Asian citrus psyllid, Diaphorina citri, was first detected in Cyprus in 2023. This finding initiated an eradication campaign and a classical biological control program based on the introduced ectoparasitoid Tamarixia radiata. Recent integrative taxonomic work revealed the presence of a second primary parasitoid species, Tamarixia citricola, associated with D. citri nymphs in Cypriot citrus orchards. Here, we provide a post-release monitoring update on this biological control program based on field surveys conducted in late October 2025 at citrus sites selected because of previous records of D. citri or recent reports of psyllid activity from the Cypriot Plant Health Services. Overall, adult D. citri abundance was very low; adults were absent from nearly all orchards and were detected at clearly higher levels in only one site, where 60% of tap-sampled flush shoots were positive for adult presence. Nymphs showed a similar pattern, with only sparse, low-density colonies detected across the surveyed orchards. Importantly, the site with the highest D. citri density also showed the highest nymph parasitism (96.2%), consistent with strong parasitoid-associated suppression. Of the primary parasitoids that emerged, 96.9% were T. radiata, whereas 3.1% were T. citricola. For the first time in Cyprus, we also recorded the hyperparasitoid Marietta leopardina, which reached 26.7% hyperparasitism and showed a marked association with primary parasitoids developing in third-instar nymphs. These results support the important role of T. radiata in the current classical biological control program in Cyprus and underscore the need for continued monitoring to understand better emerging trophic interactions that may influence long-term outcomes.

As core constituents of healthy diets, fruits are often cultivated in temporally stable and structurally complex ecosystems that harbor high levels of biodiversity. However, high-intensity orchard management can lessen the human and environmental health benefits of fruticulture. In the present article, we argue that increased emphasis on biological control could contribute to preventative management of fruit pests, weeds, and diseases, resulting in pesticide phasedown. Carefully calibrated orchard management can increase the provision of ecosystem services by above- and belowground biota, improve soil health, and store atmospheric carbon. When tactically integrated with agroecological measures, behavior-modifying chemicals, or digital tools, biological control helps to conserve pollinator or soil fauna, protect vertebrate communities, and improve vegetation restoration outcomes. Its implementation can, however, give rise to scientific and social challenges that will need to be explored. By resolving the adoption hurdles for biological control at scale, human society could enjoy the myriad benefits of nature-friendly fruit production.

Adult parasitoids require carbohydrate-rich foods to support longevity and nitrogenous compounds for sustained egg production. In many agroecosystems, honeydew, a sugar-rich excretion from phloem-feeding insects, serves as the most abundant carbohydrate source, especially where nectar is scarce. Although the benefits of honeydew to parasitoid fitness are well documented, it remains unclear how environmental exposure affects its nutritional quality. Here, we examined whether honeydew loses its nutritional value for parasitoids when it is exposed to the greenhouse environment during long periods of time. For this aim, we used the parasitoid Anagyrus vladimiri, a synovigenic parasitoid widely used in the biological control of Planococcus citri, an abundant phloem-feeding mealybug in citrus. We collected honeydew from P. citri and exposed it to greenhouse conditions for zero (fresh), seven, or 14 days. We then evaluated parasitoid survival and egg load of parasitoid wasps fed on the different honeydews as well as the feeding behaviour and preference across honeydews. Survival was highest on honeydew exposed to the environment for 14 days, while fresh honeydew supported the greatest egg load, suggesting that environmental exposure differentially affects the nutrients present in honeydew. Parasitoids significantly preferred honeydew exposed to the environment for 14 days over honeydew exposed for seven days. These results indicate that even after 14 days of exposure to the environment, honeydew remains as a good food source to maintain parasitoid longevity. Moreover, time of environmental exposure alters honeydew’s nutritional properties in ways that affect key parasitoid traits. These findings provide novel insights into how environmental conditions alter honeydew quality over time and indirectly influence biological control effectiveness.

