroecosystems.

Warming temperate winters are resulting in increased insect winter activity. With modern agroecosystems largely homogenous, characterised by low floral diversity, competitive interactions may arise between flower-visiting species, with potential implications for the ecosystem services they provide (e.g. biological control and pollination). Flower strips may be implemented during winter months to support flower-visiting insects and enhance ecosystem service provision. Employing field trials conducted in Brittany, France between 2019 and 2021 and laboratory cage experiments, the current study examined the impact of winter flower strips on aphid biological control performed by parasitoid wasps and the potential for competitive interactions between winter-active parasitoids and pollinators. Results revealed that parasitism rate was not enhanced by the presence of winter flower strips. This lack of effect was not the consequence of pollinator presence, and the current study found no effect of pollinator abundance on parasitism rate. Flower strips may thus be implemented during winter months to support nectar-feeding insects when floral resources are scarce, with no evidence of exploitative competition between pollinators and parasitoids, nor a detrimental impact on biological control provision.

Parasitoid ecology along geographic gradients: lessons for climate change studies.

One method to study the impact of climate change on host-parasitoid relationships is to compare populations along geographical gradients in latitude, altitude, or longitude. Indeed, temperatures, which vary along geographic gradients, directly shape the life traits of parasitoids and indirectly shift their populations through trophic interactions with hosts and plants. We explored the pros and cons of using these comparisons along gradients. We highlighted that the longitudinal gradients, although understudied, are well correlated to winter warming and summer heat waves and we draw attention to the impact of the increase in extreme events, which will probably be the determining parameters of the effect of climate change on host-parasitoid relationships.

Composition and Food Web Structure of Aphid-Parasitoid Populations on Plum Orchards in Chile.

By increasing plant diversity in agroecosystems, it has been proposed that one can enhance and stabilize ecosystem functioning by increasing natural enemies' diversity. Food web structure determines ecosystem functioning as species at different trophic levels are linked in interacting networks. We compared the food web structure and composition of the aphid- parasitoid and aphid-hyperparasitoid networks in two differentially managed plum orchards: plums with inter-rows of oats as a cover crop (OCC) and plums with inter-rows of spontaneous vegetation (SV). We hypothesized that food web composition and structure vary between OCC and SV, with network specialization being higher in OCC and a more complex food web composition in SV treatment. We found a more complex food web composition with a higher species richness in SV compared to OCC. Quantitative food web metrics differed significantly among treatments showing a higher generality, vulnerability, interaction evenness, and linkage density in SV, while OCC presented a higher degree of specialization. Our results suggest that plant diversification can greatly influence the food web structure and composition, with bottom-up effects induced by plant and aphid hosts that might benefit parasitoids and provide a better understanding of the activity, abundance, and interactions between aphids, parasitoids, and hyperparasitoids in plum orchards.

Landscape effects on the thermotolerance of carabid beetles and the role of behavioral thermoregulation.

Physiological thermotolerance and behavioral thermoregulation are central to seasonal cold adaptation in ectothermic organisms. For species with enhanced mobility, behavioral responses may be of greater importance in the cold stress response. Employing the carabid beetles as a study organism, the current study compared physiological thermotolerance and behavioral thermoregulation in carabid species inhabiting cereal fields in different landscape contexts, from fine grain heterogeneous "complex" landscapes to homogenous "simple" landscapes. Physiological thermotolerance was determined via measurement of the CTmin and chill coma temperature. Behavioral responses to cold temperature exposure were determined employing a purpose built arena, and thoracic temperature measured to estimate the efficacy of the behavior as a form of behavioral thermoregulation. Results revealed an influence of landscape composition on the cold tolerance of carabid beetles, although species differed in their sensitivity to landscape intensification. A reduced effect of landscape on the thermotolerance of larger carabid beetles was observed, thought to be the consequence of greater mobility preventing local acclimation to microclimatic variation along the landscape intensification gradient. Investigation into behavioral thermoregulation of the 3 largest species revealed burrowing behavior to be the main behavioral response to cold stress, acting to significantly raise carabid body temperature. This finding highlights the importance of behavioral thermoregulation as a strategy to evade cold stress. The use of behavioral thermoregulation may negate the need to invest in physiological thermotolerance, further offering explanation for the lack of landscape effect on the physiological thermotolerance of larger carabids.