Asian citrus psyllid, Diaphorina citri, is a major global pest because it is the primary vector of Candidatus Liberibacter spp., the causal agents of huanglongbing (HLB), a lethal citrus disease. Following the detection of D. citri in Cyprus in 2023, the first record of this pest in the European Union, a classical biological control program targeting this pest was initiated in spring 2024 using the parasitoid Tamarixia radiata imported from California, USA. During field surveys in summer 2024, parasitized D. citri nymphs were found both in release and non-release orchards. T. radiata was recovered from release sites, confirming its establishment. In contrast, parasitism in non-release orchards suggested the presence of native or unintentionally introduced parasitoids, or a rapid spread of T. radiata into new areas. To determine the identity of parasitoids associated with D. citri in Cyprus, an integrative approach was adopted combining field observations, molecular analyses of the COI gene, and morphological analyses. T. radiata recovered from Cyprus field sites matched reference sequences of parasitoids from California. However, other specimens were genetically and morphologically distinct and represented a new species. The new species is described here as Tamarixia citricola Hansson and Guerrieri sp. nov. Taxonomic diagnoses and characters for separating both Tamarixia species associated with D. citri are provided. Results presented here indicate the coexistence of both T. radiata (introduced) and T. citricola (likely autochthonous) in Cyprus citrus orchards. This finding has important implications for future biological control strategies and quarantine measures for D. citri in the Mediterranean basin.

Domestication often alters plant traits, leading to cascading effects on ecological interactions, particularly in tri-trophic relationships among plants, herbivores, and their natural enemies. While recent studies have investigated the influence of domestication on plant-derived food sources, its effect on guttation-a nutrient-rich exudate produced by many plants-remains unexplored. In this study, we examined the effects of guttation droplets from wild and cultivated highbush blueberries on the fitness (longevity and fecundity) and feeding preferences of three insect species from different trophic guilds: an herbivore (Drosophila suzukii), a parasitoid (Trichopria drosophilae), and a predator (Chrysoperla carnea). Additionally, we analyzed and compared the size and nutritional composition-specifically, total sugar and protein content-of guttation droplets between wild and cultivated blueberry plants. Our results indicated that guttation from wild plants enhanced the longevity of all three insect species, often surpassing that of cultivated plants and diets containing only sugar or sugar plus protein. In choice assays, all three insect species consistently preferred guttation from wild plants over that from cultivated ones. Although the guttation droplets from cultivated plants were larger, those from wild plants contained higher concentrations of sugars (six times more) and proteins (five times more), which likely contributed to the insects' enhanced fitness and preference for wild plant guttation. These findings indicate that domestication has reduced the ecological functionality of guttation in blueberries by potentially influencing tri-trophic interactions. Understanding how domestication affects plant-derived food sources like guttation could have important implications for the conservation of natural enemies in agricultural landscapes.

Crop expansion is generally followed by an increase of herbivores. Some of these herbivore species can be favoured by mutualisms with ants that can also colonise introduced crops. Here, we hypothesized that the invasive mealybug Pseudococcus longispinus has established mutualistic relationships with native or exotic ants in Mediterranean persimmon crop. This relationship could negatively impact mealybug natural enemies and facilitate the colonisation of this new crop by the mealybug. To test this hypothesis, we sampled 17 persimmon orchards from the main producing area of the Mediterranean basin across two consecutive years and three different seasons. We identified the ant complex attending Ps. longispinus, calculated the ant attendance ratios and assessed the density dependence between tending ants and Ps. longispinus at mealybug colony level. Finally, we evaluated the effect of ants on the density of Ps. longispinus and its natural enemies at tree level. Fifteen ant species were found foraging in persimmon trees, out of which 11 attended Ps. longispinus colonies. The Mediterranean native species Lasius grandis, Pheidole pallidula and Plagiolepis spp. were the most abundant ant species in the canopies, and La. grandis was the most abundant species attending Ps. longispinus. This ant tended to attend large colonies of the mealybug in spring and summer. Pseudococcus longispinus density was positively correlated with ant activity in persimmon trees, which can be partially explained by the negative effect of ants on the mealybug parasitism and the density of potential predators. This study suggests that the density and damage caused by Ps. longispinus in Mediterranean persimmon are exacerbated by the attendance of native ants that disrupt mealybug biological control. Our results reinforce the hypothesis that ant attendance is a driving factor determining the density of honeydew-producing herbivores and highlight that native ant species may facilitate the establishment and spread of exotic honeydew-producing pests in new crops.

The Asian citrus psyllid ( Diaphorina citri , Hemiptera: Psyllidae) was first detected in Cyprus in August 2023 in an orchard of Citrus x aurantium var. sinensis (orange trees) in the Phassouri area in Limassol District. Follow-up monitoring revealed the presence of D. citri in citrus orchards across all districts of the island, namely Nicosia, Larnaca, Limassol, Ammochostos and Paphos. Several demarcated areas were defined, each including an infested zone (the infested citrus orchard) and a buffer zone, encompassing the village, town or community where the infested orchard was located. The measures taken in the infested and buffer zones were mainly the application of chemical insecticides. An extensive awareness campaign was initiated by the NPPO of Cyprus, which included training all plant health inspectors in the identification of D. citri and the symptoms of huanglongbing, also known as citrus greening, a disease caused by three species of bacteria in the genus ‘ Candidatus Liberibacter’, vectored by the psyllid. Surveys continue throughout Cyprus, and in March 2024, the parasitoid wasp Tamarixia radiata , a biological control agent, was imported as part of the eradication program. Releases of T. radiata were initiated in April 2024. Citrus fruits are the second most economically important fresh product exported from Cyprus. Citrus trees can be found not only in orchards but also in urban areas, including private gardens, which complicates eradication and control efforts. Additionally, the NPPO has initiated an extensive survey to confirm the absence of huanglongbing.

Ants and phloem-feeding hemipterans have established one of the most widespread and best-known mutualisms on Earth. In this mutualism, known as trophobiosis, ants feed on honeydew excreted by phloem-feeding hemipterans and, in exchange, protect hemipterans from their antagonists. Parasitoid wasps are among the main groups of antagonists of phloem-feeding hemipterans. Like trophobiosis, the interaction between trophobiotic ants and parasitoids of phloem-feeding hemipterans has evolved over millions of years and is widely distributed both geographically and phylogenetically. Ants protect phloem-feeding hemipterans from their parasitoids in many different ways, with outcomes for parasitoids that vary from altered reproduction or development to death. Consequently, parasitoids have evolved a series of behavioural, chemical, and morphological adaptations that reduce or limit the impact of trophobiotic ants. Our review shows that research on these interactions is asymmetric and strongly biased towards certain taxa and ecosystems, mostly aphids that feed on temperate crops. It will be necessary to broaden the range of taxa and ecosystems studied to evaluate how these interactions have shaped the evolution of phloem-feeding hemipterans, their parasitoids, and trophobiotic ants. While, in general, the presence of trophobiotic ants reduces the top-down regulation of phloem-feeding hemipterans by parasitoids, recent findings suggest that the mechanisms that explain this reduction are more complex than expected. By reviewing these interactions, the limitations of past research, and the advantages of current techniques, we provide perspectives to understand: (i) the mechanisms that ants use to protect hemipterans from parasitoids; (ii) the strategies evolved by parasitoids to counteract these ants; and (iii) the multiple factors that modulate the effects of trophobiotic ants on parasitoids of hemipterans. We suggest that a better understanding of these interactions will improve the management of phloem-feeding hemipterans, which constitute one of the most damaging groups of pests to global agriculture.

Huanglongbing (HLB), also known as citrus greening disease, is one of the most destructive diseases affecting the global citrus industry. It is caused by bacteria of the Candidatus Liberibacter genus, primarily C. L. asiaticus (CLas), C. L. africanus (CLaf), and C. L. americanus (CLam), transmitted by Asian citrus psyllid [Diaphorina citri (vectors CLas)] and African citrus psyllid (Trioza erytreae [CLaf]), which likely can vector all three C.L. species. HLB causes severe symptoms in citrus trees, including leaf mottling, deformed fruits, and tree decline, leading to significant economic losses and tree death in citrus-producing regions across Asia, the Americas, and Africa. Controlling HLB progression remains challenging due to the lack of effective curative treatments and the ongoing global spread of psyllid vectors. This review provides a comprehensive overview of biological and epidemiological aspects of HLB and its vectors, focusing on recent advances in understanding of the biology of the pathogen and vector-host interactions. Additionally, we explore the case of Florida (USA), one of the most adversely affected regions, where HLB has drastically impacted commercial citrus production. The review highlights recent technological and biological advances aimed at mitigating HLB's impacts and discusses current research focused on enhancing vector control, improving plant resilience, and advancing detection methods. Finally, we address future challenges, including the need for sustainable management strategies, international cooperation, and the integration of new biotechnological tools to manage HLB-causing pathogens and disease progression, all of which are necessary to ensure the long-term sustainability of the global citrus industry.