Composition and structure of winter aphid-parasitoid food webs along a latitudinal gradient in Chile.

All species interact in complex antagonistic or mutualistic networks that may be driven by turnover in species composition due to spatiotemporal environmental filtering. Therefore, studying differences in insect communities along environmental gradients may improve our understanding of the abiotic and biotic factors that shape the structure of trophic networks. Parasitoids are interesting models to do so, due to their intimate eco-evolutionary relationship with their hosts. We explored the differences in cereal aphid-parasitoid food webs during the winter among nine localities in Chilean central-south valley, along a gradient of 1200 km from north (29° S) to south (40° S). We hypothesized that diapause incidence would increase in the coldest areas, resulting in a lower number of parasitoid species active during the winter. Consequently, network specialization, generality, and vulnerability indexes should increase with decreasing latitude, which implies fewer and more weakly connected links per parasitoid species through an increased fraction of basal host species. Based on the severity of winter, three areas along the explored gradient were distinguished, but clustering did not follow a clear north-south latitudinal gradient. Instead, few differences were observed in overwintering strategies, with very low levels of diapause in all localities, and no major differences in food-web composition. The major differences along the gradient were the relative abundances of the different aphid, parasitoid and hyperparasitoid species, with higher levels of spatial and temporal variation observed for the less abundant species. Our results provide a better understanding of the activity and abundance of aphid parasitoids during winter in relation to climatic conditions.

Effect of a cover crop on the aphid incidence is not explained by increased top-down regulation.

Background: Cover crops can be used as a habitat management strategy to enhance the natural enemies and their temporal synchronization with a target pest. We examined the effect of winter oat intercropping within organic plum orchards on the natural enemy abundance and seasonal dynamics on the biological control of plum aphids in spring in Central Chile. Methods: We compared the incidence and abundance of natural enemies and aphid pests from winter to the end of spring using two treatments: (1) plum trees with an oat cover crop (OCC) and (2) plum trees without a cover crop but with spontaneous vegetation (SV). We hypothesized that cover crops allow the development of winter cereal aphids, promoting the early arrival of natural enemies in spring, resulting in an earlier control of plum aphids. Results: Winter cereal aphids developed well on the OCC, and as a result, a lower plum aphid incidence in spring was observed when compared to the SV. However, the abundance of natural enemies and the parasitism rates cannot explain the positive impacts of the oat cover crop on the aphid populations as there were no differences between treatments. A potential effect of the oat due to chemical and/or physical stimuli (bottom-up effects) could help to explain these results.

Multi-scale approach to biodiversity proxies of biological control service in European farmlands.

Intensive agriculture has profoundly altered biodiversity and trophic relationships in agricultural landscapes, leading to the deterioration of many ecosystem services such as pollination or biological control. Information on which spatio-temporal factors are simultaneously affecting crop pests and their natural enemies is required to improve conservation biological control practices. We conducted a study in 80 winter wheat crop fields distributed in three regions of North-western Europe (Brittany, Hauts-de-France and Wallonia), along intra-regional gradients of landscape complexity. Five taxa of major crop pests (aphids and slugs) and natural enemies (spiders, carabids, and parasitoids) were sampled three times a year, for two consecutive years. We analysed the influence of regional (meteorology), landscape (structure in both the years n and n-1) and local factors (hedge or grass strip field boundaries, and distance to boundary) on the abundance and species richness of crop-dwelling organisms, as proxies of the service/disservice they provide. Firstly, there was higher biocontrol potential in areas with mild winter climatic conditions. Secondly, natural enemy communities were less diverse and had lower abundances in landscapes with high crop and wooded continuities (sum of interconnected crop or wood surfaces), contrary to slugs and aphids. Finally, field boundaries with grass strips were more favourable to spiders and carabids than boundaries formed by hedges, while the opposite was found for crop pests, with the latter being less abundant towards the centre of the fields. We also revealed temporal modulation-and sometimes reversion-of the impact of local elements on crop biodiversity. To some extent, these results cause controversy because they show that hedgerows and woodlots should not be the unique cornerstones of agro-ecological landscape design strategies. We point out that combining woody and grassy habitats to take full advantage of the features and ecosystem services they both provide (biological pest control, windbreak effect, soil stabilization) may promote sustainable agricultural ecosystems. It may be possible to both reduce pest pressure and promote natural enemies by accounting for taxa-specific antagonistic responses to multi-scale environmental characteristics.

Wolbachia manipulate fitness benefits of olfactory associative learning in a parasitoid wasp.

Upon encountering a host, a female parasitoid wasp has to decide whether to learn positive or negative cues related to the host. The optimal female decision will depend on the fitness costs and benefits of learned stimuli. Reward quality is positively related to the rate of behavioral acquisition in processes such as associative learning. Wolbachia, an endosymbiotic bacterium, often plays an impressive role in the manipulation of its arthropod host's biology. Here, we studied the responses of two natural Wolbachia infected/uninfected Trichogramma brassicae wasp populations to theoretically high- and low-reward values during a conditioning process and the consequences of their responses in terms of memory duration. According to our results, uninfected wasps showed an attraction response to high-value rewards, but showed aversive learning in response to low-value rewards. The memory span of uninfected wasps after conditioning by low-value rewards was significantly shorter than that for high-value rewards. As our results revealed, responses to high-quality hosts will bring more benefits (bigger size, increased fecundity and enhanced survival) than those to low-quality hosts for uninfected wasps. Infected wasps were attracted to conditioned stimuli with the same memory duration after conditioning by both types of hosts. This was linked to the fact that parasitoids emerging from both types of hosts present the same life-history traits. Therefore, these hosts represent the same quality reward for infected wasps. According to the obtained results, it can be concluded that Wolbachia manipulates the learning ability of its host, resulting in the wasp responding to all reward values similarly.

Dealing with predictable and unpredictable temperatures in a climate change context: the case of parasitoids and their hosts.

The Earth's climate is changing at a rapid pace. To survive in increasingly fluctuating and unpredictable environments, species can either migrate or evolve through rapid local adaptation, plasticity and/or bet-hedging. For small ectotherm insects, like parasitoids and their hosts, phenotypic plasticity and bet-hedging could be critical strategies for population and species persistence in response to immediate, intense and unpredictable temperature changes. Here, we focus on studies evaluating phenotypic responses to variable predictable thermal conditions (for which phenotypic plasticity is favoured) and unpredictable thermal environments (for which bet-hedging is favoured), both within and between host and parasitoid generations. We then address the effects of fluctuating temperatures on host-parasitoid interactions, potential cascading effects on the food web, as well as biological control services. We conclude our review by proposing a road map for designing experiments to assess if plasticity and bet-hedging can be adaptive strategies, and to disentangle how fluctuating temperatures can affect the evolution of these two strategies in parasitoids and their hosts.

Measuring the evolutionary potential of a winter-active parasitic wasp to climate change.

In temperate climates, as a consequence of warming winters, an increasing number of ectothermic species are remaining active throughout winter months instead of diapausing, rendering them increasingly vulnerable to unpredictable cold events. One species displaying a shift in overwintering strategy is the parasitoid wasp and biological control agent Aphidius avenae. The current study aimed to better understand the consequence of a changing overwintering strategy on the evolutionary potential of an insect population to adapt to the cold stress events, set to increase in frequency, even during milder winters. Using a parental half-sibling breeding design, narrow-sense heritability of the cold tolerance, morphology and longevity of A. avenae was estimated. The heritability of cold tolerance was estimated at 0.07 (CI95% = [0.00; 0.25]) for the Critical Thermal Minima (CTmin) and 0.11 (CI95% = [0.00; 0.34]) for chill coma temperature; estimates much lower than those obtained for morphological traits (tibia length 0.20 (CI95% = [0.03; 0.37]); head width 0.23 (CI95% = [0.09; 0.39]); wing surface area 0.28 (CI95% = [0.11; 0.47])), although comparable with the heritability estimate of 0.12 obtained for longevity (CI95% = [0.00; 0.25]). The heritability estimates obtained thus suggest that A. avenae possesses low adaptive potential against cold stress. If such estimates are indicative of the evolutionary potential of A. avenae cold tolerance, more emphasis may be placed on adaptive phenotypic plasticity at the individual level to persist in a changing climate, with potential implications for the biological control function they provide.