Defensive chemicals of prey can be sequestered by some coevolved predators, which take advantage of prey toxins for their own defence. The increase in the number of invasive species in the Anthropocene has resulted in new interactions among non-coevolved predator and prey species. While novelty in chemical defence may provide a benefit for invasive prey against non-coevolved predators, resident predators with the right evolutionary pre-adaptations might benefit from sequestering these novel defences. Here, we chose a well-known system of invasive species to test whether non-coevolved predators can sequester and use toxins from exotic prey. Together with the invasive prickly pear plants, cochineal bugs ( Dactylopius spp.) are spreading worldwide from their native range in the Americas. These insects produce carminic acid, a defensive anthraquinone that some specialized predators sequester for their own defence. Using this system, we first determined whether coccinellids that prey on cochineal bugs in the Mediterranean region tolerated, sequestered, and released carminic acid in reflex bleeding. Then, we quantified the deterrent effect of carminic acid against antagonistic ants. Our results demonstrate that the Australian coccinellid Cryptolaemus montrouzieri sequestered carminic acid, a substance absent in its coevolved prey, from exotic cochineal bugs. When attacked, the predator released this substance through reflex bleeding at concentrations that were deterrent against antagonistic ants. These findings reveal that non-coevolved predators can sequester and use novel toxins from exotic prey and highlights the surprising outcomes of novel interactions that arise from species invasions.

Plant guttation is an exudation fluid composed of xylem and phloem sap secreted at the margins of leaves of many agricultural crops. Although plant guttation is a widespread phenomenon, its effect on natural enemies remains largely unexplored. A recent study showed that plant guttation can be a reliable nutrient-rich food source for natural enemies, affecting their communities in highbush blueberries. This review highlights the potential role of plant guttation as a food source for natural enemies, with a particular emphasis on its nutritional value, effects on insect communities, and potential use in conservation biological control. We also discuss possible negative implications and conclude with some open questions and future directions for research.

Honeydew is the excretion of plant-feeding hemipterans and it is one of the most abundant source of carbohydrates for parasitoids and predators in agroecosystems. Being so abundant, honeydew mediates direct and indirect interactions that affect biological control. We describe these interactions and identify honeydew-management strategies to reduce pest pressure. First, the presence of nondamaging honeydew producers in cover crops and hedges increases the efficacy of parasitoids and predators. Second, breaking the mutualism between ants and honeydew-producing pests with alternative sugar sources promotes biological control of these pests. Third, we propose to explore honeydew volatiles to attract biological control agents and repel pests, as well as to induce plant defenses. Finally, we urge reducing the use of systemic pesticides that contaminate honeydew and negatively affect biological control agents that feed on it. Overall, we propose that honeydew management is integrated in pest management programs to contribute to sustainable agriculture.

Exclusion of ground-nesting ants from tree canopies is an ant-management technique used in fruit crops. This strategy aims to disrupt the symbiotic relationship between ants and honeydew-producing herbivores, which are protected by ants, and enhance the biological control of these herbivores. Here, we evaluated this strategy to enhance the biological control of mealybugs in persimmon, but obtained an unexpected outcome. Using a randomized block design in a commercial persimmon orchard during two consecutive years, we demonstrated that physical barriers based on sticky bands excluded native ground-nesting ant species from persimmon trees and reduced the protection that these ants provided to mealybugs. The exclusion of native ground-nesting ants also increased the density and efficacy of the biological control agents of the mealybug. However, physical barriers favoured the establishment of the exotic canopy-nesting ant species Cardiocondyla obscurior. This ant species established its colonies under the sepals of persimmon fruit, and its abundance increased by more than 10 times in trees with physical barriers. Our results show that the exclusion of native ground-nesting from fruit trees can promote the control of invasive mealybugs, but may also facilitate the establishment and spread of exotic canopy-nesting ant species, hence increasing their invasive potential. Further research should evaluate alternative ant management strategies to enhance biological control of honeydew-producing herbivores without benefiting exotic canopy-nesting ant species.

BACKGROUND Chaetanaphothrips orchidii is an invasive thrips of tropical origin that was detected in 2016 in Spanish citrus, where it can damage up to 70% of the fruit. Pupation site and emergence rates are key biological traits for thrips management that are unknown for C. orchidii. Here, we determined the pupation site and period of C. orchidii in citrus and evaluated the effect of soil moisture on adult emergence. RESULTS A two-year field study showed that C. orchidii pupated in the soil from May to December in commercial citrus orchards. Chaetanaphothrips orchidii emergence was very low compared to other harmful thrips species in citrus. Using D/E traps, we demonstrated that the thrips emerged mainly from wet areas near drip irrigation emitters during the summer, and its emergence was strongly related to the soil water content. A laboratory experiment confirmed that C. orchidii did not emerge at RH below 70%, and its emergence peaked at 97% RH. CONCLUSIONS Our results have important implications for the sustainable management of C. orchidii because soil moisture is very low in Mediterranean citrus during summer, except in areas near drip irrigation emitters. Therefore, these and other potential high-humidity areas should be considered crucial targets to manage this pest in citrus during summer, reducing cost and labor.

Herbivore feeding often increases secondary metabolite production in plants. These herbivore-induced plant proteins might end up in honeydew excreted by phloem-feeding insects. This is important because honeydew is one of the most abundant and accessible carbohydrate sources for natural enemies in many agroecosystems and these proteins can thus mediate many tri-trophic interactions. Here, we hypothesized that defensive metabolites induced in the phloem by herbivory accumulate in the honeydew excreted by phloem-feeding insects and, consequently, affect the fitness of the herbivores' natural enemies that feed on it. We used a tri-trophic system consisting of citrus plants, the mealybug Planococcus citri and its primary parasitoid Anagyrus vladimiri. First, we assessed A. vladimiri fitness when fed on P. citri honeydew. We then collected honeydew of seven phloem-feeding insects, including P. citri, and analysed their protein content. Finally, we analysed the effect of superoxide dismutase (SOD), an antioxidant enzyme associated with plant defences that was commonly found in the analysed honeydews, on A. vladimiri fitness. The fitness of A. vladimiri increased when fed on honeydew compared to a sucrose-based diet, demonstrating that honeydew can contain compounds that benefit natural enemies. Proteomic analyses showed that defence-related plant proteins were present in honeydew of seven phloem-feeding insects analysed. Among these, the enzyme SOD was present in honeydew of all of them. Moreover, the levels of SOD were 10-fold higher in the phloem of plants infested by P. citri than in that of uninfested plants. SOD was also actively excreted in P. citri honeydew, and we proved that it increases the fecundity of the parasitoid A. vladimiri. We conclude that enzymatic proteins induced by herbivory in the phloem and involved in plant defence, accumulate in the honeydew excreted by phloem-feeding insects and, contrary to the current paradigm, at least some of these can have positive effects on the third trophic level.Read the free Plain Language Summary for this article on the Journal blog. Al alimentarse de las plantas, los insectos herb & iacute;voros suelen inducir en & eacute;stas la producci & oacute;n de metabolitos secundarios. Estos compuestos pueden terminar en la melaza que excretan los propios insectos que se han alimentado del floema. Esto es importante porque la melaza es una de las fuentes de carbohidratos m & aacute;s abundantes y accesibles para los enemigos naturales en la mayor & iacute;a de agroecosistemas y, por tanto, estos metabolitos pueden modular numerosas interacciones tritr & oacute;ficas. En este estudio, se plante & oacute; si los metabolitos inducidos por los insectos herb & iacute;voros que se alimentan del floema se acumulan en la melaza que excretan y, en consecuencia, afectan los par & aacute;metros biol & oacute;gicos de los enemigos naturales que se alimentan de esta melaza. Las especies seleccionadas para este estudio fueron los c & iacute;tricos como plantas, la cochinilla Planococcus citri como herb & iacute;voro que se alimenta de floema y excreta melaza y su parasitoide primario Anagyrus vladimiri. Primero, se evalu & oacute; la longevidad y fecundidad de A. vladimiri cuando se aliment & oacute; de melaza de P. citri. Luego, se recolect & oacute; melaza de siete especies de insectos que se alimentan del floema, incluyendo P. citri, y se analiz & oacute; su contenido de prote & iacute;nas. Finalmente, se evalu & oacute; el efecto de la prote & iacute;na super & oacute;xido dismutasa (SOD) en la longevidad y fecundidad de A. vladimiri. Esta enzima, de reconocido efecto antioxidante y asociada con las defensas de las plantas, se encontr & oacute; en todas las melazas analizadas. La longevidad y fecundidad de A. vladimiri fue mayor cuando se aliment & oacute; de melaza que cuando lo hizo de una dieta basada s & oacute;lo en az & uacute;cares, demostrando que la melaza puede contener otros compuestos que benefician a los enemigos naturales. Los an & aacute;lisis prote & oacute;micos mostraron que hab & iacute;a prote & iacute;nas vegetales relacionadas con la defensa en la melaza excretada por las siete especies de insectos. Entre ellas, la enzima SOD, con niveles expresi & oacute;n diez veces m & aacute;s altos en el floema de plantas infestadas por P. citri que en plantas no infestadas. Al a & ntilde;adir SOD a una dieta basada en az & uacute;cares, la fecundidad del parasitoide A. vladimiri aument & oacute; de forma significativa. Este estudio concluye que las prote & iacute;nas enzim & aacute;ticas inducidas en la defensa de las plantas, se acumulan en la melaza excretada por insectos que se alimentan del floema y, contrariamente al paradigma actual, algunas de estas enzimas tienen efectos positivos en el tercer nivel tr & oacute;fico. Read the free Plain Language Summary for this article on the Journal blog.image