Suitability and Profitability of a Cereal Aphid for the Parasitoid Aphidius platensis in the Context of Conservation Biological Control of Myzus persicae in Orchards.

The use of cover crops can promote the abundance and early arrival of populations of natural enemies. Cereal cover crops between orchards rows could encourage the early arrival of the parasitoid Aphidius platensis, as they offer alternative winter hosts (e.g., Rhopalosiphum padi), enhancing the control of Myzus persicae in spring. However, the preference for and suitability of the alternative host must be addressed beforehand. To evaluate the potential of this strategy, we assessed host preference using behavioural choice tests, as well as no-choice tests measuring fitness traits, when developing on both host species. One source field for each aphid population from the above hosts was chosen. There was a clear choice for R. padi compared to M persicae, independently of the source, probably due to more defensive behaviours of M. persicae (i.e., kicks and escapes). Nevertheless, both aphid species were suitable for parasitoids' development. The female progeny developed on R. padi were larger in size, irrespective of their origin. According to our results, in peach orchards with cereals sown between peach trees during the autumn, where we expect when R. padi populations will no longer be available during spring, A. platensis should be able to switch to M. persicae.

Interspecific competition among aphid parasitoids: molecular approaches reveal preferential exploitation of parasitized hosts.

When a guild of species exploit the same limited resources, interspecific competition induces the exclusion of inferior competitors, in which case, interspecific recognition mechanisms are needed. Here, we address resource partitioning and interspecific competition among three main solitary parasitoid species attacking the same host resource, the aphid Sitobion avenae in wheat fields. Optimal host acceptance models predict that parasitoid species should prefer attacking unparasitized hosts when they are available in order to maximize their fitness, as already parasitized hosts are less valuable for laying eggs, especially for inferior competitors. Therefore, we expected the level of competition (multiparasitism) in the field to increase at low host density. By using a combination of taxonomical (determination) and molecular (PCR-based) approaches, we assessed the species of all parasitoid adults and immature stages within aphid hosts. Our results demonstrate that, early in the season, the multiparasitism rates were low, whereas they were high in the mid-late season, corresponding to an aphid density decrease over time. Moreover, parasitoid species could not have been exploiting host resources randomly and the better competitor, Aphidius ervi, seemed to be foraging preferentially on hosts already parasitized by the inferior competitor A. rhopalosiphi, even when unparasitized hosts were still available. This could be due to differences in their host detection capability, as species with a narrow host range may be better at detecting their hosts in comparison with species with a greater host range, such as A. ervi, with a greater host range within the guild. Our study suggests differences in the host exploitation of two prevalent parasitoid species through the main period of aphid colonization, which still allowed the coexistence of a third inferior competitor (A. rhopalosiphi) within the assemblage, in spite of some negative interactions (multiparasitism) and redundancies.

Plasticity in a changing world: behavioural responses to human perturbations.

Most insect species are affected by Human Induced Rapid Environmental Changes (HIREC). Multiple responses to HIREC are observed in insects, such as modifications of their morphology, physiology, behavioural strategies or phenology. Most of the responses involve phenotypic plasticity rather than genetic evolution. Here, we review the involvement of behavioural plasticity in foraging, reproduction, habitat choice and dispersal; and how behavioural plasticity modifies social behaviour and inter-specific interactions. Although important, behavioural plasticity is rarely sufficient to cope with HIREC. An increasing number of studies find species to respond maladaptively or insufficiently to various anthropogenic disturbances, and less often is large degree of plasticity linked to success.

The effect of landscape complexity and microclimate on the thermal tolerance of a pest insect.

Landscape changes are known to exacerbate the impacts of climate change. As such, understanding the combined effect of climate and landscape on agroecosystems is vital if we are to maintain the function of agroecosystems. This study aimed to elucidate the effects of agricultural landscape complexity on the microclimate and thermal tolerance of an aphid pest to better understand how landscape and climate may interact to affect the thermal tolerance of pest species within the context of global climate change. Meteorological data were measured at the landscape level, and cereal aphids (Sitobion avenae, Metopolophium dirhodum and Rhopalosiphum padi) sampled, from contrasting landscapes (simple and complex) in winter 2013/2014 and spring 2014 in cereal fields of Brittany, France. Aphids were returned to the laboratory and the effect of landscape of origin on aphid cold tolerance (as determined by CTmin ) was investigated. Results revealed that local landscape complexity significantly affected microclimate, with simple homogenous landscapes being on average warmer, but with greater temperature variation. Landscape complexity was shown to impact aphid cold tolerance, with aphids from complex landscapes being more cold tolerant than those from simple landscapes in both winter and spring, but with differences among species. This study highlights that future changes to land use could have implications for the thermal tolerance and adaptability of insects. Furthermore, not all insect species respond in a similar way to microhabitat and microclimate, which could disrupt important predator-prey relationships and the ecosystem service they provide.

Intraspecific maternal competition induces summer diapause in insect parasitoids.

Organisms often live in unpredictable environments and have to adopt life history strategies that optimize their fitness under these conditions. According to bet-hedging theory, individuals can reduce variation in fitness outcomes by investing in different strategies at the same time. For arthropods, facultative summer diapause enables survival during dry and hot periods of the year, and can be triggered by a decrease in resource abundance. However, the effect of resource depletion on diapause induction has never been disentangled from the effect of the perception of the presence of competitors. Using two solitary parasitoid species of cereal aphids as a model system, Aphidius avenae (Haliday) and Aphidius rhopalosiphi (De Stefani-Perez) (Hymenoptera: Braconidae), we tested whether (i) low absolute host density and/or (ii) high levels of parasitoid females' competition lead to maternal-induced summer diapause in parasitoid offspring. Under summer-like climatic conditions, emerging parasitoid females were (i) reared alone and exposed to different host densities (from 5 to 130 aphids), or (ii) reared together with competing females (from 2 to 20 females) and then exposed individually to 50 aphids. For both parasitoid species, low aphid densities did not induce summer diapause. However, the incidence of summer diapause increased up to a maximum of 11% with increasing levels of competition experienced by female parasitoids. More than 60% of the females produced both diapausing and nondiapausing offspring after being kept at the two highest competition densities. Such a "spreading-the-risk" strategy has likely evolved to optimize parasitoid fitness by preventing the following generation from exposure to low populations of suitable hosts and high mortality from superparasitism. These results provide the first experimental evidence of direct maternal competition-induced diapause in insects, and may change the way we apprehend the evolution of arthropod seasonal ecology, by considering intraspecific competition.

Response of life-history traits to artificial and natural selection for virulence and nonvirulence in a Drosophila parastitoid, Asobara tabida.

Co-evolution of host-parasitoid interactions is determined by the costs of host resistance, which received empirical evidence, and the costs of parasitoid virulence, which have been mostly hypothesized. Asobara tabida is a parasitoid, which mainly parasitizes Drosophila melanogaster and D. subobscura, the first species being able to resist to the parasitoid development while the second species is not. To parasitize resistant hosts, including D. melanogaster, A. tabida develops sticky eggs, which prevent encapsulation, but this virulence mechanism may be costly. Interindividual and interpopulation variation in the proportion of sticky eggs respectively allowed us to (i) artificially select and compare life-history traits of a virulent and a nonvirulent laboratory strain, and (ii) compare a virulent and a nonvirulent field strain, to investigate the hypothetical costs of virulence. We observed strong differences between the 2 laboratory strains. The nonvirulent strain invested fewer resources in reproduction and walked less than the virulent one but lived longer. Concerning the field strains, we observed that the nonvirulent strain had larger wings while the virulent one walked more and faster. All together, our results suggest that virulence may not always be costly, but rather that different life histories associated with different levels of virulence may coexist at both intra- and interpopulation levels.

Why is there no impact of the host species on the cold tolerance of a generalist parasitoid?