1. The simplification of agricultural landscapes has been associated with an increase in pest pressure. While monocultures increase the resources available for pests and may facilitate their dispersion, the lack of non-crop habitats may reduce the resources available for pest natural enemies. Herein, we tested which of these hypotheses, namely 'resource concentration' and 'natural enemies', can better explain the abundance of invasive mealybug pests in two subtropical fruit crops.2. For this aim, 17 persimmon orchards and 16 citrus orchards were sampled during three different seasons across two consecutive years. Using a model selection approach, we assessed the effects of the surrounding landscape (proportion of focal crops and semi-natural habitats at different distances) and inter-row ground cover vegetation on the abundance of mealybugs and their natural enemies.3. The proportion of focal crop in the landscape increased the abundance of mealybugs attacking both crops. This effect was found at closer distances (up to similar to 600 m) in citrus and at both closer and further distances (up to 1250 m) in persimmon.4. Non-crop habitats, both surrounding semi-natural habitats and ground cover vegetation, decreased the abundance of mealybugs by increasing the activity of their parasitoids in persimmon. Conversely, non-crop habitats did not decrease the abundance of the main mealybug species attacking citrus, likely because this mealybug species was not attacked by native or naturalized parasitoids.5. Synthesis and applications. Our models show that the increase in habitat heterogeneity at local and landscape scales can reduce the abundance of invasive mealybugs in subtropical crops via 'resource concentration' and 'natural enemies' mechanisms. Therefore, habitat diversification strategies should be considered in the conservation biological control of invasive mealybugs. Importantly, our findings also show that the presence of efficient natural enemies is critical to maximize their control through habitat diversification strategies.

Mealybugs (Hemiptera: Pseudococcidae) are the main pest of persimmon in Spain, the second producer in the world. In order to develop an Integrated Pest Management (IPM) program, it is necessary to identify the main mealybug species, determine their phenology, and develop tools to predict damage. To do this, we sampled 17 orchards from the main persimmon producing area in Spain over two years. Pseudococcus longispinus (Targioni-Tozzeti) was the most abundant and widely distributed species. This mealybug species completed three generations per year and reached peak density just before harvest. Fruit infestation at harvest was highly correlated with mealybug density in spring and summer. The estimated thermal constants to complete development and one generation were 512.5 and 956.3 degree days, respectively. Based on climate change predictions, crop damage caused by the third generation of P. longispinus will increase in 2040 and the mealybug will complete a fourth generation by 2080. Pseudococcus longispinus has become the main pest for Mediterranean persimmon and damage produced by this mealybug may be exacerbated by climate change. This work pro -vides essential data to design a sampling protocol and determine intervention times and thresholds against this mealybug.

•Natural parasitism of Pseudococcus longispinus was highly variable among Mediterranean persimmon orchards.•We tested whether hyperparasitoids affected the biological control of this invasive mealybug.•Three hyperparasitoid species attacked the main parasitoid Anagyrus fusciventris in persimmon.•Hyperparasitism did not affect the population growth rate of P. longispinus within the same year.•Several strategies are proposed to enhance biological control of P. longispinus.