For generalist parasitoids such as those belonging to the Genus Aphidius, the choice of host species can have profound implications for the emerging parasitoid. Host species is known to affect a variety of life history traits. However, the impact of the host on thermal tolerance has never been studied. Physiological thermal tolerance, enabling survival at unfavourable temperatures, is not a fixed trait and may be influenced by a number of external factors including characteristics of the stress, of the individual exposed to the stress, and of the biological and physical environment. As such, the choice of host species is likely to also have implications for the thermal tolerance of the emerging parasitoid. The current study aimed to investigate the effect of cereal aphid host species (Sitobion avenae, Rhopalosiphum padi and Metopolophium dirhodum) on adult thermal tolerance, in addition to sex and size, of the aphid parasitoids Aphidius avenae, Aphidius matricariae and Aphidius rhopalosiphi. Results revealed no effect of host species on the cold tolerance of the emerging parasitoid, as determined by CTmin and Chill Coma, for all parasitoid species. Host species significantly affected the size of the emerging parasitoid for A. rhopalosiphi only, with individuals emerging from R. padi being significantly larger than those emerging from S. avenae, although this did not correspond to a difference in thermal tolerance. Furthermore, a significant difference in the size of male and female parasitoids was observed for A. avenae and A. matricariae, although, once again this did not correspond to a difference in cold tolerance. It is suggested that potential behavioural thermoregulation via host manipulation may act to influence the thermal environment experienced by the wasp and thus wasp thermal tolerance and, in doing so, may negate physiological thermal tolerance or any impact of the aphid host.

Are aphid parasitoids from mild winter climates losing their winter diapause?

Temperature is both a selective pressure and a modulator of the diapause expression in insects from temperate regions. Thus, with climate warming, an alteration of the response to seasonal changes is expected, either through genetic adaptations to novel climatic conditions or phenotypic plasticity. Since the 1980s in western France, the winter guild of aphid parasitoids (Hymenoptera: Braconidae) in cereal fields has been made up of two species: Aphidius rhopalosiphi and Aphidius matricariae. The recent activity of two other species, Aphidius avenae and Aphidius ervi, during the winter months suggests that a modification of aphid parasitoid overwintering strategies has taken place within the guild. In this study, we first performed a field survey in the winter of 2014/15 to assess levels of parasitoid diapause incidence in agrosystems. Then, we compared the capacity of the four parasitoid species to enter winter diapause under nine different photoperiods and temperature conditions in the laboratory. As predicted, historically winter-active species (A. rhopalosiphi and A. matricariae) never entered diapause, whereas the species more recently active during winter (A. avenae and A. ervi) did enter diapause but at a low proportion (maximum of 13.4 and 11.2%, respectively). These results suggest rapid shifts over the last three decades in the overwintering strategies of aphid parasitoids in Western France, probably due to climate warming. This implies that diapause can be replaced by active adult overwintering, with potential consequences for species interactions, insect community composition, ecosystem functioning, and natural pest control.

Decrease of memory retention in a parasitic wasp: an effect of host manipulation by Wolbachia?

Several factors, such as cold exposure, aging, the number of experiences and viral infection, have been shown to affect learning ability in different organisms. Wolbachia has been found worldwide as an arthropod parasite/mutualist symbiont in a wide range of species, including insects. Differing effects have been identified on physiology and behavior by Wolbachia. However, the effect of Wolbachia infection on the learning ability of their host had never previously been studied. The current study carried out to compare learning ability and memory duration in 2 strains of the parasitoid Trichogramma brassicae: 1 uninfected and 1 infected by Wolbachia. Both strains were able to associate the novel odors with the reward of an oviposition into a host egg. However, the percentage of females that responded to the experimental design and displayed an ability to learn in these conditions was higher in the uninfected strain. Memory duration was longer in uninfected wasps (23.8 and 21.4 h after conditioning with peppermint and lemon, respectively) than in infected wasps (18.9 and 16.2 h after conditioning with peppermint and lemon, respectively). Memory retention increased in response to the number of conditioning sessions in both strains, but memory retention was always shorter in the infected wasps than in the uninfected ones. Wolbachia infection may select for reduced memory retention because shorter memory induces infected wasps to disperse in new environments and avoid competition with uninfected wasps by forgetting cues related to previously visited environments, thus increasing transmission of Wolbachia in new environments.